Overview of Rust Fungi - Plant Management Network€¦ · •ca. 7000 species ... and comprise 22...

Overview of Rust Fungi

M.C. Aime Department of Plant Pathology and Crop Physiology

Louisiana State University Agricultural Center

Overview

What defines the rusts? How are they different from other pathogen groups? Molecular phylogenetics.

What are rusts?

• Pucciniales (Uredinales) in Pucciniomycotina (Basidiomycota) • ca. 7000 species (Kirk et al. 2008) – the largest group of phytopathogens ca. 25% of all described Basidiomycota ca. 8% of all described Fungi • all parasites on plants including many cultivated crops varying degrees of host specificity • Rusts figure prominently as exotic invasives (e.g., Soybean rust, UG99,

sugarcane rust) and comprise 22 of the 52 fungi on the USDA Animal and Plant Health Inspection Service Regulated Plant Pests list (2000)

• only a few have been successfully cultured in vitro on artificial media (i.e., obligate biotrophs)

• Huge genomes [ca. 90 Mbp (haploid) Cronartium1 – 400+ Mbp (e.g. Uromyces2)]

• germination of basidia from teliospores (=probasidium) • lack basidiocarps; lack clamp connections; simple septal pores • complex life cycle (most complex in Fungi?) up to 5 stages (& 7 morphologically distinct spores)

most species need two different plants (taxonomically unrelated) to complete their life cycle (i.e., heteroecious)

life cycle data missing for many

What are rusts?

1Andersen et al. 2010. Mycologia 102(6):1295-1302; 2 Eilam et al. 1994. Phytopathology 84:728-735.

Dictionary of the Fungi, 8th (1996):Dictionary of the Fungi, 8th (1996): Teliomycetes

Uredinales (14 families, 7000 spp.) Septobasidiales

Dictionary of the Fungi, 9th (2001):Dictionary of the Fungi, 9th (2001): Urediniomycetes

Uredinales (15 families, 6929 spp.) Septobasidiales & 3 additional orders (Agaricostilbales, Atractiellales, Microbotryales)

Dictionary of the Fungi, 10th (2008):Dictionary of the Fungi, 10th (2008): Pucciniomycotina

Pucciniales (14 families, 7798 spp.) 9 classes, 18 orders, 36 families

Classification

•• “Primitive” ascomycete “Primitive” ascomycete TaphrinaTaphrina E.g., D.B.O. Savile 1971 and earlier authors “The rust fungi are

fully parasitic and arose from parasitic ancestors.”

• Septobasidium Septobasidium and and UredinellaUredinella E.g., J.N. Couch 1937 – similarities in probasidia

• AuriculariaceousAuriculariaceous “rusts” (“rusts” (Eocronartium, Eocronartium, InsolibasidiumInsolibasidium, , JolaJola, ,

HerpobasidiumHerpobasidium, , PlatycarpaPlatycarpa, , and and XeonogloeaXeonogloea)) E.g., Atkinson 1902; Y. Hiratsuka 1990 – probasidia & simple

septa; basidiocarp formation and lack of key anamorphs

What are rust relatives?

Image: R.J. Bandoni. 1995. Studies in mycology 38:13-27

Pucciniomycotina Combined nuc-lsu & nuc-ssu

8 classes, 18 orders

Aime et al. 2006. An overview of the higher-level classification of Pucciniomycotina based on combined analyses of nuclear large and small subunit rDNA sequences. Mycologia 98:896-905

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Photo: Daniel Henk

Photos: © Matthias Lutz: Bio Pro website




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Oberwinkler F. 1982. In: Wells (Ed). Basidium and Basidiocarp. Pp. 9-35.

Photo: David McLaughlin in Aime et al. 2006

PlatygloealesPlatygloeales phytoparasites Eocronartium Jola Insolibasidium Herpobasidium Platygloea

• ca. 150 genera • 13 families sensu Cummins & Hiratsuka (2003) • Spermagonia, teliospores, basidium germination

Pucciniales

George Baker Cummins 1904-2007

Photo: © Mycological Society of America 2011.

Pucciniastraceae (9) Most fern rusts; Naohidemyces

Coleosporiaceae (3) Coleosporium, Chrysomyxa

Cronartiaceae (2) Cronartium, Endocronartium

Mikronegeriaceae (3) Bllastospora, Mikronegeria

Melampsoraceae (1) Melampsora

Phakopsoraceae (13) Phakopsora, Cerotelium, Dasturella

Chaconiaceae (10) Hemileia, Olivea, Ochropsora

Uropyxidaceae (14) Tranzschelia, Prospodium, Desmella

Pileolariaceae (4) Uromycladium, Pileolaria

Raveneliaceae (22) Sphenospora, Triphragmium

Phragmidiaceae (11) Gymnoconia, Gerwasia, Kuehneola

Pucciniaceae (16) Uromyces, Gymnosporangium

Pucciniosiraceae (9) Dietelia, Ceatocoma

Puccinia mariae-wilsoniae U-005 Claytonia (Portulaceae I-III) MD Puccinia mariae-wilsoniae MCA2497 Claytonia (Portulaceae) MD

Puccinia lagenophorae U-384 (#569) Asteraceae (Osteocarpum) South Africa Puccinia lagenophora U-128 Senecio (I-III Asteraceae) Tunisia Puccinia lagenophorae U-107 Senecio (I, III Asteraceae) Argentina Puccinia violae U-023 Viola (Violaceae II-III) Norway

Puccinia violae U-269 Viola (Violaceae II-III) NY Uromyces sp. MCA 2870 Juncaceae (Juncus? III) VA Puccinia? U397 Asteraceae MD

Puccinia? U-390 Asteraceae MD Puccinia cf. violae U-4 Violaceae (Viola cucullata )-I, autoecious) MD Uromyces vignae? U-015 Macroptilium (Fabaceae II) Argentina

Uromyces vignae JRH-472 Fabaceae (Vigna II-III) Puerto Rico Uromyces vignae JRH-468 Fabaceae (Vigna II-III) Puerto Rico

Uromyces vignae U-125 Vigna (Fabaceae II) C.R. Uromyces vignae U-126 Vigna (Fabaceae II) C.R.

Uromyces appendiculatus U-122 Phaseolus (II Fabaceae) C.R. Uromyces appendiculatus U-123 Phaseolus (II Fabaceae) Mexico Uromyces appendiculatus U-00123 Phaseolus (Fabaceae) Mexico

Uromyces neotropicalis sp.nov MCA2533 Cucurbitaceae II,III Ecuador Dietelia mesoamericana sp.nov. U-320 (W1909) Asteraceae (M. micrantha microcyclic I) Mexico

Dietelia portoricensis u-322 (no #) Asteraceae (Mikania micrantha I-microcyclic) Costa Rica Puccinia granularis U-443 Geraniaceae (Pelargonium O,I,II,III) South Africa

Puccinia granularis U-444 (#581) Geraniaceae (Pelargonium II,III) South Africa Puccinia granularis U-444 (#581) Geraniaceae (Pelargonium II,III) South Africa

Puccinia granularis U-443 (#580) Geraniaceae (Pelargonium O,I,II,III) South Africa Puccinia spegazzinii JRH-226 Asteraceae (Mikania micrantha III) Honduras

Puccinia spegazzinii U-319 (W2102) Asteraceae (Mikania micrantha microcyclic III) Peru Puccinia spegazzinii U-315 (W2020) Asteraceae (Mikania micrantha IIImicrocyclic) Argentina

Dietelia mesoamericana sp. nov. U-321 (mislabelled as U143) Mikania Mexico Puccinia hydrocotyles U-079 Hydrocotyle (II-III Apiaceae) AL Puccinia? MCA-2864 Asteraceae (Helianthus) VA Puccinia pimpinellae? MCA2576 Apiaceae( Osmorhiza I) ID

Puccinia pimpinellae U-43 Osmorhiza (Apiaceae III) UT Puccinia pimpinellae MCA2604 Osmorhiza (Apiaceae III) TN Puccinia berkheyicola U-314 Berkheya (Asteraceae) South Africa

Puccinia notobasidis U-229 Cirsium (Asteraceae) Turkey Puccinia variabilis var. lapsanae U-37 Asteraceae (Lapsana II,III) UT

?? U-412 Asteraceae (Youngia) Guam ?? JRH199 Wedelia (Asteraceae) Hondorus (inc seq)

Puccinia menthae MCA 2601 Monarda (Lamiaceae II) TN Puccinia menthae U-343 Origanum (Lamiaceae II) Argentina Puccinia menthae MCA 2601 Monarda (Lamiaceae II) TN

Puccinia marylandica MCA2602 Sanicula (Apiaceae II-III) TN Puccinia marylandica MCA2606 Sanicula (Apiaceae II-III?) TN

Puccinia commelinae U-049 Commelina (Commelinaceae II) Guyana Puccinia synedrellae JRH152 Synedrella (III Asteraceae) Costa Rica Puccinia cnici-oleracei GUY-6 Asteraceae (Emilia sonchifolia III) Guyana Puccinia synedrellae JRH-152 Asteraceae (Synedrella nodiflora III) Costa Rica Puccinia cnici-oleracei JRH-204 Asteraceae (Emilia fosbergii III) Honduras Puccinia cnici-oleracei JRH-490 Asteraceae (Emilia fosbergii III) Puerto Rico

Puccinia MCA-2864 Asteraceae (Helianthus) VA Uromyces ficariae AF426204 Ranunculus (Ranunculaceae)

Puccinia sp. MCA2575 Asteraceae Idaho Puccinia sp. MCA-2575 Asteraceae (II) ID

Puccinia chondrillina U-202 Asteraceae (Chondrilla) Turkey [BPI extraction] Puccinia balsamorrhizae U-332Asteraceae (Balsamorhiza III?) B.C. Puccinia balsamorrhizae U-354 Asteraceae (Balsamorhiza) ID Puccinia balsamorrhizae U-355 Asteraceae (Balsamorhiza) ID

Miyagia pseudosphaeria U-63 (BPI) Asteraceae (Sonchus oleraceus) Puccinia calcitrapae v. centaureae U-201 Asteraceae (Centaurea depressa) Turkey [BPI extraction]

Puccinia calcitrapae U-218 Carduus (Asteraceae) Turkey Puccinia calcitrapae U-218 Asteraceae (Carduus nutans) Turkey Puccinia carthami complex MCA-2881 Asteraceae (Cirsium?) Costa Rica

Puccinia acroptili U-224(confirmed no.) Asteraceae (Acroptilon) Turkey Puccinia acroptili U-578 Asteraceae (Acroptilon repens III) NM Puccinia acroptili U-135 (Dana Berner 02-048) Asteraceae (Acroptilon repens II-III?) Turkey Puccinia acroptili U-224 Asteraceae (Acroptilon repens) Turkey Puccinia acroptili U-577 Asteraceae (Centaurea repens) NM Puccinia U-203 Asteraceae (Cardueae II-III) Turkey [host & rust confirmed from BPI] Puccinia punctiiformis U-225 Cirsium (Asteraceae) Turkey

Puccinia punctiiformis U-225 Asteraceae (Cirsium arvense) Turkey Puccinia calcitrapae U-164 Asteraceae (Carthamus tinctorius II-III) Argentina

Puccinia centaurea U-278 (FDWSRU 04-010) Asteraceae (Centaurea hybrid) OR Puccinia calcitrapae v. centaureae U-140 (Dana Berner 03-051) Asteraceae (Centaurea macedonica) Greece

Puccinia calcitrapae v. centaurea U-209 Asteraceae (Centaurea calcitrapa) Turkey Puccinia carthami U-323 Asteraceae (Carthamus tinctorius III) Oman

Puccinia atropuncta MCA2609 Veratrum (Liliaceae III) TN ?? U492 Antirrhinum (Scrophulariaceae) Germany

?? MCA2529 Geranium (Geraniaceae II) Ecuador Endophyllum stachytarphetae GUY-55 Stachytarpheta (III Verbenaceae)

?? MCA2558 Phragmites? (Poaceae) Ecuador Endophyllum cassiae U-525 Fabaceae (Cassia III?) Nigeria

?? JRH-466 Cyperaceae Puerto Rico ?? U-265 Cyperus (Cyperaceae II?) Guam

Puccinia cnici-oleracei JRH-200 Asteraceae (Melampodium divaricatum III) Honduras Puccinia cnici-oleracei JRH-207 Asteraceae (Malampodium divaricatum III) Honduras

Puccinia ursiniae U-463 (#590) Asteraceae (Ursinia) South Africa Puccinia tanaceti U-040 Artemisia (Asteraceae II-III) UT

Puccinia rubefaciens? U-330 Galium (Rubiaceae III?) B.C. Canada Puccinia medellinensis JRH-154 Lamiaceae (Hyptis II) Costa Rica Puccinia melampodii JRH138 Emilia (Asteraceae III) Costa Rica

Uromyces rumicis U-227 Rumex (Polygonaceae) Turkey Uromyces lespedezae-procumbentis U-007 Trifolium (Fabaceae III) MD Puccinia convolvuli? MCA2778 Calystegia (Convolvulaceae II) MD

Puccinia convolvuli U-068 Calystegia (II-III Convolvulaceae) MD Puccinia scleriicola GUY-57 Scleria (I Cyperaceae)

Puccinia ibrae JRH-228 Convolvulaceae (Ipomoea I-III) Honduras Puccinia thaliae JRH141 Canna (Cannaceae II) Costa Rica Puccinia thaliae GUY-3 Canna (Cannaceae II) Guyana

Puccinia kalchbrenneriana U-312 (#560) Asteraceae (Helichrysum III?) South Africa Puccinia kalchbrenneriana U-313 (#555) Asteraceae (Helichrysum III?) South Africa

Puccinia kalchbrenneri? U-258 (#539) Asteraceae (Helichrysum III) South Africa Uromyces colombianus JRH-145 Asteraceae (Melanthera II-III) Costa Rica

Uromyces psoraleae U-029 Psoralidium (Fabaceae I-III) UT Uromyces sp. JRH-482 Cucurbitaceae (Cayaponia) Puerto Rico

Puccinia gigantea AF426198 Epilobium (Onagraceae) Puccinia enceliae JRH-166 Asteraceae (Tithonia II-III) Costa Rica Puccinia thunbergiae U-260 (543) Acanthaceae (Thunbergia O,I,III) South Africa

Puccinia hydrocotyles U-105 Hydrocotyle (Apiaceae) Argentina Uromyces galegae U-242 Galega (Fabaceae) Turkey

Uredo cajani U-119 Cajanus (Fabaceae II) Honduras Uredo cajani MCA-2895 Fabaceae (Cajanus) Costa Rica Uredo cajani JRH-489 Fabaceae (Cajanus II) Puerto Rico Uromyces dolicholi JRH-469 Fabaceae (Rhynchosia II) Puerto Rico Uromyces scutellatus U-204(presumed) Euphorbiaceae (Euphorbia) Turkey

Uromyces euphorbiae GUY-7 Euphorbia (II Euphorbiaceae) Uromyces pisi AF426201 Euphorbia

Uromyces pisi-sativi? MCA2377 Euphorbia (Euphorbiaceae) Italy Uromyces? U-471 Euphorbiaceae (Euphorbia) Germany

Endophyllum euphorbiae-sylvaticae AF426200 Euphorbia Puccinia abrupta JRH-146 Viguiera (Asteraceae II-III) Costa Rica

Uromyces viciae-fabae AF426199 Vicia (Fabaceae) Puccinia sp.nov.? U-210 Tragopagon (Asteraceae I-III) Turkey

Puccinia stoboeae U-259 (#550) Asteraceae (Berkheya O,I,II) South Africa Uromyces hedysari-paniculati U-124 Desmodium (Fabaceae II-III ) Honduras

Uromyces hedysari-paniculati U-052 Desmodium (Fabaceae II) Guyana Aecidium sp. U-211 Salsola (Chenopodiaceae I) Turkey

Uromyces ciceris-arietini U-183 Cicer (Fabaceae II-III ) SD Uromyces junci AF426203 Pulicaria (Asteraceae)

Puccinia? MCA2593 (Asteraceae O?) ID Puccinia diplachnis MCA2375 Cirsium (I Asteraceae) Switzerland {T=Poaceae} Puccinia verruca U-130 Centaurea (Asteraceae III) Tunisia

Puccinia collinsiae U-044 Collinsiae (Scrophulariaceae) UT Uromyces trifolii-repentis U-207 Trifolium (Fabaceae) Turkey

Puccinia urticae-acutiformis AF426202 Carex (Cyperaceae) Puccinia hemerocallidis U498 Hemerocallis (Lilioaceae) TX Puccinia hemerocallidis U-072 Hemerocallis (II Liliaceae) AL Puccinia hemerocallidis U-073 Hemerocallis (II Liliaceae) AL Uromyces euryopsidicola U-381 (#575) Asteraceae (Euryops III) South Africa

Uromyces euryopsidicola U-216 (#295) Asteraceae (Euryops) South Africa Uromyces euryopsidicola U-261 (#295) Asteraceae (Euryops) South Africa Uromyces euryopsidicola U-216 (#295) Asteraceae (Euryops) South Africa

Puccinia sparganioides U-368 Poaceae (Spartina II,III) NY Puccinia buxi U-418 Buxus (Buxaceae) Georgia

Puccinia tillandsiae U419 Tillandsia (Bromeliaceae) Guatemala Puccinia arechaveletae MCA2430 Serjania (Sapindaceae) Guyana Puccinia arechavelaete GUY-66 Serjania (III Sapindaceae) Puccinia arechavaletae JRH175 Unvillea (II-III Sapindaceae) Costa Rica

Puccinia physalidis U-189 Physalis (Solanaceae III) ND Uromyces ixiae U452 Romulea (Iridaceae) S. Africa Uromyces transversalis U-386 Iridaceae (Gladiolus) Mexico

Puccinia hordei MCA2391 (Poaceae) CA Puccinia? U-288 Poaceae (Pennisetum II) Guam Uromyces setaria-italicae JRH220 Melinis (II Poaceae) Hondorus

Uromyces setaria-italicae JRH205 Panicum (II Poaceae) Hondorus Uromyces setaria-italicae JRH202 Panicum (II Poaceae) Honduras

Puccinia gouaniae JRH159 Gouania (II Rhamnaceae) Costa Rica Puccinia arundinariae? MCA2494 Arundinaria (Poaceae) VA

Puccinia purpurea JRH198 Sorghum (Poaceae II-III) Honduras Puccinia purpurea JRH213 Sorghum (Poaceae II) Hondurus

Puccinia purpurea JRH218 Sorghum (Poaceae II) Hondurus Puccinia duthiae GUY8 Bothriochloa (Poaceae II) Guyana

Puccinia duthiae U-264 Dichanthium (Poaceae II) Guam Puccinia exhauriens U-538 (#600) Oleaceae (Jasminum) South Africa

Puccinia exhauriens U-539 (#605) Oleaceae (Jasminum) South Africa Puccinia moraeae U462 Moraea (Iridaceae) S. Africa Uromyces massoniae? U-446(#586) Liliaceae (Massonia II-III) South Africa Uromyces salsolae U-134 Salsola (Chenopodiaceae) Syria

Puccinia sp. U-006 Allium? (Liliaceae III) MD Uromyces aloes U-00372 (#567) Aloaceae (Aloe) South Africa

Puccinia obliquo-septata MCA2431 Olyra (Poaceae) Guyana Puccinia obliquo-septata GUY-10 Olyra (II Poaceae)

Puccinia peperomiae JRH192 Piper (II Piperaceae) Costa Rica Puccinia fallax MCA2305 Palicourea (Rubiaceae II-III) Guyana Puccinia fallax GUY-58 Palicourea (II Rubiaceae)

Sphenospora kevorkianii U-10 Orchidaceae (Stanhopea II) Peru Puccinia coronata U-583 Poaceae (Avena) ND Puccinia coronata U400 Festuca (Poaceae II, III) VA Puccinia coronata U-244 Rhamnus (Rhamnaceae O-O) ND Puccinia coronata U-245 Scolochloa (Poaceae III?) ND

Puccinia coronata AF426207 Rhamnus (Rhamnaceae) Puccinia pampeanum U-109 Salpichroa (III Solanales) Argentina Endophyllum pampeanum U-108 Salpichroa (III Solanales) Argentina

Uromyces gageae AF426208 Gagea (Liliaceae) Aecidium wiborgiae U-380 (#576) Fabaceae (Wiborgia) South Africa Aecidium wiborgiae U-379 (#573) Fabaceae (Wiborgia) South Africa

Uromyces sp. MCA2383 Lilium (III Liliaceae) Switzerland Puccinia mogiphanis U-111 Alternanthera (Amaranthaceae) Nigeria Puccinia stonemanniae U-310 Thesium (Santalaceae III?) South Africa

Puccinia sorghi U-362 Poaceae (Zea mays II, III) VA Uromyces polygoni U-214 Polygonum (Polygonaceae) Turkey Uromyces polygoni U-214 Polygonum (Polygonaceae) Turkey

Puccinia mogiphanis JRH 180 Amarantaceae (Alternanthera II) Costa Rica Puccinia cynodontis JRH 219 Cynodon (Poaceae II-III) Hondorus Aecidium kalanchoe U-18 Kalanchoe

Puccinia horiana U-391 Asteraceae (Chrysanthemum III) DE Puccinia horiana U-392 Asteraceae (Chrysanthemum III) MD Puccinia horiana U-070 Chrysanthemum (Asteraceae) NJ Aecidium sp. U-465 (#579) Fabaceae (Lebeckia) South Africa

Puccinia arenariae U142 Dianthus (Caryophyllaceae) Ecuador Puccinia graminis U-478 Berberis (Berberidaceae) Germany Puccinia.poarum FROM BRANDON on Tussilago (iii on Poaceae)

Puccinia MCA-2882 Asteraceae? Costa Rica Puccinia MCA-2886 Asteraceae? Costa Rica

Puccinia graminis (as Aecidium sp.) PDD78317 Berberis I N.Z. Puccinia graminis U-469 Agropyron (Poaceae) Germany Puccinia graminis U-470 Agropyron? (Poaceae) Germany Puccinia graminis U-470 Poaceae (Agropyron?) Germany

Uredo baruensis GUY-71 Sapotaceae (Chrysophyllum II) Guyana [IDd as Coleosporium on Merremia (Convolvulaceae)] JRH-484 P.R. (seq. confirmed 2X) Sphaerophragmium acacia JRH-486 Fabaceae (Albizia II-III) Puerto Rico

Sphaerophragmium acaciae JRH-470 Fabaceae (Albizia II) Puerto Rico Pileolaria toxicodendri AY 745698 (AFTOL ID 988)

Pileolaria toxicodendri u-607 Anacardiaceae (Toxicodendron) MN Puccinia tetragonia U-442 Aizoaceae (Tetragonia) South Africa

?? U-476 Allium (Liliaceae) Germany Uromyces albucae U-447 (#591) Liliaceae (Albuca) South Africa Puccinia dispersa U-234 Anchusa (Boraginaceae) Turkey Puccinia? MCA-2885 ?? Costa Rica

Chrysocyclus cestri U-156 Cestrum (Solanaceae III) Argentina Puccinia smilacis U-12 Smilacaceae (Smilax II-III) MD Puccinia smilacis U-393 Smilacaceae (Smilax) MD

Stereostratum corticiodes U-027 Bambusa (Poaceae II-III) China Puccinia asarina MCA2374 Asarum (III Aristolochiaceae) Italy

Puccinia pampeana JRH135 Capsicum (Solanaceae II-III aecioid) Costa Rica Endophyllum macowanii U-255 Rhamnus (Rhamnaceae III?) South Africa

Puccinia thlaspeos U-054 Boechera (III Brassicaceae) WA Uromyces bulbinis BRIP-28988 Liliaceae (Bulbine) Australia

Uromyces heteromorphae U-309 Peucedanum (Apiaceae) South Africa Uromyces trollipii U-445 Zygophyllaceae (Zygophyllum I, III) South Africa

Endophyllum osteospermi from Alan Wood South Africa Endophyllum elytropappi U254 Elytropappus (Asteraceae III?) South Africa

Puccinia magnusiana PUR-19 Ranunculaceae (Clematis O,I) Italy Puccinia festucae MCA2380 Lonicera (I Caprifoliaceae) Italy Puccinia hordei U-129 Taeniatherum (Poaceae III) Turkey Uromyces zygadeni MCA2577 Zigadenus (Liliaceae I-III) Idaho

Puccinia phragmitis U-495 Poaceae (Phragmites) Austria Puccinia phragmitis U-423 Poaceae (Phragmites III) MN

Puccinia recondita MCA2376 Aquilegia (Ranunculaceae) Italy ?? U473 Narcissus (Liliaceae) Germany

?? U473 Narcissus (Liliaceae) Germany Uredo dioscoreae-filiformis U458 Dioscorea (Discoreaceae) S. Africa

Uromyces kentaniensis U-385 (s.n.) Iridaceae (Chasmanthe O,I,II,III) South Africa Puccinia recondita U-363 Poaceae (Agrostis II,III) VA

Puccinia extensicola var. hydnoidea U-186 Dirca (Thymelaeaceae I) MN Cumminsiella mirabilissima AF426206 Mahonia (Berberidaceae) Cumminsiella U-480 Berberidaceae (Mahonia) Germany

Puccinia recondita U-364 Poaceae (Agropyron II,III) VA Puccinia myrsiphylli U-377 (#571) Liliaceae (Asparagus) South Africa

Puccinia hordei U-238 Lolium (Poaceae II-III) Turkey Uromyces geranii MCA2381 Geranium (O-I Geraniaceae) Italy

Puccinia malvacearum? U474 Alcea (Malvaceae) Germany Puccinia sorghi? U-326 Oxalidaceae (Oxalis I) MD Puccinia malvacearum AF426205 Alcea (Malvaceae)

Puccinia malvacearum JRH-225 Malvaceae (Malvastrum III) Honduras Puccinia isiacae U-340 Phragmites (Poaceae) Turkey Puccinia sp. U-139 Cardaria (Brassicaceae I) Turkey

Puccinia magnusiana? U-223 Poaceae (Phragmites) Turkey Puccinia magnusiana? U-223 Poaceae (Phragmites) Turkey

Uromycladium tepperianum U-466 Fabaceae (Acacia O,III) South Africa Uromycladium fusisporum BRIP-27608 Fabaceae (Acacia II-III) Australia

Phakopsora arthuriana U-8 Euphorbiaceae (Jatropha II) P.R. Phakopsora JRH-488 Euphorbiaceae P.R.

Phakopsora arthuriana GUY-2 Euphorbiaceae (Jatropha II) Guyana Phakopsora meibomiae JRH-456 Fabaceae (Lablab purpureus II) P.R. Phakopsora meibomiae JRH-471 Fabaceae (Lablab purpureus II) P.R. Phakopsora meibomiae JRH-475 Fabaceae (Lablab purpureus II) P.R. Phakopsora meibomiae U-102 Fabaceae (Phaseolus II) Honduras Phakopsora meibomiae JRH-284 Fabaceae (Desmodium II) Panama

Phakopsora meibomiae JRH-314 Fabaceae (Phaseolus multiflorus II) Panama Phakopsora tecta JRH-457 Commelinaceae (Commelina II) P.R. Phakopsora tecta JRH-137 Commelinaceae (Commelina II) Costa Rica Phakopsora tecta JRH-476 Commelinaceae (Commelina II) Puerto Rico Phakopsora tecta U-536 (#610) Commelinaceae (Aneilema) South Africa

Phakopsora pachyrhizi U-573 Fabaceae (Glycine max) LA Phakopsora pachyrhizi U-574 Fabaceae (Glycine max) LA Phakopsora pachyrhizi U-644 Fabaceae (Glycine max) Zimbabwe

Phakopsora gossypii BRIP-39200 Malvaceae (Gossypium II) Australia Uredopeltis sp nov? U-454 (#582) Tiliaceae (Grewia) South Africa Phakopsora nishidana U-14 (JRH 1996-051) Moraceae (Ficus II) Argentina

Batistopsora crucis-filii GUY-36 Annonaceae (Annona II-III) Guyana Phakopsora nishidana U-9 Moraceae (Ficus) P.R.

[IDd as]Ravenelia platensis U-91 Fabaceae (Erythrina O) Argentina ?? JRH160 Clytostoma (Bignoniaceae) Costa Rica

Prospodium? JRH-160 Bignoniaceae (Clytostoma) Costa Rica Prospodium? JRH-178 Bignoniaceae Costa Rica

?? JRH178 ?? (Bignoniaceae) Costa Rica ?? JRH 181 Bignoniaceae Costa Rica

Prospodium? JRH-181 Bignoniaceae Costa Rica Prospodium lippiae U-152 Verbenaceae (Aloysia II-III) Argentina

Prospodium appendiculatum JRH-251 Bignoniaceae (Tecoma) Mexica Raveneliaceae U-410 Fabaceae (Derris II) Guam Raveneliaceae U-416 Fabaceae (Derris II) Guam

Ravenellia havanensis U-19 Fabaceae (Enterolobium II-III)

Argentina Ravenelia havanensis U-114 Fabaceae (Enterlobium II-III)

Argentina Ravenelia echinata var. ectypa U-86 Fabaceae (Calliandra)

Argentina Ravenelia sp. U-00118 Acacia (Fabaceae) C.R.

Uredo sp. nov? (Ravenelia) U-121 Fabaceae (Acacia II) Costa Rica Uredo sp. nov? (Ravenelia) U-121 Acacia (Fabaceae II) C.R. Ravenelia acaciae-pennatulae U-115 Fabaceae (Acacia II) Mexico

Uredo sp.nov? U-120 Diphysa (Fabaceae II) C.R. Uredo sp.nov? U-120 Fabaceae (Diphysa II) Costa Rica

Ravenelia macrocarpa U-88 Fabaceae (Senna II) Argentina Puccinia spegazzinii U-317 (W1705) Asteraceae (M. micrantha III-microcyclic) Trinidad

Ravenelia macowaniana U-376t Fabaceae (Acacia III) South Africa Ravenelai macowaniana U-449 (#588) Fabaceae (Acacia III) South Africa

Ravenelia inornata U-376a (#570) Fabaceae (Acacia I) South Africa [from Phakopsora coca U-65 Erythroxylum (Erythroxylaceae II)] Argentina Gymnosporaingium MCA2568B Rosaceae Amelanchier I) VA

Gymnoconia peckiana U-349 Rosaceae (Rubus allegheniensis O,I) MN Gymnoconia peckiana U-349 Rosaceae (Rubus allegheniensis O,I) MN Gymnosporangium japonicum U-24 Cupressaceae (Juniperus III) CA

Gymnosporangium sp. MCA 2776 Rosaceae (Malus) VA Gymnosporangium juniperi-virginianae AY629316 AFTOL-712 Gymnosporangium juniperi-virginianae AF522167

Gymnosporangium exiguum? U-25 Rosaceae (Heteromales I) CA Gymnosporangium clavariiforme AF426211 Cupressaceae (Juniperus communis)

Gymnosporangium sabinae AY512845 Gymnosporangium sabinae AF426209 Rosaceae (Pyrus communis)

Gymnosporangium cornutum AF426210 Rosaceae (Sorbus aucuparia) enodphytic rust? U-665 Poaceae (Bouteloua eriopoda, asymptomatic) NM Puccinia podophylli U-2 Berberidaceae (Podophyllum I-III) MD Puccinia podophylli U-13 Berberidaceae (Podophyllum O-III) MD Tranzschelia discolor JRH-139 Rosaceae (Prunus domestica II) Costa Rica

Tranzschelia discolor U-507 (KR-0010962) Ranunculaceae (Anemone coronaria O,I) Greece Tranzschelia discolor U-509 (KR-0010965) Rosaceae (Prunus persica) Iran Tranzschelia discolor U-510 (KR-0010966) Rosaceae (Prunus domestica) Iran Tranzschelia discolor U-517 (KR-0010973) Rosaceae (Prunus persica) Iran Tranzschelia discolor AF426223 Rosaceae (Prunus domestica)

Tranzschelia hyrcanica U-512 (KR-0010968) Rosaceae (Prunus cerasifera) Iran Tranzschelia hyrcanica U-513 (KR-0010969) Rosaceae (Prunus cerasifera) Iran Tranzschelia hyrcanica U-514 (KR-0010970) Rosaceae (Prunus cerasifera) Iran Tranzschelia hyrcanica U-515 (KR-0010971) Rosaceae (Prunus cerasifera) Iran Tranzschelia hyrcanica U-511 (KR-0010967) Rosaceae (Prunus spinosa) Iran Tranzschelia pruni-spinosae U-516 (KR-0010972) Rosaceae (Prunus cerasifera) Iran Tranzschelia pruni-spinosae AF426224 Ranunculaceae (Anemone ranunculoides) Tranzschelia fusca AF426225 Ranunculaceae (Anemone nemorosa) Phakopsora? JRH-463 Poaceae Puerto Rico

Melampsora epitea complex U-562 Salicaceae (Salix sp.) WA melampsora epitea complex (SAL-53) Salix bebbiana ID Melampsora epitea? U-172 S. sp. Norway

Melampsora laricis-epitea f.sp. laricis-daphnoides

AY444789 (&AY444777ITS2) S. acutifolia N. Ireland Melampsora epitea complex U-559 Salicaceae (Salix sp.) WA Melampsora (SAL-2) S. melanopsis ID

Melampsora coleosporioides AY652951 (&AY652948ITS2) S.

babylonica, Japan Melampsora amygdalinae AY444782 (AY444776ITS2) S.

triandra N.Ireland Melampsora epitea U-284 [seq. inc.] S. humboldtiana

Venezuela Melampsora ribesii-purpureae AY444791 (&AY444770ITS2) S.

purpurea England Melampsora hypericorum U-394 Clusiaceae (Hypericum) WA

Melampsora euphorbiae-gerardianae? U-485 Euphorbiaceae

(Euphorbia) Germany Melampsora euphorbiae U-138 [LSU only] Euphorbiaceae

(Euphorbia II) Syria Melampsora epitea complex U-557 [multiple PPs] Salix sp. WA

Melampsora sp. (SAL-83) S. lemmonii ID Melampsora laricis AB116809 Populus davidiana, Tibet, China

Melampsora occidentalis U-395 Populus balsamifera II WA Melampsora (SAL-49) [LSU only] Pinaceae (Abies grandis I) ID

Pucciniastrum agrimoniae AF426234 Rosaceae (Agrimonia eupatoria) Cronartium quercuum U-62 Fagaceae (Quercus macrocarpa) IN Cronartium quercuum U-432 Fagaceae (Quercus muehlenbergii II) IN Cronartium quercuum U-434 Fagaceae (Quercus alba II) IN Cronartium quercuum U-435 Fagaceae (Quercus bicolor II) IN Cronartium quercuum u-436 Fagaceae (Quercus macrocarpa II) IN Cronartium ribicola U-396/U-268 Ribes VA/Pinus WY

Endocronartium harknessii AF522175 Cronartium flaccidum AF426239 Asclepiadaceae (Vincetoxicum hirundinaria)

Endocronartium harknessii AY700193 Pucciniastrum agrimoniae? U-304 Rosaceae (Leucoside) South Africa [inc. seq.]

Chrysomyxa arctostaphyli AF522163 Chrysomyxa arctostaphyli AY700192 Chrysomyxa ledi AF426246 Pinaceae (Picea abies) Chrysomyxa rhododendri AF426245 Ericaceae (Rhododendron ferrugineum)

Caeoma tsukubaensis AB19124 Ericaceae (Rhododendron kaempferi) Japan Thekopsora areolata AF426235 Rosaceae (Prunus padus)

Hyalopsora U-361 undet. fern (Woodsiaceae?) UT Hyalopsora polypodii AF426229 Dryopteridaceae (Cystopteris fragilis)

Hyalopsora polypodii AY512852 Pucciniastrum? MCA2884 ?? Costa Rica

Melampsoridium betulinum AF426228 Fagaceae (Betula pendula) Naohidemyces vaccinii MCA 2780 Ericaceae (Vaccinium) WA Naohidemyces vaccinii AF426238 Ericaceae (Vaccinium uliginosum)

Uredinopsis sp. AF522181 Uredinopsis filicina AF426237 Thelypteridaceae (Phegopteris connectilis) Milesina (as Milesia) scolopendri AF426236 Aspleniaceae (Asplenium scolopendrium)

Pucciniastrum epilobii AF426226 Onagraceae (Epilobium angustifolium) Pucciniastrum epilobii AF522178 Pucciniastrum circaeae AY745697

Pucciniastrum epilobii AF522179 Pucciniastrum circaeae AF426227 Onagraceae (Circaea lutetiana)

Melampsorella caryophyllacearum AF426232 Pinaceae (Abies alba) Pucciniastrum goeppertianum AF522180

Coleosporium ipomoeae JRH-195 Convolvulaceae (Ipomoea nil II) Costa Rica Coleosporium ipomoeae JRH-485 Convolvulaceae (Merremia quinquefolia II-III) Puerto Rico Coleosporium ipomoeae JRH-465 Convolvulaceae (Ipomoea triloba II) P.R. Coleosporium ipomoeae JRH-216 Convolvulaceae (Ipomoea cholulensis II-III) Honduras Coleosporium ipomoeae JRH-168 Convolvulaceae (Ipomoea trifida II-III) Costa Rica Coleosporium ipomoeae JRH-230 Convolvulaceae (Ipomoea anisomeres II) Honduras Coleosporium ipomoeae JRH-143 Convolvulaceae (Ipomoea neeiII) Costa Rica Coleosporium plumierae U-112 Apocynaceae (Plumeria rubra II-III) Nigeria Coleosporium plumeriae U-50 Apocynaceae (Plumeria rubra II) Guyana Coleosporium plumeriae U-266 Apocynaceae (Plumeria II) GUAM

Coleosporium (asterum or delicatulum?) U-369 Asteraceae (Solidago) NY Coleosporium asterum U-67 Asteraceae (Symphyotrichum (=Aster) racemosum II-III) MD

Coleosporium asterum AF522165 Coleosporium asterum AF426241 Asteraceae (Aster ciliolatus) Coleosporium (asterum or delicatulum?) MCA-2600 Asteraceae (Solidago II) TN Coleosporium U-430 Asteraceae (Solidago?) NJ Coleosporium U-389 Asteraceae (Solidago?) MD Coleosporium U-388 Asteraceae (Solidago?) PA Coleosporium (asterum or delicatulum?) U-428 Asteraceae (Solidago) MD Coleosporium (asterum or delicatulum?) MCA-2777 S(Asteraceae (Solidago canadensis II-III) MD

Coleosporium asterum AF522164 Coleosporium sp. U-398 Asteraceae MD Coleosporium asterum U-366 Asteraceae (Aster novae-angliae II-III?) MD Coleosporium (asterum or delicatulum?) U-347 Asteraceae (Solidago) PA

Coleosporium vernoniae U-371 Asteraceae (Vernonia) IL Coleosporium tussilaginis U-74 Asteraceae (Senecio triangularis) OR Coleosporium tussilaginis MCA-2389 Asteraceae (Sonchus?) CA Coleosporium tussilaginis MCA-2392 Asteraceae (Sonchus or Lactuca) CA Coleosporium vernoniae GUY-25 Asteraceae (Elephantopus mollis II) Coleosporium senecionis AY512840 Coleosporium tussilaginis AF426242 Asteraceae (Tussilago farfara) Coleosporium cacaliae AF426243 Asteraceae (Adenostyles glabra) Coleosporium campanulae AF426244 Campanulaceae (Campanula scheuchzeri)

Thekopsora symphyti AF426230 Boraginaceae (Symphytum officinale) Thekopsora guttata AF426231 Rubiaceae (Galium odoratum)

Gymnosporangium libocedri AF522168 Phakopsora apoda PDD-72076 Poaceae (Pennisetum II) New Zealand Phakopsora apoda PDD-68451 Poaceae (Pennisetum II) New Zealand

Dasturella divina BRIP-39289 Poaceae (Babusa II) Australia Dasturella divina MCA-2887 Poaceae (bamboo) Costa Rica

Phakopsora? MCA2617 Poaceae (Panicum) NC Phakopsora phakopsoroides JRH-184 Poaceae (Olyra II) Costa Rica

Phakopsora euvitis BRIP-27996 Vitaceae (Vitis II) Australia Phakopsora vernoniae U-256 [#368] Asteraceae (Vernonia) South Africa

Uredo alpestris AF426212 Violaceae (Viola alpestris) Kuehneola loeseneriana? MCA-2879 Rosaceae (Rubus trichomallus?) Costa Rica

Gerwasia (as Mainsia) pittieriana U-78 Rosaceae (Rubus) Guatemala Kuehneola uredinis U-429 Rosaceae (Rubus) MD Kuehneola uredinis U-303 Rosaceae (Rubus pinnatus) South Africa

Kuehneola uredinis U-348 Rosaceae (Rubus) PA Kuehneola uredinis AY745696 AFTOL-987 Kuehneola uredinis AF426218 Rosaceae (Rubus fruticosus)

Kuehneola uredinis MCA-2393 Rosaceae (Rubus III) MD Kuehneola uredinis MCA-2867 Rosaceae (Rubus) MD Kuehneola uredinis MCA-2863 Rosaceae (Rubus) VA

Frommeella mexicana var. indicae U-3 Rosaceae (Duchesnea II) MD FrommeÎlla mexicana MCA-2496 Rosaceae (Duchesnea) MD Frommeella mexicana MCA-2495 Rosaceae (Duchesnea II?) VA

Phragmidium mucronatum U-106 Rosaceae (Rosa II) Argentina Phragmidium mucronatum? U-353 Rosaceae (Rosa) OR

Phragmidium sanguisorbae AF426216 Rosaceae (Sanguisorba minor) Phragmidium violaceum AF426214 Rosaceae (Rubus fruticosus) Phragmidium violaceaum MCA-2782 Rosaceae (Rubus fruticosus? II-III) France Phragmidium violaceum U-645 Rosaceae (Rubus armeniacus II) OR

Phragmidium rubi-idaei AF426215 Rosaceae (Rubus idaeus) Phragmidium fragariae AF426217 Rosaceae (Potentilla sterilis)

Phragmidium montivagum AF426213 Rosaceae (Rosa cf. woodsii) Triphragmium ulmariae AF426219 Rosaceae (Filipendula ulmaria)

Ochropsora ariae AF426221 Ranunculaceae (Anemone nemorosa) Ochropsora ariae AF426222 Rosaceae (Aruncus dioicus)

Trachyspora intrusa AF426220 Rosaceae (Alchemilla vulgaris) Trachyspora intrusa MCA2384 Alchemilla (Rosaceae) Switzerland Caeoma torreyae AF522183

Eocronartium muscicola AY512844 Platygloea vestita AY512872 Rhodotorula fushanensis AB176591

Helicobasidium purpureum AY885168

1 change

1229 bp.; 857 incl. char.; 263 PI char 470 seq; 350 taxa (+4 outgroup)

13 of 13 families (100%) 58 of 117 genera (50%)

~700 of 7000 species (10%)

(classifications fide Cummins & Hiratsuka 2003)

Pucciniales rDNA LSU

Dataset: Aime. 2006. Mycoscience 47(3):112-122. And unpublished.

10 GROUPS

8 families supported 4 families redundant

(1 unresolved) Several “orphan genera”

Puccinia cf. violae U-4 Violaceae (Viola cucullata O-I, autoecious) MD

Uromyces appendiculatus AY123307 [Fabaceae]

Puccinia pelargonii-zonalis AY123316

Puccinia pelargonii-zonalis AY125421 Geraniaceae (Pelargonium, autoecious) Australia

Dietelia mesoamericana [as portoricensis] AY125414 Asteraceae (Mikania micrantha) Mexico

Dietelia portoricensis AY125389 Asteraceae (Mikania micrantha) Costa Rica

Puccinia lagenophorae AY125413 Asteraceae (Senecio minimus, autoecious)

Puccinia cardui-pycnocephali AY125410 Asteraceae (Carduus pycnocephali I, III)

Miyagia pseudosphaeria AY125411 [Sonchus]

Puccinia menthae AY123315 [autoecious Lamiaceae]

Puccinia hordei AY125412 (I on Thalictrum sp., III on Hordeum vulgare)

Puccinia hordei MCA2391 (AFTOL1402) Poaceae CA

Puccinia graminis f. sp. tritici AY125409 [I-Berberidaceae, III-Poaceae]

Aecidium kalanchoe U-18 Crassulaceae (Kalanchoe I) Costa Rica

Sphenospora kevorkianii U-10 Orchidaceae (Stanhopea II) Peru

Puccinia smilacis MCA-2866 Smilacaceae (Smilax III) MD

Puccinia smilacis U-393 Smilacaceae (Smilax) MD

Aecidium epimedii AB011050 [Berberidaceae (Podophyllaceae), Asia]

Nyssopsora echinata AY123312 [autoecious Apiaceae, old name Triphragmium]

Nyssopsora echinata U77061

Dasturella divina AY123288 [bambusa & oth Poaceae]

Puccinia suzutake AB011053 [Hydrangeaceae, Poaceae]

Batistopsora crucis-filii GUY-36 Annonaceae (Annona II-III) Guyana

MasseeÎlla capparis BRIP-26745 Euphorbiaceae (Flueggea III) Australia

Physopella tecta AY125397 Commeliniaceae (Commelina diffusa III) Hawaii

Phakopsora jatrophicola AY125398 Euphorbiaceae (Jatropha III) probably microcyclic

Phakopsora pachyrhizi AY125399 (no known I, III on Glycine)

Phakopsora pachyrhizi AY123301

Uredo vetus AY123318 [Selaginella--Selaginellaceae]

Pileolaria brevipes AY123314 [autoecious on Toxicodendron, Anacardiaceae]

Tranzschelia discolor AY125403 I on Anemone sp.; III on Prunus padus, heteroecious

Gymnosporangium libocedri AY123290 [I-Rosaceae; III-Calocedrus decurrens, Cupressaceae)

Puccinia podophylli U-2 Berberidaceae (Podophyllum I-III) MD

Ochropsora kraunhiae AB011051 [old name Phakopsora, Japan, Fumariaceae & Fabaceae I]

Physopella ampelopsidis AB011054 [III-Vitaceae; I- Sabiaceae]

Gymnoconia peckiana AY125407 I & III on Rubus articus, autoecious

Gymnoconia nitens AY123308

Gymnoconia nitens U41565

Kuehneola uredinis AY123310 [Rubus, autoecious]

Kuehneola uredinis AFTOL-987

Gerwasia rubi AY125388 France [Rubus]

Frommeella mexicana var. indicae U-3 Rosaceae (Duchesnea II) MD

Phragmidium barnardii AY125406 I & III on Rubus triphyllus, autoecious

Phragmidium tuberculatum AY123313 [Rosa]

Phragmidium rubi-idaei AY125405 I & III on Rubus idaeus, autoecious

Triphragmium ulmariae AY125402 I & III on Filipendula ulmaria (Rosaceae), autoecious


Trachyspora intrusa MCA-2384 Rosaceae (Alchemilla I-III) Switzerland

Gymnosporangium juniperi-virginianae AY123289 [I-Malus, III-Juniperus, Cupressaceae]

Gymnosporangium clavipes AY123309 [I-Rosaceae; III-Juniperus]

Uromycladium sp. AY125415 Fabaceae (Acacia mearnsii)

Uromycladium notabile AY 125416 Fabaceae (Acacia sp. I,III) autoecious


Uromycladium tepperianum AY125420 Fabaceae (Acacia sp. I,III) autoecious

Uromycladium tepperianum AY 125419 Fabaceae (Acacia sp. I,III) autoecious

Uromycladium tepperianum U-466 Fabaceae (Acacia O,III) South Africa

Prospodium tuberculatum AY221466 [Verbenaceae III Latin America]

Prospodium lippae U-152 (AFTOL) Verbenaceae (Aloysia II-III) Argentina

Ravenellia havanensis U-19 Fabaceae (Enterolobium II-III) Argentina

Ravenelia havanensis U-114 (AFTOL) Fabaceae (Enterolobium) Argentina

Maravalia sapotae JRH-172 Sapotaceae (Manilkara II-III) Costa Rica

Coleosporium asterum AY123286 [I-pinus; III-Asteraceae]

Coleosporium asterum AY123287

Coleosporium paederiae AY125390 Rubiaceae (Paederia foetida) [I-Pinus; III-Rubiaceae]

Chrysomyxa arctostaphyli AY123285 [I-Pinaceae, Picea; III-Ericaceae, Acrtostaphylos]

Chrysomyxa arctostaphyli AY657009

Hyalospora aspidiotus AY123291 [I-Abies (Pinaceae); III-Dryopteridaceae Europe]

Hyalopsora polypodii AB011016 Dryopteridaceae[heteroecious but I host unknown; III on ferns]

Hyalopsora polypodii AB011017 Dryopteridaceae (Athyrium yokoscense)

Melampsoridium sp. AY123299

Melampsoridium betulinum AY125391 I Pinaceae (Larix sp.) III Fagales (Betula pubescens); heteroecious

Cronartium ribicola M94338 I-Pinus; III-Grossulariaceae, Scrophulariaceae, Saxifragaceae]

Endocronartium harknessii AY665785 [Pinus, autoecious]

Endocronartium (as Peridermium) harknessii M94339 I

Milesina leviuscula AY123300 [Abies & ferns]

Uredinopsis intermedia AB011018 Dryopteridaceae [I-Abies]

Uredinopsis sp. (pteridis or hashiokai) AY123306

Melampsora epitea AY123293 [1-Abies or Ribes; III Salix]

Melampsora lini AY125396 Linaceae (Linum marginale I, & III-autoecious)

Melampsora laricis-populina AY125395 [I-Larix; III-Populus canadensis]

Melampsora euphorbiae AY125393 Euphorbiaceae (Euphorbia peplus III?)

Melampsora hypericorum AY123311 [Clusiaceae]

Melampsora euphorbiae-gerardianae AY123294 [Euphorbiaceae]

Melampsora occidentalis AY123295 [I-Pinaceae; III-Populus]

Pucciniastrum areolatum AY125392 I Pinaceae (Picea abies), III Rosaceae (Prunus padus); heteroecious

Melampsorella caryophyllacearum AY123298 [I-Abies; III-Carophyllaceae]

Pucciniastrum goeppertianum AY123305 [montypic Calyptospora? I-Abies; III-Vaccinium]

Pucciniastrum epilobii AY123303 [I-Abies; III-Onagraceae]

Peridermium ephedrae AY123317 [Ephadraceae I (ii, III)]

Hemileia vastatrix AY 125422 Rubiaceae (Coffea) South Africa

Naohidemyces vaccinii MCA 2780 Ericaceae (Vaccinium) WA [I-Tsuga, Pinaceae]

Maravalia cryptostegiae AY125404 [Asclepiadaceae (Cryptostegia) Australia]

Caeoma torreyae AY123284 [Taxaceae (Torreya) CA]

Racospermyces koae AY125387 Fabaceae (Acacia koa I, III) (autoecious,functionally microcyclic)

Atelocauda digitata AY125400 Fabaceae (Acacia koa I,III) (autoecious,functionally microcyclic)

Helicobasidium purpureum D85648

Eocronartium muscicola AY123323

Septobasidium canescens AY123320

Platygloea vestita AY124480

Rhodotorula fushanensis AB176530


rDNArDNA SSUSSU

PucciniaceaePucciniaceae PucciniaceaePucciniaceae

Nyssopsora Nyssopsora & & KwelingiaKwelingia

Phakopsoraceae p.p.Phakopsoraceae p.p.

PhragmidiaceaePhragmidiaceae inclincl. . TrachysporaTrachyspora & & TriphragmiumTriphragmium

GymnosporangiumGymnosporangium

UromycladiumUromycladium

RaveneliaceaeRaveneliaceae p.p.p.p.

MelampsoraceaeMelampsoraceae (incl. (incl. ColeosporiaceaeColeosporiaceae & & PucciniastraceaePucciniastraceae))

MikronegeriaceaeMikronegeriaceae + + C.C. torreyaetorreyae,, HemileiaHemileia

PileolariaPileolaria ((PileolariaceaePileolariaceae)), , OchropsoraOchropsora && TranzscheliaTranzschelia

Pucciniales
















































































































PucciniaceaePucciniaceae excl.excl.



rDNArDNA SSUSSU

Pucciniales
















































































































PhakopsoraceaePhakopsoraceae not monophyleticnot monophyletic


rDNArDNA SSUSSU

Pucciniales

















































































































rDNArDNA SSUSSU

Pucciniales

PhragmidiaceaePhragmidiaceae s.s.s.s. &&

RaveneliaceaeRaveneliaceae s.s.s.s. HostHost--specific familiesspecific families


0.01 substitutions/site

Phakopsoraceae p.p

Phragmidiaceae

Coleosporiaceae s.s.

Mikronegeriaceae

Melampsoraceae

Tranzschelia

Raveneliaceae

Pileolariaceae

Gymnosporangium

Cronartiaceae s.s.

Caeoma torreyae

Pucciniaceae Aecial Host

Basal lineages (I-Taxaceae, Podocarpaceae, Araucariaceae)

Gymnosperm rusts (I-Pinaceae)

Angiosperm rusts (I-Angiosperms)


rDNArDNA LSU LSU & SSU& SSU

Pucciniales

What about heteroecism?

•• “Primitive” hosts harbor “primitive” rusts (e.g., “Primitive” hosts harbor “primitive” rusts (e.g., HyalopsoraHyalopsora & other fern rusts)& other fern rusts) Heteroecism is ancestral

• Rusts are evolved from “shortRusts are evolved from “short--cycled” (unexpanded) cycled” (unexpanded) tropical rusts with simple tropical rusts with simple teliosporesteliospores ((ChaconiaceaeChaconiaceae) ) (e.g., Ono 2002)(e.g., Ono 2002) Heteroecism is derived

Sjamsuridzal et al. 1999. Mycoscience 40:21-27.

Maier et al. 2003. Can. J. Bot. 81:12-23.

Uromyces setaria-italicae JRH202 Panicum (II Poaceae) Honduras Puccinia gouaniae JRH159 Gouania (II Rhamnaceae) Costa Rica

Puccinia arundinariae? MCA2494 Arundinaria (Poaceae) VA Puccinia purpurea JRH198 Sorghum (Poaceae II-III) Honduras Puccinia purpurea JRH213 Sorghum (Poaceae II) Hondurus

Puccinia purpurea JRH218 Sorghum (Poaceae II) Hondurus Puccinia duthiae GUY8 Bothriochloa (Poaceae II) Guyana Puccinia duthiae U-264 Dichanthium (Poaceae II) Guam

Puccinia exhauriens U-538 (#600) Oleaceae (Jasminum) South Africa Puccinia exhauriens U-539 (#605) Oleaceae (Jasminum) South Africa

Puccinia moraeae U462 Moraea (Iridaceae) S. Africa Uromyces massoniae? U-446(#586) Liliaceae (Massonia II-III) South Africa Uromyces salsolae U-134 Salsola (Chenopodiaceae) Syria




Sphenospora kevorkianii U-10 Orchidaceae (Stanhopea II) Peru Puccinia coronata U-583 Poaceae (Avena) ND Puccinia coronata U400 Festuca (Poaceae II, III) VA Puccinia coronata U-244 Rhamnus (Rhamnaceae O-O) ND Puccinia coronata U-245 Scolochloa (Poaceae III?) ND Puccinia coronata AF426207 Rhamnus (Rhamnaceae)

Puccinia pampeanum U-109 Salpichroa (III Solanales) Argentina Endophyllum pampeanum U-108 Salpichroa (III Solanales) Argentina Uromyces gageae AF426208 Gagea (Liliaceae) Aecidium wiborgiae U-380 (#576) Fabaceae (Wiborgia) South Africa Aecidium wiborgiae U-379 (#573) Fabaceae (Wiborgia) South Africa Uromyces sp. MCA2383 Lilium (III Liliaceae) Switzerland

Puccinia mogiphanis U-111 Alternanthera (Amaranthaceae) Nigeria Puccinia stonemanniae U-310 Thesium (Santalaceae III?) South Africa

Puccinia sorghi U-362 Poaceae (Zea mays II, III) VA Uromyces polygoni U-214 Polygonum (Polygonaceae) Turkey Uromyces polygoni U-214 Polygonum (Polygonaceae) Turkey Puccinia mogiphanis JRH 180 Amarantaceae (Alternanthera II) Costa Rica

Puccinia cynodontis JRH 219 Cynodon (Poaceae II-III) Hondorus Aecidium kalanchoe U-18 Kalanchoe Puccinia horiana U-391 Asteraceae (Chrysanthemum III) DE Puccinia horiana U-392 Asteraceae (Chrysanthemum III) MD Puccinia horiana U-070 Chrysanthemum (Asteraceae) NJ Aecidium sp. U-465 (#579) Fabaceae (Lebeckia) South Africa

Puccinia arenariae U142 Dianthus (Caryophyllaceae) Ecuador Puccinia graminis U-478 Berberis (Berberidaceae) Germany Puccinia.poarum FROM BRANDON on Tussilago (iii on Poaceae) Puccinia MCA-2882 Asteraceae? Costa Rica Puccinia MCA-2886 Asteraceae? Costa Rica Puccinia graminis (as Aecidium sp.) PDD78317 Berberis I N.Z. Puccinia graminis U-469 Agropyron (Poaceae) Germany Puccinia graminis U-470 Agropyron? (Poaceae) Germany Puccinia graminis U-470 Poaceae (Agropyron?) Germany

Uredo baruensis GUY-71 Sapotaceae (Chrysophyllum II) Guyana [IDd as Coleosporium on Merremia (Convolvulaceae)] JRH-484 P.R. (seq. confirmed 2X) Sphaerophragmium acacia JRH-486 Fabaceae (Albizia II-III) Puerto Rico Sphaerophragmium acaciae JRH-470 Fabaceae (Albizia II) Puerto Rico

Pileolaria toxicodendri AY 745698 (AFTOL ID 988) Pileolaria toxicodendri u-607 Anacardiaceae (Toxicodendron) MN

Puccinia tetragonia U-442 Aizoaceae (Tetragonia) South Africa ?? U-476 Allium (Liliaceae) Germany Uromyces albucae U-447 (#591) Liliaceae (Albuca) South Africa Puccinia dispersa U-234 Anchusa (Boraginaceae) Turkey Puccinia? MCA-2885 ?? Costa Rica


Maravalia sapotae JRH-172 Sapotaceae (Manilkara II-III) Costa Rica Stereostratum corticiodes U-027 Bambusa (Poaceae II-III) China Puccinia asarina MCA2374 Asarum (III Aristolochiaceae) Italy Puccinia pampeana JRH135 Capsicum (Solanaceae II-III aecioid) Costa Rica Endophyllum macowanii U-255 Rhamnus (Rhamnaceae III?) South Africa Puccinia thlaspeos U-054 Boechera (III Brassicaceae) WA Uromyces bulbinis BRIP-28988 Liliaceae (Bulbine) Australia Uromyces heteromorphae U-309 Peucedanum (Apiaceae) South Africa Uromyces trollipii U-445 Zygophyllaceae (Zygophyllum I, III) South Africa




Puccinia recondita MCA2376 Aquilegia (Ranunculaceae) Italy ?? U473 Narcissus (Liliaceae) Germany ?? U473 Narcissus (Liliaceae) Germany Uredo dioscoreae-filiformis U458 Dioscorea (Discoreaceae) S. Africa Uromyces kentaniensis U-385 (s.n.) Iridaceae (Chasmanthe O,I,II,III) South Africa

Puccinia recondita U-363 Poaceae (Agrostis II,III) VA Puccinia extensicola var. hydnoidea U-186 Dirca (Thymelaeaceae I) MN

Cumminsiella mirabilissima AF426206 Mahonia (Berberidaceae) Cumminsiella U-480 Berberidaceae (Mahonia) Germany




Puccinia malvacearum JRH-225 Malvaceae (Malvastrum III) Honduras Puccinia isiacae U-340 Phragmites (Poaceae) Turkey Puccinia sp. U-139 Cardaria (Brassicaceae I) Turkey Puccinia magnusiana? U-223 Poaceae (Phragmites) Turkey Puccinia magnusiana? U-223 Poaceae (Phragmites) Turkey


Phakopsora arthuriana U-8 Euphorbiaceae (Jatropha II) P.R. Phakopsora JRH-488 Euphorbiaceae P.R. Phakopsora arthuriana GUY-2 Euphorbiaceae (Jatropha II) Guyana

Phakopsora meibomiae JRH-456 Fabaceae (Lablab purpureus II) P.R. Phakopsora meibomiae JRH-471 Fabaceae (Lablab purpureus II) P.R. Phakopsora meibomiae JRH-475 Fabaceae (Lablab purpureus II) P.R. Phakopsora meibomiae U-102 Fabaceae (Phaseolus II) Honduras Phakopsora meibomiae JRH-284 Fabaceae (Desmodium II) Panama Phakopsora meibomiae JRH-314 Fabaceae (Phaseolus multiflorus II) Panama Phakopsora tecta JRH-457 Commelinaceae (Commelina II) P.R. Phakopsora tecta JRH-137 Commelinaceae (Commelina II) Costa Rica Phakopsora tecta JRH-476 Commelinaceae (Commelina II) Puerto Rico Phakopsora tecta U-536 (#610) Commelinaceae (Aneilema) South Africa Phakopsora pachyrhizi U-573 Fabaceae (Glycine max) LA Phakopsora pachyrhizi U-574 Fabaceae (Glycine max) LA Phakopsora pachyrhizi U-644 Fabaceae (Glycine max) Zimbabwe



[IDd as]Ravenelia platensis U-91 Fabaceae (Erythrina O) Argentina Prospodium sp nov? JRH-183 Bignoniaceae (Cydista II-III) Costa Rica

?? JRH160 Clytostoma (Bignoniaceae) Costa Rica Prospodium? JRH-160 Bignoniaceae (Clytostoma) Costa Rica

Prospodium? JRH-178 Bignoniaceae Costa Rica ?? JRH178 ?? (Bignoniaceae) Costa Rica ?? JRH 181 Bignoniaceae Costa Rica



Ravenellia havanensis U-19 Fabaceae (Enterolobium II-III) Argentina Ravenelia havanensis U-114 Fabaceae (Enterlobium II-III) Argentina Ravenelia echinata var. ectypa U-86 Fabaceae (Calliandra) Argentina Ravenelia sp. U-00118 Acacia (Fabaceae) C.R.



Ravenelia macrocarpa U-88 Fabaceae (Senna II) Argentina Puccinia spegazzinii U-317 (W1705) Asteraceae (M. micrantha III-microcyclic) Trinidad Ravenelia macowaniana U-376t Fabaceae (Acacia III) South Africa Ravenelai macowaniana U-449 (#588) Fabaceae (Acacia III) South Africa




Gymnosporangium exiguum? U-25 Rosaceae (Heteromales I) CA Gymnosporangium clavariiforme AF426211 Cupressaceae (Juniperus communis) Gymnosporangium sabinae AY512845 Gymnosporangium sabinae AF426209 Rosaceae (Pyrus communis)

Gymnosporangium cornutum AF426210 Rosaceae (Sorbus aucuparia) enodphytic rust? U-665 Poaceae (Bouteloua eriopoda, asymptomatic) NM Puccinia podophylli U-2 Berberidaceae (Podophyllum I-III) MD Puccinia podophylli U-13 Berberidaceae (Podophyllum O-III) MD Tranzschelia discolor JRH-139 Rosaceae (Prunus domestica II) Costa Rica Tranzschelia discolor U-507 (KR-0010962) Ranunculaceae (Anemone coronaria O,I) Greece Tranzschelia discolor U-509 (KR-0010965) Rosaceae (Prunus persica) Iran Tranzschelia discolor U-510 (KR-0010966) Rosaceae (Prunus domestica) Iran Tranzschelia discolor U-517 (KR-0010973) Rosaceae (Prunus persica) Iran Tranzschelia discolor AF426223 Rosaceae (Prunus domestica) Tranzschelia hyrcanica U-512 (KR-0010968) Rosaceae (Prunus cerasifera) Iran Tranzschelia hyrcanica U-513 (KR-0010969) Rosaceae (Prunus cerasifera) Iran Tranzschelia hyrcanica U-514 (KR-0010970) Rosaceae (Prunus cerasifera) Iran Tranzschelia hyrcanica U-515 (KR-0010971) Rosaceae (Prunus cerasifera) Iran Tranzschelia hyrcanica U-511 (KR-0010967) Rosaceae (Prunus spinosa) Iran Tranzschelia pruni-spinosae U-516 (KR-0010972) Rosaceae (Prunus cerasifera) Iran Tranzschelia pruni-spinosae AF426224 Ranunculaceae (Anemone ranunculoides) Tranzschelia fusca AF426225 Ranunculaceae (Anemone nemorosa) Phakopsora? JRH-463 Poaceae Puerto Rico

Melampsora epitea complex U-562 Salicaceae (Salix sp.) WA melampsora epitea complex (SAL-53) Salix bebbiana ID Melampsora epitea? U-172 S. sp. Norway Melampsora laricis-epitea f.sp. laricis-daphnoides AY444789 (&AY444777ITS2) S. acutifolia N. Ireland Melampsora epitea complex U-559 Salicaceae (Salix sp.) WA Melampsora (SAL-2) S. melanopsis ID

Melampsora coleosporioides AY652951 (&AY652948ITS2) S. babylonica, Japan Melampsora amygdalinae AY444782 (AY444776ITS2) S. triandra N.Ireland

Melampsora epitea U-284 [seq. inc.] S. humboldtiana Venezuela Melampsora ribesii-purpureae AY444791 (&AY444770ITS2) S. purpurea England

Melampsora hypericorum U-394 Clusiaceae (Hypericum) WA Melampsora euphorbiae-gerardianae? U-485 Euphorbiaceae (Euphorbia) Germany Melampsora euphorbiae U-138 [LSU only] Euphorbiaceae (Euphorbia II) Syria

Melampsora epitea complex U-557 [multiple PPs] Salix sp. WA Melampsora sp. (SAL-83) S. lemmonii ID

Puccinia mariae-wilsoniae U-005 Claytonia (Portulaceae I-III) MD Puccinia mariae-wilsoniae MCA2497 Claytonia (Portulaceae) MD Puccinia lagenophorae U-384 (#569) Asteraceae (Osteocarpum) South Africa Puccinia lagenophora U-128 Senecio (I-III Asteraceae) Tunisia Puccinia lagenophorae U-107 Senecio (I, III Asteraceae) Argentina Puccinia violae U-023 Viola (Violaceae II-III) Norway Puccinia violae U-269 Viola (Violaceae II-III) NY Uromyces sp. MCA 2870 Juncaceae (Juncus? III) VA Puccinia? U397 Asteraceae MD Puccinia? U-390 Asteraceae MD

Puccinia cf. violae U-4 Violaceae (Viola cucullata )-I, autoecious) MD Uromyces vignae? U-015 Macroptilium (Fabaceae II) Argentina Uromyces vignae JRH-472 Fabaceae (Vigna II-III) Puerto Rico Uromyces vignae JRH-468 Fabaceae (Vigna II-III) Puerto Rico Uromyces vignae U-125 Vigna (Fabaceae II) C.R. Uromyces vignae U-126 Vigna (Fabaceae II) C.R. Uromyces appendiculatus U-122 Phaseolus (II Fabaceae) C.R. Uromyces appendiculatus U-123 Phaseolus (II Fabaceae) Mexico Uromyces appendiculatus U-00123 Phaseolus (Fabaceae) Mexico



Puccinia granularis U-444 (#581) Geraniaceae (Pelargonium II,III) South Africa Puccinia granularis U-444 (#581) Geraniaceae (Pelargonium II,III) South Africa Puccinia granularis U-443 (#580) Geraniaceae (Pelargonium O,I,II,III) South Africa

Puccinia spegazzinii JRH-226 Asteraceae (Mikania micrantha III) Honduras Puccinia spegazzinii U-319 (W2102) Asteraceae (Mikania micrantha microcyclic III) Peru Puccinia spegazzinii U-315 (W2020) Asteraceae (Mikania micrantha IIImicrocyclic) Argentina Dietelia mesoamericana sp. nov. U-321 (mislabelled as U143) Mikania Mexico

Puccinia hydrocotyles U-079 Hydrocotyle (II-III Apiaceae) AL Puccinia? MCA-2864 Asteraceae (Helianthus) VA Puccinia pimpinellae? MCA2576 Apiaceae( Osmorhiza I) ID






Puccinia commelinae U-049 Commelina (Commelinaceae II) Guyana Puccinia synedrellae JRH152 Synedrella (III Asteraceae) Costa Rica Puccinia cnici-oleracei GUY-6 Asteraceae (Emilia sonchifolia III) Guyana Puccinia synedrellae JRH-152 Asteraceae (Synedrella nodiflora III) Costa Rica Puccinia cnici-oleracei JRH-204 Asteraceae (Emilia fosbergii III) Honduras Puccinia cnici-oleracei JRH-490 Asteraceae (Emilia fosbergii III) Puerto Rico Puccinia MCA-2864 Asteraceae (Helianthus) VA Uromyces ficariae AF426204 Ranunculus (Ranunculaceae)


Puccinia chondrillina U-202 Asteraceae (Chondrilla) Turkey [BPI extraction] Puccinia balsamorrhizae U-332Asteraceae (Balsamorhiza III?) B.C. Puccinia balsamorrhizae U-354 Asteraceae (Balsamorhiza) ID Puccinia balsamorrhizae U-355 Asteraceae (Balsamorhiza) ID Miyagia pseudosphaeria U-63 (BPI) Asteraceae (Sonchus oleraceus) Puccinia calcitrapae v. centaureae U-201 Asteraceae (Centaurea depressa) Turkey [BPI extraction]


Puccinia acroptili U-224(confirmed no.) Asteraceae (Acroptilon) Turkey Puccinia acroptili U-578 Asteraceae (Acroptilon repens III) NM Puccinia acroptili U-135 (Dana Berner 02-048) Asteraceae (Acroptilon repens II-III?) Turkey Puccinia acroptili U-224 Asteraceae (Acroptilon repens) Turkey Puccinia acroptili U-577 Asteraceae (Centaurea repens) NM Puccinia U-203 Asteraceae (Cardueae II-III) Turkey [host & rust confirmed from BPI] Puccinia punctiiformis U-225 Cirsium (Asteraceae) Turkey Puccinia punctiiformis U-225 Asteraceae (Cirsium arvense) Turkey Puccinia calcitrapae U-164 Asteraceae (Carthamus tinctorius II-III) Argentina Puccinia centaurea U-278 (FDWSRU 04-010) Asteraceae (Centaurea hybrid) OR Puccinia calcitrapae v. centaureae U-140 (Dana Berner 03-051) Asteraceae (Centaurea macedonica) Greece Puccinia calcitrapae v. centaurea U-209 Asteraceae (Centaurea calcitrapa) Turkey Puccinia carthami U-323 Asteraceae (Carthamus tinctorius III) Oman Puccinia atropuncta MCA2609 Veratrum (Liliaceae III) TN ?? U492 Antirrhinum (Scrophulariaceae) Germany


?? MCA2558 Phragmites? (Poaceae) Ecuador Endophyllum cassiae U-525 Fabaceae (Cassia III?) Nigeria ?? JRH-466 Cyperaceae Puerto Rico ?? U-265 Cyperus (Cyperaceae II?) Guam Puccinia cnici-oleracei JRH-200 Asteraceae (Melampodium divaricatum III) Honduras Puccinia cnici-oleracei JRH-207 Asteraceae (Malampodium divaricatum III) Honduras

Puccinia ursiniae U-463 (#590) Asteraceae (Ursinia) South Africa Puccinia tanaceti U-040 Artemisia (Asteraceae II-III) UT Puccinia rubefaciens? U-330 Galium (Rubiaceae III?) B.C. Canada

Puccinia medellinensis JRH-154 Lamiaceae (Hyptis II) Costa Rica Puccinia melampodii JRH138 Emilia (Asteraceae III) Costa Rica Uromyces rumicis U-227 Rumex (Polygonaceae) Turkey

Uromyces lespedezae-procumbentis U-007 Trifolium (Fabaceae III) MD Puccinia convolvuli? MCA2778 Calystegia (Convolvulaceae II) MD Puccinia convolvuli U-068 Calystegia (II-III Convolvulaceae) MD

Puccinia scleriicola GUY-57 Scleria (I Cyperaceae) Puccinia ibrae JRH-228 Convolvulaceae (Ipomoea I-III) Honduras

Puccinia thaliae JRH141 Canna (Cannaceae II) Costa Rica Puccinia thaliae GUY-3 Canna (Cannaceae II) Guyana

Puccinia kalchbrenneriana U-312 (#560) Asteraceae (Helichrysum III?) South Africa Puccinia kalchbrenneriana U-313 (#555) Asteraceae (Helichrysum III?) South Africa Puccinia kalchbrenneri? U-258 (#539) Asteraceae (Helichrysum III) South Africa Uromyces colombianus JRH-145 Asteraceae (Melanthera II-III) Costa Rica





Uromyces euphorbiae GUY-7 Euphorbia (II Euphorbiaceae) Uromyces pisi AF426201 Euphorbia Uromyces pisi-sativi? MCA2377 Euphorbia (Euphorbiaceae) Italy Uromyces? U-471 Euphorbiaceae (Euphorbia) Germany Endophyllum euphorbiae-sylvaticae AF426200 Euphorbia Puccinia abrupta JRH-146 Viguiera (Asteraceae II-III) Costa Rica


Puccinia stoboeae U-259 (#550) Asteraceae (Berkheya O,I,II) South Africa Uromyces hedysari-paniculati U-124 Desmodium (Fabaceae II-III ) Honduras Uromyces hedysari-paniculati U-052 Desmodium (Fabaceae II) Guyana Aecidium sp. U-211 Salsola (Chenopodiaceae I) Turkey

Uromyces ciceris-arietini U-183 Cicer (Fabaceae II-III ) SD Uromyces junci AF426203 Pulicaria (Asteraceae) Puccinia? MCA2593 (Asteraceae O?) ID Puccinia diplachnis MCA2375 Cirsium (I Asteraceae) Switzerland {T=Poaceae} Puccinia verruca U-130 Centaurea (Asteraceae III) Tunisia







Puccinia hordei MCA2391 (Poaceae) CA Puccinia? U-288 Poaceae (Pennisetum II) Guam Uromyces setaria-italicae JRH220 Melinis (II Poaceae) Hondorus Uromyces setaria-italicae JRH205 Panicum (II Poaceae) Hondorus

Melampsora laricis AB116809 Populus davidiana, Tibet, China Melampsora occidentalis U-395 Populus balsamifera II WA

Melampsora (SAL-49) [LSU only] Pinaceae (Abies grandis I) ID Pucciniastrum agrimoniae AF426234 Rosaceae (Agrimonia eupatoria)

Cronartium quercuum U-62 Fagaceae (Quercus macrocarpa) IN Cronartium quercuum U-432 Fagaceae (Quercus muehlenbergii II) IN Cronartium quercuum U-434 Fagaceae (Quercus alba II) IN Cronartium quercuum U-435 Fagaceae (Quercus bicolor II) IN Cronartium quercuum u-436 Fagaceae (Quercus macrocarpa II) IN Cronartium ribicola U-396/U-268 Ribes VA/Pinus WY Endocronartium harknessii AF522175

Cronartium flaccidum AF426239 Asclepiadaceae (Vincetoxicum hirundinaria) Endocronartium harknessii AY700193 Pucciniastrum agrimoniae? U-304 Rosaceae (Leucoside) South Africa [inc. seq.] Chrysomyxa arctostaphyli AF522163 Chrysomyxa arctostaphyli AY700192 Chrysomyxa ledi AF426246 Pinaceae (Picea abies) Chrysomyxa rhododendri AF426245 Ericaceae (Rhododendron ferrugineum) Caeoma tsukubaensis AB19124 Ericaceae (Rhododendron kaempferi) Japan

Thekopsora areolata AF426235 Rosaceae (Prunus padus) Hyalopsora U-361 undet. fern (Woodsiaceae?) UT

Hyalopsora polypodii AF426229 Dryopteridaceae (Cystopteris fragilis) Hyalopsora polypodii AY512852

Pucciniastrum? MCA2884 ?? Costa Rica Melampsoridium betulinum AF426228 Fagaceae (Betula pendula)

Naohidemyces vaccinii MCA 2780 Ericaceae (Vaccinium) WA Naohidemyces vaccinii AF426238 Ericaceae (Vaccinium uliginosum) Uredinopsis sp. AF522181 Uredinopsis filicina AF426237 Thelypteridaceae (Phegopteris connectilis) Milesina (as Milesia) scolopendri AF426236 Aspleniaceae (Asplenium scolopendrium)




Coleosporium ipomoeae JRH-195 Convolvulaceae (Ipomoea nil II) Costa Rica Coleosporium ipomoeae JRH-485 Convolvulaceae (Merremia quinquefolia II-III) Puerto Rico Coleosporium ipomoeae JRH-465 Convolvulaceae (Ipomoea triloba II) P.R. Coleosporium ipomoeae JRH-216 Convolvulaceae (Ipomoea cholulensis II-III) Honduras Coleosporium ipomoeae JRH-168 Convolvulaceae (Ipomoea trifida II-III) Costa Rica Coleosporium ipomoeae JRH-230 Convolvulaceae (Ipomoea anisomeres II) Honduras Coleosporium ipomoeae JRH-143 Convolvulaceae (Ipomoea neeiII) Costa Rica Coleosporium plumierae U-112 Apocynaceae (Plumeria rubra II-III) Nigeria Coleosporium plumeriae U-50 Apocynaceae (Plumeria rubra II) Guyana Coleosporium plumeriae U-266 Apocynaceae (Plumeria II) GUAM Coleosporium (asterum or delicatulum?) U-369 Asteraceae (Solidago) NY Coleosporium asterum U-67 Asteraceae (Symphyotrichum (=Aster) racemosum II-III) MD Coleosporium asterum AF522165 Coleosporium asterum AF426241 Asteraceae (Aster ciliolatus) Coleosporium (asterum or delicatulum?) MCA-2600 Asteraceae (Solidago II) TN Coleosporium U-430 Asteraceae (Solidago?) NJ Coleosporium U-389 Asteraceae (Solidago?) MD Coleosporium U-388 Asteraceae (Solidago?) PA Coleosporium (asterum or delicatulum?) U-428 Asteraceae (Solidago) MD Coleosporium (asterum or delicatulum?) MCA-2777 S(Asteraceae (Solidago canadensis II-III) MD Coleosporium asterum AF522164

Coleosporium sp. U-398 Asteraceae MD Coleosporium asterum U-366 Asteraceae (Aster novae-angliae II-III?) MD Coleosporium (asterum or delicatulum?) U-347 Asteraceae (Solidago) PA Coleosporium vernoniae U-371 Asteraceae (Vernonia) IL Coleosporium tussilaginis U-74 Asteraceae (Senecio triangularis) OR Coleosporium tussilaginis MCA-2389 Asteraceae (Sonchus?) CA Coleosporium tussilaginis MCA-2392 Asteraceae (Sonchus or Lactuca) CA Coleosporium vernoniae GUY-25 Asteraceae (Elephantopus mollis II) Coleosporium senecionis AY512840 Coleosporium tussilaginis AF426242 Asteraceae (Tussilago farfara) Coleosporium cacaliae AF426243 Asteraceae (Adenostyles glabra) Coleosporium campanulae AF426244 Campanulaceae (Campanula scheuchzeri) Thekopsora symphyti AF426230 Boraginaceae (Symphytum officinale) Thekopsora guttata AF426231 Rubiaceae (Galium odoratum)






Gerwasia (as Mainsia) pittieriana U-78 Rosaceae (Rubus) Guatemala Kuehneola uredinis U-429 Rosaceae (Rubus) MD Kuehneola uredinis U-303 Rosaceae (Rubus pinnatus) South Africa Kuehneola uredinis U-348 Rosaceae (Rubus) PA Kuehneola uredinis AY745696 AFTOL-987 Kuehneola uredinis AF426218 Rosaceae (Rubus fruticosus) Kuehneola uredinis MCA-2393 Rosaceae (Rubus III) MD Kuehneola uredinis MCA-2867 Rosaceae (Rubus) MD Kuehneola uredinis MCA-2863 Rosaceae (Rubus) VA Frommeella mexicana var. indicae U-3 Rosaceae (Duchesnea II) MD

FrommeÎlla mexicana MCA-2496 Rosaceae (Duchesnea) MD Frommeella mexicana MCA-2495 Rosaceae (Duchesnea II?) VA



Phragmidium rubi-idaei AF426215 Rosaceae (Rubus idaeus) Phragmidium fragariae AF426217 Rosaceae (Potentilla sterilis) Phragmidium montivagum AF426213 Rosaceae (Rosa cf. woodsii)

Triphragmium ulmariae AF426219 Rosaceae (Filipendula ulmaria) Ochropsora ariae AF426221 Ranunculaceae (Anemone nemorosa) Ochropsora ariae AF426222 Rosaceae (Aruncus dioicus)




Mikronegeria alba PUR-41 (Fagaceae) Argentina

Mikronegeria fuchsiae U-872 (Onagraceae III) New Zealand


Caeoma peltatum U-870 (Podocarpaceae I) New Zealand

Blastospora smilacis PUR-1 (Smilacaceae) Japan

Uredo fadogiae (=Hemileia fadogiae) U-900 (Rubiaceae) South Africa

Hemileia wrightiae U-1110 (Apocynaceae) India

Hemileia woodii U-897 (Rubiaceae) South Africa

Hemileia sp. MCA-3261 (Rubiaceae) Uganda

Hemileia gardeniae-thunbergiae U-461 (Rubiaceae) South Africa

Hemileia vastatrix MCA-2877 (Rubiaceae) Costa Rica

Hemileia vastatrix U-127 (Rubiaceae II) Mexico

Hemileia vastatrix JRH-236 (Rubiaceae II) Honduras

Hemileia scholzii U-1345 (Verbenaceae) South Africa

Maravalia sapotae JRH-172 (Sapotaceae II-III) Costa Rica

Caeoma torreyae AF522183

Caeoma torreyae U-808 (Taxaceae) CA

Caeoma torreyae U-1168 (Taxaceae I) CA

Helicobasidium purpureum

Eocronartium muscicola

Platygloea vestita

























































































































Basal rust lineages Mikronegeriaceae

+ Hemileia & Caeoma

torreyae

28S MP >50% support MP & ML >50% support MP or ML

L. Bonar. 1951. Two new fungi on Torreya. Mycologia 43:62-66

Data for Torreya from the Global Biodiversity Information Facility Network (http://zipcodezoo.com/Key/Plantae/Torreya_Genus.asp)

R. Peterson. 1974. Rust fungi with Caeoma-like sori on conifers. Mycologia 66: 242-255.

Caeoma torreyae

Torreya californica (Taxaceae s.l.) is endemic to restricted areas in CA

Caeoma torreyae on Taxaceae probably belongs to an undescribed primitive genus of Uredinales that is not derived from the supposedly ancestral Pucciniastraceae

Roger Peterson 1974





















Platygloea vestita

Ritschel A. 2005. Monograph of the genus Hemileia (Uredinales). Bibliotheca Mycologica 200:1-132.

Hemileia (ca. 42 species)

• No 0-I known • II-III on 10 host families (many

Rubiaceae) • Suprastomatal sori • Chaconiaceae (Cummins &

Hiratsuka 2003)





















Platygloea vestita

G.B. Cummins & Y. Hiratsuka. 2003. Illustrated genera of rust fungi, 3r Ed.

Mikronegeriaceae (3 genera, 13 species)

• All heteroecious (incl. newly described

M. fuchsiae) • II-III on Fagaceae & other families • O-I on conifers (Araucariaceae,

Podocarpaceae)

Rust life cycle variations

IV 0 I II III

0 = spermagonial I = aecial II = uredinial III = telial IV = basidiospore

macrocyclic

(hetero-/auto-)


IV 0 I II III


demicyclic

macrocyclic

(hetero-/auto-)

(hetero-/auto-)


IV 0 I II III

microcyclic


demicyclic

macrocyclic

(hetero-/auto-)

(hetero-/auto-)

Species A (heteromacro-form,

ancestral)

Aecial host (0, I; systemic)

e.g., Ranunculaceae

Telial host (II, III, IV)

e.g., Prunoideae

Species B (micro-form,

derived)

Former aecial host (0, III; systemic)

e.g., Ranunculaceae

Tranzschel’s Law (Tranzschel 1904)

reduction

species species A + A + B = B = correlated correlated speciesspecies

Species considered related to each other due to similarities in III morphology and a shared host, but with different life cycles. • Microcyclic species are derived • No microcyclic species in more “primitive” families (except for Chrysomyxa abietis) • Mostly confined to genera in the Pucciniaceae

























































































































Tranzschelia fusca (= T. anemones) AF426225 (Anemone nemorosa) Germany Tranzschelia fusca (KR 12289) (Anemone nemorosa) Germany

Tranzschelia pruni-spinosae AF426224 (Anemone ranunculoides) Tranzschelia pruni-spinosae (KR-0010972) (Prunus cerasifera) Iran

Tranzschelia pruni-spinosae (KR 12356) (Prunus cerasifera) Iran Tranzschelia pruni-spinosae (KR/ H. Jage 450/04) (Anemone ranunculoides) Germany Tranzschelia pruni-spinosae (KR/ H. Jage 1508/04) (Prunus spinosa) Germany

Tranzschelia fusca (MIN 885891) (Anemone quinquefolia) USA Tranzschelia fusca (MIN 885887) (Anemone quinquefolia) USA Tranzschelia fusca (Anemone quinquefolia) USA

Tranzschelia pruni-spinosae (MIN 885892) (Anemone quinquefolia) USA Tranzschelia pruni-spinosae s. l. (KR 12108) (Prunus) Japan

Tranzschelia pulsatillae (KR 24825)(Pulsatilla halleri) Slovakia Tranzschelia hyrcanica U-512 (KR-0010968) (Prunus cerasifera) Iran Tranzschelia hyrcanica U-513 (KR-0010969) (Prunus cerasifera) Iran Tranzschelia hyrcanica U-514 (KR-0010970) (Prunus cerasifera) Iran Tranzschelia hyrcanica U-515 (KR-0010971) (Prunus cerasifera) Iran

Tranzschelia sp. (MIN 885888) (Anemone quinquefolia I) USA

Tranzschelia spec. (KR 12357) (Prunus cf. serotina) USA

Tranzschelia arthurii? MCA-2865 (Prunus serotina) USA Tranzschelia arthuri? U-179 Rosaceae USA

Tranzschelia arthuri? U-179 Rosaceae USA Tranzschelia sp. MCA-3588 Rosaceae USA

Tranzschelia arthuri? U-179 Rosaceae USA Tranzschelia pr.-sp. var. americana (MIN 885890) (Prunus pensylvanica) USA

Tranzschelia pr.-sp. var. americana (Anemone quinquefolia) USA Tranzschelia pr.-sp. var. americana (MIN 885890) (Prunus pensylvanica) USA

Tranzschelia sp. (DAOM 223134) (Anemone quinquefolia) Canada Tranzschelia discolor AF426223 (Prunus domestica) Germany

Tranzschelia discolor JRH-139 (Prunus domestica) Costa Rica Tranzschelia discolor U-884 (Prunus persica) Oman Tranzschelia discolor (KR-0010962) (Anemone coronaria) Greece Tranzschelia discolor (KR-0010965) (Prunus persica) Iran Tranzschelia discolor (KR-0010966) (Prunus domestica) Iran Tranzschelia discolor (KR-0010973) (Prunus persica) Iran Tranzschelia discolor (KR-0010833) (Prunus domestica) Germany

Tranzschelia discolor (KR-0010963) (Anemone coronaria) Greece Tranzschelia discolor (KR-0010974) (Prunus cerasifera) Iran Tranzschelia discolor (KR 13368) (Prunus dulcis) Iran Tranzschelia discolor (KR-0010961) (Anemone coronaria) Greece Tranzschelia discolor (KR-0010960) (Prunus domestica) Germany Tranzschelia iranica (KR 12352) (Prunus spec.) Iran Tranzschelia iranica (KR 12353) (Prunus spec.) Iran

Tranzschelia arasbaranica (KR 0012355) (Prunus mahaleb) Iran Tranzschelia microcerasi (KR 12351) (Prunus microcarpa) Iran

Tranzschelia thalictri (DAOM 187607) (Thalictrum polygamum) Canada Tranzschelia thalictri (KR 12349) (Thalictrum aquilegiifolium) Germany

Tranzschelia thalictri (KR V. Kummer 156/4) (Thalictrum minus) Germany Tranzschelia asiatica (KR 12106) (Prunus grayana) Japan

Leucotelum cerasi (KR-24931) (Eranthis hyemalis O,I) Austria Puccinia podophylli U-803 (Podophyllum peltatum) USA

Ochropsora ariae U-1036 (Anemone O,I) Germany Ochropsora ariae U-1036 (Anemone O,I) Germany

100

98

98

91

71

75

82

94

53 94

98

89

69

Tranzschelia

micro-form: 0,III,IV

macro-form on aecial host: 0,I (Ranunculaceae) on telial host: II,III,IV (Prunoideae)

Life Cycle Types

Dataset: Scholler & Aime, unpublished.


























































































































Tranzschelia pruni-spinosae AF426224 (Anemone ranunculoides) Germany Tranzschelia pruni-spinosae (KR-10972) (Prunus cerasifera) Iran




Tranzschelia pulsatillae (KR 24825) (Pulsatilla halleri) Slovakia

53 94

98

89

69

Tranzschelia



Life Cycle Types


2 corr. species

Correlated species:

T. fusca s. l. T. pruni-spinosae s. l.

Europe/SW Asia


























































































































Tranzschelia pruni-spinosae AF426224 (Anemone ranunculoides) Germany Tranzschelia pruni-spinosae (KR-10972) (Prunus cerasifera) Iran




Tranzschelia pulsatillae (KR 24825) (Pulsatilla halleri) Slovakia

53 94

98

89

69

Tranzschelia


2 corr. species

2 corr. species

Correlated species:

T. fusca s. l. T. pruni-spinosae s. l.

North America



Life Cycle Types

























































































































Uromyces plumbarius

(autoecious on Onagraceae)

and Puccinia dioicae

(heteroecious on

Onagraceae/Cyperacea)

Uromyces plumbarius

Correlated species (as P. heteranthae)? Orton. 1912. Mycologia 4:203. These data: Kaur et al. submitted; Dixon et al. 2011. Mycol. Prog. 9:459-468.

























































































































Data: Dixon et al. 2011. Mycol. Prog. 9:459-468.

Sugarcane rusts

Sugarcane rusts

























































































































Data: Dixon et al. 2011. Mycol. Prog. 9:459-468.

P. miscanthi – heteromacrocyclic; aecial

host is Plantago spp. (Plantaginaceae)

P. substriata & P. agrophila –

heteromacrocyclic; aecial host is Solanum spp.

(Solanaceae)

Robert StackRobert Stack Dave & Ellen Dave & Ellen FarrFarr Tom BrunsTom Bruns Roger PetersonRoger Peterson BernaBerna TunaliTunali Walter KaiserWalter Kaiser Curt W. Curt W. RoaneRoane Reinhard BerndtReinhard Berndt Ed LickeyEd Lickey Dana Dana BernerBerner John John McKemyMcKemy Alan WoodAlan Wood Michael DeadmanMichael Deadman Amy RossmanAmy Rossman William BruckartWilliam Bruckart Harry EvansHarry Evans Linley SmithLinley Smith Gunter Gunter ArnoldArnold RadamesRadames UrtiagaUrtiaga James James GinnsGinns Larry LittlefieldLarry Littlefield R. Gopal R. Gopal Kapooria Kapooria John HartJohn Hart Andrea RomeroAndrea Romero John OliveJohn Olive George Newcombe George Newcombe ChandalinChandalin Bennett Bennett TaloTalo PastorPastor--CorralesCorrales Min Min RayamajhiRayamajhi Steve Steve RehnerRehner Libby RobertsLibby Roberts AndrzejAndrzej ChlebickiChlebicki MishaMisha SogonovSogonov John John PlischkePlischke Allison Allison Kennedy Kennedy Don FerrinDon Ferrin Patricia BollichPatricia Bollich Chris ClarkChris Clark

••Markus Scholler, Markus Scholler, StaatlichesStaatliches Museum f. Museum f. NaturkundeNaturkunde, , Karlsruhe, Germany (Karlsruhe, Germany (TranzscheliaTranzschelia)) ••David Hibbett, Brandon David Hibbett, Brandon Matheny, Daniel Henk, David Matheny, Daniel Henk, David McLaughlin (Pucciniomycotina)McLaughlin (Pucciniomycotina) ••Aime Lab: Cindy Park, Aime Lab: Cindy Park, Ramandeep Kaur, Merje Toome, Ramandeep Kaur, Merje Toome, Alistair McTaggart, Andrew Alistair McTaggart, Andrew RodriguezRodriguez

Collaborators:Collaborators: Collections:Collections:

••Assembling the Fungal Tree Assembling the Fungal Tree of Life National Science of Life National Science Foundation DEB 0732968Foundation DEB 0732968 ••USDAUSDA--APHIS Cooperative APHIS Cooperative agreement 10agreement 10--81008100--15301530--CACA ••Louisiana Board of Regents Louisiana Board of Regents LEQSF (2009LEQSF (2009--12)_RD_A12)_RD_A--0101

Funding:Funding:

Overview of Rust Fungi - Plant Management Network€¦ · •ca. 7000 species ... and comprise 22...

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