A bird’s eye view of the fungal Kingdom A quest for nomenclatural stability

Sybren de Hoog CBS-KNAW Fungal Biodiversity Centre, Utrecht, The Netherlands Histoplasma capsulatum on Ajellomyces capsulatum routine media: anamorph on 1/10 SGA with salts: teleomorph

Histoplasma capsulatum

Ascomycota

Higher fungi Higher

Coelomycetes Hyphomycetes

Basidiomycota

Glomerulomycota Zoopagomycotina Entomophthoromycotina Mucoromycotina

Chytridiomycota lower fungi lower

Ascomycota

Higher fungiHigher Coelomycetes

Basidiomycota

Glomerulomycota Zoopagomycotina Entomophthoromycotina Mucoromycotina

Chytridiomycota Lower fungi Lower E urotiales

Onygenales Sporothrix

Chaetothyrial es

Sordariales Ophiostomatales Diaporthales X ylarial es

Hypocreales

Microascales

Lecanorales Ascomycota Ascomycota Dothideales Mycocal ici um albonigrum

Pleosporales “Dematiaceae” Molecular siblings Leotiales Geomyces pannorum Leotiales

P ezizales

Calicium tricolor

Hemiascomycetes

Pneumocystis carini i Schizosaccharomyces pombe

Uredinales

Sporidiales Til letial es Quambalaria Ustilagi nales

Basidiomycota Tremel lales Basidiomycota

S tereales

A garicales

Glom ales Linderina pennispora Glomerulomycota Cantharellus tubaeformis Harpellales Zoopagomycotina Entomophthorales Entomophthoromycotina Zygomycota Mucorales Mucoromycotina Mortierellales B asidi obolus ranarum Chytridiomycota Chytridiales Chytridium confervae Chytridiomycota EF081388 Passalora bellynckii GQ852618 Dothistroma septosporum CBS 383 74 EF081387 Passalora bellynckii Mycosphaerella microsora EU167599 EF081386 Mycosphaerella microsora Pseudocercosporella bakeri CPC 17570 61 Passalora fulva DQ008163 GU137910 Passalora fulva Passalora ageratinae EF081431 Dothistroma septosporum CBS 112498 Mycosphaerella pini GQ852597 EF081448 Dothistroma septosporum GQ852597 56 Dothistroma septosporum CBS 543 74 EF081435 Dothistroma pini GQ852596 Dothistroma pini EF081438 Mycosphaerella africana Mycosphaerella aurantia DQ246256 HQ454509 22 Mycosphaerella buckinghamiae EU707856 Mycosphaerella ellipsoidea HQ454514 61 Mycosphaerella ellipsoidea GQ852602 Passalora brachycarpa 71 Passalora brachycarpa GQ852619 EU704083 Passalora sp GQ852623 Passalora daleae AY350578 754 Passalora daleae EU040236 No No medical/No of total of 1000 ( Bayesian,LSU Mycosphaerella keniensis AY571381 Mycosphaerella graminicola CBS 115943 1/0.96/86 Ramularia pratensis var pratensis HQ454510 Ramularia brunnea 8265 Ramularia aplospora EF081382 61 Ramularia nagornyi Ramularia miae 37 Mycosphaerella punctiformis CBS 113265 FJ469668 57 81 Ramularia endophylla 66 Lecanosticta acicola HQ454519 76 Phaeophleospora eugeniicola 65 Phaeophleospora eugeniae JF499854 Penidiella nectandrae 59 Mycosphaerella endophytica CBS 114662 JF499856 63 Mycosphaerella gregaria 44 Mycosphaerella stromatosa EU019261 Mycosphaerella pseudoendophytica DQ246253 Zasmidium aerohyalinosporum JF499859 96 Brunneosphaerella protearum CPC 13905 Passalora vaginae EU343546 71 Mycosphaerella rosigena 14 Ramichloridium pini 25 Phaeophleospora concentrica HQ693526 67 Mycosphaerella marksii GQ852614 96 Mycosphaerella marksii CBS 110942 JF770463 80 Mycosphaerella intermedia DQ246248 Zasmidium nabiacense JF770462 Mycosphaerella parkii Ramichloridium biverticillatum JF499858 84 13 Mycosphaerella madeirae 84 bootstrap ( Mycosphaerella madeirae DQ204756 DQ204745 50 Microcyclosporella mali CPC 16171 Periconiella arcuata Verrucisporota daviesiae DQ204753 85 47 83 48 Verrucisporota daviesiae CBS 116002 Periconiella velutina DQ204746 85 57 Verrucisporota proteacearum Rasutoria pseudotsugae DQ204755 Rasutoria tsugae 92 Rasutoria tsugae ratstk DQ204748 12 Ramichloridium musae 1/0.86/97 82 Zasmidium anthuriicola DQ204750 Zasmidium citri 8673 Ramichloridium australiense DQ204768 30 Ramichloridium strelitziae Zasmidium cellare DQ204756 47 Mycosphaerella aleuritidis 1/0.92/90 66 Zasmidium nocoxi GU214400 64 Ramichloridium cerophilum Ramichloridium cerophilum CBS 103 59 Polythrincium trifolii EU167613 DQ678050 Passalora dalbergiae Pseudocercospora vitis DQ073923 dH13831 Pseudocercospora sphaerulinae Pseudocercospora vitis CPC 11595 dH13868 61 Pseudocercospora bixae Pseudocercospora robusta dH13840 Pseudocercospora pseudoeucalyptorum 1/0.94/99 49 Pseudocercospora gracilis 18 dH13836 Pseudocercospora eucalyptorum Pseudocercospora basitruncata dH13862 14 Pseudocercospora crousii 62 Pseudocercospora fori dH13880 78 Pseudocercospora natalensis DQ267576 Pseudocercospora platani 68 Pseudocercospora fijiensis DQ678060 Pseudocercospora fijiensis OSC 100622 82 Mycosphaerella milleri DQ767658 6649 Pseudocercospora basiramifera 41 Pseudocercospora paraguayensis GQ852634 1/0.97/94 DQ678094 Pseudocercospora schizolobii 73 Pseudocercospora tereticornis DQ678095 Pseudocercospora punctata Mycosphaerella pyri 98 AY016365 74 Pseudocercospora griseola f griseola Phaeocryptopus gaeumannii dH13845 Phaeocryptopus gaeumannii CBS 267 37 Mycosphaerella heimii CBS 110682 GU048607 Mycosphaerella konae 88 Mycosphaerella heimioides Mycosphaerella heimii HQ026799 Mycosphaerella irregulariramosa 74 45 Mycosphaerella holualoana HQ026798 Mycosphaerella crystallina Mycosphaerella acaciigena 624 GU238160 86 Mycosphaerella colombiensis Passalora graminis CBS181.58 Phaeophleospora atkinsonii CBS 124565 S. arenarium Sonderhenia eucalypticola AY849958 89 Sonderhenia eucalyptorum Cercosporella virgaureae GQ852585 Passalora eucalypti Kittipan AY849959 61 Ramulispora sorghi GQ852653 Mycosphaerella latebrosa CBS142.60S. salinum 85 Mycosphaerella latebrosa CBS 687 94 85 Septoria berberidis AY849955 Mycosphaerella ribis 74 Mycosphaerella harthensis AY787937 Mycosphaerella flageoletiana Septoria populicola AY849954 86 Septoria apiicola GQ852674 1/0.93/97 69 Pseudocercosporella capsellae GU214662 Cercospora zebrinae AY849946 61 Cercospora beticola CBS 116456 AY787931 78 Cercospora beticola Cercospora apii Septoria rubi AY849944 50 Septoria convolvuli GQ852675 Septoria senecionis GQ852678 AY849945 64 Septoria leucanthemi GQ852677 bp Mycosphaerella coacervata AY849956 69 Mycosphaerella linorum 1/0.74/93 Schizothyrium pomi CBS 228 57 CBS734.96 Zygophiala cryptogama S. salinum 97 Ramichloridium apiculatum CBS 400 76 AY849957 Dissoconium aciculare CBS 204 89 18 87 Dissoconium australiensis EU673244 86 64 Dissoconium dekkeri CBS 111282 86 Dissonium dekkeri CBS 111282 EU673245 51 Dissoconium commune CBS 110747 83 Dissoconium commune EU019267 Cladosporium cladosporioides CBS112388 EU673243 Cladosporium hillianum CBS125988 ) of of ) Cladosporium iridis CBS 138 40 AY004336 Cladosporium herbarum CBS 399 80 74 Davidiella allicina EU673229 Davidiella macrospora Cladosporium iridis CBS138 40 EU673230 42 Cladosporium herbarum CBS121621 0.96/-/- Sphaerulina polyspora JF441219 59 Cladosporium bruhnei Cladosporium allicinum CBS18854 JF441205 71 Melanodothis caricis 83 Cladosporium cladosporioides CBS 170 54 Cladosporium fusiforme CBS119414 91 GU301815 Toxicocladosporium chlamydosporum 1/0.79/100 Toxicocladosporium irritans GU301867 78 Toxicocladosporium rubrigenum Samerpitak 39 CCFEE 5398 FJ469672 59 Rachicladosporium cboliae CBS 119432 5211 EF175664 91 CCFEE 5395 88 82 CCFEE 5458 EF175658 Rachicladosporium luculiae 1/0.98/98 14 Verrucocladosporium dirinae EF175662 Graphiopsis chlorocephala

324 83 CCFEE 5388 CCFEE 5456 EF175666 98 CCFEE 5527 AY541493 Anisomeridium consobrinum 59 Devriesia strelitziicola GU214417 CBS118321 TRN287 GU301855 93 CCFEE 5413 Mycosphaerella eurypotami JK 5586J CBS850.73 92 Sporidesmium pachyanthicola Hortaea acidophila CBS 113389 CBS100438 92 CBS118302 TRN142 Devriesia hilliana GU214414 CBS130.65 Devriesia lagerstroemiae GU214415 73 64 Devriesia lagerstroemiae CBS125422 CBS228.66 CCFEE 5681 Passalora sp GQ852622 31 Devriesia queenslandica CBS129527 CBS383.81 Devriesia agapanthi CBS132689 69 CBS118285 TRN81 CBS510.71 61 Teratosphaeria knoxdavesii 8161 Teratosphaeria knoxdavesii EU707865 CBS225.77 CCFEE 5672 . . 76 Devriesia strelitziae CBS 122379 CBS203.27 45 Tripospermum myrti CCFEE 5582 CBS175.65 57 Cystocoleus ebeneus EU048571 Cystocoleus ebeneus EU048573 CBS131815 99 Cystocoleus ebeneus L315 CBS118287 TRN77 43 99 CBS118301 TRN138 CBS284.64 CBS118305 TRN62 CBS118608 TRN139b

CBS131816 TRN61 86 CBS118283 TRN124 CBS435.76 CBS118296 TRN114 97 CCFEE 5488 CBS211.53 8 CBS118284 TRN104 CCFEE 5489 CBS202.27 CCFEE 5459 44 CBS118294 TRN111 CBS239.78 Devriesia americana EU040227 94 Staninwardia suttonii 1049 CBS102491 Staninwardia suttonii DQ923535 39 Devriesia pseudoamericana CPC 16174 48 Ramichloridium brasilianum CBS116655 66 Pseudoramichloridium henryi GQ303320 61 CBS118292 TRN96 CBS100442 34 CBS118300 TRN137 CCFEE 451 0.80/-/- CBS729.95 89 CCFEE 5207 88 CCFEE 5312 CBS100486 98 Phacellium paspali Phacellium paspali CBS 113093 CBS412.51 74 Aulographina pinorum CBS 174 90 Catenulostroma abietis CBS 459 93

). 97 CBS172.74 Catenulostroma microsporum EU019255 68 Catenulostroma germanicum Aulographina pinorum CBS 302 71 CBS475.80 Xenomeris juniperi CCFEE 5522 CBS780.83 47 99 CCFEE 5523 Pseudocercosporella fraxini GU214682 1/0.98/99 CBS423.64 61 Capnobotryella renispora CBS 215 90 Penidiella strumelloidea EU019277 83 CBS536.69 Microcyclospora tardicrescens CPC 16187 81 Microcyclospora malicola CPC 16172 76 CBS118.91 70 Microcyclospora pomicola CPC 16173 Penidiella kurandae EU040214 CBS437.64 Devriesia thermodurans EU040229 75 81 Devriesia acadiensis CBS 115874 CBS867.95 92 Devriesia staurophora 375 81 50 Devriesia shelburniensis CBS 115876 CBS547.81 75 Devriesia shelburniensis EU040228 1/0.88/51 90 Recurvomyces mirabilis CCFEE 5264 Recurvomyces mirabilis CCFEE 5475 65 CBS864.95 95 CCFEE 5575 CCFEE 5401 CBS125818 Catenulostroma protearum CPC 15368 Readeriella eucalypti CBS125817 64 Readeriellaa readeriellophora Readeriella pseudocallista CBS119775 33 Readeriella callista 1/0.97/84 86 Readeriella dimorphospora HQ634611 Readeriella angustia 58 Readeriella minutispora 67 Readeriella nontingens HQ634610 Readeriella eucalyptigena 75 Readeriella mirabilis DQ885904 Readeriella dendritica 95 60 Readeriella patrickii DQ885905 Readeriella tasmanica 11 Readeriella novaezelandiae DQ885906 Readeriella menaiensis Teratosphaeria mexicana Sympoventuriaceae 89 Teratosphaeria pseudosuberosa EU035424 86 Teratosphaeria suberosa CPC 11032 1/0.97/99 85 EU035423 66 Teratosphaeria suberosa DQ246235 Phaeothecoidea intermedia CBS469.95 Phaeothecoidea eucalypti S. excentricum 93 Phaeothecoidea minutispora 59 45 CCFEE 502 CBS588.66 Teratosphaeria parva Veronaeopsis simplex Baudoinia compniacensis EU035435 74 69 Catenulostroma elginense CBS 111030 53 CBS119304 TRN440 EU035464 79 Teratosphaeria associata CBS 112224 89 Teratosphaeria jonkershoekensis EU035456 Friedmanniomyces endolithicus CCFEE 670 Meristematico nero 5176 group5282 1/0.94/94 DQ470974 Friedmanniomyces endolithicus CCFEE 5310 CCFEE 5385 EU035447 80 21 Friedmanniomyces endolithicus CCFEE 5305 Friedmanniomyces endolithicus CCFEE 524 CCFEE 5530 GU323212 39 CCFEE 5393 CCFEE 5328 GU456336 61 CCFEE 5283 CCFEE 5396 EF114712 Friedmanniomyces endolithicus CCFEE 5273 93 93 Friedmanniomyces endolithicus CCFEE 5180 31 EU035433 72 Friedmanniomyces endolithicus CCFEE 5199 43 Friedmanniomyces endolithicus CCFEE 5311 EU035473 Friedmanniomyces endolithicus CCFEE 5195 31 Piedraia hortae var hortae CBS 374 71 Piedraia quintanilhae

Phylogeny of GU348998 90 Penidiella pseudotasmaniensis GQ852625 HQ599579 998 Penidiella tasmaniensis DQ246233 22 CBS118768 TRN232 Penidiella columbiana EU035468 CBS117941 TRN262 86 Penidiella eucalypti EF114694 47 Penidiella tenuiramis 73 Teratosphaeria ohnowa JQ036232 81 Teratosphaeria flexuosa 94 71 Teratosphaeria secundaria EU035462 Venturiaceae CBS118356 Capnodiales TRN125 CBS117950 TRN431 GU301814 92 CBS117930 TRN79 64 CBS118766 TRN159 1/0.71/97 CBS117937 TRN211 EU035443 28 CCFEE 5499 67 CCFEE 5492 EF114692 CCFEE 5394 6269 CCFEE 5389 GU301878 Elasticomyces elasticus CCFEE 5526 Elasticomyces elasticus CCFEE 5505 EU035454 66 Elasticomyces elasticus CCFEE 5506 Elasticomyces elasticus CCFEE 5537 JQ036233 71 Elasticomyces elasticus CCFEE 5538 Elasticomyces elasticus CCFEE 5525 EF114695 18 Elasticomyces elasticus CCFEE 5474 46 Elasticomyces elasticus CCFEE 5490 JF263534 Elasticomyces elasticus CCFEE 5543 71 Elasticomyces elasticus CCFEE 5547 74 Batcheloromyces proteae 0.57/-/- HQ613813 69 Batcheloromyces proteae CBS 110696 18 Teratosphaeria toledana HQ599589 Teratosphaeria blakelyi 78 Teratosphaeria suttonii CBS210.95 75 Teratosphaeria zuluensis S. fusarioideum 34 CBS117932 TRN123 AF050275 96 Teratosphaeriaceae sp TRN 123 Teratosphaeria veloci EU041874 81 Teratosphaeria gauchensis Teratosphaeria stellenboschiana EU019295 EU041872 89 Teratosphaeria stellenboschiana GQ852716 Teratosphaeria stellenboschiana CBS 1 Teratosphaeria nubilosa EU041873 66 Teratosphaeria molleriana 7836 Teratosphaeria angophorae EU041875 CCFEE 5500 EU107312 Teratosphaeria considenianae 93 Teratosphaeria majorizuluensis 96 Teratosphaeria dimorpha AY004338 96 Teratosphaeria profusa Teratosphaeria maculiformis AFTOL668 84 Teratosphaeria corymbiae 28 Teratosphaeria proteaearboreae CBS769.83 33 Teratosphaeria fibrillosa CBS 121707 1/0.78/94 S. cateniphorum 43 Teratosphaeria fibrillosa EU019282 92 Teratosphaeria fibrillosa CPC 1876 FJ358231 Teratosphaeria hortaea FJ790300 71 Teratosphaeria miniata FJ358225 20 Teratosphaeria hortaea FJ790299 31 Teratosphaeria complicata AF050263 76 Teratosphaeria brunneotingens Mycosphaerella cryptica DQ329021 89 Teratosphaeria cryptica 87 Teratosphaeria cryptica CBS 110975 EF643750 80 Teratosphaeria juvenalis Teratosphaeria verrucosa AY300828 Teratosphaeria viscidus 81 Teratosphaeria destructans 65 AY584644 83 Teratosphaeria eucalypti Teratosphaeria alboconidia 1/0.70/93 DQ329014 87 Teratosphaeria macowanii Teratosphaeria maxii 97 DQ782913 Teratosphaeria ovata 80 CCFEE 5734 1/0.88/65 67 Hortaea werneckii EU019270 EF105141 99 61 Hortaea werneckii CBS 100496 CBS117931 TRN122 EF105138 23 Stenella araguata Penidiella venezuelensis EF105132 69 CCFEE 5457 8678 CCFEE 5507 DQ329015 CCFEE 5508 Hortaea thailandica EF411063 57 CBS118355 TRN500 55 CBS118764 TRN126 AF279407 86 CCFEE 5569 1/0.94/95 5 Antennariella placitae AY607737 68 Condioxyphium gardeniorum CPC 14327 78 Microxyphium aciculiforme CBS 892 73 Condioxyphium gardeniorum AY640962 82 Capnodium coffeae CBS 147 52 1/0.85/59 69 Microxyphium theae CBS 202 30 9 DQ361098 86 TRN450 Polychaeton citri DQ361104 44 Leptoxyphium fumago CBS 123 26

Ascomycota Microxyphium citri CBS 451 66 DQ361102 81 TRN87 Phaeotheca fissurella 99 Comminutispora agavaciensis U93304 45 Comminutispora agavaciensis CBS 619 95 TRN437 DQ239894 97 Capnodium salicinum CBS 131 34 85 Fumagospora capnodioides DQ239886 CCFEE 5772 Scorias spongiosa CBS 325 33 0.95/0.92/86 AB075328 9 80 Racodium rupestre EU048576 1/0.92/82 47 Racodium rupestre L423 AB075330 Racodium rupestre EU048575 98 Racodium rupestre EU048577 73 AB075326 70 Racodium rupestre L424 CCFEE 5559 AB075327 99 CCFEE 5578 Phaeotheca triangularis CCFEE 5579 AB075329 CCFEE 5540 TRN444 AY004346 69 Elsinoe centrolobi CBS 222 50 78 Elsinoe phaseoli CBS 165 31 GQ911144 45 Elsinoe veneta CBS 150 27 TRN235 1/-/- 74 U29571 TRN236 94 Phaeosclera dematioides CBS 157 81 78 U29570 98 Endosporium aviarium UAMH 10530 40 Endosporium populitremuloidis UAMH 10529 Myriangium duriaei CBS 260 36 U29572 45 29 99 Myriangium hispanicum CBS 247 33 CCFEE 5409 GU808560 81 CCFEE 5545 1/0.98/100 CCFEE 5469 AY213700 CCFEE 5496 68 CCFEE 5491 AY176751 CCFEE 5470 CCFEE 5466 FJ864710 11 46 CCFEE 5460 CCFEE 5416 AY853379 CCFEE 5462 99 CCFEE 5461 AY796003 CCFEE 5477 68 CCFEE 5465 CCFEE 5495 AY796004 97 TRN11 87 TRN258 1/0.97/75 AY796001 TRN272 94 97 TRN275 AY796002 65 TRN37 CCFEE 5560 EF413632 69 TRN131 95 62 Columnosphaeria fagi CBS 171 93 AF356689 64 Aureobasidium pullulans CBS 584 75 96 Kabatiella caulivora CBS 242 64 DQ832334 CCFEE 5558 97 CCFEE 5552 34 CCFEE 5553 1/1/100 DQ832330 83 TRN9 99 Dothiora cannabinae CBS 737 71 AF356693 Dothiora ellyptica CBS 736 71 11 Phaeocryptopus nudus CBS 268 37 DQ119558 96 Delphinella strobiligena CBS 735 71 95 Sydowia polyspora CBS 116 29 DQ119564 72 Dothidea insculpta CBS 189 58 97 Dothidea sambuci DAOM 231303 DQ119562 87 Dothidea insculpta Stylodothis puccinioides CBS 193 58 DQ119571 68 TRN24 24 TRN278 DQ119570 TRN15 62 TRN16 94 TRN268 DQ119569 Astrothelium cinnamomeum DUKE 0000007 Laurera megasperma AFTOL 2094 CBS549.74 99 49 Trypethelium nitidiusculum AFTOL 2099 Neta Delitschia winteri CBS 225 62 DQ075680 Delitschia didyma UME 31411 76 98 Delitschia didyma UME 31411v2 JN036557 Zopfia rhizophila CBS 207 26 salmonicolor 99 Aigialus grandis 2Q 78 AY489721 76 Aigialus grandis JK 5244A 45 Aigialus parvus A6 AY489723 Ascocratera manglicola JK 5262C 0.99/0.94/59 Rimora mangrovei JK 5246A 90 Anteaglonium globosum ANM 925 2

U68127 62 7 Anteaglonium globosum SMH 5283 58 Anteaglonium parvulum SMH 5210 U68128 87 Anteaglonium abbreviatum ANM 925 1 80 Anteaglonium abbreviatum GKM 1029 AY882374 Anteaglonium latirostrum L100N 2 Tetraplosphaeria sasicola MAFF 239677 AY882373 68 Gibbera conferta CBS 191 53 0.99/-/57 80 29 Venturia inaequalis CBS 815 69 63 AY882372 Tyrannosorus pinicola CBS 124 88 94 Venturia populina CBS 256 38 AY882371 31 Apiosporina collinsii CBS 118973 41 45 Apiosporina morbosa dimosp AY882377 Astrosphaeriella aggregata MAFF 239486 Astrosphaeriella bakeriana CBS 115556 AY882375 87 Schismatomma decolorans DUKE 0047570 48 92 Wicklowia aquatica AF289 1 1/0.98/99 69 Wicklowia aquatica CBS 125634 49 EF151316 Mauritiania rhizophorae BCC 28867 Quintaria lignatilis CBS 117700 EU862178 Quintaria submersa CBS 115553 68 Lindgomyces ingoldianus ATCC 200398 66 HM449060 47 Lindgomyces rotundatus HHUF 27999 64 Lindgomyces breviappendiculata HHUF 2 HM449058 50 Massariosphaeria grandispora CBS 613 86 47 Ulospora bilgramii CBS 110020 GU563359 78 Lepidosphaeria nicotiae CBS 101341 15 Amniculicola parva CBS 123092 HM449063 Massariosphaeria typhicola CBS 123126 1/0.83/51 48 Lophiotrema bruneusporum CBS 123095 6 Lophiotrema lignicola CBS 122364 1/0.97/96 GU563363 93 Lophiotrema nucula CBS 627 86 Verruculina enalia JK 5253A 43 EF100924 Byssolophis sphaerioides IFRDCC2053 21 Quadricrura septentrionalis CBS 125429 EF100957 47 Polyplosphaeria fusca MAFF 239685 1/1/100 66 Triplosphaeria maxima MAFF 239682 AJ580945 Pseudotetraploa curviappendiculata MA 89 79 Westerdykella angulata CBS 610 74 AJ579686 97 Westerdykella cylindrica CBS 454 72 15 Westerdykella ornata CBS 379 55 DQ268616 90 Preussia lignicola CBS 264 69 82 Sporormiella minima CBS 524 50 Floricola striata JK 5678I DQ268615 76 Preussia funiculata CBS 659 74 18 86 89 Preussia terricola DAOM 230091 DQ268617 77 Lophiostoma alpigenum GKM 1091b Lophiostoma fuckelii GKM 1063 DQ294353 Lophiostoma compressum IFRD 2014 99 Lophiostoma arundinis CBS 621 86 AB363791 98 Lophiostoma caulium CBS 624 86 69 Lophiostoma caulium CBS 623 86 10 DQ821538 7 Lophiostoma crenatum CBS 629 86 1/0.80/84 Massariosphaeria phaeospora CBS 611 86 DQ821537 89 Falciformispora lignatilis BCC 21117 76 Falciformispora lignatilis BCC 21118 Halomassarina thalassiae JK 5262D HQ413647 97 75 Roussoella pustulans MAFF 239637 82 Roussoellopsis sp MAFF 239638 DQ368627 43 Roussoella hysterioides CBS 125434 AB237812 95 Arthopyrenia salicis 1994 Coppins 45 86 Arthopyrenia salicis CBS 368 94 Roussoella hysterioides MAFF 239636 AB237811 34 Melanomma rhododendri ANM 73 Pleomassaria siparia CBS 279 74 GU198953 30 Melanomma pulvispyrius CBS 371 75 44 Beverwykella pulmonaria CBS 283 53 2650 AB237826 57 Monotosporella tuberculata CBS 256 84 2352 Thyridaria rubronotata CBS 419 85 AB197135 59 Julella avicenniae BCC 20173 36 Corynespora cassiicola CBS 100822 99 Corynespora smithii CABI 5649b AB237820 Massaria platani CBS 221 37 Katumotoa bambusicola MAFF 239641 AB237819 95 Lentithecium aquaticum CBS 123099 Keissleriella cladophila CBS 104 55 AB237816 92 Ophiosphaerella sasicola MAFF 239644 Lentithecium fluviatile CBS 122367 1/0.98/100 69 AB237813 Asteromassaria pulchra CBS 124082 37 68 Lentithecium arundinaceum CBS 619 86 AB237829 6961 Neottiosporina paspali CBS 331 37 Morosphaeria ramunculicola BCC 18405 AB237835 Kirschsteiniothelia elaterascus HKUCC 82 Karstenula rhodostoma CBS 690 94 0.81/0.87/- 81 Paraconiothyrium minitans CBS 122788 AB237815 63 Letendraea helminthicola CBS 884 85 AB237814 83 Letendraea padouk CBS 485 70 70 Montagnula opulenta CBS 168 34 15 Bimuria novaezelandiae CBS 107 79 FJ495190 Phaeodothis winteri CBS 182 58 71 Didymocrea sadasivanii CBS 438 65 EU833992 78 Kalmusia scabrispora NBRC 106237 56 Phaeosphaeria brevispora MAFF 239276 FJ495192 68 Phaeosphaeria brevispora NBRC 106240 Massarina igniaria CBS 845 96 AB512358 794 Byssothecium circinans CBS 675 92 0.82/0.75/75 Corynespora olivacea CBS 114450 Pleospora herbarum CBS 191 86 GQ253099 75 Pyrenophora phaeocomes DAOM 222769 AB512352 40 Cochliobolus sativus DAOM 226212 25 Cochliobolus heterostrophus CBS 134 39 1/0.93/98 48 Alternaria alternata CBS 916 96 86 FJ495191 Setosphaeria monoceras AY016368 Pyrenophora triticirepentis OSC 100066 AB512357 Massarina eburnea CBS 473 64 Pyrenochaeta nobilis CBS 407 76 AB512355 6736 CCFEE 5390 Leptosphaeria doliolum CBS 505 75 JN673046 Phoma complanata CBS 268 92 16 Phaeosphaeria ammophilae CBS 114595 FJ005153 89 Phaeosphaeria luctuosa CBS 308 79 Phaeosphaeria caricis CBS 120249 2857 Ophiosphaerella herpotricha CBS 620 86 FJ005155 70 Phoma heteromorphospora CBS 115 96 FJ005157 65 Leptosphaeria biglobosa CBS 303 51 0.82/0.79/67 49 Neophaeosphaeria filamentosa CBS 102202 28 Phoma exigua CBS 431 74 FJ005156 CCFEE 5494 Ascochyta pisi CBS 126 54 CBS586.82 Phoma glomerata CBS 528 66 S. fusiforme Didymella bryoniae CBS 133 96 0.92/0.94/100 FJ005154 Saccothecium sepincola CBS 278 32 Leptosphaerulina australis CBS 317 83 EF105421 Phoma herbarum CBS 276 37 8564 Pleospora ambigua CBS 113979 EF105422 Clathrospora elynae CBS 196 54 Phoma zeaemaydis CBS 588 69 Camarosporium quaternatum CBS 483 95 EF105423 Phoma betae CBS 109410 Leptosphaeria dryadis CBS 643 86 DQ431914 3180 Monascostroma innumerosum CBS 345 50 1/0.98/99 Leptosphaerulina argentinensis CBS 56 AY756353 Loratospora aestuarii JK 5535B Phoma radicina CBS 111 79 AY300841 36 Phaeosphaeria avenaria DAOM 226215 Phaeosphaeria eustoma CBS 573 86 DQ431915 57 Phaeosphaeria nodorum Broad Phaeosphaeriopsis musae CBS 120026 Chaetosphaeronema hispidulum CBS 216 75 CBS380.87 74 S. tropicum 88 Entodesmium rude CBS 650 86 Otthia spiraeae CBS 114124 AY902790 90 Neofusicoccum ribis CBS 115475 91 Otthia spiraeae CBS 113091 AY902789 Botryosphaeria dothidea CBS 115476 Macrophomina phaseolina CBS 227 33 89 AY902800 20 Spencermartinsia viticola CBS 117009 84 Macrovalsaria megalospora 178149 EU107318 Macrovalsaria megalospora 178150 34 Diplodia mutila CBS 431 82 65 Botryosphaeria tsugae CBS 418 64 HQ110696 91 0.86/0.87/100 49 Phaeobotryosphaeria visci DQ377868 EU107319 81 Saccharata proteae CBS 115206 Guignardia citricarpa CBS 102374 60 Guignardia bidwellii CBS 237 48 AY902802 83 59 89 Guignardia gaultheriae CBS 447 70 Bagnisiella examinans CBS 551 66 EU107316 86 Cryomyces CCFEE 5476 Cryomyces CCFEE 5554 HQ110697 cryomyces minteri CCFEE 5187 99 Cryomyces antarcticus CCFEE 453 EF026115 54 Cryomyces antarcticus CCFEE 536 1/0.76/57 9099 Aglaospora profusa CBS 123109 HQ110700 Massaria anomia CBS 591 78 Kirschsteiniothelia maritima CBS 221 60 HQ110699 90 Lophium elegans EB 0366 95 Lophium mytilinum CBS 114111 HQ110698 94 Mytilinidion californicum EB 0385 24 Mytilinidion mytilinellum CBS 303 34 36 59 Mytilinidion resinicola CBS 304 34 HQ110701 71 Mytilinidion rhenanum EB 0341 14 Mytilinidion scolecosporum CBS 305 34 HQ110704 96 47 Mytilinidion acicola EB 0349 Mytilinidion tortile EB 0377 HQ110703 18 Mytilinidion andinense CBS 123562 83 97 Mytilinidion thujarum EB 0268 HQ110702 61 Jahnula aquatica R68 1 62 93 Jahnula seychellensis SS2113 1 DQ247805 Aliquandostipite khaoyaiensis CBS 118232 Jahnula bipileata F49 1 NG027655 85 CCFEE 5317 1/0.99/100 91 CCFEE 5533 83 CCFEE 5326 DQ220398 CCFEE 5586 91 CCFEE 5397 DQ220397 94 49 20 CCFEE 5550 CCFEE 5532 0.67/-/100 AJ276064 83 CCFEE 5572 CCFEE 5331 AY548292 96 Hysterium barrianum ANM 1442 28 39 Hysterium barrianum ANM 1495 DQ470973 16 Hysterium angustatum CBS 123334 Psiloglonium araucanum CBS 112412 NG027620 3 Hysterobrevium smilacis CBS 114601 1/1/100 88 Psiloglonium clavisporum CBS 123338 85 AY640972 96 Psiloglonium clavisporum GKM L172A Oedohysterium sinense CBS 123345 Oedohysterium insidens ANM 1443 AF353612 96 91 Oedohysterium insidens CBS 238 34 1/1/100 66 Psiloglonium simulans CBS 206 34 NG027574 Hysteropatella clavispora CBS 247 34 88 Patellaria cf atrata BCC 28877 FJ171872 55 Helicomyces roseus CBS 283 51 1/0.73/100 86 54 Tubeufia paludosa CBS 120503 U42695 Tubeufia paludosa CBS 245 49 25 Tubeufia cerea CBS 254 75 NG027573 97 Glonium circumserpens CBS 123343 Glonium stellatum CBS 207 34 Z19578 34 Glonium circumserpens CBS 123342 76 Rhytidhysteron rufulum CBS 306 38 93 78 Rhytidhysteron rufulum GKM 361A AF047831 Gloniopsis arciformis GKM L166A 34 Gloniopsis subrugosa CBS 123346 HQ262394 Gloniopsis praelonga CBS 112415 88 73 Gloniopsis praelonga CBS 123337 1/1/100 HQ262392 Acrogenospora sphaerocephala CBS 164 76 TRN529 HQ262396 CCFEE 5284 99 CCFEE 5176 EU545185 43 CCFEE 5318 CCFEE 5504 AF313370 66 CCFEE 5327 Basidiomycetes CCFEE 5303 0.09 0.09 substitution/site 0.02 EF081388 EF081387 EF081386 GU137910 EF081431 EF081448 EF081435 EF081438 HQ454509 HQ454514 EU704083 0/1600 AY350578 No No medical/No of total of 1000 ( Bayesian,LSU 1/0.96/86 AY571381 HQ454510 EF081382 FJ469668 HQ454519 JF499854 JF499856 EU019261 JF499859 EU343546 HQ693526 JF770463 JF770462 JF499858 bootstrap ( DQ204745 DQ204753 DQ204746 DQ204755 1/0.86/97 DQ204748 DQ204750 DQ204768 8/7300 1/0.92/90 DQ204756 GU214400 DQ678050 dH13831 dH13868 1/0.94/99 dH13840 dH13836 dH13862 dH13880 DQ678060 DQ767658 1/0.97/94 DQ678094 DQ678095 11/350 AY016365 dH13845 GU048607 HQ026799 HQ026798

624 GU238160 CBS181.58S. arenarium AY849958

Kittipan AY849959 0/100 CBS142.60S. salinum AY849955 AY787937 1/0.93/97 AY849954 AY849946 AY787931 AY849944 AY849945 bp 1/0.74/93 AY849956 CBS734.96S. salinum AY849957 EU673244 43/4800 EU673245 EU673243 ) of of ) AY004336 EU673229 0.96/-/- EU673230 JF441219 JF441205 1/0.79/100 GU301815 GU301867 Samerpitak FJ469672 EF175664 0/500 1/0.98/98 EF175658 EF175662 324 EF175666 AY541493 GU301855 CBS850.73 CBS100438 CBS130.65 0/15 CBS228.66 CBS383.81 CBS510.71 CBS225.77 . . CBS203.27 CBS175.65 0/25 CBS131815 CBS284.64

CBS131816 CBS435.76 CBS211.53 CBS202.27 0/30 CBS239.78 CBS102491 CBS116655 CBS100442 0.80/-/- CBS729.95 CBS100486 CBS412.51

). CBS172.74 CBS475.80 CBS780.83

1/0.98/99 CBS423.64 CBS536.69 4/25 CBS118.91 CBS437.64 CBS867.95 1/0.88/51 CBS547.81 CBS864.95 CBS125818 CBS125817 1/0.97/84 CBS119775 HQ634611 HQ634610 DQ885904 DQ885905 DQ885906 Sympoventuriaceae 1/0.97/99 EU035424 EU035423 CBS469.95S. excentricum CBS588.66 Veronaeopsis simplex EU035435 EU035464 EU035456 1/0.94/94 DQ470974 EU035447 GU323212 GU456336 EF114712 EU035433 EU035473

Phylogeny of GU348998 HQ599579 0/130 EU035468 EF114694 JQ036232 EU035462 Venturiaceae 1/0.71/97 GU301814 EU035443 EF114692 GU301878 EU035454 JQ036233 EF114695 JF263534 0.57/-/- HQ613813 HQ599589 CBS210.95S. fusarioideum AF050275 EU041874 EU041872 EU041873 EU041875 EU107312 AY004338 AFTOL668 1/0.78/94 CBS769.83S. cateniphorum FJ358231 FJ358225 AF050263 DQ329021 EF643750 AY300828 AY584644 44/250 1/0.70/93 DQ329014 1/0.88/65 DQ782913 EF105141 EF105138 EF105132 DQ329015 EF411063 1/0.94/95 AF279407 AY607737 1/0.85/59 AY640962 DQ361098 DQ361104 Ascomycota DQ361102 U93304 DQ239894 DQ239886 1/0.92/82 0.95/0.92/86 AB075328 AB075330 AB075326 AB075327 AB075329 AY004346 1/-/- GQ911144 U29571 76/1000 U29570 U29572 1/0.98/100 GU808560 AY213700 AY176751 FJ864710 AY853379 AY796003 AY796004 1/0.97/75 AY796001 AY796002 50/90 EF413632 AF356689 DQ832334 1/1/100 DQ832330 AF356693 DQ119558 DQ119564 DQ119562 DQ119571 DQ119570 DQ119569 CBS549.74 Neta DQ075680 JN036557 salmonicolor AY489721 0.99/0.94/59 AY489723

U68127 U68128 AY882374 0.99/-/57 AY882373 33/2700 AY882372 AY882371 AY882377 1/0.98/99 AY882375 EF151316 EU862178 HM449060 HM449058 GU563359 1/0.83/51 HM449063 1/0.97/96 GU563363 EF100924 19/400 1/1/100 EF100957 AJ580945 AJ579686 DQ268616 DQ268615 DQ268617 DQ294353 29/850 AB363791 1/0.80/84 DQ821538 DQ821537 HQ413647 DQ368627 AB237812 AB237811 GU198953 AB237826 AB197135 AB237820 AB237819 6/300 1/0.98/100 AB237816 AB237813 AB237829 0.81/0.87/- AB237835 AB237815 AB237814 FJ495190 EU833992 FJ495192 0.82/0.75/75 AB512358 GQ253099 1/0.93/98 AB512352 0/350 FJ495191 AB512357 AB512355 JN673046 FJ005153 FJ005155 FJ005157 0.82/0.79/67 FJ005156 CBS586.82 S. fusiforme 0.92/0.94/100 FJ005154 0/30 EF105421 EF105422 EF105423 1/0.98/99 DQ431914 AY756353 0/2000 AY300841 DQ431915 CBS380.87S. tropicum AY902790 AY902789 AY902800 EU107318 0.86/0.87/100 HQ110696 EU107319 AY902802 0/15000 EU107316 HQ110697 1/0.76/57 EF026115 HQ110700 HQ110699 HQ110698 HQ110701 HQ110704 0/300 HQ110703 HQ110702 DQ247805 1/0.99/100 NG027655 DQ220398 DQ220397 0.67/-/100 AJ276064 1/1700 AY548292 DQ470973 1/1/100 NG027620 AY640972 1/1/100 AF353612 NG027574 1/0.73/100 FJ171872 5/5 U42695 NG027573 Z19578 AF047831 HQ262394 1/1/100 HQ262392 HQ262396 50/1000 EU545185 AF313370 Basidiomycetes 0.09 0.09 substitution/site 1. Changing genus names

CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi Madurella mycetomatis Madurella grisea

Sarah Abdalla Ahmed

CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi Sole clinical picture: mycetoma = fibrosis, grains, fistels, osteomyelitis

CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi E urotiales

Onygenales

Chaetothyrial es

Sordariales Ophiostomatales Madurella mycetomatis Diaporthales X ylarial es

Hypocreales

Microascales

Lecanorales Ascomycota Ascomycota Dothideales Mycocal ici um albonigrum

Pleosporales Madurella grisea “Dematiaceae” Dikarya Leotiales Geomyces pannorum Leotiales

P ezizales

Calicium tricolor

Hemiascomycetes

Pneumocystis carini i Schizosaccharomyces pombe

Uredinales

Sporidiales

Til letial es Ustilagi nales

Basidiomycota Tremel lales Basidiomycota

S tereales

A garicales

Glom ales Linderina pennispora Glomerulomycota Cantharellus tubaeformis Harpellales Zoopagomycotina Entomophthorales Entomophthoromycotina Zygomycota Mucorales Mucoromycotina Mortierellales B asidi obolus ranarum Chytridiomycota Chytridiales Chytridium confervae Chytridiomycota Madurella mycetomatis: endemic in arid climate zone northeastern Africa

Mycetoma Research Center, Khartoum

CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi Order Sordariales

Nearest neighbor: Chaetomium Natural habitat coprophilic Often no anamorph

CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi Order Pleosporales

Madurella grisea = Trematosphaeria grisea Natural habitat waterborne

CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi Trematosphaeria grisea

Delayed sporulation with coelomycetous anamorph

CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi meningitis Distribution of cerebral infections primary E urotiales (aspergillus) rhinoorbital rhinocerebral near-drown Onygenales

Chaetothyrial es (black yeasts) Sordariales Ophiostomatales Diaporthales X ylarial es

Hypocreales Microascales (scedosporium) Lecanorales Ascomycota Dothideales Mycocal ici um albonigrum

Pleosporales

“Dematiaceae”

Leotiales Geomyces pannorum Leotiales

P ezizales

Calicium tricolor

Hemiascomycetes

Pneumocystis carini i Schizosaccharomyces pombe

Uredinales

Sporidiales

Til letial es Ustilagi nales

Tremel lales Basidiomycota (cryptococcus) S tereales

A garicales

Glom ales Linderina pennispora Cantharellus tubaeformis Harpellales Entomophthorales Zygomycota Mucorales (mucorales) Mortierellales B asidi obolus ranarum Chytridiales Chytridium confervae Chytridiomycota Orders with intrinsic antifungal resistance

E urotiales

Onygenales

Chaetothyrial es

Sordariales Ophiostomatales Diaporthales X ylarial es

Hypocreales

Microascales

Lecanorales Ascomycota Dothideales Mycocal ici um albonigrum

Pleosporales

“Dematiaceae”

Leotiales Geomyces pannorum Leotiales

P ezizales

Calicium tricolor

Hemiascomycetes

Pneumocystis carini i Schizosaccharomyces pombe

Uredinales

Sporidiales

Til letial es Ustilagi nales

Tremel lales Basidiomycota

S tereales

A garicales

Glom ales Linderina pennispora Cantharellus tubaeformis Harpellales Entomophthorales Zygomycota Mucorales Mortierellales B asidi obolus ranarum Chytridiales Chytridium confervae Chytridiomycota Conclusion: phylogeny places fungi in logical arrangement. Natural behaviour significantly different between groups. - Virulence potential and safety issues. - Infection route and course. - Therapy.

Temporary problems: - Sampling effects. - Taxonomic level undetermined.

CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi 1. Sampling effect: New species added, old species sequenced; ‘final’ tree does not exist

Pyrenochaeta mackinonii 1976 = Nigrograna mackinonii 2012 = Biatriospora mackinonii or B. marina 2014

CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi 2. Clade definitions: No operational criterium for clade delimitation, generic concepts may vary

CBS 297.58 CBS 385.87 87 CBS 124169 ATCC 26490 Petriella sordida 98 CBS 184.73 T sordida CBS 520.72 Petriella clade CBS 362.61 ST guttulata 72 Petriella guttulata CBS 265.64 CBS 347.64 89 99 CBS 391.75 Petriella setifera CBS 395.69 CBS 710.96 CBS 311.72 T africanum Petriellopsis africana Petriellopsis CBS 100391 88 clade CBS 116906 Scedosporium prolificans S. prolificans clade CBS 452.89 T inflatum CBS 102083 ET putredinis 96 91 CBS 118694 CBS 100392 CBS 108.10 T lesnei Parascedosporium Parascedosporium CBS 127.84 T tectonae putredinis clade CBS 102085 CBS 116910 T aurantiacum 89 Scedosporium CBS 489.72 T desertorum aurantiacumScedosporium CBS 116911 T minutispora desertorumPseudallescheria minutispora CBS 117406 T dehoogi Scedosporium dehoogii Pseudallescheria CBS 119707 T boydii Pseudallescheria boydii clade CBS 254.72 T angusta Pseudallescheria angusta CBS 117407 T apiosperma Pseudallescheria CBS 987.73 apiosperma CBS 129.78 T fimeti Lophotrichus fimeti Lophotrichus CBS 320.72 T basitruncatum Graphium basitruncatum clade 72 CBS 102632 ET penicillioides 100 98 CBS 102630 Graphium penicillioides JCM 10499 CBS 422.94 98 CMW 5605 T fimbriasporum Graphium fimbriisporum CBS 421.94 84 AsaP.18 Graphium clade 92 CMW 5611 CMW 503 T pseudormiticum Graphium CMW 12285 pseudormiticum CMW 5598 CBS 116195 CMW 5603 Graphium laricis CMW 5601 T laricis CBS 665.71Microascus trigonosporus

0.02 CBS CBS - KNAW - KNAW Fungal Fungal Biodiversity Biodiversity Centre Centre - Clinical- Clinical and and Extremophilic Extremophilic Fungi Fungi Alternaria = Allewia 1990 = Chalastospora 2007 = Embellisia 1971 = Lewia 1986 = Nimbya 1989 = Pseudoalternaria 2013 = Sinomyces 2009 = Teretispora 2007 = Ulocladium 1851 = Undifilum 2009 = Ybotromyces 1986

Woudenberg et al. – Stud. Mycol. 2013

CBS CBS - KNAW - KNAW Fungal Fungal Biodiversity Biodiversity Centre Centre - Clinical- Clinical and and Extremophilic Extremophilic Fungi Fungi Proposed solution for temporary problems at the genus level: Follow changes with some delay, until established.

CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi CBS - KNAW Fungal Biodiversity Centre - Clinical and Extremophilic Fungi