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et et B. Ascomycota > Wallemiomycetes G. hirsutum et al.!"E6 Basidioascus, B. persicus et al.56!7@ Key words: Agaricomycotina Basidiomycota Geminibasidiomycetes Wallemiomycetes Geminibasidiales !5 Basidioascus, Basidioascus Saccharomycetes IMA FUNGUS · 6(1): 215–231 (2015) 215–231 · 6(1): FUNGUS IMA G. donsium Geminibasidiales Basidioascus Geminibasidiales B. undulatus et al. 56!_V [ % 2 /@ 7 Wallemiomycetes Geminibasidium @ <>/? magus/ . undulatus [ al. 56!_ V N > 9% < > $ . V [ U 566_ . U % al. 56!_ V > Basidiomycota Agaricomycotina B. V [ Ainsworth Ainsworth !5 Geminibasidiales [ % [$ N . B. undulatus Agaricomycotina : genome, origins, and sexuality origins, : genome, $ % et al.566E $ %W , which he B. undulatus, which he undulatus % 9 3 et al.56!_ Geotrichum B. undulatus Geminibasidiales Geminibasidiomycetes Basidioascus

< G Geminibasidiomycetes !5 V Basidioascus undulatus Wallemiomycetes Basidioascus undulatus et al. 56!_

[ Basidioascus O . B. undulatus . $ < % G Wallemia U G S G Q G / / .WG "F6G /%Q Q 2!/ @* @ / O @ Z % V/E45E#.!F6!Z@/ + % US G Q G / / .WG "F6G /%Q Q < + W @ Z % Q _6U G Q Q 2!>F>4G K Article info:@ [!FU 56!4K/ [!_`56!4K; [!"`56!4 ? V $ % [ Wallemiomycetes %$ N [ $ %% Z _4 . $ [ V $ _5U O % 57 F!5_ ; % % 6GFG Abstract: 2!/6GFG . [ !"E7^ © 2015 International Mycological Association © 2015 International Mycological conditions: distribute and transmit the work, under the following are free to share - to copy, You Attribution: 3 7 5 9 <V> ! Basidioascus undulatus Basidioascus Non-commercial:No derivative works:Any of the above conditions can be waived if you get For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Nothing in this license impairs or restricts the author’s moral rights. permission from the copyright holder. > & Bisby’s Dictionary of the Fung 2 & Bisby’s B. undulatus in Saccharomycetes $ Galactomyces Dipodascus $ G V > ; / U 566_ Z U 566_ INTRODUCTION % O V P O % P U 566_ / O U+ [!644"E\ 56!46F6!!7 VOLUME 6 · NO. 1 Nguyen et al.

Wallemiales % "_ ? De novo Wallemiomycetes > et al. 56!_ Wallemiales @;/%_6+ % et al.56!5 Wallemia with three + 9 > et al. 56!_ [W. sebi, W. muriae W. ichthyophaga et [ .. al.5664V Wallemiomycetes 5! _! 7! 4! F! #! E! "! W

ARTICLE [ U et @@;/GS%56+ O et al. al. 566F 56!! [.$!.74.!6. 6. 6#.4. 6#. !6.O5666.V!F.! 5666 % 9% / Z/@V %5_] % et al.56!_V + 5 % Agaricomycotina %5!6 * I @ O 56!5/ ; et al.56!5 et al.56!_ + 5 @/U % +/U N [ % @/U %6!!"* et al.566" $ B. undulatus % %6E!] . % / et al.56!5V [ > et al. 56!_ W GS]U/ % 54 ; et al. 566# $ % % GS] W ? B. undulatus ] %% 9 et al. 56!! ?. [ Q % % U % 764 [\\ . % Geminibasidiales in Wallemiomycetes [ . . _4 [\\ \ V . B. undulatus U/2S 5%5!69 I % 56!! N U/2S 5 B. undulatus Wallemia . The ]U .S@%5_+ %I* O % % 56!!ab initio >G+N % [ @? ? . $ %556@ I+ 5664; Agaricomycotina % < et al. 566" $ % $ B. undulatus, G. donsium, W. sebi $ ? . ?G<@; % [ Geminibasidiales $ V ? . N ; @ 4 GF ` et al. 56!7 [ MATERIALS AND METHODS ... . . . Z ; \@ .; Growth, DNA extraction and sequencing %555EV U* % V$. Basidioascus undulatus/ $ \\ >/R Q ; ].+ @ * UQ >/ % !5 * et al. 566# < R <>/ ? ? [ % >G+N ]/ > % [ * V + G `V*@66666666% `V*@6!666666K+ ; / .? !6! . >; `>/57#""5 `]NUG ] % et N 9 @?5466 V @?X_ al.56!7 > G G @ @ #" +*/@V [ Basidioascus undulatus Genome assembly and annotation V ? V ? % . % W G%6!6! [\\ \ +*/@V \ ?\ Z $ W/@V. %555EG et al.566" V % 66!_K [\\ \ $ \ ] % %% 4R in Saccharomyces cerevisiae Cryptococcus neoformans

216 IMA FUNGUS Basdioascus genome, origins, and sexuality

9 et al.56!! ? .% % ;9*N; @ X ARTICLE ¢!6S.64 B. undulatus +*/@V %74_]et al.56!6V % 555E Saccharomyces cerevisiae C. O % neoformans ;+ _ %!7 * et al. 56!_ * . [ et al. 566" G/V.]V * I ; 9 et al.56!! 5667 / % U % O{!66 $ ¢6! [ N ; @ % 555E [.$!66646 ! 5 Saccharomyces cerevisiae _ [.$!666 ! 5 _ ? .% ¢!6S.64 / Q $R Phylogenomics and molecular dating . O% ;+ _ %_5 ; ? Agaricus bisporus . % bisporus 9"# % 56 U et al 56!5 Alternaria % + % [ . brassicicola Q et al 56!5 Arthrobotrys oligospora .V . $ . /VGG 57"5# et al 56!! Aureobasidium pullulans $ . . ! Rhizophagus irregularisGlomeromycota % pullulans SW.!46 ] et al 56!7 Auricularia [ < % subglabra % 56 W et al 56!5 Botrytis cinerea% 9 et al. !""# !6 @ % 2 56!5 Coccidioides immitis @ @ et al. 5667V I + 566# + IV @ et al 566" Coprinopsis cinerea @ et al 56!6 ; I 56!_K $ 56!6 Cryptococcus neoformans % neoformans `SG5! % [ ? G * et al 5664 Dacryopinax <`U #_! @@;! % % [.$!66646 !6 W et al 56!5 Fomitiporia mediterranea % !6 ! ;+ _ W et al 56!5 Fomitopsis pinicola W;.4E45# @@! %_6 W et al 56!5 Malassezia globosa et al Confocal laser scanning microscopy 566# Mixia osmundae N/U !7_57 % !6 V et al V % 56!7 Monacrosporium haptotylumG+@56646U Basidioascus undulatus/ [@ V et al56!_Rhizophagus irregularisTremella 756M#66 Tuber melanosporum @VQ"N S> U et al56!6Ustilago maydis2 et al566F 566"N @ G U N Wallemia ichthyophaga SW.""7 et al 56!_ W. sebi % !6 ; et al 56!5 Transmission electron microscopy `]N UG ] % et al. 56!7 / % Basidioascus undulatus Wallemia sebiG+@F__FF WZ> U et al.566E . 9 Q ? O {"6zU et al.566E % 5_:GV $ % [ O $ 4 & 4 ? V ? _6 76 % %555E ? .% ¢!6S. % . 64 [$ ? M!EF `]N UG % ; ? :G ? 9 !"EF I W V.G%!666 !F!_> et al. !"EEU< I 5666/ O 5666 [ ? . O. !z \% !z \% %E4 G [ #5/ %E4 G

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VOLUME 6 · NO. 1 217 Nguyen et al. ARTICLE

Fig. 1. Basidioascus undulatus

!66& $ 4EzV[ 5""5 . % "# 2 V % 9 IQR<!"E# . % 5E % @ * Z % 57V>46 * U + N* !42 / GS]U/E!z EEz . !6 5z GS] % 46z 4 @ R / !_"_4 ab initio 9 et al.!"EF@ $ ]U .S@ F!5_ `SQ*!566S`SQ*V % >G+N ? @N/ *_G GG< @N/ @? F!5_ < ]/ U % N `@ et al.56!5 Q_FE! F6z RESULTS 577576z Genome sequencing, assembly and annotation @ . N ? $ !5 ? $ ! V %$ [ $ % 5 >et al.56!_ !6! / "_ !!] R de novo O W !V % _5 U V ]G Basidioascus undulatus [

218 IMA FUNGUS Basdioascus genome, origins, and sexuality

N NN . ARTICLE < I ] !"#5 % W !/ W !+ NW !G N

W !<W NNW !S . W !WV W !] W !9 % V W !N % W !`

Mating genes ]. 9U] <>/ . % U/;2U/;22U/;222 ]V; ] ]! ]" ` et al. 56!_ Saccharomyces cerevisiae U V 5 B. undulatus/ $ KSS! FUS_ . 5E_# Fig. 2. V V A–B. Basidioascus undulatus

Septal pore morphology B. undulatus G. Wallemia sebi, Basidioascus donsium æ6!5 ) W. sebi æ66F ) undulatus, Gemmibasidium donsium / W. sebi [ Wallemiomycetes [$ [[ % W 5. Basidioascus B. undulatus G. donsium undulatus, G. donsium W. sebi % W. sebi, B. Agaricomycotina V undulatus G. donsium B. undulatus G. donsium W. sebiV $ W. sebi were Phylogenomic analysis and molecular dating % W 5SMW + Basidioascus [ Wallemio- in B. undulatus W 5/M+ G. donsium W 5GM< mycetes [ S . % W. sebi than in B. $ undulatus G. donsium/ . \ et al.5664U $ % et al. 566F V . ? _4 . W. sebi % [ V !"#7 ; et al. 56!5 W. sebi W _/ Q /WVQ* @ + < <>/ > et al. 56!_ G et al.566#V B. undulatus Basidioascus Wallemia, with G. donsium !66V [ W. sebi W ; et al.56!5 et al.56!_

VOLUME 6 · NO. 1 219 Nguyen et al. G\ G\ undulatus 550A163039 1512E55FF2 102794F55C 1586D154C9 S/ >/ >/ >/ >/ >/ >/ >/ >/ >/ >/ >/ <4!56/+6S_ !#! 66 _7#F 7EFF <4!56/+6S_ /4G!#+#<__ #"_ 66 76 475" /4G!#+#<__ Basidioascus undulatus , Cryptococcus neoformans vs. Saccharomyces cerevisiae Basidioascus undulatus 47_ 6666 5#7E 7_4! >/ Saccharomyces cerevisiae Basidioascus undulatus Locus ID Score E-value % identity % positive Locus ID Score E-value % identity % positive Probable Locus ID > > > > S/ <4!56/+6S_ b / / > / / > > / / / > / > / > / / > / > / / > > ;4#!!4E! W6++5F6/*_!5 ;##F!7"! G6F*5!6 *546 ;##!"5"! !G//<45WW6 47_ 66 _6EE 45!F !G//<45WW6 EF" 6 _E"! F!!F U S!! <947 + 6#_ SG!!7 U !__ SG!65 SG!67 * _5" 9 !4# Gene name @5\>+@! < _F"G @/S5\GQU! ]*!#4G 9WU!\US _ ]*54!G >/UE\U S5 9 6EF ;4FE5!4! 6!5GGF!5/! EEF 66 5#57 7F57 G_5E7E57<+ 5!! 66 #44# E_5! SG!6#\USN5 ` 65!G Regulation of checkpoint DNA damage DNA crossover frequency Strand invasion Spo11 Removal of Table 1. * % Table Process DSB generation

220 IMA FUNGUS Basdioascus genome, origins, and sexuality ARTICLE G\ undulatus 5756414F12 E449052477 122F99619E 4AE1080675 A74C037F01 DB366D3081 7F6ED5B406 D659917BB2 29F5BE17B9 EG6_/5"6GS 8C03A290CE 36DFC1EC56 13E0CFCD07 2C075CC8D6 5G6#4GGE > vs. Cryptococcus neoformans Basidioascus 4!E 66 4#"4 #6"! >/ >/ >/ >/ >/ >/ > > > vs. Saccharomyces cerevisiae Basidioascus undulatus Basidioascus undulatus Locus ID Score E-value % identity % positive Locus ID Score E-value % identity % positive Probable Locus ID > > > EG6_/5"6GS EF# 66 _!"_ 4_F! >/ 5G6#4GGE/ b / / > / / > / / > / > / > / > 4#554##4 EG6_/5"6GS !#F 66 7EF4 F#6_ EG6_/5"6GS 5_F 66 F64! #4_E ;###_F6! S44S#/+6+< E7 66 _67" 7"#E S44S#/+6+< !6! 66 _5_" 46# ;##5446! 4#4F7!7W!5 F5 66 56"7 7_4" 4#4F7!7W!5 !!4 66 _#!_ 4!64 ;4FF#_E! 7!"44W+<+< 57E 66 7767 F#4! 7!"44W+<+< _44 66 47#4 #6E5 neoformans ;##4E57! !_S6GWG<6# _76 66 5#4E 7#_5 !_S6GWG<6# 7#F 66 5""E 46F4 Cryptococcus a Locus ID < 554 ;4F#"F5! 5G6#4GGE<`! Q*!67G VQ;5VQ;_ >*6EE * 5_7 ;4FF#66! 7/S!6E6F#4 ;##_6_4! +GW!#/7S5/ !!"# 5FF 6 66 45"# 7_F# #!7# F647 7/S!6E6F#4 +GW!#/7S5/ !7"! 6 FF7E #"57 VQ;! Q*66FG ;4#5"54! <+_FF<_6E! 7_" 66 7"#E FFE" <+_FF<_6E! 4!E 66 4F#! #64F U*95 * 6_4G SQ! Q 6__G ;###6_7! _F*6E5 9Q;5 ]*6__ 465447__ > 9Q;! N*6#5 4654446#

VOLUME 6 · NO. 1 221 Nguyen et al. G\ undulatus 1FC448F440 45_F"/!G75 68E0241B20 130D4D9937 GEE56/EF7+ C71F05C621 AE68F812D8 6453C9DC91 AD4E738F04 82DC381A41 062A151CC3 10D8B5FBE4 ARTICLE / / / / / > vs. Cryptococcus neoformans Basidioascus >/ >/ >/ >/ > !_6<7<""_# F4! 66 _E67 4"#E !_6<7<""_# vs. Saccharomyces cerevisiae Basidioascus undulatus Basidioascus undulatus Locus ID Score E-value % identity % positive Locus ID Score E-value % identity % positive Probable Locus ID > > 45_F"/!G75 457 6666 _4" 7E#5 GEE56/EF7+ 5#_ 66 5E"4 77F4 >/ b / / / > / > / > / / > 4E5FF766 !_6<7<""_# FEF 66 _464 4##_ !_6<7<""_# !75 66 _5F! 7FF4 !_76"6476 > ;4#!!FE! G#!W64GF5! E"4 6 7!7! F6_! G#!W64GF5! !_F# 6 4FF #_F! ;4#6_7E! F74_G"

a

Locus ID cerevisiae Saccharomyces *NW! ]*6"6 @*E S !!FG @*4 <*6!_ <>*7 Q 664G ;4#5F65! "56F"4E!E" 55_ 66 5#4 7__E SE<5"++4#6 "65 6 #!4 E_7" @ @5 `*6"5 ;<@4 675F7! SGE ; 66#G @GG_ ;7647!! U;@_ `*6!" U@GF Q _47G U@G#U@G_ 9 6_"G * 5!" ;##_7E!! /SFEWE!5G+N; [\\ %\ \ end joining Other Nonhomologous Table 1. G Table Process a

222 IMA FUNGUS Basdioascus genome, origins, and sexuality

Table 2. * ARTICLE Process Gene name Saccharomyces Basidioascus undulatus vs. Saccharomyces cerevisiae cerevisiaea Locus ID Locus ID Score E-value % identity % positive Mating response @VS_ 2*!#EG F6!6#6/""5 E7# 66 54!# 77_# 2@@! ] 676 6_<<_4/5_7 _7! 66 4E4 #574 WZ@_ +*6!F 6_<<_4/5_7 _44 66 4#7E #7E_ @VS# <*!4" G4FS/!!W"_ 56" 66 7"_5 FF6F @VS!! * _F5 /#5F/E#S4E _!" 66 45F F#EF ];/! 9 664G 66 @VS56 9*66#G /+W7!< W/ ! `*!4#G > @VS4 < !6_ > Pheromone 2S! ]*56_G G!S+E_57*5_E ! @VS57 ` !!# > @VS!7 < 7!6G GG"6W7/"7F ""E 66 7474 4EFE /*! ; !55 "E<555!+4E !5" 66 _6_E 4!" @VSF 2*56"G S

the Wallemiomycetes % sebi76z W. ichthyophaga74zV _+ Agaricomycotina % Basidioascus EEz 9 et al. !""# @ et al. 5667 GS] + I V 56!6 ; % I 56!_ W _+ Z B. undulatus Wallemia in B. undulatus. 546/5"U W. sebi W. ichthyophaga U % ]U . __/FU S@ % V . 9% et al.566E]U .S@ DISCUSSION ? % Genome sequencing and annotation + >/ Basidioascus undulatus [ Geminibasidiales ? ]U .S@ ?V!!] ? ? >/ >% % % 57 [ ? 5""5 W >/ G<@ % ? @ et al.56!7Wallemia sebi ? ]VM/] W. ichthyophaga % V ? . B. undulatus < % ]GM/] !6M!5 z O B. undulatus _5 U et al.566F _z W. sebi"EU ; et al.56!5 U I9 566EV W. ichthyophaga"FU et al.56!_/ ]G >/ >G+N @? ? B. undulatus 4Ez W. !_ U 56!7

VOLUME 6 · NO. 1 223 Nguyen et al. ARTICLE Saccharomycotina Pezizomycotina Agaricomycotina Saccharomyces cerevisiae Orbiliomycetes Dothideomycetes Sordariomycetes Eurotiomycetes Tremellomycetes Taphrinomycotina Ustilaginomycotina Leotiomycetes Pezizomycetes Wallemiomycetes Agaricomycetes Pucciniomycotina Geminibasidiomycetes ureobasidium pullulans Neurospora crassa lternaria brassicicola Trichoderma atroviride Trichoderma A rthrobotrys oligospora A Taphrina deformans Taphrina Dacrymycetes Malassezia globosa Penicillium chrysogenum A Monacrosporium haptotylum Coccidioides immitis Tremella mesenterica Tremella Cryptococcus neoformans Tuber melanosporum Tuber Botrytis cinerea Sclerotinia sclerotiorum Tilletiaria anomala Tilletiaria Wallemia ichthyophaga Wallemia Puccinia striiformis uricularia subglabra Pyronema confluens 1.00 Wallemia sebi Wallemia garicus bisporus A 1.00 A Coprinopsis cinerea 1.00 Fomitiporia mediterranea Fomitopsis pinicola Mixia osmundae 1.00 Dacryopinax sp 1.00 Ustilago maydis 1.00 1.00 1.00 Basidioascus undulatus 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 Glomeromycota 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 0.99 1.00 1.00 Rhizophagus irregularis 0.2 A

224 IMA FUNGUS Basdioascus genome, origins, and sexuality

ARTICLE

garicomycotina A Pezizomycotina Glomeromycota Orbiliomycetes garicomycetes Tremellomycetes Saccharomycotina A Geminibasidiomycetes Eurotiomycetes Dothideomycetes Leotiomycetes Rhizophagus irregularis Glomeromycota Wallemiomycetes Sordariomycetes Pezizomycetes Ustilaginomycotina Taphrinomycotina Pucciniomycotina Dacrymycetes rthrobotrys oligospora ureobasidium pullulans lternaria brassicicola uricularia subglabra garicus bisporus Saccharomyces cerevisiae Monacrosporium haptotylum atroviride Trichoderma Neurospora crassa Sclerotinia sclerotiorum Botrytis cinerea Penicillium chrysogenum Coccidioides immitis Mixia osmundae ichthyophaga Wallemia Basidioascus undulatus mesenterica Tremella Dacryopinax sp Fomitiporia mediterranea Rhizophagus irregularis deformans Taphrina melanosporum Tuber Pyronema confluens A A A Puccinia striiformis anomala Tilletiaria Ustilago maydis Malassezia globosa sebi Wallemia Cryptococcus neoformans A Fomitopsis pinicola Coprinopsis cinerea A 0 33±6 Cenozoic Era 100 76±12 96±17 94±22 121±16 Mesozoic Era Jurassic Cretaceous 200 167±10 179±18 Triassic 212±19 233±3 U 234±28 249±3 Permian A. G ? _4 250±29 253±10 253±22 262±30 300 307±26 312±11 Carboniferous 321±20 327±13 400 Devonian 377±36 379±31 Paleozoic Era 383±18 Silurian 405±18 409±32 Ordovician 447±35 500 474±34 485±21 B. G $ . V % Cambrian 537±25 600 . V Precambrian Eon Million years ago (Mya) 700 661±26 684±25 B Fig. 3. ; !6!5" ] %/ G/V.]V ; @ $ G

VOLUME 6 · NO. 1 225 Nguyen et al.

Table 3.] Basidioascus undulatus Wallemia sebi W. Ichthyophaga Descriptive statistic B. undulatus W. sebi W. icthyophaga Genome assembly @? N 9 .@? N 9 .@?I747 N 9 .@? V !5U a —— ARTICLE "_ — — < O !!] — _#] / O _5U c "EU "FU S ; ]G 4Ezc 76z 74z > % 57 #! {5#6 > _64E !!7 "4 > 5""5 4F E5 * O "#2 c "662 #"62 >46 !42 c _762 7762 Genome annotation ; EEz —— / !_"_4e 45E7 7EF_ S% F!5_e —— S% _FE! — — @ [ W G ? Z/@VGS]U/ U/2S

? >et al.56!_V B. undulatus O$ P N U/2S 5 % @ ? % Q% R $ / S% < V% % O /S< $ /S< S et al. 566" V % $ /S< Q % ]U .S@ % W ! % % $ % % V ; . ? S et [ % O al.566"Q% !_"_4 ]U . / % S@ F!5_ [ % W F!5_ V ! HOP5 !6 +*/@V [ V HOP5 %% Z ; \@ .; % . N ; @ Z 745 . U et al.566E [ W V HOP5 % B. undulatus >/? B. undulatus, % EEz % % % GS] ! % !6 Meiosis Basidioascus undulatus >et $ al.56!_V . Basidioascus undulatus W $ W !2M* $ B. % undulatus] % B. undulatus $ % $ $ ; ? N B. undulatus Mating V U/V [ $ % *et al. % > et al. 56!6N % 56!_ $ Saccharomyces cerevisiae V % % S. cerevisiae

226 IMA FUNGUS Basdioascus genome, origins, and sexuality

B. undulatus Basidioascus ARTICLE Agaricomycotina W 9 9< _/ V \ $ ]G O B. > O undulatus Wallemia %/ 9< 9Exobasidiomycetes Ustilaginomycetes ; Malasseziomycetes5F5U / KEX! KEX5 STE!_ RAM! RAM5 STE!7 AXL!! STEF % 546 U N B. undulatus$ RCE!\STE57 Wallemia B. undulatus % V 5 % % Z % ; V N S $ %% . % U/;2 % $ % @ I+ 566EV ` I + 56!! W. sebi W. ichthyophaga __/F U/;2 % U % K 9U]<>/ !!"U et al.56!_ % U/V $ 9 et al.!""FU V STE3, KSS! FUS3, STE# STE!! GPA! $ STE7 STE56 V 5 $ ]. " STE!E STE!5 % % * et U/;2 STE4 FAR! al.566" [ 9U]<>/ +6!_!!5#75 + ? GE+##<6!/ > Wallemiales Geminibasidiales % et al. 56!_ Wallemiomycetes > et al. 56!_ V W 5 Geminibasidium % Basidioascus; % Wallemiomycetes Basidiomycota U et al. 566F et al. 5664 $ % U % Agaricomycotina; et al.56!5>et al.56!_ % et al.56!_V [ Wallemiomycetes Q incertae sedis in the Basidiomycota in the % <>/ Wallemia >G+N $ W 56!5 U+

VOLUME 6 · NO. 1 227 Nguyen et al.

et al. 56!_ S * [\\ Notes[ V $ N $ Geminibasidiales % Agaricomycotina Wallemiomycetes > et al. 56!_ N Geminibasidiomycetes Wallemiomycetes Agaricomycetes, Tremellomycetes ; et al.56!5

ARTICLE Dacrymycetes Agaricomycotina9 566#+ Geminibasidiomycetes9<V>I@ cl.

Geminibasidiomycetes Wallemiomycetes nov. Agaricomycotina U+ U+E!!FE6 $ V WallemiomycotinaU+ U+ 446_F7 Description[G $ Wallemia < 56!7 V Agaricomycotina Basidiomata Basidia Wallemiomycetes . the Geminibasidiomycetes Basidiospores Arthroconidial TAXONOMY \ . $ @ % . Agaricomycotina<Prosyllabus[$$$% 566! . U+ U+4F644_ @ Homonym[Agaricomycotina + et al., Mycol. Prog. 5[ 74566F Type order[Geminibasidiales

Description[ U Basidiomycota % ACKNOWLEDGEMENTS $ % + 4@ >/ + et al. 566F / ; / W [ U * % Agaricomycetes, Dacrymycetes, Tremellomycetes, U/2S 5G * Wallemiomycetes Geminibasidiomycetes ;+ 2 + GS]U/ G ] 2 W VSU`G % Notes[/ Agaricomycotina<566! N ] % + et al.566F `]NUG Hymenomycetes sensu @ I V !""4 <>/ V * > [ % $ G G @ AgaricomycotinaV N ? V Z@ > @ W >@W B. persicus % % $> et al.56!7 Geminibasidiomycetes V Wallemia species REFERENCES

Wallemiomycetes et al., Antonie van Leeuwenhoek / ` G G/ % 2 `/ X U + S; et al. 87[_555664 56!!] U+ U+46!7"F Sclerotinia sclerotiorum Botrytis cinereaPLoS Genetics 7 E[!6655_6 Description[ G $ + % / > @ / % < ] % // <% U et al. Agaricomycotina V 56!5 @;/[

$ .? Journal of Computational @ % Biology 19[744M7## $ + + +@ `; UQ W566F V V . [ Mycological V . Progress 5[7!MFF . + U*V `56!6< M % Fungal Biology Reviews 24[!M!F + O U 9 GX ` 9` + < ; % Type order[Wallemiales 56!! @ . @@;/GS Bioinformatics 27[4#EM4#"

228 IMA FUNGUS Basdioascus genome, origins, and sexuality

+ %U* O/56!!S 9 @ U @+ * * @ % G9 9 U et al. 56!! ARTICLE ]U .S ]U .S@ Current Protocols in G % Glomus % Bioinformatics[ M7F!6 $ Genome Biology and Evolution G G G ] /% X U > ; ` 3["46M"4E et al. 566" +*/@V[ BMC 9 V@ VN VU % . `9 Uet Bioinformatics 10[75! al.!"EF/ [ @ R Journal G <]% U2O / UGet al.56!! of Electron Microscopy 35[_67M_6F >$ ? % 9 9/2 + U!""FV Puccinia striiformistritici [ PLoS ONE 6 E[575_6 Ustilago maydisEMBO Journal 15[!F_5M!F7! G ]` ; U < +V + U* <` 9 < ] < < U !""# W 566#O566FP~/ W V * [ U G % Mycologia 98[E46ME4" Homobasidiomycetes American Journal of Botany 84[ "E!M G Q9/ `U W / 2 // @ % ` et al. ""! 56!_ ] ? Taphrina 9 <@ 566# O566FP~ / % % deformans mBio 4 _[ AgaricomycotinaMycologia 98["!#M"54 66644.666!_ 9 9G !"EF W O. N[ Ultrastructure G /] O@] .]O`UV `V Uet al.5664 Techniques for Microorganisms / 9G V ` [ + 5]Q[ % % O !E_M5!5> [;; Bioinformatics 21[ 9 G U 56!! U/2S 5[ _F#7M_F#F . . < 29 ] / V9 !"E6 Compendium of Soil BMC Bioinformatics 12[7"! Fungi*[/ ; 9 ` QR< 2* !"E# W O </+56!7WallemiomycotinaIndex Fungorum 73[! Experimental Mycology 11[546M5F" < S] ] U9 !"#5 ; . % ` V@@* 9 `29Get al.56!_Polyporales HomobasidiomycetidaeTransactions of the British Mycological % Society 58[_E#M_"5 Mycologia 105[!_#7M!_"6 S 2U +9 G U566" % ` ; + < G 9 W U * et al. 56!7 N ; @ 4[ . [ BMC Bioinformatics 10[F# Bioinformatics 30[!5_FM!576 W @ 56!5 V >G+N V $ Nucleic Acids ` @2 + ` 56!! W [ Research 40< [< et Ustilago maydisNature 444["#M!6! al. 566_ V ? [ 2 @ 2 `!"E5/ % Neurospora crassaNature 422[E4"MEFE Mycotaxon 15[!6_M ] ./ WQ %2G `G O*U] O@ !56 et al.56!5 [% $ . ? 2 ;U G ;WU < @ `/ 566E Ainsworth & Bioinformatics 28[5F#EM5F#" Bisby’s Dictionary of the Fungi. !6 [ G/+N ] /+ +]+9< % <+et al.!""F ; * F666Science 274[47F4F_M4F# 2 G; 9 .S / @ .@ X U O / [ ? >/Nucleic Acids Research 35[_!66M_!6E Molecular Biology and Evolution * +@ O @*56!5W . 27[55!M557 + 5Nature Methods 9[_4#M_4" ] % NX > 9 @ 2 / Q et al. 56!7 * >; 95667/+ $ . UG [ !666 Nucleic . Acids Research 42< [V ]56!_Z/@V[? * >* V+ ? @566";+ _[ + Bioinformatics 29E[ !6#5M!6#4 Bioinformatics 25[55EFM55EE 9 +`; U<GG/ ` 56!! * > >@ < `56!_;+ U;N[ / Mycology 2[ !!E. [ $ [ !7! % @ Biology 62[F!!MF!4

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initio % Genome Research % < 2]9 +UX /`@ Ù 9et al. ARTICLE 18[!"#"M!""6 566"@ $ Agaricomycotina V WG!"#7W Wallemia sebi Basidiomycota Cantharellales Canadian Journal of Botany 52[54E#M54"6 HymenochaetalesMycological Research 113[44"M4#F V SU U2/2 /@ /et al.56!_ ` @ G 9 ; ] / + V et al. 566# ] O % < . Malassezia % % Proceedings of the National % % Academy of Sciences, USA 110[56!!#M56!55 Proceedings of the National Academy of Sciences, VU2/9 +* O/V /et al.56!7 < USA 104[!E#_6M!E#_4 ? % Tilletiaria anomala ` * ` W * et al. 56!! ] Z+G"4!Genome Announcements 2_[664_".!7 Arthrobotrys oligospora VUQ / ` V* O 2*et al.56!7 % . PLoS Pathogens ] ? % % 7"[!665!#" Mixia ` * < 9 ` et al. 56!_ ] osmundaeNew Phytologist 202[447M4F7 ? Wallemia V @/ W W U ] V 2 W et al. ichthyophaga[ BMC 56!_V % . Genomics 14[F!# % [ % PLoS ; 9 ]@ @ 9` W Ù ].G Genetics 9"[!66_E56 > 5664 V $ $ % + U// / 2+ `9< Ùet WallemiaWallemiomycetes Wallemiales % al.566E]? [ Antonie van Leeuwenhoek 87[_!!M_5E Penicillium chrysogenum Nature Biotechnology 26[ !!F!M!!FE

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