ARTICLE 49 -like
DQG et al Key words: Key words: Anthracostroma Camarosporula Cladosporium Mycosphaerella phylogeny Sphaerulina taxonomy Teratosphaeria
Anthracostroma and Mycosphaerella RQ VSSLQ +DZDLL C. silenes , the basionym Camarosporula Persoonia
1 WHOHRPRUSK IMA FUNGUS · VOLUME 2 · NO 1: 49–64 · NO 2 · VOLUME FUNGUS IMA IURP 6RXWK $IULFD 7KH within the Capnodiales ZKLFKRFFXUVRQOHDYHVRI Etlingera elatior RQ LQWURGXFHG LQ WKLV VWXG\ LQFOXGH VS$XVWUDOLD DQG Sphaerella agapanthi Camarosporula One of the largest barriers to understanding host One of the largest barriers to understanding ivecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get Any of the above conditions can be waived ivecommons.org/licenses/by-nc-nd/3.0/legalcode. S. myriadea MATERIALS AND METHODS MATERIALS Isolates /HDI DQG VWHP WLVVXH EHDULQJZDWHU DVFRPDWD IRU DSSUR[LPDWHO\ ZHUH VRDNHG K DIWHU LQ ZKLFK WKH\ ZHUH SODFHG RUEOHPLVKHVVSRWVDQGVSHFNV %DW]HU VSHFL¿FLW\ DQG VSHFLDWLRQ LQ WKLV JURXS RI RUJDQLVPV KDV been the relative unavailability of authentic cultures, as most been the relative unavailability of authentic described without of the species known to date, have been WKHGHSRVLWRIDVVRFLDWHGH[W\SHFXOWXUHVRU'1$7KHUHVXOW species remains is that the ecological behaviour of many REVFXUH &URXV *URHQHZDOG ZKLOHWKHSK\ORJHQHWLF position of many other pathogens and genera remains position of many other pathogens and XQFHUWDLQ 'XULQJ WKH FRXUVH RI WKH SUHVHQW VWXG\ VHYHUDO potentially novel Mycosphaerella species were collected, while potential epitype specimens ZHUH DOVR FROOHFWHG IRU ROGHU ZHOOHVWDEOLVKHG QDPHV 7KH Sphaerulina aim of this study was thus to describe these species, and also elucidate the phylogenetic relationship of genera such as +DZDLL (SLW\SHVDUHDOVRGHVLJQDWHGIRU Mycosphaerellaceae M. etlingerae , and for Agapathus umbellatus
Grevillea RQ ZKLFKLVDOHDISDWKRJHQRI EDVHGRQ RQ DQG-RKDQQHV=*URHQHZDOG et al Mycosphaerellaceae persooniae
LQ(XURSH 3RUWXJDO )XUWKHUPRUHWZRVH[XDOVSHFLHVRI C. grevilleae et al. E DQGEHORQJVWRWKH Davidiellaceae Mycosphaerella $SWURRW 7KH1HWKHUODQGV DQG Teratosphaeriaceae , and Sphaerulina A. umbellatus Mycosphaerella &URXV complex are ecologically is actually polyphyletic Anthracostroma persooniae %UHWW$6XPPHUHOO et al. EF species are also among the has in the past been recognised et al complex Mycosphaerella Schizothyriaceae DUHGHVFULEHGQDPHO\ FRPE QRY Teratosphaeriaceae Hedychium coronarium is newly reported on Teratosphaeriaceae , teleomorph , the basionym of Camarosporula Dissoconiaceae and et al UHFHQWSK\ORJHQHWLFVWXGLHV Mycosphaerella-
et al IUXLWOHVLRQV 3UHWRULXV Cerastium semidecandrum .D]XDNL7DQDND 1 VSHFLHV 1HZ VSHFLHV RI Mycosphaerella 8. )LQDOO\WKHSK\ORJHQHWLFSRVLWLRQRIWZRJHQHUDDUHQHZO\FRQ¿UPHGQDPHO\ Davidiellaceae RQ et al &URXV %UDXQ 6SHFLHVDUH Species in the present study were compared based on their morphology, growth characteristics in culture, study were compared based on their morphology, Species in the present C. persooniae
Carpinus, Castanopsis, Fagus, Quercus 2 LQWKHERWWRPRI3HWULGLVKOLGVZLWKWKHWRSKDOIRIWKH GLVK FRQWDLQLQJ PDOW H[WUDFW DJDU 0($ 2[RLG +DPSVKLUH 8. &URXV et al. F $VFRVSRUH JHUPLQDWLRQ SDWWHUQV ZHUH H[DPLQHG DIWHU K DQG single ascospore and conidial cultures established as ARTICLE GHVFULEHGHDUOLHU &URXVet al &URXV 7R isolate asexual fungi, host tissues were incubated in PRLVWFKDPEHUVIRUXSWRZNDQGLQVSHFWHGGDLO\IRU microfungi, and single conidial colonies established on 0($ &URXV &RORQLHV ZHUH VXEFXOWXUHG RQWR SRWDWRGH[WURVH DJDU 3'$ V\QWKHWLF QXWULHQW SRRU DJDU 61$ 0($ DQG RDWPHDO DJDU 2$ Accession number (ITS, LSU, TEF, ACT) Accession number (ITS, LSU, TEF, &URXVet al F DQGLQFXEDWHGXQGHUFRQWLQXRXV QHDUXOWUDYLROHWOLJKWDW&WRSURPRWHVSRUXODWLRQ -)-)²² -)-)²² -)-) ²² -)-)²² -)-)²² -)-)²² -)-)²² () -)() () -)-)²² GenBank GenBank 1RPHQFODWXUDO QRYHOWLHV DQG GHVFULSWLRQV ZHUH GHSRVLWHG LQ 0\FR%DQN ZZZ0\FR%DQNRUJ &URXV et al. D $OO FXOWXUHV REWDLQHG LQ WKLV VWXG\ DUH maintained in the open collection of the Centraalbureau YRRU6FKLPPHOFXOWXUHV &%6.1$: DQGWKHZRUNLQJ FROOHFWLRQ &3& RI3:&URXVLQ&%6 7DEOH DNA phylogeny *HQRPLF '1$ ZDV H[WUDFWHG IURP P\FHOLD RI IXQJDO TM 3:&URXV 3:&URXV .7DQDND :*DPV :*DPV :*DPV $$SWURRW 3:&URXV %$6XPPHUHOO -)-)-)-) 3:&URXV %$6XPPHUHOO -) -)²² 3:&URXV %$6XPPHUHOO -)-)²² 3:&URXV %$6XPPHUHOO -)-) ²² colonies cultivated on MEA using the UltraClean 0LFURELDO '1$ ,VRODWLRQ .LW 0R %LR /DERUDWRULHV 6RODQD%HDFK&$86$ 7KH3ULPHUV9 * GH+RRJ *HUULWVYDQGHQ(QGH DQG/5 9LOJDO\V +HVWHU ZHUHXVHGWRDPSOLI\SDUWRIWKHQXFOHDUU'1$ South Africa South Africa 3RUWXJDO 3:&URXV Japan Hawaii Hawaii 1HWKHUODQGV $$SWURRW Hawaii 8. Australia Australia Australia Australia Country Collector(s) RSHURQ VSDQQLQJ WKH ¶ HQG RI WKH 6 QU51$ JHQH 668 WKH¿UVWLQWHUQDOWUDQVFULEHGVSDFHU ,76 WKH 6QU51$JHQHWKHVHFRQG,76UHJLRQ ,76 DQGWKH ¶HQGRIWKH6QU51$JHQH /68 7KHSULPHUV,76 Cerastium VS VS VS :KLWHet al. DQG/68)G &URXVet al E ZHUH XVHG DV LQWHUQDO VHTXHQFH SULPHUV WR HQVXUH VS Silene maritima JRRG TXDOLW\ VHTXHQFHV RYHU WKH HQWLUH OHQJWK RI WKH Hedychium coronarium Etlingera elatior Etlingera elatior Agapanthus umbellatus Agapanthus umbellatus Agapanthus umbellatus Persoonia Persoonia Persoonia DPSOLFRQ7RKHOSUHVROYHVSHFLHVRICladosporium, the ,76UHJLRQZDVVXSSOHPHQWHGZLWKSDUWLDOVHTXHQFHV Quercus dentata Grevillea of the translation elongation factor 1-_JHQH ()_ XVLQJWKHSULPHUV()) &DUERQH .RKQ /HDIVSRWVRQ Dead leaves of Dead leaves of Dead leaves and stems of semidecandrum Dead leaves of Stems of exposed DQG() 2¶'RQQHOOet al. DQGWKHDFWLQJHQH Substrate $&7 XVLQJ WKH SULPHUV $&7) DQG $&75 &DUERQH .RKQ 7KH3&5DPSOL¿FDWLRQVZHUH SHUIRUPHGRQD*HQH$PS3&56\VWHP $SSOLHG %LRV\VWHPV)RVWHU&LW\&$86$ LQDWRWDOYROXPHRI /VROXWLRQFRQWDLQLQJ±QJRIWHPSODWH'1$ 1 î3&5EXIIHUP00J&OSPROIRUHDFKSULPHU 0RIHDFKG173 0IRU()Į DQG8Taq '1$SRO\PHUDVH %LROLQH/XFNHQZDOGH*HUPDQ\ )RU &3& &%6 &3& /HDIVSRWVRQ &%6 &3& /HDIVSRWVRQ &%6 -&0 /HDYHVRI &%6 &3& &3& &%6 &3& &%6 &%6 &3& &%6 &%6 &3& &%6 /HDYHVRI &%6 &3&&%6 /HDIVSRWVRQ &%6 &3&&%6 /HDIVSRWVRQ Strain no. &3&&%6 /HDIVSRWVRQ ()Į/GLPHWK\OVXOIR[LGH '062 ZDVDGGHGWR WKHDPSOL¿FDWLRQUHDFWLRQ3&5DPSOL¿FDWLRQFRQGLWLRQV were set as follows: an initial denaturation temperature RI &IRUPLQIROORZHGE\F\FOHVRIGHQDWXUDWLRQ WHPSHUDWXUHRI &IRUVSULPHUDQQHDOLQJDW& &ROOHFWLRQGHWDLOVDQG*HQ%DQNDFFHVVLRQQXPEHUVRILVRODWHVIRUZKLFKQRYHOVHTXHQFHVZHUHJHQHUDWHGLQWKLVVWXG\ &IRU()Į IRUVSULPHUH[WHQVLRQDW&IRU VDQGD¿QDOH[WHQVLRQVWHSDW&IRUPLQ7KH UHVXOWLQJ DPSOLFRQV ZHUH VHTXHQFHG XVLQJ WKH 3&5 &%6 &%6.1$:)XQJDO%LRGLYHUVLW\&HQWUH8WUHFKW7KH1HWKHUODQGV &3& &XOWXUHFROOHFWLRQRI3HGUR&URXVKRXVHGDW&%6 QIDFWRUDOSKDJHQH $&7 SDUWLDODFWLQJHQH ,76 ,QWHUQDOWUDQVFULEHGVSDFHUVDQGWRJHWKHUZLWK6QU '1$ /68 SDUWLDO6QU'1$ 7() SDUWLDOWUDQVODWLRQHORQJDWLR Teratosphaeria agapanthi Teratosphaeria Sphaerulina myriadea Mycosphaerella holualoana Mycosphaerella cerastiicola Mycosphaerella etlingerae Cladosporium silenes Cladosporium grevilleae Table 1. Table Species Camarosporula persooniae 1 SULPHUV DQG D %LJ'\H 7HUPLQDWRU &\FOH 6HTXHQFLQJ 50 IMA FUNGUS Additions to the Mycosphaerella complex .LWY $SSOLHG%LRV\VWHPV DQGDQDO\VHGRQDQ$%,3ULVP XVHG LQ WKH SK\ORJHQHWLF DQDO\VLV ZHUH SDUVLPRQ\ ARTICLE [O'1$6HTXHQFHU 3HUNLQ(OPHU1RUZDON&186$ LQIRUPDWLYHZHUHYDULDEOHDQGSDUVLPRQ\XQLQIRUPDWLYH 7KH JHQHUDWHG VHTXHQFHV ZHUH FRPSDUHG ZLWK RWKHU DQGZHUHFRQVWDQW1HLJKERXUMRLQLQJDQDO\VLVXVLQJWKH IXQJDO '1$ VHTXHQFHV IURP 1&%,¶V *HQ%DQN VHTXHQFH WKUHHVXEVWLWXWLRQPRGHOVRQWKHVHTXHQFHGDWD\LHOGHGWUHHV GDWDEDVH XVLQJ D PHJDEODVW VHDUFK VHTXHQFHV ZLWK KLJK ZLWKVLPLODUWRSRORJ\DQGERRWVWUDSVXSSRUWYDOXHV7ZHQW\ VLPLODULW\ZHUHDGGHGWRWKHDOLJQPHQW /68 RUGLVFXVVHGLQ ¿YHHTXDOO\PRVWSDUVLPRQLRXVWUHHVZHUHREWDLQHGIURPWKH WKHVSHFLHVQRWHVZKHUHDSSOLFDEOH ,76()ĮDQG$&7 KHXULVWLFVHDUFKWKH¿UVWRIZKLFKLVVKRZQLQ)LJ 7/ 7KHDGGLWLRQDO*HQ%DQNVHTXHQFHVZHUHPDQXDOO\DOLJQHG &, 5, 5& 7KHSK\ORJHQHWLF XVLQJ6HTXHQFH$OLJQPHQW(GLWRUYD 5DPEDXW WUHHRIWKH/68UHJLRQ )LJ VKRZHGWKUHHGLVWLQFWJURXSV 7KH SK\ORJHQHWLF DQDO\VHV RI WKH DOLJQHG VHTXHQFH GDWD RI IXQJDO LVRODWHV WKH ¿UVW ZDV WKH Davidiellaceae clade ZHUH SHUIRUPHG XVLQJ 3$83 3K\ORJHQHWLF$QDO\VLV 8VLQJ ERRWVWUDSVXSSRUW WKHVHFRQGWKHTeratosphaeriaceae 3DUVLPRQ\ Y E 6ZRIIRUG DQG FRQVLVWHG RI FODGH ERRWVWUDS VXSSRUW DQG WKH WKLUG ZDV WKH QHLJKERXUMRLQLQJ DQDO\VHV ZLWK WKH XQFRUUHFWHG µS¶ WKH MycosphaerellaceaeFODGH ERRWVWUDSVXSSRUW .LPXUDSDUDPHWHUDQGWKH+.<VXEVWLWXWLRQPRGHOVDQG SDUVLPRQ\DQDO\VHV$OLJQPHQWJDSVZHUHWUHDWHGDVPLVVLQJ GDWDDQGDOOFKDUDFWHUVZHUHXQRUGHUHGDQGRIHTXDOZHLJKW TAXONOMY $Q\ WLHV ZHUH EURNHQ UDQGRPO\ ZKHQ HQFRXQWHUHG )RU SDUVLPRQ\DQDO\VHVDOLJQPHQWJDSVZHUHWUHDWHGDVD¿IWK During the present study several novel taxa were delineated, ³QHZ´ FKDUDFWHU VWDWH DQG DOO FKDUDFWHUV ZHUH XQRUGHUHG RU QHZO\ FROOHFWHG FXOWXUHV OLQNHG WR HVWDEOLVKHG QDPHV DQG RI HTXDO ZHLJKW 0D[LPXP SDUVLPRQ\ DQDO\VLV ZDV These species are treated per family below: performed using the heuristic search option with 100 random simple taxa additions and tree bisection and reconstruction Davidiellaceae 7%5 DVWKHEUDQFKVZDSSLQJDOJRULWKP%UDQFKHVRI]HUR OHQJWKZHUHFROODSVHGDQGDOOPXOWLSOHHTXDOO\SDUVLPRQLRXV Cladosporium grevilleae &URXV Summerell, sp. WUHHVZHUHVDYHG7KHUREXVWQHVVRIWKHWUHHVREWDLQHGZDV nov. HYDOXDWHGE\ERRWVWUDSUHSOLFDWLRQV +LOOLV %XOO 0\FR%DQN0% 7UHHOHQJWK 7/ FRQVLVWHQF\LQGH[ &, UHWHQWLRQLQGH[ 5, )LJV DQG UHVFDOHG FRQVLVWHQF\ LQGH[ 5& ZHUH FDOFXODWHG DQG WKHUHVXOWLQJWUHHVZHUHSULQWHGZLWK7UHH9LHZY 3DJH Etymology 1DPHGDIWHUWKHKRVWRQZKLFKLWZDVFROOHFWHG 1HZ VHTXHQFHV ZHUH ORGJHG LQ *HQ%DQN 7DEOH Grevillea and the alignments and phylogenetic trees in TreeBASE ZZZWUHHEDVHRUJ Asci fasciculati, bitunicati, subsessiles, obovoidei vel late ellipsoidei, RFWRVSRUL ± × ± P $VFRVSRUDH WUL DG SOXULVHULDWDV Morphology hyalinae, guttulatae cum inclusionibus angularibus, crassitunicatae, Culture characteristics were recorded from colonies grown on UHFWDHYHOOHQLWHUFXUYDWDHIXVRLGHVHOOLSVRLGHDHXWULQTXHREWXVDH 0($2$DQG3'$SODWHVDIWHU±ZNLQFXEDWLRQLQGDUNQHVV PHGLDQHVHSWDWDH ± ± ± ×± ± P DW&&RORQ\FRORXUV VXUIDFHDQGUHYHUVH ZHUHDVVHVVHG using the colour charts RI5D\QHU 3UHSDUDWLRQVZHUH Typus: Australia: New South Wales: Mount Annan Botanical PRXQWHGLQODFWLFDFLGDQGVWXGLHGXQGHUDOLJKWPLFURVFRSH î *DUGHQRQOHDYHVRIGrevilleaVS$XJ P.W. Crous PDJQL¿FDWLRQ 7KH FRQ¿GHQFHLQWHUYDOVRIVSRUHV & B.A. Summerell JT 974 '$5±KRORW\SXV FXOWXUHV ZHUHGHULYHGIURPREVHUYDWLRQV XQOHVVVWDWHGRWKHUZLVH H[W\SH&3& ± &%6 ZLWKH[WUHPHVJLYHQLQSDUHQWKHVHV In vivo: Leaf spotsDEVHQWAscomata occurring in leaf litter, DPSKLJHQRXVEODFNVXEHSLGHUPDOHUXPSHQWWRVXSHU¿FLDO RESULTS JORERVHWRPGLDPZLWKFHQWUDOSHULSK\VDWHRVWLROH ± P GLDP ZDOO RI ± OD\HUV RI EURZQ WH[WXUD DNA phylogeny DQJXODULV Asci aparaphysate, fasciculate, bitunicate with $SSUR[LPDWHO\ EDVHV VSDQQLQJ WKH ,76 DQG /68 ¿VVLWXQLFDWH GLVFKDUJH VXEVHVVLOH RERYRLG WR EURDGO\ UHJLRQVZHUHREWDLQHGIRULVRODWHVOLVWHGLQ7DEOH7KHVH HOOLSVRLG VOLJKWO\ FXUYHG VSRUHG ± × ± P ZLWK WZR UHJLRQV ZHUH DQDO\VHG VHSDUDWHO\ ,76 WR GHWHUPLQH YLVLEOH DSLFDO DSLFXOXV Ascospores tri- to multi-seriate, VSHFLHV OHYHO UHODWLRQVKLSV SUHVHQWHG LQ WKH VSHFLHV QRWHV hyaline, guttulate with angular inclusions, thick-walled, EHORZZKHUHDSSOLFDEOH DQG/68IRUWKHJHQHULFSODFHPHQW straight to slightly curved, fusoid-ellipsoid with obtuse ends, $SSUR[LPDWHO\DQGEDVHVZHUHGHWHUPLQHGIRU() medianly 1-septate, widest in middle of apical cell, slightly ĮDQG$&7UHVSHFWLYHO\DQGWKHLUUHVSHFWLYHEODVWUHVXOWV constricted at the septum, tapering towards both ends, but are discussed under the CladosporiumVSHFLHVQRWHVEHORZ VOLJKWO\PRUHWRORZHUHQG ± ± ± ×± ± P 7KHPDQXDOO\DGMXVWHG/68DOLJQPHQWFRQWDLQHG WD[D ascospores surrounded with a thin sheath when mounted in LQFOXGLQJWKHRXWJURXSVHTXHQFH DQGRIWKHFKDUDFWHUV ZDWHUEHFRPLQJEURZQDQGYHUUXFXORVHZLWKDJH DVFRVSRUH VOLUME 2 · NO. 1 51 Crous et al. Saccharomyces cerevisiae Z73326 Cladosporium sp. EU167586 70 Cladosporium bruhnei GU214408 100 Cladosporium grevilleae CPC 2913 Davidiellaceae 90 Cladosporium silenes CBS 109082 ARTICLE “Pseudocercospora” sp. GU214681 Teratosphaeria cryptica GU214505 98 90 CPC 18332 CPC 18304 Teratosphaeria agapanthi 2x 95 100 CPC 18266 63 Dothideomycetes sp. GU323983 Teratosphae- 54 Teratosphaeria encephalarti FJ372414 riaceae Teratosphaeriaceae sp. GU323219 Phaeothecoidea melaleuca HQ599595 CBS 116258 55 CBS 112494 Camarosporula persooniae 100 CBS 112302 89 Mycosphaerella colombiensis DQ204745 100 10 changes CPC 12277 Mycosphaerella etlingerae CPC 12274 100 Mycosphaerella irregulariramosa GQ852609 Mycosphaerella holualoana GU214440 90 CPC 12286 Mycosphaerella holualoana Mycosphaerella crystallina EU167579 Mycosphaerella waimeana AY260083 Mycosphaerella acaciigena GU214432 93 Mycosphaerella heimioides GU214439 77 Mycosphaerella heimii GU214438 Pseudocercospora eucommiae GU253742 98 Septoria obesa GU214493 50 69 CBS 124646 Sphaerulina myriadea Mycosphae- Mycosphaerella latebrosa GU214444 rellaceae 58 Septoria aceris GQ852673 77 Mycosphaerella berberidis EU167603 Mycosphaerella harthensis EU167602 Mycosphaerella flageoletiana EU167597 Ramularia endophylla AY490776 93 Ramularia proteae EU707899 Ramularia aplospora EU040238 57 Ramularia acroptili GU214689 79 Ramularia sp. GU214693 CBS 115913 Mycosphaerella cerastiicola 63 Cercosporella centaureicola EU019257 Mycosphaerella fragariae GU214691 Ramularia miae DQ885902 Ramularia pratensis var. pratensis EU019284 Ramularia sp. GU214690 Ramularia coleosporii GU214692 Ramularia uredinicola GU214694 Fig. 1. 7KH¿UVWRIHTXDOO\PRVWSDUVLPRQLRXVWUHHVREWDLQHGIURPDKHXULVWLFVHDUFKZLWKUDQGRPWD[RQDGGLWLRQVRIWKH/68VHTXHQFH DOLJQPHQWXVLQJ3$83YE7KHVFDOHEDUVKRZVFKDQJHVDQGERRWVWUDSVXSSRUWYDOXHVIURPUHSOLFDWHVDUHVKRZQDWWKHQRGHV 7KLFNHQHGOLQHVLQGLFDWHWKHVWULFWFRQVHQVXVEUDQFKHVDQGQRYHOVHTXHQFHVDUHSULQWHGLQEROGIDFH7KHGLIIHUHQWIDPLOLHVDUHLQGLFDWHGZLWK FRORXUHGEORFNV7KHWUHHZDVURRWHGWRSaccharomyces cerevisiae *HQ%DQNQR= 52 IMA FUNGUS Additions to the Mycosphaerella complex ARTICLE Fig. 2. Cladosporium grevilleae '$5 A, B.$VFLC.2VWLRODUUHJLRQ DUURZHG D.$VFXVZLWKDVFRVSRUHV%DUV P Fig. 3. Cladosporium grevilleae '$5 $VFLZLWKDVFRVSRUHV$VFRVSRUHVVKRZLQJVKHDWKDQGDQJXODULQFOXVLRQV%DU P germination with germ tubes parallel to the long axis of the Notes: The genus CladosporiumLVOLQNHGWRWHOHRPRUSKV VSRUHEXWGLVWRUWLQJSURPLQHQWO\ RULJLQDOVSRUHFHOOVXSWR that are placed in Davidiella %UDXQet al.6FKXEHUW P ZLGH JHUPLQDWLQJ ZLWK QXPHURXV JHUP WXEHV IRUPLQJ et al ,QPRYLQJWRDVLQJOHQRPHQFODWXUHIRUSOHRPRUSKLF dense clusters of hyphae, but mostly remaining hyaline after fungi, we give preference to the oldest genus name, namely KRQ0($ Cladosporium Cladosporium grevilleae only forms the VH[XDO VWDWH RI WKH OLIHF\FOH 7KH DVFRVSRUHV DUH W\SLFDO RI Culture characteristics &RORQLHV DIWHU ZN DW & a Cladosporium teleomorph, having thick walls, and angular VSUHDGLQJ UHDFKLQJ ± PP GLDP 2Q 0($ HUXPSHQW LQFOXVLRQV $SWURRW EHFRPLQJ EURZQ DQG YHUUXFXORVH ZLWKHYHQOREHGPDUJLQV VXUIDFHIROGHGZLWKVSDUVHDHULDO ZLWKDJH3UHVHQWO\ZHDUHQRWDZDUHRIDQ\VH[XDORUDVH[XDO P\FHOLXP ROLYDFHRXVJUH\ UHYHUVH LURQJUH\ 2Q 2$ ÀDW species on Grevillea that represent this fungus, and thus with sparse aerial mycelium, and lobed, somewhat feathery ZH GHVFULEH LW KHUH DV QHZ$OWKRXJK DOO IRXU ORFL VXSSRUWHG PDUJLQV VXUIDFHSDOHROLYDFHRXVJUH\LQPLGGOHLURQJUH\LQ the association of the species with Cladosporium, it did not RXWHUUHJLRQ2Q3'$HUXPSHQWZLWKOREHGIHDWKHU\PDUJLQV match any of the CladosporiumVHTXHQFHVFXUUHQWO\DYDLODEOH VXUIDFHIROGHGZLWKVSDUVHDHULDOP\FHOLXPJUH\ROLYDFHRXV RQWKH*HQ%DQNQXFOHRWLGHGDWDEDVH FORVHVWPDWFKRQ,76 UHYHUVHLURQJUH\ was Davidiella macrospora *HQ%DQN (8 ZLWK VOLUME 2 · NO. 1 53 Crous et al. ARTICLE Fig. 4. Cladosporium silenes &%6+ A.$VFRPDWDRQKRVWWLVVXH DUURZV B.$VFLC.$VFRVSRUHV DUURZGHQRWHVPXFRLGDSSHQGDJH $±&IURP%HQVFKet al. D–F.&RQLGLRSKRUHVZLWKFRQLGLDOFKDLQV%DUV P LGHQWLW\RQ()ĮZDVCladosporium myrtacearum*HQ%DQN discharge, subsessile, obovoid to broadly ellipsoid, straight +0ZLWK LGHQWLW\DQGRQ$&7LWZDVCladosporium WRVOLJKWO\FXUYHGVSRUHG±×±PZLWKYLVLEOH iranicum*HQ%DQN+0 ZLWK LGHQWLW\ DSLFDODSLFXOXVAscospores tri- to multiseriate, hyaline, non- guttulate with angular inclusions, thick-walled, straight to Cladosporium silenes Crous, sp. nov. slightly curved, fusoid-ellipsoid with obtuse ends, medianly 0\FR%DQN0% 1-septate, widest in middle of apical cell, not to slightly )LJ constricted at the septum, tapering towards both ends, but VOLJKWO\ PRUH WR ORZHU HQG ± ± ± × ± ± Etymology 1DPHGDIWHUWKHKRVWRQZKLFKLWZDVFROOHFWHG P WXUQLQJ EURZQ RQFH GLVFKDUJHG DQG VRPH FRQWDLQLQJ Silene maritima UHPQDQWV RI D PXFRLG OD\HU JHUPLQDWLQJ IURP ERWK HQGV GLVWRUWLQJEHFRPLQJEURZQDQG¿QHO\YHUUXFXORVH Cladosporii cladosporioidis similis, sed conidiophoris brevioribus, Mycelium consisting of branched, septate, pale to medium non ramosis, cellulis conidiogenois longioribus, ramo-conidiis et EURZQ ± P ZLGH K\SKDH ZLWKRXW DQ\ VZHOOLQJV DQG FRQLGLLVLQWHUFDODULEXVEUHYLRULEXVGLVFHUQLWXU FRQVWULFWLRQVVPRRWKWRPLQXWHO\YHUUXFXORVHZDOOVXQWKLFNHQHG Conidiophores solitary, macronematous or micronematous, Typus: UK: Pembrokeshire: Skomer Island, stems of exposed arising terminally from ascending hyphae or laterally, straight Silene maritima $XJ A. Aptroot &%6 WR VRPHZKDW ÀH[XRXV QDUURZO\ F\OLQGULFDO WR F\OLQGULFDO + ±KRORW\SXV FXOWXUHH[W\SH&%6 REORQJQRQQRGXORVHQRWJHQLFXODWHVLQXRXVí í × í í PXQEUDQFKHGSOXULVHSWDWHXVXDOO\QRWFRQVWULFWHG In vivo: Ascomata occurring in exposed twigs, amphigenous, at septa, pale to medium olivaceous-brown, smooth to minutely EODFN WR GDUN EURZQ VXEHSLGHUPDO WR P GLDP YLVLEOH verruculose, especially towards the base, walls unthickened, E\ DQ HUXPSHQW FHQWUDO SHULSK\VDWH RVWLROH ± P EDVH VRPHWLPHV VZROOHQ XS WR P ZLGH PLFURQHPDWRXV ZDOO RI ± OD\HUV RI UHGEURZQ textura angularis Asci FRQLGLRSKRUHV VKRUWHU XQEUDQFKHG í ×íP DSDUDSK\VDWH IDVFLFXODWH ELWXQLFDWH ZLWK ¿VVLWXQLFDWH Conidiogenous cells integrated, usually terminal, sometimes 54 IMA FUNGUS Additions to the Mycosphaerella complex ARTICLE Fig. 5. Mycosphaerella cerastiicola &%6 A, B. /HDYHV ZLWK EODFN DVFRPDWD DQG FRQLGLRPDWD C, D.$VFL E, F.$VFRVSRUHV G. &RQLGLRSKRUHVJLYLQJULVHWRFRQLGLDH.&RQLGLD%DUV P intercalary, cylindrical-oblong, not geniculate, non-nodulose, Notes 6WUDLQ&%6 UHSUHVHQWVDQDVFRVSRUHLVRODWH í íPORQJZLWKXSWRWKUHHORFLFURZGHGDWWKHDSH[ obtained from material of Silene maritima and initially VXEGHQWLFXODWH WR GHQWLFXODWH SURWXEHUDQW í P GLDP LGHQWL¿HGDVMycosphaerella tassianaYDUarthopyrenioides FHQWUDO GRPH PRVWO\ ÀDW VRPHZKDW WKLFNHQHG DQG GDUNHQHG Morphologically the CBS strain is similar to C. cladosporioides, UHIUDFWLYH Ramoconidia straight to slightly curved, cylindrical- EXW LV D PHPEHU RI D GLVWLQFW FODGH C. cladosporioides REORQJí ± ×í ± PDVHSWDWHSDOHROLYDFHRXV s.lat /LQHDJH VHH ¿J SDUW$ LQ %HQVFK et al. brown, concolorous with tips of conidiophores, smooth, base not Cladosporium sileniae LVWKH¿UVWVH[XDOVSHFLHVNQRZQIURP FODGRVSRULRLGíPZLGHWKLFNHQHGVRPHZKDWUHIUDFWLYH the cladosporioidesFRPSOH[*LYHQWKDWWKHVHDVFRPDWDDUH Secondary ramoconidia aseptate, smooth, pale olivaceous UDWKHULQFRQVSLFXRXV LPPHUVHGDSSUR[PGLDP LWV EURZQF\OLQGULFDOREORQJ ± ± ± ×±PConidia not surprising that they have been largely overlooked in the QXPHURXV FDWHQDWH LQ EUDQFKHG FKDLQV RI XS WR LQ WKH SDVWCladosporium sileniae differs from C. cladosporioides XSSHUXQEUDQFKHGSDUWEUDQFKLQJLQDOOGLUHFWLRQVIntercalary in having shorter, unbranched conidiophores, longer conidia OLPRQLIRUP HOOLSVRLGRYRLG í í × í í conidiogenous cells, shorter ramo- and intercalary conidia P DVHSWDWH ZLWK XS WR GLVWDO KLOD Small terminal conidia %HQVFKet al DVHSWDWH VXEJORERVH RERYRLG RYRLG WR OLPRQLIRUP í í × í í P KLOD GDUNHQHG DQG VRPHZKDW WKLFNHQHG Mycosphaerellaceae ±PGLDP Mycosphaerella cerastiicola Crous, sp. nov. Culture characteristics &RORQLHVDIWHUZNDW &VSUHDGLQJ 0\FR%DQN0% with moderate aerial mycelium and smooth, lobate margins, )LJ UHDFKLQJ PP GLDP DIWHU ZN 2Q 0($ VXUIDFH SDOH Anamorph: Septoria-like ROLYDFHRXVJUH\WRROLYDFHRXVJUH\UHYHUVHLURQJUH\1R2$ VXUIDFHJUH\ROLYDFHRXVWRROLYDFHRXVJUH\2Q3'$VXUIDFH Etymology 1DPHGDIWHUWKHKRVWRQZKLFKLWZDVFROOHFWHG JUH\ROLYDFHRXVUHYHUVHROLYDFHRXVJUH\ Cerastium semidecandrum VOLUME 2 · NO. 1 55 Crous et al. Ascosporae imbricatae, hyalinae, guttulatae, tenuitunicatae, rectae PLGGOHVDOPRQLQRXWHUUHJLRQ&RORQLHVRQ61$GHYHORSLQJ vel leniter curvatae, fusoides-ellipsoideae, constrictae ad septa DGLIIXVHUHGSLJPHQWLQDJDU PHGLDQDXWULQTXHDWWHQXDWDH ± ± ± × ± ± P Notes: Mycosphaerella cerastiicola is presently the Typus: Netherlands: Flevoland 1RRUGRRVWSROGHU QHDU only Mycosphaerella species known from Cerastium ARTICLE 8UN NP EORFN FRRUGLQDWLRQV RQ Caryophyllaceae $SWURRW %RWK ,76 DQG /68 GDWD dyke of former sea, on dead leaves and stems of Cerastium )LJ VXSSRUW WKH SODFHPHQW RI WKLV JHQXV ZLWKLQ WKH semidecandrum 0D\ A. Aptroot &%6 + ± Ramularia FODGH $ EODVW VHDUFK XVLQJ WKH ,76 VHTXHQFH KRORW\SXV FXOWXUHVH[W\SH&3& &%6 &3& REWDLQV DV FORVHVW KLWV PD[LPXP LGHQWLW\ RI ± DQ undescribed species of “Pseudocercosporella” VWUDLQV.$&& .$&& &3& *HQ%DQN () In vivo: Leaf spots DEVHQW Ascomata amphigenous on () *8 ZKLFK DSSHDUV WR EHORQJ WR WKLV leaves and stems, black, subepidermal, becoming erumpent, JHQXV7KHJHQXVRamularia has thus far been assumed to VROLWDU\ JORERVH XS WR P GLDP FHQWUDO RVWLROH ± be monophyletic, and thus the description of M. cerastiicola P GLDP ZDOO FRQVLVWLQJ RI ± OD\HUV RI PHGLXP EURZQ with its Septoria-like to Pseudocercosporella-like anamorph textura angularisAsci aparaphysate, fasciculate, bitunicate LVDQHQLJPD3UHVHQWO\WKHUHLVQRERRWVWUDSVXSSRUWIRULW ZLWK¿VVLWXQLFDWHGLVFKDUJHVXEVHVVLOHQDUURZO\HOOLSVRLGWR to represent another genus than Mycosphaerella/Ramularia, VXEF\OLQGULFDOVWUDLJKWWRVOLJKWO\FXUYHG ± VSRUHG± UDLVLQJ WKH TXHVWLRQ LI WKH DQDPRUSK FRXOG UHSUHVHQW D ×±P DSLFDOFKDPEHU±PGLDPAscospores possible Ramularia synanamorph? Further collections would bi- to tri-seriate, overlapping, hyaline, guttulate, thin-walled, EHUHTXLUHGWRDGGUHVVWKLVLVVXH straight to somewhat curved, fusoid-ellipsoidal with acutely URXQGHGHQGVZLGHVWDWVHSWXPZKHQLPPDWXUHMXVWDERYH Mycosphaerella etlingerae Crous, sp. nov. median septum when mature, constricted at the septum, 0\FR%DQN0% WDSHULQJ WRZDUGV ERWK HQGV ± ± ± × ± ± )LJ P DVFRVSRUHVJHUPLQDWHIURPERWKHQGVZLWKJHUPWXEHV SDUDOOHOWRWKHORQJD[LVRIWKHVSRUH Etymology 1DPHGDIWHUWKHKRVWRQZKLFKLWZDVFROOHFWHG SeptoriaOLNH VWDWH GHYHORSLQJ RQ 61$ DQG 2$ Etlingera elatior Conidiomata pycnidial on host, pycnidial to sporodochial on agar, black under the dissecting scope, globose, immersed Ascosporae imbricatae, hyalinae, granulatae, tenuitunicatae, rectae LQ DJDU WR VXSHU¿FLDO FRORQLHV DOVR VSRUXODWLQJ SURIXVHO\ vel leniter curvatae, fusoides-ellipsoideae, mediane 1-septatae, RQ DHULDO P\FHOLXP YLD VROLWDU\ ORFL P GLDP P OHQLWHUFRQVWULFWDHDGVHSWD ± ± ± × ± P WDOO WKLFNHQHG DORQJ WKH ULP Stromata dark brown under WKH FRPSRXQG PLFURVFRSH WR P GLDP P WDOO Typus: USA: Hawaii: on dead leaves of Etlingera elatior consisting of dark brown, thickened, ovoid to globose cells, $XJW. Gams &%6+±KRORW\SXV FXOWXUHVH[ ± P GLDP XSSHU OD\HUV IHUWLOH JLYLQJ ULVH WR D GHQVH W\SH&%6 &3&± network of intermingled brown, hyphal-like elements, WKDW LQ WXUQ JLYH ULVH WR DQ XSSHU OD\HU RI FRQLGLRSKRUHV In vivo: Leaf spots DEVHQW Ascomata amphigenous, black, ConidiophoresF\OLQGULFDO±VHSWDWHSDOHEURZQWRK\DOLQH VXEHSLGHUPDO EHFRPLQJ HUXPSHQW JORERVH XS WR P at base, hyaline at apex, smooth, rarely branched at bulbous GLDP FHQWUDO RVWLROH ± P GLDP ZDOO FRQVLVWLQJ RI ± EDVH±×PConidiogenous cells terminal, hyaline, layers of medium brown textura angularisAsci aparaphysate, ± × ± P SUROLIHUDWLQJ V\PSRGLDOO\ ZLWK ÀDWWHQHG IDVFLFXODWHELWXQLFDWHZLWK¿VVLWXQLFDWHGHKLVFHQFHVXEVHVVLOH ORFL±PGLDPXQWKLFNHQHGDQGQRWGDUNHQHG RQKRVW narrowly ellipsoid to subcylindrical, straight to slightly curved, PDWHULDO FRQLGLRJHQRXV FHOOV DPSXOOLIRUP ± P ORQJ VSRUHG ± ×±PAscospores bi- to tri-seriate, ± P ZLGH DW EXOERXV EDVH V\PSRGLDO RU SKLDOLGLF ZLWK overlapping, hyaline, granular, thin-walled, straight to curved, SHULFOLQDOWKLFNHQLQJYLVLEOHConidia solitary, hyaline, smooth, fusoid-ellipsoidal with obtuse ends, widest in middle of apical guttulate, cylindrical with obtuse apex and truncate base, cell, medianly 1-septate, slightly constricted at the septum, at times somewhat long obconically subtruncate on host tapering towards both ends, but more prominently towards the PDWHULDO±VHSWDWHVWUDLJKWWRÀH[XRXV ± ± ± ORZHUHQG ± ± ± × ± P DVFRVSRUHVJHUPLQDWH × ± P DY×P in vitro from both ends, with germ tubes growing parallel to the long D[LV GHYHORSLQJ ODWHUDO EUDQFKHV DVFRVSRUHV UHPDLQLQJ Culture characteristics &RORQLHV DIWHU ZN DW & K\DOLQHEHFRPLQJFRQVWULFWHGWRPZLGH spreading, erumpent, with folded surface, splitting agar, with sparse aerial mycelium and smooth, lobate margins, Culture characteristics: &RORQLHVDIWHUZNDW &VSUHDGLQJ UHDFKLQJ±PPGLDP2Q0($VXUIDFHVDOPRQWRÀHVK erumpent, surface irregular, folded, margin smooth, lobate, WRGLUW\ZKLWHUHYHUVHFKHVWQXW FHQWUH WRDSULFRW PDUJLQ ZLWKVSDUVHDHULDOP\FHOLXPUHDFKLQJPPGLDP2Q0($ 2Q2$VXUIDFHROLYDFHRXVEXIIIHUWLOH2Q3'$VXUIDFHGLUW\ surface pale mouse grey with patches of olivaceous grey ZKLWH ZLWK SDWFKHV RI ROLYDFHRXV JUH\ UHYHUVH LURQJUH\ LQ DQGVFDUOHWZLWKZKLWHPDUJLQ UHYHUVHLURQJUH\ UHGFU\VWDOV 56 IMA FUNGUS Additions to the Mycosphaerella complex ARTICLE Fig. 6. Mycosphaerella etlingerae &3& A.&RORQ\RQPDOWH[WUDFWDJDUB, C.$VFLD.$VFRVSRUHVE–G.*HUPLQDWLQJDVFRVSRUHV%DUV P SURGXFHGLQDJDU2Q2$VXUIDFHROLYDFHRXVJUH\2Q3'$ Mycosphaerella holualoana Crous et al., Mycotaxon surface smoke-grey with patches of pale olivaceous-grey and 78 VFDUOHW )LJ Notes 1R Mycosphaerella species have previously been In vivo: Leaf spots DEVHQW Ascomata amphigenous, black, reported from Etlingera elatior Zingiberaceae WKRXJK VL[ subepidermal, becoming erumpent, aggregated in clusters, species have been reported on Zingiberaceae7KHVHLQFOXGH JORERVH WR P GLDP FHQWUDO RVWLROH ± P GLDP M. alpiniae and M. alpiniicola on Alpinia in China, M. amomi ZDOO FRQVLVWLQJ RI ± OD\HUV RI PHGLXP EURZQ textura on Amomum in China, M. hedychii on HedychiumLQ%UD]LOM. angularis Asci aparaphysate, fasciculate, bitunicate with zingiberi on ZingiberIURP&KLQDDQG.RUHDDQGM. zingiberis on ¿VVLWXQLFDWH GHKLVFHQFH VXEVHVVLOH QDUURZO\ HOOLSVRLG WR Zingiber IURP -DSDQ ZZZQWDUVJULQJRYIXQJDOGDWDEDVHV VXEF\OLQGULFDOVWUDLJKWWRVOLJKWO\FXUYHGVSRUHG±× Other than for M. hedychii DVFRVSRUHV±×P 6RDUHV ± P Ascospores bi- to tri-seriate, overlapping, hyaline, %DUUHWR WKHRWKHUVSHFLHVDUHVRPHZKDWREVFXUHDQG prominently guttulate, thin-walled, straight, fusoid-ellipsoidal $SWURRW FRXOG QRW WUDFH DQ\ PDWHULDO IRU FRPSDULVRQ ZLWK REWXVH HQGV ZLGHVW MXVW DERYH VHSWXP PHGLDQO\ More than one species of Mycosphaerella was observed on the 1-septate, constricted at the septum, tapering towards both leaves of Etlingera elatior collected in Hawaii, but ascospores HQGVEXWPRUHSURPLQHQWO\WRZDUGVWKHORZHUHQG ± ± of M. etlingerae could be distinguished from other taxa by ± × ± ± P DVFRVSRUHVJHUPLQDWHIURPERWK being IXVRLGHOOLSVRLGDODQGZLGHVWLQPLGGOHRIWKHDSLFDOFHOO ends, with several germ tubes that are irregular in width and Mycosphaerella etlingerae LV LGHQWLFDO WR VHTXHQFHV JURZWKGLUHFWLRQ DVFRVSRUHVEHFRPLQJVOLJKWO\FRQVWULFWHGDW of M. thailandica and M. colombiensisIRUERWK,76DQG/68 VHSWXPWRPZLGHUHPDLQLQJK\DOLQHVPRRWK Mycosphaerella colombiensis, described from Eucalyptus leaf spots in Colombia, has ascospores that are obovoid, ± ± Culture characteristics &RORQLHVDIWHUZNDW &VSUHDGLQJ ± ×± P &URXV WKXVTXLWHGLVWLQFWIURP erumpent, surface folded with sparse aerial mycelium and M. etlingerae. Mycosphaerella thailandica has ascospores HYHQOREDWHPDUJLQUHDFKLQJPPGLDP2Q0($VXUIDFH WKDW DUH IXVRLGHOOLSVRLGDO ± ± ± × ± ± P SDOHROLYDFHRXVJUH\ZLWKSDWFKHVRIROLYDFHRXVJUH\ UHYHUVH thus being very similar to those of M. etlingerae. However, ROLYDFHRXVJUH\WRLURQJUH\2Q2$VXUIDFHROLYDFHRXVJUH\ M. thailandica causes a disease of Acacia in Thailand, and UHYHUVH ROLYDFHRXVJUH\ ZLWK SDWFKHV RI VFDUOHW 2Q 3'$ has a Pseudocercospora state in vivo and in vitro &URXVet al. surface grey-olivaceous to olivaceous grey, reverse iron-grey F ZKLFKLVGLIIHUHQWIURPWKDWREVHUYHGLQM. etlingerae. ZLWKSDWFKHVRIVFDUOHWLQFHQWUH VOLUME 2 · NO. 1 57 Crous et al. ARTICLE Fig. 7. Mycosphaerella holualoana &3& A, B.$VFLC.$VFRVSRUHVD, E.*HUPLQDWLQJDVFRVSRUHV%DUV P Specimen examined: USA: Hawaii: on dead leaves of Hedychium EHORZ VWUDLJKW RU VOLJKWO\ FXUYHG VHSWDWH ZLWK D SULPDU\ coronarium$XJW. Gams &%6+ FXOWXUHV&%6 VHSWXPQHDUO\PHGLDQ ±DY K\DOLQHVPRRWK &3& ZLWKRXWVKHDWKRUDSSHQGDJHV ± ± ± × ± ± ± P DY×P /: ± DY Notes: Mycosphaerella holualoana, which is part of the M. heimii- FRPSOH[ &URXV EDVHGRQLWV/68DQG,76VHTXHQFHV Culture characteristics &RORQLHVDIWHUZNDW &VSUHDGLQJ was initially described from leaf spots of Leucospermum VSS erumpent, with sparse aerial mycelium and feathery margins, FROOHFWHG LQ +DZDLL 7D\ORU et al $VFRVSRUHV ZHUH UHDFKLQJ PP GLDP 2Q 0($ VXUIDFH VDIIURQ UHYHUVH VRPHZKDW ODUJHU WKDQ LQ WKH SUHVHQW FROOHFWLRQ ± × OXWHRXV2Q2$VXUIDFHVDIIURQ2Q3'$VXUIDFHGLUW\ZKLWH ± P EXW WKH JHQHUDO VKDSH FXOWXUH FKDUDFWHULVWLFV DQG WRSDOHOXWHRXV UHYHUVHSDOHOXWHRXV germination patterns suggest that this is the same species, occurring on dead leaves of Hedychium coronarium$QRWKHU Specimens examined: UK: sine loc., on leaves of Quercus robur, species of Mycosphaerella, M. hedychii, is also known from J.E. Vize>0LFURIXQJL%ULW([1R @ H[,0,. 0 WKLVKRVWLQ+DZDLLDQG%UD]LO 6WHYHQV 6RDUHV %DUUHWR ± Japan: Aomori 7VXJDUX .LGXNXUL %HQVHPDUVK ¶¶¶ 1 ,W GLIIHUV IURP M holualoana by being associated with ¶¶¶( RQOHDYHVRIQuercus dentata$SUK. Tanaka EURZQOHDIVSRWV±PPGLDPKDYLQJVKRUWHUDVFL ±× 2243 ++8) VLQJOHDVFRVSRUHFXOWXUH&%6 -&0 ±P DQGVPDOOHUDVFRVSRUHV ±×P ±Germany: Driesen, Lasch >5DEHQKRUVW)XQJL( X U QR @ / ±USA: California 6HTXRLD1DWLRQDO3DUNDOW PRQ Sphaerulina myriadea '& 6DFFMichelia 1 leaves of Castanopsis sempervirens-XQ H.E. Parks %3, /DNH&R+REHUJ¶V5HVRUWRQOHDYHVRIQuercus kelloggii, )LJ 0D\ V. Miller %3, Maryland: Marlboro, on leaves of Quercus alba, $SU C.L. Shear %3, Texas: Leaf spots epiphyllous, round to irregular shaped, grey to Houston, on leaves of Quercus alba, $SU H.W. Ravenel %3, EODFN±PPGLDP DYPP Ascomata pseudothecial, LPPHUVHGVXEHSLGHUPDOHUXPSHQWDWWKHWRSVLQJOHWR± grouped, globose in longitudinal section, glabrous, without Notes: The genus Sphaerulina, which is based on S. SURPLQHQWEHDN±PWDOO DYP ±P myriadea ZLWK VHSWDWH DVFRVSRUHV ZDV GLVWLQJXLVKHG GLDP DY P Ostiole FHQWUDO ± P ZLGH ZLWK from MycosphaerellaZLWKVHSWDWHDVFRVSRUHV7KHVHWZR K\DOLQHLQQHUSHULSK\VHVRI±PZLGHAscomatal wall genera have traditionally been separated on the basis of textura angularis LQ VXUIDFH YLHZ LQ ORQJLWXGLQDO VHFWLRQ DVFRVSRUHVHSWDWLRQDORQH+RZHYHUVHYHUDOVSHFLHVZLWKD ± P WKLFN DW VLGH DQG EDVH FRPSRVHG RI ± OD\HUV Sphaerulina DVFRPDWDO DQDWRP\ DQG VHSWDWH DVFRVSRUHV RISRO\JRQDOWRVXEJORERVHEURZQFHOOV ±×± P are known that do not belong to Sphaerulina s. str., suggesting ± P WKLFN DURXQG DSLFDO RVWLROH ,QWHUDVFDO ¿ODPHQWV WKDWDVFRVSRUHVHSWDWLRQDORQHLVLQVXI¿FLHQWO\UREXVWWRLQIHU QRWVHHQAsciELWXQLFDWHLQVWUXFWXUHGLVFKDUJH¿VVLWXQLFDWH SK\ORJHQHWLFUHODWHGQHVV &URXVet al 7KHPDWWHULV FOXVWHUHGDULVLQJIURPWKHFHQWUXPEDVHRI±PWKLFN further complicated as S. myriadea, which occurs on hosts in cylindrical to obclavate, rounded at apex, with or without a the FagaceaeDSSHDUVWREHDVSHFLHVFRPSOH[%HFDXVHRI VKDOORZDSLFDOFKDPEHU caPKLJK ZLWKDNQREOLNH this, no epitype is designated here, pending further collections VWLSH ±PORQJ RUVHVVLOHZLWKWULVHULDWHWRELVHULDWH of authentic European material on QuercusIURP)UDQFH DVFRVSRUHV±×± ± P DY ×P In our study, S. myriadea clusters in Mycosphaerellaceae, Ascospores cylindrical, rounded at apex, slightly tapered at as sister to Septoria s.str. )LJ $OWKRXJK WKH QDPH 58 IMA FUNGUS Additions to the Mycosphaerella complex ARTICLE Fig. 8. Sphaerulina myriadea ++8) A./HDIVSRWRIQuercus dentataB.&ORVHXSRIHUXPSHQWDVFRPDWDC, D./RQJLWXGLQDOVHFWLRQ RIDVFRPDWDE.)DVFLFXODWHDVFLF. )LVVLWXQLFDWHDVFXVG, H.$VFLI.$SLFHVRIDVFLJ–N.$VFRVSRUHVO.*HUPLQDWLQJDVFRVSRUH%DUV $ PP %& P '(2 P )±1 P VOLUME 2 · NO. 1 59 Crous et al. ARTICLE Fig. 9. Camarosporula persooniae &%6 A./HDIVSRWZLWKDVFRVWURPDWDB, C.$FHUYXOLD, E.$VFLF.$VFRVSRUHVG, H.&RQLGLD%DUV P Sphaerulina SUHGDWHV WKDW RI Mycosphaerella Syll. Fung. 17 Mycosphaerellaceae have recently been shown to represent Pseudosphaerella persooniae +HQQ +DQVI Proc. Linn. D JHQHULF FRPSOH[ &URXV et al D E LQ ZKLFK Soc. New South Wales 79 Sphaerulina DSSHDUV WR UHSUHVHQW D GLVWLQFW OLQHDJH Mycosphaerella is restricted to species with Ramularia Leaf spots amphigenous, irregular, dark brown to black, DQDPRUSKV 9HUNOH\ et al DQG UHPDLQV GLVWLQFW IURP VSHFNVRIPPFRDOHVFLQJWRODUJHUVSRWVWRFPGLDP Sphaerulina$PHJDEODVWVHDUFKXVLQJWKH,76VHTXHQFHRI Ascostromata amphigenous, predominently epiphyllous, S. myriadea places it with members of Mycosphaerella, but subcuticular, uni-to multi-locular, erumpent, solitary to ZLWKDKLJKHVWTXHU\FRYHUDJHRI IRUMycosphaerella DJJUHJDWHGWRPGLDPZLWKVLQJOHFHQWUDOSHULSK\VDWH brassicicola *HQ%DQN (8 ,GHQWLWLHV RVWLROH ± P GLDP ZDOO FRQVLVWLQJ RI EURZQ WKLFN *DSV walled, textura angularis EDVDO VWURPD RQ WKH HSLGHUPLV FRQVLVWLQJ RQ D VLQJOH OD\HU RI EURZQ WKLFNZDOOHG FHOOV Teratosphaeriaceae ,QWHUDVFDO ¿ODPHQWV pseudoparaphysoids, ¿ODPHQWRXV branched, anastomosed, hyaline, indistinct, constricted at Camarosporula persooniae +HQQ 3HWUSydowia 8: VHSWD ± P Asci VHVVLOH VWLSLWDWH RERYRLG VSRUHG ± × ± P ZLWK ZHOO GHYHORSHG DSLFDO FKDPEHU )LJ ± P GLDP DQG PXOWLOD\HUHG HQGRWXQLFD Ascospores Basionym: Hendersonia persooniae+HQQHedwigia 40 fusoid-ellipsoid, hyaline, smooth, thick-walled, prominently JXWWXODWHZLGHVWMXVWDERYHWKHPHGLDQVHSWXPSURPLQHQWO\ Synonym: Dichomera persooniae +HQQ +HQQ Hedwigia FRQVWULFWHGDWVHSWXP ± ± ± × ± ± ± P 42 EHFRPLQJSDOHEURZQLQROGHUDVFL Conidiomata acervular, amphigenous, subcuticular, Teleomorph: Anthracostroma persooniae +HQQ 3HWU HSLGHUPDOWRVXEHSLGHUPDOVHSDUDWHRUFRQÀXHQWWRPP Sydowia 8 diam, composed of dark brown, thick-walled, globose to Basionym: Mycosphaerella persooniae+HQQHedwigia 42: angular cells, dehiscing irregularly at the apex, visible as small EODFNVSRWVRQVXSHU¿FLDOVWURPDWDConidiophores reduced Synonyms: Sphaerella persooniae +HQQ 6DFF '6DFF WR FRQLGLRJHQRXV FHOOV Conidiogenous cells determinate, 60 IMA FUNGUS Additions to the Mycosphaerella complex integrated, cylindrical to doliiform, hyaline to pale brown, RI EURZQ WH[WXUD DQJXODULV ,QWHUDVFDO ¿ODPHQWV DEVHQW ARTICLE VPRRWKWR¿QHO\YHUUXFXORVHOLQLQJWKHFRQLGLRPDWDOFDYLW\ Asci IDVFLFXODWH ELWXQLFDWH ZLWK ¿VVLWXQLFDWH GHKLVFHQFH ±×±P PLQXWHSHULFOLQDOWKLFNHQLQJYLVLEOHDWDSH[ subsessile, fusoid-ellipsoid, straight to slightly curved, ZKLFKLQVRPHFDVHVDSSHDUVWRDOVRSUROLIHUDWHSHUFXUUHQWO\ VSRUHG±×±PZLWKDVPDOODSLFDODSLFXOXV Conidia GLFW\RVHSWDWH GLVWRVHSWDWH ZLWK WUDQVYHUVH Ascospores multi-seriate, hyaline, thick-walled, with angular VHSWDDQG±YHUWLFDORUREOLTXHVHSWDWKLFNZDOOHG¿QHO\ cellular inclusions, fusoid-ellipsoid, medianly 1-septate, verruculose, irregular or clavate to obclavate, with a broad becoming constricted at the septum, widest in middle of WUXQFDWHEDVH ± ± ± × ± ± ± P DSLFDO FHOO ZLWK URXQGHG HQGV JUDQXODU ± ± ± ×± ± P DVFRVSRUHJHUPLQDWLRQLUUHJXODUIURPERWK Culture characteristics &RORQLHVDIWHUZNDW &VSUHDGLQJ HQGVRUPLGGOHRIWKHFHOOSDUDOOHORUDWDQJOHWRWKHORQJD[LV erumpent with folded surface, and smooth, lobate margins, VSRUH EHFRPLQJ EURZQ YHUUXFXORVH ± P ZLGH EXW QRW UHDFKLQJPPGLDP2Q0($VXUIDFHXPEHUWRROLYDFHRXV VZHOOLQJDQGGLVWRUWLQJDIWHUKRQ0($ JUH\ UHYHUVH LURQJUH\ 2Q 2$ VXUIDFH ROLYDFHRXV UH\ 2Q 3'$VXUIDFHDQGUHYHUVHROLYDFHRXVJUH\ Culture characteristics &RORQLHV DIWHU G DW & spreading, with sparse aerial mycelium and even, smooth, Specimens examined: Australia: Western Australia: 3HUWK RQ OREDWH PDUJLQV UHDFKLQJ PP GLDP 2Q 0($ VXUIDFH Persoonia elliptica Pritzel 104a %±KRORW\SHRIHendersonia GDUN PRXVHJUH\ UHYHUVH JUHHQLVK EODFN 2Q 2$ VXUIDFH persooniae New South Wales %RXGKL 1DWLRQDO 3DUN FRDVWDO ROLYDFHRXV JUH\ 2Q 3'$ VXUIDFH ROLYDFHRXV JUH\ UHYHUVH understory, on leaves of Persoonia levis6HSW C. Liddell LURQJUH\ . 0 H[,0, WHOHRPRUSKDQGDQDPRUSKSUHVHQW .XULQJJDL &KDVH 1DWLRQDO 3DUN RQ OHDYHV RI Persoonia VS Specimens examined: South Africa: Western Cape Province: on 1RY P.W. Crous & B.A. Summerell HSLW\SH &%6 + upper surface of dead leaves of Agapanthus umbellatus Alliaceae ±epitypus hic designatus of Hendersonia persooniae FXOWXUHVH[ Kalchbrenner 1342 . 0 ± KRORW\SH .LUVWHQERVFK %RWDQLFDO HSLW\SH&3& &%6&3& &%6 *DUGHQDWHQWUDQFHWRORZHUJDWHRQOHDIVSRWVRIA. umbellatus 0D\P.W. Crous &%6+±epitypus hic designatus Notes: The teleomorph genus Anthracostroma is based on a FXOWXUHVH[HSLW\SH&3&± &%6 ±Portugal: species initially described in Mycosphaerella as M. persooniae, Braga, Ria do Souto, on dead leaf of A. umbellatus,-XQHP.W. which is in accordance to its MycosphaerellaOLNHPRUSKRORJ\ Crous &%6+ FXOWXUH&3&± &%6 :LWKLQWKHTeratosphaeriaceae this genus is distinct in that the ascomata are situated in a subcuticular stroma, and the Notes: Ascospores in the holotype specimen were similar DVFLDUHLQWHUPLQJOHGDPRQJQXPHURXVSVHXGRSDUDSK\VHV ± × ± P WR WKRVH REVHUYHG LQ WKH SUHVHQW The Camarosporula anamorph is reminiscent of Dichomera, FROOHFWLRQV $SWURRW UHJDUGHG WKLV DV D VSHFLHV RI which is again allied to Botryosphaeriales %DUEHUet al Davidiella, but probably only observed immature asci, Crous et al. D Camarosporula is thus distinct within given that his observations referred to somewhat smaller Teratosphaeriaceae %RWK WKH SK\ORJHQHWLF DQDO\VLV RI WKH DVFRVSRUHV ±×±P )XUWKHUPRUHKHDOVRUHIHUV /68VHTXHQFHV )LJ DQGWKHPHJDEODVWVHDUFKHVRIWKH to other African collections, which again match the ascospore ,76VHTXHQFHVSODFHPhaeothecoidea melaleuca as closest GLPHQVLRQVRQWKHW\SH7KHVHREVHUYDWLRQVWRJHWKHUZLWKLWV sister species to Camarosporula persooniae. occurrence on AgapanthusLQ3RUWXJDOVXJJHVWWKLVVSHFLHV WR SUREDEO\ EH KRVWVSHFL¿F Mycosphaerella agapanthi- Teratosphaeria agapanthi .DOFKEU &RRNH &URXV umbellati, described from the same host in India, may be a comb. nov. ODWHUV\QRQ\PZLWKDVFL±×PDQGDVFRVSRUHV± 0\FR%DQN0% ×±P &RUOHWW %RWK/68DQG,76GDWDSODFHT. )LJ agapanthi in Teratosphaeria )LJ ZLWK,76VHTXHQFHVRIT. Basionym: Sphaerella agapanthi .DOFKEU &RRNHGrevillea considenianae *HQ%DQN*4 T. miniata *HQ%DQN 9 *4 DQGT. stellenboschiana *HQ%DQN*4 Synonym: Mycosphaerella agapanthi .DOFKEU &RRNH KDYLQJWKHKLJKHVWLGHQWLW\ LQDPHJDEODVWVHDUFKRI /LQGDXin(QJOHU 3UDQWO1DWUOLFKHQ3ÀDQ]HQI 1 WKH*HQ%DQNQXFOHRWLGHGDWDEDVH In vivo: Leaf spots amphigenous, ellipsoid, large, developing DISCUSSION where plants are grown in wet, shady areas, pale to medium brown, with a red-brown border, coalescing, becoming visible The present study resolves the phylogenetic position of DVOHDIWLSEOLJKWV\PSWRPVAscomata amphigenous, black, VHYHUDOJHQHUDRIZKLFKWKHFODVVL¿FDWLRQKDVEHHQWKHWRSLF substomatal, erumpent, predominantly arranged in tight, RISDVWVSHFXODWLRQ7KHJHQXVCamarosporula WHOHRPRUSK URXQGFOXVWHUV±PPGLDP DVFRPDWD±PGLDP Anthracostroma LVVKRZQWREHORQJWRWKHTeratosphaeriaceae, ZLWKFHQWUDOSHULSK\VDWHRVWLROH±PGLDP SHULSK\VHV which is not totally surprising given the fact that its teleomorph ±VHSWDWH±×±P DVFRPDWDOZDOORI±OD\HUV was originally described as Mycosphaerella persooniae VOLUME 2 · NO. 1 61 Crous et al. ARTICLE Fig. 10. Teratosphaeria agapanthi &3& A, B./HDIVSRWVDQGEOLJKWRQAgapanthusC.&LUFXODUDJJUHJDWLRQRIDVFRPDWDD. Evenly GLVWULEXWHGDVFRPDWDZLWKRVWLRODUUHJLRQVYLVLEOHE, G–J.$VFLF.2VWLROHZLWKSHULSK\VHV DUURZ K.$VFRVSRUHVL.*HUPLQDWLQJDVFRVSRUHV %DUV P +RZHYHUEDVHGRQLWVXQLTXHDVFRVWURPDWDZLWKHUXPSHQW Furthermore, we managed to recollect a species associated ascomata, and acervular conidiomata with muriformly with leaf spots of Agapanthus in South Africa, M. agapanthi, septate, brown conidia, Camarosporula appears to represent ZKLFK $SWURRW VXVSHFWHG WR UHSUHVHQW D SRVVLEOH a distinct genus within this family, which is also supported species of Davidiella$VFDQEHVHHQIURPLWVSK\ORJHQHWLF E\LWV'1$SK\ORJHQ\ )LJ 7KHSK\ORJHQHWLFSRVLWLRQRI SRVLWLRQ )LJ KRZHYHULWLVDVSHFLHVRITeratosphaeria, Sphaerulina, and potential taxonomic implications thereof which appears to be closely associated with its host, has KDVEHHQGLVFXVVHGLQGHWDLOHOVHZKHUH &URXVet al. EHHQLQWURGXFHGLQWR(XURSH 3RUWXJDO DQGPD\DOVRRFFXU as the genus predates Mycosphaerella5HVXOWVREWDLQHGLQ LQRWKHUFRQWLQHQWVZKHUHWKLVSODQWLVFXOWLYDWHG this study, however, have shown that while Mycosphaerella- Two species of Mycosphaerella known from leaf litter OLNH WD[D ZLWK VHSWDWH DVFRVSRUHV KDYH HYROYHG PRUH FROOHFWHG LQ +DZDLL KDYH EHHQ WUHDWHG M. etlingerae than once in the Mycosphaerellaceae &URXV et al. on Etlingera elatior, and M. holualoana on Hedychium Sphaerulina s.str W\SL¿HG E\ S. myriadea, represents a coronarium7KHODWWHUVSHFLHVKDVXQWLOQRZEHHQDFFHSWHG lineage embedded in the PseudocercosporaSeptoriaFODGH as a leaf pathogen of LeucospermumLQ+DZDLL 7D\ORUet al. Although Sphaerulina may represent a genus in its own right, DQGLWVRFFXUUHQFHRQOHDIOLWWHURIDQRWKHUKRVWPD\ WKHSUHVHQWGDWDDUHVWLOOLQVXI¿FLHQWWRUHVROYHLWVSK\ORJHQ\ suggest that more stringent plant hygiene practices need within the Mycosphaerellaceae to be followed in Protea¿HOGVLQ+DZDLLDVGHDGOHDYHVRI This study also introduces two species of Cladosporium HedychiumDSSHDUWRDFWDVDQDOWHUQDWHKRVWIRUWKLVIXQJXV that have teleomorphs, C silenes which is a member of Although this study resolved several long standing the C. cladosporioides FRPSOH[ %HQVFK et al. DQG TXHVWLRQV UHODWHG WR WKH SK\ORJHQ\ RI JHQHUD VXFK DV & grevilleae ZKLFK GRHV QRW IRUP DQ DQDPRUSK LQ FXOWXUH CamarosporulaAnthracostroma and Sphaerulina, it also 62 IMA FUNGUS Additions to the Mycosphaerella complex UDLVHGVRPHQHZTXHVWLRQV7KLVLVVSHFL¿FDOO\WUXHIRUWKH &RUWLQDV01&URXV3::LQJ¿HOG%':LQJ¿HOG0- 0XOWL ARTICLE species described here as M. cerastiicola, collected on gene phylogenies and phenotypic characters distinguish Cerastium semidecandrum LQ 7KH 1HWKHUODQGV :KLOH WKH two species within the Colletogloeopsis zuluensis complex teleomorph is rather odd in the sense that it can have less than associated with EucalyptusVWHPFDQNHUVStudies in Mycology eight ascospores, the anamorph is very peculiar in having a 55 ± Septoria- or PseudocercosporellaOLNHPRUSKRORJ\ S\FQLGLD &URXV 3: Mycosphaerella VSS DQG WKHLU DQDPRUSKV to sporodochia, and sympodial to phialidic proliferation, with associated with leaf spot diseases of Eucalyptus Mycologia FRQLGLD DOVR IRUPHG LQGLYLGXDOO\ RQ DHULDO P\FHOLXP 7KH Memoirs 21 ± oddity of this species lies in the fact that it clusters in the middle &URXV 3: Taxonomy and Pathology of Cylindrocladium of the Ramularia clade, which has hitherto been accepted (Calonectria) and allied genera.6W3DXO01 $363UHVV DVPRQRSK\OHWLF &URXVet al. DE :KHWKHURamularia &URXV 3: 7D[RQRP\ DQG SK\ORJHQ\ RI WKH JHQXV would eventually be reavealed as paraphyletic, or if this MycosphaerellaDQGLWVDQDPRUSKVFungal Diversity 38 ± Septoria-like anamorph in fact represents a synanamorph &URXV3:$SWURRW$.DQJ-&%UDXQ8:LQJ¿HOG0- 7KH of a Ramularia species, can only be resolved once more genus MycosphaerellaDQGLWVDQDPRUSKVStudies in Mycology Mycosphaerella species with Ramularia-like anamorphs 45 ± KDYHEHHQFROOHFWHGDQGVXEMHFWHGWR'1$DQDO\VLV &URXV3:%UDXQ8 MycosphaerellaDQGLWVDQDPRUSKV 1DPHVSXEOLVKHGLQCercospora and PassaloraCBS Biodiversity Series 1 ± ACKNOWLEDGEMENTS &URXV 3: %UDXQ 8 *URHQHZDOG -= Mycosphaerella is SRO\SK\OHWLFStudies in Mycology 58 ± :H WKDQN WKH WHFKQLFDO VWDII $ULHQ YDQ ,SHUHQ FXOWXUHV 0DUMDQ &URXV 3: *DPV : 6WDOSHUV -$ 5REHUW 96WHJHKXLV * D 9HUPDDV SKRWRJUDSKLF SODWHV DQG 0LHNH 6WDULQN:LOOHPVH 0\FR%DQN DQRQOLQHLQLWLDWLYHWRODXQFKP\FRORJ\LQWRWKHVW '1$ LVRODWLRQ DPSOL¿FDWLRQ DQG VHTXHQFLQJ IRU WKHLU LQYDOXDEOH FHQWXU\Studies in Mycology 50 ± DVVLVWDQFH :DOWHU *DPV DQG $QGUp $SWURRW ERWK IRUPHUO\ &URXV 3: *URHQHZDOG -= +RVWV VSHFLHV DQG JHQRW\SHV &%6 DUH WKDQNHG IRU VHYHUDO RI WKH FROOHFWLRQV WUHDWHG KHUH 7KH RSLQLRQVYHUVXVGDWDAustralasian Plant Pathology 34 ± FXUDWRUVRI% %RWDQLVFKH*DUWHQXQG%RWDQLVFKHV0XVHXP%HUOLQ &URXV3:*URHQHZDOG-=0DQVLOOD-3+XQWHU*&:LQJ¿HOG0- 'DKOHP*HUPDQ\ %3, 86'$%HOWVYLOOH86$ DQG.HZ 8. DUH E 3K\ORJHQHWLFUHDVVHVVPHQWRIMycosphaerellaVSSDQG acknowledged for making several valuable specimens available for their anamorphs occurring on Eucalyptus. Studies in Mycology VWXG\ 50 ± &URXV 3: *URHQHZDOG -= 3RQJSDQLFK . +LPDPDQ :$U]DQORX 0:LQJ¿HOG0- F &U\SWLFVSHFLDWLRQDQGKRVWVSHFL¿FLW\ REFERENCES among Mycosphaerella VSS RFFXUULQJ RQ $XVWUDOLDQ Acacia VSHFLHVJURZQDVH[RWLFVLQWKHWURSLFVStudies in Mycology 50: $SWURRW$ MycosphaerellaDQGLWVDQDPRUSKV &RQVSHFWXV ± of MycosphaerellaCBS Biodiversity Series 5 ± &URXV 3: *URHQHZDOG -= :LQJ¿HOG 0- $SWURRW $ 7KH %DUEHU3$%XUJHVV7+DUG\*6W-6OLSSHUV%.HDQH3-:LQJ¿HOG value of ascospore septation in separating Mycosphaerella from 0- Botryosphaeria species from Eucalyptus in Australia Sphaerulina in the Dothideales: a Saccardoan myth? Sydowia are pleoanamorphic, producing Dichomera synanamorphs in 55 ± FXOWXUHMycological Research 109 ± &URXV 3: .DQJ -& %UDXQ 8 $ SK\ORJHQHWLF UHGH¿QLWLRQ %DW]HU -& 0HUFHGHV 'LD] $ULDV 0 +DUULQJWRQ 7& *OHDVRQ 0/ of anamorph genera in Mycosphaerella EDVHG RQ ,76 U'1$ *URHQHZDOG -= &URXV 3: )RXU VSHFLHV RI Zygophiala VHTXHQFHDQGPRUSKRORJ\Mycologia 93 ± Schizothyriaceae, Capnodiales DUHDVVRFLDWHG ZLWK WKHVRRW\ &URXV 3: 6FKRFK &/ +\GH .' :RRG $5 *XHLGDQ & +RRJ EORWFKDQGÀ\VSHFNFRPSOH[RQDSSOHMycologia 100 ± *6 GH *URHQHZDOG -= D 3K\ORJHQHWLF OLQHDJHV LQ WKH %HQVFK . *URHQHZDOG -= 'LMNVWHUKXLV - 6WDULQN:LOOHPVH 0 CapnodialesStudies in Mycology 64 ± Andersen B, Summerell BA, Shin H-D, Dugan FM, Schroers H-J, &URXV 3: 6OLSSHUV % :LQJ¿HOG 0- 5KHHGHU - 0DUDVDV :)2 %UDXQ8&URXV3: 6SHFLHVDQGHFRORJLFDOGLYHUVLW\ZLWKLQ 3KLOLSV $-/ $OYHV $ %XUJHVV 7 %DUEHU 3 *URHQHZDOG -= the Cladosporium cladosporioides FRPSOH[ Davidiellaceae, D 3K\ORJHQHWLF OLQHDJHV LQ WKH Botryosphaeriaceae Capnodiales Studies in Mycology 67 ± Studies in Mycology 55 ± %UDXQ8&URXV3:'XJDQ)*URHQHZDOG-*+RRJ6*GH &URXV 3: 6XPPHUHOO %$ &DUQHJLH $- :LQJ¿HOG 0- +XQWHU 3K\ORJHQ\ DQG WD[RQRP\ RI Cladosporium-like hyphomycetes, *& %XUJHVV7,$QGMLF 9%DUEHU 3$*URHQHZDOG -= E including DavidiellaJHQQRYWKHWHOHRPRUSKRICladosporium Unravelling Mycosphaerella: do you believe in genera? VVWUMycological Progress 2 ± Persoonia 23 ± &DUERQH,.RKQ/0 $PHWKRGIRUGHVLJQLQJSULPHUVHWVIRU &URXV3:9HUNOH\*-0*URHQHZDOG-=6DPVRQ5$ HGV F VSHFLDWLRQ VWXGLHV LQ ¿ODPHQWRXV DVFRP\FHWHV Mycologia 91: Fungal Biodiversity >CBS Laboratory Manual Series QR @ ± &HQWUDDOEXUHDXYRRU6FKLPPHOFXOWXUHV8WUHFKW &RUOHWW 0 $Q DQQRWDWHG OLVW RI WKH SXEOLVKHG QDPHV LQ &URXV3::LQJ¿HOG0-0DQVLOOD-3$OIHQDV$&*URHQHZDOG-= Mycosphaerella and Sphaerella, Mycologia Memoir 18 ± E 3K\ORJHQHWLFUHDVVHVVPHQWRIMycosphaerellaVSSDQG VOLUME 2 · NO. 1 63 Crous et al. their anamorphs occurring on Eucalyptus,,Studies in Mycology 6FKXEHUW.*URHQHZDOG-=%UDXQ8'LMNVWHUKXLV-6WDULQN0+LOO 55 ± &)=DODU3+RRJ*6GH&URXV3: %LRGLYHUVLW\LQWKH &URXV3::LQJ¿HOG0-3DUN5) Mycosphaerella nubilosa a Cladosporium herbarumFRPSOH[ Davidiellaceae, Capnodiales synonym of M. mollerianaMycological Research 95 ± with standardisation of methods for Cladosporium taxonomy and )DUU')%LOOV*)&KDPXULV*35RVVPDQ$< Fungi on plants GLDJQRVWLFVStudies in Mycology 58 ± ARTICLE and plant products in the United States6W3DXO01 $363UHVV 6RDUHV '- %DUUHWR 5: )XQJDO SDWKRJHQV RI WKH LQYDVLYH +LOOLV '0 %XOO -- $Q HPSLULFDO WHVW RI ERRWVWUDSSLQJ DVD riparian weed Hedychium coronarium IURP %UD]LO DQG WKHLU PHWKRG IRU DVVHVVLQJ FRQ¿GHQFH LQ SK\ORJHQHWLF DQDO\VLV SRWHQWLDOIRUELRORJLFDOFRQWUROFungal Diversity 28 ± Systematic Biology 42 ± 6WHYHQV )/ +DZDLLDQ IXQJL Bernice P. Bishop Museum +RRJ*6GH*HUULWVYDQGHQ(QGH$+* 0ROHFXODUGLDJQRVWLFV Bulletin 19 ± RI FOLQLFDO VWUDLQV RI ¿ODPHQWRXV Basidiomycetes Mycoses 41: 6ZRIIRUG'/ PAUP*: phylogenetic analysis using parsimony ± (*and their methods). 9HUVLRQ 6XQGHUODQG 0$ 6LQDXHU 2¶'RQQHOO . .LVWOHU +& &LJHOQLN ( 3ORHW] 5& 0XOWLSOH $VVRFLDWHV HYROXWLRQDU\ RULJLQV RI WKH IXQJXV FDXVLQJ 3DQDPD GLVHDVH RI 7D\ORU -( &URXV 3: 3DOP 0( )ROLDU DQG VWHP IXQJDO banana: concordant evidence from nuclear and mitochondrial pathogens of ProteaceaeLQ+DZDLLMycotaxon 78 ± JHQH JHQHDORJLHV Proceedings of the National Academy of 9HUNOH\ *-0 &URXV 3: *URHQHZDOG -= %UDXQ 8 $SWURRW $ Sciences, USA 95 ± Mycosphaerella punctiformis revisited: morphology, 3DJH5'0 75((9,(: $QDSSOLFDWLRQWRGLVSOD\SK\ORJHQHWLF SK\ORJHQ\DQGHSLW\SL¿FDWLRQRIWKHW\SHVSHFLHVRIWKHJHQXV WUHHV RQ SHUVRQDO FRPSXWHUV Computer Applications in the Mycosphaerella Dothideales, Ascomycota Mycological Biosciences 12 ± Research 108 ± 3UHWRULXV0&&URXV3:*URHQHZDOG-=%UDXQ8 3K\ORJHQ\ 9LOJDO\V 5 +HVWHU 0 5DSLG JHQHWLF LGHQWL¿FDWLRQ DQG of some cercosporoid fungi from CitrusSydowia 55 ± PDSSLQJRIHQ]\PDWLFDOO\DPSOL¿HGULERVRPDO'1$IURPVHYHUDO 5DPEDXW$ Sequence Alignment Editor9HUVLRQ2[IRUG CryptococcusVSHFLHVJournal of Bacteriology 172 ± 'HSDUWPHQWRI=RRORJ\8QLYHUVLW\RI2[IRUG :KLWH7- %UXQV7/HH -7D\ORU- $PSOL¿FDWLRQ DQG GLUHFW 5D\QHU5: A Mycological Colour Chart.HZ &RPPRQZHDOWK VHTXHQFLQJ RI IXQJDO ULERVRPDO 51$ JHQHV IRU SK\ORJHQHWLFV 0\FRORJLFDO,QVWLWXWH In ,QQLV 0$ *HOIDQG '+ 6QLQVN\ -- :KLWH 7- HGV PCR 6FKRFK&/6KRHPDNHU5$6HLIHUW.$+DPEOHWRQ66SDWDIRUD-: Protocols: a guide to methods and applications ±6DQ &URXV3: $PXOWLJHQHSK\ORJHQ\RIWKHDothideomycetes 'LHJR&$ $FDGHPLF3UHVV XVLQJIRXUQXFOHDUORFLMycologia 98 ± 64 IMA FUNGUS