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AR TICLE Advances in Glomeromycota Taxonomy And GRLLPDIXQJXV IMA FUNGUS · VOLUME 2 · NO 2: 191–199 Advances in GlomeromycotaWD[RQRP\DQGFODVVL¿FDWLRQ ARTICLE )ULW]2HKO(ZDOG6LHYHUGLQJ-DYLHU3DOHQ]XHOD.XUW,QHLFKHQDQG*ODGVWRQH$OYHVGD6LOYD $JURVFRSH5HFNHQKRO]7lQLNRQ5HVHDUFK6WDWLRQ $57 (FRORJLFDO)DUPLQJ6\VWHPV5HFNHQKRO]VWUDVVH&+=ULFK6ZLW]HUODQG FRUUHVSRQGLQJDXWKRUHPDLOIULW]RHKO#DUWDGPLQFK ,QVWLWXWHRI3ODQW3URGXFWLRQDQG$JURHFRORJ\LQWKH7URSLFVDQG6XEWURSLFV8QLYHUVLW\RI+RKHQKHLP*DUEHQVWUDVVH'6WXWWJDUW *HUPDQ\ 'HSDUWDPHQWR GH 0LFURELRORJtD GHO 6XHOR \ 6LVWHPDV 6LPELyWLFRV (VWDFLyQ ([SHULPHQWDO GHO =DLGtQ &6,& 3URIHVRU$OEDUHGD *UDQDGD6SDLQ %DVHO=ULFK3ODQW6FLHQFH&HQWHU,QVWLWXWHRI%RWDQ\8QLYHUVLW\RI%DVHO+HEHOVWUDVVH&+%DVHO6ZLW]HUODQG 'HSDUWDPHQWR GH 0LFRORJLD &&% 8QLYHUVLGDGH )HGHUDO GH 3HUQDPEXFR$Y 3URI 1HOVRQ &KDYHV VQ &LGDGH 8QLYHUVLWDULD 5HFLIH3(%UD]LO Abstract: &RQFRPLWDQWPRUSKRORJLFDODQGPROHFXODUDQDO\VHVKDYHOHGWRPDMRUEUHDNWKURXJKVLQWKHWD[RQRPLF Key words: organization of the phylum Glomeromycota)XQJLLQWKLVSK\OXPDUHNQRZQWRIRUPDUEXVFXODUP\FRUUKL]DDQGVR Archaeosporomycetes IDUWKUHHFODVVHV¿YHRUGHUVIDPLOLHVDQGJHQHUDKDYHEHHQGHVFULEHGSensu latoVSRUHIRUPDWLRQLQRI endomycorrhizas the arbuscular mycorrhiza-forming genera is exclusively glomoid, one is gigasporoid, seven are scutellosporoid, evolution IRXUDUHHQWURSKRVSRURLGWZRDUHDFDXORVSRURLGDQGRQHLVSDFLVSRURLG6SRUHELPRUSKLVPLVIRXQGLQWKUHH Gigasporales JHQHUDDQGRQHJHQXVLVDVVRFLDWHGZLWKF\DQREDFWHULD+HUHZHSUHVHQWWKHFXUUHQWFODVVL¿FDWLRQGHYHORSHGLQ Glomerales VHYHUDOUHFHQWSXEOLFDWLRQVDQGSURYLGHDVXPPDU\WRIDFLOLWDWHWKHLGHQWL¿FDWLRQRIWD[DIURPJHQXVWRFODVVOHYHO Glomeromycetes Paraglomeromycetes phylogeny 9$P\FRUUKL]D Article info:6XEPLWWHG1RYHPEHU$FFHSWHG1RYHPEHU3XEOLVKHG1RYHPEHU INTRODUCTION UDGLDOJORPRLG DQG VFXWHOORVSRURLG 'LIIHUHQFHV LQ VSRUH ZDOOVWUXFWXUHZHUHXVHGDWWKHVSHFLHVOHYHO Glomeromycota taxonomy was largely morphologically driven 7RGD\ ZH DFFHSW WKUHH FODVVHV Archaeosporomycetes, XS WR WKH HQG RI WKH ODVW PLOOHQQLXP$OO JORPHURP\FRWHDQ Glomeromycetes, and Paraglomeromycetes ¿YH RUGHUV fungi, except one genus, are known to form arbuscular Archaeosporales, Diversisporales, Gigasporales, Glomerales P\FRUUKL]D 7KHLU LGHQWL¿FDWLRQ ZDV EDVHG RQ VSRUH and Paraglomerales IDPLOLHVJHQHUDDQGDSSUR[LPDWHO\ morphology, spore formation, and spore wall structure VSHFLHV HJ 0RUWRQ 5HGHFNHU 6FKler et al HJ*HUGHPDQQ 7UDSSH:DONHU 6DQGHUV 2HKO 6LHYHUGLQJ :DONHU 6FKOHU 0RUWRQ %HQQ\ 6FKHQFN 3pUH] +RZHYHU 6LHYHUGLQJ 2HKO6SDLQet al2HKOet al as soon as molecular phylogenetic tools became available, D±G3DOHQ]XHODet al WKH\ZHUHLQFOXGHGLQWD[RQRPLFDQDO\VHV HJ6LPRQet al Until recently, it was unclear whether glomoid and DQG VRRQ EHFDPH WKH GULYHUV RI WKH HVWDEOLVKPHQW gigasporoid species could be further divided into different RIDQHZWD[RQRP\ 0RUWRQ 5HGHFNHU6FKler et morphological groups congruent with the major phylogenetic al ,QZLWKRXWWKHEHQH¿WRIPROHFXODUDVSHFWV FODGHVREWDLQHGE\PROHFXODUDQDO\VHV$¿UVWUHYLVLRQRIWKH the arbuscular mycorrhiza-forming fungi were organized VSRURJHQRXVFHOOIRUPLQJ JLJDVSRURLGDQGVFXWHOORVSRURLG LQ WKUHH IDPLOLHV Acaulosporaceae, Gigasporaceae, and Glomeromycetes according to concomitant morphological Glomeraceae DQGVL[JHQHUD Acaulospora, Entrophospora, DQGSK\ORJHQHWLFIHDWXUHV 2HKOHWal ZDVQRWDFFHSWHG Gigaspora, Glomus, Sclerocystis, and Scutellospora E\ DOO P\FRORJLVWV 0RUWRQ 0VLVND +RZHYHU ODWHU within one order, Glomerales 0RUWRQ %HQQ\ RIWKH VWXGLHVZLWKDEURDGHUGDWDEDVH HJ*RWRet al fungal phylum Zygomycota 7KDW FODVVL¿FDWLRQ ZDV EDVHG 2HKO et al E FRQ¿UPHG WKDW WKH UHYLVHG JHQXV on spore morphology and spore formation characteristics Scutellospora, as well as the new Racocetra, Cetraspora, DFDXORVSRURLG HQWURSKRVSRURLG JLJDVSRURLG JORPRLG Dentiscutata, and Orbispora,DUHPRQRSK\OHWLF © 2011 International Mycological Association You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: <RXPXVWDWWULEXWHWKHZRUNLQWKHPDQQHUVSHFL¿HGE\WKHDXWKRURUOLFHQVRU EXWQRWLQDQ\ZD\WKDWVXJJHVWVWKDWWKH\HQGRUVH\RXRU\RXUXVHRIWKHZRUN Non-commercial: <RXPD\QRWXVHWKLVZRUNIRUFRPPHUFLDOSXUSRVHV No derivative works: <RXPD\QRWDOWHUWUDQVIRUPRUEXLOGXSRQWKLVZRUN 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. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. VOLUME 2 · NO. 2 191 Oehl et al. $ ODUJH JURXS RI VSHFLHV IRUPV JORPRLG VSRUHV DQG LW VWDLQ EOXH WR GDUN EOXH LQ WU\SDQ EOXH ,Q Glomeromycetes, had been believed that there were too few morphological Gigasporales species do not form intraradical vesicles but FKDUDFWHUVRIVLJQL¿FDQFHWRGLIIHUHQWLDWHWKHP7D[RQRPLVWV auxiliary cells in soils, which clearly distinguish them from have consequently started basing groupings of the glomoid Glomerales and Diversisporales species almost exclusively on molecular phylogenetic Gigasporales exhibit gigasporoid or scutellosporoid spore ARTICLE FKDUDFWHUV $ UHFHQW UHYLVLRQ RI WKHVH JORPRLG VSHFLHV IRUPDWLRQ 2HKOet al.E LHVSRUHVIRUPHGWHUPLQDOO\ has, however, shown that molecular phylogeny is actually on sporogenous cells and with either germ warts on the congruent with the morphological characteristics of these LQQHU VXUIDFH RI WKH PRQRZDOOHG VSRUH ZDOO JLJDVSRURLG IXQJL 2HKOet al.F )XQJDOVSHFLHVZLWKHQWURSKRVSRURLG Gigasporaceae RU D GLVFUHWH JHUPLQDWLRQ VKLHOG RQ WKH VSRUH IRUPDWLRQ ZHUH DOVR UHYLVHG 2HKO et al. G LQQHUPRVW µJHUPLQDOZDOO¶ RI±ZDOOV VFXWHOORVSRURLG 7KHREMHFWLYHRIWKLVSDSHULVWRSUHVHQWWKHFXUUHQWRYHUDOO 7KHUHDUHWKUHHIDPLOLHVZLWKVFXWHOORVSRURLGVSRUHIRUPDWLRQ FODVVL¿FDWLRQ V\VWHP RI Glomeromycota that has emerged sensu lato Dentiscutataceae, Racocetraceae and from these recent studies, and to summarize the major Scutellosporaceae 2HKOet al Scutellosporaceae form PRUSKRORJLFDOIHDWXUHVLQWKHSK\OXPGRZQWRJHQXVOHYHO PRQROREHG Orbispora RUELOREHG Scutellospora K\DOLQH JHUPLQDWLRQVKLHOGV )LJV± Racocetraceae species form wavy-like, multiply lobed, hyaline germination shields and MATERIALS AND METHODS KDYHHLWKHUWZR Racocetra RUWKUHH Cetraspora VSRUHZDOOV )LJV ± Dentiscutataceae species form yellow-brown to 7KH PRUSKRORJLFDO PROHFXODU DQG SK\ORJHQHWLF DQDO\VHV EURZQJHUPVKLHOGVWKDWDUHELOREHG Fuscutata)LJ RU performed are presented in a series of recent publications ZLWK PXOWLSOH FRPSDUWPHQWV Dentiscutata, triple-walled; dealing with different species groups of Glomeromycota HJ QuatunicaIRXUZDOOHG)LJV± 2HKO et al D E G I 6LHYHUGLQJ 2HKO In Archaeosporales and Diversisporales, four genera 6LOYDet al.6SDLQet al.3DOHQ]XHODet al. have spore formation laterally on the neck of terminal or LQWHUFDODU\ VSRULIHURXV VDFFXOHV DFDXORVSRURLG sensu lato 7DEOH Acaulospora, Otospora, and the bi-morphic Ambispora and Archaeospora7KHVHJHQHUDFDQHDVLO\EH RESULTS AND DISCUSSION separated on spore wall number and spore wall structure 3DOHQ]XHOD et al. 7ULSOHZDOOHG Acaulospora )LJXUH LV D VFKHPDWLF WUHH IRU Glomeromycota based on VSHFLHV KDYH D FKDUDFWHULVWLF JUDQXODU µEHDGHG¶ LQQHU ZDOO PROHFXODUSK\ORJHQHWLFDQDO\VHVRIWKH668,76UHJLRQSDUWLDO VXUIDFH 0RUWRQ %HQQ\ ZKLFKLVDEVHQWLQDFDXOR /68RIWKHU51$JHQHDQGSDUWLDOȕWXEXOLQJHQH HJ2HKOet ambisporoid spores of triple-walled Ambispora species alD±G ,Q7DEOHWKHPDMRUPRUSKRORJLFDO 6SDLQet al3DOHQ]XHODet al 7KHZDOOVWUXFWXUH features of all higher level taxa are presented, with the taxa of the bi-walled Otospora is more complex than that of bi- DUUDQJHG DFFRUGLQJ WR WKHLU WD[RQRPLF UDQN GRZQ WR JHQXV walled ArchaeosporaVSHFLHV 3DOHQ]XHODet al 7KUHHJORPHURP\FRWHDQFODVVHV¿YHRUGHUVIDPLOLHVDQG In Archaeosporales, Diversisporales, and Glomerales, JHQHUDKDYHEHHQUHFRJQL]HGWRGDWH 7DEOH Sensu lato, WKHUH DUH ¿YH JHQHUD ZLWK VSRUH IRUPDWLRQ ZLWKLQ WKH VSRUH IRUPDWLRQ LQ RI WKH DUEXVFXODU P\FRUUKL]DIRUPLQJ QHFN RI WHUPLQDO RU LQWHUFDODU\ VSRULIHURXV VDFFXOHV LH genera have exclusively glomoid, one has gigasporoid, seven entrophosporoid sensu lato 7DEOH Entrophospora, have scutellosporoid, four have entrophosporoid, two genera Kuklospora, Sacculospora, Tricispora, and bimorphic have acaulosporoid, and one has pacisporoid spore formation, Intraspora 2HKOet al.G 7ULSOHZDOOHGKuklospora has while three genera show spore bimorphism, and one genus WKH FKDUDFWHULVWLF JUDQXODU µEHDGHG¶ LQQHU ZDOO VXUIDFH RI LV DVVRFLDWHG ZLWK F\DQREDFWHULD WKH RQO\ RQH QRW IRUPLQJ Acaulosporaceae 6LHYHUGLQJ 2HKO ZKLFKLVDEVHQW DUEXVFXODUP\FRUUKL]DV in spores of triple-walled Sacculospora 2HKOet alG Hitherto, Paraglomeromycetes DUH PRQRJHQHULF 7DEOH 7KHZDOOVWUXFWXUHRIELZDOOHGEntrophospora and Tricispora DUH FKDUDFWHUL]HG E\ PRQRZDOOHG VSRUHV IRUPHG is more complex than that of bi-walled, bimorphic Intraspora WHUPLQDOO\ RQ K\SKDH LH JORPRLG VSRUHV sensu lato DQG VSHFLHV 6LHYHUGLQJ 2HKO 2HKO et al G JHUPLQDWH GLUHFWO\ WKURXJK WKH VSRUH ZDOO7KHLU DUEXVFXODU Entrophospora and Tricispora can be distinguished through mycorrhizal structures do not or only faintly stain in trypan WKHWZRFLFDWULFHV VFDUV DQGSRUHVWUXFWXUHVSUR[LPDODQG EOXH Archaeosporomycetes includes organisms that are distal to the sporiferous saccule: the proximal pore is wide exclusively bimorphic since they form either acaulosporoid or in Tricispora and closed by a septum, while it is narrow and entrophosporoid spores simultaneously with glomoid spores, closed by a plug in Entrophospora7KHGLVWDOSRUHDQGVFDU RU DUH DVVRFLDWHG ZLWK F\DQREDFWHULD 7KH P\FRUUKL]DO is absent in Entrophospora from the structural layer, and
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