ARTICLE 191
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ler al OHU 6WXWWJDUW et Cetraspora , Glomerales , al ¿YH RUGHUV et $P\FRUUKL]D Key words: Key words: Archaeosporomycetes endomycorrhizas evolution Gigasporales Glomerales Glomeromycetes Paraglomeromycetes phylogeny 9 Archaeosporomycetes Racocetra Gigasporales al2HKO al ZDVQRWDFFHSWHG , et DUHPRQRSK\OHWLF DQGVR Paraglomeromycetes al IDPLOLHV JHQHUDDQGDSSUR[LPDWHO\ 6SDLQ according to concomitant morphological Orbispora, , and IMA FUNGUS · VOLUME 2 · NO 2: 191–199 · NO 2 · VOLUME FUNGUS IMA 6LHYHUGLQJ :DONHU 6FK , as well as the new al E FRQ¿UPHG WKDW WKH UHYLVHG JHQXV latoVSRUHIRUPDWLRQLQRI et Paraglomerales 7RGD\ ZH DFFHSW WKUHH FODVVHV Until recently, it was unclear whether glomoid and Until recently, ivecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get Any of the above conditions can be waived ivecommons.org/licenses/by-nc-nd/3.0/legalcode. Sensu Archaeosporales, Diversisporales Glomeromycetes UDGLDOJORPRLG DQG VFXWHOORVSRURLG 'LIIHUHQFHV LQZDOOVWUXFWXUHZHUHXVHGDWWKHVSHFLHVOHYHO VSRUH
and 2HKO VSHFLHV HJ 0RUWRQ 5HGHFNHU 6FK Scutellospora Dentiscutata, and Glomeromycetes DQGSK\ORJHQHWLFIHDWXUHV 2HKOHW et D±G3DOHQ]XHOD VSRURJHQRXVFHOOIRUPLQJ JLJDVSRURLGDQGVFXWHOORVSRURLG E\ DOO P\FRORJLVWV 0RUWRQ 0VLVND +RZHYHU ODWHU VWXGLHVZLWKDEURDGHUGDWDEDVH HJ*RWR 2HKO 6LHYHUGLQJ 2HKO gigasporoid species could be further divided into different gigasporoid species could be further divided into different morphological groups congruent with the major phylogenetic FODGHVREWDLQHGE\PROHFXODUDQDO\VHV$¿UVWUHYLVLRQRIWKH DQG*ODGVWRQH$OYHVGD6LOYD , et al and et ler .XUW,QHLFKHQ Entrophospora , Gigasporaceae, PROHFXODUDQDO\VHVKDYHOHGWRPDMRUEUHDNWKURXJKVLQWKHWD[RQRPLF
)XQJLLQWKLVSK\OXPDUHNQRZQWRIRUPDUEXVFXODUP\FRUUKL]D and Scutellospora Acaulospora 0RUWRQ %HQQ\ RIWKH -DYLHU3DOHQ]XHOD Sclerocystis, Glomeromycota , Acaulosporaceae,
Glomerales taxonomy was largely morphologically driven taxonomy was largely morphologically driven Zygomycota 7KDW FODVVL¿FDWLRQ ZDV EDVHG DQGVL[JHQHUD ZLWKRXWWKHEHQH¿WRIPROHFXODUDVSHFWV (ZDOG6LHYHUGLQJ &RQFRPLWDQWPRUSKRORJLFDODQG
$ 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 structures of Archaeosporaceae are similar to those of formed only on the overlying, hyaline, evanescent layer, Paraglomeraceae, while Ambisporaceae form vesicular- while, in light microscopy, the distal pore with a distal scar is arbuscular mycorrhizal structures staining pale blue in trypan obvious in Tricispora 6LHYHUGLQJ 2HKO3DOHQ]XHODet EOXH,QFRQWUDVWP\FRUUKL]DOVWUXFWXUHVLQGlomeromycetes al.2HKOet al.G
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VOLUME 2 · NO. 2 193 Oehl et al. ARTICLE 0\FRUUKL]DOVWUXFWXUHV VWDLQLQJLQ7U\SDQEOXH Hyphae 9$+ 9$+ 9$+ 9$+ 9$+ 9$+ 9$+ 9$+ 9$+ 9$+ 9$+ 9$+ 9$+ 9$+ 9$+ 9$+ RUE Unknown? Unknown? Unknown? *HUPWXEH JW 9HVLFOHV$UEXVFOHV gt through wall? gt through hypha gt through hypha gt through hypha gt through hypha gt through hypha gt through hyphae gt through hypha? gt through hypha? gt through hypha? JHUPVKLHOG RUE lobed germ structure *HUPLQDWLRQ VSHFL¿F germination structure gt through wall; mono- PXOWLSO\ OREHGK\DOLQH gt through wall; multiply JWWKURXJKZDOO PRQR WR lobed, hyaline germ shield
spore walls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lomeromycota Funneliformis Septoglomus Simiglomus Albahypha Viscospora Entrophospora Redeckera Otospora Tricispora Acaulospora Entrophosporaceae Claroideoglomus Sacculosporaceae Sacculospora Pacisporaceae Pacispora Acaulosporaceae Kuklospora )DPLO\ *HQXV 6SRUHIRUPDWLRQ 1XPEHURI Diversisporaceae Diversispora Glomeromycetes Table 1. 0DMRUPRUSKRORJLFDOFKDUDFWHUVIRUKLJKHUOHYHOWD[DRI Table Class 2UGHU Diversisporales Glomerales Glomeraceae Glomus
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VOLUME 2 · NO. 2 195 Oehl et al. ARTICLE
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D 7KH PRUSKRORJLFDO GLIIHUHQWLDWLRQ RI K\SKDOZDOOOD\HUEXWWKHVXEWHQGLQJK\SKDHDUHK\DOLQH )LJV the glomoid species is mainly based on the morphology of the ± ,QFRQWUDVWDiversispora and Redeckera form spores VXEWHQGLQJ K\SKDH RI WKH VSRUHV DQG VSRUH ZDOO VWUXFWXUH whose structural wall layer is not obviously continuous with 6SRUHVRIFunneliformis, Glomus, Septoglomus, and Simiglomus WKHK\SKDOZDOOOD\HU )LJV± FRQVHTXHQWO\VXFKVSRUHV species have subtending hyphae that are concolorous or slightly DSSHDUWRKDYHLQFOXGHGµHQGRVSRUHV¶ VOLUME 2 · NO. 2 197 Oehl et al. Funneliformis, Glomus, Septoglomus, and Simiglomus )85%6DQWD&DWDULQD6WDWH%UD]LO *,1&2%(/LQ/RXYDLQ can be separated by the structure of the spore base and /D1HXYH Glomeromycota,Q9LWUR&ROOHFWLRQDWWKH&DWKROLF VXEWHQGLQJ K\SKDH sh Glomus species often have an 8QLYHUVLW\ RI /RXYDLQ %HOJLXP RU 6$) LQ =XULFK 6ZLVV LQWURYHUWHG ZDOO WKLFNHQLQJ 2HKO et al D )LJV ± &ROOHFWLRQ RI $UEVXFXODU 0\FRUUKL]DO )XQJL DW $JURVFRSH which is only otherwise seen in Viscospora Funneliformis $57 6ZLW]HUODQG ZLOO IDFLOLWDWH IXUWKHU SURJUHVVHV LQ WKH ARTICLE species generally have an easily visible septum in the area taxonomy of glomeromycotean fungi that were thought to of the spore base, and their sh are regularly funnel-shaped to have not enough criteria to morphologically separate them F\OLQGULFDO )LJ Septoglomus species have constricted unequivocally into the higher level taxa they phylogenetically to cylindrical sh, and usually there is a septum at the spore EHORQJ WR &XUUHQWO\ VHYHUDO DUEXVFXODU P\FRUUKL]DO IXQJL EDVH )LJ ,QSimiglomus, sh are cylindrical and thick- are being described as new to science each year by an walled, and they have several septa some distance from the LQFUHDVLQJ QXPEHUV RI UHVHDUFK JURXSV$ VLPSOH EXW ZHOO VSRUH EDVH )LJ Claroideoglomus has funnel- to bird- MXVWL¿HGFRQFOXVLRQLVWKDWDVDUHVXOWRIIXWXUHFRQFRPLWDQW bill-shaped sh, with sh and shZDOOVWKDWDUH!WLPHVZLGHU morphological and molecular analyses, yet more higher level DW WKH VSRUH EDVH WKDQ VRPH GLVWDQFH IURP WKH EDVH )LJ taxa will be proposed in this ancient fungal phylum, at all Albahypha has slightly funnel to bill-shaped sh and sh OHYHOVIURPFODVVGRZQWRJHQXV ZDOOV WKDW DUH WLPHV ZLGHU DW WKH VSRUH EDVH WKDQ DW VRPHGLVWDQFHIURPWKHEDVH )LJ DQGViscospora has cylindrical sh )LJ ZLWKDQsh wall that may be thickened ACKNOWLEDGEMENTS over large distances and may bear septa in the hyphae ZLWKLQWURYHUWHGZDOOWKLFNHQLQJVLQWKHDUHDRIWKHVHSWXP :HDFNQRZOHGJHWKHKHOSRIPDQ\FXUUHQWDQGIRUPHUVWXGHQWVDQG In Diversispora, the sh are usually quite fragile and hyaline, WHFKQLFLDQVLQ6ZLW]HUODQG6SDLQDQG%UD]LOIRURXWVWDQGLQJVXSSRUW distal to the pore closure at the spore base or in the sh )LJ HVSHFLDOO\'DYLG6FKQHLGHU5REHUW%|VFK*LDFRPR%XVFR/RXLV Redeckera species have a broad septum at the spore /DZRXLQ 'RPLQJR $OYDUH] 'DQLHOOH 6LOYD 1DWiOLD 6RXVD DQG EDVH )LJ DQGWKHVWUXFWXUDOZDOOOD\HUGRHVQRWFRQWLQXH 'DQLHOH 0DJQD 7KLV VWXG\ KDV EHHQ VXSSRUWHG ZLWKLQ WKH 6ZLVV PRUHWKDQ±PLQWRWKHVXEWHQGLQJK\SKDDQGWKXVWKH 1DWLRQDO 6FLHQFH )RXQGDWLRQ 616) 3URMHFW B shPD\LQÀDWHDWWKLVGLVWDQFHIURPWKHVSRUHEDVH E\WKH616)3URJUDPPH1)3µ/DQGVFDSHV DQGKDELWDWVRIWKH 7KHUH DUH WKUHH ELPRUSKLF JHQHUD ZLWK JORPRLG VSRUH $OSV¶E\WKH6SDQLVK0LQLVWU\RI(QYLURQPHQW 00$2$31SURMHFW IRUPDWLRQ *ORPRDPELVSRURLG VSRUHV KDYH D VXEK\DOLQH DQG)$(&$ -XQWDGH$QGDOXFtD6SDLQ3URMHFW to ochraceous, evanescent outer wall layer continuous with DQGE\WKH)XQGDomRGH$PSDURj&LrQFLDH7HFQRORJLDGR(VWDGR the outer acaulo-ambisporoid spore wall, while the second, GH 3HUQDPEXFR )$&(3( DQG WKH 8)3( ZKLFK SURYLGHG JUDQWV structural layer is hyaline and continuous with the middle wall WR)2HKODVµYLVLWLQJSURIHVVRU¶DQG)$&(3(ZKLFKDOVRSURYLGHG RIDFDXORDPELVSRURLGVSRUHV 6SDLQet al3DOHQ]XHOD ¿QDQFLDOVXSSRUWIRU*$6LOYD et al. *ORPRDUFKDHRVSRURLGDQG*ORPRLQWUDVSRURLG spores are among the smallest within Glomeromycota ca P DQGWKXVGLI¿FXOWWRREVHUYH REFERENCES %áDV]NRZVNL-.RYiFV*0 %DOi]V7. 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D E D E DQG LW LV GLI¿FXOW WR SUHGLFW KRZ E Glomus africanum and G. iranicum, two new species of PRUSKRORJLFDO LGHQWL¿FDWLRQ ZLOO GHYHORS LQ WKRVH IXQJL DUEXVFXODUP\FRUUKL]DOIXQJL Glomeromycota Mycologia 102: 2WKHU UHFHQW SURJUHVV KDV EHHQ PDGH RQ Acaulospora – species with pitted surface ornamentation, where several *HUGHPDQQ-:7UDSSH-0 7KHEndogonaceaeRIWKH3DFL¿F VSHFLHV WKDW VXSHU¿FLDOO\ DOO UHVHPEOHG A. scrobiculata, 1RUWKZHVWMycologia Memoir 5 – have been separated through extensive morphological and *RWR%76LOYD*$0DLD/&2HKO) Dentiscutata colliculosa, PROHFXODUVSRUHDQDO\VHV HJ2HKOet al. 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