AR TICLE Validation and Justification of the Phylum Name
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GRLLPDIXQJXV IMA FUNGUS · VOLUME 2 · NO 2: 173–175 9DOLGDWLRQDQGMXVWL¿FDWLRQRIWKHSK\OXPQDPHCryptomycota phyl. nov. ARTICLE 0HUHGLWK'0-RQHV7KRPDV$5LFKDUGV'DYLG/+DZNVZRUWKDQG'DYLG%DVV 6FKRRORI%LRVFLHQFHV8QLYHUVLW\RI([HWHU([HWHU(;4'8.FRUUHVSRQGLQJDXWKRUHPDLOGEDVV#QKPDFXN 'HSDUWPHQWRI=RRORJ\1DWXUDO+LVWRU\0XVHXP&URPZHOO5RDG/RQGRQ6:%'8. 'HSDUWDPHQWRGH%LRORJtD9HJHWDO,,)DFXOWDGGH)DUPDFLD8QLYHUVLGDG&RPSOXWHQVHGH0DGULG3OD]D5DPyQ\&DMDO0DGULG 6SDLQDQG'HSDUWPHQWRI%RWDQ\1DWXUDO+LVWRU\0XVHXP&URPZHOO5RDG/RQGRQ6:%'8. Abstract: 7KHUHFHQWO\SURSRVHGQHZSK\OXPQDPHCryptomycotaSK\OQRYLVYDOLGO\SXEOLVKHGLQRUGHUWRIDFLOLWDWHLWV Key words: XVHLQIXWXUHGLVFXVVLRQVRIWKHHFRORJ\ELRORJ\DQGSK\ORJHQHWLFUHODWLRQVKLSVRIWKHFRQVWLWXHQWRUJDQLVPV7KLVQDPH chitin LVSUHIHUUHGRYHUWKHSUHYLRXVO\WHQWDWLYHO\SURSRVHG³5R]HOOLGD´DVQHZGDWDVXJJHVWWKDWWKHOLIHVW\OHDQGPRUSKRORJ\ chytrid of Rozella is not representative of the large radiation to which it and other CryptomycotaEHORQJ)XUWKHUPRUHWD[DDW )XQJL higher ranks such as phylum are considered better not based on individual names of included genera, but rather on some phylogeny special characteristics – in this case the cryptic nature of this group and that they were initially revealed by molecular Rozella PHWKRGVUDWKHUWKDQPRUSKRORJLFDOGLVFRYHU\,IWKHJURXSZHUHODWHUYLHZHGDVDPHPEHURIDGLIIHUHQWNLQJGRPWKH 5R]HOOLGD QDPHVKRXOGEHUHWDLQHGWRLQGLFDWHLWVIXQJDODI¿QLWLHVDVLVWKHSUDFWLFHIRURWKHUIXQJDOOLNHSURWLVWJURXSV Article info:6XEPLWWHG6HSWHPEHU$FFHSWHG1RYHPEHU3XEOLVKHG1RYHPEHU INTRODUCTION )XQJL XQLFHOOXODU ]RRVSRUHV VLQJOHFHOOHG ZLWK D VLQJOH 7KH GHVLJQDWLRQ ³FU\SWRP\FRWD´ ZDV LQWURGXFHG E\ -RQHV PLFURWXEXODU ÀDJHOOXP DQG F\VWV ZLWKRXW D FKLWLQFHOOXORVH et al. WR DFFRPPRGDWH D ZHOOVXSSRUWHG FODGH XVLQJ FHOOZDOO)RUPLQJHSLELRQWLFDVVRFLDWLRQV ULERVRPDO '1$ U'1$ SK\ORJHQLHV RI RUJDQLVPV SXWDWLYHO\ EUDQFKLQJGHHSZLWKLQWKHIXQJDOUDGLDWLRQ7KHUDQNRISK\OXP Representatives *HQ%DQN DFFHVVLRQ QRV $- is the most appropriate for this group as current results show $-$-)-)-DQG)- that it has fungal characteristics but is distinct from other fungi and Rozella in not having a chitin-rich cell wall in the major stages of its life- F\FOHVRIDULGHQWL¿HGLQFOXGLQJSXWDWLYHWURSKLFLQWHUDFWLRQV Illustrations: Jones et al ¿JVGD±H However, Cryptomycota was not validly published as a VFLHQWL¿FQDPHLQWKDWZRUNDVQR/DWLQGLDJQRVLVZDVSURYLGHG 0F1HLOOet al$UW $/DWLQGLDJQRVLVLVSURYLGHGKHUH DISCUSSION LQRUGHUWRIRUPDOO\HVWDEOLVKWKHQDPH,QDGGLWLRQFRPPHQWV are made on our decision to introduce this name rather than It has been demonstrated that Rozella occupies a deep WDNH XS WKH HDUOLHU LQIRUPDO QDPH ³5R]HOOLGD´ DQG RQ WKH branching position in phylogenetic analyses of kingdom GLVWLQFWLYHIHDWXUHVRIWKHSK\OXPDQGLWVSRVLWLRQ Fungi -DPHVet al.DE DOWKRXJKERRWVWUDSVXSSRUWIRU this relationship is inconsistent and often weak in the most FRPSUHKHQVLYHO\ VDPSOHG SK\ORJHQLHV -DPHV et al. D TAXONOMY b, Jones et al 7KH QDPH ³5R]HOOLGD´ ZDV FRLQHG E\ /DUDet al WRDFFRPPRGDWHRozella and a number of Cryptomycota 0'0-RQHV 7$5LFKDUGVphyl. environmental sequences that form a distinct clade, but we nov. refer to this group henceforth as Cryptomycota for reasons 0\FR%DQN0% LQGLFDWHGEHORZ-RQHVet al. VKRZHGWKDWCryptomycota Etymology: crypto- – hidden; and -mycotaDSK\OXPRIIXQJL are more diverse than previously recognised and that the molecular diversity of this group may be as diverse as the rest )XQJLXQLFHOOXODUHV]RRVSRULVXQLFHOOXODULVXQLÀDJHOODWLEXVÀDJHOOLV of the known FungiDFFRUGLQJWRU'1$JHQHPDUNHUV PLFURWXEXODULVF\VWHVVLQHWXQLFDFKLWLQRVDYHOFHOOXORVD&RQVRUWLD 0HPEHUV RI Cryptomycota are found in freshwater, soil, HSLELRQWLFDIRUPDWD VHGLPHQWDQGVRPHPDULQHKDELWDWV-RQHVet al XVHG © 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. 2 173 Jones et al. OLQHDJHVSHFL¿F ÀXRUHVFHQFH in situ K\EULGL]DWLRQ ),6+ FHOO F\FOHFKDUDFWHULVWLF±ZRXOGEHPLVOHDGLQJ/DUDet al. ZDOO VWDLQV DQG LPPXQRÀXRUHVFHQFH VWDLQLQJ WR VKRZ WZR were also hesitant commending the use of the proposed distinct lineages within Cryptomycota, which comprised ovoid QDPH³EHWZHHQTXRWDWLRQPDUNVXQWLOPRUSKRORJLFDODQGRU cells of ca.PGLDPH[LVWLQJLQDWOHDVWWKUHHPRUSKRORJLHV XOWUDVWUXFWXUDOV\QDSRPRUSKLHVDUHGH¿QHGWRGLDJQRVHDQG LQ IUHVKZDWHU HQYLURQPHQWV XQLÀDJHOODWH ]RRVSRUHV PRUH YDOLGDWHWKLVHQWLUHJURXS´-RQHVet al GHPRQVWUDWH ARTICLE YDULDEO\VKDSHGFHOOVZLWKRXWÀDJHOODDWWDFKHGWRRWKHUHXNDU\RWLF that this key characteristic of Rozella does not seem to extend PLFURVFRSLFRUJDQLVPV HJGLDWRPKRVWV DQGQRQÀDJHOODWH DFURVVWKHZKROHJURXSDQGWKHUHIRUHWKHQDPH³5R]HOOLGD´LV F\VWV1RQHRIWKHVHVWDJHVZHUHVKRZQWRSRVVHVVDFKLWLQRU QRWUHSUHVHQWDWLYHRIWKHJURXSDVDZKROH FHOOXORVHZDOODOWKRXJKRWKHUOLIHF\FOHSKDVHVZLWKDFKLWLQDQG ,WLVLPSRUWDQWWRUHFRJQL]HWKDWRXUFXUUHQWNQRZOHGJH RUFHOOXORVHFHOOZDOOPD\UHPDLQXQGHWHFWHG$FKLWLQFHOOZDOOLV of the life stages of the newly discovered Cryptomycota VRPHWLPHVFLWHGDVGH¿QLQJIHDWXUHRINLQJGRPFungi, although and of Rozella LV YHU\ LQFRPSOHWH $V -RQHV et al. we note that this is not a reliable diagnostic feature as distantly suggest, chitin may be present in the walls of some currently related protist groups also possess chitin on their cell surface unknown CryptomycotaOLIHF\FOHVWDJH V DQGRUSUHVHQWLQ HJ.QHLSSet al. uncharacterized lineages within Cryptomycota, and even in 7KHQDPH³5R]HOOLGD´ZDVDSSOLHGWRWKLVSK\OXPE\/DUD currently unknown stages in Rozella,WZRXOGEHSUHPDWXUH et al EXWLQDQLQIRUPDOZD\EHWZHHQLQYHUWHGFRPPDV therefore, to separate Cryptomycota from the kingdom Fungi DQGZLWKQRIRUPDOGLDJQRVLV7KH,&=1 ,QWHUQDWLRQDO&RGH on the single character that they do not possess chitin walls RI =RRORJLFDO 1RPHQFODWXUH ,QWHUQDWLRQDO &RPPLVVLRQ RQ ZKLFKDVPHQWLRQHGDERYHLVQRWGLDJQRVWLFIRUFungi =RRORJLFDO 1RPHQFODWXUH GRHV QRW DSSO\ WR QDPHV Cryptomycota have some strong resemblances to above the rank of family-group, but if it were in those ranks Chytridiomycota µFK\WULGV¶ LQ ERWK VWUXFWXUH HJ ÀDJHOODU it would be viewed as unavailable as a conditional name DSSDUDWXV DQGHFRORJ\LIQRWLQFHOOZDOOFKHPLVWU\7KHUHLV $UW )RU QDPHV LQWURGXFHG XQGHU WKH ,&=1 ZKLFK QRDJUHHGGH¿QLQJQRQPROHFXODUFKDUDFWHULVWLFIRULGHQWLI\LQJ later are found to belong to Fungi WKH ,&1 ,QWHUQDWLRQDO the boundaries of kingdom Fungi7KHUHIRUHDVVHYHUDORWKHU &RGH RI 1RPHQFODWXUH IRU DOJDH IXQJL DQG SODQWV QRZ key characteristics are shared by Cryptomycota and some DFFHSWVWKHPDVDYDLODEOHXQGHU$UW DVUHYLVHGDWWKH Fungi, the former are most sensibly and parsimoniously 0HOERXUQH&RQJUHVVLQ-XO\0F1HLOOet al 7KXV considered as belonging to the latter as they form the closest QR/DWLQGLDJQRVLVZDVUHTXLUHGDVLWZDVIRUIXQJDOQDPHV EUDQFKHV RQ SK\ORJHQHWLF WUHHV -DPHV et al.DE LQWURGXFHGEHWZHHQDQG-DQXDU\ +RZHYHUZH /DUDet al. -RQHVet al 7KLVVWDQFHLVHQWLUHO\ DUHLQFOLQHGQRWWRDFFHSW³5R]HOOLGD´EHFDXVHRIWKHXVHRI consistent with the historical position regarding Rozella: for the inverted commas suggesting the usage was a tentative WKH ODVW \HDUV OHDGLQJ P\FRORJLVWV KDYH FODVVL¿HG WKLV suggestion and in any case note that it is not mandatory to genus within Fungi HJ+HOG.LUNet al follow the principle of priority of publication for names above Cryptomycota LQFOXGLQJRozella FRQVLVWHQWO\EUDQFK WKHUDQNRIIDPLO\ ,&1 RUIDPLO\JURXS ,&=1 ,QGHHGWKH with FungiLQDOOSK\ORJHQLHVVRIDUFRQVWUXFWHG+RZHYHUWKHLU ,&=1GRHVQRWFRYHUUDQNVKLJKHUWKDQIDPLO\JURXS position as the primary branch within fungi is much weaker :H GHFLGHG WKDW LW ZRXOG EH EHWWHU QRW WR GH¿QLWHO\ HJ-DPHVet al.-RQHVet al. ,QGHHGWKH\FRXOG establish a name based on Rozella for several reasons: actually occupy a higher branching position within Fungi,I 7KHIXQJDOWHUPLQDWLRQWREHXVHGIRUQDPHVLQWKH this is the case, their lack of some traditionally diagnostic UDQNRISK\OXPLV³mycota´XQGHUWKH,&1 0F1HLOOet al fungal features such as a chitin cell wall may be the result of $UW DQGWKDWWHUPLQDWLRQKDVDOVREHHQXVHGIRUSK\OD secondary losses, which would not preclude them from being traditionally studied by mycologists but which are no longer considered Fungi,QWKLVFDVHH[FOXGLQJCryptomycota from considered Fungi EXW SODFHG LQ RWKHU NLQJGRPV ([DPSOHV the Fungi could potentially make the rest of fungi paraphyletic include Hyphochytriomycota 5+:KLWWDNHU :KLWWDNHU ±DKLJKO\XQGHVLUDEOHDQGQRWORJLFDOO\VXVWDLQDEOHVLWXDWLRQ QRZSODFHGLQStraminipila0:'LFNMyxomycota In the absence of a strong morphological argument to exclude %ROG %ROG IRUVOLPHPRXOGVLQWKHProtozoa, and this group from the fungal kingdom – we must therefore look Oomycota $U[ $U[ IRU IXQJDO DQDORJXHV LQ WKH to the only available data, which is phylogenetic, and argues Straminipila 7KLV SUDFWLFH KDV EHHQ HPSOR\HG LQ VWDQGDUG that Cryptomycota are most reasonably considered to be UHIHUHQFH ZRUNV HJ .LUN et al. DQG DOVR WKH PRVW ZLWKLQ)XQJL UHFHQWWH[WERRNV HJ0RRUHet al. &RQVHTXHQWO\ZHDJUHHZLWK/DUDet al. WKDW Cryptomycota represent a very diverse radiation, there are sound reasons for considering Rozella DQG QRZ potentially equivalent