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Molecular Phylogenetic Studies on the Lichenicolous Xanthoriicola Physciae

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Molecular Phylogenetic Studies on the Lichenicolous Xanthoriicola Physciae GRLLPDIXQJXV IMA FUNGUS · VOLUME 2 · NO 1: 97–103 Molecular phylogenetic studies on the lichenicolous Xanthoriicola physciae ARTICLE reveal Antarctic rock-inhabiting fungi and Piedraia species among closest relatives in the Teratosphaeriaceae Constantino Ruibal1$QD00LOODQHVDQG'DYLG/+DZNVZRUWK 1'HSDUWDPHQWRGH%LRORJtD9HJHWDO,,)DFXOWDGGH)DUPDFLD8QLYHUVLGDG&RPSOXWHQVHGH0DGULG3OD]D5DPyQ\&DMDO0DGULG6SDLQ 'HSDUWDPHQWRGH%LRORJtD\*HRORJtD(6&(78QLYHUVLGDG5H\-XDQ&DUORV0yVWROHV0DGULG6SDLQ 'HSDUWPHQWRI%RWDQ\7KH1DWXUDO+LVWRU\0XVHXP&URPZHOO5RDG/RQGRQ6:%'8.FRUUHVSRQGLQJDXWKRUHPDLOGKDZNVZRUWK# QKPDFXN Abstract: The phylogenetic placement of the monotypic dematiaceous hyphomycete genus Xanthoriicola Key words: ZDVLQYHVWLJDWHG6HTXHQFHVRIWKHQ/68UHJLRQZHUHREWDLQHGIURPVSHFLPHQVRIX. physciae, which Ascomycota IRUPHGDVLQJOHFODGHVXSSRUWHGERWKE\SDUVLPRQ\ DQGPD[LPXPOLNHOLKRRG ERRWVWUDSV Capnodiales DQG%D\HVLDQ3RVWHULRU3UREDELOLWLHV 7KHFORVHVWUHODWLYHVLQWKHSDUVLPRQ\DQDO\VLVZHUHVSHFLHV Friedmanniomyces of Piedraria, while in the Bayesian analysis they were those of Friedmanniomyces7KHVHWKUHHJHQHUD hyphomycetes along with species of Elasticomyces, Recurvomyces, TeratosphaeriaDQGVHTXHQFHVIURPXQQDPHGURFN lichenicolous fungi LQKDELWLQJIXQJL 5,) ZHUHDOOPHPEHUVRIWKHVDPHPDMRUFODGHZLWKLQCapnodiales with strong support Piedrariaceae in both analyses, and for which the family name Teratosphaeriaceae can be used pending further studies rock inhabiting fungi RQDGGLWLRQDOWD[D Article info:6XEPLWWHG0D\ $FFHSWHG0D\3XEOLVKHG-XQH INTRODUCTION RI WKH FROODUHWWH RI WKH FRQLGLRJHQRXV FHOOV 7KLV LV D UDUH situation in hyphomycetous conidial fungi, and is otherwise The generic name Xanthoriicola was introduced for the species seen only in Craspedodidymum, Cystodendron, Lambinonia, X. physciae +DZNVZRUWK 3XQLWKDOLQJDP 7KLVIXQJXV Metacapnodium, and some groups of Phialophora s. lat appears to be obligately lichenicolous on Xanthoria parietina (OOLV6HLIHUWet al ,QDGGLWLRQVRPHVWXGLHVE\ LQ(XURSHDQGLVDOVRUHSRUWHGIURP$IULFDDQG$VLD 6LHIHUW Raman spectroscopy suggested that Xanthoriicola physciae et al 7KH IXQJXV SULPDULO\ RFFXUV LQ WKH DSRWKHFLD might form scytonemin, a protective pigment only otherwise growing through the hymenium, with broad cupulate NQRZQLQF\DQREDFWHULD 3UHHFH ±DOWKRXJKWKDWUHSRUW enteroblastic conidiogenous cells generating conidia at the now seems likely to have been a result of contamination from VXUIDFH )LJ 7KHFRQLGLDDUHGDUNEURZQVSKHULFDOVLQJOH F\DQREDFWHULDJURZLQJRQWKHVXUIDFHRIWKHK\PHQLXP FHOOHG DQG KDYH D FRDUVH ZDUWHG VXUIDFH RUQDPHQWDWLRQ Xanthoriicola physciae FRQVHTXHQWO\ DSSHDUHG RQ 7KLVIXQJXVZDVLOOXVWUDWHGE\OLQHGUDZLQJVLQ+DZNVZRUWK morphological grounds to occupy an isolated position 3XQLWKDOLQJDP DQG SKRWRPLFURJUDSKV DQG VFDQQLQJ DPRQJVWWKHFRQLGLDOIXQJL7KLVLQYHVWLJDWLRQZDVLQLWLDWHGLQ HOHFWURQ PLFURJUDSKV DUH SUHVHQWHG LQ +DZNVZRUWK RUGHUWRGHWHUPLQHLWVSKO\RJHQHWLFUHODWLRQVKLSV The surfaces of infected apothecia become sooty black and VR DUH HDVLO\ VHHQ LQ WKH ¿HOG :KROH VZDUGV RI WKH KRVW lichen are rarely affected, so while deleterious to the host it MATERIALS AND METHODS GRHV QRW GHVWUR\ WKHLU SRSXODWLRQV ,W KDV QRW EHHQ UHSRUWHG as growing in isolated culture, and experiments to inoculate Choice of additional taxa and outgroup IUHVK VSHFLPHQV RI WKH KRVW KDYH SURYHQ XQVXFFHVVIXO 7) In addition to the Xanthoriicola VSHFLPHQV VHTXHQFHG 3UHHFHXQSXEOGDWD )XUWKHUPRUHQRVH[XDOVWDWHKDVEHHQ VSHFLPHQV RI Dothideomycetes were included in the discovered or postulated by association with other fungi that PROHFXODU VWXG\ )LJV ± 7KH VDPSOLQJ ZDV VHOHFWHG RFFXURQWKHVDPHKRVWOLFKHQ WR LQFOXGH WD[D WKDW ZHUH FORVH WR RXU QHZ VHTXHQFHV The fungus is particularly unusual in that the conidia are LQ *HQ%DQN LH PDLQO\ FRQLGLDO PHPEHUV RI WKH IRUPHG ³VHPLHQGRJHQRXVO\´ WKDW LV ZLWKLQ WKH ORZHU SDUW Teratosphaeriaceae, together with other representatives of © 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. 1 97 Ruibal, Millanes, and Hawksworth ;) ¶*$*$**$7*&77&7***&$¶ DQG ;5 ¶&&*$$*&7&&&$&&7&&*77¶ 3ULPHU FRPELQDWLRQV XVHGZHUH/55/5/55;5DQG;)/53&5 3&5 DPSOL¿FDWLRQV ZHUH SHUIRUPHG XVLQJ ,OOXVWUD +RW 6WDUW 3&5 EHDGV DFFRUGLQJ WR WKH PDQXIDFWXUHU¶V ARTICLE instructions, and using the settings in Hawksworth et al %HIRUHVHTXHQFLQJWKH3&5SURGXFWVZHUHSXUL¿HGXVLQJ WKH 9LRJHQH 3&50 &OHDQXS 6\VWHP RU WKH HQ]\PDWLF PHWKRG([RVDS,7 Sequence alignment and phylogenetic analysis 6HTXHQFHVZHUHDOLJQHGXVLQJ0$))7Y .DWRKet al. .DWRK 7RK XVLQJWKHSURFHGXUHVGHVFULEHGLQ :HGLQet al 7KHDPELJXRXVUHJLRQVLQWKHDOLJQPHQW ZHUH LGHQWL¿HG DQG HOLPLQDWHG XVLQJ *EORFNV Y E &DVWUHVDQD Maximum parsimony and parsimony bootstrap analyses ZHUHSHUIRUPHGXVLQJ3$83YE 6ZRIIRUG ZLWKWKH IROORZLQJVHWWLQJVJDSVZHUHWUHDWHGDV³PLVVLQJGDWD´ UDQGRPDGGLWLRQVHTXHQFHUHSOLFDWHV7%5EUDQFKVZDSSLQJ steepest descent off, collapse branches if minimum length is 0, MulTrees on, and with 1000 trees allowed to be saved LQ HDFK UHSOLFDWH )RU WKH ERRWVWUDS DQDO\VHV )HOVHQVWHLQ ZH XVHG KHXULVWLF VHDUFK VHWWLQJV LGHQWLFDO ZLWK WKH above analysis, but with ten random addition replicates, 1000 bootstrap replicates, a full heuristic search, and retained JURXSV ZLWK D IUHTXHQF\ ! 3DUVLPRQ\XQLQIRUPDWLYH FKDUDFWHUVZHUHH[FOXGHGIURPWKHVHDQDO\VHV 0D[LPXP OLNHOLKRRG DQDO\VHV 0/ ZHUH DFKLHYHG Fig. 1. Xanthoriicola physciae ,0, A. Apothecia of XVLQJWKHSURJUDP*DUOLY =ZLFNO 5XQVZHUH Xanthoria parietinaLQIHFWHGE\WKHIXQJXVB. Conidiogenous cells WHUPLQDWHG DIWHU JHQHUDWLRQV ZLWK QR VLJQL¿FDQW LQWKHXSSHUSDUWRIWKHK\PHQLXPC. &RQLGLD%DUV$ PP%±& LPSURYHPHQWLQ±OQ/,PSURYHPHQWYDOXHVZHUHVHWWR P ZLWKDWRWDOLPSURYHPHQWORZHUWKDQFRPSDUHGWRWKHODVW WRSRORJ\UHFRYHUHG%RRWVWUDSVXSSRUWZDVDVVHVVHGXVLQJ WUHHUHSOLFDWHVXQGHUWKHVDPHSDUDPHWHUVDVDERYH Capnodiales and DothidealesDothidea insculpta was used :H XVHG WKH %D\HVLDQ PHWKRG RI +XHOVHQEHFN et al DVRXWJURXS WR DQDO\VH WKH GDWD E\ 0DUNRY &KDLQ 0RQWH &DUOR 9RXFKHU LQIRUPDWLRQ DQG *HQ%DQN DFFHVVLRQ QXPEHUV 0&0& VDPSOLQJDVLPSOHPHQWHGLQWKHVRIWZDUH0U%D\HVY RIQHZO\VHTXHQFHGWD[DDUHSURYLGHGLQ7DEOH +XHOVHQEHFN 5RQTXLVW /LNHOLKRRGPRGHOVZHUH selected for each of the three gene regions using the Akaike DNA extraction ,QIRUPDWLRQ &ULWHULRQ $,& DQG WKH %D\HVLDQ ,QIRUPDWLRQ '1$ ZDV H[WUDFWHG GLUHFWO\ IURP GULHG VSHFLPHQV )XQJL &ULWHULRQ %,& DVLPSOHPHQWHGLQM0RGHOWHVW 3RVDGD growing in the host apothecia were carefully excised with :HXVHGIXOOOLNHOLKRRGRSWLPL]DWLRQDQGVHOHFWHGIURPDPRQJ WKHSRLQWRIDVWHULOHVFDOSHOEODGHWRPLQLPL]HDVPXFKDV RQO\ WKH PRGHOV LPSOHPHQWHG LQ 0U%D\HV )ROORZLQJ WKLV SRVVLEOHWKHREWDLQLQJRIKRVWWLVVXH7RWDO'1$ZDVH[WUDFWHG VFKHPHD*75,*PRGHOZDVFKRVHQIRUWKHQXFOHDU/68 XVLQJ WKH 4LDJHQ '1HDV\ 3ODQW 0LQL.LW DFFRUGLQJ WR WKH U'1$GDWDXVLQJERWKFULWHULD7KHQXPEHURIGLVFUHWHJDPPD PDQXIDFWXUHU¶VLQVWUXFWLRQV FDWHJRULHVZDVNHSWDWGHIDXOWIRXU%D\HVLDQSULRUGLVWULEXWLRQV LQFOXGHGWUHDWLQJDOOWUHHWRSRORJLHVDVHTXDOO\OLNHO\DXQLIRUP $PSOL¿FDWLRQDQGVHTXHQFLQJ GLVWULEXWLRQIRUWKHJDPPDVKDSHSDUDPHWHUDXQLIRUP A fragment of caESLQWKHQ/68ZDVDPSOL¿HGXVLQJ GLVWULEXWLRQIRUWKHSURSRUWLRQRILQYDULDEOHVLWHVDQGD WKH SULPHUV /55 5 9LOJDO\V ZZZELRORJ\GXNHHGX ÀDW 'LULFKOHWIRUWKHUDWHPDWUL[7ZRSDUDOOHO IXQJLP\FRODESULPHUVKWP /5 9LOJDO\V +HVWHU UXQVZHUHSHUIRUPHGHDFKZLWK¿YHFKDLQVIRXURIZKLFKZHUH DQG DOVR RQHV VSHFL¿FDOO\ GHVLJQHG LQ RXU ODERUDWRU\ WR LQFUHPHQWDOO\KHDWHGZLWKDWHPSHUDWXUHRI7KHDQDO\VLV VHOHFWLYHO\ DPSOLI\ WKH '1$ RI Xanthoriicola physciae, was diagnosed for convergence every 100 000 generations, DYRLGLQJ WKDW RI WKH KRVW 7KH SULPHUV ZH GHVLJQHG ZHUH measured as the average standard deviation of splits across 98 IMA FUNGUS Molecular phylogeny of Xanthoriicola physciae GU214428 Hortaea acidophila 72 ARTICLE 100 EU436763 Devriesia strelitziae GU214414 Devriesia hilliana EU019248 Capnobotryella renispora EU019256 Teratosphaeria pseudosuberosa EU019274 Penidiella columbiana EU019278 Penidiella venezuelensis 88 EU019270 Hortaea werneckii 100 GU301818 Hortaea werneckii GU323969 TRN122 EU019250 Stenella araguata 95 AY016366 Piedraia hortae GU214468 Piedraia quintanilhae Xanthoriicola physciae CR4 81 Xanthoriicola physciae CR19 Xanthoriicola physciae CR44 Xanthoriicola physciae CR20 Xanthoriicola physciae CR21 91 Xanthoriicola physciae CR22 Xanthoriicola physciae CR23 Xanthoriicola physciae CR45 Xanthoriicola physciae CR50 Xanthoriicola physciae CR51 Xanthoriicola physciae CR58 100 GU250364 Friedmanniomyces endolithicus GU250367 Friedmanniomyces endolithicus GU250366 Friedmanniomyces endolithicus GU250368 Friedmanniomyces simplex 95 GU301874 Teratosphaeria jonkershoekensis GU323956 TRN5 GU250376 Elasticomyces elasticus GU250375 Elasticomyces elasticus GU323978 TRN211 GU250398 CCFEE5499 FJ790299 Teratosphaeria hortaea 80 EU019295 Teratosphaeria stellenboschiana GU214702 Teratosphaeria destructans FJ493217 Teratosphaeria juvenalis GQ852580 Baudoinia compniacensis 100 DQ678092 Catenulostroma abietis EU019255 Catenulostroma microsporum GU323983 TRN279
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