Racoleus, a new genus of sterile filamentous lichen-forming fungi from the tropics, with observations on the nomenclature and typification of Cystocoleus and Racodium

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© 2011 International Mycological Association

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INTRODUCTION

7KH HQLJPDWLF VWHULOH ¿ODPHQWRXV OLFKHQV SODFHG LQ Cystocoleus and Racodium DUH FKDUDFWHUL]HG E\ IXQJDO K\SKDH ZKLFK VXUURXQG D ¿ODPHQW RI WKH JUHHQ DOJD Trentepohlia7KHDOJDO¿ODPHQWVGHWHUPLQHWKHVKDSHDQGWKH enveloping hyphal layer is generally only single, the hyphae being parallel to the axis with elongate rectangular cells in Racodium, DQGLUUHJXODUO\WZLVWHGDURXQGWKHDOJDO¿ODPHQW in Cystocoleus. Both genera comprise a single species, and occur, often together, on inclined to vertical siliceous rocks in recesses where it is cool and there is no direct rain but a high KXPLGLW\7KH\DUHVFDUFHO\GLVWLQJXLVKDEOHPDFURVFRSLFDOO\ 1HYHUWKHOHVV GHVSLWH WKHLU DQDWRPLFDO VLPLODULW\ PROHFXODU data have now shown that the two genera are not part of a VLQJOH PRQRSK\OHWLF JURXS 0XJJLDet al. %RWK ZHUH found to belong to Capnodiales, with Racodium rupestre basal to the clade containing Cystocoleusebeneus, which is close to Mycosphaerellaceae 7KHVH UHVXOWV KDYH EHHQ FRQ¿UPHG E\ VXEVHTXHQW PROHFXODU SK\ORJHQHWLF DQDO\VHV &URXVet al.5XLEDOet al.ZLWKCystocoleus now being recognised as a member of Teratosphaeriaceae and the more basal Racodium as incertae sedis.

$VLPLODU¿ODPHQWRXVPHWKRGRIIRUPLQJDOLFKHQVWUXFWXUH LV VHHQ RWKHUZLVH RQO\ LQ D IHZ JHQHUD ,QCoenogonium, a leaf and bark dwelling member of the Gyalectaceae, the photobiont is a species of Trentepohlia in most species but FDQ DOVR EH D ¿ODPHQWRXV VSHFLHV RIPhysolinum LQ WKH corticolous sterile genus Pyrenothrix V\Q Lichenothrix WKH ¿ODPHQWRXV SKRWRV\QWKHWLF SDUWQHU EHORQJV WR WKH cyanobacterial genus ScytonemaEphebe, a genus mainly of riverside rocks and belonging to the Ephebaceae, where the photosynthetic partner is StigonemaDQGDOVRLQ some species of the mainly tropical basidiomycete genus Dictyonema where the photosynthetic partner also belongs to Scytonema$Physolinum forming lichen-like threads in which the algal cells can be in two or more rows, has also been documented from a dimly lit limestone cave, but the IXQJXVLQYROYHGKDVQRWEHHQLGHQWL¿HG'DYLVet al.,Q JHQHUDO¿ODPHQWRXVOLFKHQDVVRFLDWLRQVDUHH[WUHPHO\UDUH DQGQRQHZJHQXVRI¿ODPHQWRXVOLFKHQVKDVEHHQGHVFULEHG VLQFHWKHWKFHQWXU\

+HUH ZH GHVFULEH D WKLUG JHQXV RI VWHULOH ¿ODPHQWRXV lichens, Racoleus, for a tropical bark-inhabiting species NQRZQ WR 56 IRU RYHU \HDUV DQG GLIIHULQJ LQ WKH development of long hair-like lateral spines which are not seen

Racoleus

DQHZJHQXVRIVWHULOH¿ODPHQWRXVOLFKHQIRUPLQJIXQJLIURP

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Cystocoleus and Racodium

'DYLG/+DZNVZRUWK1_{, Rolf SantessonDQG/HLI7LEHOO}

1_{'HSDUWDPHQWRGH%LRORJtD9HJHWDO,,)DFXOWDGGH)DUPDFLD8QLYHUVLGDG&RPSOXWHQVHGH0DGULG3OD]D5DPyQ\&DMDO&LXGDG8QLYHUVLWDULD}

(0DGULG6SDLQDQG'HSDUWPHQWRI%RWDQ\1DWXUDO+LVWRU\0XVHXP/RQGRQ6:%'8.FRUUHVSRQGLQJDXWKRUHPDLOGKDZNVZRUWK# QKPDFXN

_{0XVHXPRI(YROXWLRQ8SSVDOD8QLYHUVLW\1RUE\YlJHQ6(8SSVDOD6ZHGHQ}

_{'HSDUWPHQWRI6\VWHPDWLF%LRORJ\(YROXWLRQDU\%LRORJ\&HQWUH8SSVDOD8QLYHUVLW\1RUE\YlJHQ'6(8SSVDOD6ZHGHQ}

Abstract:Racoleus trichophorusJHQVSQRYLVGHVFULEHGIRUDWURSLFDOVWHULOH¿ODPHQWRXVOLFKHQL]HG IXQJXV ZKLFK RYHUJURZV YDULRXV FUXVWRVH OLFKHQV RQ EDUN ,W VKDUHV VRPH IHDWXUHV ZLWKCystocoleus and RacodiumEXWLVXQLTXHLQKDYLQJQRQOLFKHQL]HGORQJODWHUDOVSLQHV7KHJHQXVZKLFKLVNQRZQ IURP&KLQDWKH,YRU\&RDVWDQG3HUXLVRIXQFHUWDLQV\VWHPDWLFSRVLWLRQRQWKHEDVLVRIPRUSKRORJLFDO

similarities, however, it may be referred to “? Capnodialesincertae sedis´ad interim,QDGGLWLRQWKH

QRPHQFODWXUHDQGW\SL¿FDWLRQRIWKHPRQRW\SLFJHQHUDCystocoleus and Racodium are reviewed, and

OHFWRW\SHV VHOHFWHG IRU WKH W\SH RI HDFK 7KH DYDLODEOH LQIRUPDWLRQ RQ WKH HFRORJ\ DQG GLVWULEXWLRQ RI WKHVHWZRJHQHUDLVDOVRVXPPDUL]HGDQGVFDQQLQJHOHFWURQPLFURJUDSKV6(0RIDOOWKUHHVSHFLHVDUH SUHVHQWHGIRUWKH¿UVWWLPH

Article info:6XEPLWWHG$SULO$FFHSWHG$SULO3XEOLVKHG0D\

Key words:

Ascomycota Capnodiales Coenogonium lichens

Hawksworth, Santesson, and Tibell

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in Cystocoleus and Racodium, and also in the arrangement of the hyphae and the way they interlock. In addition, we take the opportunity to present some other observations aimed DW FODULI\LQJ DQG ¿[LQJ WKH QRPHQFODWXUH DQG W\SL¿FDWLRQ RI Cystocoleus ebeneus and Racodium rupestre.

MATERIALS AND METHODS

0LFURVFRSLFH[DPLQDWLRQVZHUHPDGHZLWKHLWKHUD:LOGRUDQ 2OPSXV%+UHVHDUFKPLFURVFRSHERWK¿WWHGZLWKGUDZLQJ WXEHVDQGWKHODWWHUZLWKD1LNRQ&RROSL[GLJLWDOFDPHUD DQG1RPDUVNLLQWHUIHUHQFHFRQWUDVWRSWLFV$OOPHDVXUHPHQWV ZHUHPDGHLQZDWHUPRXQWV

Scanning electron micrographs were prepared from air-dried specimens which had been gold-coated during rotation XQGHU YDFXXP DQG H[DPLQHG LQ D 6WHUHRVFDQ &DPEULGJH 6FLHQWL¿F,QVWUXPHQWVRSHUDWLQJDWNY

'1$H[WUDFWLRQZDVDWWHPSWHGRQDIUDJPHQWRIWKH,YRU\ Coast isotype of Racoleus trichophorus LQ *=8 FDUHIXOO\ removed under a dissecting microscope, using the method of Cubero et alEXWZDVXQVXFFHVVIXO0ROHFXODUPHWKRGV and results obtained with Cystocoleus ebeneus and Racodium rupestreKDYHEHHQUHSRUWHGVHSDUDWHO\0XJJLDet al

Specimen citations for Cystocoleus ebeneus and Racodium rupestre are restricted to those discussed in UHODWLRQWRW\SL¿FDWLRQVRUGLVWULEXWHGLQH[VLFFDWDH

TAXONOMY

Racoleus

56DQW '+DZNVZ

gen. nov.

0\FR%DQN0%

Etymology: From the generic names Rac>odium@DQG>Cysto@ coleusZLWKZKLFKWKHJHQXVKDVVRPHIHDWXUHVLQFRPPRQ

Similis Racodiis rupestris, sed differt in cellulis verrucosis et in

VSLQXOLVODWHUDOLVQRQOLFKHQLEXVLQVWUXFWLV

Typus: Racoleus trichophorus 56DQW '+DZNVZ

Thallus VXSHU¿FLDO ÀXII\ EURZQ ¿ODPHQWRXVPhotobiont Trentepohlia, VLQJOH ¿ODPHQWV RI ZKLFK DUH HQVKHDWKHG E\ IXQJDOK\SKDHFilaments suberect to decumbent or spreading, sympodially branched, outer wall undulating and irregularly FRUUXJDWHGZLWKQXPHURXVODWHUDOVSLQHVHyphae in a single OD\HU VXUURXQGLQJ WKH DOJDO ¿ODPHQW RULHQWDWHG YHUWLFDOO\ DORQJDQGDOZD\VSDUDOOHOWRWKHD[LVRIWKH¿ODPHQWEURZQ septate, thick-walled, uneven and undulate to corrugated, not RUQDPHQWHGSpines arising at broadly acute to almost right angles to the vertical axis, brown, stiff, thick-walled, smooth-ZDOOHGQRWRUQDPHQWHGRUFRUUXJDWHGConidiogenous cells and conidiaXQNQRZQ

Observations:KLOHLQWKHDEVHQFHRIDQ\VH[XDOVWDWHDQG PROHFXODUGDWDQRGH¿QLWHRSLQLRQRQWKHV\VWHPDWLFSRVLWLRQ

of the new genus can be expressed, in view of the similarities to both Cystocoleus and Racodium it seems likely that it will also prove to belong to Capnodiales ZH WKHUHIRUH VXJJHVW that it is listed as “? Capnodialesincertae sedis´XQWLOIUHVK GDWDEHFRPHDYDLODEOH

Racoleus trichophorus

5 6DQW ' +DZNVZ

sp.

nov.

0\FR%DQN0%

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((

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EtymologyWKHHSLWKHWUHFDOOVWKHVSLQ\KDLUOLNHRXWJURZWKV

7KDOOLV OLFKHQLEXV FXP ¿ODPHQWLV —P ODWLV HW VSLQLV ODWHUDOLV DUFXDWLVQRQOLFKHQLEXVXVTXH±[±—PLQVWUXFWLV

Typus: Ivory Coast: $ELGMDQ LQ WKH IRUHVW RI %DQFR ca NP QRUWK RI$ELGMDQ o _{¶ 1} o _{¶ : RQ WUXQN RI D ODUJH}

tree in a very dark rainforest, overgrowing Dichosporidium brunnthaleri, -XO\ R. Santesson 10344a 836 KRORW\SXV*=8LVRW\SXV

Thallus VXSHU¿FLDO IRUPLQJ GHQVH ÀXII\ SDWFKHV UHFDOOLQJ FRWWRQZRRO WR PP GLDP SDOH WR IXVFRXV EURZQ ¿ODPHQWRXVPhotobiont Trentepohlia, VLQJOH ¿ODPHQWV RI ZKLFKDUHHQVKHDWKHGE\IXQJDOK\SKDHFilaments suberect to decumbent or spreading on the surface, sympodially EUDQFKHG±—PZLGHRXWHUZDOOXQGXODWLQJDQGLUUHJXODUO\ FRUUXJDWHGUHÀHFWLQJWKHPRUSKRORJ\RIWKHIXQJDOK\SKDH ZLWK QXPHURXV ODWHUDO VSLQHVHyphae in a single layer VXUURXQGLQJWKHDOJDO¿ODPHQWRULHQWDWHGYHUWLFDOO\DORQJDQG DOZD\V SDUDOOHO WR WKH D[LV RI WKH ¿ODPHQW EURZQ ± —P ZLGHVHSWDWHVHSWDJHQHUDOO\±—PDSDUWWKLFNZDOOHG uneven and undulate to corrugated, corrugations tending WRLQWHUOLQNZLWKWKRVHRIDGMDFHQWK\SKDHQRWRUQDPHQWHG Spines arising at broadly acute to almost right angles to the vertical axis, brown, stiff, thick-walled, smooth-walled, not ornamented or corrugated, arcuate to straight, directed RXWZDUGVDQGXSZDUGVPDLQO\± —PLQOHQJWKDQG± —P ZLGH JUDGXDOO\ WDSHUHG WRZDUGV WKH WLS ZKLFK LV ± —P ZLGH WKH EDVH H[SDQGHG LQWR D IRRWOLNH FHOO DGKHULQJ WR WKH DOJDO ¿ODPHQW DQG PHDVXULQJ ± —P LQ OHQJWK Conidiogenous cells and conidiaXQNQRZQ

Ecology: All collections are on tree trunks in dense shade in tropical rain forests and on whitish crustose lichens, notably Dichosporidium brunnthaleri, D. nigrocinctum, Pyrgillus indicus, DQXQLGHQWL¿HGDUWKRQLRLGOLFKHQSUREDEO\DVSHFLHVRI Cryptothecia7KHRacoleus overgrows the crustose lichens DQGKDVQRLQWLPDWHFRQWDFWZLWKWKHPDQGLVHDVLO\UHPRYHG :H GR QRW FRQVLGHU LW OLFKHQLFRORXV DQG LWV RFFXUUHQFH RQ whitish lichens is perhaps a sampling artefact, possibly due to LWEHLQJPRUHHDVLO\YLVLEOHDJDLQVWDZKLWHEDFNJURXQG

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Observations: This new genus differs from both Cystocoleus and Racodium LQWKHSUHVHQFHRIODWHUDOVSLQHV)LJ&±( &(DVZHOODVLQLWVHFRORJ\DQGGLVWULEXWLRQ,QDGGLWLRQ WKHK\SKDHVXUURXQGLQJWKHDOJDO¿ODPHQWGLIIHULQWKDWWKH\ DUH RULHQWDWHG SDUDOOHO WR WKH ¿ODPHQW D[LV ZLWK LQWHUORFNLQJ corrugations, the surface of the hyphae is smooth in the 6(0,QCystocoleus, similar corrugation occurs but is less pronounced and the hyphae are more irregularly arranged, WHQGLQJ WR ZUDS DURXQG WKH DOJDO ¿ODPHQW UDWKHU WKDQ EH strictly orientated along its axis, giving it a more knobbly DSSHDUDQFH )LJV $ $ WKH VXUIDFH RI WKH K\SKDH DOVR DSSHDURUQDPHQWHGLQWKH6(0)LJ$,QRacodium the hyphae lack interlocking corrugations, are thicker-walled than in the other genera, and fused to form elongated rectangular FHOOVRULHQWDWHGYHUWLFDOO\DORQJWKHD[LVRIWKHDOJDO¿ODPHQW )LJ % JLYLQJ DQ RYHUDOO VPRRWK UDWKHU WKDQ D NQREEO\

DSSHDUDQFHWKHK\SKDOZDOOVDUHFRPSOHWHO\VPRRWKLQWKH 6(0)LJ%7KHGLIIHUHQFHVEHWZHHQWKHVHWKUHHJHQHUD DUHVXPPDUL]HGLQ7DEOH

1RGHVFULSWLRQRIDIXQJXVUHFDOOLQJRacoleus trichophorus could be found in the lichenological or wider mycological OLWHUDWXUH ZH H[DPLQHG +RZHYHU DV ZH DUH OHVV IDPLOLDU with phycological publications, we cannot totally exclude the possibility that the dual organism has been given a name in DQROGDOJRORJLFDOZRUN

6DQWHVVRQ KDG QRWHG WKDW 'RGJH PHQWLRQHGD¿ODPHQWRXVOLFKHQZLWKEURZQK\SKDHIURP Costa Rica, and it is conceivable that could have been this species, but no illustration was provided and the material has QRW EHHQ UHH[DPLQHG 'RGJH WUHDWHG WKLV OLFKHQ XQGHU WKH name Coenogonium heterotrichum 0OO$UJ 0OOHU EXWDV'RGJHQRWHGWKDWVSHFLHVKDVFRORXUOHVVK\SKDH

Fig. 1. Racoleus trichophorusA, B. Habit, overgrowing Dichosporidum brunnthaleri RQ EDUNC, D. 'HWDLO RI OLFKHQL]HG ¿ODPHQWVE. SEM

PLFURJUDSKVKRZLQJWKHGHQWDWHZDOOVRIWKHK\SKDHRYHUWKHDOJDO¿ODPHQWDQGDOVRWKHFKDUDFWHULVWLFODWHUDOVSLQHVA-D Santesson 10344a,

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7KLV ZDV FRQ¿UPHG LQ WKH W\SH PDWHULDO Costa Rica: San José:6DQ0DUFRVGH'RWD7RQGX]DOWPRQWKDOOXVRI PyxineVSPitt 6115*±KRORW\SHZKLFK KDV DOJDO ¿ODPHQWV ZLWK VRPH HQFUXVWLQJ K\DOLQH K\SKDH VRPH RI ZKLFK JURZ RXW DZD\ IURP WKH ¿ODPHQWV EXW QR regular structure is evident with no brown hyphae or any IRUPLQJMLJVDZOLNHSDWWHUQVRQWKHVXUIDFH

Other specimens examined: China: Yunnan Province;LVKXDQJEDQQD

'LVWULFW-LQJKRQJ&R0HQJOXQ(OHFWULF6WDWLRQ0RQVXQIRUHVWYDOOH\

o_¶1o_{¶(DOWca PRQDWUHHLQDUDWKHUGDUN}

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Cryptothecia6HSWR. Santesson 32036b 836± Peru:

Dept. Loreto,TXLWRV([SORUDPD/RGJHcaNP1(RI,TXLWRV/DNH

7UDLO o_¶6 o_{¶:DOWca 100 m, on a tree trunk in a tropical}

rainforest, overgrowing Dichosporidium nigrocinctum,-DQ R.

& B. Santesson P7:4 .06836Dept. San Martin3URY

/DPDV&HUUR%ODQFRcaNPRQURDG::1:RI7DUDSRWRca

o_¶6 o_{¶:DOWca PRQDWUHHWUXQNLQDGDUNUDLQIRUHVW}

overgrowing Pyrgillus indicus, 0DUR. Santesson & G. Thor

P77:116836

Cystocoleus

Thwaites,

Ann. Mag. nat. Hist., ser. 2

3

:

Type: Cystocoleus ebeneus 'LOOZ7KZDLWHV

Cystocoleus ebeneus

'LOOZ 7KZDLWHV

Ann. Mag.

nat. Hist., ser. 2

3

Basionym: Conferva ebenea 'LOOZBr. Confervæ SO

Synonyms: Croolepus ebeneus'LOOZ&$JDUGKSyst. Alg

Racodium ebeneum'LOOZ)USumma Veg. Scand1

Coenogonium ebeneum 'LOOZ$/6PMongr. Br. Lich2:

Coenogonium germanicum *OFNFlora 82 Coenogonium schmidlei Simmer, Allgem. Bot. Zeit5

Byssus nigra DXFWSSQRQ +XGVFl. Anglica Conferva nigra DXFWSSQRQ +XGV5RWKCatal. Bot3:

Cystocoleus nigra DXFWSSQRQ +XGV+DULRWJ. Bot4 DV³niger´

Coenogonium nigrum DXFWSSQRQ +XGV=DKOEUAnn. Naturhist. Mus. Wien 25

Fig. 2ACystocoleus ebeneus ¿ODPHQW

Hawksworth .0 B. Racodium

rupestre¿ODPHQWDalhem80(

C-ERacoleus trichophorus Santesson,

.0 C )LODPHQWD 'HWDLO RI

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Type: United Kingdom: Morayshire: “On the stump of a tree >sic@LQ0DFEHWK¶V:RRG>%URGLH@QU)RUUHV´$XJW. J. Hooker%0±lectotypus hic designatus

Descriptions and illustrations (selected) *OFN -¡UJHQVHQ6FKDGH6NXMD 2UH6PLWK

6PLWKet alDQG:LUWK

Exsiccatae$Q]LLich. Langob. QR %0 836Krypt. Exs. Vindob. QR %0 836 0RXJHRW 1HVWOHU Stirps Crypt. QR %0 836 ZLWKRacodium rupestre Rabenhorst, Lich. Eur. QR %0 836 ZLWKRacodium rupestre DQG 5lVlQHQLichenotheca Fenn. QR %0 836

Fig. 36(0PLFURJUDSKVRI¿ODPHQWVA-BCystocoleus ebeneus Santesson 22339, 836C-D. Racodium rupestre Santesson 14386836

Hawksworth, Santesson, and Tibell

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Number of species0RQRW\SLF

Ecology: On vertical or somewhat inclined or underhanging siliceous rocks out of direct rain, but also on soil or eroded PRVVFXVKLRQVLQWKHVXEDQWDUFWLFLVODQGV-¡UJHQVHQ The species often grows mixed with Racodium rupestre. (FRORJLFDO LQFOXGLQJ TXDGUDW GDWD DUH SURYLGHG E\ VHYHUDO authors, including James et al6FKDGHDQG :LUWK7KHFRPPXQLWLHVIRUPHGEODFNIHOWOLNHSDWFKHV over extensive areas of rock, are so conspicuous that they have been given the phytosociological name Racodietum rupestris 6FKDGH 7KH PRVW FRPPRQO\ DVVRFLDWHG lichens are species of Lepraria and Leproloma.

Distribution (XURSH SDUWO\ PDSSHG E\ :LUWK 1RUWK $PHULFD&DQDGD86$6RXWK$PHULFD$UJHQWLQD%ROLYLD &KLOH &RORPELD 3HUX $IULFD .HQ\D $VLD 0RQJROLD $XVWUDODVLD 1HZ =HDODQG 7DVPDQLD DQG $QWDUFWLFD VXEDQWDUFWLFLVODQGV

Observations :H ZLVK WR GUDZ DWWHQWLRQ WR WKH RYHUORRNHG and painstaking work on the culture of this fungus reported by 6NXMD 2UHDQGLOOXVWUDWHGE\FRORXUSODWHV$IWHUWZR months in pure water, fungal hyphae grew free from the algal ¿ODPHQWV VSUHDGLQJ LUUHJXODUO\ EUDQFKLQJ DQG UHWDLQLQJ DQRGXORVHDSSHDUDQFHloc. cit.WDEOH,¿JTXLWHXQOLNH the lateral hairs in Racoleus. These workers also found that the Trentepohlia also grew out separately when cultured on %HQHFNHtVDJDUloc. cit.WDEOH,¿JV,QIUHVKPDWHULDOLQ WKH¿HOGWXIWVRIK\SKDHVLPLODUWRWKRVHUHSRUWHGE\6NXMD 2UHDUHRFFDVLRQDOO\HQFRXQWHUHG

There is also an interesting observation recorded in an DQQRWDWLRQ E\ : :DWVRQ RQ D PL[HG FROOHFWLRQ with Racodium rupestre from near Shepley in BM that ³:KHQ WUHDWHG ZLWK VWURQJ QLWULF DFLG WKHCoenogonium >LH Cystocoleus@DSSHDUVUHGGLVKZLWKWKH¿ODPHQWK\SKDHWZLVWHG whilst the RacodiumUHPDLQVGDUNZLWKK\SKDHSDUDOOHO´

‘YVWHGDO 6PLWK FRPPHQW WKDW ³7KH $QWDUFWLF SRSXODWLRQVGLIIHUVRPHZKDWIURPWKH1RUWK(XURSHDQRQHV DQG PD\ EH DQ XQGHVFULEHG WD[RQ´ EXW GR QRW HODERUDWH IXUWKHU DQG QR GLYHUJHQFHV ZHUH QRWHG E\ /LQGVD\ Material from Antarctica was not included in the molecular study of Muggia et al

*OFN LQ D FULWLFDO EXW OLWWOHFLWHG VWXG\ GLVWLQ guished Coenogonium germanicum from Cystocoleus rupestris on the basis of the differences in the arrangement

RIWKHK\SKDHZKLFKKHDOVRLOOXVWUDWHGLQWUDQVYHUVHVHFWLRQV He cultured the algal partners, which he referred to different species of Trentepohlia+RZHYHU*OFN¶VFULWLFDOO\H[HFXWHG illustrations leave no doubt as to the application of his species QDPHRQHRIWKHVHFRXOGEHGHVLJQDWHGDVOHFWRW\SHHJ3O ¿JV±LIQRRULJLQDOPDWHULDOFDQEHORFDWHGLQ+(,'RU0 ZKHUHVSHFLPHQVFRXOGEHORFDWHG6LPPHULOOXVWUDWHG Coenogonium germanicum and compared it with his newly described C. schmidlei ERWK KDG WKH LUUHJXODU K\SKDO arrangements typical of Cystocoleus ebeneus, and he seems to have separated them because of the proliferation of non-OLFKHQL]HG K\SKDH LQCoenogonium germanicum 2ULJLQDO material of C. schmidleiH[KHUE5HLPHUVLQ%H[DPLQHGE\ 56DQG/7LVLQGHHGCystocoleus ebeneus, VRFRQ¿UPLQJ WKHV\QRQ\P\

/LQGDX XVHG *OFN¶V QDPH LQ WKH VHQVH of Cystocoelus ebeneus, according to his description, and =DKOEUXFNQHU OLVWHG LW DV D V\QRQ\P RI WKH SUHVHQW VSHFLHV

/LQGVD\ FRPPHQWHG WKDW Coenogonium kerguelense GHVFULEHG E\ 'RGJH IURP .HUJXHOHQ Island, could either be a Coenogonium or the Cystocoleus 7KURXJK WKH FRXUWHV\ RI $ODQ )U\GD\ '/+ ZDV DEOH to H[DPLQHSKRWRPLFURJUDSKVRIRQHRIWKHWZRV\QW\SHV&RWH WDSLVVDQWOHVDOYHROHVGHODIDFH1(GX0RQW&DPSEHOO 1RYAlbert de la Rue # 64LQ)+7KLVFRPSULVHG a creamy yellowish buff felted colony which microscopically FRPSULVHGDOJDO¿ODPHQWVca—PZLGHZLWKDQHQFUXVWHG VXUIDFH 1R GLVWLQFWLYHCystocoleus or other dematiaceous hyphae were apparent in the photomicrographs, and the ZLGWKRIWKH¿ODPHQWVLVRXWVLGHWKHQRUPDO—PUDQJH of C. ebeneus 'RGJH¶V QDPH FRQVHTXHQWO\ DSSHDUV WR EH EDVHGRQDFRORQ\RIDQRQOLFKHQL]HGTrentephohliaVSHFLHV

Nomenclature: The name Byssus nigra is of uncertain DSSOLFDWLRQDQGQHYHUDSSHDUVWRKDYHEHHQIRUPDOO\W\SL¿HG but some authors have used the epithet for Cystocoleus ebeneusLQWKHSDVW7KHQDPHZDVLQWURGXFHGE\+XGVRQ IRUDVWLII¿ODPHQWRXVRUJDQLVPIRXQGRQFDOFDUHRXV URFNV QHDU ,QJOHERURXJK DQG 6HWWOH LQ <RUNVKLUH D PRVW LPSUREDEOHKDELWDWIRUWKLVOLFKHQ+RZHYHUKHVXEVHTXHQWO\ amended the notes on ecology and distribution to “in rupibus HWVD[LVJUDQGLRULEXVLQERUHDOLSDUWH$QJOLDHHWLQ:DOOLD´DQG UHIHUUHG WR LW DV WKH ³$QJOLV EODFN E\VVXV´ +XGVRQ Byssus nigraZDVIHDWXUHGDV³%ODFNURFNE\VVXV´LQEnglish Botany 6PLWK 6RZHUE\ ZLWK D UHIHUHQFH EDFN WR

Table 1. Main anatomical characters distinguishing the genera Cystocoleus, Racodium, and Racoleus

Character Cystocoleus Racodium Racoleus Hyphal arrangement Twisted Vertical Vertical Hyphal wall corrugation 3UHVHQW Absent 3UHVHQW

Hyphal wall ornamentation :DUWHG Smooth Smooth

/DWHUDOVSLQHV Absent Absent 3UHVHQW

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+XGVRQEXWQRW+XGVRQIURPWKHSDJHQXPEHU FLWHG LH S DQG QRW DQG VRPH HDUOLHU XVDJHV DQG SRO\QRPLDOV7KH RULJLQDO PDWHULDO RQ ZKLFK 6RZHUE\¶V illustration was based could not be located in BM, but an index to the original plates and specimens held there gives the modern name as Racodium rupestre and not Cystocoleus ebeneusRUDQ\RILWVV\QRQ\PV

+XGVRQ¶VELQRPLQDOZDVOLVWHGDVDV\QRQ\PRIConferva ebeneaE\'LOOZ\QZKHQLQWURGXFLQJWKDWQDPHEXW LQWHUHVWLQJO\IURPKLVFLWLQJ³S´KHZDVDOVRUHIHUULQJWR WKHVHFRQGHGLWLRQRI+XGVRQ¶VZRUNDVKDG6PLWK±ZKRLV QRWPHQWLRQHGE\'LOOZ\Q6PLWK¶VWH[WLQGLFDWHVWKDWLWLVPRVW likely he was dealing with either C. ebeneus or Racodium rupestre from his description of the habitat and comment WKDW³LWLVDOZD\VIRXQGRQDPLFDHRXVRUTXDUW]RVHVWRQH´ It appears to have been applied to both species by early DXWKRUV DQG VR LV EHVW OLVWHG DV D ³SUR SDUWH´ XVDJH XQGHU HDFKSHQGLQJDQ\IRUPDOW\SL¿FDWLRQ

A complication in the nomenclatural situation arises from the existence of a nomenclaturally independent name Conferva nigra+XGVEDVHGRQDGLIIHUHQWW\SH±LQGHHG that taxon was described as abundant on the seashore in <RUNVKLUH'LOOZ\QLQGLFDWHGWKDWKHKDGVHHQDXWKHQWLF specimens of that taxon, and considered to represent a seaweed which he called Conferva atro-rubescens 'LOOZ ±DUHGDOJDIRUZKLFKWKHFXUUHQWQDPHLVPolysiphonia nigra +XGV %DWWHUV IROORZLQJ QHRW\SL¿FDWLRQ RI WKLV ELQRPLQDORI+XGVRQ¶VE\0DJJV +RPPHUVDQG7KH H[LVWHQFHRIWKLVQDPHPHDQVWKDW'LOOZ\Q¶VFLWLQJRIByssus nigra as a synonym does not render Conferva ebenea as VXSHUÀXRXVDQGLOOHJLWLPDWHXQGHU$UWDVDFRPELQDWLRQ of Byssus nigra into Conferva would have created a homonym WREHUHMHFWHGXQGHU$UW

In introducing the generic name Cystocoleus, “with the VDQFWLRQ RI P\ IULHQG WKH 5HY 0 - %HUNHOH\´ 7KZDLWHV OLVWHG ³Byssus nigra, E.B W ´ DV D V\QRQ\P However, as he did not refer to Hudson at all and it is clear that he was listing the English Botany XVDJH VHH DERYH UDWKHUWKDQWUHDWLQJWKHQDPHDVDV\QRQ\P,QWKLV7KZDLWHV ZDV SHUKDSV IROORZLQJ +RRNHU ZKR GLG QRW mention Hudson and attributed the binominal Byssus nigra RQO\WR³6P´LQKLVLQGH[+RRNHU7KHOHJLWLPDF\ RI 7KZDLWHV¶ FRPELQDWLRQ LV QRW WKHUHIRUH WKUHDWHQHG E\ +XGVRQ¶VQDPHDVKHGLGQRWWUHDWLWDVDV\QRQ\PPHUHO\ listing the usage in English Botany

:H VHH QR QRPHQFODWXUDO REVWDFOH WR WKH FRQWLQXHG use of the name Cystocoleus ebeneus,I+XGVRQ¶VRULJLQDO PDWHULDORUDODWHUW\SL¿FDWLRQRIKLVQDPHZHUHGLVFRYHUHG and that indeed proved to belong to this taxon or to Racodium rupestreWKHHSLWKHWVKRXOGEHSURSRVHGIRUUHMHFWLRQLQRUGHU to maintain whichever of the two currently used names was WKUHDWHQHG

7\SL¿FDWLRQ:,QWKHRULJLQDO DFFRXQWRI'LOOZ\QWKUHH

VSHFLPHQV ZHUH PHQWLRQHG ³Dillenius. – On Rocks in WKH+LJKODQGVJames Brodie, Esq.´2QWKHVWXPSRID GHDGWUHHLQ0DFNEHWK¶V:RRGDW%URGLHQHDU)RUUHV1%

W.J. Hooker, Esq.´DQG³2Q%LUFKWUHHVDW&RIWHI\QHDU 1RUZLFKMr. S. Wilkins.´7KHVHFRQGFROOHFWLRQLVSUHVHQWLQ %0ODEHOOHG³2QWKHVWXPSRIDWUHHLQ0DFEHWK¶V:RRGQU )RUULHV >VLF@$XJ ´ DQG GHVSLWH WKH XQXVXDO KDELWDW UHSRUWHG LW LV DQ DSSURSULDWH OHFWRW\SH IRU 'LOOZ\Q¶V QDPH However, the specimen is not on a piece of bark or wood, does not have any evident adhering woody fragments, but GRHVKDYHVRPHJUDQLWLFFU\VWDOVLQWHUPL[HGWKLVFDXVHVXV WR GRXEW WKDW LW ZDV JURZLQJ GLUHFWO\ RQ D WUHHVWXPS 7KDW 7KZDLWH¶VSO¿JV±DFWXDOO\LOOXVWUDWHGDVSHFLPHQ of Racodium rupestre does not affect the application of the generic name as “Conferva ebenea, Dillw. W´ZDVJLYHQ as the basionym of Cystocoleus ebeneus.

Racodium

)U

Syst. Mycol.

3

nom. cons.

Synonyms: Rhacodium 6SUHQJLinn. Syst. Veg., 16th _{edn, 4:}

orth. varnom. illegit$UWV

Rhacodiopsis Donk, Persoonia 8 nom. illegit $UW

1RQRacodium3HUVNeues Mag. Bot. 1)U Syst. Mycol. 1[OYLnom. rej.

Type: Racodium rupestre 3HUV

Racodium rupestre

3HUV

Neues Mag. Bot.

1

Synonyms: Byssus rupestris 3HUV '&in /DPDUFN DeCandolle, Fl. Françrd _{edn 2}

Dematium rupestre 3HUV1HHVSyst. Pilze Cystocoleus rupestris 3HUV5DEHQKKrypt.-Fl. Sachsen 2:

Rhacodiopsis rupestris 3HUV 'RQN Persoonia 8

Byssus nigra DXFWSSQRQ +XGVFl. Anglica Conferva nigra DXFWSSQRQ +XGV5RWKCatal. Bot3:

Cystocoleus nigra DXFWSSQRQ +XGV+DULRWJ. Bot4 DV³niger´

Coenogonium nigrum DXFWSSQRQ +XGV=DKOEUAnn. Naturhist. Mus. Wien 25

Type: United Kingdom: Wales: “Racodium rupestre 3 Byssus nigra (QJO %RW :DOHV +E 3HUV´ / SURSDUWH±lectotypus hic designatus

Descriptions and illustrations (selected): Brodo et al

*OFN 6PLWK 6PLWKet al 7KZDLWH¶V

DVCystocoelus ebeneus

Hawksworth, Santesson, and Tibell

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Number of species0RQRW\SLF

Ecology: As for Cystocoleus ebeneus.

Distribution (XURSH PRVW FRXQWULHV 1RUWK $PHULFD &DQDGD 86$ 6RXWK $PHULFD $UJHQWLQD &KLOH $IULFD 6RXWK$IULFD$VLD-DSDQDQG$XVWUDOLD7DVPDQLD

Observations =DKOEUXFNQHU 9DLQLR 'RGJH DQG &KULVWLDQVHQ PHQWLRQ WKH DOJD LQ Racodium as belonging to the genus Cladophora rather than to TrentepohliaEXWWKHEDVLVIRUWKLVLVREVFXUH,QRUGHUWR resolve this matter, the alga was isolated into pure culture by .RFKZKRIRXQGLWWREHDPHPEHURIWKHTrentepohlia aureaJURXS

Nomenclature: The nomenclatural issues surrounding this QDPH DUH FRPSOH[ 5LHGO DQG +DZNVZRUWK independently noted that “Racodium´ZDVEHLQJXVHGIRUD VWHULOH ¿ODPHQWRXV OLFKHQ E\ OLFKHQRORJLVWV DQG IRU D TXLWH GLIIHUHQW QRQOLFKHQL]HG K\SKRP\FHWH E\ P\FRORJLVWV These authors both concluded that the name should be W\SL¿HG E\ WKH OLFKHQL]HG HOHPHQW LHR. rupestre UDWKHU WKDQ WKH FRQLGLDO IXQJXV LHR. cellare 3HUV EXW LQ D SRVWKXPRXV SXEOLFDWLRQ 'RQN GLVDJUHHG ,Q RUGHU WRUHVROYHWKHPDWWHU+DZNVZRUWK 5LHGOSURSRVHG WKH QDPH IRU FRQVHUYDWLRQ IRU WKH OLFKHQL]HG IXQJXV WKHLU proposal was accepted and it is now list as conserved in the Code7KHQDPHRhinocladiella ellisii '+DZNVZ ZDV LQWURGXFHG IRU WKH FRQLGLDO VWDWH RI WKH ³FHOODU IXQJXV´ However, sterile material of the same species, which does not form conidia, had been referred to as Zasmidium cellare 3HUV )U DQG ZDV FRQVLGHUHG D V\QDQDPRUSK 'H +RRJGLGQRWFRQVLGHUWKHVHQDPHVDVV\QDQDPRUSKV as the conidiogenous cells were micronematous and not PDUNHGO\GLIIHUHQWIURPWKHVWHULOHK\SKDHDQGFRQVHTXHQWO\ commend the use of Zasmidium cellareIRUWKLVIXQJXV7KLV interpretation has been followed in the molecular phylogenetic VWXG\ RI$U]DQORXet al ZKR IRXQG WKDW LW FOXVWHUHG with Ramichloridum species and published photographs of WKHFRQLGLRJHQRXVFHOOVDQGFRQLGLD

'RQNQRWHGWKDW³Rhacodium´ZDVRUWKRJUDSKLFDOO\ the more correct spelling of the generic name, as derived IURPWKH*UHHN³ȡĮȤȠȢ´UDJEXWWKHIRUP³Racodium´LVWKDW FRQVHUYHGLQDFFRUGDQFHZLWKJHQHUDOXVDJH

9DLQLROLVWHGCoenogonium germanicum as a synonym of Racodium rupestre, but the original illustrations of *OFNDQGWKRVHRI6LPPHUDUHRICystocoleus ebeneusDVLQWHUSUHWHGKHUH

+HQVVHQ -DKQVHYLGHQWO\UHJDUGHGWKLVVSHFLHV as congeneric with Cystocoleus EXW GLG QRW H[SODLQ ZK\ PROHFXODUGDWDVKRZWKDWYLHZWREHXQVXSSRUWDEOH

7\SL¿FDWLRQ7KHW\SL¿FDWLRQRIWKHQDPHRacodium rupestre

KDVQRWEHHQDGGUHVVHGDQGDIRUPDOW\SL¿FDWLRQSXEOLVKHG ,Q RUGHU WR FODULI\ WKH LVVXH WKH VSHFLPHQV LQ 3HUVRRQ¶V FROOHFWLRQ LQ / ZHUH VWXGLHG E\ 56 LQ ZKR GLG QRW

SXEOLVK KLV UHVXOWV DW WKDW WLPH 56 IRXQG WKDW WKHUH ZHUH four specimens under this name:

/ ³Racodium rupestre 3HUV 6\Q LQ

UXSLEXV XPEURVLV 0RXJ LQ KE 3HUV´ 7KLV LVCystocoleus ebeneus ZLWKVSDUVHR. rupestre

/ ³Racodium rupestre 3Byssus nigra (QJO%RW:DOHV+E3HUV´7KLVLVR. rupestre ZLWKVSDUVH C. ebeneus

/³Racodium rupestre 3HUV+E3HUV´7KLV is C. ebeneus with no Racodium

/³1R$Q6DQGKDOHQ0$XJRacodium

rupestre. +E 3HUV´ 7KLV FRQVLVWV RQO\ RI QRQOLFKHQL]HG IXQJDOK\SKDH

Three of the collections were mixed with Cystocoleus ebeneus, DQG LW LV HYLGHQW WKDW LQ SUDFWLFH 3HUVRRQ DSSOLHG Racodium rupestre LQDVHQVHHPEUDFLQJERWKJHQHUD7KLV is hardly surprising as the species commonly grow mixed WRJHWKHUDQGUHTXLUHPLFURVFRSLFVWXG\WRVHSDUDWHWKHPZLWK FRQ¿GHQFH,QRUGHUWR¿[WKHDSSOLFDWLRQRI3HUVRRQ¶VQDPH LQWKHVHQVHLQZKLFKLWLVFXUUHQWO\XVHG/LV FRQVHTXHQWO\GHVLJQDWHGDVOHFWRW\SHKHUH as that is the only one in which the Racodium SUHGRPLQDWHV

ACKNOWLEDGEMENTS

:H DUH LQGHEWHG WR WKH FXUDWRUV RI DOO FROOHFWLRQV FLWHG LQ WKH WH[W

and others from which we have examined material during this study

IRUDFFHVVWRFROOHFWLRQVLQWKHLUFDUH%%0*./0$)80( 836DQG6*|UDQ7KRU6ZHGLVK$JULFXOWXUDO8QLYHUVLW\8SSVDOD IRU LGHQWL¿FDWLRQ RI WKH OLFKHQV EHLQJ RYHUJURZQ E\ WKHRacoleus,

DQG $ODQ 0 )U\GD\ 0LFKLJDQ 6WDWH 8QLYHUVLW\ ZKR NLQGO\ PDGH

available photomicrographs of Coenogonium kerguelense.:RUNRQ

the text and the drawings included in this paper commenced while

'/+ZDVDYLVLWLQJUHVHDUFKHUDWWKH8QLYHUVLW\RI8PHnLQ DQGKHLVJUDWHIXOWR2YH((ULNVVRQIRUWKHIDFLOLWLHVSURYLGHGDWWKDW WLPH,WZDVFRPSOHWHGZKLOH'/+ZDVLQUHFHLSWRIVXSSRUWIURP WKH0LQLVWHULRGH(GXFDFLyQ\&LHQFLDRI6SDLQ3UR\HFWRV,'&*/

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%URZQ*UHHQDQG/RQJPDQV

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+XGVRQ:Flora Anglicand_{HGQ/RQGRQ-1RXUVH}

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.RFK:'LH*RQLGLHYRQRacodium rupestre3HUVVorträge

aus dem Gesamtgebiet der Botanik, neue folge 1±

/LQGDX*Die Flechten>.U\SWRJDPHQÀRUDI|U$QIlQJHUQR@

%HUOLQ-XOLXV6SULQJHU

/LQGDX*Die Flechtennd_{HGQ>.U\SWRJDPHQÀRUDI|U$QIlQJHU}

QR@%HUOLQ-XOLXV6SULQJHU

/LQGVD\'&1RWHVRQ$QWDUFWLFOLFKHQV,,,Cystocoleus niger

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5LHGO + 'LH *DWWXQJVQDPHQ Racodium 3HUVRRQ XQG

Zasmidium )ULHVTaxon 17±

5XLEDO & *XHLGDQ & 6HOEPDQQ / *RUEXVKLQD $$ &URXV 3: *URHQHZDOG-=0XJJLD/*UXEH0,VROD'6FKRFK&/6WDOH\ -7/XW]RQL)GH+RRJ*63K\ORJHQ\RIURFNLQKDELWLQJ

fungi related to DothideomycetesStudies in Mycology 64±

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Upsaliensis 12±

6FKDGH >)@$ 'LH 9HUEULHWXQJ YRQRacodium rupestre 3HUV

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±

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Latviensis 8±

6PLWK$/ A Monograph of British Lichens. 9RO QG HGQ

/RQGRQ%ULWLVK0XVHXP1DWXUDO+LVWRU\

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/HLS]LJ*%RUQWUDHJHU

=DKOEUXFNQHU$ >/LFKHQHV )OHFKWHQ@ % 6SH]LHOOHU7HLODie