ARTICLE 373 . et al. et al. (Saale), hyphomycetes taxonomy Key words: Ascomycota Cercospora s. lat sp. sp. RUVH\RXRU\RXUXVHRIWKHZRUN nom. comb. IMA FUNGUS IMA (2015) 373–469 · 6(2): . The species P. viburni-nudi P. C. celosiigena WKH¿UVWSDUWRIWKLV P. depazeoides P. C. thunbergiigena nom. nov., nom. nov., Amaranthaceae sp. nov., and sp. nov., ). The fourth contribution of this The fourth contribution ). comb. nov., comb. nov., nom. nov., nom. nov., , and ivecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get ivecommons.org/licenses/by-nc-nd/3.0/legalcode. ) 4. Species on dicots on dicots ) 4. Species MATERIALS AND METHODS MATERIALS on our papers The present work is a compilation based global literature. and unpublished data, as well as the papers cited under Details of methods are given in the are concerned, As far as new examinations references. fungal structures have been examined by standard methods using an Olympus BX50 microscope, of light microscopy, with distilled water and lactic acid as media, but without any staining. If possible, measurements of 30 conidia and other VWUXFWXUHV KDYH EHHQ PDGH DW DThe following All illustrations have been prepared by UB. PDJQL¿FDWLRQ RI î abbreviations are used: author names follow Brummit & Powell (1992), journals Bridson (2004a, b), and exsiccatae http://www.botanischestaatssammlung.de/DatabaseClient/ Taxonomy IndExs/index.jsp (IndExs – Index of Exsiccatae). and nomenclature of plant families, genera and species (http:// are based on the “Angiosperm Phylogeny Website” database Tropicos www.mobot.org/mobot/research/apweb/), The Plant List (http://www. and (http://www.tropicos.org/), theplantlist.org). follows the principles circumscribed in part 1 (Braun follows the principles circumscribed in 2013). Ascomycota ( P. viburni-erosi P. , Altingiaceae KRURUOLFHQVRU EXWQRWLQDQ\ZD\WKDWVXJJHVWVWKDWWKH\HQG WFKDUDFWHUVZHUHSXEOLVKHGLQ , C. sambucicola 3 ) . 2013); . 2013); . 2015). . 2015). sp. nov., sp. nov., Cercospora blepharidicola et al et al Aizoaceae , nom. nov., nom. nov., Aizoaceae , Pseudocercospora cyathulae Mycosphaerellaceae and allied genera, and allied genera, (Braun Mycosphaerellaceae Mycosphaerellaceae P. viburnicola P. Adoxaceae , . The structure of part 4 The structure of part 4 . Mycosphaerellaceae Adoxaceae comb. nov., comb. nov., C. justiciigena , var. nov., nov., var. Poaceae , and Chiharu Nakashima , and Chiharu Cercospora 2 ) and comprise a very large ) and comprise a very large Amaranthaceae sp. nov., sp. nov., DGLVFXVVLRQRIWD[RQRPLFDOO\UHOHYDQ .), belonging to the Actinidiaceae to , . 2014); and part three with a treatment . 2014); and part three with a treatment . 2013) with the aim of working towards . 2013) with the aim of working towards Amaranthaceae Actinidiaceae et al et al viburni-sargentii Ascomycota
/HVLRQVIRUPHGDVEURZQOHDIVSRWVZLWKLQGH¿QLWHPDUJLQ RQAsystasia spp...... C. asystasiana ARTICLE Leasion visible as greyish white leaf spots; on other hosts ...... 16
&RQLGLRJHQRXVORFL±PZLGH RQJusticia adhatoda ...... C. adhatodae &RQLGLRJHQRXVORFLVRPHZKDWQDUURZHU±PZLGH RQStrobilathes ...... C. strobilanthis
6WURPDWDZHOOGHYHORSHGWRPGLDP RQBarleria and Blepharis spp...... 18 6WURPDWDODFNLQJRUVPDOO±PGLDP RQRWKHUKRVWV ...... 19
&RQLGLRSKRUHVORQJ±P RQBarleria spp...... C. barleriicola &RQLGLRSKRUHVPXFKVKRUWHU±P RQBlepharis spp...... C. blepharidicola
&RQLGLDYHU\QDUURZDERXW±P RQRuellia spp...... C. ruellina &RQLGLDZLGHU±P RQRWKHUKRVWV...... 20
&RQLGLRSKRUHV±PORQJ RQJusticia and Rhytoglossa spp...... C. diantherae &RQLGLRSKRUHVORQJHUWRP RQRWKHUKRVWV ...... 21
21 (20) Leaf spots yellowish brown to dark brown, later greyish white with dark border, 0.5–4 mm diam; onPeristrophe spp...... C. peristrophes Leaf spots dark brown to black, vein-limited, 3–10 mm diam; on Thunbergia ...... C. thunbergiigena
Tabular key to Cercospora species on Acanthaceae according to host genera
Acanthus A single species ...... C. acanthi
Adhatoda, see Justicia
Andrographis &RQLGLDDFLFXODUEDVHWUXQFDWH±î±P...... C. andrographidicola Conidia acicular to obclavate, base truncate to obconically truncate, shorter and narrower, ±î±P...... C. andrographidis
Asystasia A single species ...... C. asystasiana
Barleria A single species ...... C. barleriicola
Blepharis A single species ...... C. blepharidicola
Crossandra A single species ...... C. crossandrae
Hygrophila A single species ...... C. hygrophilae
Justicia &RQLGLDLQFKDLQV±î±P ± ± ± VHSWDWHK\DOLQH onJusticia adhatoda [Adhatoda vasica] ...... Cercospora justiciae-adhatodae Conidia formed singly, much longer and pluriseptate ...... 2
6WURPDWDODUJH±PGLDP FRQLGLRSKRUHVVKRUW±î±P±VHSWDWH FRQLGLDVKRUW DFLFXODUVXEF\OLQGULFDO±î±P±VHSWDWH...... C. balaghatensis 6WURPDWDODFNLQJRUVPDOOHU±PGLDP FRQLGLRSKRUHVPXFKORQJHU± P SOXULVHSWDWHWKURXJKRXW FRQLGLDORQJHU±PSOXULVHSWDWH ...... 3
VOLUME 6 · NO. 2 375 376 ARTICLE FRQLGLD±PORQJ RQ 6WURPDWDVPDOOHU±PGLDP FRQLGLRSKRUHVPXFKVKRUWHU on ...... (recordsfrom Africa and Asia rareandunproven) 6WURPDWD±PGLDP FRQLGLRSKRUHVORQJDQGDVHSWDWH &RQLGLDQDUURZHU±î±P RQ &RQLGLD±î±P RQ 3 Conidiaaciculartoobclavate-cylindrical,basetruncateobconically ...... 4 (3) &RQLGLDDFLFXODUQDUURZHU±î±PEDVHWUXQFDWH Cercospora acanthi-longifolii base truncate to somewhat obconically truncate ...... basetruncatetosomewhatobconically &RQLGLDDFLFXODUWRVRPHZKDWREFODYDWH±î ± ±P Thunbergia A...... singlespecies Strobilanthes A...... singlespecies Rungia A...... singlespecies Ruellia A...... singlespecies Rhytiglossa A...... singlespecies Peristrophe A...... singlespecies Pachystachys A...... singlespecies Lepidagathis Cercosporella compacta Synonyms (Fig. 1) Cercospora acanthi Cercospora specieson Acanthaceae ...... mixedwithshorterobclavate-cylindricalconidia Conidiaconsistentlyacicular, basetruncate,oratleastlongerconidiaacicular, 3 (2) u.13 (BUCM)]. Jul. 1937 Distr. Severin,nearCazane,on 0HP6HFĠùWL Mycoth. Ital.1580,e.g.BPI420644]. Mycoth. Ital.1580](B700016201); Padova, botanicalgarden,Jun.1904, [ no. 2273(1876). Exs. Micol. ‘FloraVeneta Critt.’ lectotype oii ossetyaiua,bs rnae ...... Conidia consistentlyacicular, basetruncate oii blvt-yidia ihocnclytuct ae cclrcndalcig ...... Conidia obclavate-cylindricalwithobconicallytruncatebase,acicularconidialacking , EdnNov., Ser. Sec.,Cent.3(Resp.23), : Cercosporellaacanthi (designatedbyCrous&Braun2003): Justicia betonica ., ser. 3, Traverso, Pass., inRabenh., 15 : 484(1941)[ 6ăYXO 6DQGX I : 93(1899). sa(ni) ...... (India) , Asia (Pass.)D.Sacc., Hedwigia Acanthus longifolius Justicia spicigera isolectotypes Justicia adhatodae holotype A. Pigal 43 Acad. Române : 422(1904) Justicia Fungi Eur. : Romania: [P. Sacc., : Sacc., Suppl. Italy spp.,commoninNorth America Braun , 16 , Central ...... America (Guatemala) : [ Adhatoda vasica et al conidiation, scattered. punctiform, darkbrown,later greyishwhitebyabundant EURZQ occasionally somewhatraised. JUH\LVK ¿QDOO\ EURZQ to greyishwhite,usuallywith anarrowdarkerborder, RFKUDFHRXV WR \HOORZLVK DQGODUJHUVRPHWLPHV]RQDWHDW¿UVWSDOHJUHHQLVKODWHU FRQÀXHQW VRPHWLPHV GLDP PP ± DQJXODULUUHJXODU WR Description Ital. 191,1580. Exsiccatae Illustration Crous &Braun(2003:40). Lindau (1910:133),Chupp(1954:21),Braun(1995:100), Literature . ±î±P FRQLGLD±PORQJ ...... ±î±P±VHSWDWH : Saccardo(1886:448;1906:562;1972:1367), &KXSS ¿J : Rabenh.,FungiEur. Exs.2273.D.Sacc.,Mycoth. ,Ai Ida ...... (India) ], Asia : Leaf spots ...... Mycelium amphigenous, circular, subcircular Caespituli internal...... 5 ...... 4 ...... C. thunbergiigena C. lepidagathidis .... C. thunbergiana C. jacobiniicola C. peristrophes C. strobilanthis Stromata C. justiciigena C. justiciicola C. justiciicola C. justiciicola amphigenous, C. diantherae C. diantherae C. adhatodae IMA FUNGUS C. ruellina .....6 ...... well- Cercosporoid fungi 4
F\OLQGULFDO VWUDLJKW WR FXUYHG ± î ± P UDUHO\ ARTICLE longer, shorter conidia 1–5-septate, longer ones indistinctly pluriseptate, hyaline, thin-walled, smooth, apex pointed, base truncate to short obconically truncate, (1.5–)2–2.5(–3) PZLGHKLODWKLFNHQHGDQGGDUNHQHG
Lectotype (designated here, MycoBank, MBT202779): Italy: Parma, Botanical Garden, on Acanthus spinosus, Jul. 1875, G. Passerini [Rabenh., Fungi Eur. Exs. 2273] (HAL). Isolectotypes: e.g., B, BPI 432370, HBG, LE 40368.
Host range and distribution: On Acanthus (hungaricus [balcanicus], mollis [longifolius, niger], spinosus [spinosissimus], Acanthus spp., Ruttya fruticosa), Acanthaceae, Africa (Algeria, Ethiopia), Europe (Germany, Hungary, Italy, Romania).
Notes 7KH ¿UVW YDOLG GHVFULSWLRQ RI Cercospora acanthi dates back to 1876 (in Rabenh., Fungi Eur. Exs. 2273), with a brief description on the label. The secondary description in “Hedwigia 16: 123 (1877)”, usually cited as the original description, is younger; this is a consequence of Saccardo not accepting publication in exsiccate labels as acceptable. This species is a true Cercospora readily distinguishable from the C. apii s. lat. complex by its short, aseptate or sparingly septate, pale conidiophores and acicular to obclavate conidia. Records of this species on Peristrophe bicalyculata from Myanmar (Thaung 1984) are very doubtful and excluded.
Cercospora adhatodae S. Chowdhury, Lloydia 18: 84 “1955” (1956). (Fig. 2)
Literature: Vasudeva (1963: 32), Crous & Braun (2003: 44), Kamal (2010: 13).
Illustration: &KRZGKXU\ ¿J
Description: Leaf spots amphigenous, 0.5–8 mm diam, RIWHQ FRQÀXHQW IRUPLQJ ODUJHU SDWFKHV VXEFLUFXODU WR Fig. 1. Cercospora acanthi (Rabenh., Fungi Eur. Exs. 2273, HAL, irregularly shaped, greyish white. Caespituli amphigenous, lectotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. ¿QH GDUN Mycelium internal. Stromata lacking or small, %DU P substomatal, brown. Conidiophores in small, divergent fascicles, arising from internal hyphae or stromatic hyphal aggregations, through stomata or erumpent, erect, straight, GHYHORSHG±PGLDPVXEVWRPDWDOWRLPPHUVHGODUJH VXEF\OLQGULFDOWRJHQLFXODWHLQWKHXSSHUKDOI± î± stromata often rupturing the stomata, somewhat erumpent, P±VHSWDWHEURZQWRGDUNEURZQSDOHUWRZDUGVWKHWLS pale, subhyaline to yellowish brown, later dark brown, thin-walled, smooth; conidiogenous cells integrated, terminal, FRPSRVHGRIVZROOHQK\SKDOFHOOV±PGLDPFLUFXODU FRQLGLRJHQRXV ORFL FRQVSLFXRXV DERXW ± P ZLGH to somewhat irregular in shape. Conidiophores in small thickened and darkened. Conidia solitary, acicular, straight to large fascicles, dense, arising from stromata, through WRFXUYHG±î±P±VHSWDWHK\DOLQHWKLQ stomata or erumpent, erect, straight to curved or somewhat ZDOOHGVPRRWKDSH[SRLQWHGEDVHWUXQFDWHDERXW±P JHQLFXODWHVLQXRXV XQEUDQFKHG ± ± î ± P wide, hila thickened and darkened. 0–2-septate, subhyaline, yellowish to pale olivaceous, paler towards the tip, somewhat darker in mass, thin- Holotype: India: Asom (Assam): Kokilamukh, on Justicia walled, smooth; conidiogenous cells integrated, terminal adhatoda, 12 Feb. 1951, S. Chowdhury (HCIO). or conidiophores reduced to conidiogenous cells, 10–30 PORQJPRVWO\ZLWKDVLQJOHRUWZRRFFDVLRQDOO\VHYHUDO Host range and distribution: On Justicia adhatoda [Adhatoda FRQVSLFXRXVFRQLGLRJHQRXVORFL±PGLDPConidia vasica], Acanthaceae, Asia (India, Asom, Karnataka, Uttar solitary, acicular, smaller conidia narrowly obclavate- Pradesh, West Bengal).
VOLUME 6 · NO. 2 377 378 ARTICLE Literature Cercospora andrographidicola 293593 and330436). and UttarPradeshhavebeenexamined(K(M)IMI135861, Several IndiancollectionsfromKarnataka,West Bengal Notes fascicle. Fig. 2. (Fig. 3) “ Chi, andrographicola : This isaspeciesofthe Cercospora adhatodae B. J. S.China Agric. Univ : Chi(1994:94). Conidiophore tips. ”. C. (K(M) IMI135861). &RQLGLD%DU P Cercospora apiis.lat. . 11 S.Q. Chen&P.K. (3): 61(1990);as A. Conidiophore complex. Braun et al 86). Fig. 3. ± ±VSDHRLDHX FRQLGLRJHQRXV conidiogenous locus, thickenedanddarkened. ROLYDFHRXV ±VHSWDWH cells integrated,terminal,usually withasingleterminal î P ± ± XQEUDQFKHG QRQJHQLFXODWH VXEF\OLQGULFDO loose fascicles,4–15,arising fromstromata,erect,straight, EURZQ GLDP P ± ZHOOGHYHORSHG below. to irregular, centrepale,marginindistinct,greyishgreen Description Illustration . A. Conidiophore fascicle. Cercospora andrographidicola Caespituli &KL ¿J : Leaf spots amphigenous. B. amphigenous, circular, elliptical &RQLGLXP%DU P Mycelium EDVHGRQ&KL ¿J internal. Conidiophores IMA FUNGUS Stromata Conidia in Cercosporoid fungi 4
VROLWDU\ DFLFXODU VWUDLJKW WR VRPHZKDW FXUYHG ± î ARTICLE ±PK\DOLQHWKLQZDOOHGVPRRWKDSH[SRLQWHGEDVH truncate, hila thickened and darkened.
Holotype: China: Guangdong: Gaoyao, on Andrographis paniculata, Oct. 1986, S. G. Chen 123 (Hb. S. China Agric. Univ., Guangzhou).
Host range and distribution: On Andrographis paniculata, Acanthaceae, Asia (China).
Notes: Belonging to the Cercospora apii s. lat. complex, but type material was not available for a re-examination. To-anun et al. (2011) described and illustrated “C. andrographidicola” on Andrographis paniculata from Thailand. This material is, however, morphologically dinstinct by lacking or small VWURPDWDXSWRPGLDPVKRUWHUJHQLFXODWHFRQLGLRSKRUHV and much shorter, acicular to obclavate conidia, and rather belongs to C. andrographidis.
Cercospora andrographidis Thirum. & Govindu, Sydowia 7: 310 (1953). (Fig. 4)
Literature: Vasudeva (1963: 36), Braun & Crous (2003: 57), Kamal (2010: 16), To-anun et al. (2011: 30), as “C. andrographidicola”.
Illustrations: Thirumalachar & Govindu (1953: plate 9, ¿JV ± 7RDQXQ et al ¿J DV ³C. andrographidicola”.
Description: Leaf spots amphigenous, circular to somewhat DQJXODULUUHJXODU ± PP GLDP DW ¿UVW GLQJO\ JUHHQLVK olivaceous or brown, later greyish white, surrounded by a narrow brown margin or marginal line, slightly raised or margin broader, pinkish brown. Caespituli amphigenous, mainly epiphyllous, not very conspicuous. Mycelium internal. Stromata lacking or inconspicuous and small, only a few VZROOHQK\SKDOFHOOV±PGLDPXVXDOO\LQWUDHSLGHUPDO LI SUHVHQW WR P GLDP EURZQ FHOOV ± P GLDP ZDOO slightly thickened. Conidiophores solitary or in small divergent fascicles, arising from internal hyphae or aggregations of swollen hyphal cells, erumpent, erect, straight to geniculate, Fig. 4. Cercospora andrographidis (BPI 432654). A. Conidiophore XQEUDQFKHG ± î ± P ± VHSWDWH OLJKW WR fascicle. B. Solitary conidiophore. C.&RQLGLD%DU P medium dark brown, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal, occasionally LQWHUFDODU\ ± P ORQJ V\PSRGLDO FRQLGLRJHQRXV ORFL Note: The conidia of this species were described to be FRQVSLFXRXVWKLFNHQHGDQGGDUNHQHG±PGLDPConidia “acicular” with obconically truncate base, i.e. they are, solitary, acicular to somewhat obclavate, straight to curved, at least partly, obclavate as depicted in Thirumalachar & ±î±P±VHSWDWHK\DOLQHWKLQZDOOHGVPRRWK *RYLQGX SODWH 9, ¿J 7KLV FRXOG EH FRQ¿UPHG apex subacute, base truncate to somewhat obconically on the base of an examined collection from India (West WUXQFDWH±PZLGHKLODWKLFNHQHGDQGGDUNHQHG Bengal, Midnapur, Daspur, 20 Mar. 1967, M. Mandal, BPI 432654). To-anun et al. (2011) described and illustrated Holotype: India: Bihar: Patna, on Andrographis sp., 20 Jan. “C. andrographidicola” on Andrographis paniculata from 1951, M. J. Thirumalachar (not traced). Thailand. This material, characterised by lacking or small stromata and acicular to obclavate conidia, is quite distinct Host range and distribution: On Andrographis (paniculata, from true C. andrographidicola and rather belongs to C. Andrographis sp.), Acanthaceae, Asia (India, Bihar, Andhra andrographidis. An additional sample from India (Daspur, BPI Pradesh, West Bengal; Thailand). 432654) has been examined.
VOLUME 6 · NO. 2 379 380 ARTICLE nemorum and [records of Host rangeanddistribution chelonoides Holotype and darkened. RUVOLJKWO\REFRQLFDOO\WUXQFDWH±PZLGHKLODWKLFNHQHG ±VHSWDWH hyaline, thin-walled,smooth,apexsubacute,basetruncate P ± î ± VLJPRLG VRPHZKDW RU GLDP P solitary, aciculartosomewhatobclavate,straightcurved ± DERXW ORFL GDUNHQHG FRQLGLRJHQRXV DQG ORQJ WKLFNHQHG P ± DERXW LQWHUFDODU\ RU thin-walled, smooth;conidiogenouscellsintegrated,terminal ± ± î±P±VHSWDWHEURZQWRGDUNEURZQ subcylindrical todistinctlygeniculate-sinuous,unbranched, through stomata,erect,straighttosomewhatcurved, internal hyphaeorsmallsubstomatalhyphalaggregations, solitary orinsmalldivergentfascicles,2–5,arisingfrom it iscurrentlyimpossibletoresolve thecladeconcerned. species namesof possible toexcludethatseveral otherhostspeciesandolder hitherto littleknownhostrange ofthistaxon,itisnotyet from sequences basedonNorth American collectionsretrieved malloti internal. Caespituli VFDWWHUHGVXEFLUFXODU±PPGLDPEURZQPDUJLQLQGH¿QLWH Description ¿J Illustration Nakashima Literature (Fig. 5) Cercospora asystasiana tentatively denominatedas gangetica complex. Recordsof Notes – seenotes]. species, including derived fromseveralcollectionsonvariousunrelatedhost from northern Thailand usingmolecularmethods.Sequences of thiscladecomprisingplurivorous material infecting true malloti common (2013: 157). Taxa belongingtothiscladerepresentedthemost are unclearbutmightbelongto 180 (1967). Nguanhom C. C. malloti Mallotus : This speciesispartofthe This : LVQHLWKHUVHWWOHGQRU¿QDOO\SURYHQVLQFHFXOWXUHVDQG cf. is morphologicallypartofthe : Cercospora ), : Yen &Lim(1980:155),CrousBraun(2003:70), Stromata malloti H ¿ H L /LP Fig. 7. Cercospora barleriicola (K(M) IMI 102433). A. Conidiophore fascicles. B. Conidiophores. C.&RQLGLD%DU P Cercospora barleriicola Payak & Thirum., Indian Phytopathol. 2: 191 (1949); as “barlericola” Fig. 6. Cercospora balaghatensis (K(M) IMI 150920a, holotype). A. (Fig. 7) Conidiophore fascicles. B. Conidiophores. C. &RQLGLD%DU P Synonym: Cercospora barleriae-cristatae Govindu & Thirum., Sydowia 10: 273 “1956” (1957) [holotype: India: loose to usually dense fascicles, arising from stromata, erect, Karnataka: Bangalore, on Barleria cristata, 10 Dec. 1953, straight, subcylindrical, barely geniculate, unbranched, about H. C. Govindu (not traced)]. ±î±P±VHSWDWHSDOHROLYDFHRXVWREURZQLVK thin-walled, smooth; conidiogenous cells integrated, Literature: Chupp (1954: 22), Vasudeva (1963: 46–47), terminal, conidiogenous loci conspicuous, thickened and Crous & Braun (2003: 76), Kamal (2010: 21), Meeboon et al. GDUNHQHGDERXW±PGLDPConidia solitary, acicular or (2007a,b), To-anun et al. (2011: 31) VXEF\OLQGULFDO VWUDLJKW WR VOLJKWO\ FXUYHG ± î ± P 0–5-septate, hyaline, thin-walled, smooth, apex subacute or Illustrations *RYLQGX 7KLUXPDODFKDU SODWH9,,,¿J VXEREWXVHEDVHWUXQFDWH±PZLGHKLODWKLFNHQHGDQG 9DVXGHYD ¿J 7RDQXQet al. (2011: 31, darkened. ¿J Holotype: India: Madhya Pradesh: Balaghat, on Justicia sp., Description: Leaf spots amphigenous, subcircular to angular- Acanthaceae, Jun. 1970, S. M. Singh (K(M) IMI 150920a). irregular, 2–8 mm diam, diffuse discolorations, yellowish to dark reddish brown, later pale brownish to greyish brown or Host range and distribution: Only known from the type greyish white with darker border. Caespituli amphigenous, collection. subeffuse to punctiform, dark brown. Mycelium internal. Stromata almost lacking or small to moderately large, 10–60 Notes: A true Cercospora s. str. distinct from C. apii s. lat. P GLDP ROLYDFHRXVEURZQ WR GDUN EURZQ VXEVWRPDWDO by having very large stromata, short, densely fasciculate to immersed. Conidiophores in small to moderately large conidiophores and short, 0–4-septate conidia. fascicles, loose to dense, arising from stromata, through stomata or erumpent, erect, straight, subcylindrical to VOLUME 6 · NO. 2 381 382 ARTICLE Holotype WUXQFDWH±PZLGHKLODWKLFNHQHGDQGGDUNHQHG smooth, apexpointed,basetruncatetosomewhatobconically FXUYHG±î±P±VHSWDWHK\DOLQHWKLQZDOOHG obclavate-cylindrical withobconicallytruncatebase,straightto longer conidiaacicularwithtruncatebase,shorteronesmaybe Basionym (Fig. 8) MycoBank MB814564 Cercospora blepharidicola FRQVSLFXRXV ORFL WKLFNHQHGDQGGDUNHQHG ± ±PGLDP FRQLGLRJHQRXV ORQJ P ± WHUPLQDO to slightlythickened,smooth;conidiogenouscellsintegrated, brown orthroughoutpalertowardsthetip,wallthin PWRSOXULVHSWDWHWKURXJKRXWSDOHWRPHGLXPROLYDFHRXV PRGHUDWHO\JHQLFXODWHVLQXRXVXQEUDQFKHG±î± homonym of Notes collection. Host rangeanddistribution Acanthaceae Division, Toda Khurai,on Holotype truncate orsubtruncate,hilathickenedanddarkened. WR P 10-septate, hyaline,thin-walled,smooth,apexpointed,base ± î ± FXUYHG WR VWUDLJKW DFLFXODU VROLWDU\ cylindrical, cicatrized,thickenedanddarkened. thin-walled, smooth;conidiogenouscellsintegrated,terminal, EDVHDERXW±î±P±VHSWDWHOLJKWROLYDFHRXV VXEF\OLQGULFDOÀH[XRXVXQEUDQFKHGDULVLQJIURPDVZROOHQ olivaceous. internal. amphigenous, effuse, uniformlydistributed. small tolarge,scattered,darkbrown. Description Illustration cryptic speciesareinvolved. remains unclearifasinglepolymorphousspeciesorseveral lat. Barleria Host rangeanddistribution traced). University, on variable have beenexaminedandprovedtobelongasingle on Notes Thailand), West Indies(Jamaica). Karnataka, Maharashtra,MadhyaPradesh,UttarPradesh; Barleria illeg. Mycol. Pl.Pathol. complex. Sincephylogeneticdataarenotyetavailable,it : The name : The : SeveralIndiancollectionsandasamplefromJamaica sp.), Cercospora s.str. (Art. 53.1),non : : Stromata : : Dubey India India : spp. (K(M)IMI102433,163693,226980,265815) Cercospora blepharidis Conidiophores C. blepharidis Leaf spots , Dec.1996, Acanthaceae Barleria cristata : : Uttar Pradesh et al Madhya Pradesh C. blepharidis HOGYOSGWGD GDUN GLDP P WR ZHOOGHYHORSHG 41 ¿J C. blepharidis : 514(2011), as“ amphigenous, mainlyepiphyllous, .K ue AR-58 R. K.Dubey , Asia (India,JammuandKashmir, species belongingtothe Chidd.(Chiddarwar 1960). in loosefascicles,erect,straight, : On : Onlyknownfromthetype , 9Dec.1949, Blepharis maderaspatensis : Varanasi, BaranaresHindu U. Braun, Barleria R.K.Dubey : NorthSagarForest R.K. Dubeyetal., Chidd, 1960. ( cristata nom. nov. M. Payak blepharia (HCIO439724). Conidia et al , Caespituli Mycelium prionitis C. apiis. Conidia solitary, ”, . isa nom. (not Braun J. , , et al Description Illustration Literature Cercospora crossandrae Fig. 8. and darkened. EDVH truncate toshortobconically truncate,probablythickened SRLQWHG DSH[ K\DOLQH ±VHSWDWH P ± ± DERXW î GHVFULSWLRQ RULJLQDO WKH WR DFFRUGLQJ ³¿OLIRUP 4–7-septate, brown. IDVFLFXODWHJHQLFXODWHXQEUDQFKHGDERXW±îP with darkbrownmargin. \HOORZ ERUGHU EURZQ sometimes withconcentricrings. XS GU\LQJ ¿QDOO\ IROLDJH FRQÀXHQW ¿J . Madras Agric. J Madras Agric. A. Cercospora blepharidicola Conidiophore fascicle. : Crous&Braun(2003:143),Kamal(2010:36). : Jaganathan : Leaf spots . Conidia 59 et al circular toirregular, 2–5mmdiam, : 672(1972). Mycelium ¿J B. Jagan., Palan.&Narayans., solitary, aciculartoobclavate (based onDubay &RQLGLD%DU P internal. Colonies Conidiophores et al IMA FUNGUS white, later . 2011: 516, Cercosporoid fungi 4 Holotype: India: Tamil Nadu: Coimbatore, College Orchard ARTICLE of the Tamil Nadu Agricultural University, on Crossandra infundibuliformis, Acanthaceae (T.N. Agric. Univ., Coimbatore, Pl. Pathol. Herb. No. 231). Host range and distribution: Only known from the type collection. Notes: It is unknown if type material of this species is maintained. It was not available for examination. Based on the original description and illustration, we suppose that this species belongs to Cercospora s. str. although details of the conidiogenous loci and hila were not described. Cercospora diantherae Ellis & Kellerm., J. Mycol. 1: 2 (1885). (Fig. 9) Synonym: Cercospora jacobinae Mendoza, Philipp. J. Sci. 75: 169 (1941) [holotype: Philippines: Manila, on Justicia carnea, Mendoza, no. 7124 (not traced)]. Literature: Saccardo (1886: 448), Chupp (1954: 23), Crous & Braun (2003: 159). Illustration &KXSS ¿J Exsiccatae: Ellis & Everh., Fungi Columb. 695. Ellis & Everh., N. Amer. Fungi 1750. Kellerm. & Swingle, Kansas Fungi 33. Roum., Fungi Sel. Gall. Exs. 5190. Description: Leaf spots circular to somewhat angular- LUUHJXODU±PPGLDPRFFDVLRQDOO\FRQÀXHQWDQGODUJHU slightly zonate, centre greyish white with narrow to often broad brown border. Caespituli amphigenous, dark brown. Mycelium internal. Stromata ODFNLQJ WR VPDOO ± P diam, composed of a few swollen hyphal cells, brown, substomatal to intraepidermal. Conidiophores in small to moderately large fascicles, 2–15, divergent to moderately dense, arising from internal hyphae or stromatic hyphal aggregations, through stomata or erumpent, erect, straight, subcylindrical to sinuous or somewhat geniculate, XQEUDQFKHG RU UDUHO\ EUDQFKHG ± î ± P aseptate to pluriseptate throughout, pale to medium brown, Fig. 9. Cercospora diantherae (CUP 39680, lectotype). A. wall thin to slightly thickened, smooth; conidiogenous cells Conidiophore fascicles. B. Conidiophores, C. &RQLGLD%DU P LQWHJUDWHGWHUPLQDO±PORQJZLWKDVLQJOHWRVHYHUDO FRQLGLRJHQRXV ORFL ± P GLDP Conidia solitary, acicular or subacicular, straight to curved or occasionally Delaware, Florida, Illinois, Indiana, Kansas, Maryland, VRPHZKDWVLJPRLG± ± î±PSOXULVHSWDWH Missouri, Oklahoma, Texas, Washington, West Virginia), GLVWDQFH EHWZHHQ VHSWD ± P K\DOLQH WKLQZDOOHG South America (Venezuela). smooth, apex acute to subobtuse, base usually truncate, RFFDVLRQDOO\VRPHZKDWREFRQLFDOO\WUXQFDWH±PZLGH Notes: This species belongs to the Cercospora apii (s. hila thickened and darkened. lat.) complex. Type material (collected in Sep. 1884) is not preserved at NY. There are numerous topotypes from 1886, Lectotype (designated here, MycoBank, MBT202780): USA: e.g. B; BPI 435404, 435692, 435700, 435701; NY 270701. Kansas: on Justicia americana, Sep. 1884, W. A. Kellerman However, the type material collected in Sep. 1884 was traced (CUP 39680). Isolectotype: PH 1798. in CUP and FH. The CUP material is designated as lectotype. Records of C. diantherae on Jacobinia spp. are doubtful. The Host range and distribution:On Justicia (americana, carnea, hosts concerned probably refer to Justicia spp. in the current ovata, Justicia sp.), Rhytiglossa humilis [Justicia humilis], sense. Acanthaceae, Asia (Philippines), North America (USA, VOLUME 6 · NO. 2 383 384 ARTICLE Description Illustration Literature (Fig. 10) Cercospora hygrophilae Conidiophores. Fig. 10. or immersed, globosetosomewhatirregular, brown. RIVRPHVZROOHQK\SKDOFHOOV±PGLDPVXEVWRPDWDO internal. blackish. oblong, ovaltoirregular, 5–15mmdiam, darkbrownto Acad. Sci Stromata : Crous&Braun(2003:222),Kamal(2010:52). 3RQQDSSD ¿J Caespituli Cercospora hygrophilae : Leaf spots B. ., sect.B, &RQLGLD%DU P lacking orsmall,formedasaggregations PKJQX Q GDUN ¿QH DPSKLJHQRXV 67 amphigenous, oftenmarginal, : 31(1968). Ponnappa, (K(M) IMI126157). Proc. Indian Mycelium Braun A. et al Holotype WKLFNHQHG and darkened. KLOD ZLGH P ± WUXQFDWH EDVH SRLQWHG DSH[ VPRRWK WKLQZDOOHG K\DOLQH ±VHSWDWH P ± ± î Conidia ORFLFRQVSLFXRXVWKLFNHQHGDQGGDUNHQHG±PGLDP FRQLGLRJHQRXV ORQJ P ± DERXW WHUPLQDO LQWHJUDWHG EURZQ WKURXJKRXW SOXULVHSWDWH WR wall somewhatthickened,smooth;conidiogenouscells P ± î ± subcylindrical tosomewhatgeniculateabove,unbranched, aggegations, erumpentorthroughstomata,erect,straight, 2–15, arisingfrominternalhyphaeorstromatichyphal Conidiophores Mycelium amphigenous, mainlyhypophyllous,punctiform,brown. with somewhatraisednarrowborderline. irregular, 2–6mmdiam,lead-colouredtoblackish, Description Literature Cercospora jacobiniicola 42 [ Holotype darkened. RFFDVLRQDOO\WUXQFDWH±PZLGHEDUHO\WKLFNHQHGDQG apex obtusetosubacute,baseshortobconicallytruncate, to verypalegreenishorolivaceous,thin-walled,smooth, ± ± î ± ± ± P± ± VHSWDWHK\DOLQH to obclavate-subcylindrical,straightslightlycurved, darkened (paracercosporoid). FLUFOHVDERXWPGLDPRQO\PDUJLQVOLJKWO\WKLFNHQHGDQG long, conidiogenouslociconspicuous,formedasminute FHOOVRUFRQLGLRJHQRXVFHOOVLQWHJUDWHGWHUPLQDO±P walled, smooth;conidiophoresreducedtoconidiogenous ± î P±VHSWDWHVXEK\DOLQHWRSDOHROLYDFHRXVEURZQWKLQ ± XQEUDQFKHG JHQLFXODWHVLQXRXV VOLJKWO\ EURZQ WR through stomata,erect,straight,subcylindrical-conicalto ROLYDFHRXV GDUN divergent todense,arisingfrominternalhyphaeorstromata, GLDP P Conidiophores ± DERXW (Fig. 11) [ Host rangeanddistribution auriculata belonging tothe Notes Bengal). [ conidiophores andconidia. GHVFULSWLRQ RULJLQDO is misleadingandpossiblybasedonnotfullydeveloped 7KH P ± î ± ORQJ YHU\ DUH Jacobinia spicigera spinosa quadrivalvis . (CUP 40082). 1 : 174(1950);as“ : This speciesisatypicalmemberof solitary, acicular, straight,curvedtosigmoid,50–250 , : : : Chupp(1954:24). , 21Feb.1967, Guatemala internal. India Asteracantha longifolia : ]), Leaf spots Acanthaceae in divergenttomoderatelydensefascicles, : C. apiis.lat. in smalltomoderatelylargefascicles, Karnataka Stromata ], Acanthaceae : Chimaltenango,on jacobinicola K. M.Ponnappa : Bannerghatta,on amphigenous, subcircularto , Asia (India,Karnataka,West complex, buttheconidiophores lacking orsmall,substomatal, : On A.S. Mull.&Chupp, Conidia , 2Oct.1941, ], ”. Hygrophila polysperma, ringens solitary, cylindrical (K(M) IMI126157). Cercospora s.str. Justicia spicigera IMA FUNGUS A. S.Muller Hygrophila ( Caespituli auriculata Ceiba Cercosporoid fungi 4 ARTICLE Fig. 12a. Cercospora justiciae-adhatodae (BPI 1103659, holotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 P Fig. 11. Cercospora jacobiniicola (CUP 40082, holotype). A. Description: Leaf spots amphigenous, circular, subcircular Conidiophore fascicles. B. Conidiophore, C. &RQLGLD%DU P WR VOLJKWO\ DQJXODULUUHJXODU± PP GLDP DW ¿UVWEURZQ but soon turning greyish white to white, margin narrow, somewhat raised, dark, brown, dark violet to almost Host range and distribution: Only known from the type blackish. Caespituli amphigenous, punctiform, scattered to collection. dense, brown to dark brown. Mycelium internal. Stromata ODFNLQJ RU DOPRVW VR WR ZHOOGHYHORSHG ± P GLDP Note: Tentatively maintained in Cercospora. The conidia VXEVWRPDWDO WR LQWUDHSLGHUPDO EURZQ FHOOV ± P GLDP are colourless or almost so, the conidiogenous loci are ZDOO DW ¿UVW WKLQ ODWHU VOLJKWO\ WKLFNHQHG Conidiophores in conspicuous, but minute, ca P ZLGH DQG VRPHZKDW small, loose to moderately large and dense fascicles, arising paracercospora-like. Cultures and results based on molecular from substomatal or intraepidermal hyphae or stromata, sequence analyses are necessary to resolve the true generic emerging through stomata or erumpent, erect, straight to DI¿QLW\RIWKLVVSHFLHV curved, subcylindrical or somewhat attenuated towards the WLSWRPRGHUDWHO\JHQLFXODWHVLQXRXVXQEUDQFKHG±î Cercospora justiciae-adhatodae U. Braun, sp. nov. ±P±VHSWDWHSDOHROLYDFHRXVWRROLYDFHRXVEURZQ MycoBank MB814556 paler towards the tip, thin-walled, smooth; conidiogenous (Fig. 12a,b) cells integrated, terminal or conidiophores reduced to FRQLGLRJHQRXVFHOOV±PORQJSUROLIHUDWLRQV\PSRGLDO Diagnosis: Differs from Cercospora adhatodae and all FRQLGLRJHQRXV ORFL FRQVSLFXRXV ± P ZLGH VRPHZKDW other Cercospora species on hosts of the Acanthaceae in thickened and darkened. Conidia solitary and catenate, IRUPLQJ VPDOO FDWHQDWH FRQLGLD ± ± ± î ± P in simple or occasionally branched chains, narrowly (0–)1–3(–4)-septate. F\OLQGULFDOIXVLIRUP VKRUW REFODYDWH ± ± ± î ± VOLUME 6 · NO. 2 385 386 ARTICLE Holotype slightly thickenedanddarkened. VPRRWK VXEWUXQFDWHWRVKRUWREFRQLFDOO\WUXQFDWH±PZLGHKLOD WKLQZDOOHG K\DOLQH apex subacute,subobtuseorshortconicallytruncate,base ± ± ± VHSWDWH P Symptoms onleaves. Fig. 12b. Phaeoramularia inclined toputthisspecies in hosts belongingtothe species fromallother formed inchains. These characteristicsdistinguishthenew by muchshorter, narrower, usually 1–3-septateconidia the latterspecies,whichbelongsto as Notes collection. Host rangeanddistribution 1103659). 10 Jan.1969, Hills, on Cercospora adhatodae : Type materialofthisspecies wasoriginallydeposited Justicia adhatoda : Cercospora justiciae-adhatodae India V. S.Seshadri ), butcatenate, colourlessconidiaarerather : Karnataka Acanthaceae Cercospora [ Adhatoda vasica , butitisquitedistinctfrom : ChikkaballapurDistrict,Nandi & : Onlyknownfromthetype K. A. Lucy Channnamma Lucy K. A. Passalora s.lat speciesdescribedon . Previouslyonewere (BPI 1103659, holotype). C. apii ], Acanthaceae . (including complex, (BPI Braun , et al Basionym (Fig. 13) MycoBank MB814565 Cercospora justiciigena ¿J± Fig. 13. although culturesandmoleculardataarenotyetavailable. new speciesis,atleastfortheinterim,placedin hyphomycetes (seeBraun shown inthecourseofphylogeneticstudiescercosporoid in favourof P Literature . Cercospora macrospora Karan &Manohar., Cercosporajusticiigena : Crous&Braun (2003:40),Kamal(2010:12). : Cercosporaacanthacearum Cercospora s.str. A. Conidiophore fascicle. Botanique et al Bres., 1886. (based onKaran&Manoharachary U. Braun, . 2013). Therefore, thepresent , whichhasrecentlybeen 7 : 159“1976”(1978),non nom. nov. var. B. Conidia. Bar=10 IMA FUNGUS macrospora Cercospora Cercosporoid fungi 4 Illustration .DUDQ 0DQRKDUDFKDU\ ¿J± ARTICLE Description: Leaf spots epiphyllous, zonate, 4–6 mm diam, white, surrounded by a pinkish border. Mycelium internal. Stromata DERXW ± P GLDP GDUN EURZQ pseudoparenchymatic. Conidiophores fasciculate, arising from stromata, erect, straight to curved, unbranched, JHQLFXODWH LQ WKH XSSHU KDOI DERXW ± î ± P aseptate, brown, colourless towards the tip; conidiophores reduced to conidiogenous cells, apex rounded with dark annular conidiogenous loci. Conidia solitary, obclavate, VWUDLJKWDERXW±î±PSOXULVHSWDWHK\DOLQHWKLQ walled, smooth, tips pointed, base short obconically truncate, hila somewhat thickened and darkened. Holotype: India: Andhra Pradesh: Hyderabad, University, green house, on Justicia betonica, Acanthaceae, 10 Dec. 1964, D. Karan & C. Manoharachary (HY 197). Host range and distribution: Only known from the type collection. Notes: This fungus was introduced as variety of C. acanthacearum, although quite different from the latter species by its much longer conidiophores and much wider conidia. Owing to the description and illustration of conspicuous, dark conidiogenous loci and hila as well as colourless conidia formed singly, C. acanthacearum var. macrospora is undoubtedly a true Cercospora (s. str.) species. The generic DI¿QLW\RIC. acanthacearum is unclear, but this species might rather be a member of Pseudocercospora. Cercospora justiciicola F.L. Tai, Lloydia 11: 47 (1948); as “justiciaecola”. (Fig. 14) Literature: Chupp (1954: 24), Ellis (1976: 244), Guo et al. (2005: 17), Kamal (2010: 55). Fig. 14. Cercospora justiciicola (based on Guo et al ¿J A. Conidiophore fascicle. B. &RQLGLD%DU P Illustrations 7DL ¿J (OOLV ¿J$ Guo et al ¿J Holotype: China: Sichuan: Chengtu, on Justicia sp., 15 Oct. Description: Leaf spots amphigenous, subcircular, 3–10 mm 1944, H. C. Lin (HMAS 12127). GLDP \HOORZLVK RFKUDFHRXV SDOH EURZQ WR ¿QDOO\ JUH\LVK white, often somewhat zonate, sometimes with yellowish Host range and distribution: On Justicia (betonica, carnea halo. Caespituli amphigenous, punctiform, brown. Mycelium [Jacobinia obtusior], diffusa,ÀDYD, gendarussa, procumbens, internal. Stromata DOPRVW ODFNLQJ WR GHYHORSHG ± P simplex, Justicia sp.), Pachystachys lutea [Justicia lutea], diam, brown, substomatal to immersed. Conidiophores in Rungia (pectinata [Justicia pectinata, 5XQJLD SDUYLÀRUD], small to moderately large fascicles, divergent, arising from repens [Justicia repens], Rungia sp.), Acanthaceae, Africa stromata, through stomata or erumpent, erect, straight, (Guinea, Mauritius), Asia (Brunei, China; India, Andhra subcylindrical to usually geniculate-sinuous, unbranched, Pradesh, Madhyar Pradesh, Uttar Pradesh; Nepal), North ± î ± P ±VHSWDWH ROLYDFHRXVEURZQ WKLQ America (USA, Florida). walled, smooth; conidiogenous cells integrated, terminal WR LQWHUFDODU\ DERXW ± P ORQJ FRQLGLRJHQRXV ORFL Notes: A true Cercospora s. str. FORVHWRRUFRQVSHFL¿FZLWK WKLFNHQHG DQG GDUNHQHG ± P GLDP Conidia solitary, C. apii s. lat. Records of C. justiciicola on Asystasia gangetica acicular to obclavate-subcylindrical, straight to curved, [coromandeliana] (Crous & Braun 2003) are unclear, doubtful ± ± ± î± ± P ± ± ± VHSWDWH and might belong to C. asystasiana hyaline, thin-walled, apex subacute, base truncate to short REFRQLFDOO\ WUXQFDWH ± P ZLGH KLOD WKLFNHQHG DQG Cercospora lepidagathidis Govindu & Thirum., darkened. Sydowia 9: 222 (1955). VOLUME 6 · NO. 2 387 388 ARTICLE Synonyms (Fig. 15) Cercospora peristrophes Holotype truncate, hilathickened. walled, smooth,apexpointed,baseshortobconically WKLQ K\DOLQH ±VHSWDWH P ± î ± FXUYHG Conidia intercalary, withseveralconspicuousconidiogenousloci. tips colourless;conidiogenouscellsintegrated,terminalor EURZQ WR ROLYDFHRXV SDOH ±VHSWDWH P ± î erect, mostlyunbranched,stronglygeniculate-sinuous,33– brown. lacking orsmall,composedofafewswollenhyphalcells, brown. 4–7 mmdiam,centregreyishwhite,marginpinkishor Description 9DVXGHYD ¿J Illustrations 248), Kamal(2010:59). Literature Host rangeanddistribution traced). Lepidagathis cuspidata Cercospora peristrophigena true description andillustration,thisspeciesisundoubtedlya could neitherbetracedatBPInorIMI.Basedontheoriginal deposited atBPI,HCIOandIMI.However, typecollections (1955) mentionedthattypematerialofthisspecieswas Notes Maharashtra, UttarPradesh;Pakistan). incurva Pseudocercospora lepidagathidis. alopecuroidea Indies (PuertoRico,andVirgin Islands)on 9DVXGHYD ¿J &KDXGKDU\ Illustrations 316), Kamal(2010:73). Literature Description ¿J 5DR Sydowia (1999), al bicalyculata Uttar Pradesh et al [ 41989); bicalyculata Peristrophe holotype Cercospora s.str. ., : Intheoriginalpublication,Govindu& Thirumalachar Caespituli Sugarcane Pathology Conidiophores ., : [ solitary, obclavate-cylindrical,straighttoslightly : Vasudeva (1963:133–134),Crous&Braun(2003: : Vasudeva (1963:162),Crous&Braun(2003: hyalina : J. LivingWorld : *RYLQGX 7KLUXPDODFKDU SODWH¿J 7KLUXPDODFKDU *RYLQGX SO¿JV± nom. inval : India isotype Leaf spots : Leaf spots Cercospora peristrophigena 7 India DUHUHVXOWVRIPLVLGHQWL¿FDWLRQVDQGEHORQJWR : 47(1953). , Nov. 1993, et al ]), : amphigenous. : Nichlaul,Maharajganj,on Acanthaceae : GPU5070];as“ : Karnataka ¿J Uttar Pradesh . (Art.39.1)and , 20May1953, amphigenous, circularorsubcircular, in well-developed,densefascicles, amphigenous, circular tosomewhat Records ofthisspeciesfromWest , Nov. 1993,S.Narayan(HCIO S. Narayan 2 : On , Vol. 1: S. Narayan , Mysore,Bababudans,on : 38(1995)[ Mycelium , Asia (India,Karnataka, : Nichlaul,Maharajganj,on Thirum. &Govindu, Lepidagathis peristrophegena nom. illeg Fungal Diseases (HCIO41989)]. H. C.Govindu internal. et al. R.K. Chaudhary holotype Lepidagathis ( , inRao et al . (Art.53.1) Peristrophe cuspidata Stromata : . (1995: ”. India (not : 82 Braun et : , et al Conidiophore fascicle. Fig. 15. P surrounded byadarkerborder. WR JUH\ GLQJ\ JUH\LVKZKLWHPDUJLQLQGH¿QLWHPDUJLQDOVOLJKWO\UDLVHGRU ¿QDOO\ FHQWUH EURZQ GDUN WR EURZQ EURZQ angular-irregular, scattered,0.5–8mmdiam,yellowish ± PHWW DH ROLYDFHRXV SDOH ±VHSWDWH P ± ± î ± somewhat curvedtogeniculate-sinuous, unbranched, internal hyphaeorstromata,through stomata,erect,straight, solitary orinsmallfascicles, 2–7,divergent,arisingfrom substomatal, olivaceous-brown orbrown. Stromata hyphae branched,septate,paleolivaceousorbrownish. E\ ZKLWH JUH\LVK HIIXVH abundant conidialformationorbrownish. WR SXQFWLIRUP ¿QHO\ VFDWWHUHG . Cercospora peristrophes ORW DNQRVDO± P GLDP P ± VPDOO RU ODFNLQJ DOPRVW B. Conidiophore tips. (BPI 439429,isolectotype). Caespituli C. Mycelium Conidia. Bar=10 Conidiophores amphigenous, IMA FUNGUS internal; A. Cercosporoid fungi 4 to olivaceous brown, paler towards the tip, thin-walled, ARTICLE smooth; conidiogenous cells integrated, terminal and LQWHUFDODU\ ± P ORQJ FRQLGLRJHQRXV ORFL WKLFNHQHG DQGGDUNHQHG±PGLDPConidia solitary, acicular or subacicular to somewhat obclavate-subcylindrical when VKRUWHU± ± î±P±VHSWDWHK\DOLQH thin-walled, smooth, apex subacute to obtuse, base truncate RUDOPRVWVR±PZLGHKLODWKLFNHQHGDQGGDUNHQHG Lectotype (designated here, MycoBank, MBT202781): India: Bihar: Patna, on Peristrophe bicalyculata, 25 Nov. 1952, M. J. Thirumalachar (K(M) IMI 55521). Isolectotypes: BPI 439429, CUP 40770. Host range and distribution: On Peristrophe bicalyculata, Acanthaceae, Asia (India, Bihar, Maharashtra, Uttar Pradesh; Myanmar), Africa (Somalia). Notes: This species belongs to the Cercospora apii s. lat. complex. Records from “Nepal” (Crous & Braun 2003, MycoBank and Index Fungorum) refer to the type of C. peristrophigena (Rao et al. 1999), which is, however, incorrect since this species was described from India (Uttar Pradesh). Cercospora ruellina R.K. Chaudhary et al., J. Living World 2(2): 43 (1995). (Fig. 16) Synonym: Cercospora ruelliae S. Narayan et al., in Rao et al., Sugarcane Pathology, Vol. 1: Fungal Diseases: 86 (1999), nom. inval. (Art. 39.1) [holotype: India: Uttar Pradesh: Gorakhpur, Maharajganj, on Ruellia prostrata, Dec. 1993, S. Narayan (HCIO 41994); isotype: GPU 5077]. Literature: Crous & Braun (2003: 360), Kamal (2010: 83). Illustration: Rao et al. ¿J Description: Leaf spots amphigenous, circular or almost so, 1–6 mm diam, brown to blackish on the upper leaf surface, whitish green with olivaceous margin below. Caespituli amphigenous, effuse. Mycelium internal; hyphae branched, septate, hyaline to pale olivaceous. Stromata Fig. 16. Cercospora ruellina (based on Rao et al. ¿J VXEVWRPDWDO ± P GLDP ROLYDFHRXV WR ROLYDFHRXV A. Conidiophore fascicle. B. C.&RQLGLD%DU P brown. Conidiophores solitary or in small fascicles, 2–5, arising from stromata, through stomata, erect, straight to JHQLFXODWHVLQXRXV XQEUDQFKHG DERXW ± î ± Host range and distribution: Only known from the type P± VHSWDWHOLJKWROLYDFHRXVWRROLYDFHRXVEURZQWKLQ collection. walled, smooth; conidiogenous cells integrated, terminal and intercalary; conidiogenous loci conspicuous, thickened and Notes: This species is a typical member of the Cercospora darkened. Conidia solitary, narrowly acicular, shorter conidia apii s. lat. complex with acicular conidia. A morphologically sometimes subcylindrical to somewhat obclavate, straight well agreeing North American sample on Ruellia ciliosa has to curved, often with short lateral germ tubes which may been examined (USA, Illinois, Chandlerville, 18 Aug. 1886, JLYHULVHWRVHFRQGDU\FRQLGLDDERXW±î±P A. B. Seymour, BPI 435186), although the conidiophores 3–18-septate, hyaline, thin-walled, smooth, apex obtuse to in the latter collection are much longer and somewhat subacute, base truncate, hila thickened and darkened. EURDGHU ± î ± P +RZHYHU LI ,QGLDQ DQG 1RUWK $PHULFD FROOHFWLRQV DUH FRQVSHFL¿F UHTXLUHV SK\ORJHQHWLF Holotype: India: Uttar Pradesh: Gorakhpur, Maharajganj, H[DPLQDWLRQ DQG FRQ¿UPDWLRQ ,QGLDQ DV ZHOO DV 1RUWK on Ruellia prostrata, Dec. 1993, S. Narayan (HCIO 41994). American samples seems to pertain to plurivorous species Isotype: GPU 5077. of the C. apii complex. VOLUME 6 · NO. 2 389 390 ARTICLE (Fig. 17) Illustration Guo Literature Cercospora strobilanthis white orwhite, with darkerborder, narrowtomoderatelywide, PPGLDPDW¿UVWSDOHJUHHQLVKODWHUEURZQLVK¿QDOO\JUH\LVK Description ¿J P Conidiophore fascicle. Fig. 17. Appl et al Cercospora strobilanthis . . (2005:18),Kamal(2010:89). : Crous&Braun(2003:390),Guo(2001:343–344), 17 &KLGGDUZDU SO,,¿JV± *XR : Leaf spots : 77(1962);as“ B. amphigenous, circulartoirregular, 2–10 Conidiophore tips. Chidd., strobilanthidis (K(M) IMI83190,holotype). Mycopathol. Mycol. C. Conidia.Bar=10 ”. et al Braun A. . et al QRWYHU\GLVWLQFWRU¿QHO\SXQFWLIRUPGDUN symptoms. VRPHWLPHV ERUGHU EURZQ ZLWKGLIIXVHEURZQLVKKDOR¿QDOO\VRPHWLPHVZLWKVKRWKROH GDUN ZLWK ZKLWH JUH\LVK ¿QDOO\ or somewhatirregular, 1–7mmdiam,brown,greyish Description ¿J *XR Illustrations al 403), Braun&Urtiaga(2008:485),Kamal(2010:93),Guo (1991: 291),Braun Literature Cercospora thunbergiana hila thickenedanddarkened. EDVHWUXQFDWHRUVOLJKWO\REFRQLFDOO\WUXQFDWH±PZLGH somewhat constrictedatsepta, hyaline,thin-walled,smooth, RFFDVLRQDOO\ ±VHSWDWH P ± ± î ± FXUYHG Conidia ZLGH P ± ± GDUNHQHG DQG WKLFNHQHG FRQVSLFXRXV WHUPLQDODQGLQWHUFDODU\±PORQJFRQLGLRJHQRXVORFL WRZDUGV the tip,thin-walled,smooth;conidiogenouscellsintegrated, SDOHU EURZQ ± VHSWDWH P ± ± î ± stomata, erect,straighttogeniculate-sinuous,unbranched, arising frominternalhyphaeorsmallstromata,through in smallfascicles,2–15,divergent,occasionallysolitary, VZROOHQK\SKDOFHOOV±PGLDPEURZQ Stromata RLLJQXOFWLNQG QGUHH ±P ± GDUNHQHG diam. DQG WKLFNHQHG ORFL FRQLGLRJHQRXV conidiogenous cellsintegrated,terminalandintercalary, olivaceous-brown, wallthinorslightlythickened,smooth; ±î±P±VHSWDWHWKURXJKRXWSDOHEURZQRU subcylindrical todistinctlygeniculate,1–6times,unbranched, arising fromstromata,throughstomata,erect,straight, Conidiophores DEVHQWRUVPDOOPDLQO\VXEVWRPDWDO±PGLDPEURZQ GDUN ¿QH SXQFWLIRUP brown toalmostblack. (Fig. 18) Holotype WUXQFDWH ±PKLODWKLFNHQHGDQGGDUNHQHG somewhat attenuatedattheverybase(veryshortobconically thin-walled, smooth,apexpointed,basetruncatetousually K\DOLQH ±VHSWDWH P ± ± î ± FXUYHG WR lat. somewhat attenuatedattheverybase. Note Acanthaceae Host rangeanddistribution IMI 83190). Strobilanthes . (2005:18). . complex,butwellcharacterisedbyconidiathatareoften 198 (1965);as“ : Atrue Conidia solitary, aciculartosomewhatobclavate,straight : : Yen &Lim(1980:165),BraunCastañedaRuiz lacking orsmall,formingsmallaggregationsof : Fig. 18. Cercospora thunbergiana (PC, lectotype). A. Conidiophore Fig. 19. Cercospora thunbergiigena (K(M) IMI 345300, holotype). A. fascicle. B. Conidophore tips. C.&RQLGLD%DU P Conidiophore fascicle. B. &RQLGLD%DU P Lectotype (designated here, MycoBank, MBT202782): Literature: Crous & Braun (2003: 403), Kamal (2010: 93). Singapore: Bukit Timah (Hwa Chung College), on Thunbergia alata, 3 Aug. 1964, S. H. Yen 73 (PC). Isolectotype: K(M) IMI Illustration: Srivastava et al ¿J 120995. Description: Leaf spots amphigenous, often vein-limited, Host range and distribution: On Thunbergia (alata, erecta, mostly marginal, 3–10 mm diam, dark brown to blackish. JUDQGLÀRUD, Thunbergia sp.), Acanthaceae, Asia (Brunei; Caespituli DPSKLJHQRXV ¿QH Mycelium internal. Stromata India, Andhra Pradesh, Uttar Pradesh; Singapore), South DEVHQW RU VPDOO VXEVWRPDWDO ± P GLDP EURZQ America (Venezuela), West Indies (Cuba). Conidiophores in small, loose fascicles, arising from small stromata, through stomata, erect, straight to moderately Cercospora thunbergiigena U. Braun & Crous, nom. JHQLFXODWH XQEUDQFKHG DERXW ± î ± P nov. 3–10-septate, brown, thin-walled, smooth; conidiogenous MycoBank MB814567 cells integrated, terminal and intercalary, conidiogenous loci (Fig. 19) FRQVSLFXRXV WKLFNHQHG DQG GDUNHQHG DERXW P GLDP Basionym: Cercospora thunbergiana R.K. Srivast et al., Conidia solitary, narrowly acicular, straight to curved, 45–155 Kavaka 20/21: 42 “1992/1993” (1995), nom. illeg. î±P±VHSWDWHK\DOLQHWKLQZDOOHGVPRRWKDSH[ (Art. 53.1), non C. thunbergiana J.M. Yen, 1965; as SRLQWHG EDVH WUXQFDWH ± P ZLGH KLOD WKLFNHQHG DQG “thumbergiana”. darkened. VOLUME 6 · NO. 2 391 392 ARTICLE Illustration Literature Synonym on Basionym Passalora acanthicola Passalora specieson Acanthaceae &RQLGLDVROLWDU\REFODYDWHVKRUW±î±P± ± Key toPassaloraspecieson Acanthaceae Passalora border. centre greyishwhite,surroundedbyapinkishorlightbrown Description Illustration Kamal (2010:12). Literature Cercospora acanthacearum species NQRZQ LQVXI¿FLHQWO\ DQG H[FOXGHG 'RXEWIXO Notes collection). Host rangeanddistribution Pandey Holotype EOYW î ± ±HWW \OQ WKLQ K\DOLQH ±VHSWDWH P ± î ± REFODYDWH reduced toconidiogenouscells. aseptate, palebrown,thin-walled,smooth;conidiophores WRVLQXRXVJHQLFXODWHXQEUDQFKHG±î±P arising fromstromatichyphalaggregations,erect,straight GLDP P ± FHOOV K\SKDO Stromata narrower conidia. It resembles on ± î±P±VHSWDWHSDOHROLYDFHRXV RQ &RQLGLRSKRUHVPXFKVKRUWHU±P FRQLGLD±î±P &RQLGLRSKRUHV±PORQJSOXULVHSWDWH FRQLGLDF\OL &RQLGLDLQFKDLQVHLWKHUPXFKORQJHU± PRUPXFKQDUUR (Fig. 20) Trans. Brit.Mycol.Soc London KQHJD JUDQGLÀRUD 7KXQEHUJLD Sydowia Mycosphaerella and Anam : This speciesbelongstothe (K(M)IMI345300). Caespituli : : : Chupp(1954:22). : Vasudeva (1963:30),Crous&Braun(2003:40). : Phaeoramulariaacanthicola 'HLJKWRQ ¿J *RYLQGX 7KLUXPDODFKDU SO¿J India small, onlycomposedofafewbrownswollen Cropr acanthicola Cercospora : 156 Leaf spots 8 C. thunbergiana : 221(1954). Adhatoda vasica : 121(1944). : Uttar Pradesh mostly epiphyllous. circular toirregular, 2–4mmdiam, . , 88 (Hansf.) U.Braun&Crous, Acanthaceae : Onlyknownfromthetype : 385(1987). , butdiffers inhavingmuch : Gorakhpur, Ramgarharea, Govindu& Thirum., Conidiophores ...... see ...... Conidia 1 : 40(2003). Hansf., C. apiis.lat (Hansf.)Deighton, Mycelium , Dec.1989, solitary, narrowly Proc. Linn.Soc. fasciculate, . complex. internal. V. P. P. V. Braun cnhsarboreus Acanthus et al QGULFDORUVRPHZKDWF\OLQGULFDOREFODYDWH REFODYDWHF\OLQGULFDOVWUDLJKWRUVOLJKWO\FXUYHG± î± GLDP P in simpleorbranchedchains, cylindricalorsomewhat ± DERXW GDUNHQHG DQG and intercalary, conidiogenouslociconspicuous, thickened thin-walled, smooth;conidiogenous cellsintegrated,terminal pluriseptate throughout,moderately brownorreddishbrown, JHQLFXODWHVLQXRXVVLPSOHRUEUDQFKHG±î±P of swollenhyphalcells,emergingthroughstomata,erect, to 10,arisingfrominternalhyphaeorsmallaggregations FRORXUOHVV ZLGH P lacking oralmostso. ± K\SKDH LQWHUQDO deep reddishbrown,moreorlessvein-limited. 5–8 mmdiam,brown. Description answer thisquestion. agreeing withtheoriginaldescriptionarenecessaryto but are-examinationoftypematerialornewcollections Cercospora s.str original descriptionthisspeciesdoesprobablynotbelongto by Govindu& Thirumalachar (1954),butbasedonthe of thestructureconidiogenouslociwerenotprovided could notbetraced,neitherinBPInorIMI(nowK).Details deposited atBPI,IMI,andHCIO.However, typematerial material ofnewspeciesdescribedintheirpaperhadbeen Notes collection. Host rangeanddistribution:Onlyknownfromthetype traced). betonica Holotype truncate. walled, smooth,apexpointed,baseshortobconically C. peristrophes Cercospora peristrophes Note Literature . VHSWDWH RQ Tropicale dei FunghidellaSomalia 1953. (Art. 53.1),non : ZHU±P RQRWKHUKRVWV : According toGovindu& Thirumalachar (1954),type Cercospora peristrophes ± ± ± VHSWDWHK\DOLQH , : : Crous&Braun(2003:316). Acanthaceae India : Leaf spots ): 65(1988),as“ Barleria lupulina Thirum. &Govindu,1953. : ...... Itmightbeaspeciesof Karnataka C. peristrophes Conidiophores DW¿UVWGLIIXVH\HOORZLVKODWHUDQJXODU , 31Jan.1953, Cercospora justiciae-adhatodae Caespituli : Mysore,NandiHills,on E. Castell., E. Castell.isahomonymof ...... peristrophis ...... 2 ...... in smallloosefascicles, hypophyllous, effuse, ( Thirum. &Govindu, Biblioteca Agraria H. C.Govindu Pseudocercospora Elenco Annotato Conidia P. barleriigena P. acanthicola ”, IMA FUNGUS nom. illeg Mycelium catenate, Stromata Justicia (not . , Cercosporoid fungi 4 ARTICLE Fig. 21. Passalora barleriigena (based on Meeboon et al. 2007a: ¿J A. Conidiophore fascicle. B. Conidiophores. C. Conidia. %DU P Fig. 20. Passalora acanthicola (K(M) IMI 4557a, holotype). A. Base of conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 Illustration: Meeboon et al D ¿J P Description: Leaf spots amphigenous, subcircular to irregular, ±PPGLDPSDOHWRSDOHEURZQPDUJLQDW¿UVWLQGH¿QLWH P±VHSWDWHSDOHROLYDFHRXVWKLQZDOOHGVPRRWKDSH[ later conspicuous, dark. Caespituli amphigenous. Mycelium obtuse or subtruncate, base short obconically truncate, 1.5–2 internal. Stromata VXEVWRPDWDO VPDOO ± P GLDP PZLGHKLODWKLFNHQHGDQGGDUNHQHG composed of 3–7 swollen hyphal cells, brown. Conidiophores in dense fascicles, 4–7, arising from stromata, through stomata, Holotype: Uganda: Kampala, Kawandra, on Acanthus erect, straight, subcylindrical, somewhat attenuated towards arboreus, Acanthaceae, Jan. 1943, C. G. Hansford 3144 WKHWLSXQEUDQFKHG±WLPHVJHQLFXODWHDERXW±î± (K(M) IMI 4557a). P±VHSWDWHGDUNHUEURZQEHORZSDOHUWRZDUGVWKHWLS thin-walled, smooth; conidiogenous cells integrated, terminal, Host range and distribution: Only known from the type conidiogenous loci conspicuous, thickened and darkened, collection. ±PGLDPConidia solitary, mostly obclavate, occasionally oblong cylindrical, straight to occasionaly somewhat curved, Passalora barleriigena Meeboon & Hidayat, ±î±P± ± VHSWDWHVXEK\DOLQHWRSDOHEURZQ Mycotaxon 102: 140 (2007). thin-walled, smooth, apex obtuse, base short obconically (Fig. 21) WUXQFDWH±PZLGHKLODWKLFNHQHGDQGGDUNHQHG VOLUME 6 · NO. 2 393 394 ARTICLE on &RQLGLDDFLFXODUWRREFODYDWHF\OLQGULFDOEDVHWUXQFDW &RQLGLRSKRUHVVKRUW±P± ± VHSWDWH RQ &RQLGLRSKRUHVORQJHUWRPDQGRU LIVKRUWHU ±VHSWD on 6WURPDWDGHYHORSHG DQGRUFRQLGLRSKRUHVVKRUWHUWRPDQ other hosts ...... 6WURPDWDODFNLQJRUDOPRVWVR FRQLGLRSKRUHV±î 6WURPDWDGHYHORSHG DQGRUFRQLGLDORQJHUWRPKLODEUR 6WURPDWDODFNLQJ FRQLGLD±î±PKLOD±P &RQLGLRSKRUHVWRPORQJ±VHSWDWH RQ &RQLGLRSKRUHV±î±P± ± VHSWDWH RQ Conidialbasetruncatetoobconicallytruncate;on 10 (9) 8 Conidiaobclavate-cylindrical tosubacicular;conidiawithpointedapex;on 9 (8) &RQLGLRSKRUHV±PORQJ±VHSWDWH RQ &RQLGLRSKRUHVVKRUWHU±î±P FRQLGLDOKLOD±P on &RQLGLDREFODYDWHF\OLQGULFDOEDVHFRQVLVWHQWO\REFRQLFDOO\W KLOD±PZLGH RQ &RQLGLRSKRUHV±î±P±VHSWDWH FRQLGLDUDWKHU FRQLGLRSKRUHVWRPORQJ RQ Conidiacylindricalor subcylindrical, basetruncatetosomewhatobconicallytruncate; &RQLGLRSKRUHV± ± î±P±VHSWDWH RQ 6 (4) &RQLGLRSKRUHVVKRUW±î±P±VHSWDWH RQ puiett) nohrhss ...... (pluriseptate);onotherhosts &RQLGLRSKRUHVORQJHUDQGEURDGHUWRî±PORQJHUFR &RQLGLRSKRUHVUHODWLYHO\VKRUW± ± P 6WURPDWDGHYHORSHG±PGLDP ...... Stromata lackingoralmostso 3 (2) DQGRUFRQLGLDPXFKQDUURZHU±P 6WURPDWDGHYHORSHG DQGRUFRQLGLRSKRUHVVKRUWHUWRDERXW 6WURPDWDODFNLQJ FRQLGLRSKRUHV±î±P FRQLGL 0\FHOLXPLQWHUQDO VXSHU¿FLDOK\SKDHZLWKVROLWDU\FRQLGLRSKRU 0\FHOLXPLQWHUQDODQGH[WHUQDO VXSHU¿FLDOK\SKDHZLWKVROLWD Key toPseudocercosporaspecieson Acanthaceae Pseudocercospora Meeboon (CMU 28212). National Park,on Holotype Conidial base consistently obconically truncate ...... Conidial baseconsistentlyobconicallytruncate Conidia obclavate-cylindrical, subacicular conidia lacking; on other hosts ...... Conidia obclavate-cylindrical,subacicularconidialacking;onotherhosts oiipoe –-ett;o te ot ...... Conidiophores 0–3-septate;onotherhosts : & Thailand I. Araki Rhinacanthus Paratype Cynarospermum Lepidagathis alopecuroidea Barleria lupulina (CMU28213). : Chiang MaiProvince : thesamelocality, 10Dec.2006, , Ruellia ...... , 30Dec.2005, ...... Thunbergia alata : DoiSuthep-Pui ...... 7 ...... Strobilanthes cusia J. Meeboon ...... 5 ...... 3 ...... Rhinacanthus nasutus Justicia japonica Braun Odontonema callistachyum J. Cynarospermum HWRORQJREFRQLFDOO\WUXQFDWH±PZLGH Rhinacanthus nasutus Justicia galapagana et al Thunbergia ±P RQ ZLGH RQ Acanthaceae Host rangeanddistribution:On ...... D±PORQJ RQ ORQJ±î±P±VHSWDWH UXQFDWH FRQLGLRSKRUHVVKRUWHUWRPORQJ . U\FRQLGLRSKRUH HV WH DGHU±P RQRWKHUKRVWV PDQGRUQDUURZHU±P ZLGH RQRWKHUKRVWV GDERYHDOOQDUURZHU±PZLGH QLGLRSKRUHVZLWKPRUHWKDQWZRVHSWD in vivo ...... 17 ...... Blechum pyramidatum Barleria cristata p...... spp. , Dyschoriste ...... ntdvlpd ...... 12 ...... notdeveloped , Asia (Thailand). , Asia Acanthus guineensis ...... in vivo ...... , Justicia Justicia dvlpd ...... 2 ...... developed ...... 8 ...... 14 ...... 4 ...... P. consociata ...... p...... spp. , ...... 6 ...... 13 ...... 11 ...... 9 ...... Barleria lupulina P. odontonematis P. thunbergiicola P. lepidagathidis P. baphiacanthi P. thunbergiae P. rhinacanthi P. rhinacanthi var. P. blepharidis P. justiciicola ...... 10 ...... IMA FUNGUS P. barleriae ...... 15 ...... P. justiciae P. acanthi dimorpha P. blechi , Cercosporoid fungi 4 &RQLGLDREFODYDWHF\OLQGULFDODFLFXODUFRQLGLDQRWIRUPHGEDVH±ZLGH RQRWKHUKRVWV ...... 16 ARTICLE &RQLGLRSKRUHV±î±P± ± VHSWDWH H[WHUQDOP\FHOLXPODFNLQJ onDyschoriste, Justicia, Ruellia ...... P. consociata var. consociata &RQLGLRSKRUHVWRPORQJ±VHSWDWH H[WHUQDOP\FHOLXPXVXDOO\GHYHORSHG onRhinacanthus nasutus ...... P. rhinacanthi &RQLGLRSKRUHVORQJ±P RQAsystasia gangetica ...... P. asystasiae &RQLGLRSKRUHVVKRUWHUWRPDYHUDJH P RQRWKHUKRVWV (see “Tabular key to Pseudocercospora species on Acanthaceae according to host genera” ±IXUWKHULGHQWL¿FDWLRQMXVWEDVHGRQPRUSKRORJ\EDUHO\SRVVLEOH Tabular key to Pseudocercospora species on Acanthaceae according to host genera Acanthus A single species ...... P. acanthi Asystasia A single species ...... C. asystasiae Baphicacanthus, see Strobilanthes Barleria A single species ...... P. barleriae Blechum A single species ...... P. blechi Blephris, see Cynarospermum Cynarospermum A single species ...... P. blepharidis Dicliptera A single species ...... P. diclipterae Dyschoriste A single species ...... P. consociata var. consociata Ecbolium A single species ...... P. ecbolii Justicia 0\FHOLXPLQWHUQDOVXSHU¿FLDOK\SKDHZLWKVROLWDU\FRQLGLRSKRUHVODFNLQJ...... P. consociata var. consociata 0\FHOLXPLQWHUQDODQGH[WHUQDOVXSHU¿FLDOK\SKDHZLWKVROLWDU\FRQLGLRSKRUHVGHYHORSHG ...... 2 6WURPDODFNLQJ FRQLGLRSKRUHV±î±P FRQLGLDF\OLQGULFDOWRREFODYDWHVXEF\OLQGULFDO ±î±PKLOD±PZLGH...... P. justiciae 6WURPDWDGHYHORSHG±PGLDP FRQLGLRSKRUHVVKRUWHUDQGQDUURZHU±î±P FRQLGLDQDUURZHUREFODYDWH±î±PKLOD±PZLGH...... P. justiciicola Lepidagathis A single species ...... P. lepidagathidis Odontonema A single species ...... P. odontonematis Rhinacanthus A single species ...... P. rhinacanthi VOLUME 6 · NO. 2 395 396 ARTICLE ±î±P±VHSWDWHSDOHROLYDFHRXVEURZQ 6WURPDWDWRPGLDP FRQLGLRSKRUHVORQJHUDQGEURDGHU± –-ett,sbyln,pl elws re ovr aeoiaeu ...... 0–8-septate,subhyaline,paleyellowishgreentoveryolivaceous 6WURPDWDODFNLQJ FRQLGLRSKRUHVVKRUWDQGQDUURZ±î± Thunbergia A...... singlespecies Strobilanthes A...... singlespecies Rungia (Fig. 23) collection. Host rangeanddistribution 1937 (K(M)IMI7695). (Fig. 22) Basionym Pseudocercospora asystasiae guineensis Holotype URXQGHG±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG olivaceous, apexpointed,baseobconicallytruncatetoalmost RUVRPHZKDWVLJPRLG± î±P±VHSWDWHSDOH obclavate-subcylindrical tosubacicular, straighttocurved but alwaysunthickenedandnotdarkened. long, conidiogenouslociinconspicuoustosubdenticulate, RUFRQLGLRSKRUHVUHGXFHGWRFRQLGLRJHQRXVFHOOV±P thin-walled, smooth;conidiogenouscellsintegrated,terminal XQEUDQFKHG±î±P± ± VHSWDWHSDOHEURZQ VWUDLJKW HUHFW K\SKDH subcylindrical-conical tosomewhatgeniculate-sinuous, VXSHU¿FLDO IURP DULVLQJ VROLWDU\ dense fascicles,8–50,arisingfromstromata,oroccasionally brown. smooth. VXSHU¿FLDO VXEK\DOLQH K\SKDH±PZLGHSDOHROLYDFHRXVVHSWDWHWKLQZDOOHG ZLGH P ± K\SKDH LQWHUQDO punctiform, scattered. raised. diam, grey-brown,withnarrowdarkbrownborder, somewhat Description Illustration Pseudocercospora acanthi Pseudocercospora specieson Acanthaceae A...... singlespecies Ruellia Nakashima Literature Gard. Bull Mycol. Soc. 178 (1967). Conidiophores Caespituli Stromata : : Yen &Lim(1980:169),CrousBraun(2003:70), : 'HLJKWRQ ¿J , iraLeone Sierra Cercospora asystasiae : et al Acanthaceae Leaf spots ., . (2010). Singapore 88 ZHOOGHYHORSHG±PGLDPFRPSDFW amphigenousratherpaleolivaceous, : 381(1987). in smalltosometimeslargeand amphigenous, subcircular, to5mm Mycelium : Gorahun(Tunkia), on , 3 Apr. 1939, 33 : Onlyknownfromthetype : 169(1980). Deighton, J.M. Yen, internal andexternal; (J.M. Yen)J.M. Yen, F. C.Deighton Conidia Rev. Mycol Trans. Brit. Acanthus solitary, . Braun , M 32 : et al Fig. 22. A. LUUHJXODU±PPGLDPDW¿UVWSDOHJUHHQLVKJUH\LVKJUHHQ VXSHU¿FLDOK\SKD Description ¿J Illustrations . Conidiophore fascicles. P FRQLGLD± î±P ...... P. consociata î±P FRQLGLDDOVRORQJHUDQGEURDGHU Pseudocercospora acanthi : ODWHU \HOORZLVK EURZQ WR EURZQLVK JUH\LVK EURZQ ¿QDOO\ Host range and distribution: On Asystasia (gangetica ARTICLE greyish white, occasionally zonate, margin indistinct to distinct, [coromandeliana], nemorum), Acanthaceae, Africa (Ivory dark brown to blackish. Caespituli amphigenous, indistinct to Coast), Asia (Indonesia, Japan, Singapore). SXQFWLIRUPEURZQVFDWWHUHGWRFRQÀXHQWDQGGHQVHUMycelium internal. Stromata ODFNLQJRUVPDOOVXEVWRPDWDO±PGLDP Note: A second collection on Asystasia gangetica from Ivory subglobose, brown. Conidiophores in small to moderately large Coast (29 Dec. 1974, G. Gilles 47), deposited at PC, has fascicles, loose to moderately dense, arising from stromata, been examined. emerging through stomata, erect, straight, subcylindrical to ÀH[XRXVVLQXRXVRUVRPHZKDWJHQLFXODWHVLQXRXVXQEUDQFKHG Pseudocercospora baphicacanthi W.H. Hsieh & ±î±P±VHSWDWHROLYDFHRXVEURZQWKLQZDOOHG Goh, Bot. Bull. Acad. Sin. 30: 123 (1989). smooth; conidiogenous cells integrated, terminal, about 10–40 (Fig. 24) P ORQJ FRQLGLRJHQRXV ORFL LQFRQVSLFXRXV WR VXEGHQWLFXODWH neither thickened nor darkened. Conidia solitary, obclavate- Literature: Hsieh & Goh (1990: 12), Guo & Hsieh (1995: 1), F\OLQGULFDO VWUDLJKW WR FXUYHG ± ± î ± ± ± Guo et al. (1998: 11). P± ± VHSWDWHVXEK\DOLQHWRSDOHROLYDFHRXVEURZQWKLQ walled, smooth, apex obtuse to subacute, base short obconically WUXQFDWH±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG Holotype: Singapore: Botanical Garden, on Asystasia gangetica, 5 Jan. 1966, J. M. Yen 731 (PC). Fig. 24. Pseudocercospora baphicacanthi (based on Hsieh & Goh ¿J A. 6ROLWDU\ FRQLGLRSKRUH DULVLQJ IURP VXSHU¿FLDO Fig. 23. Pseudocercospora asystasiae (PC, holotype). A. hyphae or emerging through stomata. B. Conidiophores, C. Conidia. Conidiophore fascicle. B. Conidiophores. C. &RQLGLD%DU P %DU P VOLUME 6 · NO. 2 397 398 ARTICLE Holotype short obconicallytruncate,hilaunthickened,notdarkened. walled, smooth,apexobtuseorsubobtuse,basetruncateto î±P±VHSWDWHJUHHQLVKWRSDOHROLYDFHRXVWKLQ cylindrical orsubcylindrical,straighttosomewhatcurved,30– collection. Host rangeanddistribution W. H.Hsieh cusia DERXWPZLGHXQWKLFNHQHGQRWGDUNHQHG conidiogenous lociinconspicuousorvisibleastruncatetips, walled, smooth;conidiogenouscellsintegrated,terminal, septate, palebrownthroughoutorpalertowardsthetip,thin- JHQLFXODWHVLQXRXVVLPSOHRUEUDQFKHG±î±P hyhae, lateral,erecttodecumbent,straightcurved, HPHUJLQJWKURXJKVWRPDWDRUVROLWDU\DULVLQJIURPVXSHU¿FLDO WUXQFDWH±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG walled, smooth,apexsubacute toobtuse,baseobconically ±VHSWDWH subhyaline topaleolivaceous orolivaceous-brown,thin- P ± î ± ± FXUYHG VRPHZKDW WR Conidia or somewhatrefractive,butneither thickenednordarkened. ORFL FRQLGLRJHQRXV ORQJ inconspicuous orsubconspicuousbybeingsubdenticulate P ± WHUPLQDO LQWHJUDWHG brown, tipspaler, thin-walled,smooth;conidiogenous cells ROLYDFHRXV WR ROLYDFHRXV ± ±VHSWDWH P ± î RU occasionally branched,somewhatgeniculate-sinuous,20– VLPSOH ÀH[XRXV GHFXPEHQW WR HUHFW VWRPDWD WKURXJK arising frominternalhyphalorsmallaggregations, Conidiophores velvety. Caespituli blackish brownpatches,coveringlargeleafsegments. leaf surface,darkbrownbelow, sometimesforminglarge ±PPGLDPRUFRQÀXHQWDQGODUJHUEODFNLVKRQWKHXSSHU Description %UDXQ ¿J Illustrations Literature 3DFL¿TXH Basionym Pseudocercospora barleriae VWRPDWD lacking. WKURXJK HPHUJLQJ K\SKDH VXSHU¿FLDO Caespituli WKH RQ SDWFKHV XSSHUOHDIVXUIDFH\HOORZLVKEURZQEHORZPDUJLQLQGH¿QLWH \HOORZLVK IRUPLQJ FRQÀXHQW VRPHWLPHV Description ¿J *XR Illustration (Fig. 25) Braun, [ Baphicacanthus cusia Mycelium Conidiophores solitary, obclavatetoobclavate-cylindrical,straight : : Yen &Lim(1980:156),CrousBraun(2003:76). : 17 +LK *K ¿ *R +VLHK *XR ¿J *RK +VLHK Cercosporabarleriae Taiwan : : hypophyllous. Fig. 26. Pseudocercospora blechi (CUP-VZ-002240, holotype). A. Conidiophore fascicles. B. Conidiophore. C. &RQLGLD%DU P Description: Leaf spots mainly epiphyllous, indistinct to irregular and somewhat reddish, 0.5–3 mm diam, sometimes FRQÀXHQW PDUJLQ LQGH¿QLWH Caespituli hypophyllous, subeffuse, dark olivaceous to blackish. Mycelium internal. Stromata lacking or small, composed of a few swollen hyphal cells, brown. Conidiophores in small, loose fascicles, arising from internal hyhae or hyphal aggregations, emerging through VWRPDWDHUHFWÀH[XRXVJHQLFXODWHVLQXRXVVLPSOHRUUDUHO\ EUDQFKHG ± î ± P ±VHSWDWH SDOH ROLYDFHRXV or olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to FRQLGLRJHQRXV FHOOV ± P ORQJ FRQLGLRJHQRXV ORFL Fig. 27. Pseudocercospora blepharidis (K(M) IMI 83163, holotype). inconspicuous, unthickened, not darkened. Conidia solitary, A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 cylindrical to obclavate-cylindrical, straight to slightly curved, P ±î±P±VHSWDWHVXEK\DOLQHWRSDOHROLYDFHRXV apex obtuse to subacute, base obconically truncate, 1–1.5 PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG Description: Leaf spots amphigenous, subcircular, ovoid to irregular, 3–18 mm diam, pale to dark brown, sometimes Holotype: Venezuela: Caracas, on Blechum pyramidatum, somewhat zonate, above all on the lower leaf surface. 28 Jul. 1938, A. S. Muller 2240 (CUP-VZ-002240). Isotype: Caespituli amphigenous, punctiform, brown. Mycelium VIA. internal. Stromata well-developed, substomatal, 20–30 P GLDP ROLYDFHRXVEURZQ Conidiophores in small to Host range and distribution: On Blechum pyramidatum moderately large fascicles, arising from stromata, through [brownei], Acanthaceae, Central and South America stomata, divergent to dense, straight, subcylindrical-conical (Panama, Venezuela), West Indies (Cuba, Puerto Rico, WRVOLJKWO\JHQLFXODWHVLQXRXVXQEUDQFKHG±î±P Virgin Islands). 0–1(–2)-septate, pale to medium brown or olivaceous-brown, wall thin or only slightly thickened, smooth; conidiogenous Pseudocercospora blepharidis (Chidd.) U. Braun & cells integrated, terminal or conidiophores reduced to Crous, Mycosphaerella and Anam.: 82 (2003). FRQLGLRJHQRXV FHOOV ± P ORQJ FRQLGLRJHQRXV ORFL (Fig. 27) inconspicuous. Conidia solitary, acicular to obclavate- Basionym: Cercospora blepharidis Chidd., Sydowia 13: 154 F\OLQGULFDO VWUDLJKW WR FXUYHG ± î ± P “1959” (1960). 3–8-septate, subyhaline to pale olivaceous, thin-walled, smooth, apex subacute to obtuse, base truncate to long Literature: Kamal (2010: 155). REFRQLFDOO\ WUXQFDWH ± P ZLGH KLOD XQWKLFNHQHG QRW darkened. Illustration &KLGGDUZDU SODWH9¿JV± VOLUME 6 · NO. 2 399 400 ARTICLE Conidiophore fascicles. b. Fig. 28a,b.a. Basionym (Fig. 28a) Pseudocercospora consociata Holotype asperrimum collection. Host rangeanddistribution 1956, P. consociata var. & X.J.Liu, 70 (1883). .P hdawr2 P. P. Chiddarwar consociata : : India Cercospora consociata Pseudocercospora consociata [ Blepharis asperrima : var. Mycosystema Maharashtra dimorpha D. Conidiophores. (K(M)IMI83163). (HAL 2190F, holotype). : Khandala,on : Onlyknownfromthetype 2 : 232(1989). ], G. Winter, (G. Winter) Y.L. Guo Acanthaceae E. &RQLGLD%DU P var. Cynarospermum consociata Hedwigia A. , 9Nov. 6XSHU¿FLDOK\SKDH (BPI435187,neotype). Braun 22 : et al to diffuse yellowishtobrownishdiscolorations,orforming Description Exsiccatae ¿J ¿J *XR Illustration (2010: 166),Phengsintham & Braun(2003:136),Freire(2006:236),Kamal (1965: 7),Guo&Hsieh(1995:1), Literature . B. 6ROLWDU\FRQLGLRSKRUHVDULVLQJIURPVXSHU¿FLDOK\SKDH : Saccardo(1886:470),Chupp(1954:23),Katsuki A. &KXSS ¿J *XR +VLHK : Ellis&Everh.,N. Amer. Fungi2477. etal. : Conidiophore fascicles. Leaf spots ¿J 3KHQJVLQWKDP lacking or almost so, inconspicuous lacking oralmostso,inconspicuous et al . 2013:110). B. Conidiophores. etal. (1998:11), Crous IMA FUNGUS et al C. Conidia. . 2013: C. Cercosporoid fungi 4 subcircular to angular-irregular lesions, 1–10(–15) mm diam, terminal or conidiophores often reduced to conidiogenous ARTICLE EURZQLVK ¿QDOO\ JUH\LVK EURZQ WR JUH\LVK ZKLWH PDUJLQ FHOOV ± P ORQJ FRQLGLRJHQRXV ORFL LQFRQVSLFXRXV LQGH¿QLWH RU GDUNHU EURZQ RFFDVLRQDOO\ UHGGLVK EURZQ otherwise as in var. consociata. sometimes slightly raised, narrow, often only formed as marginal line. Caespituli amphigenous, punctiform to effuse, Holotype: Venezuela: Lara State: Quíbor, on Justicia dark brown to blackish, later greyish by abundant conidiation. galapagana, Acanthaceae, Jan. 2007, R. Urtiaga (HAL 2190 F). Mycelium internal; hyphae branched, septate, occasionally FRQVWULFWHGDWVHSWD±PZLGHVXEK\DOLQHWREURZQLVK Host range and distribution: Only known from the type thin-walled, smooth. Stromata variable, almost lacking, small collection. to medium in size, subglobose to somewhat irregular, 10–30 PGLDPRFFDVLRQDOO\ODUJHUWRPGLDPVXEVWRPDWDOWR Notes: Pseudocercospora consociata, known from Asia, immersed, olivaceous-brown to dark brown, cells subglobose North and South America on hosts of various genera of the WRVRPHZKDWDQJXODULUUHJXODULQRXWOLQH± ± PGLDP Acanthaceae, is probably taxonomically heterogeneous. This ZDOOWRPZLGHConidiophores in small to moderately large species is morphologically rather variable in all basic traits. fascicles, occasionally in large fascicles, divergent to dense, However, a further splitting needs cultures and molecular arising from substomatal swollen hyphal cells or stromata, sequence analyses based on material from all continents and emerging through stomata or erumpent, erect, straight, host genera involved. Type material of this species, collected subcylindrical or attenuated towards the tip, slightly to distinctly by Seymour before 1884, could not be traced. Records of geniculate-sinuous, unbranched or rarely branched, 5–50(– Pseudocercospora consociata on Dicliptera chinensis from î±P±VHSWDWHORQJHURQHVSODLQO\SOXULVHSWDWH China (Guo & Hsieh 1995, Guo et al. 1998) are undoubtedly subhyaline to pale or medium olivaceous, olivaceous-brown incorrect and belong to P. diclipterae. The two species are or brown, thin-walled, smooth; conidiogenous cells integrated, morphologically barely distinguishable. terminal or conidiophores reduced to conidiogenous cells, ± P ORQJ FRQLGLRJHQRXV ORFL LQFRQVSLFXRXV RU YLVLEOH Pseudocercospora diclipterae (A.K. Kar & M. as truncate tips or shoulders, always unthickened and not Mandal) Deighton, Trans. Brit. Mycol. Soc. 88: 388 darkened. Conidia solitary, obclavate-cylindrical, straight to (1987). FXUYHG ± ± î ± ± P ± ± VHSWDWH (Fig. 29) subhyaline to pale olivaceous, thin-walled, smooth, apex acute Basionym: Cercospora diclipterae A.K. Kar & M. Mandal, to obtuse, base short to sometimes long obconically truncate, Trans. Brit. Mycol. Soc. 53: 337 (1969). ±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG Literature: Crous & Braun (2003: 159), Kamal (2010: 171). [Holotype: USA: Illinois: on Ruellia ciliosa, A. B. Seymour]. Neotype (designated here, MycoBank, MBT202783): USA: Illustration .DU 0DQGDO ¿J Missouri: near Emma, on Ruellia ciliosa, Aug. 1889, C. H. Demetrio [Ellis & Everh., N. Amer. Fungi 2477] (BPI 435187). Description: Leaf spots amphigenous, subcircular, 1–6 mm Isoneotypes: Ellis & Everh., N. Amer. Fungi 2477, e.g. DAOM, GLDPVFDWWHUHGVRPHWLPHVFRQÀXHQWXQLIRUPO\\HOORZLVKWR FH. GXOO\HOORZLVKEURZQODWHUROLYDFHRXVE\DEXQGDQWIUXFWL¿FDWLRQ Caespituli amphigenous, mostly hypophyllous, punctiform Host range and distribution: On Dyschoriste oblongifolia, to effuse, brown or deep olivaceous. Mycelium internal. Justicia (gendarussa, procumbens), Ruellia (ciliosa, Stromata ODFNLQJ RU VPDOO ± P GLDP VXEJORERVH RU prostrata, strepens, tuberosa), Acanthaceae, Asia (China; somewhat oblong, substomatal, brown. Conidiophores in India, Uttar Pradesh, West Bengal; Japan, Thailand), North small to moderately large fascicles, usually 2–15, divergent, America (USA, Alabama, Florida, Iowa, Illinois, Mississippi, arising from substomatal hyphae or stromata, through Missouri, Oklahoma), South America (Brazil, Venezuela). stomata, erect, straight to curved or geniculate-sinuous, XQEUDQFKHG±î±P±VHSWDWH\HOORZLVKEURZQ var. dimorpha U. Braun & Urtiaga, Feddes Repert. thin-walled, smooth; conidiogenous cells integrated, terminal 119: 489 (2008). or conidiophores sometimes reduced to conidiogenous (Fig. 28b) FHOOV ± P ORQJ FRQLGLRJHQRXV ORFL LQFRQVSLFXRXV WR subdenticulate, unthickened, not darkened. Conidia solitary, Illustration %UDXQ 8UWLDJD ¿J REFODYDWHVXEF\OLQGULFDOVWUDLJKWWRFXUYHG±î±P 2–16-septate, pale yellowish brown, olivaceous, thin-walled, Description: Differs in vivo from var. consociata by the smooth, apex subacute or subobtuse, base short obconically IRUPDWLRQ RI VXSHU¿FLDO K\SKDH ZLWK VROLWDU\ FRQLGLRSKRUHV WUXQFDWH±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG VXSHU¿FLDO K\SKDH HPHUJLQJ WKURXJK VWRPDWD EUDQFKHG VHSWDWH± ± PZLGHVXEK\DOLQHWRSDOHROLYDFHRXV Holotype: India: West Bengal, Darjeeling, Sevak Road, on thin-walled, smooth; solitary conidiophores arising from Dicliptera chinensis [roxburghiana], 11 Nov. 1967, M. Mandal VXSHU¿FLDO K\SKDH ODWHUDO HUHFW VWUDLJKW VXEF\OLQGULFDO (K(M) IMI 135116). Isotype: BPI 435712. conical to somewhat geniculate-sinuous, unbranched, 5–30 î±P± ± VHSWDWHVXEK\DOLQHWRSDOHROLYDFHRXV Host range and distribution: On Dicliptera chinensis, brown, thin-walled, smooth; conidiogenous cells integrated, Acanthaceae, Asia (?China; India, West Bengal). VOLUME 6 · NO. 2 401 402 ARTICLE Basionym (Fig. 30) Pseudocercospora ecbolii chinensis Note P A. Fig. 29. notes under Conidiophore fascicles. Deighton, Brit. Mycol.Soc : Recordsof Pseudocercospora diclipterae , fromChinabelongpossiblyto : Cercosporaecbolii P. consiciata Trans. Brit.Mycol.Soc Pseudocercosora consociata . 53 : 338(1969). B. ). Conidiophores. A.K. Kar&M.Mandal, (A.K. Kar&M.Mandal) (K(M) IMI135116, holotype). . 88 C. P. diclipterae : 388(1987). Conidia. Bar=10 on Dicliptera Trans. (see Braun et al Conidiophore fascicles. Fig. 30. P LPVEOEVWROQ DN OYFRV WR ROLYDFHRXV GDUN large fascicles, mostly2–15,arisingfromstromata, through REORQJ WR olivaceous-brown. VXEJORERVH GLDP P internal. IUXFWL¿FDWLRQ DEXQGDQW Caespituli E\ EHORZ ROLYDFHRXV GHHS ¿QDOO\ limited, yellowishdiscolorations,4–10mmdiam,scattered, Description Illustration Literature . Pseudocercospora ecbolii : Crous&Braun(2003:168),Kamal(2010:172). Stromata .DU 0DQGDO ¿J : hypophyllous, effuse, olivaceous. Leaf spots Conidiophores lacking orsmall,substomatal, 5–25 B. Conidiophores. amphigenous, formedasvein- (K(M) IMI135117, holotype). in smalltomoderately C. &RQLGLD%DU P IMA FUNGUS Mycelium A. Cercosporoid fungi 4 stomata, divergent to moderately dense, erect, straight, ARTICLE subcylindrical-conical to geniculate-sinuous, unbranched RU RFFDVLRQDOO\ ± WLPHV EUDQFKHG ± î ± P 0–6-septate, pale olivaceous to olivaceous-brown, thin- walled, smooth; conidiogenous cells integrated, terminal or conidiophores sometimes reduced to conidiogenous FHOOV±PORQJFRQLGLRJHQRXVORFLLQFRQVSLFXRXVWR subdenticulate, unthickened, not darkened. Conidia solitary, narrowly obclavate-cylindrical, straight to curved, 20–100 î ± P ±VHSWDWH SDOH ROLYDFHRXV WR YHU\ SDOH olivaceous-brown, thin-walled, smooth, apex subacute or VXEREWXVH EDVH REFRQLFDOO\ WUXQFDWH ± P ZLGH KLOD neither thickened nor darkened. Holotype: India: West Bengal: 24-Parganas, Rajarhat, on Ecbolium ligustrinum, Acanthaceae, 16 Jul. 1967, M. Mandal (K(M) IMI 135117). Isotype: BPI 436048. Host range and distribution: Only known from the type collection. Pseudocercospora justiciae (F.L. Tai) Y.L. Guo & X.J. Liu, Mycosystema 4: 103 (1991). (Fig. 31) Basionym: Cercospora justiciae F.L. Tai, Lloydia 11: 47 (1948). Literature: Chupp (1954: 24), Guo & Hsieh (1995: 2), Guo et al. (1998: 11). Illustrations 7DL ¿J *XR +VLHK ¿J 3), Guo et al. ¿J Description: Leaf spots lacking or only formed as yellowish to yellowish brown discolorations on the upper leaf surface, circular, 2–5 mm diam, grey to greyish brown below, margin LQGH¿QLWHCaespituli hypophyllous, effuse, sooty. Mycelium LQWHUQDODQGH[WHUQDOVXSHU¿FLDO K\SKDHHPHUJLQJWKURXJK VWRPDWDEUDQFKHGVHSWDWH±PZLGHSDOHROLYDFHRXV to olivaceous-brown, thin-walled, smooth. Stromata not developed. Conidiophores in loose fascicles, 2–11, arising from internal hyphae, through stomata or solitary, arising from VXSHU¿FLDOK\SKDHHUHFWVXEF\OLQGULFDOÀH[XRXVFXUYHG Fig. 31. Pseudocercospora justiciae (based on Guo & Hsieh 1995: 4, sinuous to somewhat geniculate-sinuous, unbranched or ¿J A. Conidiophore fascicle. B. &RQLGLD%DU P EUDQFKHG ± î ± P ±VHSWDWH ROLYDFHRXV brown to pale brown, thin-walled, smooth; conidiogenous cells integrated, terminal, conidiogenous loci inconspicuous, Notes: A record of P. justiciae on Justicia sp. from Florida, occasionally subdenticulate. Conidia solitary, cylindrical to 86$ $O¿HUL et al. 1984) is unclear and doubtful. Material REFODYDWHVXEF\OLQGULFDOVWUDLJKWWRFXUYHG±î± could not be traced. P±VHSWDWHSDOHROLYDFHRXVWRROLYDFHRXVWKLQ walled, smooth, apex obtuse to subacute, base obconically Pseudocercospora justiciicola P.N. Singh et al., WUXQFDWHDERXW±PZLGHKLODQHLWKHUWKLFNHQHGQRU Mycol. Res. 100: 1129 (1996). darkened. (Fig. 32) Holotype: China: Sichuan: Chengtu, on Justicia procumbens, Illustration: Singh et al ¿JV± 1943, Lee Ling 125 (HMAS 12126). Description: Leaf spots lacking or almost so, lesions Host range and distribution: On Justicia (procumbens, indistinct, occasionally formed as greyish discolorations. Justicia sp.), Acanthaceae, Asia (China), ?North America Caespituli hypophyllous, greyish, discrete to effuse. Mycelium (USA, Florida). LQWHUQDODQGH[WHUQDO VXSHU¿FLDOK\SKDHEUDQFKHGVHSWDWH VOLUME 6 · NO. 2 403 404 ARTICLE VXSHU¿FLDOK\SKD A. Holotype WUXQFDWHDERXW±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG smooth, apexacutetosubobtuse, baseshortobconically WKLQZDOOHG ROLYDFHRXV ±VHSWDWH P ± î ± straight tocurved,occasionallywithshortlateralbranchlets, loci unthickened,notdarkened. VPRRWK conidiogenous cellsintegrated,terminal, WKLQZDOOHG ROLYDFHRXVEURZQ ±VHSWDWH ± P î ± ÀH[XRXV XQEUDQFKHG WR JHQLFXODWHVLQXRXV VWUDLJKW VRPHZKDW HUHFW WHUPLQDO RFFDVLRQDOO\ ODWHUDO DULVLQJIURPVWURPDWDRUVROLWDU\DULVLQJIURPVXSHU¿DOK\SKDH WR VXEJORERVH GLDP somewhat oblong,brown. P ± DERXW VPDOO VPRRWK GHYHORSHG WKLQZDOOHG GLDP P ± VXEK\DOLQH Fig. 32. 366368). [ simplex Conidiophore fascicle. Pseudocercospora justiciicola ], : Isotype Acanthaceae Nepal C. : GPU4004. : Kathmanduvalley, on Conidiophore. B. , Nov.–Dec. 1993, Solitary conidiophoresarisingfroma Conidiophores D. Conidia &RQLGLD%DU P (K(M) IMI366368,holotype). solitary, obclavate, in loosefascicles, Justicia japonica Kamal (K(M) IMI Stromata Braun et al which is,however, agenuinespeciesof (Stevenson 1975,Minter previous falselyreferredtoas Note collection. Host rangeanddistribution 16490 alopecuroidea Holotype WUXQFDWH±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG walled, smooth,apexobtusetosubacute,baseobconically WKLQ ROLYDFHRXV SDOH WR VXEK\DOLQH ± VHSWDWH P ± î straight tocurved,narrowlyobclavate-cylindrical,20–120 WKLFNHQHG QHLWKHU ORFL nor darkened,occasionallysubdenticulate. FRQLGLRJHQRXV ORQJ P ± FHOOV integrated, terminalorconidiophoresreducedtoconidiogenous RU olivaceous-brown, thin-walled,smooth;conidiogenouscells XQEUDQFKHG SDOH ±VHSWDWH P ± î JHQLFXODWHVLQXRXV ± EUDQFKHG UDUHO\ RQO\ ÀH[XRXV HUHFW WHUPLQDO WR RU ODWHUDO VWUDLJKW K\SKDH VXSHU¿FLDO IURP DULVLQJ aggregations, throughstomata,orconidiophoressolitary, fascicles, arisingfrominternalhyphaeorstromatichyphal aggregations, brown. Stromata WR VXEK\DOLQH VHSWDWH VWRPDWD pale olivaceousorolivaceous-brown,thin-walled,smooth. ZLGH P WKURXJK ± HPHUJLQJ EUDQFKHG VSDULQJO\ K\SKDH VXSHU¿FLDO H[WHUQDO subeffuse, ratherinconspicuous. purple tinge,marginindistinct. dingy olivaceous-browntoblackishbrownorwith subcircular tosomewhatirregularorevendiffuse, brownish, Description Illustration (Fig. 33) Pseudocercospora lepidagathidis Basionym Pseudocercospora odontonematis discussion under 1986, original labelas paratype materialof pertains to Notes collection. Host rangeanddistribution Literature (Fig. 34) . Mycotaxon (2003). Braun &Crous, Cercospora : The materialon : There isasecondcollectionfromNepalthatprobably (BPI437637). R. K.Verma : : Chupp(1954: 25). lacking or only with very small substomatal hyphal lacking oronlywithverysmallsubstomatalhyphal : %UDXQ &URXV ¿J : Cercospora odontonematis P. justiciicola uroRico Puerto Leaf spots , : 25(1954);as“ Acanthaceae 92 Justicia P. rungiae : 399(2005). (GPU,KK221,K(M)IMI303480)].See P. rungiae Conidiophores Mycosphaerella and Anam. sp.),Nepal,KathmanduValley, Jan. . Verma &Kamal(1991)citeditas et al : SanJuan,on Lepidagathis alopecuroidea amphigenous, 1–5 mm diam, amphigenous, 1–5mmdiam, . , 10Feb.1962, : Onlyknownfromthetype : Onlyknownfromthetype . 2001,Crous&Braun2003), [on “ odontonemae Cercospora lepidagathidis Caespituli Ruellia prostrata Mycelium in small, usually loose in small,usuallyloose Chupp, U.Braun&Crous, Cercospora s.str Conidia ”. internal and internaland (Chupp) U. Monograph of hypophyllous, hypophyllous, Lepidagathis IMA FUNGUS H. L.Rubin ” (onthe solitary, : 296 was . Cercosporoid fungi 4 ARTICLE Fig. 33. Pseudocercospora lepidagathidis (BPI 437637, holotype). A. 6ROLWDU\ FRQLGLRSKRUHV DULVLQJ IURP VXSHU¿FLDO K\SKDH B. Conidiophore fascicles. B. Conidiophores. C. &RQLGLD%DU P Fig. 34. Pseudocercospora odontonematis (CUP 40422, lectotype). A. 6XSHU¿FLDO K\SKD B. Solitary conidiophores arising from VXSHU¿FLDOK\SKDHC. Conidiophore fascicle. D. Conidiophores. E. Description: Leaf spots amphigenous, circular to somewhat &RQLGLD%DU P angular-irregular, 2–5 mm diam, brown, later with greyish brown to dingy grey centre and darker border. Caespituli hypophyllous, punctiform to effuse. Mycelium LQWHUQDODQGH[WHUQDO VXSHU¿FLDO FHOOVRULQWHJUDWHGWHUPLQDO±PORQJFRQLGLRJHQRXVORFL K\SKDH EUDQFKHG VHSWDWH ± P ZLGH VXEK\DOLQH RU SDOH inconspicuous to subdenticulate, but always unthickened and olivaceous, thin-walled, smooth; stromata substomatal, 10–40 not darkened. Conidia solitary, narrowly cylindrical to obclavate- P GLDP EURZQ Conidiophores in small to moderately large F\OLQGULFDO ± î ± P ± ±VHSWDWH YHU\ SDOH fascicles, loose to dense, arising from stromata, through olivaceous, thin-walled, apex obtuse to subacute, base more or VWRPDWDRUHUXPSHQWDOVRVROLWDU\DULVLQJIURPVXSHU¿DOK\SKDH OHVVWUXQFDWHWRGLVWLQFWO\REFRQLFDOO\WUXQFDWH±PZLGH lateral, rarely terminal, erect, straight, subcylindrical-conical to hila unthickened, not darkened. VWURQJO\ JHQLFXODWHVLQXRXV XQEUDQFKHG ± î ± P 0–2-septate, very pale or pale olivaceous to olivaceous-brown, Lectotype (designated here, MycoBank, MBT202784): thin-walled, smooth; conidiophores recuced to conidiogenous Mexico: Veracruz: Cordoba, on Odontonema callistachyum, VOLUME 6 · NO. 2 405 406 ARTICLE (Fig. 35) Basionym Pseudocercospora rhinacanthi Acanthaceae collection. Host rangeanddistribution Isolectotype et al. Literature HPQODRW OQ RLLJQXORFL FRQLGLRJHQRXV ORQJ shoulders. P ± inconspicuous orvisibleastruncate,subdenticulatetips DERXW WHUPLQDO conidiophores reducedtoconidiogenouscellsorintegrated, pale olivaceoustoolivaceous-brown,thin-walled,smooth; RURFFDVLRQDOO\EUDQFKHG± ± î±P±VHSWDWH subcylindrical tomoderatelygeniculate-sinuous,unbranched VWUDLJKW ODWHUDO K\SKDH VXSHU¿FLDO IURP DULVLQJ VROLWDU\ RU arising fromstromata,throughstomata,erecttodecumbent, in loosetomoderatelydense,smallfascicles,usually2–6, ±PGLDPROLYDFHRXVEURZQWREURZQ ROLYDFHRXVEURZQ WR VXEK\DOLQH thin-walled, smooth. ZLGH P ± VHSWDWH Mycelium on dryleaves. diam, brownorlaterwithdingygreycentre,ratherindistinct with yellowishhalotocircularorangular-irregular, 2–10mm yellowish tobrowndiscolorationsordiffuse brownspots Description Exsiccatae Synonym Rhinacanthus Indonesia Lectotype unthickened, notdarkened. EDVHXVXDOO\ORQJREFRQLFDOO\WUXQFDWH±PZLGHKLOD SDOH olivaceous, thin-walled,smooth,apexmoreorlesspointed, WR ± VXEK\DOLQH î ± ± ± VHSWDWH LQGLVWLQFWO\ ± ± ± P FXUYHG VRPHZKDW WR VWUDLJKW Host rangeanddistribution Fungi Imp.Exs.847,e.g.,K(M)IMI89002(slide),HBG,W. Imp. Exs.847](BPI440801). two speciesaremorphologically barelydistinguishable. are unclear, unprovenandmightbelongto procumbens Notes (Indonesia, Philippines, Thailand). [ Justicia nasuta Mycol. Pap 1 11) s“ 414 (1912);as Wiss Kaiserl. Akad. Fung (1998:376),Crous&Braun(2003:351). : Chineserecordsof . : : Chupp(1954:25),Guo&Hsieh(1995:357), : 25 LQWHUQDODQGH[WHUQDO VXSHU¿FLDOK\SKDHEUDQFKHG : : Kabát&Bubák,FungiImp.Exs.847. : Cercosporina rhinacanthi ( Conidia : CUP 60863. Java : 917(1931). designated here ecsoarhinacanthi Cercospora (Tai 1979,Guo&Hsieh1995, , 25Jul.1932, Leaf spots sp.,1907, Caespituli . ], : Bogor(Buitenzorg),BotanicGarden,on 140 Rhinacanthus rhynacanthi solitary, narrowlyobclavate-cylindrical, Stromata : 152(1976). ., amphigenous, mainlyhypophyllous. F. v. Höhnel amphigenous, formedasdiffuse Math.-Naturwiss. Cl : Onlyknownfromthetype : On Isolectotypes O. A. Plunkett O. A. , MycoBank,MBT202785): ”. lacking orsmall,substomtal, P. rhinacanthi sp.), Rhinacanthus [Kabát&Bubák,Fungi (Höhn.) Sacc., Höhn., (Höhn.) Deighton, Acanthaceae P. consociata : Kabát&Bubák, (CUP 40422). Conidiophores on ., Sitzungsber. etal. Wien ( nasutus Justicia 1998) , Asia , . The 121 Syll. Braun : et al yellowish. angular-irregular ordiffuse, 3–5mm diam,greenishwhite, Description ±¿JV± Illustrations Literature Synonym Pseudocercospora rungiae A. Fig. 35. (Fig. 36) VXSHU¿FLDOK\SKD . Conidiophore fascicles. Indian Phytopathol. (HCIO 32003)]. Kolhapur, on 8 : 69“1977”(1978)[ Pseudocercospora rhinacanthi : : Crous&Braun(2003:361),Kamal(2010:216). ecsoarungiae Cercospora : 3DWLO ¿J 9HUPD .DPDO Caespituli Leaf spots C. Rungia repens Conidiophores. B. hypophyllous, scattered, rather 44 mainly hypophyllous,circular to Solitary conidiophoresarisingfroma : 446(1991). holotype M.S. Patil, , 11 Dec.1975, D. R.K. Verma &Kamal, &RQLGLD%DU P : (BPI 440801,lectotype). India Botanique : Maharashtra IMA FUNGUS M. S.Patil (Nagpur) : Cercosporoid fungi 4 WUXQFDWH±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG ARTICLE Holotype: India: Uttarakhand: Kath Godam, Nainital, on Rungia pectinata, Oct. 1985, R. K. Verma (GPU, KK 125). Isotype: K(M) IMI 299176. Host range and distribution: On Rungia (pectinata [SDUYLÀRUD], repens), Acanthaceae, Asia (India, Maharashtra, Uttarakhand; Nepal). Note: The identity of Cercospora rungiae and Pseudocercospora rungiae has been proven by Kamal (2010). A record of C. rungiae from Andhra Pradesh (Braun et al. 1992: 363) refers to a true Cercospora (see C. justiciicola) and not to this species. Verma & Kamal (1991) cited a collection on “Ruellia prostrata” (on the original label as Justicia sp.) as paratype material (Nepal: Kathmandu Valley, Jan. 1986, R. K. Verma (GPU, KK 221, K(M) IMI 303480). 7KLVFROOHFWLRQFKDUDFWHULVHGE\IRUPLQJVXSHU¿FLDOK\SKDH with solitary conidiophores is excluded from P. rungiae. 7KH LGHQWL¿FDWLRQ RI WKH KRVW LV XQFHUWDLQ DQG WKH LGHQWLW\ of this fungus is not quite clear, but it probably belongs to Pseudocercospora justiciicola. Pseudocercospora thunbergiae (Boedijn) U. Braun & Sivapalan, Fungal Diversity 3: 21 (1999). (Fig. 37) Basionym: Cercospora thunbergiae Boedijn, Nova Hedwigia 3: 411 (1961). Literature: Crous & Braun (2003: 403), Kamal (2010: 225). Illustrations %RHGLMQ WDE ¿J %UDXQ 6LYDSDODQ ¿J Description: Leaf spots amphigenous, angular-irregular, often YHLQOLPLWHG±PPGLDPVRPHWLPHVFRQÀXHQWDQGODUJHU GDUNEURZQODWHUJUH\LVKEURZQWRGLQJ\JUH\PDUJLQLQGH¿QLWH Caespituli hypophyllous, inconspicuous. Mycelium internal and H[WHUQDO VXSHU¿FLDOK\SKDHVSDULQJO\EUDQFKHG±PZLGH Fig. 36. Pseudocercospora rungiae (K(M) IMI 299176, isotype). septate, subhyaline to pale olivaceous, thin-walled, smooth. A. Conidiophore fascicle. B. Conidiophore (branched). C. Solitary Stromata lacking or very small, only composed of a few FRQLGLRSKRUHVDULVLQJIURPDVXSHU¿FLDOK\SKDD.&RQLGLD%DU P VZROOHQK\SKDOFHOOV±PGLDPROLYDFHRXVWREURZQLVK Conidiophores VROLWDU\DULVLQJIURPVXSHU¿FLDOK\SKDHODWHUDO rarely terminal, occasionally emerging through stomata, inconspicuous. Mycelium internal; hyphae branched, septate, solitary or in small, loose fascicle, erect, straight, subcylindrical- FRORXUOHVV ± P ZLGH VRPHZKDW ZLGHU QHDU VWURPDWD FRQLFDOWRVRPHZKDWJHQLFXODWHVLQXRXVXQEUDQFKHG±î Stromata lacking or poorly developed, substomatal, small, ±P±VHSWDWHVXEK\DOLQHWRSDOHROLYDFHRXVEURZQ WR DERXW P GLDP Conidiophores fasciculate, 2–10, thin-walled, smooth; conidiogenous cells integrated, terminal divergent, arising from small stromata, through stomata, or conidiophores aseptate, reduced to conidiogenous cells, erect, straight, subcylindrical-conical to distinctly geniculate- ± P ORQJ FRQLGLRJHQRXV ORFL LQFRQVSLFXRXV QHLWKHU VLQXRXV XQEUDQFKHG WR EUDQFKHG ± î ± P thickened nor darkened. Conidia solitary, narrowly obclavate- 0–3(–6)-septate, pale olivaceous to olivaceous-brown, thin- F\OLQGULFDO ± î ± P ± ±VHSWDWH VXEK\DOLQH walled, smooth; conidiophores occasionally reduced to pale yellowish green to very pale olivaceous, thin-walled, conidiogenous cells, but mostly integrated, terminal, 10–30 VPRRWKDSH[VXEDFXWHEDVHREFRQLFDOO\WUXQFDWH±P PORQJFRQLGLRJHQRXVORFLLQFRQVSLFXRXVWRVXEGHQWLFXODWH wide, hila unthickened, not darkened. but always unbranched and not darkened. Conidia solitary, REFODYDWHVXEF\OLQGULFDO VWUDLJKW WR FXUYHG ± î ± Holotype: Indonesia: Jawa Barat: Bogor, botanical garden, P±VHSWDWHVXEK\DOLQHWRSDOHROLYDFHRXVWKLQZDOOHG Thunbergia alata, Apr. 1950, K. B. Boedijn (L 53879). Isotype: smooth, apex obtuse or subacute, base short obconically K(M) IMI 91586. VOLUME 6 · NO. 2 407 408 ARTICLE Basionym Pseudocercospora thunbergiicola Host rangeanddistribution hyphae. Conidiophores andhyphaeemergingthroughastoma. Fig. 37. Literature Conidiophores. Indonesia). JUDQGLÀRUD (Fig. 38) 30 on, : 195(1965);as“ C. Mycol. Pap Pseudocercospora thunbergiae : Yen &Lim(1980: 188),Crous&Braun(2003: 403). 6ROLWDU\FRQLGLRSKRUHDULVLQJIURPDVXSHU¿FLDOK\SKD : ), Cercospora thunbergiicola Acanthaceae E. &RQLGLD%DU P . 140 thunbergicola , Asia (Brunei;India,UttarPradesh; : 154(1976). : On Thunbergia ”. J.M. Yen, (L 53879,holotype). (J.M. Yen)Deight- ( alata Rev. Mycol B. VXSHU¿FLDO , erecta Braun A. D. . , et al VXSHU¿FLDOK\SKDH Conidiophore fascicles. Fig. 38. F\OLQGULFDOVWUDLJKWWRFXUYHGDERXW±î±P tips orshoulders. conidiogenous lociinconspicuous orvisibleastruncate FRQLGLRJHQRXVFHOOVLQWHJUDWHGWHUPLQDO±PORQJ ROLYDFHRXVEURZQ somewhat palertowardsthe tip,thin-walled,smooth; RU ROLYDFHRXV SDOH ±VHSWDWH P XQEUDQFKHGRURFFDVLRQDOO\RQFHEUDQFKHG±î± narrower towardsthetip,togeniculate-sinuous, hyphae, lateral,erect,straight,subcylindricalorsomewhat VXSHU¿FLDO IURP DULVLQJ VROLWDU\ RU GHQVH PRGHUDWHO\ WR hyphae orstromata,emergingthroughstomata,divergent moderately largefascicles,2–22,arisingfromsubstomatal substomatal, subglobose,brown. Stromata ±PZLGHSDOH hyphae emergingthroughstomata,sparinglybranched, conspicuous. margin darkbrown. angular-irregular, 0.5–3mmdiam,centrepale,whitish, Description ¿J Illustrations . Pseudocercospora thunbergiicola ORW DNQRVDO± P GLDP P ± VPDOO RU ODFNLQJ DOPRVW 4–11-septate, pale olivaceous-brown, thin-walled, smooth, ARTICLE apex subacute, base short to long obconically truncate, ±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG Lectotype (designated here, MycoBank, MBT202786): Singapore: Botanic Garden, on Thunbergia alata, 20 Jun. 1964, S. H. Yen 60 (PC). Isolectotype: K(M) IMI 120991. Host range and distribution: On Thunbergia alata, Acanthaceae, Asia (Singapore). Semipseudocercospora A single species. Semipseudocercospora peristrophes-acuminatae (J.M. Yen) J.M. Yen, Mycotaxon 17: 363 (1983). (Fig. 39) Basionym: Cercospora peristrophes-acuminatae J.M. Yen, Rev. Mycol. 29: 230 (1964). Literature: Yen & Lim (1980: 163), Crous & Braun (2003: 316). Illustrations Description: Leaf spots lacking or rather indistinct, forming diffuse dark brown discolorations. Caespituli hypophyllous, rarely amphigenous, effuse, brown. Mycelium internal. Stromata lacking. Conidiophores solitary or in small, divergent fascicles, arising from internal hyphae, through stomata, erect, VWUDLJKWWRFXUYHGRUVRPHZKDWÀH[XRXVORQJ¿OLIRUPQRWRU EDUHO\ JHQLFXODWH XQEUDQFKHG ± ± î ± P 3–15-septate throughout, medium brown, paler towards the tip, wall somewhat thickened, smooth; conidiogenous cells LQWHJUDWHG WHUPLQDO DERXW ± P ORQJ FRQLGLRJHQRXV ORFL VXEFRQVSLFXRXV ± P GLDP LQ IURQW YLHZ YLVLEOH as small circle with darker rim and minute central pore, unthickened, but somewhat refractive or slightly darkened- refractive, loci somewhat bulging, denticle-like. Conidia solitary, short obclavate or clavate, ellipsoid, subcylindrical RU RERYRLG XVXDOO\ VWUDLJKW ± î ± P SDOH olivaceous to olivaceous-brown, thin-walled, smooth, apex EURDGO\URXQGHGEDVHVKRUWREFRQLFDOO\WUXQFDWH±P wide, hila unthickened, but occasionally slightly refractive or darkened-refractive. Lectotype (designated here, MycoBank, MBT202787): Singapore: Katong, on Peristrophe acuminata, Acanthaceae, S. H. Yen 20 (PC). Isolectotype: K(M) IMI 122324. Host range and distribution: Only know from the type collection. Notes 7KH JHQHULF DI¿QLW\ RI Semipseudocercospora is quite unclear, and it is unknown if it is a true cercosporoid genus (Braun et al. 2013). Molecular sequence analyses are necessary to elucidate the phylogenetic position of this Fig. 39. Semipseudocercospora peristrophes-acuminatae (PC, genus. holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. %DU P VOLUME 6 · NO. 2 409 410 ARTICLE (Fig. 41) (Fig. 40) J VH R J X *XR +VLHK ¿J *XR ¿J *RK +VLHK ¿J Illustrations 13), Guo&Hsieh(1995:4–5), Guo Literature Pseudocercospora actinidiae on &RQLGLDEURDGO\REFODYDWHVXEF\OLQGULFDO± ± î± Key toPseudocercosporaspecieson Actinidiaceae Pseudocercospora specieson Actinidiaceae Illustrations Literature Cercospora actinidiae A singlespecies Cercospora specieson Actinidiaceae Actinidiaceae Note collection. Host rangeanddistribution 50000). &RQLGLDQDUURZO\REFODYDWH¿OLIRUP±î±PEDVH± Description ¿J Actinidiaceae Holotype KLOD thickened anddarkened. ZLGH P ± DWWHQXDWHG VOLJKWO\ RU WUXQFDWH EDVH pluriseptate, hyaline,thin-walled,smooth,apexpointed, P ± ± î ± ± VLJPRLG VRPHWLPHV FXUYHG GLDP P ± intercalary, conidiogenouslocithickenedanddarkened, smooth; conidiogenouscellsintegrated,terminaland PSOXULVHSWDWHPHGLXPEURZQWREURZQWKLQZDOOHG ± ± î ± DERXW XQEUDQFKHG XVXDOO\ JHQLFXODWH hyphae orswollenhyphalcells,erect,straight,curvedto or insmallfascicles,2–8,divergent,arisingfrominternal a fewswollenhyphalcells,brown. Mycelium with narrowdarkbrownborder. 2–8 mmdiam,centregreyishbrownoryellowishbrown, 140 Mycol. Sin.Suppl : Belongingtothe : 10(1976). : : Liu&Guo(1987:355),Hsieh &Goh(1990:12– : Guo inernal. 'LKR ±¿V ± /X *XR /LX ± ¿JV ± 'HLJKWRQ /LX *XR ¿J *XR : China Leaf spots , 11 Sep.1961, Actinidia et al : Stromata . (2005:23). Jiangsu Conidia . C. apiis.lat. amphigenous, subcirculartoirregular, 1 p...... spp. : 353“1986”(1987). et al : Nanjing,on X.J. Liu& Y.L. Guo, X. J.Liu lacking orsmall,composedof : Onlyknownfromthetype solitary, acicular, straightto ±¿J Caespituli et al Deighton, complex. & Conidiophores Q. M.Ma249 . (1998:15–16). amphigenous. Actinidia Mycol. Pap. et al (HMAS . (2005: solitary Acta Braun sp., et al UQKG P LHVPWPV DQDVWRPRVLQJ VRPHWLPHV ZLGH P VWRPDWD ± WKURXJK HPHUJLQJ EUDQFKHG K\SKDH VXSHU¿FLDO H[WHUQDO PKJQX DQ\ \RKORV¿H\ XFLRP WR ÀRFFRVH ROLYDFHRXV SXQFWLIRUP GHHS ¿QHO\ HIIXVH K\SRSK\OORXV PDLQO\ DPSKLJHQRXV LQGH¿QLWH PDUJLQ EURZQ JUH\LVK WR EURZQ SDOH FRQÀXHQW VRPHWLPHV GLDP and larger, browntodarkbrown,later with palercentre, PP ± DQJXODULUUHJXODU Description Exsiccatae Conidiophore fascicle. Fig. 40. . PEDVH±PZLGH PZLGH Cercospora actinidiae : ReliquiaeFarlowianae829. : Leaf spots ...... B. Conidiophore tip. W UW DNQ DHVELFOU WR VXEFLUFXODU ODWHU ODFNLQJ ¿UVW DW (based onGuo Mycelium C. et al &RQLGLD%DU P ...... P. hangzhouensis ¿J P. actinidiae IMA FUNGUS internal and Caespituli A. A. Cercosporoid fungi 4 ARTICLE Fig. 41. Pseudocercospora actinidiae (FH, holotype). A. Stroma. B. Conidiophore fascicles. C. Solitary FRQLGLRSKRUHV DULVLQJ IURP VXSHU¿FLDO hyphae. D. Conidiophores. E. Conidia. %DU P pale olivaceous to pale brown, thin-walled, smooth. Holotype: China: Guizhou (Kweichow): Fan Ching Shan, Stromata lacking below to well-developed on the upper on Actinidia callosa, 21 Sep. 1931, S. Y. Cheo 570 (FH). OHDI VXUIDFH ± P GLDP VXEVWRPDWDO VXEJORERVH Isotypes: BPI 437420, 876607; HMAS 12125 and Reliquiae pale brown. Conidiophores in loose to dense fascicles, to Farlowianae 829. 100 conidiophores or even more, arising from stromata, through stomata, above all when epiphyllous, and solitary, Host range and distribution: On Actinidia (callosa, chinensis, DULVLQJIURPVXSHU¿FLDOK\SKDHODWHUDORFFDVLRQDOO\WHUPLQDO Actinidia sp.), Actinidiaceae, Asia (China, Anhui, Fujian, differentiation between hypophyllously formed erect to Guizhou; Korea, Japan, Taiwan). GHFXPEHQWEUDQFKHGFRQLGLRSKRUHVDQGVXSHU¿FLDOK\SKDH GLI¿FXOWIHUWLOHWKUHDGVWRPORQJVLPSOHRUEUDQFKHG Pseudocercospora hangzhouensis X.J. Liu & Y.L. individual solitary conidiophores formed as lateral branchets Guo, Acta Mycol. Sin. Suppl. 1: 357 “1986” (1987). ±PORQJ±PZLGHDVHSWDWHWRSOXULVHSWDWHSDOH (Fig. 42) to medium olivaceous, thin-walled, smooth; conidiogenous Synonym: Pseudocercospora actinidiicola Goh & W.H. Hsieh, cells integrated, terminal, conidiogenous loci inconspicuous Bot. Bull. Acad. Sin. (Taipei) 30: 122 (1989) [holotype: WRVXEGHQWLFXODWH±PGLDPXQWKLFNHQHGQRWGDUNHQHG Taiwan: Taichung, NCHU Campus, on Actinidia chinensis, Conidia solitary, obclavate-subcylindrical, straight to curved 14 Aug. 1984, T. K. Goh (NCHUPP-49); isotype: K(M) IMI RUVRPHZKDWVLJPRLG± ± î±P± VHSWDWH 312075]. pale to medium olivaceous, thin-walled, smooth, apex obtuse, base short or rarely long obconically truncate, about 2–2.5 Literature: Hsieh & Goh (1990: 12–13), Guo & Hsieh (1995: PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG 5–6), Guo et al. (1998: 16–17). VOLUME 6 · NO. 2 411 412 ARTICLE A...... singlespecies Viburnum Basionym (Fig. 43) MycoBank MB814566 Cercospora sambucicola Cercospora specieson Adoxaceae A...... singlespecies Sambucus Tabular keytoCercosporaspecieson Adoxaceae accordingtohostgenera Cercospora Adoxaceae LQFRQVSLFXRXV 2–11-septate, hyalinetopaleolivaceous,thin-walled,smooth, ORFL FRQLGLRJHQRXV ORQJ Conidia P ± FHOOV SDOH integrated, terminalorconidiophoresreducedtoconidiogenous WR VXEK\DOLQH ±VHSWDWH olivaceous-brown, thin-walled,smooth;conidiogenouscells P ± ± î ± curved, subcylindricaltogeniculate-sinuous,unbranched, WR VWUDLJKW HUHFW K\SKDH VXSHU¿FLDO IURP DULVLQJ VROLWDU\ RU fascicles, arisingfromstromata,throughstomataorerumpent, to oblong,mainlyepiphyllous,brown. thin-walled, smooth. K\SKDHEUDQFKHG±PZLGHVXEK\DOLQHWRROLYDFHRXV amphigenous. dark margin,sometimeswithyellowishbrownhalo. brown, yellowishbrownbelow, latercentregreyishwhitewith irregular andvein-limited,1–18mmdiam,browntodark Description ¿J ¿J *XR +VLHK ¿J *XR Illustrations in small,loose fascicles,2–13,arisingfrom internalhyphae WKLFNHQHG VOLJKWO\ ZDOO GLDP ± P FHOOV EURZQ LQWUDHSLGHUPDO RU VXEVWRPDWDO GLDP P aggregations ofafewswollen hyphalcells,globose,10–25 Illustrations Literature Mycelium to palegreyishbrownbelow. by agreyishblackmarginalline,paleyellowishbrown greyish whitecentreontheupperleafsurface,surrounded 2–8 mmdiam,paleyellowishbrowntobrown,laterwith Description ¿J sambuci Mycotaxon VROLWDU\QDUURZO\REFODYDWH¿OLIRUP±î±P : Guo : internal. : : /X *R J VH *RK +VLHK ¿J *XR /LX *XR -LDQJ ¿J *XR Cercospora sambuci Leaf spots Leaf spots F. Stevens&C.J. King,1927. 74 Mycelium et al : 262(2000), Stromata . (2005:55–56). Stromata amphigenous, subcirculartoangular- amphigenous, circularorsubcircular, QHQO QHWUD VXSHU¿FLDO H[WHUQDO DQG LQWHUQDO lacking orsmall,onlyformed as Y.L. Guo, GD VXEJORERVH GLDP P ± nom. illeg. Caespituli Conidiophores Y.L. Guo& Y. Jiang, Conidiophores nom. nov. (Art.53.1),non amphigenous. et al solitary or Caespituli Caespituli in dense in dense . (2005: et al Braun C. . et al Clethra and Guo javanica Host rangeanddistribution williamsii Holotype wide, hilathickenedanddarkened. VPRRWKDSH[SRLQWHGRUVXEREWXVHEDVHWUXQFDWH±P ±î±PWRSOXULVHSWDWHK\DOLQHWKLQZDOOHG acicular, straight tosomewhatcurvedorslightlysigmoid, Japan, Korea, Taiwan). Actinidia Host rangeanddistribution 28 Sep.1961, complex. Type materialof Notes Jilin, Zhejiang),?North America (USA, Alabama). POIVLOVW P RJ Q P LHFRQLGLD ZLGH P ± DQG ORQJ P WR IDVFLFOHV VPDOO barely distinguishablefrom second cercosporoidhyphomycete whichismorphologically actinidioides from Chinaon“ Holotype ±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG apex subacute,basesubtruncatetoshortobconicallytruncate, WKLFNHQHGDQGGDUNHQHG± ± PGLDP cells integrated,terminalandintercalary, conidiogenousloci narrower towardsthetip,thin-walled,smooth;conidiogenous at thebase,1–6(–13)-septate,mediumbrown,palerand ZLGH P WR P ± î ± ± LUUHJXODU ZLGWK straight, subcylindricaltogeniculate-sinuous,unbranched, or hyphalaggregations,throughstomataerumpent,erect, Notes: Pseudocercospora clematoclethrae species NQRZQ LQVXI¿FLHQWO\ DQG H[FOXGHG 'RXEWIXO . Guo, : This speciesbelongstothe sp.( Guo &Liu(1992)described , , : : et al. sp.), Mycosystema williamsii Adoxaceae China China Clethraceae ] ( FKDQJHGWKHLGHQWL¿FDWLRQRIWKHKRVWWR X.-J. Liu Actinidiaceae Actinidiaceae : Clematoclethra fabri : Zhejiang [ ...... Jilin Province buergeriana , Sep.1962, & ). 5 Q.-M. Ma671 Passalora catenospora Hangzhou,on : : 100(1992). C. sambucicola : On ), butlaterGuo&Hsieh(1995) , Asia (China, Anhui, Zhejiang; : On ]), Actinidia J. K.Bai : Yongji,on Sambucus Adoxaceae ” [ Cercospora apiis.lat (HMAS50001). C. scandens P. clematoclethrae ( X.J. Liu& Y.L. arguta (HMAS77346). (conidiophoresin C. sambucicola Actinidia arguta Conidia C. viburnicola (? , Asia (China, , Asia IMA FUNGUS canadensis , Sambucus contains a chinensis solitary, subsp. . , , , Cercosporoid fungi 4 ARTICLE Fig. 42. Pseudocercospora hangzhouensis (based on Guo & Hsieh ¿J A. Conidiophore fascicle. B. Solitary conidiophores DULVLQJIURPVXSHU¿FLDOK\SKDHC. &RQLGLD%DU P8%UDXQ del. Fig. 43. Cercospora sambucicola (HMAS 77346, holotype). A. DFLFXODUK\DOLQH±PZLGH 7KHFRQVSHFL¿W\RI&KLQHVH Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 collections and material from the USA remains to be proven P via phylogenetic analyses. Cercospora viburnicola W.W. Ray, Mycologia 33: erumpent, erect, straight, subcylindrical to geniculate- 174 (1941). VLQXRXVXQEUDQFKHG±î±PVHSWDWHWKURXJKRXW (Fig. 44) uniformly pale olivaceous-brown or somewhat paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, Literature: Chupp (1954: 106), Crous & Braun (2003: 421). WHUPLQDODQGLQWHUFDODU\DERXW±PORQJFRQLGLRJHQRXV ORFL FRQVSLFXRXV WKLFNHQHG DQG GDUNHQHG ± P GLDP Illustration:Guoet al ¿J Conidia solitary, acicular, shorter conidia narrowly obclavate, VWUDLJKW WR FXUYHG ± î ± P SOXULVHSWDWH K\DOLQH Description: Leaf spots amphigenous, subcircular to angular- thin-walled, smooth, apex acute or subacute, base truncate irregular, 2–12 mm diam, different shades of brown, pale to or somewhat obconically truncate in obclavate conidia, 2–2.5 medium brown, reddish brown to greyish brown. Caespituli PZLGHKLODWKLFNHQHGDQGGDUNHQHG DPSKLJHQRXV¿QHGDUNMycelium internal. Stromata lacking RU VPDOO DERXW ± P GLDP EURZQ Conidiophores in Holotype: USA: Oklahoma: Payne County, Stillwater, college small to moderately large fascicles, mostly dense, arising garden, on Viburnum opulus, 18 Aug. 1939, W. W. Ray 286 from internal hyphae or stromata, through stomata or (CUP 29236). VOLUME 6 · NO. 2 413 414 ARTICLE tomentosum collected atthetypelocality (21 Aug. 1942,on on aconfusionwith on of the QPVGQLFWR KGVUEG QLOVUWG REFODYDWH LOOXVWUDWHG DQG GHVFULEHG FRQIXVLRQ 7KH RQ PLVLGHQWL¿FDWLRQ DQG EDVHG XQGRXEWHGO\ LV 0XáHQNR %XUHN of 442266, CUP-C.H.OK-0032,ILL 29523,NY 945736). A record Notes Oklahoma, Wisconsin). ?Asia (China,Jiangsu,Shaanxi,Sichuan),USA (Florida, [ odoratissimum Host rangeanddistribution Conidiophore fascicle. Fig. 44. tomentosum C. viburnicola Viburnum cylindricum : Cercosporaviburnicola Cercospora apiis.lat Cercospora viburnicola sdpstdi eea ebra(BPI442264 ) isdepositedinseveralherbaria ], , opulus fromPolandon suspensum C. viburnicola B. &RQLGLD%DU P from China are incorrect and based fromChinaareincorrectandbased , plicatum . complex. Reords of this species . complex.Reordsofthisspecies , Viburnum W.W. Rayismorphologicallypart Viburnum opulus : On (CUP 29236,holotype). F.L. Tai. material Authentic Viburnum var. spp.), tomentosum Adoxaceae ĝZLGHUVND Viburnum Viburnum ( carlesii Braun A. – , , et al FODVVL¿HGDV Americal samples,theChinesecollectionsarecurrentlybetter PORQJ:LWKRXWDQ\PROHFXODUGDWDEDVHGRQ$VLDQDQG1RUWK DERXWPORQJDQGWKHFRQLGLDDUHDFLFXODUDERXW± The conidiophoresareformedinsmall,divergentfascicles,to VSHFLPHQVDUHSUREDEO\QRWFRQVSHFL¿FZLWKWKHODWWHUVSHFLHV to as & Jiang2000:264;Guo buddleifolium Viburnum opulus to concerned hasbeenre-examinedandturnedouttobelong PDUHQRWFRQVLVWHQWZLWK ± ± î ± ± EDVH WUXQFDWH REFRQLFDOO\ ZLWK FRQLGLD adoxae RIIUXFWL¿FDWLRQ7KHQDPH material havebeenexamined,butallwerewithoutanytrace described (Chupp1954).Numerousduplicatesoftheoriginal Notes Adoxaceae Host rangeanddistribution Gall. Exs.1871(e.g.,B,HBG,PC). recorded anddescribed fromBulgariaon and belongsto and conidiaaswellcercosporelloid conidiogenousloci Notes California, Minnesota,Missouri, NewMexico,Oregon). melanocarpa Host rangeanddistribution BPI 439986,NY 234138. cerulea San BernadinoValley, SanBernadinoCounty, on Lectotype Exsiccatae Illustration (1995: 80). Literature Type Literature Cercospora adoxae species NQRZQ LQVXI¿FLHQWO\ DQG H[FOXGHG 'RXEWIXO Synonym SUROL¿FDQV &HUFRVSRUD . Pseudocercospora opuli Syll. Fung XIX, no.1871, Toulouse 1882, Hist. IowaStateUniv : C. viburnicola : Although listedinliterature,thisspecieswasnever : and also known from Poland. Chinese collections on andalsoknownfromPoland.Chinesecollectionson France &HUFRVSRUDSUROL¿FDQV , Aug. 1893, Aug. , (Rabenh.) P. Karst. : : Saccardo(1895:606),Chupp(1954:104),Braun : Chupp(1954:98),Crous&Braun(2003:46). %UDXQ ¿J (designated byBraun1995:80): &HUFRVSRUHOODSUROL¿FDQV : Calif.Fungi403. , Europe(France). Cercospora apiis.lat ), . : on 15 Cercosporella Adoxaceae : 84(1901). Adoxa moschatellina , represent typical represent , aih2735 Parish Roum., and et al C. adoxae Ellis &Holw., . II, , North America (USA, Alaska, : On Viburnum which iscommonon (Braun1995). This specieswas J UHIHUUHG ¿J ± : On has colourlessconidiophores Fungi Sel.Gall.Exs 3 (NY 234139). C. viburnicola Sambucus : 42(1896). . mightreferto sp. (Guo 1996: 92; Guo sp.(Guo1996:92;Guo nom. nud. C. apiis.lat Adoxa moschatellina (Ellis &Holw.) Sacc., Roum.,FungiSel. , Sambucus ebulus Bull. Lab.Nat. ( USA cerulea . The collection collection . The Isolectotypes IMA FUNGUS (Art. 38.1). : ., but these ., butthese Sambucus Ramularia California Viburnum Viburnum , ., glauca Cent . : , : , Cercosporoid fungi 4 (Bakalova & Borisova 2010: 49–50). This record is, however, ±PZLGHZKLFKLVLQFRQWUDGLFWLRQZLWKWKHPXFKEURDGHU ARTICLE doubtful and probably wrong. Conidia were described to be FRQLGLD ±PZLGH RIWUXH&SUROL¿FDQV. Passalora Key to Passalora species on Adoxaceae &RORQLHVLQSDOHUHGGLVKEURZQWRIXOLJLQRXVSDWFKHV P\FHOLXPLQWHUQDODQGH[WHUQDOVXSHU¿FLDO FRQLGLRSKRUHVIDVFLFXODWHDVZHOODVVROLWDU\DULVLQJIURPVXSHU¿FLDOK\SKDH conidia subhyaline to reddish brown; on Sambucus spp...... P. lateritia &RORQLHV FDHVSLWXOL ZLWKRXWUHGGLVKWLQJH P\FHOLXPLQWHUQDOVXSHU¿FLDOK\SKDHODFNLQJin vivo; conidia not reddish brown ...... 2 &RQLGLRSKRUHVVKRUW±î±P±VHSWDWH FRQLGLD±î±P±VHSWDWH onViburnum spp...... P. viburni &RQLGLRSKRUHVPXFKODUJHU±î±P±VHSWDWH FRQLGLDPXFKORQJHUDQGVRPHZKDWEURDGHU ± ± î±P±VHSWDWH RQSambucus spp...... P. catenospora Passalora catenospora (G.F. Atk.) U. Braun & Crous, Lectotype (designated here, MycoBank, MBT202788): USA: Mycosphaerella and Anam.: 112 (2003). Alabama: Lee County, Auburn, on Sambucus canadensis, 11 (Fig. 45) Oct. 1891, G. F. Atkinson (CUP-A-002255a). Isolectotype: Basionym: Cercospora catenospora G.F. Atk., J. Elisha CUP-A-002255b. Mitchell Sci. Soc. 8: 66 “1891” (1892). Synonyms: Phaeoramularia catenospora (G.F. Atk.) Deighton, More Dematiaceous Hyphomycetes: 317 (1976). &HUFRVSRUD DI¿QLV G. Winter, in herb. [USA: Pennsylvania, on Sambucus canadensis, ex herb. G. Winter (B 700016202)]. Literature: Saccardo (1892: 645), Chupp (1954: 100). Illustration (OOLV ¿J$ Exsiccatae: Barthol., Fungi Columb. 2309. Description: Leaf spots indistinct. Caespituli forming effuse, irregular patches, hypophyllous, ochraceous, dark olivaceous to brownish. Mycelium internal. Stromata lacking, only formed as small aggregations of swollen hyphal cells or small, 10– P GLDP VXEVWRPDWDO \HOORZLVK WR RFKUDFHRXVEURZQ Conidiophores in small to moderately large fascicles, usually 3–12, arising from swollen hyphal cells or small stromata, emerging through stomata, erect, straight, subcylindrical or attenuated towards the tip, not or only slightly geniculate, VRPHZKDW FXUYHGVLQXRXV XQEUDQFKHG ± î ± P UDUHO\ ORQJHU XVXDOO\ ±VHSWDWH VXEK\DOLQH SDOH ochraceous to golden brown or olivaceous-brown, thin- walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–30 P ORQJ FRQLGLRJHQRXV ORFL FRQVSLFXRXV WKLFNHQHG DQG GDUNHQHG ± ±PZLGHConidia solitary to catenate, in simple or branched chains, cylindrical, subcylindrical, occasionally almost obclavate or broadly fusiform, straight WR FXUYHG RU VOLJKWO\ VLQXRXV ± ± î ± P 1–8-septate, subhyaline, pale olivaceous to brownish, thin- walled, smooth, apex obtuse, rounded (in primary conidia) or short conically truncate in catenate conidia (with a single or WZRKLOD EDVHVKRUWREFRQLFDOO\WUXQFDWH ± ±PZLGH hila somewhat thickened and darkened. Fig. 45. Passalora catenospora (CUP-A-002255a, lectotype). A. Conidiophore fascicles. B. Conidiophore. C.&RQLGLD%DU P VOLUME 6 · NO. 2 415 416 ARTICLE javanica 3(AL)). Recordsofthisspeciesfrom Taiwan on Other syntypesarefrom August 1891(CUP-A-002245#1– Note Oklahoma, Pennsylvania, Texas), West Indies(Haiti). USA, Alabama, Florida,Kansas,Mississippi,NorthCarolina, America (DominicanRepublic),North America (Canada; diam, aswellhyaline,acicularconidiaformedsingly. GDUNHUFRQLGLRSKRUHVZLWKODUJHUFRQLGLRJHQRXVORFL±P is readilydistinguishablefrom from theChinese s. lat. confusion. The secondfungusbelongstothe a secondcercosporoidhyphomycetewhichmightcause in Chupp(1954). Type materialof was erroneouslyreducedtosynonymywith Pseudocercospora ebulicola B. Fig. 46. canadensis Host rangeanddistribution DULVLQJIURPVXSHU¿FLDOK\SKDH Conidiophore fascicles. : The lectotypeisacollectionfromOctober1891. complex andismorphologicallybarelydistinguishable Passalora lateritia [ formosana , intermedia Cercospora sambucicola ] (Tai 1979)areincorrectandreferto (NY 838232,lectotype). , Sambucus C. Conidiophore. E. &RQLGLD%DU P ( Cercospora ebulicola : on P. catenospora spp.), C. catenospora D. Sambucus Adoxaceae Solitary conidiophores Solitary conidiophores A. Cercospora apii C. catenospora 6XSHU¿FLDOK\SKD . This species . This byitsmuch Sambucus contains ( , Central , Central cerulea, cerulea, ) which Braun et al simple chains, cylindrical or subcylindrical or somewhat simple chains,cylindricalorsubcylindricalsomewhat GLDP P ± DERXW conidiogenous lociconspicuous,thickenedanddarkened, FRQLGLRSKRUHVUHGXFHGWRFRQLGLRJHQRXVFHOOV±PORQJ walled, smooth;conidiogenouscellsintegrated,terminalor î ± WHUPLQDO ±P±VHSWDWHVXEK\DOLQHWRSDOHUHGGLVKEURZQWKLQ RU ODWHUDO K\SKDH VXSHU¿FLDO IURP DULVLQJ hyphae orstromata,throughstomataconidiophoressolitary, divergent tomoderatelydense,arisingfromsubstomatal brown. UHGGLVK GLDP P ± VXEVWRPDWDO FHOOV K\SKDO VZROOHQ Stromata VHSWDWH ZLGH concolorous withconidiophoresorpale,thin-walled,smooth. P ± EUDQFKHG VSDULQJO\ RU WKURXJK XQEUDQFKHG HPHUJLQJ SUHVHQW RU stomata, occasionallyclimbingleafhairs,rarelyanastomosing, ODFNLQJ K\SKDH VXSHU¿FLDO to ferruginouspatches. hypophyllous, effuse, formingreddish,palereddishbrown Description Swingle, KansasFungi6. Exsiccatae Literature Basionym Passalora lateritia fascicle. Fig. 47. (Fig. 46) . (1888). Mycosphaerella and Anam. Conidiophores Passalora viburni B. : Saccardo(1892:646),Chupp(1954:102). : Conidiophores. lacking or formed as stromatic aggregations of lacking orformedasstromaticaggregationsof Cercosporalateritia : Ellis&Everh.,N. Amer. Fungi1994.Kellerman& : Leaf spots Conidia (Ellis &Halst.)U.BraunCrous, in small to moderately large fascicles, in smalltomoderatelylargefascicles, (NY 830558,lectotype). C. Mycelium &RQLGLD%DU P DNQRLQGH¿QLWH RU ODFNLQJ solitary to catenate, usually in solitary tocatenate,usuallyin Ellis &Halst., 4 (2003). : 244 internal and external; internal andexternal; A. J. Mycol IMA FUNGUS Conidiophore Colonies Colonies . 4 : 7 Cercosporoid fungi 4 REFODYDWHF\OLQGULFDOVWUDLJKWWRVOLJKWO\FXUYHG± ± î Description: Leaf spots subcircular to angular-irregular, ARTICLE ±P ± ±VHSWDWHRFFDVLRQDOO\FRQVWULFWHGDWWKHVHSWD sometimes vein-limited, 2–10 mm diam, greyish green to subhyaline to pale reddish brown, thin-walled, smooth, apex brown, later centre greyish white, with darker margin. Caespituli obtuse, rounded in solitary conidia, short conically truncate in amphigenous, punctiform, greyish white to brownish. Mycelium catenate conidia, base obconically truncate, (1–)1.5–2(–2.5) internal. Stromata intraepidermal, occasionally substomatal, 10– PZLGHKLODVRPHZKDWWKLFNHQHGDQGGDUNHQHG PGLDPRFFDVLRQDOO\FRQÀXHQWDQGODUJHUSDOH\HOORZLVK brown to medium brown, composed of swollen hyphal cells, Lectotype (designated by Crous & Braun 2003: 244): USA: ±PGLDPConidiophores in small to rather large fascicles, Iowa: Story County, Ames, on Sambucus pubens, Sep. 1887, arising from stromata, erumpent, occasionally emerging B. D. Halsted (NY 838232). Isolectotypes: BPI 437793, CUP- through stomata, erect, straight, subcylindrical to geniculate- 040123, NY 838230, 838233; Ellis & Everh., N. Amer. Fungi VLQXRXV XQEUDQFKHG ± î ± P ±VHSWDWH K\DOLQH 1994, e.g., BPI 437794, MICH 15321. yellowish, greenish to pale olivaceous or yellowish brown, thin-walled, smooth; conidiogenous cells integrated, terminal Host range and distribution: On Sambucus (canadensis, or conidiophores mostly reduced to conidiogenous cells, 5–20 nigra, pubens, racemosa), Adoxaceae, North America (USA, P ORQJ FRQLGLRJHQRXV ORFL FRQVSLFXRXV PLQXWH ± P Iowa, Kansas). diam, barely or very slightly thickened, somewhat darkened- refractive. Conidia solitary or catenate, in simple or rarely Notes: This is a typical mycovellosielloid member of Passalora, branched chains, shape and size variable, narrowly ellipsoid- with fasciculate and solitary conidiophores arising from ovoid, fusiform, subcylindrical, straight to slightly curved, 8–45 VXSHU¿FLDOK\SKDHWKLFNHQHGGDUNHQHGFRQLGLRJHQRXV ORFL î±P±VHSWDWHK\DOLQHWRSDOH\HOORZLVKRUJUHHQLVK and obclavate-cylindrical, pigmented conidia. This species smooth to faintly verruculose, thin-walled, ends attenuated, is very characteristic by its effuse, reddish to reddish brown VKRUWREFRQLFDOO\WUXQFDWHZKHQLQFKDLQVDERXWPZLGHKLOD colonies and reddish tinge of conidiophores and conidia. minute, very slightly thickened and darkened-refractive. Passalora viburni (Ellis & Everh.) U. Braun & Crous, Lectotype (designated by Braun 1993): USA: Wisconsin: Mycosphaerella and Anam.: 474 (2003). Racine, on Viburnum lentago, 17 Jun. 1888, J. J. Davis (NY (Fig. 47) 830558). Isolectotype: NY 830559, WIS. Basionym: Ramularia viburni Ellis & Everh., J. Mycol. 5: 69 (1889). Host range and distribution: On Viburnum (lentago, Synonym: Phaeoramularia viburni (Ellis & Everh.) U. Braun, prunifolium), Adoxaceae, North America (USA, Tennessee, Mycotaxon 48: 293 (1993). Wisconsin). Literature: Saccardo (1892: 554), Braun (1998: 387). Notes 7KHJHQHULFDI¿QLW\RIWKLVVSHFLHVZKLFKLVLQWHUPHGLDWH between Passalora and Ramularia, is not quite clear. The general Illustrations %UDXQ ¿J %UDXQ habit of this species is rarther ramularioid, but the stromata and ¿J conidiophores are pigmented. Results of molecular sequence DQDO\VHVZRXOGEHKHOSIXOWRHOXFLGDWHWKHWUXHJHQHULFDI¿QLW\ Pseudocercospora Key to Pseudocercospora species on Adoxaceae 6WURPDWDODFNLQJ VXSHU¿FLDOK\SKDHGHYHORSHGEXWZLWKRXWVROLWDU\FRQLGLRSKRUHV FRQLGLRSKRUHVIDVFLFXODWH± PORQJSOXULVHSWDWH RQSambucus javanica, Asia ...... P. ebulicola 6WURPDWDGHYHORSHG±PGLDP VXSHU¿FLDOK\SKDHODFNLQJRULISUHVHQW ZLWKFRQLGLRSKRUHVZKLFKDUHPXFKVKRUWHU±PDQGRQO\±VHSWDWH ...... 2 6XSHU¿FLDOK\SKDHZLWKVROLWDU\FRQLGLRSKRUHVGHYHORSHG±PORQJ±VHSWDWH geniculate, i.e. proliferation sympodial; on Viburnum spp...... P. tinea 6XSHU¿FLDOK\SKDHZLWKVROLWDU\FRQLGLRSKRUHVODFNLQJ FRQLGLRSKRUHVORQJHU±P aseptate to pluriseptate (if short and mostly aseptate, then non-geniculate, proliferation percurrent, with annellations ...... 3 &RQLGLRSKRUHVVXEF\OLQGULFDOQRQJHQLFXODWHSUROLIHUDWLRQSHUFXUUHQWZLWK¿QHDQQHOODWLRQV UDWKHUVKRUW± ± î± ± P ± VHSWDWH FRQLGLDF\OLQGULFDO¿OLIRUPRURFFDVLRQDOO\ somewhat obclavate; on Viburnum spp...... P. viburnigena Conidiophores at least partly geniculate-sinuous, proliferation sympodial, without annellations RURFFDVLRQDOO\ZLWKDVLQJOHHQWHUREODVWLFUHMXYHQDWLRQPXFKORQJHU± ± P 0–3-septate or pluriseptate throughout; conidia obclavate-cylindrical to somewhat fusiform ...... 4 VOLUME 6 · NO. 2 417 418 ARTICLE &RQLGLDZLGHU±P±VHSWDWH RQ &RQLGLDREFODYDWHF\OLQGULFDO±PORQJ±VHSWDWH &RQLGLD±PZLGH±VHSWDWH RQ on orevendarkened-refractive;conidiophoresfascicluate;on Conidiogenouslociinconspicuous tosubconspicuous,i.e.unthickenedbutsomewhatrefractive 9 (6) &RQLGLDF\OLQGULFDO¿OLIRUP±î±P±VHS Asiancollectionson North American collections=var. orevendarkened-refractive;on Conidiogenouslociinconspicuoustosubconspicuous,i.e.unthickenedbutsomewhatrefractive 8 (7) ( butsomewhatrefractiveorevenslightlydarkened-refractive;on ordisarticulating,conidiogenouslociinconspicuoustosubconspicuous,i.e.unthickened, &RQLGLDK\DOLQHRUVXEK\DOLQH¿QDOO\VRPHWLPHSDOHROLYD 0–3-septate or pluriseptate throughout; conidia obclavate-cylindrical to somewhat fusiform ...... 0–3-septateor pluriseptatethroughout;conidia obclavate-cylindricaltosomewhat fusiform RURQO\ZLWKDVLQJOHHQWHUREODVWLFUHMXYHQDWLRQPXFKORQJH &RQLGLRSKRUHVORQJHUDERXW±PDYHUDJH!P ...... orsubhyalineand/orthin-walled,conidiaatleastparlycatenatedisarticulating &RQLGLRSKRUHVUHODWLYHO\VKRUWXVXDOO\±PDYHUDJH &RQLGLDQDUURZHU±PLIZLGHU>± ± P@FRQLGLDFRQV rocsoal oehtocaae ...... oroccasionallysomewhatobclavate UDWKHUVKRUW± ± î± ± P ± VHSWDWH FRQLGLD &RQLGLRSKRUHVVXEF\OLQGULFDOQRQJHQLFXODWHSUROLIHUDWL WRP RQ usuallypaletomediumolivaceousorolivaceous-brown,wallthinsomewhatthickened, &RQLGLDEURDGO\REFODYDWHRUREFODYDWHF\OLQGULFDO± &RQLGLDORQJHU±P± ± VHSWDWH RQRWKHUKRVWV aseptatetopluriseptate (ifshortandmostlyaseptate,thennon-geniculate,proliferationpercurrent, with annellations ...... geniculate,i.e. proliferationsympodial 6XSHU¿FLDOK\SKDHZLWKVROLWDU\FRQLGLRSKRUHVODFNLQJ FRQLGLR ...... 6XSHU¿FLDOK\SKDHZLWKVROLWDU\FRQLGLRSKRUHVGHYHORSHG± Viburnum unbranched; 6WURPDWDGHYHORSHG±PGLDP VXSHU¿FLDOK\SKDHODFNLQJ on on 6WURPDWDODFNLQJ VXSHU¿FLDOK\SKDHGHYHORSHG FRQLGLRSKRUHV Sambucus Tabular keytoPseudocercosporaspecieson Adoxaceae accordingtohostgenera rejuvenationleavingacoarseannellation;on ZDOOVWKLFNHQHGWRPPHGLXPWRGDUNEURZQRFFDVLRQDOO\ &RQLGLDVKRUW±î±P±VHSWDWHSLJPHQWHG FR Conidiogenous lociconsistentlyinconspicuous;conidiophoresfasciculatetocoremioid; Conidiogenous lociinconspicuous(conidiophoresusuallyinsmallfascicles,2–15,divergent, Conidia formedsingly...... Conidiophores atleastpartly geniculate-sinuous, proliferationsympodial,withoutannellations Viburnum Viburnum Sambucus javanica sect. spp...... spp. Viburnum Sambucus Opulus Viburnum sargentii sa ...... , Asia spp., North America ...... spp.,North America ), mainly spp., northern hemisphere ...... spp.,northernhemisphere Sambucus V. opuluss.lat varia Viburnum cylindricum Viburnum ; conidiophoresinlarger, sensefascicles,to50perfascicle, =var. sp ...... spp. Braun Viburnum nudum viburni-sargentii s. ni ...... sp.,India . (includingsubsp. RQSHUFXUUHQWZLWK¿QHDQQHOODWLRQV et al WDWH RQ FHRXVVROLWDU\RULQVKRUWFKDLQV î±P±VHSWDWH P ...... 11 ...... 9 ...... QLGLRSKRUHVLQGLYHUJHQWIDVFLFOHV . SKRUHVORQJHU±P , China ...... , China LVWHQWO\SDOHXVXDOO\K\DOLQH PORQJ±VHSWDWH ...... 5 ...... Sambucus ORRVHO\IDVFLFXODWHVLPSOHWREUDQFKHG U± ± P F\OLQGULFDO¿OLIRUP Viburnum erosum ZLWKDVLQJOHHQWHUREODVWLF FRQLGLRSKRUHVPRVWO\LQGHQVHUIDVFLFOHV , North ...... America ...... 7 ...... ); on Viburnum calvescens Viburnum spp...... spp. spp. and p...... spp...... trilobum ...... 8 ...... 2 ...... ) ...... P. viburni-cylindrici P. caprifoliacearum ...... 10 ...... P. viburni-erosi P. depazeoides P. depazeoides P. depazeoides P. viburni-nudi P. viburnigena P. viburnicola IMA FUNGUS P. ebulicola .... 6 ...... P. opuli P. varia P. tinea . 3 Cercosporoid fungi 4 &RQLGLDEURDGO\REFODYDWHRUREFODYDWHF\OLQGULFDO±î±P±VHSWDWH ARTICLE usually pale to medium olivaceous or olivaceous-brown, wall thin to somewhat thickened, WRP RQVibrunum spp., North America ...... P. viburnicola &RQLGLDQDUURZHU±PLIZLGHU>± ± P@FRQLGLDFRQVLVWHQWO\SDOHXVXDOO\K\DOLQH or subhyaline and thin-walled ...... 4 &RQLGLRSKRUHVUHODWLYHO\VKRUWXVXDOO\±PDYHUDJH P ...... 5 &RQLGLRSKRUHVORQJHUDERXW±PDYHUDJH!P...... 7 &RQLGLDVKRUW±î±P± ± VHSWDWHSLJPHQWHG FRQLGLRSKRUHVLQGLYHUJHQWIDVFLFOHV ZDOOVWKLFNHQHGWRPPHGLXPWRGDUNEURZQRFFDVLRQDOO\ZLWKDVLQJOHHQWHUREODVWLF rejuvenation leaving a coarse annellation; on Viburnum nudum, North America ...... P. viburni-nudi &RQLGLDORQJHU±P± ± VHSWDWH RQRWKHUKRVWV...... 6 6 (5) Conidia solitary, in short chains or disarticulating, conidiogenous loci inconspicuous to subconspicuous, i.e. unthickened, but somewhat refractive or even slightly darkened-refractive; on Viburnum spp. (Viburnum sect. Opulus), mainly V. opulus s. lat. (including subsp. calvescens and trilobum) ...... P. opuli Conidia solitary; conidiogenous loci inconspicuous, not darkened-refractive (conidiophores usually in small fascicles, 2–15, divergent, North American collections = var. varia; conidiophores in larger, sense fascicles, to 50 per fascicle, Asian collections on Viburnum sargentii = var. viburni-sargentii); onViburnum spp...... P. varia &RQLGLDF\OLQGULFDO¿OLIRUP±î±P±VHSWDWH RQViburnum erosum ...... P. viburni-erosi &RQLGLDREFODYDWHF\OLQGULFDO±PORQJ±VHSWDWH...... 8 &RQLGLD±PZLGH±VHSWDWH RQViburnum cylindricum, China ...... P. viburni-cylindrici &RQLGLDZLGHU±P±VHSWDWH RQViburnum sp., India ...... P. caprifoliacearum Pseudocercospora caprifoliacearum (C. Gupta et WRPZLGHSOXULVHSWDWHWKURXJKRXWROLYDFHRXVEURZQZDOO al.) Kamal, Cercosporoid Fungi of India: 159 (2010). WKLQZDOOHGWRVRPHZKDWWKLFNHQHGWRPDWWKHYHU\EDVH (Fig. 48) RFFDVLRQDOO\WRPVPRRWK FRQLGLRJHQRXVFHOOVLQWHJUDWHG Basionym: Phaeoisariopsis caprifoliacearum C. Gupta et al., terminal, occasionally intercalary, sympodially proliferating, with Perspectives in Mycological Research, Prof. G.P. Agarwal several conidiogenous loci, inconspicuous or visible as truncate Festschrift Volume 1: 9 (1987). tips or shoulders, subdenticulate, but not thickened. Conidia Synonym: Pseudocercospora khasiana B.K. Gupta & solitary, obclavate-cylindrical, broadly fusiform, straight to slightly Kamal, Perspectives in Mycological Research, Prof. G.P. FXUYHG DERXW ± î ± P ±VHSWDWH VXEK\DOLQH WR Agarwal Festschrift Volume 1: 25 (1987) [holotype: India: pale olivaceous or olivaceous-brown, thin-walled, smooth, Meghalaya, Shillong, on Viburnum sp., B. K. Gupta KB 58 DSH[REWXVHEDVHREFRQLFDOO\WUXQFDWH±PZLGHKLOD (K(M) IMI 274850)]. unthickened, not darkened. Literature: Braun (1992: 219), Kamal (2010: 230). Holotype: India: Uttar Pradesh: Gorakhpur, on Viburnum sp., Adoxaceae, Jan. 1985, C. Gupta (K(M) IMI 294111). Isotype: Illustrations: Gupta et al ¿J *XSWD .DPDO GPU, KC 62. ¿J Host range and distribution: On Viburnum spp., Adoxaceae, Description: Leaf spots amphigenous, subcircular to angular- Asia (India, Meghalaya, Uttarakhand, Uttar Pradesh). irregular, 2–20 mm diam, brown, olivaceous-brown, later JUH\LVKEURZQWRGLQJ\JUH\PDUJLQLQGH¿QLWHRU with narrow to Notes: Pseudocercospora caprifoliacearum and P. khasiana, broad darker border or halo. Caespituli amphigenous, scattered, both described from India on Viburnum sp., are morphologically ¿QHO\ SXQFWLIRUP EURZQ WR EODFNLVK Mycelium immersed, EDUHO\GLVWLQJXLVKDEOHDQGXQGRXEWHGO\FRQVSHFL¿F7KHWZR composed of subhyaline to pale brown, septate, thin-walled, species have been simultaneously published in a single smooth hyphae. Stromata DERXW±PGLDPGDUNEURZQ book. It is proposed to give precedence to Phaeoisariopsis VXEVWRPDWDO FHOOV ± P GLDP Conidiophores in loose to caprifoliacearum, and to reduced P. khasiana to synonymy. dense, almost coremoid fascicles, 7–18, arising from stromata, Braun (1992) examined type material of Pseudocercospora though stomata, erect, when coremioid then dense below and khasiana and compared it with P. viburni-cylindrici, described ORRVHUWRZDUGVWKHDSH[VWUDLJKWWRÀH[XRXVXQEUDQFKHGRURQO\ from China. The two species are morphologically very similar. UDUHO\EUDQFKHGVXEF\OLQGULFDO¿OLIRUPZLGWKXQLIRUPWKURXJKRXW Therefore, Braun (1992) proposed to reduce P. khasiana or tips somewhat swollen, geniculate-sinuous in the upper half, to synonymy with P. viburni-cylindrici, a treatment followed DERXW±PORQJDQG±PZLGHDSH[VRPHWLPH by Crous & Braun (2003) and Kamal (2010). Differences VOLUME 6 · NO. 2 419 420 ARTICLE %DU P holotype). Fig. 48. Basionym (Fig. 49) MycoBank MB814570 Pseudocercospora depazeoides molecular datawillbeavailable. as separatespecies,atleastfortheinterimuntilculturesand prefer tomaintain caprifoliacearum dense orevencoremioidconidiophorefascicles,similarto of be variationwithinasinglespecies. The Chinese typematerial between ChineseandIndiancollectionswereconsideredto Cercospora penicillata Synonyms P. viburni-cylindrici Crous, Rabenh., Herb.Viv. Mycol.587(HAL)]. Italy [ Viv. Mycol Bot lectotype ., : Vercellis, on sér Pseudocercospora caprifoliacearum A. : : Passalorapenicillata Exosporium depazeoides comb. nov. Conidiophore fascicle. . 3, ., no.587(1857),withdescription onlabel ( designated here 11 EXWQDUURZHUFRQLGLD±PZLGH:H : 364(1849). P. caprifoliacearum ischaracterisedbyhavingloosetooften (Ces.) Fresen., Sambucus nigra B. , MycoBank,MBT202789): Conidiophore tips. Ces., inRabenh., Beitr. Mykol Desm., (inclusing (Desm.) U.Braun& , 1856, (K(M) IMI294111, Ann. Sci.Nat., V. deCesati P .: 93(1863). . C. khasiana Conidia. Herb. Braun P. P. ) , et al Cercospora sambucina Cercospora depazeoides Cercospora ticinensis Cercospora depazeoides Cercospora depazeoides Cercospora sambuci Pseudocercospora sambucigena Phaeoramularia penicillata Cercospora depazeoides H DF \RK,D 6YO +HUE 6ăYXO ,WDO 0\FRWK 6DFF ' 9HQ 2070. Roum.,FungiSel.Gall. Exs.158,3596.Sacc.,Mycoth. & Scheuer, Reliqu.Petr. 943.Rabenh.,FungiEur. Exs.879, Petr., FungiPolon.Exs.19.Petr., Mycoth.Gen.1414.Poelt Alban. Bosn.Exs.7.Petr., Fl.Bohem.Morav. Exs.Pilze508. Krypt. Exs.727.Linh.,Fungi Hung.Exs.294.Petr., Fungi Exs. 898,899.Krieger, Schädl.PilzeKulturgew. 193,898. Maurilaun, Fl.Exs. Austr.-Hung. 788.Krieger, FungiSaxon. 1749. Kabát&Bubák,FungiImperf.Exs.97.Kernervon Everh., FungiColumb.599.Ellis&North Amer. Fungi Ess. 336.Cavara,FungiLongobard.Exs.245,984.Ellis & Exsiccatae 4), Crous &URXV ¿J %DNDORYD %RULVRYD SO 8WLO(VV¿JV±(OOLV Illustrations (2010: 39–40),Crous (2003: 156),Guo Ellis (1976:247),Braun&Mel’nik(1997:55),Crous (1937: 235),Chupp(1954,100,103),Katsuki(1965:16), Gonzáles Fragoso(1927:252),Vassiljevsky &Karakulin Literature . MBT202790): 17 III, no.280(1875). Revista Agron OSU 35059,PAD]. Util. Ess.336,e.g.BPI441927,CUP, FH,K,MICH15376, isolectotypes Cavara, FunghiParas.PianteColt.Util.Ess.336(HAL); Pavia, BotanicGarden,on ( Paras Sacc., CUP 41160, NY 2343037]. 1883, nigra [Gagry], upperwaterfallZhoekvaryRiver, on Rast Pereira (Traz-os-Montes), on Monogr 21 Aug. 1921, 21 Aug. Pennsylvania Schub., Phytopathol. Sin Stevens on designated here : 1166 (1883)[lecotype( Sambucus canadensis . , 9 Aug. 1912, . : Saccardo(1886:468 W. A. Kellerman401 et al 6 Syll. Fung ]. : Briosi&Cav., FunghiParas.PianteColt. Util. Piante Colt.Util.Ess . : 108(1912)[ : Briosi&Cavara,FunghiParas.PianteColt. (ILL 15175); 11 Mycotaxon ¿J : 59(1927)[ : Briosi&Cavara,FunghiParas.PianteColt. : DauphonCounty, on et al. USA O. E.Jennings . . . 1 12 4 (2005:279–280),Bakalova&Borisova et al : 59(1903)[ : : 469(1886). A. Elenkin Cavara, inBriosi& , MycoBank,MBT202792): : 13(1982). F. Stevens&King, Ohio isotypes Ellis &Kellerm., 92 var. (Desm.) Sacc., var. . (2013:105–106). var. Sambucus nigra type (Ces.) X.J.Liu& Y.L. Guo, : 400(2005)[ holotype , on gagrensis amphigena designated here [ : sambucina mexicana (NY 2343036); ., no.336(1900)[ Georgia – , Acc. 6736(NY 142641)]. Sambucus canadensis : CUP 14660,NY 2343299]. Sambucus nigra (LE158291)]. 469), Lindau(1910:134), U. Braun,Crous&K. ¿J' %UDXQ type : ot Rica Costa Elenkin &Ohl, : Sambucus pubens Mycoth. Ven ], 7Jul.1923, : Sousa daCâmara, Portugal (Ellis &Kellerm.) Abkhazia , Dec.1902, Amer. Naturalist holotype isolectotypes IMA FUNGUS Illinois Biol. , MycoBank, Sambucus , Briosi& : Cartago, : Chaves lectotype : Gagra : ., Funghi Bolez. Cent USA Italy , Jul. Acta F. L. L. F. A. . : : : , Cercosporoid fungi 4 ARTICLE Fig. 49. Pseudocercospora depazeoides (Rabenh., Herb. Viv. Mycol., no. 587, lectotype of Passalora penicillata, HAL). A. Conidiophore fascicles. B. Conidia. %DU P Mycol. Rom. 195. Scheuer, Mycoth. Graec. 433. Seym. & PGLDPLPPHUVHGWRVRPHZKDWHUXPSHQWPHGLXPWRGDUN Earle, Econ. Fungi 476. Syd., Mycoth. March. 1290, 2177. EURZQ FRPSRVHG RI VZROOHQ K\SKDO FHOOV ± P GLDP Syd., Mycoth. Germ. 1045, 1774, Thüm., Mycoth. Univ. 984. Conidiophores in small to large fascicles, loose to very dense, Vill, Fungi Bav. 996. Zahlbruckner, Krypt. Exs. 727 A, B. occasionally even almost coremioid, arising from stromata, emerging through stomata or erumpent, erect, straight, Description: Leaf spots amphigenous, subcircular to VXEF\OLQGULFDO WR ÀH[XRXV VRPHZKDW JHQLFXODWHVLQXRXV DQJXODULUUHJXODU ± PP GLDP VRPHWLPHV FRQÀXHQW XQEUDQFKHG± ± î±PVKRUWFRQLGLRSKRUHV DQG ODUJHU DW ¿UVW SDOH JUHHQLVK RU JUH\LVK JUHHQ ODWHU 0–1-septate, longer ones pluriseptate throughout, pale EURZQLVK ¿QDOO\ JUH\ WR JUH\LVK ZKLWH RIWHQ ]RQDWH olivaceous to dark olivaceous-brown or brown, tips often margin narrow, often formed as marginal line, somewhat paler, thin-walled, smooth or almost so; conidiogenous cells raised, dark olivaceous-brown to blackish or dark purplish integrated, terminal, occasionally conidiophores reduced violet. Caespituli amphigenous, punctiform, dark brown to WR FRQLGLRJHQRXV FHOOV ± P ORQJ V\PSRGLDOO\ RU blackish, scattered. Mycelium internal; hyphae branched, occasionally percurrently proliferating, conidiogenous VHSWDWH ± P ZLGH VXEK\DOLQH WR SDOH ROLYDFHRXV loci inconspicuous, unthickened or almost so and not thin-walled. Stromata well-developed, substomatal to darkened to subconspicuous by being subdenticulate, intraepidermal, subglobose to somewhat irregular, 10–100 WLSV WUXQFDWH RU VXEWUXQFDWH ± P ZLGH RU VOLJKWO\ VOLUME 6 · NO. 2 421 422 ARTICLE Culture characteristics darkened-refractive. RU XQWKLFNHQHG KLOD almost so,notdarkenedtosomewhatrefractiveorslightly ZLGH P ± ± ± WUXQFDWH apex obtuse,baseshorttooccasionallylongobconically thin-walled, smooth,occasionallysomewhatrough-walled, hyaline, subhyalinetopaleolivaceousorolivaceous-brown, ± ± î ± ± ± P ± ± ± VHSWDWH Conidia SDUDFHUFRVSRURLGYLVLEOHDVPLQXWHFLUFOH±PGLDP refractive todarkened-refractive,infrontviewoccasionally [ Host rangeanddistribution W. Crous Netherlands Epitype Isolectotypes Desmazières France Lectotype olivaceous-grey (surface)andolivaceous-black(reverse). with moderateaerialmycelium,palegreenishgreyto with smoothtoslightlyirregularmargin,prominentlyconvex, diam after30dinthedarkat24°C,circulartosubcircular, glauca [ species isinvolved whichbelongsin fungi on collections involvedledtoa reassessment ofcercosporoid et al. of the griseola are reminiscentofthelocuscharacters morphological rangeoftheconidiogenouslocithisspecies quite indistinctconidiogenousloci. The variabilityand Pseudocercospora sambucigena Cercospora s.str & Braun(2003)maintained colourless oralmostsotopaleolivaceousconidia,Crous circle infrontview(paracercosporoid),aswellusually cell bybeingrefractive,sometimesalsovisibleasminute darker thanthesurroundingwallofconidiogenous conspicuous bybeingveryslightlythickenedorsomewhat the conidiogenousloci,rangingfrominconspicuoustomore been confusedandmisinterpreted.Owingtothestructureof Notes West Virginia, Wisconsin). New Jersey, Ohio,Pennsylvania,SouthCarolina,Washington, Kansas, Louisiana,Maine,Maryland,Mississippi,New York, (Canada; Mexico;USA,Connecticut,Florida,Illinois,Iowa, America andWest Indies(CostaRica,Haiti),North America Spain, UK,Ukraine,former Yugoslavia), NewZealand,Central Netherlands, Poland,Portugal,Romania,Russia,Slovakia, Republic, Denmark,Germany, Greece,Hungary, Italy, Latvia, Georgia), Europe(Austria,Belgium,Bulgaria,Cyprus,Czech Russia, FarEast), Australia, Caucasus(Armenia, Azerbaijan, intermedia var. insularis buergeriana 2013). Re-examinationsoftypes andnumerousother 7KHJHQHULFDI¿QLW\RI , Paracercospora : on ( Sambucus kamtschatica (now solitary, obclavate-cylindrical,straighttocurved, designated here (CBSH-20391). ( ], designated here : Milingerwaard,on , Desm.,Pl.Crypt.France1849(NY 2343300). : Desm.,Pl.Crypt.France1849,e.g.FH,G,PC. Sambucus Pseudocercospora griseola Sambucus nigra ., andBraun&Crous(2005)introduced spp.Itemergedveryclearlythat asingle , nigra : complex(Braun Colonies sp.), , Ex-epitype culture , MycoBank,MBT202794). , mexicana pubens Adox C. depazeoides : On Cercospora depazeoides , MycoBank,MBT202793): , autumn, Sambucus nigra on MEA reaching16mm aceae, Asia (China,Japan, forcollectionswith Sambucus , ], racemosa chinensis Pseudocercospora et al ) andthespecies Phaeoisariopsis : CBS126000. J. B.H. . 2013,Crous as speciesof ( canadensis canadensis , williamsii williamsii , 2007, , ebulus The Braun has P. P. , et al WRVRPHZKDWWRUWXRXVVLPSOHRUEUDQFKHG± î± stomata, erect,subcylindrical, curved,geniculate-sinuous internal hyphaeorsmallhyphal aggregations,through Conidiophores small, onlyafewsubstomatal swollenhyphalcells,brown. P ± wide, septate,paleolivaceous. EUDQFKHG K\SKDH VXSHU¿FLDO H[WHUQDO SDUWO\ RU brownish, formingirregularpatches. surface. ODWHUJUH\LVKZKLWHPDUJLQLQGH¿QLWHRUEURZQRQWKHXSSHU spots, 1–8mmdiam,palebrown,yellowishbrownto discolorations, yellowishtobrownish,orcircularirregular Description ¿J *XR Illustrations Braun (2003:168). & Hsieh(1995:47–48),Guo Literature Basionym (Fig. 50) Pseudocercospora ebulicola the allocationof genus penicillata serious consequencesongeneticlevelsince reallocation of ¿[WKHSK\ORJHQHWLFSRVLWLRQDQGDI¿QLW\RIWKLVVSHFLHV7KH culture andsequencesobtainedfromthismaterialhelpto Veneta 280(1875).Thedesignatedepitype,theex-epitype ´EXWLWZDV¿UVWYDOLGO\SXEOLVKHGLQ0\FRWKHFD combination isusuallycitedas“ FRQ¿QHWKLVQDPHWRWKHFHUFRVSRURLGHOHPHQW6DFFDUGR¶V introduction ofthecombination we followSaccardo’s treatmentontheoccasionofhis included intheverymeagreoriginaldescription.However, cercosporoid aswellthealternarioidhyphomycetewere also analternarioidfungus.Itcannotbeexcludedthatthe contains besidesthetypicalcercosporoidhyphomycete by Crous the depazeoides sambucigena (Braun in thecurrentsenseandcircumscriptionofthisgenus Ramularia sambucina HAL, hasbeenexaminedandturnedouttobeamixtureof being made. with undesirable andrequiresaproposaltoconserve older heterotypicsynonymof molecular sequenceanalyses,renders based onmorphologicalreassesmentsandresultsof . Mycol. Pap Nat. Hist.Soc Lectotype materialof Eriksson, FungiParas.Scand.Exs.42,depositedat Pseudocercospora C. apii Cercospora et al : Chupp(1954:100),Hsieh&Goh(1990:50),Guo et al Caespituli : Cercosporaebulicola (i.e. +VLHK *RK ¿J *XR +VLHK : . 2013,Crous . The phylogeneticpositionofthisspeciesin as conservedtypeunder Art. 14.9,whichis mustbereducedtosynonymywith . (2013). C. depazeoides Leaf spots C. depazeoides . in loosefascicles,3–15, arising from 140 . 13 C. depazeoides (Braun 1995,Braun : 139(1934). : 143(1976). hypophyllous, effuse, olivaceousto and etal. FODGHKDVUHFHQWO\EHHQFRQ¿UPHG Cladosporium herbarum et al. ¿J etal. ) beingthetypespeciesof Cercospora depazeoides Pseudocercospora to Exosporium depazeoides W. Yamam., PKJQX LQGH¿QLWH DPSKLJHQRXV 2013). Nuovo Giorn.Bot.Ital Stromata (W. Yamam.)Deighton, (1998:60–61),Crous& Pseudocercospora to Pseudocercospora et al Pseudocercospora Mycelium Cercospora lacking orvery Trans. Sapporo . 2013). Thus, IMA FUNGUS Cercospora Cercospora , whichis . internal has and . an 8 P. P. , : Cercosporoid fungi 4 ARTICLE Fig. 51. Pseudocercospora opuli (Fuckel, Fungi Rhen. Exs. 118, HAL, lectotype). A. Conidiophore fascicles and solitary conidiophore. B. Conidiophore tips. C.&RQLGLD%DU P8 Host range and distribution: On Sambucus (javanica [formosana]), Adoxaceae, Asia (China, Sichuan, Zhejiang; Japan, Taiwan). Fig. 50. Pseudocercospora ebulicola (CUP-039732, lectotype). Notes: Chupp (1954) reduced Cercospora ebulicola A. Conidiophore fascicle. B. Solitary conidiophore arising from a to synonym with C. catenospora which is, however, VXSHU¿FLDOK\SKDC. Conidiophore tips. D. &RQLGLD%DU P incorrect. Cercospora ebulicola is a species of the genus Pseudocercospora whereas C. catenospora is a Phaeoramularia-like species of Passalora. P ±VHSWDWH ROLYDFHRXV WR ROLYDFHRXVEURZQ WKLQ walled, smooth; conidiogenous cells integrated, terminal Pseudocercospora opuli (Fuckel) U. Braun & Crous, and intercalary, conidiogenous loci inconspicuous or Mycophaerella and Anam.: 299 (2003). subdenticulate, but always unthickened, not darkened. (Fig. 51) Conidia solitary, obclavate-cylindrical, straight to curved, Basionym: Cercospora penicillata f. opuli Fuckel, Fungi ± î ± P ±VHSWDWH VXEK\DOLQH \HOORZLVK Rhen. Exs. 118 (1863). to pale olivaceous, thin-walled, smooth, apex obtuse to Synonyms: Cercospora opuli (Fuckel) Höhn., in Kabát & subacute, base subtruncate to short obconically truncate, Bubák, Fungi. Imperf. Exs. 445 (1907). ±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG Cercospora viburni Sacc., in Sydow, Mycoth. March. 2773 (1889), nom. nud. (ICN 38.1). Lectotype (designated here, MycoBank, MBT202795): Pseudocercospora viburni U. Braun, Nova Hedwigia 55: Taiwan: Sozan, on Sambucus javanica, 3 Dec. 1933, W. 219 (1992) [holotype: China: Hubei: Shennongjian, Yamamoto (CUP-039732). Isolectotype: HMAS 5197, K(M) on Viburnum opulus subsp. calvescens [sargentii var. IMI 7791. Topotypes: 20 Nov. 1933 (BPI 436018, 436019). calvescens], 5 Aug. 1984, Y. L. Guo 266 (HMAS 47828)]. Pseudophaeoramularia opuli (Fuckel) U. Braun, Trudy Bot. VOLUME 6 · NO. 2 423 424 ARTICLE Passalora viburni-sargentii Germany Lectotype not toslightlyrefractiveordarkened-refractive. XQWKLFNHQHG KLOD ZLGH P ± WUXQFDWH REFRQLFDOO\ VKRUW short conicallytruncateincatenateconidia,base to of primaryandsolitaryconidiaobtuse,rounded,truncate to subhyaline, laterpaleolivaceous,thin-walled,smooth,apex occasionally somewhatconstrictedatthesepta,hyaline, FXUYHG ± ± ± î ± ± ± P±VHSWDWH obclavate-subcylindrical tocylindrical,straightsomewhat catenate, insimple,shortacropetalchainsordisarticulating, DVPLQXWHFLUFOHVDERXW±PGLDP refractive, butunthickened,infrontviewsometimesvisible ORQJ being truncateandsomewhatrefractiveordarkened- P ± DERXW conidiogenous lociinconspicuoustosubconspicuousby FHOOV FRQLGLRJHQRXV WR UHGXFHG conidiogenous cellsintegrated,terminalorconidiophores WR ROLYDFHRXV SDOH olivaceous-brown andwallssomewhatthickened,smooth; ODWHU WKLQZDOOHG DQG VXEK\DOLQH ¿UVW P ± ± ± î aseptate orsparinglyseptate,usuallywith0–3septa,at ± ± EUDQFKOHW VLQJOH D ZLWK moderately geniculate-sinuous,unbranchedoroccasionally through stomataorerumpent,erect,straight,subcylindricalto fascicles, loosetodense,arisingfromstromata,emerging somewhat thickened. EURZQFRPSRVHGRIVZROOHQK\SKDOFHOOV±PGLDPZDOOV internal. punctiform, scattered,darkbrowntoblackish. white, withnarrowdarkmargin. GXOO ]RQDWH greenish, ochraceoustobrown,centrelatergreyish VRPHZKDW RFFDVLRQDOO\ ODUJHU DQG FRQÀXHQW somewhat angular-irregular, 1–8mmdiam,sometimes Description Mycoth. Graec.382.Smarods,FungiLat.Exs.448. 0\FRWK*HQ6ăYXO+HUE0\FRO5RP6FKHXHU Fungi Imperf.Exs.445.Liro,Mycoth.Fenn.295.Petr., Exsiccatae ¿J *XR ¿J %UDXQ ¿J %UDXQ 0HO¶QLN DSSHQGL[ Illustrations (1954: 103),Ellis(1976:247),Braun&Mel’nik(1997:19,77). (1910: 136),Vassiljevsky &Karakulin(1937:236),Chupp Literature Phaeoramularia penicillata Misapplied names (FH, G,K(M),etc). 118] (HAL). Viburnum opulus 161 (2012)[ Inst. im.V.L. Komarova on Phytopathol. Sin (1886: 468)andChupp(1954:103). calvescens Viburnum opulus Stromata : Saccardo(1886:468),Lindau(1910:136),Ferraris : (designatedbyBraun& Mel’nik 1997:20): : Fuckel,FungiRhen.Exs.118. Kabát&Bubák, : &KXSS (OOLV ¿J$ Rheinland-Pfalz Isolectotypes Leaf spots ], 5 Aug. 1984, , summer1863[Fuckel, holotype VXEVWRPDWDORULPPHUVHG±PGLDP : . Cercospora penicillatasensu 12 Conidiophores (4): 13(1982). subsp. : Fuckel, : 20 (Ces.) X.J.Liu& Y.L. Guo, : “in sylvaHostrichiensi”,on “in : amphigenous, subcircularto China Y.L. Guo, Y. L.Guo266 : 19(1997). calvescens Caespituli : Fungi Rhen.Exs Hubei in smalltofairlylarge Mycosystema Conidia Fungi Rhen.Exs : Shennongjian, (HMAS47828)]. amphigenous, [ sargentii Saccardo solitary to Mycelium . 118, Acta Braun var. 31 : . et al reduced tosynonymywith Passalora FODVVL¿HG loci andhilaofthisspeciesasintermediatebetween 0HO¶QLN %UDXQ LQ %UDXQ XQWKLFNHQHG refractive ordarkened-refractive,buttheyareconsistently inconspicuous tosubconspicuousbybeingsomewhat smaller units. The conidiogeouslocirangefrombeing are formedsinglyaswellinchainsordisarticulate is unusualandcausedadditionalconfusion. The conidia to followthesecondversion.Themorphologyofthisspecies combination atnewrankaccordingto Art. 41.4.We prefer (see Braun&Mel'nik1997,Crous2003)orasnew Höhnel's name wrong interpretationofthename penicillata Lindau (1910). The proposed combination involved hadalreadybeendiscussedandcorrectedby misinterpretation, althoughtheconfusionofnames the name be thereasonforSaccardo’s (1886)misinterpretationof on opuli morphological reassessments, based onresultsofmolecularsequenceanalysesand genus opulus on depazeoides (syn. confusions andmisinterpretations. history ofthisspeciesiscomplicatedandcharacterisedby but burejaeticum unproven andmightratherpertainto the FarEastofRussia(Egorova2007)isuncertainand Notes Mississippi, Oklahoma,Wisconsin). Rica), North America (Canada;USA,Idaho,Iowa,Kansas, Russia, Slovakia,Sweden,Ukraine),Central America (Costa Great Britain,Italy, Latvia,Poland,Portugal,Romania, Czech Republic,Denmark,Estonia,Finland,Germany, Azerbaijan, Georgia),Europe(Austria,Belarus,Bulgaria, (China, Kazakhstan,Russia, Turkey), Caucasus(Armenia, opulus are homotypicsynonyms.Chinese materialon Braun was reallocatedtothelattergenus(Crous&Braun2003, hosts (Crous &Braun2003)areadditionalunprovenanddoubtful LQ Clement FODGH VXSSRUWHG ZHOO DV phylogenetic studies(Winkworth&Donoghue2005, FRQ¿UPHG EHHQ KDV ZKLFK [ be indistinguishablefromEuropean materialof calvescens opulus Host rangeanddistribution opulus . Sambucus nigra Pseudocercospora viburni A collectionofthisspeciesfrom the Asian partof Turkey Sambucus nigra P. opuli , andFuckel(1870)treatedthefunguson : A recordofthisspecieson of P. opuli subsp. assolespeciesof et al Pseudophaeoramularia subsp. Cercospora penicillata var. et al. and C. penicillata . 2013,Crous , (Liu &Guo1982)wasalsobasedonChupp’s VHHPVWREHFRQ¿QHGWRVSHFLHVRIWKLVVHFWLRQ sargentii ) wasintroducedforacercosporoidfungus does notbelongto opulus calvescens americanum C. opuli Pseudocercospora 2014). not belongingtosect. . Fuckel(1863)added wasnotmentioned,whichseemsto var. , opulus . Chupp(1954)followedSaccardo’s iunmlantana Viburnum canbeinterpretedasnewspecies calvescens et al hasbeenexaminedandfound to : On , C. penicillata Pseudocercospora trilobum subsp. and . 2013). Viburnum now , Pseudophaeoramularia for suchspecies.However, Passalora viburni-sargentii Viburnum and proposedthenew Cercospora penicillata ], calvescens ]), opulus V. burejaeticum Passalora penicillata Opuli ( P. varia , i.e.“f. edule Pseudocercospora Adoxaceae and . The taxonomic The . Phaeoramularia C. penicillata subsp. sect. [ and P. opuli SDXFLÀRUXP IMA FUNGUS V. orientale . [ penicillata opulus Viburnum Viburnum Viburnum trilobum C. opuli Opulus , Asia . was from var. f. ], ” . , Cercosporoid fungi 4 has been examined (Kütahya, on Viburnum opulus, 9 Jul. ARTICLE 1953, ex herb. Petrak, GZU). Pseudocercospora varia is morphologically very close to and confusable with P. opuli, but the conidia are formed VLQJO\7KH ODWWHU VSHFLHV VHHPV WR EH FRQ¿QHG WR VSHFLHV on Viburnum sect. Opuli. The taxonomic meaning of conidial catenation, which occurs in varying degrees in P. opuli, is not quite clear and has to be elucidated by means of molecular methods. Other morphological characters of P. opuli and P. varia are very similar. The two species are treated as separate species, at least tentatively. Pseudocercospora tinea Y.L. Guo & W.H. Hsieh, Mycosystema 7: 124 “1994” (1995). (Fig. 52) Misapplied name: Stigmina tinea sensu Hsieh & Goh 1990. Literature: Hsieh & Goh (1990: 51–52, as Stigmina tinea), Guo et al. (1998: 63–64). Illustrations +VLHK *RK ¿J DV Stigmina tinea *XR ¿J *XR et al. ¿J Description: Leaf spots amphigenous, subcircular to angular- LUUHJXODUDQGYHLQOLPLWHG±PPGLDPRIWHQFRQÀXHQWDQG DORQJWKHOHDIPDUJLQEURZQWRGDUNEURZQPDUJLQLQGH¿QLWH or centre later greyish brown, dingy grey to greyish white, margin dark brown above and paler brown below, sometimes with diffuse yellowish to greyish brown halo. Caespituli amphigenous, punctiform, dark brown. Mycelium internal and H[WHUQDO VXSHU¿FLDOK\SKDHK\SRSK\OORXVHPHUJLQJWKURXJK stomata or arising from the base of conidiophore fascicles, branched, septate, pale olivaceous, thin-walled, smooth, ± P ZLGH Stromata well-developed, substomatal, ±PGLDPEURZQConidiophores in small to moderately large fascicles, divergent to usually dense, arising from Fig. 52. Pseudocercospora tinea EDVHGRQ*XR ¿J A. VWURPDWDWKURXJKVWRPDWDRUVROLWDU\DULVLQJIURPVXSHU¿FLDO Conidiophore fascicles. B. Conidiophores. C. Solitary conidiophores hyphae, erect, straight to curved, subcylindrical, somewhat DULVLQJIURPVXSHU¿FLDOK\SKDHD.&RQLGLD%DU P attenuated towards the tip or irregular in width, geniculate- VLQXRXV XQEUDQFKHG UHODWLYHO\ VKRUW ± î ± P 0–1-septate, pale olivaceous, olivaceous-brown to brownish, Notes: Guo and Hsieh (in Guo 1995) compared this species thin-walled, smooth; conidiogenous cells integrated, terminal with Cercospora tinea, supposed that the two species could or conidiophores often reduced to conidiogenous cells, 5–20 be synonymous, but hesitated to reduce them to synonymy. PORQJFRQLGLRJHQRXVORFLLQFRQVSLFXRXVRUVXEGHQWLFXODWH Therefore, they did not propose a new combination based but always unthickened and not darkened. Conidia solitary, on Cercospora tinea, but preferred to describe a new REFODYDWHF\OLQGULFDO±î± ± P±VHSWDWH species with a Chinese type collection. Hence, the name subhyaline to pale olivaceous or olivaceous-brown, thin- Pseudocercospora tinea is valid. However, as already walled, smooth, apex obtuse to subacute, base subtruncate stated in Braun & Hill (2002), Cercospora tinea (now WR XVXDOO\ REFRQLFDOO\ WUXQFDWH ± P ZLGH KLOD Pseudocercospora viburnigena) is clearly distinct from the unthickened, not darkened. Chinese species by having non-geniculate, only percurrently SUROLIHUDWLQJ FRQLGLRJHQRXV FHOOV ZLWK ¿QH DQQHOODWLRQV Holotype: China: Guangdong Province: Guangzhou, on 6XSHU¿FLDOK\SKDHDUHRQO\UDUHO\IRUPHGPseudocercospora Viburnum macrocephalum, 6 Nov. 1961, Q. M. Ma & X. J. Liu tinea is possibly rather common in China and Taiwan. Two 1079 (HMAS 67254). additional Taiwanese collections on Viburnum sp. have EHHQ H[DPLQHG DQG LGHQWL¿HG E\ & 1DNDVKLPD 78$ Host range and distribution: On Viburnum (luzonicum, 56), and Japanese collections on Viburnum suspensum and macrocephalum, plicatum var. tomentosum [tomentosum], V. plicatum var. tomentosum [V. tomentosum] have been suspensum, Viburnum sp.), Adoxaceae, Asia (China, Anhui, UHYLVHGDQGFRQ¿UPHGDVP. tinea by C. Nakashima (CNS567, Guangdong, Sichuan, Zhejiang; Japan, Taiwan). CNS976). VOLUME 6 · NO. 2 425 426 ARTICLE brownish halo. raised, andsometimeswithochraceous, yellowishbrownto to moderaterlywide,darkbrown margin,sometimessomewhat white, marginindistinct,formed asmarginallineorwithnarrow to mediumdarkbrownorreddish brown,latergreytogreyish LUUHJXODU±PPGLDPVRPHWLPHVFRQÀXHQWDQGODUJHUSDOH Description 280. Ellis&Everh.,N. Amer. Fungi3190. Exsiccatae Hsieh (1995:51),Crous&Braun(2003:418). Literature (Fig. 53a) Basionym Pseudocercospora varia P A. Fig. 53a. Conidiophore fascicles. State Mus.Nat.Hist var. Cheng, varia Pseudocercospora varia : Saccardo(1886:468),Chupp(1954:105),Guo& : Cropr varia Cercospora : Clements&Clements,Crypt.Format.Colorad. : Acta Mycol.Sin Leaf spots Caespituli amphigenous,subcirculartoangular- . B. amphigenous, punctiform, scattered, amphigenous, punctiform, scattered, 35 Conidiophores. : 141(1884). Peck, . 11 var. (Peck) J.K.Bai&M.Y. : 123(1992). varia Rep. (Annual)New York (BPI 442161,isotype). C. Conidia. Bar=10 Braun et al Passalora antipus specimens, e.g.on lonicericola Collections on varia rather infavourof (short petiolesdonotcoincidewith accordance with K\SKDH with solitaryconidiophoresfoundon“ 6XSHU¿FLDO IRXQG EHHQ QRW KDV LGHQWL¿FDWLRQ ¿QDO 3'' EXWVXI¿FLHQWIUXFWL¿FDWLRQIRUD Holotype KLOD ZLGH P ± unthickened, notdarkened. WUXQFDWH REFRQLFDOO\ ORQJ RU VKRUW WR smooth, apexobtuse,occasionallysubacute,basetruncate WKLQZDOOHG VXEK\DOLQH WR K\DOLQH ±VHSWDWH P ± î subacicular, straighttosomewhatcurved,(15–)20–85(–90) circle. occasionally somewhatrefractiveorlocivisibleasminute or subdenticulate,butalwaysunthickenedandnotdarkened, conidiogenous lociinconspicuousorvisiblebybeingtruncate FRQLGLRSKRUHVUHGXFHGWRFRQLGLRJHQRXVFHOOV±PORQJ P WR thick, smooth;conidiogenouscellsintegrated,terminalor VRPHWLPHV EHORZ DOO DERYH P WR WKLFNHQHG mostly palertowardsthetip,wallthintousuallysomewhat olivaceous-brown oryellowishbrown,aboveallbelow, DW¿UVWYHU\SDOHK\DOLQHWRVXEK\DOLQHODWHUSDOHROLYDFHRXV XQEUDQFKHG ± ± ± î± PXVXDOO\±VHSWDWH occasionally somewhatgeniculate-sinuousintheupperhalf, or attenuatedtowardsthetip,mostlynotgeniculate-sinuous, erumpent, erect,straighttosomewhatcurved,subcylindrical rarely dense,arisingfromstromata,throughstomataor P or inusualysmallfasciles,mostly2–15,usuallydivergent, WR WKLFNHQHG ZDOO GLDP P cells globoseorsubglobosetosomewhatirregular, 3–10 subglobose, paletomediumbrownoryellowishbrown, GLDP P ± ZHOOGHYHORSHG WR VPDOO LQWUDHSLGHUPDO RU VPRRWK WKLQZDOOHG ZLGH P ± IRUPHG SUREDEO\ K\SKDH by conidialgermination);hyphaebranched,septate,pale, VXSHU¿FLDO ZLWK FROOHFWLRQV VRPH brown, darkbrowntoalmostblackish. unclear. The collectionsconcernedhavebeenexamined (Gagdil 2005)on to as short chainsandlateralbranchlets. They arereferred opulus Notes Wisconsin, Wyoming). South Carolina, Texas, Vermont, Virginia, West Virginia, Massachusetts, Mississippi,New York, Oklahoma,Oregon, Alabama, Alaska, Florida,Idaho,Illinois,Iowa,Kansas, suspensum Viburnum acerifolium alnifolium Host rangeanddistribution 442161. prunifolium . on : Collectionson P. opuli Conidia and : Lonicera , cassinoides USA , , pubescens V. trilobum (material examined:BPI 442155), other Adoxaceae . RecordsofthisspeciesfromNewZealand solitary, obclavate-cylindrical,cylindrical,rarely : L. japonica New York P. varia VSSDUHDOVREDVHGRQPLVLGHQWL¿FDWLRQV Viburnum lantata (Ellis &Holw.) U.Braun&Crous. L. ciliata V. burejaeticum , Aug., , Viburnum edule , dilatatum have atendencytoformconidiain UD¿QHVTXLDQXP . The hostidentity isalsouncertain , North America (Canada;USA, : Albany County, EastBerne,on C. H.Peck (BPI 442168–442170),refer to belong to : On , lentago and V. lantana Viburnum . Otherrecordsof Conidiophores Stromata V. lantana Mycelium (NYS). [ V. carlesii SDXFLÀRUXP 9 Pseudocercospora var. , odoratissimum DI¿QH IMA FUNGUS but theyare Isotype ( substomatal substomatal ” arenotin acerifolium internal (in internal (in arequite [ solitary solitary DI¿QH : BPI ], C. V. V. ], , , Cercosporoid fungi 4 having much larger, dense fascicles of conidiophores. The ARTICLE taxonomic status of the Asian collections is not quite clear DQG LQ XUJHQW QHHG RI PROHFXODU FRQ¿UPDWLRQ ,W FDQQRW EH excluded that this is a separate species, but for the interim we prefer a conservative treatment as variety. Pseudocercospora viburnicola U. Braun, sp. nov. MycoBank MB814559 (Fig. 54) Diagnosis: Differs from Pseudocercospora viburni-nudi in KDYLQJ PXFK ORQJHU XVXDOO\ REFODYDWH FRQLGLD ± î ± P ZLWK ± VHSWD DQG VRPHZKDW WKLFNHQHG ZDOOV and subconspicuous conidiogenous loci. Pseudocercospora caprifoliacearum, known from India on Viburnum sp., has ORQJHUFRQLGLRSKRUHV±PDQGWKHFRQLGLDDUH± PZLGHDQGWKLQZDOOHG Description: Leaf spots amphigenous, subcircular to angular- LUUHJXODU±PPGLDPPHGLXPWRGDUNEURZQ¿QDOO\ZLWK pale centre, greyish brown to dingy greyish white, margin LQGH¿QLWHRUQDUURZGDUNHURUVXUURXQGHGE\GDUNHUYHLQV occasionaly somewhat raised. Caespituli amphigenous, punctiform, scattered, blackish. Mycelium internal. Stromata ODFNLQJ RU VXEVWRPDWDO WR LQWUDHSLGHUPDO ± P GLDP EURZQ VZROOHQ K\SKDO FHOOV ± P GLDP ZDOOV VOLJKWO\ thickened. Conidiophores in small, divergent fascicles, mostly 3–12, arising from substomatal hyphae or stromata, through stomata or erumpent, erect, straight, subcylindrical to distinctly geniculate or geniculate-sinuous in the upper fertile portion, XQEUDQFKHG±î± ± P ± ± ± VHSWDWH Fig. 53b. Pseudocercospora varia var. viburni-sargentii (KUS 14011, medium to dark brown throughout or somewhat paler holotype). A. Conidiophore fascicles. B. Conidia. %DU P WRZDUGVWKHWLSZDOOVRPHZKDWWKLFNHQHGWRPVPRRWK FRQLGLRJHQRXV FHOOV LQWHJUDWHG WHUPLQDO ± P ORQJ proliferation sympodial, occasionally with enteroblastic, var. viburni-sargentii U. Braun & H.D. Shin, var. nov. monopodial proliferation leaving coarse annellations, MycoBank MB814558 conidiogenous loci inconspicuous to conspicuous by being (Fig. 53b) GHQWLFOHOLNH ± P GLDP RIWHQ VRPHZKDW UHIUDFWLYH RU even appearing darkened, in front view sometimes visible as Literature: Guo & Hsieh (1995: 51, as Pseudocercospora minute circle with dark rim (caused by the relatively thick and varia), Guo et al. (1998: 65, as Pseudocercospora varia), Kim dark walls of the conidiophores and conidiogenous cells), & Shin (1999), Shin & Kim (2001: 226, as Pseudocercospora but loci not thickened. Conidia solitary, obclavate, often with varia). almost rostrate apex, shorter conidia subcylindrical, 25–125 î ± P ±VHSWDWH RFFDVLRQDOO\ VXEK\DOLQH WR SDOH Illustrations *XR +VLHK ¿J *XR et al. (1998: olivaceous, but usually medium olivaceous or olivaceous- 6KLQ .LP ¿J EURZQZDOOWKLQWRVRPHZKDWWKLFNHQHG WRP VPRRWK DSH[REWXVHWRVXEDFXWHEDVHREFRQLFDOO\WUXQFDWH±P Diagnosis: Morphologically close to North American wide, hila unthickened, not darkened, but often somewhat collections of var. varia, but characterised by having refractive or ultimate rim slightly darker. conidiophores in larger, dense fascicles, about 10–30; FRQLGLD±î±P±VHSWDWH Holotype: USA: Wisconsin: Madison, Arboretum, on Viburnum sp. [?UD¿QHVTXLDQXP](asViburnum “lentago”), 5 Holotype: Korea: Pyeongchang, on Viburnum sargentii, 25 Sep. 1953, W. W. Diehl (BPI 442174). Aug. 1997, H. D. Shin (KUS-F 14011). Host range and distribution: On Viburnum (cassinoides, Host range and distribution: On Viburnum sargentii, Viburnum sp.), Adoxaceae, North America (USA, Wisconsin, Adoxaceae, Asia (China, Heilongjian, Liaoning; Korea). West Virginia). Notes: The Asian collections on Viburnum sargentii from Notes: An additional collection of this species has been China and Korea differ from North American specimens in examined [USA: West Virginia: Pocahontas County, on VOLUME 6 · NO. 2 427 428 ARTICLE (Braun which haveturnedouttobelong Basionym Pseudocercospora viburni-cylindrici in thisspeciesarereminiscentofformer thickened wallsandthestructureofconidiogenousloci loci andconidialhila. The shapeoftheconidiawithsomewhat mostly subhyalineconidiaandunthickenedconidiogenous Pseudocercospora varia comparable cercosporoidfungion much widerconidiawithsomewhatthickenedwalls.Other of the distinguishable fromall L. Sheldon Viburnum cassinoides Synonyms (Fig. 55) Pseudocercospora viburni-cylindrici Braun, (1948), Sin Zhao, Ray, 1941. .: 907(1979). Adoxaceae et al : Acta Mycol.Sin : Cercospora viburni-cylindrici Nova Hedwigia nom. illeg. 3815(BPI439059)]. This species iseasily . 2013). Cercospora viburnicola byitsconsistentlysolitary, pigmented, , 6 Aug. 1909, (Art. 53.1),non Pseudocercospora . , arequitedistinctbynarrower, 12 55 : 198(1993). : 219(1992). (FL. Tai) Y.L. Guo&W.X. F.L. Tai, Viburnum C. viburnicola F.L. Tai, Prathigada B. Brooks Pseudocercospora species onhosts Lloydia (F.L. Tai)U. spp.,e.g. Syll. Fung. , hb. species 11 W.W. W.W. : 54 Braun J. et al ±PGLDPEURZQ developed, subglobosetosomewhatirregular, substomatal, ±î±P±VHSWDWHSDOHROLYDFHRXVWKLQZDOOHG solitary, obclavate-cylindrical,straightoronlyslightly curved, loci inconspicuousunthickened andnotdarkened. FHOOVLQWHJUDWHGWHUPLQDODERXW±PORQJFRQLGLRJHQRXV to brown,wallsomewhatthickened, smooth;conidiogenous ±î±P±VHSWDWHPHGLXPROLYDFHRXVEURZQ straight, subcylindricaltogeniculate-sinuous,unbranched, fascicles, arisingfromstromata,throughstomata,erect, large, somewhatdivergenttodenseorevencoremioid dark, browntodarkbrown. halo. ZLWK ZKLWH JUH\LVK darker margin,brown,sometimeswithadiffuse olivaceous RU EURZQ JUH\LVK EURZQ FHQWUH ¿QDOO\ EURZQWRGDUNEURZQRUHYHQEODFNLVKPDUJLQLQGH¿QLWHRU LUUHJXODUVL]HYDULDEOH±PPGLDPVRPHWLPHVFRQÀXHQW Description ¿J ¿J *XR +VLHK ¿J *XR Illustrations Crous &Braun(2003:421). Literature . Caespituli : Guo&Hsieh(1995:52), : 7DL ¿J %UDXQ SODWH Leaf spots hypophyllous, punctiform,scattered,effuse, %DU P fascicles. (BPI 442174,holotype). Fig. 54. amphigenous, subcirculartoangular- Conidiophores Mycelium B. Pseudocercospora viburnicola Conidiophoretips. internal. in smalltomoderately etal. etal. A. Stromata IMA FUNGUS Conidiophore (1998: 65); (1998: 67, C. Conidia Conidia. well- Cercosporoid fungi 4 smooth, apex obtuse to subacute, base short obconically Pseudocercospora viburni-erosi U. Braun & H.D. ARTICLE WUXQFDWH±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG Shin, sp. nov. MycoBank MB814560 Holotype: China: Yunnan Province: Kunming, on Viburnum (Fig. 56) cylindricum, Adoxaceae, Jun. 1938, Xu Ren (HMAS 01929). Literature: Shin & Kim (2001: 228). Host range and distribution: Only known from the type collection. Illustration: 6KLQ .LP ¿J Notes: Braun (1992) reduced Pseudocercospora khasiana to Diagnosis: Differs from P. viburni-cylindrici in having large, synonymy with P. viburni-cylindrici, although the conidia in the dense, often coremoid conidiophore fascicles and above all Indian type material are distinctly wider and the conidiophore PXFK ORQJHUSOXULVHSWDWH F\OLQGULFDO¿OLIRUP FRQLGLD OHQJWK fascicles are smaller and not coremioid. Therefore, we prefer ±PZLWKWUXQFDWHEDVH to consider P. khasiana (= P. caprifoliacearum) a distinct species of its own, at least for the interim (see notes under the latter species). Korean material on Viburnum erosum, referred to as P. viburni-cylindrici by Shin & Kim (2001: 228), differs in having much longer, pluriseptate, cylindrical conidia with truncate base. The material concerned is described as a new species, Pseudocercospora viburni-erosi. Fig. 55. Pseudocercospora viburni-cylindrici (HMAS 01929, holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Fig. 56. Pseudocercospora viburni-erosi (KUS-F14126, holotype). %DU P A. Conidiophore fascicle. B. &RQLGLD%DU P VOLUME 6 · NO. 2 429 430 ARTICLE wide, hilaunthickened,notdarkened. VPRRWKDSH[REWXVHEDVHWUXQFDWHRUDOPRVWVR±P P ± î ± 3–12-septate, subhyalinetoverypaleolivaceous,thin-walled, FXUYHG PRGHUDWHO\ WR VR DOPRVW RU not darkened. terminal, conidiogenouslociinconspicuous,unthickened, the tip,thin-walled,smooth;conidiogenouscellsintegrated, ± P±VHSWDWHROLYDFHRXVEURZQWREURZQSDOHUWRZDUGV î ± XQEUDQFKHG SRUWLRQ XSSHU WKH LQ DOO DERYH straight, subcylindricaloronlyslightlygeniculate-sinuous, coremioid, arisingfromstromata,throughstomata,erect, EURZQ VXEJORERVH in well-developedfascicles,to30,denseverydense, RU JORERVH GLDP P Stromata ZLGH P ± K\DOLQH VHSWDWH EUDQFKHG K\SKDH LQWHUQDO Caespituli centre turninggreyishwhite,withraisedbrownborderline. FHQWUHVXUURXQGHGE\DEURZQWRGDUNEURZQERUGHU¿QDOO\ ¿UVW DW GLDP PP brown todarkbrown,laterwithgreyishgrey WR ODUJHU DQG FRQÀXHQW VRPHWLPHV circular oralmostsotoangular-irregular, 3–10mmdiam, Description substomatal, smalltowell-developed,15–30 : hypophyllous, brown,punctiform. Leaf spots Conidia DPSKLJHQRXVVFDWWHUHGWRFRQÀXHQW RLD\FOQUFO¿LRPVWUDLJKW F\OLQGULFDO¿OLIRUP VROLWDU\ Conidiophores Mycelium Braun et al separate species. better excludedfrom REYLRXV WKHVH RQ differences, theKoreanmaterialon %DVHG EDVH WUXQFDWH ZLWK P ± OHQJWK FRQLGLD F\OLQGULFDO¿OLIRUP ORQJHUSOXULVHSWDWH PXFK dense, oftencoremoidconidiophorefasciclesandaboveall Korean materialon 2001). The twospeciesaremorphologicallyveryclose,but assigned to Notes collection. Host rangeanddistribution taquetii Holotype Basionym (Fig. 57) Pseudocercospora viburnigena . Mycol. Progr 1994. non : Type materialofthenewspecieswaspreviously , 4Sep.1997, Pseudocercospora tinea : : Korea Cercospora tinea Pseudocercospora viburni-cylindrici &RQLGLD%DUV P fascicles. Mycoth. Ven Fig. 57. . : Kwachon,on 1 Viburnum erosum : 23(2002). H. D.Shin P. viburni-cylindrici Pseudocercospora viburnigena B. . 1252,HAL,lectotype). Stroma cells. : Onlyknownfromthetype Sacc., (KUS-F14126). Y.L. Guo&W.H. Hsieh Viburnum erosum Michelia differs inhavinglarge, U. Braun&Crous, Viburnum erosum C. Conidiophores. andtreatedas 1 A. : 268(1878), IMA FUNGUS (Shin&Kim Conidiophore (Sacc., var. is D. Cercosporoid fungi 4 Synonyms: Cercoseptoria tinea (Sacc.) Deighton, Mycol. Mycoth. Ven. 1252, e.g. B, BPI 441941, FH, HBG. Epitype ARTICLE Pap. 140: 167 (1976). (designated in Crous et al. 2013: 108): The Netherlands: Stigmina tinea (Sacc.) M.B. Ellis, More Dematiaceous Bilthoven, Sweelincklaan 87, on Viburnum davidii, 26 May Hyphomycetes: 118 (1976). 2008, M. K. Crous (CBS H-20393). Ex-epitype culture: CBS Cercostigmina tinea (Sacc.) U. Braun, Cryptog. Bot. 4: 108 125998. (1993). Host range and distribution: On Viburnum (davidii, plicatum Literature: Chupp (1954: 105), Katsuki (1965: 16), Kirk var. plicatum, tinus, Viburnum sp.), Adoxaceae, Europe (1999), Crous & Braun (2003: 421), Crous et al. (2013: 108). (Germany, Great Britain, Italy, The Netherlands, Portugal, Spain), North America (USA, Louisiana). Illustrations (OOLV ¿J % .LUN ¿J unnumbered), Crous et al ¿J Notes $OHFWRW\SL¿FDWLRQRICercospora tinea is necessary. Type material or any other samples are not listed for PAD Description: Leaf spots amphigenous, subcircular to in Gola (1930). Therefore, one of the syntypes distributed angular-irregular, sometimes vein-limited, 2–15 mm diam, as “Sacc., Mycoth. Ven. 1252” is designated as lectotype. pale to darker brown or centre later greyish brown to dingy 7KHSK\ORJHQHWLFDI¿QLW\RIWKLVVSHFLHVSUHYLRXVO\UHIHUUHG JUH\ PDUJLQ LQGH¿QLWH RU GDUNHU VRPHWLPHV ZLWK UHGGLVK to as Stigmina and Cercostigmina, was elucidated by Crous tinge, occasionally slighty raised. Caespituli amphigenous, et al. (2013). Based on molecular sequence analyses, it punctiform, olivaceous to dark or blackish brown, scattered. could be demonstrated that this species clusters in a big Mycelium internal, external mycelium lacking or occasionaly Pseudocercospora clade. with some external hyphae; hyphae branched, septate, Pseudocercospora tinea (see above) is quite distinct ± P ZLGH VXEK\DOLQH WKLQZDOOHG VPRRWK Stromata from Cercospora tinea E\ KDYLQJ VXSHU¿FDO K\SKDH ZLWK VXEVWRPDWDO RU LPPHUVHG ± P GLDP RFFDVLRQDOO\ solitary conidiophores and conidiogenous cells sympodially ODUJHU WR P GLDP K\SRSK\OORXV VWURPDWD VPDOOHU proliferating. Katsuki (1965) recorded Cercospora tinea from and substomatal, epiphyllous ones larger and immersed to Japan on Viburnum dilatatum and V. suspensum. Kobayashi somewhat erumpent), subglobose, brown to dark brown, (2007) added V. davidii, V. erosum, V. plicatum and V. FRPSRVHG RI EURZQ VZROOHQ K\SKDO FHOOV ± P GLDP sieboldii as host species for Japan. Japanese collections subglobose to somewhat irregular, walls thickened, 0.5–2 on Viburnum suspensum and V. plicatum var. tomentosum PConidiophores in somewhat divergent to usually dense, [V. tomentosum] have been examined by C. Nakashima sometimes very dense fascicles, larger fascicles sporodochial, (CNS567, CNS976) and turned out to belong to P. tinea. arising from stromata, emerging through stomata or erumpent, Therefore, all other Japanese records of P. viburnigena [C. occasionally with solitary conidiophores arising from tinea] remain unproven and doubtful, i.e, they seem rather to VXSHU¿FLDO K\SKDH VWUDLJKW WR VRPHZKDW FXUYHGVLQXRXV refer to P. tinea. not geniculate, unbranched, cylindrical, subcylindrical or A single North America collection has been examined and slightly attenuated towards the tip, sometimes ampulliform, FRQ¿UPHG USA: Louisiana: Lafayette, on Viburnum plicatum, DSH[DW¿UVWURXQGHGODWHUWUXQFDWHRUVXEWUXQFDWH± ± 21 Sep. 1886, A. B. Langlois, BPI 441939, 441940). î ± ± ± VHSWDWH SDOH EURZQ RU ROLYDFHRXV brown, thin-walled, smooth, occasionally slightly rough- Pseudocercospora viburni-nudi U. Braun, sp. nov. walled; conidiophores mostly reduced to conidiogenous MycoBank MB814562 FHOOV RFFDVLRQDOO\ LQWHJUDWHG WHUPLQDO ± P ORQJ ZLWK (Fig. 58) DVLQJOHWHUPLQDOFRQLGLRJHQRXVORFXV±PZLGHQHLWKHU thickened nor darkened, proliferation percurrent, with 1–3 Diagnosis: Distinguished from all other species of not very conspicuous annellations. Conidia solitary, acicular Pseudocercosora on Viburnum spp. and other hosts of the WR QDUURZO\ F\OLQGULFDO¿OLIRUP RFFDVLRQDOO\ VRPHZKDW Adoxaceae by having much shorter, 1–4(–5)-septate conidia, obclavate, straight to curved or somewhat sigmoid, (20–)30– ±î±P ± î ± P ± ±VHSWDWH VXEK\DOLQH WR YHU\ pale olivaceous or somewhat brownish, thin-walled, smooth, Description: Leaf spots amphigenous, subcircular to angular- guttulate when fresh, apex obtuse to subacute, base truncate LUUHJXODUVRPHWLPHVYHLQOLPLWHG±PPGLDPRUFRQÀXHQW WR VRPHZKDW REFRQLFDOO\ WUXQFDWH ± P ZLGH KLOD DQGODUJHUPHGLXPWRGDUNEURZQPDUJLQLQGH¿QLWHODWHUZLWK unthickened, not darkened. paler centre, greyish brown, grey to greyish white, surroundced In vitro: Colonies on MEA reaching 23 mm diam after by a narrow to moderately wide darker border, dark brown to 30 d at 24°C in the dark, circular, convex, margin smooth, almost blackish, sometimes with diffuse reddish halo. Caespituli distinctly darker than the rest of of colony, slightly folding amphigenous, punctiform, scattered, dark brown to blackish. occurs towards the margin of the colony, with moderate to Mycelium internal. Stromata substomatal to intraepidermal, 15– profuse aerial mycelium, surface olivaceous-grey, reverse PGLDPGDUNEURZQConidiophores in small to moderately greenish black. large fascicles, 3–18, arising from stromata, through stomata or erumpent, divergent, erect, straight, subcylindrical, only Lectotype (designated here, MycoBank, MBT202797): terminal part somewhat geniculate-sinuous or subdenticulate, Italy: Padova, on Viburnum tinus, Oct. 1877, Bizzozero XQEUDQFKHG±î±PEDVHRFFDVLRQDOO\VRPHZKDW [Sacc., Mycoth. Ven. 1252] (HAL, s.n.). Isolectotypes: Sacc., ZLGHUWRPDSH[VRPHWLPHVVOLJKWO\VZROOHQWRPZLGH VOLUME 6 · NO. 2 431 432 ARTICLE &RQLGLDREFODYDWHF\OLQGULFDOWRDFLFXODUWRPORQJED Key toCercosporaspecies on Aizoaceae Cercospora Aizoaceae FRQVWULFWHG VOLJKWO\ RIWHQ ± ± VHSWDWH P ± î ± obclavate-cylindrical, fusiform,ellipsoid,straighttocurved, always unthickenedandnotdarkened. VXEGHQWLFXODWHGHQWLFOHOLNHORFLDERXW±PGLDPEXWORF conidiogenous lociinconspicuoustoconspicuousbybeing LQWHJUDWHGWHUPLQDO±PORQJV\PSRGLDOO\SUROLIHUDWLQJ a singleconspicuousannellation;conidiogenouscells FRQLGLRSKRUHV P WR occasionally withasingleenteroblasticrejuvenationleaving WKLFNHQHG VRPHZKDW ZDOO WLS WKH 1–5-septate, mediumtodarkbrownthroughoutorpalertowards &RQLGLDDFLFXODUWRPORQJEDVHWUXQFDWH RQ ...... portulacastrum Conidia solitary, short Braun Tetragonia tetragonoides L et al nudum Holotype PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG short obconicallytruncate,sometimesrounded,1.5–2.5(–3) or olivaceous-brown,thin-walled,smooth,apexobtuse,base at thesepta,subhyalinetomostlypalemediumolivaceous collection. Host rangeanddistribution VHWUXQFDWHWRREFRQLFDOO\WUXQFDWH RQ . , Adoxaceae : USA : South Carolina ...... , 15Oct.1943, : Onlyknownfromthetype tips. Conidiophore. A. nudi Fig. 58. : MurrellsInlet,on ...... D. Conidiophore fascicle. Carter 855 (BPI 442176,holotype). Pseudocercospora viburni- &RQLGLD%DU P Trianthema C. trianthematis C. C. tetragoniae (BPI442176). Conidiophore IMA FUNGUS Viburnum B. Cercosporoid fungi 4 Cercospora species on Aizoaceae mm, pale brown, dingy olivaceous, greyish brown to grey ARTICLE or dingy greyish white, margin brown to reddish brown, Cercospora tetragoniae (Speg.) Siemaszko, Mater. occasionally somewhat zonate. Caespituli amphigenous, Mikol. Fitopatol. Rossii 1(3): 40 (1915). mostly epiphyllous, punctiform, dark brown to blackish. (Fig. 59) Mycelium internal. Stromata lacking or almost so to small, Basionym: Cercosporina tetragoniae Speg., Anales Mus. VXEVWRPDWDO ± ± P GLDP EURZQ Conidiophores Nac. Hist. Nat. Buenos Aires 20: 429 (1911). solitary or fasciculate, 2–10, divergent, arising from internal Synonyms: Cercospora tetragoniae (Speg.) Vassiljevsky, hyphae or stromata, through stomata or occasionally in Vassiljevky & Karakulin, Fungi imperfecti parasitici 1. erumpent, erect, straight, subcylindrical to geniculate- Hyphomycetes: 221 (1937). VLQXRXVXQEUDQFKHG±î±P±VHSWDWHSDOH Cercospora tetragoniae (Speg.) Chupp, in Viégas, Bol. Soc. yellowish, olivaceous-brown to brown, pale towards the tip, Bras. Agron. 8: 54 (1945). thin-walled, smooth; conidiogenous cells integrated, terminal, sometimes conidiophores reduced to conidiogenous cells, Literature: Chupp (1954: 27), Katsuki (1965: 7), Sutton & ±PORQJFRQLGLRJHQRXVORFLFRQVSLFXRXVWKLFNHQHG Pons (1980: 216), Braun & Mel’nik (1997: 97), Braun (2000b: DQG GDUNHQHG ± P GLDP Conidia solitary, acicular, 78), Crous & Braun (2003: 400). VWUDLJKW WR FXUYHG ± î ± P WR SOXULVHSWDWH hyaline, thin-walled, smooth, apex pointed, base truncate, Illustration 6XWWRQ 3RQV ¿J) ±PZLGHKLODWKLFNHQHGDQGGDUNHQHG Description: Leaf spots amphigenous, circular or subcircular, ±PPGLDPVFDWWHUHGWRFRQÀXHQWDQGODUJHUXSWR Fig. 59. Cercospora tetragoniae (LPS 16153, holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 Fig. 60. Cercospora trianthematis (K(M) IMI 83193, lectotype). P A. Conidiophores fascicle. B. &RQLGLD%DU P VOLUME 6 · NO. 2 433 434 ARTICLE DUHEDVHGRQPLVLGHQWL¿FDWLRQV oftensomewhat raisedmargin;caespitulimainlyepiphyllous,conspicuous,punctiformtoalmostpustulate, /HDIVSRWVVPDOO±PPGLDPGLQJ\JUHHQLVKJUH\EUR [ Host rangeanddistribution 241730 (slide). (Fig. 60) Records fromBrazilon Notes (USA, Maryland,Indiana, Texas). America (Argentina,ElSalvador, Brazil),North America Israel, Japan),Caucasus(Georgia),CentralandSouth Sierra Leone, Tanzania, Uganda,Zimbabwe), Asia (Brunei, efsosabsentoralmost so,sometimeswithslightdiscolorationsontheupperleafsurface;myceliuminternal; Leafspots 1 Key tocercosporoidspecieson Altingiaceae Altingiaceae Synonym trianthematis Cercospora 18 Nov. 1909, Holotype Casepituli irregular, scattered,0.5–6mmdiam,darkbrowntoblackish. Description Bhartiya Illustrations Literature expansa 25 Appl isotype Aizoaceae Gorakhpur, on : This speciesbelongstothe : 45“1997”(1998)[ Leaf spotslarger, usually1–10mmdiam;caespitulilessconspicuous,neverpustulate;conidiophorespaler, With distinctleafspots;myceliuminternalandoftenalsoexternal;conidiophoresfasciculate . : et al ]), : : Crous&Braun(2003:409),Kamal(2010:14,94). 17 Cercosporaaizoacearum Argentina : GPU8073]. : Aizoaceae amphigenous, mainly hypophyllous, punctiform, amphigenous, mainlyhypophyllous,punctiform, &LGUD OW, J± ¿JV ,, SODWH &KLGGDUZDU on subhyaline orpaletomedium olivaceousorolivaceous-brown,hilaunthickened,notdarkened FRQLGLRJHQRXVORFLLQFRQVSLFXRXVQHLWKHUWKLFNHQHGQRWGDUNHQH VXEK\DOLQHWRSDOHROLYDFHRXVRUROLYDFHRXVEURZQWKLQZDOOHG somewhat darkened-refractive;on FRQLGLDWRPORQJVXEK\DOLQHRUYHU\SDOHROLYDFHRXVKLO DURXQGWKHRXWHUULPVRPHZKDWGDUNHQHGUHIDUFWLYH±PGL WKLFNHQHG WRP ± ± PORQJ FRQLGLRJHQRXVORFLFRQ dark browntoblackish;conidiophorespalemedium brown,palertowardsthetip,wallsomewhat FRQLGLDPXFKQDUURZHU±P DOVRVROLWDU\DULVLQJIURPVXSHU¿FLDOK\SKDHLISUHVHQWDVHSW XQWKLFNHQHGQRUGDUNHQHG FRQLGLD±î±P RQ VXSHU¿FLDOK\SKDHODFNLQJ FRQLGLRSKRUHVIDVFLFXODWH±VHSWD ...... : 80(1962);as“ Leaf spots ¿J , Sep.1998, C. Spegazzini iudma formosana Liquidambar Trianthema portulacastrum : LaPlata,on , Africa (Cameroon,Kenya,Malawi, Spinacia oleracea amphigenous, circular to oval or amphigenous, circulartoovalor holotype : On (LPS16153). H. D.Bhartiya trianthemae Chidd., Tetragonia Tetragonia tetragonoides : Bhartiya C. apiis.lat India and Mycopathol. Mycol. : Isotype ”. /VW\UDFLÀXD Uttar Paradesh ...... (Mendes ( et al. tetragonoides (HCIO, s.n.); /LTXLGDPEDUVW\UDFLÀXD [ monogyna . complex. ()IMI : K(M) , Kavaka et al. Braun saadNrhAeia ...... , Asia andNorth America ], , : et al longer conidia,arangesimilarto shows cylindricaltosomewhatobclavateshorterandacicular conidia weredescribedtobecylindrical,buttheillustration XQGRXEWHGO\FRQVSHFL¿F7\SHPDWHULDOZDVQRWDYDLODEOH7KH C. aizoacearum complex byhavingobclavate-cylindricaltoacicularconidia. Note Aizoaceae brown. Host rangeanddistribution 32 India Lectotype ±PZLGHKLODWKLFNHQHGDQGGDUNHQHG subobtuse, basetruncatetoshortobconicallytruncate,about EURDGHU RFFDVLRQDOO\ P (1–)3–11-septate, ± hyaline,thin-walled,smooth,apexpointedto î ± ± FXUYHG WR conidia obclavatetosubcylindrical,longeronesacicular, straight ORFL FRQLGLRJHQRXV ORQJ WKLFNHQHGDQGGDUNHQHG±PGLDP P ± FHOOV FRQLGLRJHQRXV WR cells integrated,terminal,intercalaryorconidiophoresreduced to brown,wallsomewhatthickened,smooth;conidiogenous XQEUDQFKHG±î±P±VHSWDWHGDUNROLYDFHRXV straight, subcylindrical-conicaltogeniculate-sinuous,usually to divergent,arisingfromstromata,throughstomata,erect, large fascicles,2–20,occasionallysolitary, relativelydense diam, olivaceoustobrown. Trianthema portulacastrum ZQJUH\¿QDOO\XVXDOO\JUH\LVKZKLWHZLWKGDUN . (K(M)IMI83193). , North ...... America /LTXLGDPEDUVW\UDFLÀXD : A true : A : DWHQDUURZHU±PZLGH Maharashtra Mycelium DXQWKLFNHQHGWRVOLJKWO\WKLFNHQHG VKRUWHU±PORQJ WH±PZLGHFRQLGLRJHQRXVORFL , Asia (India,Maharashtra). G FRQLGLDVKRUWHUWRP VSLFXRXVVOLJKWO\WKLFNHQHGDWOHDVW DPLQIURQWYLHZYLVLEOHDVPLQXWHFLUFOH ( designated here Cercospora s.str. , describedfromIndiaonthesamehost,is internal. : Pune, Wanowri, Pune, MilitaryHospital,on : Isolectotypes Conidiophores Pseudocercospora tuberculans Stromata , 12Oct.1956, : On , North America , MycoBank,MBT202798): distinct fromthe Cercospora liquidambaris Trianthema portulacastrum C. trianthematis : BPI441988,HCIO. XVRDD P ± VXEVWRPDWDO Conidia in small to moderately in smalltomoderately P. liquidambaricola P. P. Chiddarwar solitary, shorter IMA FUNGUS C. apiis.lat. . 2 , Cercosporoid fungi 4 ARTICLE Fig. 61. Cercospora liquidambaris (CUP- A-002227(AL), lectotype). A. Conidiophore fascicles. B. Solitary conidiophores arising IURPVXSHU¿FLDOK\SKDHC. Conidiophores. D. &RQLGLD%DU P Cercospora (s. lat.) VXEVWRPDWDO RU LQWUDHSLGHUPDO ± P GLDP EURZQ Conidiophores in small to moderately large fascicles, arising Cercospora liquidambaris Cooke & Ellis ex G.F. Atk., from substomatal hyphae or stromata, though stomata or J. Elisha Mitchell Sci. Soc. 8: 48 (1892) erumpent, divergent to moderately dense, or solitary, arising (Fig. 61) IURPVXSHU¿FLDOK\SKDHODWHUDOHUHFWVWUDLJKWVXEF\OLQGULFDO or attenuated towards the apex to usually geniculate-sinuous, Literature: Chupp (1954: 259). XQEUDQFKHG ± ± ± î ± P ± ±VHSWDWH pale to medium brown, paler towards the tip, wall somewhat Description: Leaf spots amphigenous, subcircular to angular- WKLFNHQHG DW OHDVW EHORZ WR P VPRRWK RU DOPRVW irregular, small, 0.5–4 mm diam, dingy greyish green, brown, so, proliferation sympodial and occasionally percurrent WR¿QDOO\XVXDOO\JUH\LVKZKLWHRUZKLWHVRPHWLPHVVRPHZKDW (conidiophores with enteroblastic rejuvenation leaving delicate UDLVHGPDUJLQLQGH¿QLWHRUXVXDOO\GDUNHUQDUURZCaespituli annellations); conidiogenous cells integrated, terminal, 10–25 amphigenous, usually epiphyllous, very conspicuous, PORQJFRQLGLRJHQRXVORFLFRQVSLFXRXVLQIURQWYLHZYLVLEOH punctiform to almost pustulate, scattered, dark brown to DVPLQXWHFLUFOH±PGLDPDWOHDVWULPVOLJKWO\WKLFNHQHG blackish. Mycelium LQWHUQDO DQG H[WHUQDO VXSHU¿FLDO K\SKDH somewhat darkened or refractive. Conidia solitary, narrowly lacking or almost so to well-developed, branched, septate, REFODYDWHF\OLQGULFDO ¿OLIRUPVXEDFLFXODU VWUDLJKW WR FXUYHG VXEK\DOLQH WR SDOH ROLYDFHRXV RU EURZQLVK ± P ZLGH occasionally germinating with short to moderately long lateral Stromata almost lacking or small to usually well-developed, RXWJURZWKV ± î ± ± ± P ±VHSWDWH VOLUME 6 · NO. 2 435 436 ARTICLE Pseudocercospora liquidambaris Cercoseptoria liquidambaricola Synonyms Basionym Pseudocercospora liquidambaricola Pseudocercospora specieson Altingiaceae Pseudocercospora (Fig. 62) USA Lectotype so toslightlythickened,somewhatdarkenedorrefractive. VRPHWLPHVWUXQFDWH±PZLGHKLODXQWKLFNHQHGRUDOPRVW acute tosubobtuse,baseusuallyshortobconicallytruncate, subhyaline topaleolivaceous,thin-walled,smooth,apex Pseudocercospora neoliquidambaris lat. Therefore, thisspeciesistentativelyretainedin phylogenetically quiteheterogeneous(S.I.R.Videira, unpubl.). as inthe darkened. The conidiaare cercospora-like,butpigmented The conidiogenouslociandconidialhilaarethickened RI DI¿QLW\ JHQHULF 7KH Atkinson’s herbariumatCUP isdesignatedaslectotype. collected in Alabama in1891. A correspondingsamplefrom FODUL¿HGE\OHFWRW\SL¿FDWLRQ$WNLQVRQ FLWHGDVSHFLPHQ been examined. The identity ofthetrue VW\UDFLÀXD species isalsoknownfromNorth America on correctly assignedto these hostsin Asia belongsin which is,however, incorrectsincethecercosporoidfunguson to synonymywith material causingaleafspotdisease reduced liquidambaris Notes Montana, Massachusetts,NorthCarolina, Texas). Delaware, Florida,Indiana,Louisiana,Maryland,Mississippi, Altingiaceae Host rangeanddistribution 1891, until phylogeneticdatawillbeavailable. Braun, (1990) [ & Goh, PPE, hb.Sawada)]. Liquidambar formosana Cooke &EllisexG.F. Atk., 1892[ 39.1), and Agric. Res.Inst.Taiwan Trimestriel Soc.Mycol.France Trimestriel Soc.Mycol.France Kobay., 14 Oct.1928, VXSHUÀ : : The taxonomyandnomenclatureof Alabama G. F. Atkinson C. liquidambaris ( : . SeveralcollectionsfromMexicoandtheUSA have : Cercosporaliquidambaricola Passalora ] (Art.52.1). Cercosporaliquidambaris type Schlechtendalia Cercospora andsimilarfungi fromTaiwan Mycoscience , North America (Mexico;USA, Alabama, designated here have been totally confused. Chupp (1954) havebeentotallyconfused.Chupp(1954) : Auburn, on : Auburn, nom. illeg : Taiwan K. Sawada C. liquidambaris [CUP-A-002227(AL)]. compex, which recently proved to be compex,whichrecentlyprovedtobe P. liquidambaricola : Taipei,on C. liquidambaris . (Art.53.1),non 43 Sawada,basedon Taiwanese , 14Oct.1928, 85 Liquidambar : 224(2002), (NTU-PPE, hb.Sawada)]. : On 5 Pseudocercospora : 112 (1943), , MycoBank,MBT202799): : 44(2000). (J.M. Yen)J.M. Yen, Goh &W.H. Hsieh,inHsieh Cooke &EllisexG.F. Atk., 97 94 of Liquibambarformosana Liquidambar Liquidambar formosana type : 92(1981). : 52(1978). (seebelow). The later C. Nakash.& Tak. Sawada, : VW\UDFLÀXD C. liquidambaris Taiwan K. Sawada nom. illeg. remains unclear. C. liquidambaris J.M. Yen, nom. inval (J.M. Yen)U. Cercospora s. Cercospora s. Liquibambar Liquibambar and is now andisnow Cercospora Cercospora : Taipei,on VW\UDFLÀXD Rep. Gov. , 14Oct. (NTU- [ . (Art. : 150 nom. Bull. Bull. Braun is is , , , et al ¿J Fig. 62. IURPVXSHU¿FLDOK\SKDH (mainly hypophyllous when developed), branched, 1.5–4 (mainly hypophyllous whendeveloped),branched, 1.5–4 H[WHUQDO VXSHU¿FLDOK\SKDHODFNLQJRUDOPRVWVRWRGHYHORSHG delicately punctiformtosubeffuse, brown. halo, occasionallysomewhatzonate. blackish, narrow, sometimesraised, withdiffuse yellowish greyish browntowhite, withdarkborder, brownto RIWHQ vein-limited, paleolivaceous,browntodarkorlater VSRWV DQJXODU ODUJHU DQG FRQÀXHQW RU GLDP PP ± Description ¿J .RED\DVKL ¿J *XR +VLHK ¿J *XR Illustrations Crous &Braun(2003:253). Guo &Hsieh(1995:123–124), Literature . Pseudocercospora liquidambaricola : Chupp(1954:259p.p.),Hsieh&Goh(1990:150), : P ZLGH VXEK\DOLQH WR SDOH ROLYDFHRXVEURZQ WKLQZDOOHG Alabama in 1891, which can be used as lectotype. There is ARTICLE smooth. Stromata VPDOOWRPRGHUDWHO\ODUJH±PGLDP a single specimen in Atkinson’s herbarium (now CUP) which rarely larger, substomatal to immersed, subglobose, brown. undoubtedly refers to this collection. Conidiophores in small to moderately large fascicles, loose to dense, arising from stromata, through stomata or erumpent, Pseudocercospora tuberculans (Ellis & Everh.) U. DOVRVROLWDU\DULVLQJIURPVXSHU¿FLDOK\SKDHLISUHVHQWODWHUDO Braun, Schlechtendalia 2: 27 (1999). erect, straight, subcylindrical-conical to geniculate-sinuous, (Fig. 63) XQEUDQFKHG ± î ± P ±VHSWDWH VXEK\DOLQH SDOH Basionym: Cercospora tuberculans Ellis & Everh., J. Mycol. olivaceous to olivaceous-brown, thin-walled, smooth to 4: 115 (1888). somewhat rough; conidiophores reduced to conidiogenous FHOOV RU LQWHJUDWHG WHUPLQDO ± P ORQJ FRQLGLRJHQRXV Literature: Saccardo (1892: 652), Chupp (1954: 259), Crous loci inconspicuous or visible as a truncate tip, but always & Braun (2003: 412). unthickened and not darkened. Conidia solitary, cylindrical to obclavate-cylindrical, straight to curved, occasionally sigmoid, Exsiccatae: Ellis & Everh., Fungi Columb. 168. Ellis & Everh., ±î±P±VHSWDWHVXEK\DOLQHWRSDOHROLYDFHRXV N. Amer. Fungi 2292. or olivaceous-brown, thin-walled, smooth, apex obtuse to subacute, base short obconically truncate, sometimes Description: Leaf spots absent or almost so, sometimes with WUXQFDWH±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG slight discolorations on the upper leaf surface. Caespituli hypophyllous, punctiform, on small, brown, tubercle-like Neotype (designated here, MycoBank, MBT202800): swellings, 0.5–1 mm diam, sometimes effuse between Taiwan: Taipei, on Liquidambar formosana, 14 Oct. 1928, K. such swellings. Mycelium internal. Stromata almost Sawada (NTU-PPE, hb. Sawada). DEVHQWWRZHOOGHYHORSHG±PGLDPGDUNEURZQWR blackish, substomatal to immersed, composed of swollen Host range and distribution: On Liquidambar (formosana, K\SKDOFHOOV±PGLDPConidiophores in small to large VW\UDFLÀXD), Altingiaceae, Asia (China, Japan, Taiwan), North fascicles, loose to usually dense, subcylindrical to conical, America (Mexico; USA, Alabama, Florida, Louisiana, Texas). Notes: The holotype material of Cercospora liquidambaricola (Taiwan: Taichung, on Liquidambar formosana, 29 Oct. 1971, J.-M. Yen 71277 bis) could not be traced in PC or UC. Therefore, we designate a neotype here. Collections of Pseudocercospora on Liquidambar formosana in Asia and /VW\UDFLÀXDin North America are morphologically very VLPLODU,IWKH\DUHWUXO\FRQVSHFL¿FRULIWZRPRUSKRORJLFDOO\ very close, but geographically and genetically distinct VSHFLHV DUH LQYROYHG FDQ RQO\ EH FODUL¿HG RQ WKH EDVH of molecular sequence analyses. Loropetalum chinense (Hamamelidaceae) was reported as host of this species from China (Guo & Hsieh 1995; Guo et al. 1998). The identity of Pseudocercospora on this host is unclear and needs to EH FRQ¿UPHG &KXSS ZKR SURSRVHG WKH RULJLQDO synonymy, found material from the USA and Taiwan to represent the same fungus. However, Hsieh & Goh (1990) did not examine American material. Samples from Taiwan (Taichung, 8 Aug. 1945, K. Sawda, BPI 437752, 437753) DUHSUREDEO\WRSRW\SHV*XR +VLHK FRQ¿UPHGWKH synonymy of C. liquidambaricola. They mentioned that they had examined Yen’s type material, but this material was not cited under “material examined”. The material concerned is not preserved at PC. The complicated nomenclature and taxonomy of this species has been discussed by Braun (2000a: 44) who emphasized that identity and application of the name C. liquidambaris FDQ RQO\ EH FODUL¿HG E\ OHFWRW\SL¿FDWLRQ&KXSS LQWHUSUHWHGWKLVQDPHLQWKH sense of P. liquidambaricola, which is clearly distinct from P. tuberculans by much narrower conidia, but his interpretation was based on a specimen collected by Geo. V. Nash in 1895 (USA, Florida, Lake City, Plants of Florida No. 2231, 11–19 Fig. 63. Pseudocercospora tuberculans (NY 2425377, lectotype). Jul. 1895, BPI 437755, 437761, 437812), which is non- A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 type material. Atkinson (1892) cited a specimen collected in P VOLUME 6 · NO. 2 437 438 ARTICLE (see Conidiaconsistentlyacicular, basetruncate,variousmorphologically barelydistinguishablespecies 7 (6) Conidianarrowlycylindricaltosubacicular, apexobtuse,not distinctlypointed;on 6 (5) /HDIVSRWVGHYHORSHGGLVWLQFW FRQLGLDFRQVLVWHQWO\DFLFXODU VHH³'RXEWIXOH[FOXGHGDQGLQVXI¿HQWO\NQRZQVSHFLHVRI /HDIVSRWVODFNLQJRUDOPRVWVR FRQLGLDDFLFXODUWRREFO VHH³'RXEWIXOH[FOXGHGDQGLQVXI¿HQWO\NQRZQVSHFLHVRI &RQLGLRSKRUHVK\DOLQHRURQO\SDOHROLYDFHRXVDWWKHEDVH 2 Conidiogenouslocicercospora-like, i.e.distinctlythickenedanddarkened,infrontviewvisibleassmalldarkcircle 3 (2) 6WURPDWDGHYHORSHG±PGLDP FRQLGLRSKRUHVORQJHU± 6WURPDWDODFNLQJRUDOPRVWVR FRQLGLRSKRUHVVKRUW± Conidiacylindricalorobclavate-cylindrical ...... 1 Key toCercosporaspecieson Amaranthaceae Cercospora Lectotype darkened. REFRQLFDOO\WUXQFDWH±PZLGHKLODXQWKLFNHQHGQRW subtruncate toshortobconicallytruncate,rarelylong to brownish,thin-walled,smooth,apexobtuse,base FXUYHG±î±P±VHSWDWHSDOHROLYDFHRXV to somewhatobclavate-cylindrical,straightslightly FRQLGLRJHQRXV unthickened andnotdarkened. ORQJ P loci inconspicuousorvisibleastruncatetips,always ± FHOOV FRQLGLRJHQRXV cells integrated,terminalorconidiophoresreducedto olivaceous-brown, thin-walled,smooth;conidiogenous î±PUDUHO\ORQJHU±VHSWDWHSDOHROLYDFHRXVWR straight toslightlygeniculate-sinuous,unbranched,10–35 168). CUP – A(S.M.T.[03]); HBG,Ellis&Everh.,FungiColumb. Topotypes 2292, e.g.BPI442930,MICH15380,NY 2425378–2425383. BPI 442929,442032,1102025; Ellis&Everh.,N. Amer. Fungi VW\UDFLÀXD Mississippi tlatlne oii itntyaiua ihtuct aeadaueo uaueae;o te ot ...... At leastlongerconidiadistinctly acicularwithtruncatebaseandacuteorsubacuteapex; onotherhosts Conidiophores paleolivaceous-brownthroughoutortipspaler, hyalineorsubhyaline;conidianarrowlycylindrical Conidiogenous locinotcercospora-like,i.e.unthickenedtosomewhatthickened,butdarkened,atmost Conidia consistentlyacicularoratleastlongerconidiaandonlyshorteronessomewhatobclavate- ( , 27June1888, : June1890–1893(BPI442031,442033,442034; : OktibbehaCounty, Starkville,on designated here on WRVXEDFLFXODU± î±P RQ on yln rsbyln ...... hyaline orsubhyaline proliferation; conidiophoreshyaline,onlypaleolivaceousatthebaseorolivaceous-brown,tipsoften somewhat refractive,formedoncharacteristic,bulging,convextipsandshoulderscausedbysympodial with minutecentralpore;conidiophoresdistinctlybrown;on ±PSOXULVHSWDWH 0–5-septate; on yidia ...... cylindrical ...... Achyranthes Achyranthes Tabular keytoCercospora specieson Acanthaceae accordingtohost genera S. M.Tracy , MycoBank,MBT202801): p...... spp...... spp. Celosia Conidia s...... sp. (NY 2425377)...... solitary, cylindrical Liquidambar Gomphrena Isotypes USA Braun : : p...... spp. DYDWHF\OLQGULFDOEURDG± î ± ± ± P et al î±P ± VHSWDWH FRQLGLDVKRUW±î±P FRQLGLDREFODYDWHF\OLQGULFDO±î±P Nov., tuberculans undoubtedly doesnotpertainto of thiswood-inhabitingfungusisnotquiteclear, butit wood of Guo unproven andnotincludedinGuo&Hsieh(1995) formosana Notes Missouri, Mississippi). Altingiaceae Host rangeanddistribution Chenopodiaceae in thelatterfamily( Amaranthaceae (The family Amaranthaceae s.str. DQGRUQDUURZHU±P . etal. Cercospora Cercospora Celosia : RecordsofthisspeciesfromChinaon C. Keissler P±VHSWDWH DQGRUFRQLGLDORQJHU Liquidambar formosana (Keissler &Lohwag1937, Tai 1979)aredoubtful, (1998). They refertoChinesematerialondead inPetr., Crypt.Exs.3391[Hunan,Changcha, , North America (USA,Florida,Louisiana, Chenopodiaceae sp...... sp. ”) ”) asseparatefamily, atleasttentatively.) and currentlyproposedtobeincluded & s. lat H. Handel-Mazzetti Gomphrena Cercosporella pseudachyranthis ...... ), butweprefertomaintainthe ...... 6 ...... : On ...... 5 ...... is phylogeneticallycloseto C. tuberculans ) distributed as ...... 4 ...... 2 ...... p. South Africa spp., LXGPDVW\UDFLÀXD /LTXLGDPEDU (W)]. The identity C. gorakhanathii C. pretoriensis C. pretoriensis C. achyranthis C. celosiigena . IMA FUNGUS Liquidambar Cercospora ..... 3 ...... 7 , Cercosporoid fungi 4 Conidia acicular to obclavate-cylindrical, at least shorter conidia obclavate-cylindrical, with obconically truncate base ARTICLE ...... 8 6WURPDWDZHOOGHYHORSHGODUJH±PGLDP FRQLGLRSKRUHV±PORQJ FRQLGLDOEDVH±PZLGH on Alternanthera spp...... C. alternantherae 6WURPDWDODFNLQJRU±PGLDP FRQLGLRSKRUHVWRPORQJ FRQLGLDOEDVH±PZLGH onAmaranthus spp...... C. brachiata Tabular key to Cercospora species on Acanthaceae according to host genera Achyranthes 1 Conidiophores colourless or only olivaceous at the base; conidiogenous loci conspicuous, but not cercospora-like (not darkened, only refractive, characteristically bulging, cercosporella-like) ...... Cercosporella pseudachyranthis VHH³'RXEWIXOH[FOXGHGDQGLQVXI¿FLHQWO\NQRZQVSHFLHVRICercospora”) Conidiophores brown, distinctly pigmented throughout; conidiogenous loci distinctly thickened and darkened, Cercospora type ...... 2 /HDIVSRWVODFNLQJRUDOPRVWVR FRQLGLRSKRUHVWRPORQJ FRQLGLDDFLFXODUWRREFODYDWHF\OLQGULFDO ± î ± ± ± P...... C. achyranthis /HDIVSRWVGLVWLQFWZHOOGHYHORSHG FRQLGLRSKRUHVWRPORQJ FRQLGLDXVXDOO\FRQVLVWHQWO\DFLFXODU PXFKQDUURZHU±î±P...... C. achyranthina Aerva A single species ...... C. aervae-lanatae Alternanthera A single species ...... C. alternantherae Amaranthus A single species ...... C. brachiata Celosia &RQLGLRSKRUHVVKRUW±î±P ± VHSWDWH FRQLGLDVKRUW±î±PF\OLQGULFDOWR REFODYDWHF\OLQGULFDO±VHSWDWHKLOD±PZLGH ...... C. celosiigena &RQLGLRSKRUHVPXFKORQJHU±P±VHSWDWH FRQLGLDPXFKORQJHU±P ± ±VHSWDWH KLOD±PZLGH...... 2 2 (1) Stromata lacking or almost so; conidia acicular ...... C. celosiae 6WURPDWDGHYHORSHG±PGLDP FRQLGLDF\OLQGULFDORUREFODYDWHF\OLQGULFDO ...... C. gorakhanathii Cyathula A single species ...... C. apii s. lat. (C. cf. maloti sensu Groenewald et al. 2013) Digera A single species ...... C. achyranthina Gomphrena 6WURPDWDODFNLQJRUVPDOO±PGLDP FRQLGLDDFLFXODU±PORQJDSH[SRLQWHG PDLQO\RQ Gomphrena globosa, Northeast Africa, Asia, North America, West Indies ...... C. gomphrenae 6WURPDWD±PGLDP FRQLGLRJHQRXVORFLQRWFHUFRVSRUDOLNH QHLWKHUGLVWLQFWO\WKLFNHQHGQRUGDUNHQHG conidia narrowly cylindrical to subacicular, apex obtuse; on Gomphrena spp., South Africa ...... C. pretoriensis VHH³'RXEWIXOH[FOXGHGDQGLQVXI¿HQWO\NQRZQVSHFLHVRICercospora”) Pupalia A single species ...... C. achyranthina VOLUME 6 · NO. 2 439 440 ARTICLE LUUHJXODU±PPGLDPRFFDVLRQDOO\FRQÀXHQWDQGODUJHU Description ¿J Illustrations (2010: 12),Braun&Urtiaga(2013:177). 328), Crous&Braun(2003:42),Guo El NurElamin(1974),Ellis(1976:244),Crous&Braun(2001: Literature Cercospora achyranthina Cercospora specieson Amaranthaceae (Fig. 64) P A. Fig. 64. Conidiophore fascicle. Mycologia : Chupp(1954:29),Vasudeva (1963:31),Ibrahim& Cercospora achyranthina : (OOLV ¿J% *XR Leaf spots 40 : 352(1948). B. amphigenous, circulartoangular- Conidiophore tip. Thirum. &Chupp, (BPI 4323844,holotype). et al . (2005:27),Kamal C. Conidia. Bar=10 et al . (2005: Braun et al Achyranthes Host rangeanddistribution Topotype aspera Holotype WUXQFDWH±PZLGHKLODWKLFNHQHGDQGGDUNHQHG subobtuse, basetruncate,occasionallyslightlyobconically XVXDOO\ 3–10-septate, hyaline,thin-walled,smooth,apexacuteor P ± î ± ± ± REFODYDWHF\OLQGULFDO ORQJ Conidia GLDP P P ± GDUNHQHG ± DQG WKLFNHQHG LQWHUFDODU\ ORFL FRQLGLRJHQRXV DQG WHUPLQDO LQWHJUDWHG FHOOV ROLYDFHRXVEURZQ or brownish,wallsomewhatthickened,smooth;conidiogenous SDOH ± ±VHSWDWH P ± î ± geniculate-sinuous, aboveallintheupperhalf,unbranched, stromata, throughstomataorerumpent,erect,straighttocurved, in fascicles,2–10,divergent,arisingfromswollenhyphalcellsor PGLDPVXEVWRPDWDOWRLPPHUVHGEURZQ dark. is maintainedanddepositedasCUP 37201. This specimen,whichcanbeconsideredtopotypematerial, mentioned acollectionfrom2Sep.1945astypematerial. collection from1944astype.Chupp(1954)erroneously &Chupp(1948) citedasingle Thirumalachar complex. Notes (Venezuela). Bengal; Myanmar, Pakistan,Philippines),South America Pradesh, MadhyaMaharashtra,NewDelhi,West Zimbabwe), Asia (China;India,Bihar, Karnataka, Andra [ RIWHQ KDOR GDUNHU GLIIXVH purplish. ZLWK VRPHWLPHV GDUN YHU\ ¿QDOO\ grey orgreyishwhite,marginreddishbrowntopurple-violet, WRPPGLDPEURZQWRJUH\LVKEURZQ¿QDOO\VRPHWLPHVGXOO Description Exs. 546. Exsiccatae ¿J Illustrations Guo (1965: 8),Shin&Kim(2001:24),BraunCrous(2003:42), (1937: 222),Chupp(1954:30),Vasudeva (1963: 31),Katsuki Literature (Fig. 65) Cercospora achyranthis so. septate, somewhatpigmented. dark greyorbrownish. WR JUH\LVK GDUN ÀRFFRVHYHOXWLQXRXV HIIXVH K\SRSK\OORXV ¿QDOO\ EURZQLVK WR VRPHWLPHVYHLQOLPLWHGJUH\LVKPDUJLQLQGH¿QLWH JUHHQLVK GLVFRORUDWLRQV GLIIXVH RQO\ atropurpurea . Conidiophores 7 et al : 171(1909). Mycelium : This speciesispartofthe , 20 Aug. 1944, solitary, acicular, shorterconidiaoccasionallyslightly : Caespituli . (2005:28–29),Kamal(2010:12). : 2Sep.1945(CUP 37201). : Saccardo(1913:1429),Vassiljevsky &Karakulin &LI0\FRÀ'RPLQJ([V6\G)XQJL([RW India : 6KLQ .LP ¿J *XR Leaf spots ]. sp.), Amaranthaceae internal. : in small,loosefascicles, 2–12,arisingfrom Karnataka DPSKLJHQRXVVFDWWHUHG¿QHO\SXQFWLIRUP Digera muricata,Pupalialappacea Mycelium M. J.Thirumalachar Stromata lacking oralmostso,indistinct : Bangalore,on Syd. &P. Syd., : On , Africa (Sudan, Tanzania,(Sudan, , Africa Stromata almost lackingorsmall,10– internal; hyphaebranched, Cercospora apiis.lat. Achyranthes lacking oralmost (BPI 4323844). Conidiophores Conidiophores Ann. Mycol IMA FUNGUS Achyranthes et al Caespituli ( . (2005: aspera . , Cercosporoid fungi 4 Lectotype (designated here, MycoBank, MBT202802): ARTICLE Japan: Saitama Prefecture: Ome, Musashi, on Achyranthes bidentata, 22 Sep. 1905, I. Miyake (S-F23053). Isolectotypes: LEP. Host range and distribution.: On Achyranthes (aspera [indica], bidentata [japonica]), Amaranthaceae, Asia (China; India, Andhra Pradesh, Maharashtra, Rajasthan; Japan, Korea, Pakistan, Taiwan), West Indies (Dominican Republ., Puerto Rico). Note: A true Cercospora s. str. distinct from C. apii s. lat., including C. achyranthina, by having acicular to obclavate- cylindrical conidia with obconically truncate base. Obconically WUXQFDWH FRQLGLDO EDVHV DUH QRW FRQ¿QHG WR VKRUWHU \RXQJ conidia, they are also evident in longer, fully developed conidia. In addition, different from C. achyranthina by lacking or indistinct leaf spots, effuse caepituli and somewhat wider conidia. Cercospora aervae-lanatae Raghu Ram & Mallaiah, Mycol. Res. 100: 296 (1996); as “aerva-lanatae”. (Fig. 66) Synonym: Cercospora aervae R.K. Srivast. et al., Indian Phytopathol. 54: 102 (2001); as “aeruae” [holotype: India: Uttar Pradesh: Gorakhpur, on Aerva sanguinolenta [scandens], Nov. 1990, R. K. Srivastava 123 (GPU 1398). Isotype: HCIO 30878.]. Literature: Crous & Braun (2003: 47), Kamal (2003: 13). Illustrations 5DJKX 5DP 0DOODLDK ¿J Srivastava et al ¿J Description: Leaf spots amphigenous, necrotic, scattered, subcircular to angular-irregular, 2–9(–10) mm diam, greyish, sometimes with reddish or purplish margin. Caespituli epiphyllous. Mycelium immersed; hyphae branched, septate, subhyaline. Stromata ZHOOGHYHORSHG JRERVH ± P diam, brown to black. Conidiophores in loose fascicles, to 15, arising from stromata, through stomata, erect, subcylindrical WR±WLPHVJHQLFXODWHXQEUDQFKHGDERXW±î± P ±VHSWDWHSDOH ROLYDFHRXV WR EURZQ SDOHU WRZDUGV Fig. 65. Cercospora achyranthis (LEP, isolectotype). A. Conidiophore the tip, thin-walled, smooth; conidiogenous cells integrated, fascicle. B. Conidiophore tip. C. &RQLGLD%DU P terminal and intercalary, conidiogenous loci thickened and GDUNHQHG±PGLDPConidia solitary, acicular, straight WRFXUYHG±î±P±VHSWDWHK\DOLQHWKLQ substomatal swollen hyphal cells, through stomata, erect, ZDOOHGVPRRWKDSH[SRLQWHGEDVHWUXQFDWH±PZLGH straight to usually distinctly geniculate or geniculate-sinuous, hila thickened and darkened. RIWHQVWURQJO\JHQLFXODWHXQEUDQFKHG±î±P pluriseptate, pale to medium dark olivaceous or olivaceous Holotype: India: Andhra Pradesh: Nagarjuna Nagar, throughout or paler towards the tip, wall thin to somewhat University Campus, on Aerva lanata, Nov. 1991, M. Raghu thickened, smooth; conidiogenous cells integrated, terminal Ram (K(M) IMI 351224). DQG LQWHUFDODU\ DERXW ± P ORQJ FRQLGLRJHQRXV ORFL FRQVSLFXRXV WKLFNHQHG DQG GDUNHQHG ± P GLDP Host range and distribution: On Aerva (lanata, sanguinolenta Conidia solitary, acicular to obclavate-cylindrical, straight to [scandens]), Amaranthaceae, Asia (India, Andhra Pradesh, FXUYHG ± î ± ± ± P ±VHSWDWH K\DOLQH Uttar Pradesh). or almost so, thin-walled, smooth, apex acute to subobtuse, EDVHWUXQFDWHWRVKRUWREFRQLFDOO\WUXQFDWH±PZLGH Note: This species is morphologically assignable to the hila thickened and darkened. Cercospora apii s. lat. complex. VOLUME 6 · NO. 2 441 442 ARTICLE straight, subcylindrical-conical to oblong, geniculate-sinuous, straight, subcylindrical-conical tooblong,geniculate-sinuous, arising fromstromata,through stomataorerumpent,erect, developed, mostlyratherlarge fascicles,divergenttodense, LPPHUVHGODUJH±PGLDPEURZQ to blackish. Caespituli 0.5–3 mmdiam,palegreenishtodingygrey, marginbrownish. Description Illustration Crous &Braun(2003:53),Kamal(2010:15). Literature Cercospora alternantherae (Fig. 67) A. Fig. 66. P Conidiophore fascicle. 6 : 36(1890). Cercospora aervae-lanatae : Saccardo(1892:637),Chupp(1954:30–31), &KXSS ¿J : amphigenous, punctiform, scattered, dark brown amphigenous, punctiform,scattered, darkbrown Leaf spots Mycelium B. amphigenous, circularorsubcircular, internal. Conidiophore tips. Ellis &Langl., (K(M) IMI351224,holotype). Stromata Conidiophores C. substomatal to substomatal to Conidia.Bar= J. Mycol in well- Braun . et al Jul. 1888, Louisiana Lectotype PZLGHKLODWKLFNHQHGDQGGDUNHQHG pointed orsubobtuse,basetruncatetoobconicallytruncate,1–2 î±P±VHSWDWHK\DOLQHWKLQZDOOHGVPRRWKDSH[ GLDP solitary, obclavate-cylindrical, acicular, straighttocurved,20– P ± ± ± GDUNHQHG DQG WKLFNHQHG FRQVSLFXRXV ORFL FRQLGLRJHQRXV ORQJ P ± LQWHUFDODU\ VXEK\DOLQH ±VHSWDWH smooth; conidiogenouscellsintegrated,terminal,occasionally P ± pale olivaceoustobrown,palertowardsthetip,thin-walled, î ± XQEUDQFKHG P A. Fig. 67. Host rangeand distribution BPI 432464. . Conidiophore fascicles. : St.Martinsville,on Cercospora alternantherae ( designated here A. B.Langlois1430 B. Conidiophore tips. , MycoBank,MBT202806): : On Alternanthera achyrantha (NY 830163). Alternanthera (NY 830163,lectotype). C. Conidia.Bar= IMA FUNGUS Isolectotype ( ¿FRLGHD Conidia Conidia USA , 18 : : , Cercosporoid fungi 4 halimifolia, crucis [portoricensis], pungens [achyrantha, ARTICLE repens]), Amaranthaceae, North America (USA, Louisiana, Texas), South America (Brazil, Venezuela), West Indies (Cuba, Jamaica, Puerto Rico, Virgin Islands). Notes. This is a true Cercospora s. str. close to C. apii s. lat., but GLVWLQFWE\KDYLQJODUJHVWURPDWD ±PGLDP ZLWKQXPHURXV densely arranged conidiophores, smaller conidiogenous loci (1–2 P ZLGH DQG REFODYDWHF\OLQGULFDO WR DFLFXODU FRQLGLD Indian records of this species from Uttar Pradesh and West Bengal on A. sessilis (Kamal 2010) are doubtful and in need of revision and FRQ¿UPDWLRQ7KH\PLJKWUDWKHUSHUWDLQWRWKHLOOHJLWLPDWHVSHFLHV Cercospora sessilis or to the invalid C. alternantherina, both described from India on A. sessilis. Cercospora apii s. lat. (C. cf. malloti sensu Groenewald et al. 2013: 157). Notes: Nguanhom et al. (2015) examined Cercospora species from northern Thailand by means of molecular methods. A plurivorous Cercospora species referred to as Cercospora cf. malloti in Groenewald et al. (2013) turned out to be the most common taxon found during the course of this study. The collections in the clade concerned belong to C. apii s. lat., i.e. they are characterised by having consistently acicular conidia. One of the specimens involved was collected on Cyathula prostrata. For further comments, see notes under Cercospora asystasiana. Cercospora brachiata Ellis & Everh., J. Mycol. 4: 5 (1888). (Fig. 68) Synonyms: Cercospora acnidae Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 43: 89 (1891) [lectotype (designated here, MycoBank, MBT202807): USA: Delaware: Wilmington, on Amaranthus cannabinus, 30 Sep. 1889, A. Commons 1011 (NY 2408316); isolectotypes: NY 2408317, 2408318]. Cercospora amaranthi Lobik, Bolezni Rast. 17: 193 (1928) Fig. 68. Cercospora brachiata (NY 2408319, lectotype). A. [holotype: Russia: Stavropol Krai: Pyatigorsk, station of Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 Yessentuki, garden, on $PDUDQWKXVUHWURÀH[XV, 25 Sep. P 1925, A. I. Lobik (not preserved)]. Literature: Saccardo (1892: 637), Vassiljevsky & Karakulin almost lacking or small to moderately large, substomatal (1937: 222), Chupp (1954: 30–31), Katsuki (1965: 8), Braun to immersed, subglobose to somewhat irregular, 10–50 & Mel’nik (1997: 43–44), Crous & Braun (2003: 43, 86), Guo PGLDPROLYDFHRXVEURZQWREURZQFHOOV±PGLDP et al. (2005: 29–30), Kamal (2010: 25). Conidiophores fasciculate, 2–12(–20), rarely solitary, arising from swollen hyphal cells or stromata, emerging Illustrations 7DL ¿J *XRet al. ¿J through stomata or erumpent, erect, straight, subcylindrical 11), Pirnia et al ¿J to usually distinctly geniculate or geniculate-sinuous, above all in the upper half, unbranched or tips occasionally Exsiccatae: Ellis & Everh., N. Amer. Fung. 2582. IXUFDWH ± î ± P XVXDOO\ ±VHSWDWH YHU\ long conidiophores sometimes with additional septa, pale Description: Leaf spots amphigenous, circular, subcircular to medium dark brown or olivaceous-brown throughout or WR VRPHZKDW DQJXODULUUHJXODU ± PP GLDP DW ¿UVW paler towards the tip, wall thin to slightly thickened, smooth; yellowish brown, later brown, reddish or dark brown, or conidiogenous cells integrated, terminal and intercalary, ¿QDOO\ZLWKDWDQJUH\WRJUH\LVKZKLWHFHQWUHVXUURXQGHG ± ± P ORQJ FRQLGLRJHQRXV ORFL FRQVSLFXRXV by a brown, reddish to purplish brown or almost blackish WKLFNHQHG DQG GDUNHQHG ± P GLDP Conidia margin. Caespituli amphigenous, mostly hypophyllous, solitary, acicular, shorter conidia may also be narrowly punctiform scattered, dark. Mycelium internal. Stromata REFODYDWHF\OLQGULFDOVWUDLJKWWRFXUYHG±î± ± VOLUME 6 · NO. 2 443 444 ARTICLE in 1890. material (formersyntypes)fromthetypelocality, butcollected GZU, NY, PHandnumerousotherherbaria)isauthentic (Fig. 69) Cercospora celosiae (Pirnia & Braun2004),Indonesia(Shivas included inCrous&Braun(2003)refertoGermany(Jage Type materialofthisspeciesisprobablynotmaintained. for typematerialof RI former DI¿QLW\ FORVH WKH UHÀHFWV ZKLFK VXEJHQXV DV WUHDWHG genus The hostsof XQGRXEWHGO\LQÀXHQFHGE\DVVXPHGKRVWUDQJHGLIIHUHQFHV Its introductionandrecognitioninChupp(1954)were Hsieh &Goh(1990: 15),Crous&Braun(2003: 113), Guo (1965: 9),Ibrahim&ElNurElamin (1974),Ellis(1976:244), Literature probably to [ Records of is morphologicallybarelydistinctfrom do withacomplexofcrypticspecies. if allcollectionspertaintoasinglespeciesorwehave different continentsareinvolved. Thus, itremainsunclear Many hostspeciesandawidedistributionrangecovering conidiophores aswelllengthandwidthofconidia. rather variable,aboveallwithregardtothelengthof lat. Notes Islands). Republic, Haiti,PuertoRico, Trinidad and Tobago, Virgin Texas, Wisconsin),West Indies(Barbados,Cuba,Dominican North America (USA,Delaware,Florida,Illinois,Nebraska, and South America (Brazil,Ecuador, Panama,Venezuela), Iran, Pakistan),Europe(Germany, Russia,Ukraine),Central Orissa, Maharashtra,UttarPradesh,West Bengal;Indonesia, Uganda), Asia (Brunei,China;India, Andhra Pradesh,Delhi, Amarathus polygamus USA Lectotype hila thickenedanddarkened. RUREFRQLFDOO\WUXQFDWHLQVKRUWHUFRQLGLD±PZLGH acute orsubacute,sometimessubobtuse,basetruncate DSH[ VPRRWK WKLQZDOOHG K\DOLQH ±VHSWDWH P Host rangeanddistribution NY 24083120,2408321. Aug. 1887, [ [ blitoides gangeticus paniculatus japonica complex. Ellis &Everh.,N. Amer. Fung.2582(BPI432395,FH, (1929). : : This speciesbelongs to the Delaware et al Acnida Acnidia , cannabinus ] (Crous&Braun2003)aredoubtfulandbelong : Chupp(1954:32),Vasudeva (1963:76),Katsuki sp.), , UHWURÀH[XV ], ( . 2010,Hesami A. Commons626 , designated here C. achyranthina tuberculatus hybridus C. acnidae and C. brachiata , nowasynonymof Cercospora brachiata : Faulkland,on Amaranthaceae Amaranthus Cercospora amaranthi , subsp. , spinosus , werepreviouslyassignedtothe viridis caudatus Syd., : On et al. on . Somenewerrecordsnotyet (NY 2408319). , MycoBank,MBT202808): cruentus , Africa (Kenya,South Africa, PUQKV UHWURÀH[XV $PDUDQWKXV [ Amaranthus blitum species. A carefulsearch 2011). Ann. Mycol. , Achyranthes bidentata , crassipes tamariscinus et al Amaranthus Cercospora acnidae subsp. ], Cercospora apiis. is morphologically dubius . 1996)andIran C. brachiata inLEfailed. ( Isolectotypes emarginatus albus , , hybridus 27 , cruentus usually , tricolor : 430 blitum , 18 Braun et ], . : , , et al integrated, terminal andintercalary, conidiogenousloci towards thetip,thin-walled, smooth;conidiogenouscells 0–6-septate, paletomedium brown,palerandnarrower XVXDOO\XQEUDQFKHGJHQLFXODWHVLQXRXV±î±P small hyphalaggregations,through stomata,erect,straight, fascicles, 2–15,divergent,arising frominternalhyphaeor lacking orsmall,brown,substomatal. hypophyllous, delicate,dark. sometimes causingshot-holesymptoms. pale brown,margindarker, occasionallysomewhat raised, subcircular, 1–12mmdiam,occasionallylarger, tanto Description ¿J *XRHWDO ¿J Illustrations al fascicle. Fig. 69. . (2005:30–31),Kamal(2010:31). . Cercospora celosiae B. Conidiophore tip. (OOLV ¿J& +VLHK *RK : Leaf spots (BPI 434404,holotype). C. &RQLGLD%DU P amphigenous, circularor Mycelium internal. Conidiophores Caespituli A. IMA FUNGUS Conidiophore Stromata usually in Cercosporoid fungi 4 FRQVSLFXRXV WKLFNHQHG DQG GDUNHQHG ± P GLDP ARTICLE Conidia VROLWDU\DFLFXODUVWUDLJKWWRFXUYHG±î± ± PK\DOLQH±VHSWDWHDSH[DFXWHEDVHWUXQFDWH WR VRPHZKDW REFRQLFDOO\ WUXQFDWH ± P ZLGH KLOD thickened and darkened. Holotype: China: Hubei: Wang-Chia-Shau, on Celosia argentea, 4 Aug. 1928, T. F. Yu (BPI 434404). Host range and distribution: On Celosia (argentea [argentea var. cristata, cristata, plumosa], trigyna [laxa], Celosia sp.), Amaranthaceae, Africa (Nigeria, Sudan, Uganda,), Asia (Bangladesh, Brunei, Cambodia, China, India, Indonesia, Malaysia, Myanmar, Pakistan, Papua New Guinea, South Korea, Sri Lanka, Taiwan, Thailand), Central and South America (Brazil, Panama, Venezuela), North America (Mexico; USA, Alabama, Florida, Oklahoma), West Indies (Cuba). Notes: This species is characterised by having colourless acicular conidia and thickened, darkened conidiogenous loci and hila, i.e. it belongs to the Cercospora apii s. lat. complex. Japanese records of this species (Katsuki 1965) are wrong and refer to Pseudocercospora celosiarum FRQ¿UPHGE\&K Nakashima). Cercospora celosiigena U. Braun & Bagyan., sp. nov. MycoBank MB814563 (Fig. 70) Literature: Bagyanarayana et al. (1991: 324). Illustration: Bagyanarayana et al. ¿J Diagnosis: Differs from C. celosiae in having much shorter FRQLGLRSKRUHV ± î ± P ± VHSWDWH DQG VKRUW QDUURZO\REFODYDWHF\OLQGULFDOFRQLGLD±î±P only 0–5-septate, Fig. 70. Cercospora celosiigena (HAL 2898 F, holotype). A. Description: Leaf spots amphigenous, subcircular to angular- Conidiophore fascicles. B. Conidiophores. C.&RQLGLD%DU P irregular, 1–3 mm diam, somewhat raised, greyish brown to greyish white, with a narrow purplish margin. Caespituli K\SRSK\OORXVVFDWWHUHG¿QHGDUNEURZQMycelium internal. Holotype: India: Telangana: Hyderabad, on Celosia Stromata lacking or small, formed by a few swollen hyphal argentea, Amaranthaceae, Oct. 1990, G. Bagyanarayana & cells, brown, substomatal. Conidiophores in small, divergent P. Jagadeeswar (HAL 2898 F). to dense fascicles, arising from substomatal hyphae or small stromatic hyphal aggregations, through stomata, Host range and distribution: Only known from the type erect, subcylindrial, conical, straight to somewhat curved collection. RUVOLJKWO\JHQLFXODWHVLQXRXVXQEUDQFKHG±î±P 0(–1)-septate, pale olivaceous to olivaceous-brown, thin- Notes: Bagyanarayana et al. LGHQWL¿HG WKH W\SH walled, smooth; conidiophores usually aseptate, i.e. reduced collection of this species as Cercospora celosiae which to conidiogenous cells, conidiogenous cells occasionally they considered a morphologically rather variable fungus. LQWHJUDWHG WHUPLQDO ± P ORQJ FRQLGLRJHQRXV ORFL However, C. celosiae belongs to the C. apii s. lat. complex. FRQVSLFXRXVWKLFNHQHGDQGGDUNHQHG±PGLDPConidia The conidia are consistently acicular, and the conidiophores solitary, narrowly obclavate-cylindrical, straight to slightly are much longer and septate. FXUYHG ± î ± P ±VHSWDWH K\DOLQH WKLQ walled, smooth, apex acute to subobtuse, base truncate in Cercospora gomphrenae W.W. Ray, Mycologia 36: cylindrical conidia to short obconically truncate in obclavate 172 (1944). RQHV±PZLGHKLODVOLJKWO\WKLFNHQHGDQGGDUNHQHG (Fig. 71) VOLUME 6 · NO. 2 445 446 ARTICLE P Conidiophore fascicle. Fig. 71. Cercospora gomphrenae B. Conidiophore tips. (CUP 33132,holotype). C. Conidia.Bar=10 Braun A. et al DH P ZLGH P ± SDOH Mycelium Caespituli brown, yellowishreddishbrowntopurplishbrown. JUH\LVK brown, laterdingygreytogreyishwhite,margindarker, EURZQLVK ¿UVW DW GLDP PP ± DQJXODULUUHJXODU Description ¿J Illustrations et al Crous &Braun(2003:201),ShinKim(2001:67–69),Guo Literature Synonym Cercospora gorakhanathii Iran, Nepaland Taiwan (Crous&Braun2003). areundoubtedlywrongordoubtful,e.g.thosefrom W.W. Ray Holotype and darkened. DFXWHWRVXEREWXVHEDVHWUXQFDWH±PZLGHKLODWKLFNHQHG DSH[ VPRRWK WKLQZDOOHG K\DOLQH ±VHSWDWH P ± î GLDP P ± Conidia GDUNHQHG DQG WKLFNHQHG FRQVSLFXRXVO\ ORFL FRQLGLRJHQRXV ORQJ P ± LQWHUFDODU\ DQG WHUPLQDO tip, thin-walled,smooth;conidiogenouscellsintergrated, 3–10-septate, paletomediumbrown,palertowardsthe XVXDOO\ P ± î ± XQEUDQFKHG JHQLFXODWHVLQXRXV or erumpent,erect,straight,subcylindricaltodistinctly from internalswollenhyphaeorstromata,throughstomata Conidiophores GLDP P ± FHOOV EURZQ LPPHUVHG WR VXEVWRPDWDO GLDP Illustrations Literature (Fig. 72) gomphrenaea complex. Duetoconfusionwiththename Notes Oklahoma, Texas), West Indies(Cuba). India, Andhra Pradesh;Korea),North America (USA,Georgia, [ Host rangeandditribution 39900). Isotype Gomphrena globosa and larger, lightbrowntodingy grey. LUUHJXODUVRPHWLPHVYHLQOLQLWHG±PPGLDPRUFRQÀXHQW Description ¿J decumbens . Brit. Mycol.Soc Phytopathol H. D.Bhartiya Pradesh . (2005:31),Kamal(2010:47). : This speciesispartofthe : K(M)IMI168993. solitary, acicular, straighttocurved,30–300(–450) : : : Chupp(1954:34),Ibrahim&ElNurElamin(1974), : Kamal(2010:31). Cercosporacelosiicola internal; hyphae branched, septate, subhyaline or internal; hyphaebranched,septate,subhyalineor amphigenous, punctiform, dark brown, scattered. amphigenous, punctiform,darkbrown,scattered. : : : 5DL .DPDO ¿J %KDUWL\D USA : Basti,on ]), Leaf spots 6KLQ .LP ¿J *XR Leaf spots Sawada,severalrecordsof Amaranthaceae . in small, divergent fascicles, 3–15, arising in small,divergentfascicles,3–15,arising : 56 (HCIO 42689); Oklahoma : 271(2003)[ , 18 Aug. 1942, . Stromata Celosia 89 amphigenous, circular to somewhat amphigenous, circulartosomewhat : 124(1987). amphigenous, circulartoangular- : On Topotype Stillwater, Ray’s yard,on : sp. (as“ Gomphrena , Africa (Sudan), Asia (China; A.N. Rai&Kamal, ORW DNQR± P ± RU ODFNLQJ DOPRVW isotype Bhartiya W. W. Ray W. W. : from8 Aug. 1942(CUP Caespituli Cercospora apiis.lat. holotype coronata : GPU8026]. ( globosa C. gomphrenae et al. : (CUP 33132). amphigenous, ”), Nov. 1997, IMA FUNGUS India et al Cercospora , , serrata . (2005: : Trans. Indian et al Uttar . Cercosporoid fungi 4 Holotype: India: Uttar Pradesh: Gorakhpur, University, ARTICLE botanical garden, on Celosia sp., Feb. 1981, A. N. Rai KR 536 (K(M) IMI 259305). Host range and distribution: On Celosia sp., Amaranthaceae, Asia (India, Uttar Pradesh). Note: A true Cercospora s. str. well characterised by having cylindrical to somewhat obclavate or subacicular conidia. 'RXEWIXO H[FOXGHG DQG LQVXI¿FLHQWO\ NQRZQ species Cercospora alternantherina S. Narayan et al., in Rao et al., Sugarcane Pathology 1: Fungal Diseases: 62 (1999), nom. inval. (Art. 39.1). Literature: Crous & Braun (2003: 53), Kamal (2010: 15). Illustration: Rao et al ¿J Description: Leaf spots amphigenous, circular to subcircular, 0.5–4 mm diam, olivaceous to olivaceous- brown on the upper leaf surface, pale olivaceous with brown margin below. Caespituli amphigenous, effuse. Mycelium internal; hyphae branched, septate, subhyaline WR OLJKW ROLYDFHRXV DERXW P ZLGH Stromata well- GHYHORSHG VXEVWRPDWDO DERXW P GLDP SDOH olivaceous to olivaceous. Conidiophores solitary or in fascicles of 2–6, arising from stromata, through stomata, DERXW±î±PF\OLQGULFDOVWUDLJKWWRVOLJKWO\ curved, unbranched, 1–4 times geniculate, pale to dark olivaceous, smooth; conidiogenous cells integrated, terminal and intercalary, with conspicuous conidiogenous cells, thickened and darkened. Conidia solitary, acicular, VWUDLJKWWRFXUYHGDERXW±î±P±VHSWDWH hyaline, thin-walled, smooth, apex subacute to obtuse, base truncate, hila thickened and darkened. Fig. 72. Cercospora gorakhanathii (K(M) IMI 259305, holotype). A. Holotype: India: Uttar Pradesh: without detailed locality, Conidiophores fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 on Althernanthera sessilis, Amaranthaceae, Oct. 1992, S. P Narayan (GPU 5023). Isotype: HCIO 41940. Host range and distribution: On Alternanthera sessilis, ¿QHO\ SXQFWLIRUP HIIXVH GDUN GLQJ\ ROLYDFHRXV WR EURZQ Amaranthaceae, Asia (India). Mycelium internal. Stromata developed, substomatal to LPPHUVHG ± P GLDP PHGLXP WR GDUN ROLYDFHRXV Notes: This invalid species name pertains to the Cercospora Conidiophores fasciculate, divergent, 2–12 or occasionally apii s. lat. complex. The conidia were described to be solitary, arising from stromata, through stomata or erumpent, “cylindrical” although obviously acicular conidia were depicted erect, straight, subcylindrical to geniculate-sinuous, usually in the original drawing. XQEUDQFKHGUDUHO\EUDQFKHG±î±P±VHSWDWH pale, subhyaline to pale olivaceous, thin-walled, smooth; Cercospora crassoides Davis, Trans. Wisconsin Acad. conidiogenous cells integrated, terminal or occasionally Sci. 21: 298 (1924). LQWHUFDODU\ FRQLGLRJHQRXV ORFL FRQVSLFXRXV ± P Synonym: Nimbya crassoides (Davis) E.G. Simmons, diam, thickened and darkened. Conidia solitary, cylindrical, Mycotaxon 55: 146 (1995). somewhat cylindrical-obclavate to subacicular, straight to FXUYHG ± î ± P ± ±VHSWDWH K\DOLQH Literature: Chupp (1954: 33), Crous & Braun (2003: 143). thin-walled, smooth, apex subacute or subobtuse, base WUXQFDWHWRVRPHZKDWREFRQLFDOO\WUXQFDWH±PZLGH Illustration 6LPPRQV ¿J hila thickened and darkened. VOLUME 6 · NO. 2 447 448 ARTICLE diagnosis ordescription. globosae clearly designated.Furthermore,thename from twoyears,1993and1994,butaholotypewasnot type materialwasnotavailable. The authorscitedcollections subcylindrical conidia,butavalidationisnotmadesince Notes Amaranthaceae Host rangeanddistribution 41978). 1993/1994, Pradesh Syntypes truncate, thickenedanddarkened. ±VHSWDWH XVXDOO\ P ± hyaline, thin-walled,smooth,apexsubacutetoobtuse,base î ± DERXW FXUYHG (somewhat attenuatedtowardsthetip),straighttosomewhat whitish grey, marginred. Description Illustration Literature Cercospora nothosaervae and darkened. and intercalary, conidiogenouslociconspicuous,thickened WKLQ EURZQ OLJKW walled, smooth;conidiogenouscellsintegrated,terminal WR ROLYDFHRXV ±VHSWDWH P ± î unbranched, cylindrical,1–4timesgeniculate,about45–155 ROLYDFHRXV 3–7, arisingfromstromata,erect,straightoralmostso, OLJKW GLDP P to brown. ± DERXW VXEHSLGHUPDO light olivaceous. effuse. rusty brownbelow, withmargin. 1–10 mmdiam,laterto30withishbrownabove, Description Illustration Literature Cercospora gomphrenae-globosae Holotype Texas, Wisconsin). Amaranthaceae Host rangeanddistribution 435265, CUP 39528. ÀRULGDQD Diseases (1975). 39.1). al. : This speciesiswell-characterised byhaving , inRao : Gorakhpur, on , 23Jul.1921, Mycelium : isinvalidsinceitwaspublishedwithoutanyLatin : Crous&Braun(2003:292), Kamal (2010:69). : Crous&Braun(2003:2001),Kamal(2010:47). 3DWLO ¿J : Rao : hltp notclearlyindicated): (holotype : USA Leaf spots Conidiophores S. Narayan Leaf spots : 69(1999), Conidia , Asia (India,UttarPradesh). , North America (USA,Florida,Oklahoma, : et al. et al Wisconsin internal; hyphaebranched,septate, Stromata ., ¿J circular 1–2 mm diam, centre depressed, circular 1–2mmdiam,centredepressed, SugarcanePathology solitary, subcylindricaltosubacicular amphigenous, circularorsubcicular, (GPU5051,5061;HCIO41968, Caespituli J. Davis solitary orinsmallfascicles, : LoneRock,on nom. inval M.S. Patil, Gomphrena globosa : On : On well-developed, compact, Caespituli amphigenous. (WIS). Gomphrena globosa UHLKD ÀRULGDQD )URHOLFKLD . (Arts37.3and Botanique S. Narayan C. gomphrenae- amphigenous, Isotypes India 1 Froelichia : Mycelium Mycelium Fungal , Nov. : 6 : 221 : BPI Uttar Braun et , , et al internal. ecsoapretoriensis Cercospora DH\ RÀHWGUROLYDFHRXV GDUN FRQÀXHQW EDUHO\ Caespituli RQ FRQÀXHQW reddish tored-brownborder, onthelowersurfacepaler. VRPHWLPHV GLDP the uppersurfacecentrepalebrowntodingygrey, PP with ± VXEFLUFXODU Description Crous &Braun(2003:335). Literature darkened, atmost somewhatrefractive. wide, hilaunthickenedoronly veryslightlythickened,not VXEWUXQFDWHRUVOLJKWO\VKRUWREFRQLFDOO\WUXQFDWH±P walled, smooth,apexobtuse orsubobtuse,basetruncate, WKLQ VXEK\DOLQH RU K\DOLQH SOXULVHSWDWH WR P ± î occasionally subacicular, straighttoslightly curved,15–90 somewhat darkerrim. refractive, infrontviewsometimesvisibleasminutecirclewith slightly thickened,nottodarkened,oftensomewhat sympodial proliferation(lightmicroscopy),unthickenedoronly lateral viewasconvextotruncatetiporshouldercausedby LQ YLVLEOH GLDP P ± VXEFRQVSLFXRXV RU FRQVSLFXRXV ORFL FRQLGLRJHQRXV ORQJ P ± FHOOV FRQLGLRJHQRXV WR cells integrated,terminal,sometimesconidiophoresreduced pigmented throughout,thin-walled,smooth;conidiogenous paler towardsthetip,upperpartsubhyalineorsometime 0–3-septate, rarelypluriseptate,paleolivaceous-brown, P ± î ± ± JHQLFXODWHVLQXRXV VWURQJO\ RIWHQ the tip,unbranched,straighttocurvedorusuallydistinctly, erumpent, erect,subcylindrical,somewhatattenuatedtowards GLDP densely fasciculate,fewtonumerous,arisingfromstromata, P ± DERXW DQJXODUVXEJORERVH olivaceous-brown, composedofswollenhyphalcells, Stromata Holotype prominent hilum,darkenedandthickened. ROLYDFHRXV brown, apexsubobtuse,baseobconicallytruncate,witha SDOH VHSWD WKH DW FRQVWULFWHG ±VHSWDWH P DOPRVWURVWUDWHDSH[VWUDLJKWWRVOLJKWO\FXUYHG±î± prominent. ROLYDFHRXV ±VHSWDWH brown, palertowardsthetip;conidiogenouslociconspicuous, P ± î ± VZROOHQ FHOOV ÀH[XRXV WR VWUDLJKW HUHFW non-geniculate, unbranchedoroccasionallybranched,basal HUXPSHQW ± JURXSV ORRVH (Fig. 73) was notavailable. ZDVQRWSRVVLEOHWRFRQ¿UPWKLVDVVXPSWLRQ7\SHPDWHULDO cercosporoid. Itseemstobeaspeciesof Notes collections. Host rangeanddistribution 31692). brachiata . 4 : 890(1948). : The generalcharactersofthisfungusarenot Stromata : , : Chupp(1954:35),Crous&Braun(1996:298), hypophyllous,scatteredtodense,punctiform, Amaranthaceae, LPPHUVHG±PGLDPVRPHZKDWHUXPSHQW India Conidia : Leaf spots : Maharashtra lacking. solitary, broadlyobclavatewithattenuated, Conidia amphigenous, irregularlyscattered, Conidiophores 17 Dec.1973, : Onlyknownfromthetype Chupp &Doidge, : Kolhapur, on solitary, narrowlycylindrical, Mycelium solitary or in small, solitary orinsmall, M. S.Patil Alternaria Conidiophores Nothosaerva IMA FUNGUS Bothalia internal. (HCIO , butit Cercosporoid fungi 4 s. lat. Colourless conidia are also not in favour of the latter ARTICLE genus. The conidiogenous cells and conidiogenous loci are reminiscent of Cercosporella species, but the conidiophores are pigmented, often even throughout, which would be very unusual for Cercosporella. Moreover, relations to the complex around Paracercospora and Pseudocercosporella can also not be excluded with certainty. Phylogenetic data DUH UHTXLUHG WR HOXFLGDWH WKH FRUUHFW JHQHULF DI¿QLW\ RI WKLV species. For the interim, we prefer to maintain this species in Cercospora sensu latissimo. Cercospora pseudachyranthis R.F. Castañeda & U. Braun, Cryptog. Bot. 1: 43 (1989). (Fig. 74) Synonym: Cercosporella pseudachyranthis (R.F. Cast- añeda & U. Braun) U. Braun, comb. nov. MycoBank MB814568 Basionym: Cercospora pseudachyranthis R.F. Castañeda & U. Braun, Cryptog. Bot. 1: 43 (1989). Synonyms: Cercospora centrostachydis Chupp, Monograph of Cercospora: 32 (1954), nom. inval. (Art. 39.1) [type: Puerto Rico: Mayaguez, on Achyranthes aspera var. indica, 2 Mar. 1916, Whetzel & Olive 464]. Passalora pseudachyranthis (R.F. Castañeda & U. Braun) U. Braun, Nova Hedwigia 55: 221 (1992). Misapplied name: Cercospora achyranthis sensu Solheim & Stevens (1931: 378). Literature: Solheim & Stevens (1931: 378, as C. achyranthis), Chupp (1954: 32), Crous & Braun (2003: 337). Illustration &DVWDxHGD %UDXQ SO¿J Description: Leaf spots amphigenous, subcircular to somewhat irregular, 0.5–4 mm diam, yellowish, ochraceous WREURZQLVKPDUJLQLQGH¿QLWHRUQDUURZDQGGDUNHUCaespituli hypophyllous, punctiform-effuse, greyish white. Mycelium internal. Stromata well-developed, substomatal, subglobose, ± P GLDP EURZQLVK FRPSRVHG RI VZROOHQ K\SKDO FHOOV ± P GLDP VXEJORERVHDQJXODU Conidiophores in small to large fascicles (to 50 or even more), divergent to dense, arising from stromata, emerging through stomata, Fig. 73. Cercospora pretoriensis (CUP 40641, isotype). A. HUHFWVWUDLJKWWRÀH[XRXVFXUYHGLQWKHXSSHUSDUWVOLJKWO\ Conidiophore fascicles. B. Conidiophores. C&RQLGLD%DU P to mostly distinctly geniculate-sinuous, simple, rarely EUDQFKHG ± ± ± î ± ± ± P VHSWDWH longer conidiophores pluriseptate, hyaline or subhyaline Holotype: South Africa: Transvaal: Pretoria, Arcadia, on throughout to pale olivaceous below, occasionally slightly Gomphrena globosa, 14 Apr. 1913, collector not indicated darker near the base, thin-walled, smooth; conidiogenous (PREM 6593). Isotype: CUP 40641. Paratypes: CUP40639, FHOOVLQWHJUDWHGWHUPLQDO±PORQJFRQLGLRJHQRXVORFL 40640; PREM 775, 26316, 32789. conspicuous, terminal or formed on lateral shoulders caused by sympodial proliferation, subtruncate to usually convex Host range and distribution: On Gomphrena (celosioides, (light microscopy), slightly thickened, not darkened, but often serrata [decumbens], globosa), Amaranthaceae, South VRPHZKDW UHIUDFWLYH ± P GLDP Conidia solitary, Africa. REFODYDWHF\OLQGULFDO VWUDLJKW WR VOLJKWO\ FXUYHG ± î ± P ±VHSWDWH K\DOLQH RU VXEK\DOLQH WKLQZDOOHG Notes: Cercospora pretoriensis is a cercosporoid species smooth, apex obtuse, base obconically truncate to rounded, that cannot be unequivocally assigned to any of the ±PZLGHKLODDOPRVWXQWKLFNHQHGRURQO\YHU\VOLJKWO\ currently recognised cercosporoid genera just based on thickened, not darkened, but somewhat refractive. morphology. The structure of the conidiogenous loci does not agree with scars of Cercospora s. str. and Passalora Holotype: Cuba: San Miguel de los Baños, on Achyranthes VOLUME 6 · NO. 2 449 450 ARTICLE aspera P Conidiophore fascicle. Fig. 74. Host rangeanddistribution Isotype bidentata aspera (Braun 1995). The conidiophoresarenot quitecolourless, and conidiogenouslociarein agreementwith that theconidiaandstructure ofconidiogenouscells material ofthisspecieshasbeen re-examinedandrevealed not consistentwiththecurrent conceptof the conceptsoftwogenera, andcolourlessconidiaare structure oftheconidiogenouslocidoesnotcoincidewith later reallocatedto assigned to conspicuous conidiogenousloci,thisspecieswasoriginally previously beenmisinterpreted.Duetocolourlessconidiaand Notes (Pakistan), West Indies(Cuba,PuertoRico,Virgin Islands). : The morphologicalcharacters ofthisspecieshave , 22Dec.1987, , : HAL 1648F. aspera Cercosporella pseudachyranthis , Achryranthes Cercospora var. Passalora Paratypes indica B. Conidiophore tips. R.F. Castañeda (Castañeda&Braun1989),but : On sp.), [ indica, Centrostachysindica : HAL 1761F, INIFAT C87/375. (Braun 1992).However, the Achyranthes Amaranthaceae (HAL 1648F, isotype). (INIFAT C87/365). C . Conidia.Bar=10 Passalora Cercosporella ( aspera , ?Asia . Type Braun var. A. ], et al Holotype PVHSWDWHK\DOLQHWRROLYDFHRXV VXEK\DOLQH RU SDOHU EHORZ above. EURZQ GDUNHU P î 4–10, arisingfromstromata,divergent,unbranched,18.5– EURZQ GDUN GLDP P ± brown, marginbrowntoblack,mostlyepiphyllous. Description Illustration Literature Cercospora pupaliae on theupper surface becomingchlorotic, 4–8mmdiam, of typematerial): Description Illustration Literature Cercospora sessilis 1997) isunclearandunproven. A recordof Stevens (1931),wasnottracedinChupp’s herbariuminCUP. (1954), notagreeingwiththecollectionscitedinSolheim& valid publicationin1954. The typematerialcitedinChupp a Latindescriptionordiagnosiswhichwasnecessaryfor Stevens (1931:378,as Cercospora centrostachydis (Braun 1995). known insometropical-subtropicalspeciesof above allinthelowerhalf,butthiskindofpigmentationis of revision. pupaliae VKRZVWKHPWREHREFODYDWH+HQFHWKHJHQHULFDI¿QLW\RI conidia weredescribedtobe“acicular”buttheillustration species mightbelongtothelattergenus.Furthermore, lat olivaceous conidiacouldbeanindicationfor original descriptionofconidia(hyalinetoolivaceous),although specify thesedifferences, aboveallhedidnotrefertothe he didnotprovideanydescriptionorillustrationand that thisspeciesisdistinctlydifferent from had examinedtypematerialof original descriptionismeagre.Kamal(2010)alludedthathe Notes collection. Host rangeanddistribution Amaranthaceae . ., i.e.basedontheoriginaldescriptionandillustration,this Sydowia non Mycol. Appl Chupp (1954)introducedthenewspeciesname : Type materialofthisspecies wasnotavailable. The Conidia C. sessilis isnotquiteclearandthisspeciesinurgentneed : : Crous&Braun(2003:344),Kamal(2010:79). : Crous&Braun(2003:375),Kamal(2010:85). 3DWZDUGKDQ 3DQGH ¿J 3DYJK 6LQJK ¿JV± India : (based onPavgh&Singh1964 andexamination C. centrostachydis Leaf spots 23 : , 23 Aug. 1966, : 98“1969”(1970);as“ . solitary, Maharashtra Leaf spots 23 Sorokin, 1892. : 190(1964), Pavgi &U.P. Singh, EOYW WDJW± î ± VWUDLJKW REFODYDWH circular toirregular, greytodirty C. achyranthis Patwardhan & A.K. Pande, amphigenous, greenishbrown, : Onlyknownfromthetype : Pune,on with referencetoSolheim& Conidiophores fromPakistan(Ahmad G. P. Patwardhan C. pupaliae nom. illeg. ), butfailedtoadd Pupalia orbiculata pupalae . Heemphasized Cercosporella in fasciclesof Mycopathol. Passalora s. IMA FUNGUS (Art. 53.1), (AMH 294). C. apii ”. Stromata et al , but C. . , Cercosporoid fungi 4 FRQÀXHQW Stromata VXEVWRPDWDO ± P GLDP SDOH Notes 6WDWXV DQG JHQHULF DI¿QLW\ XQFOHDU W\SH PDWHULDO ARTICLE olivaceous-brown. Conidiophores in small to moderately deposited at K(M) has been examined, but it was too meagre large fascicles, divergent to moderately dense, arising IRU D ¿QDO FRQFOXVLRQ$FFRUGLQJ WR WKH RULJLQDO SXEOLFDWLRQ from stromata, erect, straight, subcylindrical to somewhat syntype material has been deposited in HCIO, but an JHQLFXODWHVLQXRXVXQEUDQFKHG± î±PFRQWLQXRXV accession number was not cited and the material concerned to septate, light brown, thin-walled, smooth; conidiogenous was not available. Some conidiophores were found, all cells integrated, terminal. Conidia solitary, obclavate, 30–120 without any conspicuous conidiogenous loci, and a single î±P±VHSWDWHK\DOLQHRUVXEK\DOLQHWKLQZDOOHG conidium which seemed to have a slightly thickened and smooth, apex subacute, base short obconically truncate. darkened hilum. Syntypes: India: Uttar Pradesh: Varanasi, on Alternanthera sessilis, Amaranthaceae, 22 Oct. 1961, U. P. Singh (HCIO, K(M) IMI 113095). Passalora Key to Passalora species on Amaranthaceae &RQLGLDIRUPHGVLQJO\REFODYDWHF\OLQGULFDO±î±P ± ±VHSWDWHSDOHROLYDFHRXV P\FHOLXP LQWHUQDO FRQLGLRSKRUHVIDVFLFXODWHDULVLQJIURPZHOOGHYHORSHGVWURPDWD±PGLDP RQ3IDI¿D sericea ...... 3SIDI¿DH &RQLGLDDWOHDVWSDUWO\FDWHQDWHDQGRUVXSHU¿FLDOP\FHOLXPZLWKVROLWDU\FRQLGLRSKRUHVIRUPHGDQGVWURPDWDODFNLQJ ...... 2 6XSHU¿FLDOK\SKDHZLWKVROLWDU\FRQLGLRSKRUHVIRUPHG VWURPDWDODFNLQJ P\FRYHOORVLHOODOLNHVSHFLHV ...... 3 6XSHU¿FLDOK\SKDHDQGVROLWDU\FRQLGLRSKRUHVODFNLQJ FRQLGLRSKRUHVIDVFLFXODWH VWURPDWDGHYHORSHG (phaeoramularia-like species) ...... 4 &RQLGLRSKRUHVUHODWLYHO\ORQJ±î±PORQJHUFRQLGLRSKRUHVSOXULVHSWDWH RQCyathula achyranthoides ...... P. cyathulae &RQLGLRSKRUHVVKRUWHU±î±P±VHSWDWH RQIresine spp...... P. gonoclada &RQLGLDXVXDOO\F\OLQGULFDORUVXEF\OLQGULFDOK\DOLQHRUVXEK\DOLQH±î±P±VHSWDWH FRQLGLRSKRUHVQDUURZ ±PZLGH RQIresine diffusa ...... P. gilbertii &RQLGLDOVKDSHYDULDEOHF\OLQGULFDOIXVLIRUPWRREFODFDWHROLYDFHRXVWRROLYDFHRXVEURZQ±PZLGH FRQLGLRSKRUHVEURDGHU±P ...... 5 5 (4) Conidiophores occasionally branched; on Iresine diffusa ...... P. iresines Conidiophores unbranched; on Gomphrena and 3IDI¿Dspp...... P. gomphrenicola Tabular key to Passalora species on Amaranthaceae according to host genera Cyathula A single species ...... P. cyathulae Gomphrena A single species ...... P. gomphrenicola Iresine 6WURPDWDODFNLQJ FRQLGLRSKRUHVVROLWDU\DULVLQJIURPVXSHU¿FLDOK\SKDH±PORQJ±VHSWDWH ...... P. gonoclada 6WURPDWDGHYHORSHG VXSHU¿FLDOK\SKDHDQGVROLWDU\FRQLGLRSKRUHVDEVHQW FRQLGLRSKRUHVIDVFLFXODWH ± ± PORQJ±VHSWDWH ...... 2 &RQLGLDXVXDOO\F\OLQGULFDORUVXEF\OLQGULFDOK\DOLQHRUVXEK\DOLQH±î±P FRQLGLRSKRUHVQDUURZ ±PZLGHXQEUDQFKHG ...... P. gilbertii &RQLGLDOVKDSHYDULDEOHF\OLQGULFDOIXVLIRUPWRREFODFDWHROLYDFHRXVWRROLYDFHRXVEURZQ±PZLGH FRQLGLRSKRUHVEURDGHU± ± PRFFDVLRQDOO\EUDQFKHG ...... P. iresines VOLUME 6 · NO. 2 451 452 ARTICLE &RQLGLDVROLWDU\REFODYDWHF\OLQGULFDO±î±P ± Illustration Literature Basionym (Fig. 75) Passalora cyathulae Passalora specieson Amaranthaceae &RQLGLDLQFKDLQVHOOLSVRLGRYRLGEURDGO\IXVLIRUPVXEF\O 3IDI¿D Holotype collection. Host rangeanddistribution Amaranthaceae hila somewhatthickenedanddarkened. ZLGH P DERXW URXQGHG WR WUXQFDWH REFRQLFDOO\ VKRUW ±VHSWDWH P subhyaline topalebrownish,thin-walled,smooth,ends ± î ± F\OLQGULFDO WR HOOLSVRLG sometimes branchedchains,straighttosomewhatcurved, WKLQ EURZQ SDOH WR walled, smooth. VXEK\DOLQH ZLGH P ± VHSWDWH EUDQFKHG VWRPDWD WKURXJK HPHUJLQJ K\SKDH VXSHU¿FLDO effuse, dark olivaceous. LQGH¿QLWH PDUJLQ EHORZ EURZQ irregular spots,0.5–2mmdiam,brownabove,olivaceous- discolorations orsmallcircular, subcirculartosomewhat Description cited “ cyathulae confused and conidialhila,conidiaformedinchains.Chupp(1954) conidiophores, thickenedanddarkenedconidiogenousloci Passalora Notes DQGGDUNHQHG ± ± PGLDP ±PORQJFRQLGLRJHQRXVORFLFRQVSLFXRXVWKLFNHQHG sometimes conidiophoresreducedtoconidiogenouscells, SOXULVHSWDWH conidiogenous cellsintegrated,terminalorintercalary, WR DVHSWDWH P throughout, subhyalinetopalebrown,thin-walled,smooth; ± î ± YDULDEOH to geniculate-sinuous,unbranchedbranched,length ORQJ DQG GHFXPEHQWFRQLGLRSKRUHVGLI¿FXOW VWUDLJKWVXEF\OLQGULFDO K\SKDH WHUPLQDO VXSHU¿FLDO IURP RU DULVLQJ FRQLGLRSKRUHV ODWHUDO K\SKDH erect todecumbent(differentiation betweensolitary VXSHU¿FLDO IURP DULVLQJ Chupp’s (1954) descriptionof new speciesbasedon described anewIndianspecies butdidnotintroduceda which is,however, incorrect.Sydow(in Sydow & Crous, Mycologia : This speciesisatypicalmycovellosiella-like Cercospora cyathulae : : Chupp(1954:33). Ragnhildiana cyathulae : SFHZWVSUFDKSDZWVROLWDU\ ZLWK K\SKDH VXSHU¿FLDO ZLWK VSHFLHV Guyana 6ROKHLP 6WHYHQV ¿J Ragnhildianacyathulae described bySydow(1937). Furthermore,he : ...... Leaf spots Mycosphaerella and Anam. 23 , 4 Aug. 1922, 4 Aug. , : 403(1931). Stromata : Coverden,on (F. Stevens&Solheim)U.Braun Ragnhildiana cyatheae Mycelium lacking orindistinct,yellowish lacking. : Onlyknownfromthetype (F. Stevens&Solheim)Syd.” F. L.Stevens C. cyathulae and theIndian Cyathula achyranthoides Caespituli Conidiophores F. Stevens&Solheim, internal andexternal; Conidia 743(ILL 11981). : 148(2003) was basedon hypophyllous, in simpleor . Moreover, Cercospora et al. solitary, 1937) Braun , et al C. hyphae. Fig. 75. length andseptation. readily distinguishablebyobviousdifferences intheconidial characters ofbothspecies,althoughthetwospeciesare LQGULFDO ± ±î±P ± ± ± VHSWDWH Conidiophore. . ±VHSWDWH B. Passalora cyathulae 6ROLWDU\FRQLGLRSKRUHVDULVLQJIURPVXSHU¿FLDOK\SKDH D ...... &RQLGLD%DU P ...... (ILL 11981, holotype). P. gomphrenicola A. IMA FUNGUS 6XSHU¿FLDO 3SIDI¿DH Cercosporoid fungi 4 Passalora gilbertii (Speg.) U. Braun, Schlechtendalia narrow brown margin. Caespituli amphigenous, punctiform, ARTICLE 5: 62 (2000). dark brown to blackish. Mycelium internal. Stromata well- (Fig. 76) GHYHORSHG±PGLDPVXEJORERVHLPPHUVHGEURZQ Basionym: Cercospora gilbertii Speg., Anales Soc. Ci. Argent. Conidiophores numerous, in loose to very dense fascicles, 10(1): 32 (1880). DULVLQJ IURP VWURPDWD HUXPSHQW HUHFW ¿OLIRUP ÀH[XRXV Synonym: Phaeoramularia gilbertii (Speg.) U. Braun, VRPHZKDW JHQLFXODWHVLQXRXV XQEUDQFKHG ± î Schlechtendalia 2: 11 (1999). ±PDVHSWDWHWRSOXULVHSWDWHWKURXJKRXWVXEK\DOLQH to pale olivaceous, thin-walled, smooth; conidiogenous Literature: Saccardo (1886: 457), Chupp (1954: 33), Crous & FHOOV LQWHJUDWHG WHUPLQDO ± P ORQJ FRQLGLRJHQRXV Braun (2003: 197). ORFL VXEFRQVSLFXRXV PLQXWH ± P GLDP VOLJKWO\ thickened and darkened-refractive. Conidia solitary or Illustration %UDXQ ¿J catenate, in short chains, subcylindrical, occasionally QDUURZO\ REFODYDWH WR IXVLIRUP ± î ± P Exsiccatae: Syd., Fungi Exot. Exs. 1047. (0–)1–4-septate, hyaline or subhyaline, thin-walled, smooth, apex obtuse, subacute or truncate in catenate conidia, base Description: Leaf spots amphigenous, subcircular to VKRUW WR ORQJ REFRQLFDOO\ WUXQFDWH DERXW P ZLGH KLOD irregular, 1–10 mm diam, oblong patches to 20 mm in barely thickened, colourless to slightly darkened-refractive. length, brown to dingy grey, zonate, sometimes with [Holotype: Uruguay: Montevideo, on Iresine diffusa, 1876, G. Gilbert 908 (not preserved).] Neotype (designated by Crous & Braun 2003): Ecuador: Quito, Pichincha Mt., 30 Sep. 1937, H. Sydow [Syd., Fungi Exot. Exs. 1047] (HBG). Isoneotypes: Syd., Fungi Exot. Exs. 1047 (e.g. BPI 436666). Host range and distribution: On Iresine diffusa [celosia, celosioides, paniculata], Amaranthaceae, Africa (São Tomé e Príncipe), North America (Mexico), South America (Colombia, Ecuador, Uruguay), West Indies (Puerto Rico, Virgin Islands). Note: The orthographic variant “gibertii” sometimes used for this species is incorrect. The epithet was derived from the QDPHRIWKHFROOHFWRU**LOEHUW7KHSK\ORJHQHWLFDI¿QLW\RI Cercospora gilbertii, characterised by an unusual combination of morphological traits, is quite uncertain. Colourless conidia are in favour of Cercospora in its current circumscription, but catenate conidia and the structure of the conidiogenous loci argue against it. Molecular analyses are necessary to HOXFLGDWH WKH WUXH JHQHULF DI¿QLW\ RI WKLV VSHFLHV )RU WKH interim we prefer to maintain this species in Passlora s. lat. Passalora gomphrenicola (Speg.) U. Braun, Schlechtendalia 5: 64 (2000). (Fig. 77) Basionym: Cercospora gomphrenicola Speg., Anales Soc. Ci. Argent. 13: 29 (1882). Synonym: Phaeoramularia gomphrenicola (Speg.) Munt.- Cvetk., Lilloa 30: 209 (1960). Literature: Saccardo (1886: 457), Chupp (1954: 34), Ellis (1971: 308), Deighton (1979: 27), Pons & Sutton (1988: 31), Crous & Braun (2003: 201–202). Illustrations 0XQWDxROD ± ¿JV ± (OOLV ¿J 'HLJKWRQ ¿J Exsiccatae: Speg., Hongos Sud-Amer. Dec. Mycol. Argent. 45. Fig. 76. Passalora gilbertii (Syd., Fungi Exot. Exs. 1047, HBG, Description: Leaf spots lacking or only visible as yellowish neotype). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. discolorations, turning dark olivaceous by abundant %DU P formation of caespituli, patches to 12 mm diam. Caespituli VOLUME 6 · NO. 2 453 454 ARTICLE smooth; conidiogenous cellsintegrated,terminal, intercalary olivaceous toolivaceous-brown, darkerinmass,thin-walled, 1–5-septate, oftensomewhat constrictedatthesepta,pale VROLWDU\ WR ULVH FRQLGLRSKRUHV± ± î±PRIWHQLUUHJXODULQZLGWK JLYLQJ K\SKDH VXSHU¿FLDO ZLWK FRQIXVDEOH decumbent branchedconidiophores reminiscentofand somewhat geniculate-sinuous,unbranchedorbranched, decumbent, straight,subcylindrical-conicaltosinuous or or fromstromata,throughstomataerumpent,erect to or evenmoreconidiophores,arisingfrominternalhyphae loose todensefascicles,largerfasciclescomposedof50 GLDP P ± RXWOLQH LQ swollen hyphalcells,circulartosomewhatangular-irregular RI FRPSRVHG EURZQ WR SDOH VXEVWRPDWDO GLDP P ± ZLGH P ± VHSWDWH FRQÀXHQW hypophyllous, effuse, olivaceous,formingsmallpatchesor Mycelium internal, hyphaecolourless,branched, Stromata Conidiophores absent orsmall,about in smalltolarge, Braun et al GDUNHQHG±PZLGH conspicuous, sometimessubdenticulate,thickenedand sympodial, occasionallypercurrent,conidiogenousloci DQGRFFDVLRQDOO\SOHXURJHQRXV±PORQJSUROLIHUDWLRQ Holotype ± PZLGHKLODVRPHZKDWWKLFNHQHGDQGGDUNHQHG in catenateconidia,baseshortobconicallytruncate,(1.5–) P ± smooth, occasionallyfaintlyrough,apexroundedtotruncate î ± ± the septa,paleolivaceoustoolivaceous-brown,thin-walled, FXUYHG VOLJKWO\ (0–)1–4(–5)-septate, occasionallysomewhatconstricted at WR VWUDLJKW chains, ellipsoid,ovoid,broadlyfusiform,subcylindrical, glomerata 436740, 722393;K(M)IMI7706 (slide),MICH15302. Speg., HongosSud-Amer. Dec.Mycol. Argent. 45,e.g.BPI . : , Feb.1881, Argentina : Buenos Aires, Palermo,on &RQLGLD%DU P K\SKD VXSHU¿FLDO B. 914, holotype). Fig. 77. C. Spegazzini Solitaryconidiophoresarisingfrom Conidia Passalora gomphrenicola A. in simpleorbranched (LPS914). Conidiophore fascicles. C. Conidiophores. IMA FUNGUS Isotypes 3IDI¿D (LPS D : . Cercosporoid fungi 4 Host range and distribution: On Gomphrena globosa, ARTICLE 3IDI¿D (glomerata [glauca, stenophylla; Gomphrena glauca], iresinoides), Amaranthaceae, Africa (South Africa, Transvaal), South America (Argentina, Brazil, Venezuela). Passalora gonatoclada (Syd.) U. Braun & Crous, Mycosphaerella and Anam.: 202 (2003). (Fig. 78) Basionym: Cercospora gonatoclada Syd., Ann. Mycol. 23: 425 (1925). Synonyms: Ragnhildiana gonatoclada (Syd.) F. Stevens & Solheim, Mycologia 23: 403 (1931). Mycovellosiella gonatoclada (Syd.) Munt.-Cvetk., Lilloa 30: 106 & 208 (1960), nom. inval. (Art. 41.5). Mycovellosiella gonatoclada (Syd.) Deighton, Mycol. Pap. 137: 69 (1974). Literature: Chupp (1954: 35), Deighton (1974: 69), Crous & Braun (2003: 202). Illustration 0XQWDxROD ¿J$ Description: Leaf spots indistinct, formed as yellowish discolorations on the upper leaf surface. Colonies hypophyllous, effuse, brownish. Mycelium internal and H[WHUQDO VXSHU¿FLDOK\SKDHVROLWDU\RFFDVLRQDO\LQWHUWZLQHG EUDQFKHGVHSWDWHVXEK\DOLQHWRROLYDFHRXVEURZQ±P wide (sterile hyphae paler and narrow, fertile hyphae with conidiophores broader and darker), thin-walled, smooth. Stromata lacking. Conidiophores solitary, arising from VXSHU¿FLDO K\SKDH ODWHUDO RU WHUPLQDO HUHFW WR GHFXPEHQW ± î ± P ±VHSWDWH GHFXPEHQW WKUHDWV ZLWK terminal conidiophores may be much longer and pluriseptate), sometimes with intercalary cells giving rise to minute peg-like SURWXEHUDQFHVZLWKDVLQJOHWHUPLQDOVFDURQO\DERXW±P long and wide, subhyaline to pale olivaceous or olivaceous- brown, thin-walled, smooth; conidiophores mostly reduced to conidiogenous cells, conidiogenous loci conspicuous, VRPHZKDWWKLFNHQHGDQGGDUNHQHG±PGLDPConidia Fig. 78. Passalora gonatoclada (Syd., Fungi Exot. Exs. 930, CUP, catenate, in simple or branched chains, ellipsoid-ovoid to isolectotype). A. 6ROLWDU\FRQLGLRSKRUHVDULVLQJIURPVXSHU¿FLDOK\SKDH F\OLQGULFDO±î±P ± ±VHSWDWHVXEK\DOLQH and through a stoma. B. Conidiophores. C&RQLGLD%DU P to pale olivaceous-brown or brownish, thin-walled, smooth, apex rounded in terminal primary conidia or conically truncate in secondary (catenate) conidia, subdenticulate when in Passalora iresines (Munt.-Cvetk.) U. Braun & Crous, branched chains, base short obconically truncate, 1.5–2.5 Mycosphaerella and Anam.: 456 (2003) PZLGHKLODWKLFNHQHGDQGGDUNHQHG (Fig. 79) Basionym: Phaeoramularia iresines Munt.-Cvetk., Lilloa 30: Lectotype (designated here, MycoBank, MBT202803): 216 (1960). Costa Rica: San José, La Caja, on Iresine calcea, 7 Jan. 1925, H. Sydow [Syd., Fungi Exot. Exs. 930] (S-F57194). Illustrations 0XQWDxROD ¿J ¿J Isolectotypes: Syd., Fungi Exot. Exs. 930, e.g., BPI 436742, CUP , HBG, K(M) IMI 7704, MICH 15303, NY 937030. Description: Leaf spots formed as epiphyllous chlorotic Former syntypes (type locality but from 5 Jan. 1925 [H. GLVFRORUDWLRQV VFDWWHUHG WR FRQÀXHQW FLUFXODU HOOLSWLFDO Sydow, Fungi itin. Costaricensi Coll. 12]): E 417850, ILL GLIIXVH YLRODFHRXV ¿QDOO\ RFKUDFHRXV WR EURZQ Caespituli 10671. hypophyllious, velutinous, brownish to brown-olivaceous, patches subcircular or sometimes vein-limited. Mycelium Host range and distribution: On Iresine (calea, diffusa internal. Stromata moderately large, substomatal, [paniculata]), Amaranthaceae, Central America (Costa Rica), olivaceous. Conidiophores in divergent to dense fascicles, West Indies (Puerto Rico, Virgin Islands). DULVLQJ IURP VWURPDWD WKURXJK VWRPDWD HUHFW ÀH[XRXV geniculate-sinuous, simple or occasionally branched, 40– VOLUME 6 · NO. 2 455 456 ARTICLE Holotype with 1–3hila. somewhat thickenedanddarkenedhilum,apexrounded or walled, smooth,baseobconicallytruncate,withasingle WKLQ ROLYDFHRXV ±VHSWDWH P ± î ± VLJPRLG cylindrical, fusiform,obclavate,straighttocurved,rarely in simpleorbranchedchains,variableshapeandsize, loci conspicuous,thickenedanddarkened. conidiophores reducedtoconidiogenouscells, integrated, terminalorintercalary, sometimeslateral,or the tip,subhyaline,thin-walled,smooth;conidiogenouscells somewhat constrictedatsepta,olivaceous,palertowards ORQJ P base sometimesswollen,aseptatetosparinglyseptate,often ± ± EUDQFKOHWV P ± ± î ± ¿J Fig. 79. &RQLGLD%DU P Iresine diffusa her timeinSouth America, sheworkedinSerbia(University LQ 7KHIDWHRIKHUKHUEDULXPFRXOGQRW\HWEHFODUL¿HG$IWH GLHG 0XQWDxROD&YHWNRYLü W\SH DV QR KHUEDULXP Note Amaranthaceae Host rangeanddistribution : Muntañola(1960)designated acollectioninherprivate Passalora iresines : Argentina A. , 1 Aug. 1959, 1 Aug. , Conidiophore fascicles. , South America (Argentina). : Tucumán EDVHGRQ0XQWDxROD ¿J : On M. Muntañola Iresine diffusa : ElCerroSanJavier, on B. Conidiophore tips. (nottraced). Conidia [ polymorpha catenate, Braun C ], U . et al PHVGVEOEVWROQ GD GDUN GLDP P ± REORQJ WR VXEJORERVH LPPHUVHG to blackish. Caespituli dingy grey, withnarrowmarginalline,somewhatraised. angular-irregular, 1–6mmdiam,pale greyishbrownto Description Literature Basionym (Fig. 80) SIDI¿DH 3DVVDORUD Fig. 80. fascicle. curator ofBCN,Barcelona). South American collectionsarenotincluded(accordingtothe mainly fungicollectedinPortugal,arepreservedBCN,but Belgrad (J.Vukojevic, pers.comm.). A partofherherbarium, of Biology),butMuntañola’s herbariumisnotpreservedin she movedtoPortugal(UniversityofBarcelona,Faculty of Belgrade,FacultyBiology),andafterherretirement . Mycosphaerella and Anam Cercospora 3DVVDORUDSIDI¿DH B. : Chupp(1954:35). : Conidiophores. HFVRD SIDI¿DH &HUFRVSRUD amphigenous, punctiform,scattered, darkbrown : Leaf spots Mycelium : 35(1954). amphigenous, circulartosomerwhat (CUP 40533,holotype). C internal. (Chupp) U.Braun&Crous, &RQLGLD%DU P .: 318(2003). Stromata Chupp, substomatal to A. IMA FUNGUS Conidiophore Monograph Cercosporoid fungi 4 EURZQ FRPSRVHG RI VXEJORERVH FHOOV ± P GLDP Holotype: Brazil: Rio Grande do Sul: Taquari, Parque Apicola, ARTICLE Conidiophores in moderately large fascicles, divergent to on 3IDI¿D sericea, Amaranthaceae, 31 Dec. 1946, J. P. da mostly dense, arising from stromata, erect, subcylindrical, Costa Neto 2224 (CUP 40533). VLQXRXVRUVOLJKWO\JHQLFXODWHXQEUDQFKHG±î±P 0–3-septate, pale olivaceous to brown, thin-walled, smooth; Host range and distribution: Only known from the type conidiogenous cells integrated, terminal, occasionally collection. FRQLGLRSKRUHV UHGXFHG WR FRQLGLRJHQRXV FHOOV ± P long, proliferation sympodial, occasionally percurrent, with Note: Based on a combination of conspicuous conidiogenous ¿QH DQQHOODWLRQV FRQLGLRJHQRXV ORFL FRQVSLFXRXV VPDOO loci, slightly thickened and darkened, 1– P ZLGH DQG ±PGLDPVRPHZKDWWKLFNHQHGDQGGDUNHQHGConidia obclavate-cylindrical, pale olivaceous conidia, Crous & Braun solitary, obclavate-cylindrical, straight to somewhat curved, (2003) reallocated this species to Passalora. ± î ± P ± ±VHSWDWH SDOH ROLYDFHRXV WKLQ walled, smooth, apex obtuse, base short obconically truncate, ±PZLGHKLODVRPHZKDWWKLFNHQHGDQGGDUNHQHG Pseudocercospora Key to Pseudocercospora species on Amaranthaceae 6XSHU¿FLDOK\SKDHZLWKVROLWDU\FRQLGLRSKRUHVGHYHORSHG...... 2 6XSHU¿FLDOK\SKDHZLWKVROLWDU\FRQLGLRSKRUHVODFNLQJ FRQLGLRSKRUHVFRQVLVWHQWO\IDVFLFXODWH...... 4 &RQLGLDEURDG±P RQCyathula tomentosa ...... P. cyathulae &RQLGLDQDUURZHU±P...... 3 3 (2) Conidia obclavate-cylindrical, pale olivaceous-brown; on Alternanthra spp...... P. alternantherae Conidia cylindrical, subacicular to obclavate-cylindrical, subhyaline to pale olivaceous; onCelosia spp...... P. celosiarum &RQLGLRSKRUHV±PORQJ FRQLGLDQDUURZO\¿OLIRUPVXEF\OLQGULFDOVXEDFLFXODURUQDUURZO\REFODYDWHF\OLQGULFDO ± ± ± î±P±VHSWDWHKLOD±PZLGH RQGomphrena spp...... P. globosae &RQLGLRSKRUHVPXFKVKRUWHU±PORQJ FRQLGLDREFODYDWHF\OLQGULFDO± PORQJ±VHSWDWHRU LIFRQLGLRSKRUHVORQJHUFRQLGLD±PZLGH RQRWKHUKRVWV ...... 5 &RQLGLRSKRUHV±î±P±VHSWDWH FRQLGLD±î±P±VHSWDWHSDOHROLYDFHRXVEURZQ onAlternanthera spp...... P. alternantherae-nodosae &RQLGLRSKRUHVVKRUWHU±PORQJRQO\± ± VHSWDWH FRQLGLDVKRUWHUDQGQDUURZHU± î±PRU subhyaline if broader; on other hosts ...... 6 6 (5) Stromata lacking; conidiophores in small fascicles; on Gomphrena spp...... P. gomphrenae 6WURPDWDGHYHORSHG±PGLDP FRQLGLRSKRUHVPRVWO\LQODUJHUIDVFLFOHVDULVLQJIURPVWURPDWD...... 7 6WURPDWDODUJH±PGLDP FRQLGLD ± ± ± PZLGHKLOD±PZLGHVXEK\DOLQH RQGomphrena pulchella ...... P. gomphrenae-pulchellae 6WURPDWDVPDOOHU±P FRQLGLD±PZLGHKLOD±PZLGHVXEK\DOLQHWRSDOHROLYDFHRXVEURZQ on other hosts ...... 8 8 (7) Conidia mostly narrowly cylindrical with truncate base, pale olivaceous-brown; onAmaranthus spp...... P. amaranthicola Conidia obclavate-cylindrical, base short obconically truncate, subhyaline to pale olivaceous; on other hosts ...... 9 9 (8) Forming distinct leaf spots; caespituli hypophyllous, punctiform; stromata substomatal; on Chamissoa altissima ...... P. chamissoana Leaf spots lacking or only with indistinct discolorations; caespituli epiphyllous, forming sooty patches; stromata immersed; on Froelichia sp...... P. froelichiae VOLUME 6 · NO. 2 457 458 ARTICLE QDUURZO\REFODYDWHF\OLQGULFDOQDUURZHU±PVXEK\D 6WURPDWD±PGLDP FRQLGLRSKRUHV±î±P FRQ unthickened, not darkened. conidiogenous cells, lociinconspicuous, cells integrated,terminalor conidiophoresreducedto pale olivaceous-brown,thin-walled, smooth;conidiogenous ±VHSWDWH P ± î ± XQEUDQFKHG JHQLFXODWH K\SKDH VXSHU¿FLDO IURP lateral, erect,straight,subcylindrical tosinuous,slightly DULVLQJ VROLWDU\ DQG VWRPDWD 5–60, divergenttodense,arisingfromstromata,through Conidiophores JORERVHVXEVWRPDWDO±PGLDPEURZQWRGDUNEURZQ ZLGH P ± K\SKDH H[WHUQDO ZLGH P ± EURZQWKLQZDOOHGVPRRWK±PZLGHLQWHUQDOK\SKDH H[WHUQDOVXSHU¿FLDO K\SKDHEUDQFKHGVHSWDWHROLYDFHRXV JUH\LVK FRQÀXHQW hypophyllous, notveryconspicuous. RIWHQ PP ± brown, marginindistinct. î ± IXVLIRUP WR 6XSHU¿FLDOK\SKDHDQGVROLWDU\FRQLGLRSKRUHVDEVHQW FRQLGLRSK ± ± ± PZLGHVXEK\DOLQH RQ 6WURPDWD±PGLDP FRQLGLRSKRUHV±î±P 6WURPDWDGHYHORSHG±PGLDP FRQLGLRSKRUHVLQVPDOOWR ...... Stromata lacking,conidiophoresin small fascicles 1 Gomphrena A...... singlespecies Froelichia A...... singlespecies Cyathula A...... singlespecies Chamissoa A...... singlespecies Celosia A...... singlespecies Amaranthus Description Illustration Literature Pseudocercospora alternatherae Pseudocercospora specieson Amaranthaceae 6XSHU¿FLDOK\SKDHZLWKVROLWDU\FRQLGLRSKRUHVGHYHORSHG Alternanthera Tabular keytoPseudocercosporaspecieson Amaranthaceae accordingtohostgenera (Fig. 81) Mycotaxon : Kamal(2010:147). : Yen : globosa Leaf spots in smalltowell-developedfascicles,about et al 16 : 39(1982). ¿J ...... Caespituli amphigenous, scattered,oval Conidia Mycelium amphigenous, mostly J.M. Yen solitary, obclavate- internal and Gomphrena pulchella Stromata et al Braun ., et al ...... only providedwithacollectionnumber). numerous unincorporated Yen collectionsinUC(unnamed, be traced.Itispossiblethatthismaterialstillamong at UC,butthetypematerialconcernedcouldcurrentlynot holotype. The mycologicalLAMcollectionsarenowhoused Notes collection. Host rangeanddistribution 10583] (nottraced). sp., Holotype unthickened, notdarkened. WKLQZDOOHG ± ROLYDFHRXVEURZQ î smooth, apexobtuse,baseshortobconicallytruncate,hila SDOH ± ±VHSWDWH FXUYHG VOLJKWO\ P WR VWUDLJKW F\OLQGULFDO Basionym (Fig. 82) Pseudocercospora Synonyms . China ada) Goh&W.H. Hsieh, Mycopathol. Mycol. Appl Rep. Gov. Agric. Res.Inst.Taiwan FRQLGLDREFODYDWHF\OLQGULFDO Amaranthaceae RUHVRQO\LQIDVFLFOHV : Yen ...... ODUJHIDVFLFOHVDULVLQJIURPVWURPDWD OLQHWRSDOHROLYDFHRXVEURZQ RQ : LGLD¿OLIRUPVXEF\OLQGULFDOVXEDFLFXODUWR : : DOWHUQDQWKHUDHQRGLÀRUDH &HUFRVSRUD 2 India Cercosporaalternanthericola : 135(1987). et al : West Bengal . (1982)cited“LAM, Yen #10583”as ...... , 7Jan.1981, DOWHUQDQWKHUDHQRGLÀRUDH ...... : Onlyknownfromthetype 27 Trans. Mycol.Soc.Republ. : Purulea,on : 93(1965)[ P. alternantherae-nodosae P. gomphrenae-pulchellae ...... B. K.DasPcc4483 35 ...... Gomphrena : 106(1928). Pavgi &U.P. Singh, P. alternantherae P. amaranthicola P. chamissoana ...... 2 ...... P. gomphrenae holotype P. celosiarum P. froelichiae Alternanthera P. cyathulae IMA FUNGUS P. globosae Sawada, : (Saw- India [Yen : Cercosporoid fungi 4 ARTICLE Fig. 81. Pseudocercospora alternantherae (based on Yen et al. ¿J A. Conidiophore fascicles. B. Solitary conidiophores DULVLQJIURPVXSHU¿FLDOK\SKDHC&RQLGLD%DU P Uttar Pradesh: Varanasi, on Alternanthera sessilis, 17 Aug. 1963, U. P. Singh (MSP 273)]. Pseudocercospora alternanthericola (Pavgi & U.P. Singh) Deighton, Mycol. Pap. 140: 139 (1976). Literature: Chupp (1954: 31), Hsieh & Goh (1990: 16), Guo & Fig. 82. 3VHXGRFHUFRVSRUD DOWHUQDQWKHUDHQRGLÀRUDH (based Hsieh (1995: 8), Guo et al. (1998: 19), Crous & Braun (2003: RQ +VLHK *RK ¿J A. Conidiophore fascicle. B. 53), Kamal (2010: 147). Conidiophores. C&RQLGLD%DU P Illustrations 3DYJL 6LQJK SO ¿JV ± +VLHK *RK ¿J *XR +VLHK ¿J ± P ±VHSWDWH XQLIRUPO\ SDOH ROLYDFHRXVEURZQ Guo et al. ¿J to brown, thin-walled, smooth; conidiogenous cells integrated, terminal, occasionally conidiophores reduced Description: Leaf spots DW ¿UVW LQGLVWLQFW RU YLVLEOH DV to conidiogenous cells, conidiogenous loci inconspicuous small brown spots, about 2–4 mm diam, later lower leaf to visible as truncate or conically truncate tips or shoulders surface gradually turning brown, upper leaf surface also formed by sympodial proliferation, unthickened, not GLVFRORXUHG OHDYHV ¿QDOO\ GLV¿JXUHG IDGHG QHFURWLF darkened. Conidia solitary obclavate-subcylindrical, Caespituli amphigenous, forming effuse, dark olivaceous VWUDLJKWWRFXUYHG±î±P±VHSWDWHSDOH SDWFKHV ¿QDOO\ FRYHULQJ WKH ZKROH OHDI EODGH Mycelium olivaceous, thin-walled, smooth, apex obtuse, rounded to internal. Stromata VPDOO VXEVWRPDWDO ± P GLDP VXEDFXWHEDVHREFRQLFDOO\WUXQFDWH±PZLGHKLOD brown. Conidiophores in small to well-developed fascicles, unthickened, not darkened. 6–25, loose to moderately dense, arising from stromata, through stomata, erect, straight, subcylindrical to curved Lectotype (designated here, MycoBank, MBT202804): RU GLVWLQFWO\ JHQLFXODWHVLQXRXV XQEUDQFKHG ± î Taiwan: Taipei, on Alternanthera sessilis, 9 May 1924, K. VOLUME 6 · NO. 2 459 460 ARTICLE Illustrations Guo Literature Synonym Basionym reduced tosynonymy. 3VHXGRFHUFRVSRUDDOWHUQDQWKHUDHQRGLÀRUDH from Indiaon K(M) IMI31945). sessilis from Taiwan havebeenexamined[Taipei, on Notes India, UttarPradesh; Taiwan). [ Host rangeanddistribution 05136, TNS-F-220608. Pseudocercospora amaranthicola Sawada PGLDPGDUNEURZQ internal. amphigenous, punctiform,darkbrowntoblackish. diam, palebrown,marginbrown. Description ¿J (Fig. 83) latter herbarium. is stillamongthenumerous untreated Yen collectioninthe neither inPCnorUC.Itispossible thatthematerialconcerned Notes collection. Host rangeanddistribution 1971, Amaranthus tricolor Holotype truncate, hilaneitherthickenednordarkened. DSH[ obtuse orsubobtuse,basetruncatetosomewhatobconically ROLYDFHRXVEURZQ SDOH ±VHSWDWH P ± î or onlyslightlyobclavate-cylindrical,straighttocurved,15– sympodial proliferation. or visibleastruncatetipslateralshoulderscausedby conidiogenous cells,lociinconspicuous cells integrated,terminalorconidiophoresreducedto pale olivaceous-brown,thin-walled,smooth;conidiogenous RURFFDVLRQDOO\EUDQFKHG±î±P±VHSWDWH XQEUDQFKHG JHQLFXODWHVLQXRXV WR VXEF\OLQGULFDO ÀH[XRXV fascicles, arisingfromstromata,erumpent,erect,straight, QRGLÀRUD amaranticola Bull. Trimestriel Soc.Mycol.France Trimestriel Soc.Mycol.France (1978). Yen, etal. : Three duplicatesofanadditionalsyntypecollection : Type materialofthisspeciescouldnotbetraced, J. M. Yen , 5 Aug. 1907, (NTU-PPE,hb.Sawada). Stromata : ], : : Hsieh&Goh(1990:17),Guo(1995:357), Bull. Trimestriel Soc.Mycol.France : Taiwan (1998: 376),Crous&Braun(2003:55). Cercoseptoria amaranthicola : VWURPDWDWKURXJKVWRPDWDRUVROLWDU\DULVLQJIURPVXSHU¿FLDO ARTICLE hyphae, lateral, erect, straight, subcylindrical-conical, not to VWURQJO\ JHQLFXODWHVLQXRXV XQEUDQFKHG ± î ± P 0–5-septate, pale olivaceous to olivaceous-brown, thin- walled, smooth; conidiogenous cells integrated, terminal RUFRQLGLRSKRUHV UHGXFHGWRFRQLGLRJHQRXV FHOOV±P long, conidiogenous loci inconspicuous, neither thickened nor darkened. Conidia solitary, cylindrical, subacicular, long DFLFXODUWRREFODYDWHF\OLQGULFDO±î±PVWUDLJKW to curved, 0–10-septate, subhyaline to pale olivaceous, thin- walled, smooth, apex obtuse to subacute, base truncate to VKRUWREFRQLFDOO\WUXQFDWH±PZLGHKLODXQWKLFNHQHG not darkened. Holotype: India: West Bengal: Calcutta, Presidency College, on Celosia argentea, 17 Feb. 1967, A. K. Kar & M. Mandal (K(M) IMI 135869). Host range and distribution: On Celosia argentea [cristata], Amaranthaceae, Asia (China; India, West Bengal; Japan). Notes: The conidial width in this species is uniform, but the length rather variable, and relatively short in the type material. The length of conidiophores is also rather variable. Japanese FROOHFWLRQVRIWKLVVSHFLHVZHUHRULJLQDOO\ZURQJO\LGHQWL¿HG and published as Cercospora celosiae (Katsuki 1965), which ZDVFODUL¿HGLQ1DNDVKLPDet al. (2002). Pseudocercospora chamissoana R.F. Castañeda & U. Braun, Cryptog. Bot. 1: 51 (1989). (Fig. 85) Illustration &DVWDxHGD5XL] %UDXQ SO¿J Description: Leaf spots amphigenous, subcircular to irregular, ± PP GLDP EURZQLVK ODWHU SDOH PDUJLQ LQGH¿QLWH RU narrow, light to dark brown, formed as mariginal line. Caespituli hypophyllous, punctiform, dark brown to blackish, scattered. Mycelium internal. Stromata VXEVWRPDWDO ± P GLDP brown, composed of swollen hyphal cells, subglobose to DQJXODULQRXWOLQH±PGLDPConidiophores in small to moderately large fascicles, divergent to dense, arising from stromata, through stomata, erect, straight, subcylindrical- FRQLFDOWRJHQLFXODWHVLQXRXVXQEUDQFKHG±î±P 0–1-septate, pale brown, thin-walled, smooth; conidiogenous Fig. 84. Pseudocercospora celosiarum (K(M) IMI 135869, holotype). cells integrated, terminal or conidiophores reduced to A. Conidiophore fascicles. B. Solitary conidiophores arising from FRQLGLRJHQRXV FHOOV ± P ORQJ FRQLGLRJHQRXV ORFL VXSHU¿FLDOK\SKDHC. Conidiophores D. &RQLGLD%DU P inconspicuous to subdenticulate, but always unthickened and not darkened. Conidia solitary, obclavate-cylindrical to VRPHZKDWIXVRLGVWUDLJKWWRVOLJKWO\FXUYHG±î±P irregular or diffuse, size variable, 1.5–30 mm diam, brown, indistinctly 3–6-septate, subhyaline to pale olivaceous, thin- dull brown to dark greyish brown, margin narrow, darker walled, smooth, apex obtuse to subacute, base obconically brown to reddish brown. Caespituli amphigenous, punctiform, WUXQFDWH±PZLGHKLODQHLWKHUWKLFNHQHGQRUGDUNHQHG dark brown or more greyish by abundant conidiation. Mycelium LQWHUQDODQGH[WHUQDO VXSHU¿FLDOK\SKDHLISUHVHQW Holotype: Cuba: Granma: Guisa, Los Corrales, on Chamissoa HPHUJLQJWKURXJKVWRPDWDEUDQFKHGVHSWDWH±PGLDP altissima, Amaranthaceae, 15 Jun. 1987, R. F. Castañeda subhyaline, thin-walled, smooth. Stromata almost lacking to Ruiz (INIFAT, C87/167). Isotype: HAL 1651 F. well-developed, substomatal, subglobose, sometimes oblong, ±PGLDPSDOHROLYDFHRXVWREURZQConidiophores in Host range and distribution: Only known from the type small to large fascicles, divergent to very dense, arising from collection. VOLUME 6 · NO. 2 461 462 ARTICLE SDWFKHVFDXVHGE\FRORQLHVRIWKHIXQJXVPDUJLQLQGH¿QLWH diam orlarger, diffuse, belowvisibleassubcircular brownish surface onlyformedasyellowish discolorations,5–10mm Description (2010: 115–116). Literature Basionym (Fig. 86) MycoBank MB814580 Pseudocercospora cyathulae Conidiophores fascicles. Fig. 85. comb. nov. (1937). Pseudocercospora chamissoana : Chupp(1954:33),Vasudeva (1963:95),Kamal : Cercosporacyathulae : Leaf spots B. lacking oralmostso,ontheupper leaf &RQLGLD%DU P Syd., (Syd.)U.Braun, (HAL 1651F, isotype). Ann. Mycol . 35 : 239 Braun A. et al Conidiophores constrictions atsepta,thin-walled, smooth. PGLDPROLYDFHRXVWRPHGLXPROLYDFHRXVEURZQRIWHQZLWK WR RFFDVLRQDOO\ FHOOV VZROOHQ ZLGH P ± GDUNHU DQG hyphae withconidiophoresor conidiogenouscellsbroader olivaceous-brown, mostlywithout constrictionsatsepta,fertile K\SKDHQDUURZ±PVXEK\DOLQHWRSDOHROLYDFHRXVRU cover, branched,sometimesanastomosing,septate,sterile plagiotropous orclimbingleafhairs,formingadensenet or brown. Caespituli hyphae. 6XSHU¿FLDOK\SKDH Fig. 86. . C. Mycelium Pseudocercospora cyathulae Conidiophores. hypophyllous, effuse, brownishordarkolivaceous- RLD\DLLJ UP XH¿LO K\SKDH VXSHU¿FLDO IURP DULVLQJ VROLWDU\ B. QHQO QHWUD XH¿LO K\SKDH VXSHU¿FLDO H[WHUQDO DQG LQWHUQDO 6ROLWDU\FRQLGLRSKRUHVDULVLQJIURPVXSHU¿FLDO D &RQLGLD%DU P (CUP 39581,holotype). Stromata IMA FUNGUS lacking. A. Cercosporoid fungi 4 ODWHUDORUWHUPLQDOOHQJWKLQGH¿QLWH IRUPLQJGHFXPEHQWIHUWLOH ARTICLE threats, but differentiation between individual conidiophores DQG K\SKDO SRUWLRQV GLI¿FXOW RU HYHQ LPSRVVLEOH VKRUWHU lateral conidiophores arising from decumbent threats about ± P ORQJ ZLGWK DERXW ± P HUHFW WR GHFXPEHQW straight, subcylindrical to geniculate-sinuous, unbranched or branched, aseptate to pluriseptate, olivaceous, olivaceous- brown to light brown, thin-walled, smooth; conidiogenous FHOOV LQWHJUDWHG WHUPLQDO ± P ORQJ FRQLGLRJHQRXV loci inconspicuous to subconspicuous by being slightly refractive, but unthickened, occasionally visible in front view as minute circle (paracercosporoid) or subdenticulate, 1.5–2 PGLDPConidia solitary, rarely in short chains, cylindrical to vermiform, shorter conidia sometimes ovoid, obovoid, short cylindrical, broadly fusiform, straight to curved, occasionally VLJPRLG ± î ± ± P ±VHSWDWH RFFDVLRQDOO\ somewhat constricted at the septa, pale brown, thin-walled, smooth, apex broadly rounded, rarely subtruncate or somewhat attenuated, base rounded to short obconically WUXQFDWH ± P ZLGH KLOD XQWKLFNHQHG QRW GDUNHQHG sometimes somewhat darker by being refractive. Holotype: India: Uttarakhand: Dehradun, Mussoorie, Rajpur, Cyathula tomentosa, Amaranthaceae, 18 Sep. 1933, R. N. Tandon 205 (CUP 39581). Host range and distribution: Only known from the type collection. Fig. 87. Pseudocercospora froelichiae (HAL 1779 F, holotype). A. Notes: The CUP collection is the only material of this species Conidiophores fascicles. B. Conidiophores. C.&RQLGLD%DU P that could be traced and examined. Chupp (1954) confused this species with Ragnhildiana cyathulae Ł Passalora cyathulae) and used the wrong citation “Cercospora cyathulae (Stev. & Solh.) Sydow” although Sydow (1937) did not intend REFODYDWHF\OLQGULFDO ± î ± P ±VHSWDWH to introduce a new combination based on Ragnhildiana pale olivaceous, thin-walled, smooth, apex obtuse or cyathulae. He undoubtedly published a new species without VXEDFXWHEDVHVKRUWREFRQLFDOO\WUXQFDWH±PZLGHKLOD any reference to R. cyathulae. The two species are neither unthickened, not darkened. FRQVSHFL¿FQRUFRQJHQHULF Holotype: Brazil: State of Ceará: Paraipaba City, on Froelichia Pseudocercospora froelichiae U. Braun & F.O. sp., 6 Aug. 2002, F. O. Freire (HAL 1779 F). Paratype: Brazil: Freire, Cryptog. Mycol. 25: 230 (2004). Rio Grande do Norte State: Areia Branca City, on Froelichia (Fig. 87) sp., 25 Aug. 2003, F. O. Freire (HAL 1780 F). Illustrations %UDXQ )UHLUH ¿J Host range and distribution: On Froelichiasp.,Amaranthaceae, South America (Brazil, State of Ceará, Rio Grande do Norte Description: Leaf spots lacking or only with inconspicuous State). discolorations, yellowish ochraceous, brownish or occasionally purplish, 1–5 mm diam. Colonies formed on Note: Pseudocercospora froelichiae on Froelichia sp. the upper leaf surface as sooty patches caused by dense (Amaranthaceae, Gomphrenoideae) is morphologically close IUXFWL¿FDWLRQ Mycelium internal. Stromata immersed to P. chamissoana, described from Cuba on Chamissoa RU VRPHZKDW HUXPSHQW ± P GLDP ROLYDFHRXV altissima (Amaranthaceae, Amaranthoideae), which differs, EURZQ FRPSRVHG RI VZROOHQ K\SKDO FHOOV ± P GLDP however, in forming distinct leaf spots, hypophyllous caespituli walls somewhat thickened. Conidiophores in small to and substomatal stromata. moderately large fascicles, loose to dense, arising from stromata, erumpent, erect, straight, subcylindrical-conical, Pseudocercospora globosae (J.M. Yen) Deighton, VOLJKWO\ JHQLFXODWHVLQXRXV XQEUDQFKHG ± î ± P Mycol. Pap. 140: 144 (1976). 0–1(–2)-septate, pale olivaceous to olivaceous-brown, thin- (Fig. 88) walled, smooth; conidiogenous cells integrated, terminal or Basionym: Cercospora globosae J.M. Yen, Rev. Mycol. 29: FRQLGLRSKRUHV UHGXFHG WR FRQLGLRJHQRXV FHOOV ± P 224 (1964). long, conidiogenous loci inconspicuous. Conidia solitary, VOLUME 6 · NO. 2 463 464 ARTICLE RLD S[ UQDH P LHDZ\XQWKLFNHQHG and notdarkened. DOZD\V ZLGH P ± WUXQFDWH DSH[ FRQLFDO FRQLGLRJHQRXV ORQJ loci inconspicuoustodistinctly denticle-like,subcylindrical- P ± LQWHUFDODU\ RFFDVLRQDOO\ walled, smooth;conidiogenous cellsintegrated,terminal, subhyaline topaleyellowish orolivaceous-brown,thin- XQEUDQFKHGWREUDQFKHG±î±P±VHSWDWH and subcylindrical-conicaltodistinctlygeniculate-sinuous, arising fromstromata,throughstomataorerumpent,straight to moderatelylargefascicles,loosedense, EURZQ \HOORZLVK GLDP P ± subglobose tosomewhatirregular, substomataltoimmersed, to dense,velvety, greyish white. by abundantfungalcolonies. subcircular tosomewhatirregular, greyishwhitetodark grey Description ¿J Illustrations 14), Crous&Braun(2003:199). Literature Conidiophores fascicles. Fig. 88. : Yen &Lim(1980:177),BraunSivapalan(1999: Pseudocercospora globosae : Fig. 90. Pseudocercospora gomphrenae-pulchellae (HAL 1635 F, isotype). A. Conidiophores fascicles. B. Conidiophores. C. Conidia. Bar = 10 P Literature: Hsieh & Goh (1990: 18–19), Guo & Hsieh (1995: Lectotype (designated here, MycoBank, MBT202805): 10), Guo et al. (1998: 21). Taiwan: Taipei, on Gomphrena globosa, 15 Nov. 1925, K. Sawada (NTU-PPE, hb. Sawada). Isolectotype: Illustrations +VLHK *RK ¿J *XR +VLHK TNS-F-220432 ¿J *XRet al ¿J Host range and distribution: On Gomphrena globosa, Description: Leaf spots amphigenous, subcircular to elliptical, Amaranthaceae, Asia (Brunei, China, Iran, Singapore, 2–10 mm diam, centre greyish white, margin reddish, on the Taiwan). upper surface with yellowish halo, below olivaceous with dark olivaceous to brown border. Caespituli hypophyllous. Notes: Records of this species from Iran are based on Mycelium internal. Stromata lacking. Conidiophores in small Hedjaroude (1976) and Bakshi et al. (2012). fascicles, not more than six, arising from internal hyphae, erect, straight, subcylindrical to somewhat attenuated Pseudocercospora gomphrenae-pulchellae U. towards the tip, slightly geniculate-sinuous, unbranched, 20– Braun et al., Fungal Diversity 6: 28 (2001). î±P±VHSWDWHROLYDFHRXVWKLQZDOOHGVPRRWK (Fig. 90) conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, conidiogenous loci Illustration: Braun et al ¿J inconspicuous, unthickened, not darkened. Conidia solitary, cylindrical, obclavate-cylindrical to subacicular, straight Description: Leaf spots amphigenous, subcircular to irregular, WR FXUYHG ± î ± P ±VHSWDWH VXEK\DOLQH WR 1–5(–8) mm diam, centre pale, yellowish to ochraceous very pale olivaceous, thin-walled, smooth, apex obtuse to or greyish white, margin narrow, dark reddish brown to subacute, base subtruncate to obconically truncate, about blackish brown. Caespituli amphigenous, punctiform, ±PZLGHKLODXQWKLFNHQHGQRWGDUNHQHG loose to dense, blackish, later greyish white by abundant VOLUME 6 · NO. 2 465 466 ARTICLE Note collection. Host rangeanddistribution DQGQDUURZHUFRQLGLDRQO\±PZLGH has smallerstromata,muchlonger, septateconidiophores gomphrenae-pulchellae ehy1340 Delhey Gomphrena pulchella the hostsoffamiliestreatedin part4. during thecourseofmonographic studiesofcercosporoidsfungion WSP forloaningtypematerialandothercollectionsintheirkeeping NIAES, NTU,NY, NYS,PAD, PC,PDD,RREM,S, TFM, TNS and DAOM, FH,HBG,HMAS,ILL,ILLS, IUM,K,L,LE,LEP, LPS,MA, We areverygratefultothedirectorsandcuratorsofB,BPI,CUP, ACKNOWLEDGEMENTS Pseudocercospora gomphrenicola species NQRZQ LQVXI¿FLHQWO\ DQG H[FOXGHG 'RXEWIXO Holotype ZLGH hila unthickened,notdarkened. P ± WUXQFDWH REFRQLFDOO\ WR WUXQFDWH URXQGHG tinge), smooth,apexobtuse,occasionallysubacute,base P ± ±VHSWDWHVXEK\DOLQH ZLWKDYHU\SDOHJUHHQLVK Conidia darkened orthickened,infrontviewvisibleasminutecircle. RFFDVLRQDOO\ LQFRQVSLFXRXV subconspicuous, paracercosporoid,i.e.rimveryslightly ORFL FRQLGLRJHQRXV ORQJ P or conidiophoresreducedtoconidiogenouscells,10–25 walled, smooth;conidiogenouscellsintegrated,terminal î ±P±VHSWDWHVXEK\DOLQHWRSDOHROLYDFHRXVWKLQ ± XQEUDQFKHG JHQLFXODWHVLQXRXV WLS WKH WRZDUGV or erumpent.erect,straight,subcylindricalattenuated moderately dense,arisingfromstromata,throughstomata Type Literature Synonym olivaceous-brown. Stromata ±PGLDPVHSWDWHVXEK\DOLQHWRSDOHEURZQVPRRWK conidial formation. Alternaria gomphrenae Kamal (2010)reduced with cicatrizedconidiogenouscellsandrostrateconidia, based ontheoriginaldescriptionandillustrationofaspecies Note P. Chiddarwar Agric. For. Morioka 22 : : Type material ofthisspecieswasnotindicated,but : India : 511 (1957). Pseudocercospora gomphrenae RLD\RFDDHFOQUFO± ± ± ± î ± REFODYDWHF\OLQGULFDO VROLWDU\ : : : Kamal(2010:179). XVRDDWLWDSGUD P GLDP P ± LQWUDHSLGHUPDO WR VXEVWRPDWDO Argentina Alternaria gomphrenae : (BB). Maharashtra RWKHUZLVHQRWVSHFL¿HG Conidiophores Isotype Mycelium : , Prov. Buenos Aires . 9 P. gomphrenicola Amaranthaceae : 6(1926). in lackingstromata,and : Pune,on : HAL 1635F. : Onlyknownfromthetype internal; hyphaebranched, in smalltolargefascicles, Togashi, Gomphrena globosa Chidd., 17Mar. 2000, , : BahiaBlanca,on to synonymywith differs from Bull. Imp.Coll. Sci. &Cult. P. globosa Braun , R. P. P. P. 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