ARTICLE 373 et al. et al. s. lat. hyphomycetes taxonomy Key words: Ascomycota Cercospora cation of ×1000. sp. sp. nom. comb. species IMA FUNGUS · 6(2): 373–469 (2015) 373–469 · 6(2): FUNGUS IMA . The rst part of this C. thunbergiigena nom. nov., C. celosiigena nom. nov., Amaranthaceae P. viburni-nudi and P. sp. nov., P. depazeoides P. comb. nov., and nom. nov., nom. nov., , ivecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get Any of the above conditions can be waived ivecommons.org/licenses/by-nc-nd/3.0/legalcode. ) 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 and unpublished data, as well as the papers cited under Details of methods are given in the concerned, As far as new examinations are references.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 structures have been made at a magni The following All illustrations have been prepared by UB. 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). ). The fourth contribution of this The fourth contribution of this Ascomycota). ( P. viburni-erosi P. , Altingiaceae , C. sambucicola 3 ) . 2013); . 2013); . 2015). . 2015). sp. nov., sp. nov., Cercospora blepharidicola ed by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). et al et al , Aizoaceae nom. nov., nom. nov., Aizoaceae , Pseudocercospora cyathulae Mycosphaerellaceae (Braun Mycosphaerellaceae Mycosphaerellaceae rst part. The present The present rst part. Adoxaceae , . The structure of part 4 The structure of part 4 . Mycosphaerellaceae Adoxaceae comb. nov., comb. nov., C. justiciigena , P. viburnicola P. nov., var. Poaceae , and Chiharu Nakashima , and Chiharu and allied genera, Cercospora and allied genera, 2 sp. nov., sp. nov., Actinidiaceae to Amaranthaceae , t, Institut für Biologie, Bereich Geobotanik und Botanischer Garten, Herbarium, Neuwerk 21, 06099 Halle (Saale), Herbarium, Neuwerk 21, 06099 Halle Bereich Geobotanik und Botanischer Garten, t, Institut für Biologie, rst part devoted to cercosporoid fungi rst part devoted to cercosporoid fungi . 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 s. lat.), belonging to the
Amaranthaceae Actinidiaceae et al et al viburni-sargentii ) and comprise a very large Ascomycota) and comprise a very large and , ,
, var. var. , Pedro W. Crous , Pedro W. 1 Acanthaceae The present paper continues a series of comprehensive taxonomic treatments of cercosporoid fungi a series of comprehensive taxonomic The present paper continues
Cercospora speci must attribute the work in the manner You P. varia P. Cercosporella pseudachyranthis Capnodiales CBS-KNAW, Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands Utrecht, 8, 3584 CT Fungal Biodiversity Centre, Uppsalalaan CBS-KNAW, Mie 514-8507, Japan Tsu, Kurima-machiya, 1577 Mie University, Graduate School of Bioresources, Martin-Luther-Universit Submitted: 26 September 2015; Accepted: 29 October 2015; Published: 10 November 2015. Accepted: 29 October 2015; Published: Article info: Submitted: 26 September 2015; are described and illustrated in alphabetical order under the particular cercosporoid genera, supplemented by keys cercosporoid genera, supplemented by in alphabetical order under the particular are described and illustrated introduction, a survey of currently recognised cercosporoid genera, a key to detailed A to the species concerned. relevant characters were published in the the genera concerned, and a discussion of taxonomically The following taxonomic novelties are introduced: series. C. justiciae-adhatodae nov., nov., nov., nov. series initiates treatments of cercosporoid fungi on dicots and comprises species occurring on hosts belonging the species occurring on hosts belonging of cercosporoid fungi on dicots and comprises series initiates treatments the families Abstract: (formerly Germany; corresponding author e-mail: [email protected] Germany; corresponding © 2015 International Mycological Association © 2015 International Mycological conditions: distribute and transmit the work, under the following are free to share - to copy, You Attribution: Acanthaceae (Acanthaceae 2 3 1 Uwe Braun Cercosporoid fungi ( fungi Cercosporoid Non-commercial: No derivative works: For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creat transform, or build upon this work. alter, may not You for commercial purposes. may not use this work You permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. permission from the copyright holder. a comprehensive monograph of this generic complex. a comprehensive monograph of this generic complex. So far three contributions have been published: part one dealing with cercosporoid fungi on other fungi (mycophylic taxa), on ferns as well as gymnosperms (Braun ( economically group of plant pathogenic, leaf-spotting, on a wide range of relevant species that cause diseases In spite of the hosts, including numerous cultivated plants. there is no modern enormous relevance of this fungal group, comprehensive treatment of by Chupp (1954) is and the only monograph published Therefore, a monographic series with seriously outdated. treatments of cercosporoid fungi based on host families was initiated (Braun contribution is the on dicots, encompassing species on hosts of the families Acanthaceae Altingiaceae True cercosporoid fungi belong to True INTRODUCTION of cercosporoids on hosts of part two dedicated to species on monocots, excluding true part two dedicated to species on monocots, excluding true grasses (Braun General chapters with generic descriptions and keys to accepted genera are included in the doi:10.5598/imafungus.2015.06.02.09 VOLUME 6 · NO. 2 374 ARTICLE
Stromatalackingorsmall, about10–25 14 (13)Stromatawell-developed, about15–55 Conidiophoresmuch shorter, to250 13 (12)Conidiophoreslong,30–500 ×3–8 Conidianarrower, 1.5–4 12 (11)Conidia (2–)2.5–5.5(–6) if Conidiophoresandconidianarrower;about1.5–5 on atleastpartly longerthan50 11Conidiophores 40–310×4–8.5 (10) Stromatalackingorsmall,about10–45 Stromatawell-developed,10–70 10 (6) Conidiophoresmuchshorter, to62.5 8 Conidiophoresto195 Conidia wider, 3–5 9 (8) Conidianarrow, 30–150×2–4 Conidiophoreslonger, 25–155 8 (7) Conidiophoresrelativelyshort,10–30(–70) Conidia consistentlyacicular, basetruncate ...... 7 (6) Conidiaaciculartoobclavate-cylindrical,basetruncateobconically ...... 6 (3) on Stromata lackingorverysmall,<25 on Stromata30–50 5 (4) Conidiophores longer, about30–185 Central America (Guatemala) ...... 3 Conidiophoresshort,10–40×2.5–5 4 (3) mixedwithshorterobclavate-cylindricalconidia ...... Conidiaconsistentlyacicular, basetruncate,oratleastlongerconidiaacicular, pluriseptatethroughout;conidialonger, 15–360 Conidiaobclavate-cylindricalwithobconicallytruncatebase...... 3 (2) acicular-subcylindrical,20–50×2–4 Stromatalackingorsmaller, 10–30 Stromatalarge,20–85 2 (1) Conidiaformedsingly...... Conidiainchains,8–40×2–3 1 Key toCercosporaspecieson Acanthaceae Cercospora Acanthaceae Cercosporoid speciesondicots TAXONOMIC TREATMENT Lepidagathis Justicia betonica Thubergia μm diam;conidiophoreslongandaseptate,35–120×3–6 ;o te ot ...... 9 μm, average>3m;onotherhosts ...... conidiaonly2–3 spp...... μm long;on μm; conidiophoresshort,10–40×3–5 , Asia (India) ...... (India) , Asia μm wide,average>3 μm, average<3m 17 ...... μm, (0–)1–3(–4)-septate,hyaline;on μm; stromatalackingorsmall,10–30 μm, average<3mwide;on μm; conidiaratherbroad,40–360×3–8 μm diam;conidiophoresshort,10–40(–7) μm; onotherhosts ...... 11 Justicia s. lat. μm; on μm diam;and/orconidiophoresmuchlonger, 10–500 ;o te ot ...... 14 μm; onotherhosts ...... μm diam;lengthoftheconidiophoressimilar, but1–6-septate; μm, 0–1-septate;conidia25–80 μm; onCrossandra μm, withmorethantwosepta;conidialonger, about20–170 ie ...... 12 μm wide ...... μm diam;on im nohrhss ...... 15 μm diam;onotherhosts ...... μm diam;and/orconidiophoresmuchlonger, m, 0–5-septate ...... μm, 0–5-septate...... sp ...... spp...... (Acanthaceae Hygrophila μm; stromata10–70 ...... 13 ...... Braun etal. ,puiett ...... 3 μm, pluriseptate ...... Andrographis p...... spp...... spp. μm wide;onotherhostsor Andrographis to Amaranthaceae μm, 0–4-septate;conidiashort, ...... μm diam;on p...... spp...... Justicia adhatoda ;o te ot ...... 8 μm; onotherhosts ...... μm long;on μm; on spp...... μ μm; m; on Acanthus Thunbergia Justicia spicigera ...... Acanthus Cercospora justiciae-adhatodae ) [Adhatoda vasica spp...... 2 ...... μm, spp...... spp...... sp...... 10 ...... C. andrographidicola μm 5 ...... 4 ...... , ...... 6 ...... C. andrographidis C. lepidagathidis C. balaghatensis C. thunbergiana C. jacobiniicola C. crossandrae C. hygrophilae ] ...... 7 ...... C. justiciigena C. justiciicola IMA FUNGUS C. acanthi C. acanthi Cercosporoid fungi 4
15 (14) Lesions formed as brown leaf spots with inde nite margin; on Asystasia spp...... C. asystasiana ARTICLE Leasion visible as greyish white leaf spots; on other hosts ...... 16
16 (15) Conidiogenous loci 2–3 μm wide; on Justicia adhatoda ...... C. adhatodae Conidiogenous loci somewhat narrower, 1.5–2.5 μm wide; on Strobilathes ...... C. strobilanthis
17 (12) Stromata well-developed, to 60 μm diam; on Barleria and Blepharis spp...... 18 Stromata lacking or small, 10–25 μm diam; on other hosts ...... 19
18 (17) Conidiophores long, 10–210 μm; on Barleria spp...... C. barleriicola Conidiophores much shorter, 15–65 μm; on Blepharis spp...... C. blepharidicola
19 (17) Conidia very narrow, about 1.5–2.5 μm; on Ruellia spp...... C. ruellina Conidia wider, 2–4 μm; on other hosts ...... 20
20 (19) Conidiophores 10–100 μm long; on Justicia and Rhytoglossa spp...... C. diantherae Conidiophores longer, to 220 μm; on other hosts ...... 21
21 (20) Leaf spots yellowish brown to dark brown, later greyish white with dark border, 0.5–4 mm diam; on Peristrophe 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 1 Conidia acicular, base truncate, 200–240 × 3.3–5 μm ...... C. andrographidicola Conidia acicular to obclavate, base truncate to obconically truncate, shorter and narrower, 30–150 × 2–4 μm ...... 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 1 Conidia in chains, 8–40 × 2–3 μm, (0–)1–3(–4)-septate, hyaline; on Justicia adhatoda [Adhatoda vasica] ...... Cercospora justiciae-adhatodae Conidia formed singly, much longer and pluriseptate ...... 2
2 (1) Stromata large, 20–85 μm diam; conidiophores short, 10–40 × 3–5 μm, 0–4-septate; conidia short, acicular-subcylindrical, 20–50 × 2–4 μm, 0–5-septate ...... C. balaghatensis Stromata lacking or smaller, 10–30 μm diam; conidiophores much longer, 10–195 μm, pluriseptate throughout; conidia longer, 15–250 μm, pluriseptate ...... 3
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conidia 25–80 Stromatasmaller, 10–25 on (recordsfrom Africa and Asia rareandunproven) ...... Stromata30–50 6 (3) Conidianarrower, 40–180×2–4 Conidia,35–250×2.5–5 Conidiaconsistentlyacicular, basetruncate ...... 5 (4) Conidiaaciculartoobclavate-cylindrical,basetruncateobconically ...... 4 (3) Conidia acicular, narrower, 45–155×2–3 Conidia obclavate-cylindrical with obconically truncate base, acicular conidia lacking Conidiaobclavate-cylindricalwithobconicallytruncatebase,acicularconidialacking ...... Cercospora acanthi-longifolii basetruncatetosomewhatobconically ...... Conidiaaciculartosomewhatobclavate,40–360×(2–)3–8 1 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) Mycoth. Ital.1580,e.g.BPI420644]. Mycoth. Ital.1580](B700016201); isolectotypes Padova, botanicalgarden,Jun.1904, [lectotype Jul. 1937 Distr. Severin,nearCazane,on 0HP6HFĠùWL no. 2273(1876). Exs., EdnNov., Ser. Sec.,Cent. 3(Resp.Cent.23), Micol. ‘FloraVeneta Critt.’ : Cercosporellaacanthi (BUCM)]. (designatedbyCrous&Braun2003): Justicia betonica ., ser. 3,
μm diam;conidiophoreslongandaseptate,35–120×3–6 Traverso, Pass., inRabenh.,FungiEur. 15: 484(1941)[holotype μm long;on S vul. &Sandu,Acad.Române I:93(1899). , Asia (India) ...... (India) , Asia μm diam;conidiophoresmuchshorter, 10–40×2.5–5 μm; on (Pass.)D.Sacc., Hedwigia Acanthus longifolius Justicia spicigera μm; onJusticiaspp.,commoninNorth America Justicia adhatodae A. Pigal
43: 422(1904) μm, basetruncate ...... : Romania: [P. Sacc., : Sacc., Suppl. Italy Braun etal. , 16 , Central America (Guatemala) : [Adhatoda vasica yellowish toochraceous,brown, and larger, sometimeszonate,at to angular-irregular, 1–12mmdiam,sometimescon Description Ital. 191,1580. Exsiccatae: Rabenh.,FungiEur. Exs.2273.D.Sacc.,Mycoth. Illustration Crous &Braun(2003:40). Lindau (1910:133),Chupp(1954:21),Braun(1995:100), Literature conidiation, scattered. punctiform, darkbrown,later greyishwhitebyabundant occasionally somewhatraised. to greyishwhite,usuallywith anarrowdarkerborder, μm, : Saccardo(1886:448;1906:562;1972:1367), : Chupp(1954:21, ], Asia (India) (India) ...... ], Asia : Leaf spotsamphigenous,circular, subcircular μm; conidia50–100 ...... μm, 0–1-septate; ...... Mycelium g. 1). Caespituli internal. ...... 5 ...... rst palegreenish,later μm long; nally greyishbrown ...... 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- uent Cercosporoid fungi 4
cylindrical, straight to curved, 20–130 × 2–5 μm, rarely 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) μm wide, hila thickened and darkened.
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: The rst valid description of 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: asudeva (1963: 32), Crous & Braun (2003: 44), Kamal (2010: 13).
Illustration: Chowdhury (1956: 85, g. 1).
Description: Leaf spots amphigenous, 0.5–8 mm diam, often con uent, forming larger patches, subcircular to Fig. 1. Cercospora acanthi (Rabenh., Fungi Eur. Exs. 2273, HAL, irregularly shaped, greyish white. Caespituli amphigenous, lectotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. ne, dark. Mycelium internal. Stromata lacking or small, Bar = 10 μm. substomatal, brown. Conidiophores in small, divergent fascicles, arising from internal hyphae or stromatic hyphal aggregations, through stomata or erumpent, erect, straight, developed, 10–70 μm diam, substomatal to immersed, large subcylindrical to geniculate in the upper half, 40–190 × 4–6 stromata often rupturing the stomata, somewhat erumpent, μm, 0–7-septate, brown to dark brown, paler towards the tip, pale, subhyaline to yellowish brown, later dark brown, thin-walled, smooth; conidiogenous cells integrated, terminal, composed of swollen hyphal cells, 2.5–10 μm diam, circular conidiogenous loci conspicuous, about 2–3 μm wide, to somewhat irregular in shape. Conidiophores in small thickened and darkened. Conidia solitary, acicular, straight to large fascicles, dense, arising from stromata, through to curved, 35–250 × 2.5–5.5 μm, 2–22-septate, hyaline, thin- stomata or erumpent, erect, straight to curved or somewhat walled, smooth, apex pointed, base truncate, about 2–3 μm geniculate-sinuous, unbranched, 10–40(–60) × 3–7 μm, 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 μm long, mostly with a single or two, occasionally several Host range and distribution: On Justicia adhatoda [Adhatoda conspicuous conidiogenous loci, 1.5–2.5 μm diam. Conidia vasica], Acanthaceae, Asia (India, Asom, Karnataka, Uttar solitary, acicular, smaller conidia narrowly obclavate- Pradesh, West Bengal).
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Literature Cercospora andrographidicola 293593 and330436). and UttarPradeshhavebeenexamined(K(M)IMI135861, Several IndiancollectionsfromKarnataka,West Bengal Notes: This isaspeciesofthe fascicle. B.Conidiophoretips. Fig. 2. (Fig. 3) “andrographicola Chi, Cercospora adhatodae J. S.China Agric. Univ : Chi(1994:94). ”. C. Conidia.Bar=10μm. (K(M) IMI135861).A. Cercospora apiis.lat. .
11 S.Q. Chen&P.K. (3): 61(1990);as Conidiophore complex. Braun etal. 86). A.Conidiophorefascicle. Fig. 3.Cercosporaandrographidicola conidiogenous locus, thickenedanddarkened. cells integrated,terminal,usually withasingleterminal 3.3–6.7 subcylindrical, non-geniculate, unbranched,150–233× loose fascicles,4–15,arising fromstromata,erect,straight, well-developed, 16–57 below. to irregular, centrepale,marginindistinct,greyishgreen Description Illustration Caespituli μm, 2–10-septate,olivaceous; conidiogenous : Chi(1994:94, : Leaf spots amphigenous. μm diam,brown. g. 86). B. Conidium.Bar=10 amphigenous, circular, elliptical Mycelium (based onChi1994:94, internal. Conidiophores μm. IMA FUNGUS Stromata Conidia g. in Cercosporoid fungi 4
solitary, acicular, straight to somewhat curved, 200–240 × ARTICLE 3.3–5 μm, hyaline, thin-walled, smooth, apex pointed, base 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 stromata, up to 30 μm diam, shorter, geniculate conidiophores, and much shorter, acicular to obclavate conidia, and rather belongs to C. andrographidis.
Cercospora andrographidis Thirum. & Govindu, Sydowia 7: 310 (1953). (Fig. 4)
Literature: asudeva (1963: 36), Braun & Crous (2003: 57), Kamal (2010: 16), To-anun et al. (2011: 30), as “C. andrographidicola”.
Illustrations: Thirumalachar & Govindu (1953: plate I, gs 5–6), To-anun et al. (2011: 30, g. 12), as “C. andrographidicola”.
Description: Leaf spots amphigenous, circular to somewhat angular-irregular, 2–6 mm diam, at rst dingly greenish, 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 swollen hyphal cells, 5–20 μm diam, usually intraepidermal, if present to 30 μm diam, brown, cells 3–8 μm diam, wall 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 unbranched, 15–165 × 3–6.5 μm, 1–9-septate, light to fascicle. B. Solitary conidiophore. C. Conidia. Bar = 10 μm. medium dark brown, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal, occasionally intercalary, 15–30 μm long, sympodial, conidiogenous loci Note: The conidia of this species were described to be conspicuous, thickened and darkened, 2–3 μm diam. Conidia “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 & 30–150 × 2–4 μm, 1–16-septate, hyaline, thin-walled, smooth, Govindu (1953: plate I, g. 6). This could be con rmed apex subacute, base truncate to somewhat obconically on the base of an examined collection from India (West truncate, 2–2.5 μm wide, hila thickened and darkened. 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.
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and [records of nemorum Holotype and darkened. or slightlyobconicallytruncate,2–3 hyaline, thin-walled,smooth,apexsubacute,basetruncate or somewhatsigmoid,45–185×2.5–5 solitary, aciculartosomewhatobclavate,straightcurved thickened anddarkened,about2–2.5 or intercalary, about15–30 thin-walled, smooth;conidiogenouscellsintegrated,terminal 30–120(–135) ×3–6 subcylindrical todistinctlygeniculate-sinuous,unbranched, through stomata,erect,straighttosomewhatcurved, internal hyphaeorsmallsubstomatalhyphalaggregations, solitary orinsmalldivergentfascicles,2–5,arisingfrom tentatively denominatedas species, including derived fromseveralcollectionsonvariousunrelatedhost from northern Thailand usingmolecularmethods.Sequences gangetica complex. Recordsof Notes: – seenotes]. Host rangeanddistribution chelonoides internal. Caespituli scattered, subcircular, 2–5mmdiam,brown,margininde Description g. 9). Illustration Nakashima Literature (Fig. 5) Cercospora asystasiana it iscurrentlyimpossibletoresolve thecladeconcerned. species namesofCercospora possible toexcludethatseveral otherhostspeciesandolder hitherto littleknownhostrange ofthistaxon,itisnotyet from sequences basedonNorth American collectionsretrieved malloti malloti common (2013: 157). Taxa belongingtothiscladerepresentedthemost are unclearbutmightbelongto true of thiscladecomprisingplurivorous material infecting 180 (1967). Nguanhom C. C. malloti Mallotus isneithersettlednor cf. is morphologicallypartofthe This speciesispartofthe : Cercospora ), Acanthaceae : Yen &Lim(1980:155),CrousBraun(2003:70), Stromata malloti : Yen (1967:181, [coromandeliana : Singapore hypophyllous, ratherinconspicuous. et al.(2010). , 5Jan.1966, Cercospora justiciicola Leaf spots arenotyetavailable.Due to thewidebut wasbasedonNorth American et al.(2015)examinedCercospora on Mallotus japonicus $V\VWDVLDVDOLFL¿ROLD A. salicifolia lacking oralmostso. encountedinthisstudy. μm, 0–5-septate,browntodarkbrown, : BotanicalGarden,on Cercospora justiciicola , knownfromthetypecollection amphigenous, onfadedleaves, J. M. Yen 769 ] (seeCous&Braun2003:234) C. cf. nally provensinceculturesand
g. 2), Yen &Lim(1980:206, : OnAsystasia mightbeinvolved. Therefore, μm long,conidiogenousloci J.M. Yen, C. asystasiana. might belongtothisspecies malloti . The tentativeallocation on μm wide,hilathickened Cercospora apiis.lat. C. apii C. apii , clusteredinaclade inGroenewald Asystasia gangetica (PC). μm diam.Conidia μm, 4–20-septate, Rev. Mycol. -like racestoC. complex. Cercospora Conidiophores (chelonoides on Cercospora Asystasia Asystasia Mycelium species nite. et al. Braun etal. The 32: cf. ,
diam, immersed, olivaceous-brown. punctiform, dark.Mycelium irregular margin. Description Illustration Literature Cercospora balaghatensis Fig. 5.Cercosporaasystasiana fascicle. B.Conidiophore, (Fig. 6) Phytopathol : Crous&Braun(2003:74),Kamal(2010:20). : Singh(1977:18, : Leaf spots . 29:17“1976”(1977). Caespituli C. Conidia.Bar=10μm. dingy greywithlighttodark brown g. 1). internal. (PC, holotype). amphigenous, scattered, S.M. Singh,Indian Conidiophores Stromata A. IMA FUNGUS Conidiophore 20–85 in large, μm Cercosporoid fungi 4 ARTICLE
Fig. 7. Cercospora barleriicola (K(M) IMI 102433). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 μm.
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. Conidia. Bar = 10 μm. 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)]. 10–40 × 3–5 μm, 0–4-septate, pale olivaceous to brownish, thin-walled, smooth; conidiogenous cells integrated, Literature: Chupp (1954: 22), asudeva (1963: 46–47), terminal, conidiogenous loci conspicuous, thickened and Crous & Braun (2003: 76), Kamal (2010: 21), Meeboon et al. darkened, about 2–2.5 μm diam. Conidia solitary, acicular or (2007a,b), To-anun et al. (2011: 31) subcylindrical, straight to slightly curved, 20–50 × 2–4 μm, 0–5-septate, hyaline, thin-walled, smooth, apex subacute or Illustrations: Govindu & Thirumalachar (1957: plate III, g. subobtuse, base truncate, 1.5–3 μm wide, hila thickened and 10), asudeva (1963: 47, g. 16), To-anun et al. (2011: 31, darkened. g. 13).
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. μm diam, olivaceous-brown to dark brown, substomatal 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
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Holotype truncate, 1.5–3 smooth, apexpointed,basetruncatetosomewhatobconically curved, 30–220×2–4 obclavate-cylindrical withobconicallytruncatebase,straightto longer conidiaacicularwithtruncatebase,shorteronesmaybe Basionym (Fig. 8) MycoBank MB814564 Cercospora blepharidicolaU.Braun,nom.nov. thickened anddarkened,(1.5–)2–3 terminal, 10–30 to slightlythickened,smooth;conidiogenouscellsintegrated, brown orthroughoutpalertowardsthetip,wallthin μ moderately geniculate-sinuous,unbranched,10–210×2.5–6 homonym of name Notes: The collection. Host rangeanddistribution Acanthaceae Division, Toda Khurai,on Holotype truncate orsubtruncate,hilathickenedanddarkened. 10-septate, hyaline,thin-walled,smooth,apexpointed,base solitary, acicular, straighttocurved,20–112 ×1–4 cylindrical, cicatrized,thickenedanddarkened. thin-walled, smooth;conidiogenouscellsintegrated,terminal, base, about15–65×4–6 subcylindrical, olivaceous. internal. amphigenous, effuse, uniformly distributed. small tolarge,scattered,darkbrown. Description Illustration cryptic speciesareinvolved. remains unclearifasinglepolymorphousspeciesorseveral lat. Barleria Host rangeanddistribution traced). University, on variable have beenexaminedandprovedtobelongasingle on Notes: SeveralIndiancollectionsandasamplefromJamaica Thailand), West Indies(Jamaica). Karnataka, Maharashtra,MadhyaPradesh,UttarPradesh; m, 1-topluriseptatethroughout,palemediumolivaceous- Barleria illeg. Mycol. Pl.Pathol. complex. Sincephylogeneticdataarenotyetavailable,it sp.), Cercospora s.str. (Art. 53.1),non : : Stromata : : Dubeyetal.(2011: 516, g.3). India India : spp. (K(M)IMI102433,163693,226980,265815) Cercospora blepharidis Conidiophores C. blepharidis Leaf spots , Dec.1996, Acanthaceae Barleria cristata exuous, unbranched,arisingfromaswollen μ : : m wide,hilathickenedanddarkened. μ Uttar Pradesh m long, conidiogenous loci conspicuous, m long,conidiogenouslociconspicuous, Madhya Pradesh C. blepharidis well-developed, to45 μ 41: 514(2011), as“ m, 3–18-septate, hyaline, thin-walled, m, 3–18-septate,hyaline,thin-walled, C. blepharidis amphigenous, mainlyepiphyllous, R. K.Dubey μm, 1–3-septate,lightolivaceous, , Asia (India,JammuandKashmir, species belongingtothe Chidd.(Chiddarwar 1960). in loosefascicles,erect,straight, : On : Onlyknownfromthetype , 9Dec.1949, Blepharis maderaspatensis : Barleria aranasi, BaranaresHindu R.K.Dubey μ m diam. : NorthSagarForest R.K. Dubeyetal.,J.
AR-58 (HCIO439724). Chidd, 1960. (cristata, prionitis M. Payak(not blepharia μm diam,dark Conidia et al.isa Caespituli Mycelium C. apiis. Conidia solitary, ”, μm, to nom. Braun etal. , , and darkened. truncate toshortobconically truncate,probablythickened 106 ×3–6 (“ liform accordingtotheoriginal description),about53– 4–7-septate, brown. fasciculate, geniculate,unbranched, about37.5–62.5×6 with darkbrownmargin. sometimes withconcentricrings. con uent, foliage Description Illustration Literature Cercospora crossandrae g. 3). Fig. 8.Cercosporablepharidicola Madras Agric. J. 59:672(1972). Madras Agric. A. Conidiophorefascicle. : Crous&Braun(2003:143),Kamal(2010:36). : Jaganathan : μm, 5–11-septate, hyaline,apexpointed,base Leaf spots nally dryingup,brown,borderyellow, Conidia et al.(1972:671, circular toirregular, 2–5mmdiam, Mycelium B. Conidia.Bar=10
Jagan., Palan.&Narayans., solitary, acicular toobclavate (based onDubay internal. Colonies g. 1). Conidiophores et al.2011: 516, μm. IMA FUNGUS white, later μm, 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: Chupp (1954: 23, g. 4).
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- irregular, 1–5 mm diam, occasionally con uent and larger, slightly zonate, centre greyish white with narrow to often broad brown border. Caespituli amphigenous, dark brown. Mycelium internal. Stromata lacking to small, 10–25 μm 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, unbranched or rarely branched, 10–100 × 3.5–6 μm, 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. Conidia. Bar = 10 μm. integrated, terminal, 10–25 μm long, with a single to several conidiogenous loci, 2.5–3.5 μm diam. Conidia solitary, acicular or subacicular, straight to curved or occasionally Delaware, Florida, Illinois, Indiana, Kansas, Maryland, somewhat sigmoid, 40–160(–180) × 2–4 μm, pluriseptate, Missouri, Oklahoma, Texas, Washington, West irginia), distance between septa 5–15 μm, hyaline, thin-walled, South America ( enezuela). smooth, apex acute to subobtuse, base usually truncate, occasionally somewhat obconically truncate, 2–3 μm wide, 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,
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(Fig. 10) Description Illustration Literature Cercospora hygrophilae Conidiophores. Fig. 10.Cercosporahygrophilae or immersed, globosetosomewhatirregular, brown. of someswollenhyphalcells, 10–30 internal. blackish. oblong, ovaltoirregular, 5–15mmdiam,dark brownto Acad. Sci.,sect.B, Stromata : Crous&Braun(2003:222),Kamal(2010:52). : Ponnappa(1968:32, Caespituli : Leaf spots B. Conidia.Bar=10 lacking orsmall,formedasaggregations amphigenous, 67: 31(1968). amphigenous, oftenmarginal,
Ponnappa, μm. g. 1). (K(M) IMI126157). μm diam,substomatal ne, dark. Proc. Indian Mycelium Braun etal. A. Holotype and darkened. apex pointed,basetruncate,2–4 × 2–5(–6) Conidia loci conspicuous,thickenedanddarkened,2.5–4 integrated, terminal,about10–35 wall somewhatthickened,smooth;conidiogenouscells 30–500 ×3–8 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 (CUP 40082). cells orconidiogenousintegrated,terminal,10–30 walled, smooth;conidiophoresreducedtoconidiogenous μ slightly geniculate-sinuous,unbranched,10–40×2.5–5 through stomata,erect,straight,subcylindrical-conicalto divergent todense,arisingfrominternalhyphaeorstromata, Conidiophores about 10–25 [Jacobinia spicigera Holotype darkened. occasionally truncate,1–1.5 apex obtusetosubacute,baseshortobconicallytruncate, to verypalegreenishorolivaceous,thin-walled,smooth, 25–80(–105) ×(2.5–)3–4(–5) to obclavate-subcylindrical,straightslightlycurved, darkened (paracercosporoid). circles, about1 long, conidiogenouslociconspicuous,formedasminute (Fig. 11) [spinosa Host rangeanddistribution auriculata belonging tothe Notes: This speciesisatypicalmemberof Bengal). [quadrivalvis conidiophores andconidia. is misleadingandpossiblybasedonnotfullydeveloped are verylong,30–500×3–8 m, 0–1-septate,subhyalinetopaleolivaceous-brown,thin- 1: 174(1950);as“ solitary, acicular, straight,curvedtosigmoid,50–250 , : : : Chupp(1954:24). , 21Feb.1967,K.M.Ponnappa Guatemala: Chimaltenango,on internal. India Asteracantha longifolia μm, 5–22-septate,hyaline,thin-walled,smooth, : ]), Acanthaceae Leaf spotsamphigenous,subcircularto in divergenttomoderatelydensefascicles, μ : μm, 1-topluriseptatethroughout,brown, μ C. apiis.lat.complex,buttheconidiophores in smalltomoderatelylargefascicles, m diam,onlymarginslightlythickenedand m diam,darkolivaceoustobrown. Karnataka Stromata ], Acanthaceae jacobinicola : Bannerghatta,on , Asia (India,Karnataka,West μ lacking orsmall,substomatal,
: OnHygrophila m wide,barelythickenedand μ A.S. Mull.&Chupp,Ceiba m. The original description original μm. The m, 2–7(–9)-septate,hyaline Conidia , 2Oct.1941,A.S.Muller ], μm long,conidiogenous μm wide,hilathickened ”. polysperma, ringens solitary, cylindrical (K(M) IMI126157). Cercospora s.str. Justicia spicigera IMA FUNGUS Hygrophila (auriculata Caespituli μm diam. μ m
Cercosporoid fungi 4 ARTICLE
Fig. 12a. Cercospora justiciae-adhatodae (BPI 1103659, holotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 μm.
Fig. 11. Cercospora jacobiniicola (CUP 40082, holotype). A. Description: Leaf spots amphigenous, circular, subcircular Conidiophore fascicles. B. Conidiophore, C. Conidia. Bar = 10 μm. to slightly angular-irregular, 0.5–4 mm diam, at rst brown, 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 lacking or almost so to well-developed, 10–30 μm diam, Note: Tentatively maintained in Cercospora. The conidia substomatal to intraepidermal, brown, cells 2–5 μm diam, are colourless or almost so, the conidiogenous loci are wall at rst thin, later slightly thickened. Conidiophores in conspicuous, but minute, ca. 1 μm wide, and somewhat 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 af nity of this species. curved, subcylindrical or somewhat attenuated towards the tip to moderately geniculate-sinuous, unbranched, 10–50 × Cercospora justiciae-adhatodae U. Braun, sp. nov. 2–5 μm, 0–3-septate, pale olivaceous to olivaceous-brown, MycoBank MB814556 paler towards the tip, thin-walled, smooth; conidiogenous (Fig. 12a,b) cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–25 μm long, proliferation sympodial, Diagnosis: Differs from Cercospora adhatodae and all conidiogenous loci conspicuous, 1–2 μm wide, somewhat other Cercospora species on hosts of the Acanthaceae in thickened and darkened. Conidia solitary and catenate, forming small catenate conidia, (8–)12–35(–40) × 2–3 μm, in simple or occasionally branched chains, narrowly (0–)1–3(–4)-septate. cylindrical-fusiform, short obclavate, (8–)12–35(–40) × 2–3
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Holotype slightly thickenedanddarkened. subtruncate toshortobconicallytruncate,1–1.5 apex subacute,subobtuseorshortconicallytruncate,base μm, (0–)1–3(–4)-septate,hyaline,thin-walled,smooth, Symptoms onleaves. Fig. 12b.Cercosporajusticiae-adhatodae Phaeoramularia inclined toputthisspecies in hosts belongingtothe species fromallother formed inchains. These characteristicsdistinguishthe new by muchshorter, narrower, usually1–3-septateconidia the latterspecies,whichbelongsto as Notes: Type materialofthisspecieswasoriginallydeposited collection. Host rangeanddistribution 1103659). 10 Jan.1969, Hills, on Cercospora adhatodae Justicia adhatoda : 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 μm wide,hila . (including complex, (BPI Braun etal. , Basionym (Fig. 13) MycoBank MB814565 Cercospora justiciigenaU.Braun,nom.nov. 1978: 158, Fig. 13. although culturesandmoleculardataarenotyetavailable. new speciesis,atleastfortheinterim,placedin hyphomycetes (seeBraun shown inthecourseofphylogeneticstudiescercosporoid in favourof μm. Literature Cercospora macrospora Karan &Manohar., Botanique Cercosporajusticiigena : Crous&Braun (2003:40),Kamal(2010:12). : Cercosporaacanthacearum g. 1–2). Cercospora s.str. A. Conidiophore fascicle. et al.2013). Therefore, thepresent Bres., 1886. (based onKaran&Manoharachary , whichhasrecentlybeen
7: 159“1976”(1978),non var. B. Conidia. Bar=10 IMA FUNGUS macrospora Cercospora
Cercosporoid fungi 4
Illustration: Karan & Manoharachary (1978: 158, g. 1–2). ARTICLE
Description: Leaf spots epiphyllous, zonate, 4–6 mm diam, white, surrounded by a pinkish border. Mycelium internal. Stromata about 30–50 μm diam, dark brown, pseudoparenchymatic. Conidiophores fasciculate, arising from stromata, erect, straight to curved, unbranched, geniculate in the upper half, about 35–120 × 3–6 μm, aseptate, brown, colourless towards the tip; conidiophores reduced to conidiogenous cells, apex rounded with dark annular conidiogenous loci. Conidia solitary, obclavate, straight, about 50–100 × 3–4.5 μm, pluriseptate, hyaline, thin- 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 af nity of C. 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. 2005: 17, g. 1). A. Conidiophore fascicle. B. Conidia. Bar = 10 μm. Illustrations: Tai (1948: 44, g. 9), Ellis (1976: 243, g. 183 A), Guo et al. (2005: 17, g. 1). Holotype: China: Sichuan: Chengtu, on Justicia sp., 15 Oct. Description: Leaf spots amphigenous, subcircular, 3–10 mm 1944, H. C. Lin (HMAS 12127). diam, yellowish, ochraceous, pale brown to nally greyish 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 almost lacking to developed, 10–30 μm 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 25–195 × 3–6 μm, 2–10-septate, olivaceous-brown, thin- America (USA, Florida). walled, smooth; conidiogenous cells integrated, terminal to intercalary, about 15–35 μm long, conidiogenous loci Notes: A true Cercospora s. str. close to or conspeci c with thickened and darkened, 1.5–3 μm diam. 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 (15–)40–140(–155) × 3–4.5(–5) μm, (2–)3–10(–17)-septate, and might belong to C. asystasiana hyaline, thin-walled, apex subacute, base truncate to short obconically truncate, 2–3 μm wide, hila thickened and Cercospora lepidagathidis Govindu & Thirum., darkened. Sydowia 9: 222 (1955).
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Synonyms (Fig. 15) Cercospora peristrophes Holotype truncate, hilathickened. walled, smooth,apexpointed,baseshortobconically curved, 23–170×2.8–4.2 Conidia intercalary, withseveralconspicuousconidiogenousloci. tips colourless;conidiogenouscellsintegrated,terminalor 183 ×2.8–5.7 erect, mostlyunbranched,stronglygeniculate-sinuous,33– brown. lacking orsmall,composedofafewswollenhyphalcells, brown. 4–7 mmdiam,centregreyishwhite,marginpinkishor Description 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: Intheoriginalpublication,Govindu& Thirumalachar Maharashtra, UttarPradesh;Pakistan). incurva Pseudocercospora lepidagathidis. alopecuroidea Indies (PuertoRico,and 12), Illustrations 316), Kamal(2010:73). Literature Description 41, g.2),Raoetal.(1999:83, asudeva (1963:134, Sydowia (1999), al., bicalyculata Uttar Pradesh et al., [holotype 41989); Peristrophe Cercospora s.str. asudeva (1963:163, Caespituli Sugarcane Pathology Conidiophores : [hyalina solitary, obclavate-cylindrical,straighttoslightly : : : J. LivingWorld
: : Govindu& Thirumalachar (1955:plate3, : Thirumalachar &Govindu(1953:pl.2, nom. inval : India isotype asudeva (1963:133–134),Crous&Braun(2003: Leaf spots : asudeva (1963:162),Crous&Braun(2003: Leaf spotsamphigenous, circular tosomewhat Cercospora peristrophigena 7:47(1953). India are resultsofmisidenti
, Nov. 1993, μm, 1–6-septate,paleolivaceoustobrown, bicalyculata ]), Acanthaceae : amphigenous. : Nichlaul,Maharajganj,on : GPU5070];as“ : Karnataka Uttar Pradesh . (Art.39.1)and g. 92). , 20May1953,H.C.Govindu amphigenous, circularorsubcircular, in well-developed,densefascicles, Records ofthisspeciesfromWest
g. 115), Chaudhary μm, 1–14-septate,hyaline,thin- , Nov. 1993,S.Narayan(HCIO S. Narayan 2: 38(1995)[ : On irgin Islands)on , S. Narayan , Mysore,Bababudans,on g. 11). Mycelium ol. 1: , Asia (India,Karnataka, : Nichlaul,Maharajganj,on Thirum. &Govindu, Lepidagathis peristrophegena cations andbelongto nom. illeg Fungal Diseases (HCIO41989)]. internal. et al.,inRao R.K. Chaudhary holotype Lepidagathis ( et al.(1995: . (Art.53.1) Peristrophe cuspidata Stromata gs 11– : ”. g. 3), India (not : 82 Braun etal. : ,
μm. Conidiophore fascicle.B. Fig. 15.Cercosporaperistrophes surrounded byadarkerborder. greyish white,margininde brown, browntodarkcentre angular-irregular, scattered,0.5–8mmdiam,yellowish 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. abundant conidialformationorbrownish. scattered, 30–220 ×(2.5–)3–7 almost lackingorsmall, 10–25 nely punctiformtoeffuse, greyishwhiteby μ Conidiophore tips. m, 1–16-septate, paleolivaceous nite, marginalslightlyraisedor (BPI 439429,isolectotype). Caespituli nally dingygreyto C. Mycelium Conidia. Bar=10 Conidiophores amphigenous, IMA FUNGUS μ m diam, internal; A.
Cercosporoid fungi 4
to olivaceous brown, paler towards the tip, thin-walled, ARTICLE smooth; conidiogenous cells integrated, terminal and intercalary, 10–35 μm long, conidiogenous loci thickened and darkened, 2.5–3.5 μm diam. Conidia solitary, acicular or subacicular to somewhat obclavate-subcylindrical when shorter, 30–130(–150) × 2.5–4.5 μm, 1–15-septate, hyaline, thin-walled, smooth, apex subacute to obtuse, base truncate or almost so, 2–3 μm wide, hila thickened and darkened.
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, ol. 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. (1999: 87, g. 13).
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. 1999: 87, g. 13). substomatal, 10–35 μm diam, olivaceous to olivaceous- A. Conidiophore fascicle. B. C. Conidia. Bar = 10 μm. brown. Conidiophores solitary or in small fascicles, 2–5, arising from stromata, through stomata, erect, straight to geniculate-sinuous, unbranched, about 20–125 × 2.5–5 Host range and distribution: Only known from the type μm, 2–9-septate, light olivaceous to olivaceous-brown, thin- 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, give rise to secondary conidia, about 30–145 × 1.5–2.5 μm, 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. broader, 60–280 × 4–7 μm. However, if Indian and North America collections are conspeci c requires phylogenetic Holotype: India: Uttar Pradesh: Gorakhpur, Maharajganj, examination and con rmation. Indian as well as North 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.
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(Fig. 17) Illustration Guo etal.(2005:18),Kamal(2010:89). Literature Cercospora strobilanthis white orwhite, with darkerborder, narrow tomoderatelywide, mm diam,at Description (2005: 19, μm. Conidiophore fascicle.B. Fig. 17.Cercosporastrobilanthis Appl . 17:77(1962);as“ : Crous&Braun(2003:390),Guo(2001:343–344), : Chiddarwar(1962:78,pl.II, g. 2). : Leaf spots rst palegreenish,laterbrownish, amphigenous, circulartoirregular, 2–10 Conidiophore tips. Chidd., strobilanthidis (K(M) IMI83190,holotype).A. Mycopathol. Mycol. gs 1–3),Guo C. Conidia.Bar=10 ”. nally greyish et al. Braun etal. not verydistinctor symptoms. with diffuse brownishhalo, nally greyishwhite,withdarkbrownborder, sometimes or somewhatirregular, 1–7mmdiam,brown,greyish Description g. 26),Guo Illustrations al. (2005:18). 403), Braun&Urtiaga(2008:485),Kamal(2010:93),Guo (1991: 291),Braun Literature Cercospora thunbergiana hila thickenedanddarkened. base truncateorslightlyobconically truncate,2–5 somewhat constrictedatsepta, hyaline,thin-walled,smooth, curved, 40–360×(2–)3–8 Conidia conspicuous, thickenedanddarkened,(1.5–)2–5 terminal andintercalary, 10–40 the tip,thin-walled,smooth;conidiogenouscellsintegrated, 40–220 ×(3–)4–8 stomata, erect,straighttogeniculate-sinuous,unbranched, arising frominternalhyphaeorsmallstromata,through in smallfascicles,2–15,divergent,occasionallysolitary, swollen hyphalcells,10–30 Stromata diam. conidiogenous locithickenedanddarkened,1.5–2.5 conidiogenous cellsintegrated,terminalandintercalary, olivaceous-brown, wallthinorslightlythickened,smooth; 35–250 ×4–5.5μm,1–7-septatethroughout,palebrownor subcylindrical todistinctlygeniculate,1–6times,unbranched, arising fromstromata,throughstomata,erect,straight, Conidiophores absent orsmall,mainlysubstomatal,10–25 punctiform, brown toalmostblack. (Fig. 18) Holotype truncate), 1.5–2.5μm,hilathickenedanddarkened. somewhat attenuatedattheverybase(veryshortobconically thin-walled, smooth,apexpointed,basetruncatetousually to curved,30–195×3–5(–5.5) somewhat attenuatedattheverybase. lat. complex,butwellcharacterisedbyconidiathatareoften Note: Atrue Acanthaceae Host rangeanddistribution IMI 83190). Strobilanthes 198 (1965);as“ Conidia solitary, acicular to somewhatobclavate,straight : : Yen &Lim(1980:165),BraunCastañedaRuiz lacking orsmall,formingsmallaggregationsof : : Yen (1965:199, India Caespituli ne, dark. et al.(2005:20, Leaf spots sp.,12Dec.1956,P. P. Chiddarwar29 , Asia (China,Guangdong;India,Maharashtra). Cercospora s.str. belongingtotheC.apiis. solitary, aciculartoslightlyobclavate,straight in divergentfascicles,2–15,rarelysolitary, : nely punctiform,dark. Maharashtra et al.(1992:363),Crous&Braun(2003: μm, 2–9-septate,brown,palertowards thunbergiaena amphigenous, mostlyhypophyllous, Mycelium Caespituli amphigenous, circular, subcircular nally sometimeswithshot-hole μm diam,brown. μm, 3–34-septate,occasionally g. 3). g. 13), Yen &Lim(1980:223, J.M. Yen, μm long,conidiogenousloci : Mumbai,Matheran,on : On μm, 3–16-septate,hyaline, internal. hypophyllous, scattered, ”. Strobilanthes Mycelium Rev. Mycol. Stromata μm diam,brown. Conidiophores IMA FUNGUS μm wide. μm wide, internal. almost (K(M) sp., 30: μm et Cercosporoid fungi 4 ARTICLE
Fig. 18. Cercospora thunbergiana (PC, lectotype). A. Conidiophore Fig. 19. Cercospora thunbergiigena (K(M) IMI 345300, holotype). A. fascicle. B. Conidophore tips. C. Conidia. Bar = 10 μm. Conidiophore fascicle. B. Conidia. Bar = 10 μm.
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. (1995: 41, g. 4). 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 amphigenous, ne. Mycelium internal. Stromata India, Andhra Pradesh, Uttar Pradesh; Singapore), South absent or small, substomatal, 10–20 μm diam, brown. America ( enezuela), West Indies (Cuba). Conidiophores in small, loose fascicles, arising from small stromata, through stomata, erect, straight to moderately Cercospora thunbergiigena U. Braun & Crous, nom. geniculate, unbranched, about 50–220 × 4–5 μm, nov. 3–10-septate, brown, thin-walled, smooth; conidiogenous MycoBank MB814567 cells integrated, terminal and intercalary, conidiogenous loci (Fig. 19) conspicuous, thickened and darkened, about 2 μm diam. 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. × 2–3 μm, 4–14-septate, hyaline, thin-walled, smooth, apex (Art. 53.1), non C. thunbergiana J.M. Yen, 1965; as pointed, base truncate, 1.5–2 μm wide, hila thickened and “thumbergiana”. darkened.
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Illustration Literature Synonym on Basionym obclavate, 14–35.5×2–3 reduced toconidiogenouscells. aseptate, palebrown,thin-walled,smooth;conidiophores to sinuous,geniculate,unbranched,14–28.5×2.8–4.2 arising fromstromatichyphalaggregations,erect,straight hyphal cells,15–30 Stromata border. centre greyishwhite,surroundedbyapinkishorlightbrown Description Passalora acanthicola Passalora specieson Acanthaceae Conidiasolitary, obclavate,short,25–50×3.5–5 1 Key toPassaloraspecieson Acanthaceae Passalora Illustration Kamal (2010:12). Literature Cercospora acanthacearum species NQRZQ LQVXI¿FLHQWO\ DQG H[FOXGHG 'RXEWIXO narrower conidia. It resembles Notes: This speciesbelongstothe collection). Host rangeanddistribution Pandey Holotype Conidiophoresmuchshorter, 10–50 20–95 × 4–6 Conidiophores100–325 Conidiainchains,eithermuchlonger, 20–95 2 (1) on (Fig. 20) Trans. Brit.Mycol. Soc London KQHJD JUDQGLÀRUD 7KXQEHUJLD Sydowia Mycosphaerella and Anam (K(M)IMI345300). Caespituli : : Phaeoramulariaacanthicola : Chupp(1954:22). : : Cercospora : Deighton(1987:387, : Govindu& Thirumalachar (1954:pl.6, India small, onlycomposedofafewbrownswollen 156 : asudeva (1963:30),Crous&Braun(2003:40). Leaf spotscirculartoirregular, 2–4mmdiam, 8:221(1954). C. thunbergiana Adhatoda vasica : 121(1944). : Uttar Pradesh mostly epiphyllous. μ
m diam.Conidiophores acanthicola μm, 1–6-septate,paleolivaceous;on . 88:385(1987). , μ (Hansf.) U.Braun&Crous, Acanthaceae : Onlyknownfromthetype m, 1–6-septate,hyaline,thin- , butdiffers inhavingmuch : Gorakhpur, Ramgarharea, ...... see ...... μm long,pluriseptate;conidiacylindricalorsomewhatcylindrical-obclavate, Govindu& Thirum., . 1:40(2003). g. 17). Conidia Hansf., C. apiis.lat (Hansf.)Deighton, Mycelium V. P. P. , Dec.1989,V. solitary, narrowly Proc. Linn.Soc. μm; conidia8–40×2–3 fasciculate, . complex. g. 1). internal. μm, ormuchnarrower, 2–3 Braun etal. μ μm, 1–3(–4)-septate;on m, Acanthus obclavate-cylindrical, straightorslightlycurved, 20–95×4–6 in simpleorbranchedchains, cylindricalorsomewhat and darkened,about1.5–2 and intercalary, conidiogenous lociconspicuous,thickened thin-walled, smooth;conidiogenous cellsintegrated,terminal pluriseptate throughout,moderately brownorreddishbrown, geniculate-sinuous, simpleorbranched,100–325×4–6 of swollenhyphalcells,emergingthroughstomata,erect, to 10,arisingfrominternalhyphaeorsmallaggregations lacking oralmostso. internal; hyphae2.5–4 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: According toGovindu& Thirumalachar (1954),type collection. Host rangeanddistribution:Onlyknownfromthetype traced). betonica Holotype truncate. walled, smooth,apexpointed,baseshortobconically C. peristrophes Cercospora peristrophes Note: Literature μm, (0–)1–3(–4)-septate,hyaline; Tropicale dei FunghidellaSomalia 1953. (Art. 53.1),non Cercospora peristrophes arboreus , : : Crous&Braun(2003:316). Acanthaceae India : Leaf spots ): 65(1988),as“ ;o te ot ...... 2 μm; onotherhosts ...... Barleria lupulina Thirum. &Govindu,1953. : ...... Itmightbeaspeciesof Karnataka C. peristrophes Conidiophores at , 31Jan.1953, rst diffuse, yellowish,laterangular, Cercospora justiciae-adhatodae μm wide,colourless. Caespituli : Mysore,NandiHills,on E. Castell., μm diam.Conidia ...... E. Castell.isahomonymof peristrophis in smallloosefascicles, hypophyllous, effuse, (Biblioteca Agraria Thirum. &Govindu, H. C.Govindu Pseudocercospora Elenco Annotato P. barleriigena P. acanthicola ”, IMA FUNGUS nom. illeg Mycelium catenate, Stromata Justicia (not μm, . , Cercosporoid fungi 4 ARTICLE
Fig. 21. Passalora barleriigena (based on Meeboon et al. 2007a: 141, g. 1). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 μm.
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. (2007a: 141, g. 1). μm. Description: Leaf spots amphigenous, subcircular to irregular, 1–14 mm diam, pale to pale brown, margin at rst inde nite, μm, 1–6-septate, pale olivaceous, thin-walled, smooth, apex later conspicuous, dark. Caespituli amphigenous. Mycelium obtuse or subtruncate, base short obconically truncate, 1.5–2 internal. Stromata substomatal, small, 7.5–20 μm diam, μm wide, hila thickened and darkened. 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 the tip, unbranched, 1–2 times geniculate, about 20–65 × 3.5– (K(M) IMI 4557a). 5 μm, 1–3-septate, darker brown below, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, Host range and distribution: Only known from the type conidiogenous loci conspicuous, thickened and darkened, collection. 1–2 μm diam. Conidia solitary, mostly obclavate, occasionally oblong cylindrical, straight to occasionaly somewhat curved, Passalora barleriigena Meeboon & Hidayat, 25–50 × 3.5–5 μm, 1–3(–4)-septate, subhyaline to pale brown, Mycotaxon 102: 140 (2007). thin-walled, smooth, apex obtuse, base short obconically (Fig. 21) truncate, 1–2 μm wide, hila thickened and darkened.
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on 15 (14)Conidiaaciculartoobclavate-cylindrical, basetruncatetolongobconicallytruncate, 2–2.5 Conidiophoreslonger, to125 14 (13)Conidiophoresshort, 5–40 nohrhss ...... 14 onotherhosts ...... Stromatadeveloped;and/orconidiophoresshorter, to75 13 (12)Stromatalackingoralmostso;conidiophores20–130 ×4–7 Stromatadeveloped;and/orconidialonger, to150 2() Stromatalacking;conidia15–65×2–4 12 (1) Conidiophoresto60 Conidial baseconsistentlyobconicallytruncate ...... 1 Conidialbasetruncatetoobconicallytruncate;on 10 (9) Conidiaobclavate-cylindrical,subacicularconidialacking; onotherhosts ...... 8 Conidiaobclavate-cylindricaltosubacicular;conidiawithpointedapex;on 9 (8) Conidiophores 0–3-septate;onotherhosts ...... 7 Conidiophores20–50 Conidiophoresshorter, 5–50×1.5–5 8 (7) on Conidia obclavate-cylindrical,baseconsistentlyobconicallytruncate;conidiophoresshorter, to80 hila 2–2.5 Conidiophores10–80×3–5 7 (3) conidiophores to 120 Conidiacylindricalorsubcylindrical,basetruncatetosomewhatobconically truncate; Conidiophores5–50(–60)×2–5 6 (4) 4 Conidiophoresshort,5–30×1.5–3 5 (4) (pluriseptate);onotherhosts ...... Conidiophores longerandbroader, to120×2.5–6 Conidiophoresrelativelyshort,5–50(–60) Stromatadeveloped,10–40 4 (3) Stromatalackingoralmostso ...... 3 (2) Stromatadeveloped;and/orconidiophoresshorter, toabout80 Stromatalacking;conidiophores50–130×4–6.5 2 (1) and/orconidiamuchnarrower, 1.5–4 Myceliuminternal;super Myceliuminternalandexternal;super 1 Key toPseudocercosporaspecieson Acanthaceae Pseudocercospora Meeboon (CMU 28212).Paratype National Park,on Holotype 1 (10) Conidiophores 5–30×1.5–4 1 (10) : & I. Araki Thailand Rhinacanthus Cynarospermum Lepidagathis alopecuroidea Barleria lupulina (CMU28213). : Chiang MaiProvince μm wide;on : thesamelocality, 10Dec.2006, , Ruellia μm long,0–3-septate;on μm long,2–6-septate;on ...... cial hyphaewithsolitaryconidiophores , 30Dec.2005, μm long;on 15 ...... μm, 0–1(–2)-septate;on Thunbergia alata μm diam ...... μm, 0–5-septate;conidiaratherlong,60–175×3.5–5 μm, 0–1(–2)-septate;on ,ado i hre)05spae ...... 17 μm, and/or(ifshorter)0–5-septate ...... μm, 0–3-septate;on ...... μm, 0–1-septate;on : DoiSuthep-Pui μm; conidialhila0.8–2 cial hyphaewithsolitaryconidiophore Strobilanthes cusia μm, hila1–1.5 J. Meeboon ...... 3 μm ...... 5 μm ...... Rhinacanthus nasutus Justicia japonica Braun etal. μm; conidia3.5–6.5 Odontonema callistachyum μm, longerconidiophoreswithmorethantwosepta μm, hilabroader, 1.5–2.5 J. Cynarospermum Rhinacanthus nasutus Justicia galapagana μm wide;on Thunbergia μm, andaboveallnarrower, 2–5 ...... Acanthaceae Host rangeanddistribution:On μm; on μm wide;onotherhosts 8 ...... μm, and/ornarrower, 2–4 ...... ntdvlpd ...... 12 in vivonotdeveloped ...... Blechum pyramidatum Barleria cristata spp...... , DyschoristeJusticia ...... μm long;on , Asia (Thailand). , Asia ...... Acanthus guineensis ...... μm; onotherhosts 13 ...... dvlpd ...... 2 in vivodeveloped ...... Justicia μm, 4–11-septate, ...... 4 ...... P. consociata μm; ...... μm wide; μm wide; spp...... 6 ...... 11 ...... 9 ...... μm long; Barleria lupulina P. odontonematis P. thunbergiicola P. lepidagathidis P. baphiacanthi P. thunbergiae P. rhinacanthi P. rhinacanthi var. dimorpha P. blepharidis ...... 10 ...... P. justiciicola IMA FUNGUS P. barleriae P. justiciae P. acanthi P. blechi ...7 ...... , Cercosporoid fungi 4
Conidia obclavate-cylindrical, acicular conidia not formed, base 1.5–2 μ wide; on other hosts ...... 16 ARTICLE
16 (15) Conidiophores 5–30 × 1.5–4 μm, 0–1(–2)-septate (external mycelium lacking); on Dyschoriste, Justicia, Ruellia ...... P. consociata var. consociata Conidiophores to 60 μm long, 0–3-septate (external mycelium usually developed); on Rhinacanthus nasutus ...... P. rhinacanthi
17 (14) Conidiophores long, 20–125 μm; on Asystasia gangetica ...... P. asystasiae Conidiophores shorter, to 75 μm, average < 50 μm; on other hosts (see “Tabular key to Pseudocercospora species on Acanthaceae according to host genera” – further identi cation just based on morphology barely possible)
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 1 Mycelium internal, super cial hyphae with solitary conidiophores lacking ...... P. consociata var. consociata Mycelium internal and external, super cial hyphae with solitary conidiophores developed ...... 2
2 (1) Stroma lacking; conidiophores 50–130 × 4–6.5 μm; conidia cylindrical to obclavate-subcylindrical, 40–110 × 3.5–6.5 μm, hila 2–2.5 μm wide ...... P. justiciae Stromata developed, 10–25 μm diam; conidiophores shorter and narrower, 20–50 × 1–5 μm; conidia narrower, obclavate, 35–120 × 1.5–5 μm, hila 1–2 μm wide ...... P. justiciicola
Lepidagathis A single species ...... P. lepidagathidis
Odontonema A single species ...... P. odontonematis
Rhinacanthus A single species ...... P. rhinacanthi
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60–175 Stromatato30 × 3.5–5 Stromatalacking;conidiophoresshortandnarrow, 5–30×1.5–3 1 Thunbergia A singlespecies ...... Strobilanthes A singlespecies ...... Rungia 0–8-septate, subhyaline, pale yellowish green to very pale olivaceous ...... 0–8-septate,subhyaline,paleyellowishgreentoveryolivaceous ...... collection. Host rangeanddistribution 1937 (K(M)IMI7695). guineensis Holotype rounded, 1.5–2 olivaceous, apexpointed,baseobconicallytruncatetoalmost or somewhatsigmoid,30–90×2.5–4 obclavate-subcylindrical tosubacicular, straighttocurved but alwaysunthickenedandnotdarkened. long, conidiogenouslociinconspicuoustosubdenticulate, or conidiophoresreducedtoconidiogenouscells,20–30 thin-walled, smooth;conidiogenouscellsintegrated,terminal unbranched, 20–40×3–4.5 subcylindrical-conical tosomewhatgeniculate-sinuous, solitary, arisingfromsuper dense fascicles,8–50,arisingfromstromata,oroccasionally brown. Basionym Pseudocercospora asystasiae smooth. hyphae 1.5–3 internal hyphae2.5–6.5 punctiform, scattered.Mycelium raised. diam, grey-brown,withnarrowdarkbrownborder, somewhat Description Illustration (Fig. 22) Pseudocercospora acanthiDeighton, Pseudocercospora specieson Acanthaceae A singlespecies ...... Ruellia (Fig. 23) Nakashima Literature Mycol. Soc.88:381(1987). Gard. Bull., 178 (1967). Conidiophores Caespituli Stromata well-developed,20–40 : : Yen &Lim (1980:169),Crous&Braun(2003:70), : : Deighton(1987:383, , Sierra Leone:Gorahun(Tunkia), on Cercospora asystasiae : et al.(2010). Acanthaceae Leaf spots μm wide,paleolivaceous,septate,thin-walled, μm wide,hilaunthickened,notdarkened. Singapore amphigenousratherpaleolivaceous, μm diam;conidiophoreslongerandbroader, 10–80×3–5 in smalltosometimeslargeand amphigenous, subcircular, to5mm , 3 Apr. 1939, μm wide,subhyaline,super μm, 0–1(–2)-septate,palebrown, 33:169(1980). μm, 4–11-septate, paleolivaceous-brown ...... cial hyphae,erect,straight, : Onlyknownfromthetype g. 14). J.M. Yen,
internal andexternal; (J.M. Yen)J.M. Yen, μm, 3–8-septate,pale μm diam,compact, F. C.Deighton Conidia Rev. Mycol. Trans. Brit. Acanthus solitary, cial Braun etal. , M μm 32: A. Fig. 22.Pseudocercosporaacanthi irregular, 2–10mmdiam,at super Description g. 31). Illustrations Conidiophore fascicles. cial hypha μm; conidia20–95×1.5–4 P. consociata var. consociata (and ?P. justiciicola : : Yen179, (1967: Leaf spots . C.Conidia.Bar=10 μm. μm; conidiaalsolongerandbroader, ...... amphigenous, subcircular to somewhat amphigenous, subcirculartosomewhat B. Solitary conidiophorearisingfrom rst pale greenish, greyish green, rst palegreenish,greyishgreen, g. 1), Yen &Lim(1980:228, μm, (K(M) IMI7695,holotype). C. baphicacanthi P. thunbergiicola P. thunbergiae IMA FUNGUS P. rungiae ) Cercosporoid fungi 4
later yellowish brown to brownish, greyish brown, nally 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). punctiform, brown, scattered to con uent and denser. Mycelium internal. Stromata lacking or small, substomatal, 10–30 μm diam, 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 exuous, sinuous or somewhat geniculate-sinuous, unbranched, Pseudocercospora baphicacanthi W.H. Hsieh & 20–125 × 3–5 μm, 0–5-septate, olivaceous-brown, thin-walled, Goh, Bot. Bull. Acad. Sin. 30: 123 (1989). smooth; conidiogenous cells integrated, terminal, about 10–40 (Fig. 24) μm long, conidiogenous loci inconspicuous to subdenticulate, neither thickened nor darkened. Conidia solitary, obclavate- Literature: Hsieh & Goh (1990: 12), Guo & Hsieh (1995: 1), cylindrical, straight to curved, 25–90(–110) × (2.5–)3–4.5(–5) Guo et al. (1998: 11). μm, 2–8(–9)-septate, subhyaline to pale olivaceous-brown, thin- walled, smooth, apex obtuse to subacute, base short obconically truncate, 1–2 μm wide, hila unthickened, not darkened.
Holotype: Singapore: Botanical Garden, on Asystasia gangetica, 5 Jan. 1966, J. M. Yen 731 (PC).
Fig. 24. Pseudocercospora baphicacanthi (based on Hsieh & Goh 1990: 13, g. 2). A. Solitary conidiophore, arising from super cial Fig. 23. Pseudocercospora asystasiae (PC, holotype). A. hyphae or emerging through stomata. B. Conidiophores, C. Conidia. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 μm. Bar = 10 μm.
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collection. Host rangeanddistribution W. H.Hsieh(NCHUPP-162). cusia Holotype short obconicallytruncate,hilaunthickened,notdarkened. walled, smooth,apexobtuseorsubobtuse,basetruncateto 100 ×3–4 cylindrical orsubcylindrical,straighttosomewhatcurved,30– about 2 conidiogenous lociinconspicuousorvisibleastruncatetips, walled, smooth;conidiogenouscellsintegrated,terminal, septate, palebrownthroughoutorpalertowardsthetip,thin- geniculate-sinuous, simpleorbranched,20–120×3–6 hyhae, lateral,erecttodecumbent,straightcurved, emerging throughstomataorsolitary, arisingfromsuper truncate, 2–2.5 walled, smooth,apexsubacute toobtuse,baseobconically subhyaline topaleolivaceous orolivaceous-brown,thin- to somewhatcurved,(10–)15–80 ×4–5 Conidia or somewhatrefractive,butneither thickenednordarkened. inconspicuous orsubconspicuousbybeingsubdenticulate integrated, terminal,10–40 brown, tipspaler, thin-walled,smooth;conidiogenouscells 130 ×4–7 occasionally branched,somewhatgeniculate-sinuous,20– through stomata,erecttodecumbent, arising frominternalhyphalorsmallaggregations, Conidiophores velvety. Caespituli blackish brownpatches,coveringlargeleafsegments. leaf surface,darkbrownbelow, sometimesforminglarge 1–5 mmdiamorcon Description Braun (2001:58, Illustrations Literature 3DFL¿TXH Basionym Pseudocercospora barleriae lacking. super Caespituli upper leafsurface,yellowishbrownbelow, margininde sometimes con Description (1995: 2, g.1),Guo Illustration (Fig. 25) Braun, cial hyphaeemergingthroughstomata. [Baphicacanthus cusia μm wide,unthickened,notdarkened. Mycelium Conidiophores solitary, obclavate toobclavate-cylindrical,straight : 17:100(1973). : Yen &Lim(1980:156),CrousBraun(2003:76). : Cercosporabarleriae : Hsieh&Goh(1990:13, g.2),Guo μm, 2–7-septate,greenishtopaleolivaceous,thin- Taiwan: Hsitou,NantouHsien,onStrobilanthes : : hypophyllous. : Yen &Lim(1973:110, hypophyllous, effuse, darkbrown,sometimes Fungal Diversity μm, (0–)1–7-septate,olivaceoustoolivaceous- Leaf spots Leaf spots in smalltomoderatelylargefascicles,loose, μm wide,hilaunthickened, notdarkened. uent, formingyellowishpatchesonthe g. 15). internal. uent andlarger, blackishontheupper et al.(1998:10, solitary orinsmall,loosefascicles, amphigenous, angular, vein-limited, irregularly shaped,2–12mmdiam, Mycelium Stromata ], Isotype: K(M)IMI312069. : Onlyknownfromthetype 8:60(2001). μm long,conidiogenousloci Acanthaceae
g. 3;1980:207, J.M. Yen &Lim, (J.M. Yen &Lim)U. g. 1). internal andexternal; lacking oralmostso. exuous, simpleor μm, 1–8-septate, , 28Mar. 1985, Conidia Stromata solitary, g. 10), nite. Cah. cial Braun etal. μm, (Fig. 26) Holotype Synonym Pseudocercospora blechi fascicle. B.Conidiophoretips. Fig. 25. to Pseudocercospora type materialshowedthatthisspecieshastobereallocated and described“brownscars”.However, theexaminationof Notes: Yen &Lim(1980)retainedthisspecies in Asia (Singapore). Host rangeanddistribution Lim g. 8). Illustrations Literature Repert Venez. Ci.Nat 73 Pseudocercospora barleriae (PC). Topotype : : Cercosporablechi : Chupp(1954:22),Crous&Braun (2003:82). Singapore . 113 : Chupp(1954:21, g.2),Braun : 120(2002). . 8:37(1942), . : onBarleriacristata : June1972(PC). : On C. Conidia.Bar=10μm. Chupp & A.S. Mull., Bol. Soc. Barleria cristata U. Braunetal., (PC, holotype). nom. inval , 3Mar. 1972,G. . (Art.39.1). et al.(2002:121, , Acanthaceae A. IMA FUNGUS Conidiophore Cercospora Feddes , Cercosporoid fungi 4 ARTICLE
Fig. 26. Pseudocercospora blechi (CUP- -002240, holotype). A. Conidiophore fascicles. B. Conidiophore. C. Conidia. Bar = 10 μm.
Description: Leaf spots mainly epiphyllous, indistinct to irregular and somewhat reddish, 0.5–3 mm diam, sometimes con uent, margin inde nite. 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 stomata, erect, exuous, geniculate-sinuous, simple or rarely branched, 10–90 × 2–4 μm, 0–3-septate, pale olivaceous or olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–30 μm long, conidiogenous loci 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, μm. 15–65 × 2–4 μm, 1–6-septate, subhyaline to pale olivaceous, apex obtuse to subacute, base obconically truncate, 1–1.5 μm wide, hila unthickened, not darkened. 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- -002240). Isotype: Caespituli amphigenous, punctiform, brown. Mycelium IA. internal. Stromata well-developed, substomatal, 20–30 μm diam, olivaceous-brown. 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, enezuela), West Indies (Cuba, Puerto Rico, to slightly geniculate-sinuous, unbranched, 5–40 × 3–5 μm, irgin 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). conidiogenous cells, 5–25 μm long, conidiogenous loci (Fig. 27) inconspicuous. Conidia solitary, acicular to obclavate- Basionym: Cercospora blepharidis Chidd., Sydowia 13: 154 cylindrical, straight to curved, 30–80 × 2.5–3.5 μm, “1959” (1960). 3–8-septate, subyhaline to pale olivaceous, thin-walled, smooth, apex subacute to obtuse, base truncate to long Literature: Kamal (2010: 155). obconically truncate, 2–2.5 μm wide, hila unthickened, not darkened. Illustration: Chiddarwar (1960: plate , gs 4–5).
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Conidiophore fascicles. b. Fig. 28a,b.a. Basionym (Fig. 28a) Pseudocercospora consociata(G.Winter) Y.L. Guo Holotype asperrimum collection. Host rangeanddistribution 1956, P. consociatavar. var. consociata & X.J.Liu, 70 (1883). P. P. Chiddarwar : : India Cercospora consociata Pseudocercospora consociata [Blepharis asperrima : Mycosystema 2:232(1989). Maharashtra dimorpha D. Conidiophores. 2(K(M)IMI83163). (HAL 2190F, holotype). : Khandala,on : Onlyknownfromthetype ], G. Winter, Acanthaceae E. Conidia.Bar=10 μm. var. Cynarospermum consociata Hedwigia A. , 9Nov. Super (BPI435187,neotype).A.
Braun etal. 22: cial hyphae. to diffuse yellowishtobrownishdiscolorations,orforming Description Exsiccatae: Ellis&Everh.,N. Amer. Fungi2477. 111, g. 2),Guoetal.(1998:12, Illustration (2010: 166),Phengsintham & Braun(2003:136),Freire(2006:236),Kamal (1965: 7),Guo&Hsieh(1995:1), Literature g. 67). B. Solitary conidiophoresarisingfrom super : Saccardo(1886:470),Chupp(1954:23),Katsuki : Chupp(1954:21, g.3),Guo&Hsieh(1995:3, : Conidiophore fascicles.B. lacking or almost so, inconspicuous Leaf spotslackingoralmostso,inconspicuous g. 2),Phengsintham et al.2013:110). Conidiophores. etal.(1998:11), Crous cial hyphae.C. IMA FUNGUS et al.2013: C. Conidia. Cercosporoid fungi 4
subcircular to angular-irregular lesions, 1–10(–15) mm diam, terminal or conidiophores often reduced to conidiogenous ARTICLE brownish, nally greyish brown to greyish white, margin cells, 5–20 μm long, conidiogenous loci inconspicuous); inde nite or darker brown, occasionally reddish brown, otherwise as in var. consociata. sometimes slightly raised, narrow, often only formed as marginal line. Caespituli amphigenous, punctiform to effuse, Holotype: Venezuela: Lara State: Qu bor, 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 constricted at septa, 1.5–5 μm wide, subhyaline to brownish, 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 μm diam, occasionally larger, to 80 μm diam, substomatal to 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 to somewhat angular-irregular in outline, 2–6(–11) μm diam, Acanthaceae, is probably taxonomically heterogeneous. This wall to 0.8 μm wide. Conidiophores 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 80) × 2–6 μm, 0–4-septate, longer ones plainly pluriseptate, 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, 5–20 μm long, conidiogenous loci inconspicuous or visible 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). curved, 20–110(–150) × 2–4.5(–5) μm, 0–10(–12)-septate, (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). 0.8–2 μm wide, hila unthickened, not darkened. Literature: Crous & Braun (2003: 159), Kamal (2010: 171). [Holotype: USA: Illinois: on Ruellia ciliosa, A. B. Seymour]. Neotype (designated here, MycoBank, MBT202783): USA: Illustration: Kar & Mandal (1969: 338, g. 1). 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, diam, scattered, sometimes con uent, uniformly yellowish to FH. dull yellowish brown, later olivaceous by abundant fructi cation. 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 lacking or small, 10–20 μm diam, subglobose or 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, enezuela). stomata, erect, straight to curved or geniculate-sinuous, unbranched, 10–55 × 3–5 μm, 0–5-septate, yellowish brown, 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) cells, 10–20 μm long, conidiogenous loci inconspicuous to subdenticulate, unthickened, not darkened. Conidia solitary, Illustration: Braun & Urtiaga (2008: 490, g. 4). obclavate-subcylindrical, straight to curved, 30–150 × 3–4 μm, 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 formation of super cial hyphae with solitary conidiophores truncate, 1.5–2 μm wide, hila unthickened, not darkened. (super cial hyphae emerging through stomata, branched, septate, 1.5–2.5(–3) μm wide, subhyaline to pale olivaceous, Holotype: India: West Bengal, Darjeeling, Sevak Road, on thin-walled, smooth; solitary conidiophores arising from Dicliptera chinensis [roxburghiana], 11 Nov. 1967, M. Mandal super cial hyphae, lateral, erect, straight, subcylindrical- (K(M) IMI 135116). Isotype: BPI 435712. conical to somewhat geniculate-sinuous, unbranched, 5–30 × 1.5–4 μm, 0–1(–2)-septate, subhyaline to pale olivaceous- Host range and distribution: On Dicliptera chinensis, brown, thin-walled, smooth; conidiogenous cells integrated, Acanthaceae, Asia (?China; India, West Bengal).
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Basionym (Fig. 30) Pseudocercospora ecbolii chinensis Note: RecordsofPseudocercosoraconsociata μm. A. Fig. 29.Pseudocercosporadiclipterae notes under Conidiophore fascicles. Deighton, Brit. Mycol.Soc.53:338(1969). , fromChinabelongpossiblyto : Cercosporaecbolii P. consiciata Trans. Brit. Mycol.Soc B. ). Conidiophores.
A.K. Kar&M.Mandal, (A.K. Kar&M.Mandal) (K(M) IMI135116, holotype). . 88:388(1987). C. P. diclipterae Conidia. Bar=10 onDicliptera Trans. (see Braun etal. Conidiophore fascicles. Fig. 30.Pseudocercosporaecbolii large fascicles, mostly2–15,arisingfromstromata, through olivaceous-brown. μ internal. Caespituli limited, yellowishdiscolorations,4–10mmdiam,scattered, Description Illustration Literature nally deepolivaceousbelow byabundantfructi m diam,subglobosetooblong, darkolivaceousto : Crous&Braun(2003:168),Kamal(2010:172). Stromata : Kar&Mandal(1969:339, : hypophyllous, effuse, olivaceous. Leaf spotsamphigenous,formedasvein- Conidiophores lacking orsmall,substomatal, 5–25 B. Conidiophores. (K(M) IMI135117, holotype).A. in smalltomoderately g. 2). C. Conidia.Bar=10μm. IMA FUNGUS Mycelium cation. Cercosporoid fungi 4
stomata, divergent to moderately dense, erect, straight, ARTICLE subcylindrical-conical to geniculate-sinuous, unbranched or occasionally 1–2 times branched, 15–65 × 3–5 μm, 0–6-septate, pale olivaceous to olivaceous-brown, thin- walled, smooth; conidiogenous cells integrated, terminal or conidiophores sometimes reduced to conidiogenous cells, 10–25 μm long, conidiogenous loci inconspicuous to subdenticulate, unthickened, not darkened. Conidia solitary, narrowly obclavate-cylindrical, straight to curved, 20–100 × 2.5–4 μm, 1–10-septate, pale olivaceous to very pale olivaceous-brown, thin-walled, smooth, apex subacute or subobtuse, base obconically truncate, 1–2 μm wide, hila 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: Tai (1948: 44, g. 11), Guo & Hsieh (1995: 4, g. 3), Guo et al. (1998: 13, g. 3).
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 inde nite. Caespituli hypophyllous, effuse, sooty. Mycelium internal and external, super cial; hyphae emerging through stomata, branched, septate, 2–3.5 μm wide, pale olivaceous 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 super cial hyphae, erect, subcylindrical, exuous, curved- Fig. 31. Pseudocercospora justiciae (based on Guo & Hsieh 1995: 4, sinuous to somewhat geniculate-sinuous, unbranched or g. 3). A. Conidiophore fascicle. B. Conidia. Bar = 10 μm. branched, 50–130 × 4–6.5 μm, 3–8-septate, olivaceous- 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 USA (Al eri et al. 1984) is unclear and doubtful. Material obclavate-subcylindrical, straight to curved, 40–110 × 3.5– could not be traced. 6.5 μm, 3–11-septate, pale olivaceous to olivaceous, thin- walled, smooth, apex obtuse to subacute, base obconically Pseudocercospora justiciicola P.N. Singh et al., truncate, about 2–2.5 μm wide, hila neither thickened nor Mycol. Res. 100: 1129 (1996). darkened. (Fig. 32)
Holotype: China: Sichuan: Chengtu, on Justicia procumbens, Illustration: Singh et al. (1996: 1129, gs 1–5). 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). internal and external; super cial hyphae branched, septate,
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super A. Holotype truncate, about1–2 smooth, apexacutetosubobtuse, baseshortobconically 35–120 ×1.5–5μm,4–14-septate,olivaceous,thin-walled, straight tocurved,occasionallywithshortlateralbranchlets, loci unthickened,notdarkened. conidiogenous cellsintegrated,terminal, μm, 2–6-septate,olivaceous-brown,thin-walled,smooth; somewhat geniculate-sinuous,unbranched,20–50×1–5 lateral, occasionallyterminal,erect,straightto arising fromstromataorsolitarysuper somewhat oblong,brown. developed, small,about10–25 subhyaline, 1–3μmdiam,thin-walled,smooth. Fig. 32. 366368). [simplex Conidiophore fascicle. cial hypha.
Pseudocercospora justiciicola ], : Isotype: GPU4004. Acanthaceae Nepal: Kathmanduvalley, on C. Conidiophore. μm wide,hilaunthickened,not darkened. B.Solitaryconidiophoresarisingfroma , Nov.–Dec. 1993, Kamal Conidiophores D. Conidia.Bar=10 Conidia μm diam,subgloboseto (K(M) IMI366368,holotype). solitary, obclavate, in loosefascicles, Justicia japonica al hyphae, μm. (K(M) IMI Stromata exuous, Braun etal. which is,however, agenuinespeciesof (Stevenson 1975,Minter previous falselyreferredtoas Note: The materialon collection. Host rangeanddistribution 16490 alopecuroidea Holotype truncate, 1–2 walled, smooth,apexobtusetosubacute,baseobconically × 2–4μ straight tocurved,narrowlyobclavate-cylindrical,20–120 nor darkened,occasionallysubdenticulate. cells, 5–25 integrated, terminalorconidiophoresreducedtoconidiogenous olivaceous-brown, thin-walled,smooth;conidiogenouscells only rarelybranched,5–80×2.5–5.5 straight to arising fromsuper aggregations, throughstomata,orconidiophoressolitary, fascicles, arisingfrominternalhyphaeorstromatichyphal aggregations, brown. Stromata pale olivaceousorolivaceous-brown,thin-walled,smooth. sparingly branched,1–3 external; super 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 materialofP. rungiae pertains to Notes: There isasecondcollectionfromNepalthatprobably collection. Host rangeanddistribution Literature (Fig. 34) Mycotaxon (2003). Braun &Crous, Cercospora (BPI437637). R. K.Verma (GPU,KK221,K(M)IMI303480)].See m, 1–9-septate,subhyalinetopaleolivaceous,thin- : : Chupp(1954: 25). lacking or only with very small substomatal hyphal lacking oronlywithverysmallsubstomatalhyphal : : Braun&Crous(2005:402, : Cercospora odontonematis P. justiciicola Puerto Rico:SanJuan,on μ μ amphigenous, 1–5 mm diam, Leaf spotsamphigenous,1–5mmdiam, m long, conidiogenous loci neither thickened m long,conidiogenouslocineitherthickened exuous, geniculate-sinuous, unbranched or exuous, geniculate-sinuous,unbranchedor , m wide,hilaunthickened,notdarkened. : 25(1954);as“ Acanthaceae 92: 399(2005). Justicia cial hyphae emerging through stomata, cial hyphaeemergingthroughstomata, P. rungiae cial hyphae, lateral or terminal, erect, cial hyphae,lateralorterminal,erect, Conidiophores Mycosphaerella and Anam. sp.),Nepal,Kathmandu . et al.2001,Crous&Braun2003), Lepidagathis alopecuroidea μ erma &Kamal(1991)citeditas . m wide, septate, subhyaline to m wide,septate,subhyalineto , 10Feb.1962, : Onlyknownfromthetype : Onlyknownfromthetype [on “Ruelliaprostrata odontonemae Cercospora lepidagathidis Caespituli Mycelium in small, usually loose in small,usuallyloose
μ Chupp, g. 5). m, 0–4-septate, pale m, 0–4-septate,pale U.Braun&Crous, Cercospora s.str. Conidia ”. internal and internaland (Chupp) U. Monograph of hypophyllous, hypophyllous, Lepidagathis IMA FUNGUS H. L.Rubin alley, Jan. ” (onthe solitary, : 296 was
Cercosporoid fungi 4 ARTICLE
Fig. 33. Pseudocercospora lepidagathidis (BPI 437637, holotype). A. Solitary conidiophores arising from super cial hyphae. B. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 μm. Fig. 34. Pseudocercospora odontonematis (CUP 40422, lectotype). A. Super cial hypha, B. Solitary conidiophores arising from super cial hyphae. C. Conidiophore fascicle. D. Conidiophores. E. Description: Leaf spots amphigenous, circular to somewhat Conidia. Bar = 10 μm. angular-irregular, 2–5 mm diam, brown, later with greyish brown to dingy grey centre and darker border. Caespituli hypophyllous, punctiform to effuse. Mycelium internal and external; super cial cells or integrated, terminal, 10–25 μm long, conidiogenous loci hyphae branched, septate, 1–3 μm wide, subhyaline or pale inconspicuous to subdenticulate, but always unthickened and olivaceous, thin-walled, smooth; stromata substomatal, 10–40 not darkened. Conidia solitary, narrowly cylindrical to obclavate- μm diam, brown. Conidiophores in small to moderately large cylindrical, 20–90 × 2–4 μm, (2–)3–8-septate, very pale fascicles, loose to dense, arising from stromata, through olivaceous, thin-walled, apex obtuse to subacute, base more or stomata or erumpent, also solitary, arising from super al hyphae, less truncate to distinctly obconically truncate, 1.5–2 μm wide, lateral, rarely terminal, erect, straight, subcylindrical-conical to hila unthickened, not darkened. strongly geniculate-sinuous, unbranched, 10–40 × 2–4.5 μm, 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,
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Basionym (Fig. 35) Pseudocercospora rhinacanthi Acanthaceae collection. Host rangeanddistribution Isolectotype shoulders. inconspicuous orvisibleastruncate,subdenticulatetips terminal, about10–25 conidiophores reducedtoconidiogenouscellsorintegrated, pale olivaceoustoolivaceous-brown,thin-walled,smooth; or occasionallybranched,5–60(–80)×2–5 subcylindrical tomoderatelygeniculate-sinuous,unbranched or solitary, arisingfromsuper arising fromstromata,throughstomata,erecttodecumbent, in loosetomoderatelydense,smallfascicles,usually2–6, 10–25 thin-walled, smooth.Stromata septate, 1–4 Mycelium on dryleaves. diam, brownorlaterwithdingygreycentre,ratherindistinct with yellowishhalotocircularorangular-irregular, 2–10mm yellowish tobrowndiscolorationsordiffuse brownspots Description Exsiccatae: Kabát&Bubák,FungiImp.Exs.847. et al.(1998:376),Crous&Braun(2003:351). Literature Rhinacanthus Indonesia Lectotype unthickened, notdarkened. base usuallylongobconicallytruncate,1–1.5 olivaceous, thin-walled,smooth,apexmoreorlesspointed, μm, indistinctly(2–)3–10(–12)-septate,subhyalinetopale straight tosomewhatcurved,(25–)40–120(–130)×2–5 Synonym Host rangeanddistribution Fungi Imp.Exs.847,e.g.,K(M)IMI89002(slide),HBG,W. Imp. Exs.847](BPI440801). two speciesaremorphologically barelydistinguishable. are unclear, unprovenandmightbelongto [Justicia nasuta procumbens Notes (Indonesia, Philippines, Thailand). Mycol. Pap.140:152(1976). Fung. 25:917(1931). 414 (1912);as Wiss Kaiserl. Akad. : Chineserecordsof μm diam,olivaceous-browntobrown. : : Chupp(1954:25),Guo&Hsieh(1995:357), : internal andexternal;super : : Cercosporina rhinacanthi (designated here,MycoBank,MBT202785): Conidia : CUP 60863. Java: Bogor(Buitenzorg),BotanicGarden,on Cercospora (T , 25Jul.1932, Leaf spots sp.,1907,F. v. Höhnel Caespituli μm wide,subhyalinetoolivaceous-brown, ai 1979,Guo&Hsieh1995, ], “rhynacanthi Rhinacanthus solitary, narrowlyobclavate-cylindrical, .,
amphigenous, mainlyhypophyllous. amphigenous, formedasdiffuse rhinacanthi Math.-Naturwiss. Cl μm long,conidiogenousloci : Onlyknownfromthetype : On Isolectotypes O. A. Plunkett (CUP 40422). O. A. cial hyphae,lateral,straight, ”. lacking orsmall,substomtal, P. rhinacanthi sp.), Rhinacanthus [Kabát&Bubák,Fungi cial hyphaebranched, (Höhn.) Sacc.,Syll. Höhn., (Höhn.) Deighton, Acanthaceae P. consociata : Kabát&Bubák, μm, 0–3-septate, Conidiophores on ., μm wide,hila Sitzungsber. etal.1998) Wien, (nasutus Justicia , Asia . The 121 Braun etal. :
yellowish. angular-irregular ordiffuse, 3–5mmdiam,greenish white, Description 444–445, Illustrations Literature Synonym Pseudocercospora rungiae A. Fig. 35. (Fig. 36) super Conidiophore fascicles. Indian Phytopathol. (HCIO 32003)]. Kolhapur, on 8: 69“1977”(1978)[ cial hypha. Pseudocercospora rhinacanthi : : Crous&Braun(2003:361),Kamal(2010:216). gs 4–5). Cercospora : : Patil(1978:70, Caespituli Leaf spots C. Conidiophores. Rungia repens
rungiae B. hypophyllous, scattered, rather 44: 446(1991). mainly hypophyllous,circular to Solitary conidiophoresarisingfroma holotype g. 1),
M.S. Patil,Botanique D. , 11 Dec.1975,M.S.Patil R.K. Conidia. Bar=10 : (BPI 440801,lectotype). erma &Kamal(1991: India erma &Kamal, : Maharashtra IMA FUNGUS (Nagpur) μm. : Cercosporoid fungi 4
truncate, 1–2 μm wide, hila unthickened, not darkened. 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. erma & Kamal (1991) cited a collection on “Ruellia prostrata” (on the original label as Justicia sp.) as paratype material (Nepal: Kathmandu alley, Jan. 1986, R. K. erma (GPU, KK 221, K(M) IMI 303480). This collection, characterised by forming super cial hyphae with solitary conidiophores is excluded from P. rungiae. The identi cation of the host is uncertain, and the identity 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: Boedijn (1961: tab. 108, g. 1), Braun & Sivapalan (1999: 20, g. 13).
Description: Leaf spots amphigenous, angular-irregular, often vein-limited, 1–10 mm diam, sometimes con uent and larger, dark brown, later greyish brown to dingy grey, margin inde nite. Caespituli hypophyllous, inconspicuous. Mycelium internal and external; super cial hyphae sparingly branched, 1–3 μm wide, 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 conidiophores arising from a super cial hypha. D. Conidia. Bar = 10 μm. swollen hyphal cells, 1.5–4 μm diam, olivaceous to brownish. Conidiophores solitary, arising from super cial hyphae, lateral, rarely terminal, occasionally emerging through stomata, inconspicuous. Mycelium internal; hyphae branched, septate, solitary or in small, loose fascicle, erect, straight, subcylindrical- colourless, 1–2 μm wide, somewhat wider near stromata. conical to somewhat geniculate-sinuous, unbranched, 5–30 × Stromata lacking or poorly developed, substomatal, small, 1.5–3 μm, 0–1-septate, subhyaline to pale olivaceous-brown, to about 25 μm diam. 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- 5–20 μm long, conidiogenous loci inconspicuous, neither sinuous, unbranched to branched, 10–75 × 2–5 μm, thickened nor darkened. Conidia solitary, narrowly obclavate- 0–3(–6)-septate, pale olivaceous to olivaceous-brown, thin- cylindrical, 20–95 × 1.5–4 μm, (0–)1–8-septate, subhyaline, walled, smooth; conidiophores occasionally reduced to pale yellowish green to very pale olivaceous, thin-walled, conidiogenous cells, but mostly integrated, terminal, 10–30 smooth, apex subacute, base obconically truncate, 1–2.5 μm μm long, conidiogenous loci inconspicuous to subdenticulate, wide, hila unthickened, not darkened. but always unbranched and not darkened. Conidia solitary, obclavate-subcylindrical, straight to curved, 30–145 × 2–4 Holotype: Indonesia: Jawa Barat: Bogor, botanical garden, μm, 1–10-septate, subhyaline to pale olivaceous, thin-walled, Thunbergia alata, Apr. 1950, K. B. Boedijn (L 53879). Isotype: smooth, apex obtuse or subacute, base short obconically K(M) IMI 91586.
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Basionym Pseudocercospora thunbergiicola Host rangeanddistribution hyphae. Conidiophores andhyphaeemergingthroughastoma. Fig. 37.Pseudocercosporathunbergiae Literature Conidiophores. Indonesia). JUDQGLÀRUD (Fig. 38) 30: 195(1965);as“thunbergicola on, Mycol.Pap.140:154(1976). C. Solitaryconidiophorearisingfromasuper : Yen &Lim(1980:188), Crous&Braun(2003:403). : ), Acanthaceae Cercospora thunbergiicola E. Conidia.Bar=10μm. , Asia (Brunei;India,UttarPradesh; : OnThunbergia ”. J.M. Yen, Rev. Mycol. (L 53879,holotype).
(J.M. Yen)Deight- (alata, erecta cial hypha. B. super Braun etal. cial A. D. super Conidiophore fascicles. Fig. 38. cylindrical, straight tocurved,about60–175 ×3.5–5 tips orshoulders. conidiogenous lociinconspicuous orvisibleastruncate conidiogenous cellsintegrated, terminal,10–30 somewhat palertowardsthe tip,thin-walled,smooth; μ unbranched oroccasionallyoncebranched,10–80×3–5 narrower towardsthetip,togeniculate-sinuous, hyphae, lateral,erect,straight,subcylindricalorsomewhat to moderatelydense,orsolitary, arisingfrom super hyphae orstromata,emergingthroughstomata,divergent moderately largefascicles,2–22,arisingfromsubstomatal substomatal, subglobose,brown. Stromata 2–3 hyphae emergingthroughstomata,sparinglybranched, conspicuous. margin darkbrown. angular-irregular, 0.5–3mmdiam,centrepale,whitish, Description g. 68), Illustrations m, 0–5-septate,paleolivaceous orolivaceous-brown, μ cial hyphae. m wide,paleolivaceous-brown,thin-walled,smooth. Pseudocercospora thunbergiicola almost lackingorsmall,10–30 : Yen (1965:197, : Mycelium Leaf spots C. Conidiophores. Conidia Caespituli B. internal andexternal;super Solitary conidiophoresarisingfrom solitary, cylindrical orobclavate- g. 12), Yen &Lim(1980:258, amphigenous, circularto D. Conidia.Bar=10 Conidiophores amphigenous, notvery (PC, holotype). IMA FUNGUS μ in smallto m diam, μ m long, μm. μ cial cial m, A. Cercosporoid fungi 4
4–11-septate, pale olivaceous-brown, thin-walled, smooth, ARTICLE apex subacute, base short to long obconically truncate, 2–2.5 μm wide, hila unthickened, not darkened.
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: Yen (1964: 233, g. 10), Yen & Lim (1980: 219, g. 22), Yen (1983: 362, g. A–B).
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, straight to curved or somewhat exuous, long, liform, not or barely geniculate, unbranched, 140–260(–300) × 5–7 μm, 3–15-septate throughout, medium brown, paler towards the tip, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal, about 10–30 μm long, conidiogenous loci subconspicuous, 1.5–2 μm diam, in front view visible 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 or obovoid, usually straight, 30–45 × 5.5–8.5 μm, pale olivaceous to olivaceous-brown, thin-walled, smooth, apex broadly rounded, base short obconically truncate, 1.5–2 μm 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: The generic af nity of 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. Bar = 10 μm.
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(Fig. 41) Note: Belongingtothe collection. Host rangeanddistribution 50000). (Fig. 40) Hsieh (1995:6, (1987: 356, g.2),Hsieh&Goh(1990:14, Illustrations 13), Guo&Hsieh(1995:4–5), Guo Literature Pseudocercospora actinidiaeDeighton, on Conidiabroadlyobclavate-subcylindrical, 25–85(–100)×5–8.5 1 Key toPseudocercosporaspecieson Actinidiaceae Pseudocercospora specieson Actinidiaceae Illustrations Literature Cercospora actinidiae A singlespecies Cercospora specieson Actinidiaceae Actinidiaceae Actinidiaceae Holotype thickened anddarkened. base truncateorslightlyattenuated,1–2.5 pluriseptate, hyaline,thin-walled,smooth,apexpointed, curved, sometimessigmoid,40–160(–220)×2–3(–4) 2.2–2.6 intercalary, conidiogenouslocithickenedanddarkened, smooth; conidiogenouscellsintegrated,terminaland 5.5) geniculate, usuallyunbranched,about60–280×4–4.5(– hyphae orswollenhyphalcells,erect,straight,curvedto or insmallfascicles,2–8,divergent,arisingfrominternal a fewswollenhyphalcells,brown. Mycelium with narrowdarkbrownborder. 2–8 mmdiam,centregreyishbrownoryellowishbrown, Description 23, g.6). Conidianarrowlyobclavate- 140 Mycol. Sin.Suppl μ m, pluriseptate,mediumbrowntobrown,thin-walled, : 10(1976). : μ : Liu&Guo(1987:355),Hsieh &Goh(1990:12– : Guoetal.(2005:23). inernal.Stromata m diam.Conidia : Deighton(1976:11–12, : Liu&Guo(1987:354, : China Leaf spotsamphigenous,subcirculartoirregular, , 11 Sep.1961,X.J.Liu Actinidia g. 5),Guo : Jiangsu . 1:353“1986”(1987). C. apiis.lat. spp...... et al.(1998:15–16, : Nanjing,on X.J. Liu& Y.L. Guo,Acta lacking orsmall,composedof : Onlyknownfromthetype solitary, acicular, straightto Caespituli liform, 15–110 ×1–4 et al.(1998:15–16). complex. g. 1),Guo & Conidiophores gs 1–2),Liu&Guo Q. M.Ma249 amphigenous. μ g. 3),Guo& Actinidia Mycol. Pap. m wide,hila g. 5). et al.(2005: (HMAS solitary Braun etal. μ sp., ie ...... μm, base1–2mwide ...... m, branched, 1–3.5 μmwide,sometimes anastomosing, external; super Description Exsiccatae: ReliquiaeFarlowianae829. effuse, deepolivaceous, amphigenous, mainlyhypophyllous, pale browntogreyishbrown, margininde and larger, browntodarkbrown,laterwith paler centre, angular-irregular, 1–10mmdiam,sometimescon Conidiophore fascicle. Fig. 40. μm, base2–2.5 Cercospora actinidiae : Leaf spots cial hyphaeemergingthrough stomata, B. μm wide; Conidiophore tip. at (based onGuoetal.2005:23, occose. rst lacking,latersubcircularto Mycelium C. nely punctiformto ...... Conidia. Bar=10 P. hangzhouensis nite. P. actinidiae IMA FUNGUS internal and Caespituli A. g. 6).A. uent μ m. m. Cercosporoid fungi 4 ARTICLE
Fig. 41. Pseudocercospora actinidiae (FH, holotype). A. Stroma. B. Conidiophore fascicles. C. Solitary conidiophores arising from super cial hyphae. D. Conidiophores. E. Conidia. Bar = 10 μm.
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). leaf surface, 10–50 μm diam, substomatal, subglobose, 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, arising from super cial hyphae, lateral, occasionally terminal, Actinidia sp.), Actinidiaceae, Asia (China, Anhui, Fujian, differentiation between hypophyllously formed erect to Guizhou; Korea, Japan, Taiwan). decumbent branched conidiophores and super cial hyphae dif cult, fertile threads to 700 μm long, simple or branched, Pseudocercospora hangzhouensis X.J. Liu & Y.L. individual solitary conidiophores formed as lateral branchets Guo, Acta Mycol. Sin. Suppl. 1: 357 “1986” (1987). 2–45 μm long, 3–6.5 μm wide, aseptate to pluriseptate, pale (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: to subdenticulate, 1.5–2 μm diam, unthickened, not darkened. 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 or somewhat sigmoid, 25–85(–100) × 5–8.5 μm, 3–9-septate, 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: μm wide, hila unthickened, not darkened. 5–6), Guo et al. (1998: 16–17).
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A singlespecies ...... Viburnum in small,loose fascicles,2–13,arisingfrom internalhyphae μm diam,wallslightlythickened. μm diam,substomatalorintraepidermal, brown,cells2–8 aggregations ofafewswollen hyphalcells,globose,10–25 amphigenous. dark margin,sometimeswithyellowishbrownhalo. brown, yellowishbrownbelow, latercentregreyishwhitewith irregular andvein-limited,1–18mmdiam,browntodark Description (1998: 17, (1990: 15, g.4),Guo&Hsieh(1995:7,6),etal. Illustrations Literature Basionym (Fig. 43) MycoBank MB814566 Cercospora sambucicola Cercospora specieson Adoxaceae A singlespecies ...... Sambucus Tabular keytoCercosporaspecieson Adoxaceae accordingtohostgenera Cercospora Adoxaceae 2–11-septate, hyalinetopaleolivaceous,thin-walled,smooth, Conidia cells, 2–20 integrated, terminalorconidiophoresreducedtoconidiogenous olivaceous-brown, thin-walled,smooth;conidiogenouscells 2–40 ×1–3.5(–4.5) curved, subcylindricaltogeniculate-sinuous,unbranched, or solitary, arisingfromsuper fascicles, arisingfromstromata,throughstomataorerumpent, to oblong,mainlyepiphyllous,brown. thin-walled, smooth. hyphae branched,1.5–3 Illustrations Mycelium to palegreyishbrownbelow. by agreyishblackmarginalline,paleyellowishbrown greyish whitecentreontheupperleafsurface,surrounded 2–8 mmdiam,paleyellowishbrowntobrown,laterwith Description 56, g.32). sambuci Mycotaxon solitary, narrowlyobclavate- : Guoetal.(2005:55–56). : internal. g. 6). : : : Guo&Jiang(2000:263, : Liu&Guo(1987:357, Cercospora sambuciY.L. Guo& Y. Jiang, Leaf spotsamphigenous,subcirculartoangular- Leaf spots F. Stevens&C.J.King,1927. μ m long, conidiogenous loci inconspicuous. m long,conidiogenouslociinconspicuous. 74: 262(2000), Mycelium Stromata Stromata μ m, 0–3-septate, subhyaline to pale m, 0–3-septate,subhyalinetopale amphigenous, circularorsubcircular, μ internal andexternal;super m wide, subhyaline to olivaceous, m wide,subhyalinetoolivaceous, lacking orsmall,onlyformed as Y.L. Guo,nom.nov. cial hyphae, erect, straight to cial hyphae,erect,straightto 10–70 nom. illeg. Caespituli Conidiophores liform, 15–110 ×1–4 Conidiophores g. 3),Guo μ g. 3),Hsieh&Goh m diam, subglobose m diam,subglobose (Art.53.1),nonC. amphigenous. et al.(2005: solitary or Caespituli Caespituli in dense in dense Braun etal. μ cial cial m, m, Clethra and Guoetal.(1998)changedtheidenti cationofhostto Japan, Korea, Taiwan). Actinidia Host rangeanddistribution 28 Sep.1961, actinidioides from Chinaon“ small fascicles, to140 barely distinguishablefrom second cercosporoidhyphomycete whichismorphologically complex. Type material ofPassaloracatenospora Notes: This speciesbelongstothe Jilin, hejiang),?North America (USA, Alabama). javanica Host rangeanddistribution williamsii Holotype wide, hilathickenedanddarkened. smooth, apexpointedorsubobtuse,basetruncate,2–3.5 37.5–175 ×3–5 acicular, straight to somewhatcurvedorslightlysigmoid, thickened anddarkened,2–3.5(–4) cells integrated,terminalandintercalary, conidiogenousloci narrower towardsthetip,thin-walled,smooth;conidiogenous at thebase,1–6(–13)-septate,mediumbrown,palerand width irregular, 42.5–140(–400)×3.5–7 straight, subcylindricaltogeniculate-sinuous,unbranched, or hyphalaggregations,throughstomataerumpent,erect, Holotype 1–2 apex subacute,basesubtruncatetoshortobconicallytruncate, Notes: Pseudocercospora clematoclethrae species NQRZQ LQVXI¿FLHQWO\ DQG H[FOXGHG 'RXEWIXO Guo, Mycosystema5:100(1992). μ m wide,hilaunthickened,notdarkened. sp.( Guo &Liu(1992)described , , Adoxaceae : : sp.), williamsii China China Clethraceae ] ( X.-J. Liu&Q.-M.Ma671 Actinidiaceae Actinidiaceae : Clematoclethra fabri μm, 5-topluriseptate,hyaline,thin-walled, : Zhejiang ...... [buergeriana Jilin Province , Sep.1962, μm longand4–7 μmwide;conidia ). : C. sambucicola : OnActinidia ), butlaterGuo&Hsieh(1995) , Asia (China, Anhui, , Asia Hangzhou, on : On ]), J. K.Bai : Yongji,on Sambucus μm diam.Conidia Adoxaceae ” [C.scandenssubsp. Cercospora apiis.lat (HMAS50001).
P. clematoclethrae μm, to8 (arguta, X.J. Liu& Y.L. (HMAS77346). (conidiophoresin C. sambucicola Actinidia arguta C.viburnicola (?canadensis , Asia (China, , Asia IMA FUNGUS Sambucus contains a chinensis hejiang; μm wide solitary, μm . , , ,
Cercosporoid fungi 4 ARTICLE
Fig. 42. Pseudocercospora hangzhouensis (based on Guo & Hsieh 1995: 7, g. 6). A. Conidiophore fascicle. B. Solitary conidiophores arising from super cial hyphae. C. Conidia. Bar = 10 μm. U. Braun del.
Fig. 43. Cercospora sambucicola (HMAS 77346, holotype). A. acicular, hyaline, 3–4.5 μm wide). The conspeci ty of Chinese Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 collections and material from the USA remains to be proven μm. via phylogenetic analyses.
Cercospora viburnicola W.W. Ray, Mycologia 33: erumpent, erect, straight, subcylindrical to geniculate- 174 (1941). sinuous, unbranched, 25–175 × 4–6 μm, septate throughout, (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). terminal and intercalary, about 15–35 μm long, conidiogenous loci conspicuous, thickened and darkened, 2–3 μm diam. Illustration: Guo et al. (2005: 57, g. 33). Conidia solitary, acicular, shorter conidia narrowly obclavate, straight to curved, 20–90 × 2–4 μm, pluriseptate, hyaline, 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 μm wide, hila thickened and darkened. amphigenous, ne, dark. Mycelium internal. Stromata lacking or small, about 10–25 μm diam, brown. 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).
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and misidenti Burek &Mu of 442266, CUP-C.H.OK-0032,ILL 29523,NY 945736). A record tomentosum collected atthetypelocality (21 Aug. 1942,on on aconfusionwith on of the Notes: Oklahoma, Wisconsin). ?Asia (China,Jiangsu,Shaanxi,Sichuan),USA (Florida, [tomentosum odoratissimum Host rangeanddistribution Conidiophore fascicle. Fig. 44.Cercosporaviburnicola C. viburnicola Viburnum cylindricum Cercosporaviburnicola Cercospora apiis.lat ) isdepositedinseveralherbaria enko 2014) is undoubtedly based on confusion enko 2014)isundoubtedlybased onconfusion ], cation. The described and illustrated obclavate cation. The describedandillustratedobclavate , opulus fromPolandonViburnum opulus suspensum C. viburnicola B. Conidia.Bar=10 from China are incorrect and based fromChinaareincorrectandbased , plicatum . complex. Reords of this species . complex.Reordsofthisspecies , Viburnum W.W. Rayismorphologicallypart : OnViburnum (CUP 29236,holotype). F.L. Tai. material Authentic μm. var. tomentosum spp.), Adoxaceae
(BPI 442264 ( widerska- Viburnum Viburnum (carlesii Braun etal. A. – , , classi Americal samples,theChinesecollectionsarecurrentlybetter μ about 450μ The conidiophoresareformedinsmall,divergentfascicles,to specimens areprobablynotconspeci to asC.viburnicola, & Jiang2000:264;Guo Viburnum opulus to concerned hasbeenre-examinedandturnedouttobelong 5.5) conidia withobconicallytruncatebase,30–90(–205)×3–5(– recorded anddescribed fromBulgariaon and belongstoCercosporella and conidiaaswellcercosporelloid conidiogenousloci Notes: California, Minnesota,Missouri, NewMexico,Oregon). melanocarpa Host rangeanddistribution BPI 439986,NY 234138. cerulea adoxae of fructi material havebeenexamined,butallwerewithoutanytrace described (Chupp1954).Numerousduplicatesoftheoriginal Notes: Although listedinliterature,thisspecieswasnever Adoxaceae Host rangeanddistribution Gall. Exs.1871(e.g.,B,HBG,PC). San Bernadino Lectotype Exsiccatae: Calif.Fungi403. Illustration (1995: 80). Literature Type Literature Cercospora adoxae species NQRZQ LQVXI¿FLHQWO\ DQG H[FOXGHG 'RXEWIXO Synonym SUROL¿FDQV &HUFRVSRUD m long.Withoutanymoleculardatabasedon Asian andNorth Pseudocercospora opuli Syll. Fung.15:84(1901). XIX, no.1871, Toulouse 1882, Hist. IowaStateUniv μ : m, arenotconsistentwith and also known from Poland. Chinese collections on andalsoknownfromPoland.Chinesecollectionson France ed asCercosporaapiis.lat cation. The name cation. The &HUFRVSRUDSUROL¿FDQV , (Rabenh.) P. Karst. Aug. 1893,
: : Saccardo(1895:606),Chupp(1954:104),Braun : Chupp(1954:98),Crous&Braun(2003:46). buddleifolium : Braun(1995:79, (designated byBraun1995:80): &HUFRVSRUHOODSUROL¿FDQV , Europe(France). m long,andtheconidiaareacicular, about35–385 ), : on Adoxaceae alley, SanBernadinoCounty, on Adoxa moschatellina
Parish represent typical Roum., andViburnum et al.2005:56–57, C. adoxae
Ellis &Holw., . II, 2735 , North America (USA, Alaska, : OnSambucus which iscommonon (Braun1995). This specieswas g. 75). : OnAdoxamoschatellina has colourlessconidiophores Fungi Sel.Gall.Exs 3: 42(1896). (NY 234139). C. viburnicola . mightreferto sp. (Guo 1996: 92; Guo sp.(Guo1996:92;Guo c with the latter species. c withthelatterspecies. nom. nud.(Art.38.1). ., but these C. apiis.lat.,butthese (Ellis &Holw.) Sacc., , Roum., FungiSel. Sambucus ebulus Bull. Lab.Nat. (cerulea USA: . The collection collection . The g. 33), referred g. 33),referred Isolectotypes: IMA FUNGUS Sambucus Ramularia California Viburnum Viburnum , ., glauca Cent . , , :
Cercosporoid fungi 4
(Bakalova & Borisova 2010: 49–50). This record is, however, 2–4 μm wide, which is in contradiction with the much broader ARTICLE doubtful and probably wrong. Conidia were described to be conidia (3–8 μm wide) of true &SUROL¿FDQV.
Passalora
Key to Passalora species on Adoxaceae 1 Colonies in pale reddish brown to fuliginous patches; mycelium internal and external, super cial; conidiophores fasciculate as well as solitary, arising from super cial hyphae; conidia subhyaline to reddish brown; on Sambucus spp...... P. lateritia Colonies (caespituli) without reddish tinge; mycelium internal, super cial hyphae lacking in vivo; conidia not reddish brown ...... 2
2 (1) Conidiophores short, 5–25 × 2–4 μm, 0–1-septate; conidia 8–45 × 1.5–4 μm, 0–4-septate; on Viburnum spp...... P. viburni Conidiophores much larger, 15–80 × 4–6 μm, 0–4-septate; conidia much longer and somewhat broader, 20–100(–125) × 3–5.5 μm, 1–8-septate; on Sambucus 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: Ellis (1976: 318, g. 240 A).
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– 25 μm diam, substomatal, yellowish to ochraceous-brown. 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, somewhat curved-sinuous, unbranched, 15–80 × 4–6 μm, rarely longer, usually 0–4-septate, subhyaline, pale ochraceous to golden brown or olivaceous-brown, thin- walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–30 μm long, conidiogenous loci conspicuous, thickened and darkened, (1–)1.5–2 μm wide. Conidia solitary to catenate, in simple or branched chains, cylindrical, subcylindrical, occasionally almost obclavate or broadly fusiform, straight to curved or slightly sinuous, 20–100(–125) × 3–5.5 μm, 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 two hila), base short obconically truncate, (1–)1.5–2 μm wide, hila somewhat thickened and darkened. Fig. 45. Passalora catenospora (CUP-A-002255a, lectotype). A. Conidiophore fascicles. B. Conidiophore. C. Conidia. Bar = 10 μm.
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diam, aswellhyaline,acicularconidiaformedsingly. darker conidiophoreswithlargerconidiogenousloci,2–3 is readilydistinguishablefrom from theChinese s. lat.complexandismorphologicallybarelydistinguishable confusion. The secondfungusbelongstothe a secondcercosporoidhyphomycetewhichmightcause in Chupp(1954). Type materialof was erroneouslyreducedtosynonymywith Pseudocercospora ebulicola javanica 3(AL)). Recordsofthisspeciesfrom Taiwan on Other syntypesarefrom August 1891(CUP-A-002245#1– Note: The lectotypeisacollectionfromOctober1891. Oklahoma, Pennsylvania, Texas), West Indies(Haiti). USA, Alabama, Florida,Kansas,Mississippi,NorthCarolina, America (DominicanRepublic),North America (Canada; B. Fig. 46. canadensis Host rangeanddistribution arising fromsuper Conidiophore fascicles. Passalora lateritia [formosana , intermedia cial hyphae. Cercospora sambucicola ] (Tai 1979)areincorrectandreferto (NY 838232,lectotype). , Sambucus C. Conidiophore. E. Conidia. Bar=10 μ (Cercospora ebulicola : onSambucus P. catenospora spp.), C. catenospora D. Adoxaceae Solitary conidiophores Solitary conidiophores A. Cercospora apii C. catenospora Super . This species . This m. byitsmuch Sambucus contains cial hypha. cial hypha. ( , Central , Central cerulea, cerulea, ) which Braun etal. μm
simple chains, cylindrical or subcylindrical or somewhat simple chains,cylindricalorsubcylindricalsomewhat about 1–2 conidiogenous lociconspicuous,thickenedanddarkened, conidiophores reducedtoconidiogenouscells,5–30 walled, smooth;conidiogenouscellsintegrated,terminalor 3–8 arising fromsuper hyphae orstromata,throughstomataconidiophoressolitary, divergent tomoderatelydense,arisingfromsubstomatal brown. swollen hyphalcells,substomatal,10–50 Stromata concolorous withconidiophoresorpale,thin-walled,smooth. unbranched orsparinglybranched,1–7 stomata, occasionallyclimbingleafhairs,rarelyanastomosing, super to ferruginouspatches. hypophyllous, effuse, formingreddish,palereddishbrown Description Swingle, KansasFungi6. Exsiccatae: Ellis&Everh.,N. Amer. Fungi1994.Kellerman& Literature Basionym Passalora lateritia fascicle. B.Conidiophores. Fig. 47.Passaloraviburni (Fig. 46) (1888). Mycosphaerella and Anam. μ m, 0–4-septate,subhyalinetopalereddishbrown,thin- cial hyphae lacking or present, emerging through cial hyphaelackingorpresent,emergingthrough Conidiophores : Saccardo(1892:646),Chupp(1954:102). : Cercosporalateritia lacking or formed as stromatic aggregations of lacking orformedasstromaticaggregationsof : μ m diam. Leaf spotslackingorinde cial hyphae, lateral or terminal, 5–75 × cial hyphae,lateralorterminal,5–75× Conidia
(Ellis &Halst.)U.BraunCrous, in small to moderately large fascicles, in smalltomoderatelylargefascicles, C. (NY 830558,lectotype).A. Mycelium Conidia. Bar=10 solitary to catenate, usually in solitary tocatenate,usuallyin
Ellis &Halst., : 244(2003). internal and external; internal andexternal; μ μ μm. m wide, septate, m wide,septate, m diam, reddish m diam,reddish nite. J. Mycol. IMA FUNGUS Conidiophore Colonies Colonies μ m long, m long, 4: 7 Cercosporoid fungi 4
obclavate-cylindrical, straight to slightly curved, 15–70(–80) × Description: Leaf spots subcircular to angular-irregular, ARTICLE 3–7 μm, (0–)1–6-septate, occasionally constricted at the septa, 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– μm wide, hila somewhat thickened and darkened. 50 μm diam, occasionally con uent and larger, pale yellowish brown to medium brown, composed of swollen hyphal cells, Lectotype (designated by Crous & Braun 2003: 244): USA: 2–6 μm diam. Conidiophores 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 sinuous, unbranched, 5–25 × 2–4 μm, 0–1-septate, hyaline, 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, μm long, conidiogenous loci conspicuous, minute, 1–1.5 μm 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 super cial hyphae, thickened, darkened conidiogenous loci × 1.5–4 μm, 0–4-septate, hyaline to pale yellowish or greenish, 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 short obconically truncate when in chains, about 1 μm wide, hila 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: The generic af nity of this species, which is intermediate between Passalora and Ramularia, is not quite clear. The general Illustrations: Braun (1993: 292, g. 21), Braun (1998: 388, habit of this species is rarther ramularioid, but the stromata and g. 647). conidiophores are pigmented. Results of molecular sequence analyses would be helpful to elucidate the true generic af nity.
Pseudocercospora
Key to Pseudocercospora species on Adoxaceae 1 Stromata lacking; super cial hyphae developed, but without solitary conidiophores; conidiophores fasciculate, 50–190 μm long, pluriseptate; on Sambucus javanica, Asia ...... P. ebulicola Stromata developed, 10–100 μm diam; super cial hyphae lacking or, if present, with conidiophores which are much shorter, 5–35 μm, and only 0–1-septate ...... 2
2 (1) Super cial hyphae with solitary conidiophores developed, 5–35 μm long, 0–1-septate, geniculate, i.e. proliferation sympodial; on Viburnum spp...... P. tinea Super cial hyphae with solitary conidiophores lacking; conidiophores longer, 10–200 μm, aseptate to pluriseptate (if short and mostly aseptate, then non-geniculate, proliferation percurrent, with annellations ...... 3
3 (2) Conidiophores subcylindrical, non-geniculate, proliferation percurrent, with ne annellations, rather short, 5–30(–50) × 2–5(–6) μm, 0(–2)-septate; conidia cylindrical- liform or occasionally somewhat obclavate; on Viburnum spp...... P. viburnigena Conidiophores at least partly geniculate-sinuous, proliferation sympodial, without annellations or occasionally with a single enteroblastic rejuvenation, much longer, 10–200(–300) μm, 0–3-septate or pluriseptate throughout; conidia obclavate-cylindrical to somewhat fusiform ...... 4
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Conidia wider, 3–6.5 Conidiaobclavate-cylindrical,25–85 11Conidia 2.5–4 (10) on orevendarkened-refractive;conidiophoresfascicluate;on Conidiogenouslociconsistentlyinconspicuous;conidiophoresfasciculatetocoremioid; Conidiogenouslociinconspicuoustosubconspicuous,i.e.unthickenedbutsomewhat refractive 9 (6) 0() Conidiacylindrical- 10 (9) Asiancollectionson North American collections=var. orevendarkened-refractive;on Conidiogenouslociinconspicuous(conidiophoresusuallyinsmallfascicles,2–15,divergent, Conidiogenouslociinconspicuoustosubconspicuous,i.e.unthickenedbutsomewhatrefractive 8 (7) Conidiaformedsingly...... ( butsomewhatrefractiveorevenslightlydarkened-refractive;on ordisarticulating,conidiogenouslociinconspicuoustosubconspicuous,i.e.unthickened, Conidiahyalineorsubhyaline, 7 (6) 0–3-septateor pluriseptatethroughout;conidia obclavate-cylindricaltosomewhat fusiform ...... oronlywitha singleenteroblasticrejuvenation, muchlonger, 10–200(–300) Conidiophoresatleast partlygeniculate-sinuous,proliferationsympodial,withoutannellations Conidiophoreslonger, about50–155 orsubhyalineand/orthin-walled,conidiaatleastparlycatenatedisarticulating ...... Conidiophoresrelativelyshort,usually10–80 6 (5) Conidianarrower, 2–5 oroccasionallysomewhatobclavate ...... rathershort, 5–30(–50) ×2–5(–6) Conidiophores subcylindrical,non-geniculate,proliferationpercurrent,with 2 (1) usuallypaletomediumolivaceousorolivaceous-brown,wallthinsomewhatthickened, to 0.8 Conidiabroadlyobclavateorobclavate-cylindrical,25–125×4.5–8 Conidialonger, 25–150 5 (4) aseptatetopluriseptate(ifshortandmostlyaseptate,thennon-geniculate,proliferationpercurrent, with annellations ...... geniculate,i.e.proliferationsympodial ...... Super 1 Super Viburnum unbranched; Stromatadeveloped,10–100 on on Stromatalacking;super 1 Sambucus Tabular keytoPseudocercosporaspecieson Adoxaceae accordingtohostgenera rejuvenationleavingacoarseannellation;on wallsthickened,to1μm,mediumdarkbrown,occasionallywithasingleenteroblastic Conidiashort,15–40×3–7 4 (3) Viburnum cial hyphaewithsolitaryconidiophoreslacking; longer, 10–200 cial hyphaewithsolitaryconidiophoresdeveloped,5–35 Viburnum Sambucus javanica μm; onViburnum μm wide,2–5-septate;on sect. spp. 10 ...... liform, 50–130×2.5–4 Sambucus Opulus μ m, 3–7-septate;on μm, ifwider[2–6.5(–7) cial hyphaedeveloped;conidiophoreslooselyfasciculate,simpletobranched; ,11(1)spae nohrhss ...... 5 μm, 1–12(–15)-septate;onotherhosts ...... Viburnum sargentii , Asia , Asia ...... μm, 1–5-septate,pigmented;conidiophoresindivergentfascicles, spp.,North America ...... ), mainlyV. opuluss.lat.(includingsubsp. μm diam;super nally sometimepaleolivaceous,solitaryorinshortchains spp.,northernhemisphere ...... Sambucus μm long,2–7-septate ...... 9 μm, average>60m ...... varia; conidiophoresinlarger, sensefascicles,to50perfascicle, μm, 0(–2)-septate;conidiacylindrical- Viburnum cylindricum Viburnum μm, 3–12-septate;on =var. cial hyphaelacking;conidiophoresmostlyindenserfascicles, ...... 7 μm, average<60m ...... μm] conidiaconsistentlypale,usuallyhyaline spp...... Braun etal. Viburnum nudum viburni-sargentii sp.,India ...... 11 , China ...... Sambucus μm long,0–1-septate, Viburnum erosum , North America μm, 2–10-septate, ); onViburnum Viburnum calvescens spp...... ne annellations, liform spp. μm, and ...... spp...... μm, trilobum ...... 8 ...... 2 ...... ) ...... P. viburni-cylindrici P. caprifoliacearum 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
3 (2) Conidia broadly obclavate or obclavate-cylindrical, 25–125 × 4.5–8 μm, 2–10-septate, ARTICLE usually pale to medium olivaceous or olivaceous-brown, wall thin to somewhat thickened, to 0.8 μm; on Vibrunum spp., North America ...... P. viburnicola Conidia narrower, 2–5 μm, if wider [2–6(–7) μm] conidia consistently pale, usually hyaline or subhyaline and thin-walled ...... 4
4 (3) Conidiophores relatively short, usually 10–80 μm, average < 60 μm ...... 5 Conidiophores longer, about 50–155 μm, average > 60 μm ...... 7
5 (4) Conidia short, 15–40 × 3–7 μm, 1–4(–5)-septate, pigmented; conidiophores in divergent fascicles, walls thickened, to 1 μm, medium to dark brown, occasionally with a single enteroblastic rejuvenation leaving a coarse annellation; on Viburnum nudum, North America ...... P. viburni-nudi Conidia longer, 30–150 μm, 1–12(–15)-septate; on other hosts ...... 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); on Viburnum spp...... P. varia
7 (4) Conidia cylindrical- liform, 50–130 × 2.5–4 μm, 3–12-septate; on Viburnum erosum ...... P. viburni-erosi Conidia obclavate-cylindrical, 25–85 μm long, 2–7-septate ...... 8
8 (7) Conidia 2.5–4 μm wide, 2–5-septate; on Viburnum cylindricum, China ...... P. viburni-cylindrici Conidia wider, 3–6.5 μm, 3–7-septate; on Viburnum sp., India ...... P. caprifoliacearum
Pseudocercospora caprifoliacearum (C. Gupta et to 6 μm wide, pluriseptate throughout, olivaceous-brown, wall al.) Kamal, Cercosporoid Fungi of India: 159 (2010). thin-walled to somewhat thickened, to 1 μm, at the very base (Fig. 48) occasionally to 1.3 μm, smooth; conidiogenous cells integrated, 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. curved, about 30–85 × 3–6.5 μm, 1–7-septate, subhyaline to 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 apex obtuse, base obconically truncate, 1.5–2.5 μm wide, hila (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. (1987: 16, g. 2), Gupta & Kamal GPU, KC 62. (1987: 33, g. 4). 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 greyish brown to dingy grey, margin inde nite or 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 nely punctiform, brown to blackish. Mycelium immersed, barely distinguishable and undoubtedly conspeci c. The two composed of subhyaline to pale brown, septate, thin-walled, species have been simultaneously published in a single smooth hyphae. Stromata about 10–50 μm diam, dark brown, book. It is proposed to give precedence to Phaeoisariopsis substomatal, cells 2–6 μm diam. 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 looser towards the apex, straight to exuous, unbranched or only from China. The two species are morphologically very similar. rarely branched, subcylindrical- liform, width uniform throughout 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 about 60–155 μm long and 2.5–4.5 μm wide, apex sometime by Crous & Braun (2003) and Kamal (2010). Differences
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Bar =10μm. holotype). Fig. 48.Pseudocercosporacaprifoliacearum Basionym (Fig. 49) MycoBank MB814570 Pseudocercospora depazeoides(Desm.)U.Braun& molecular datawillbeavailable. as separatespecies,atleastfortheinterimuntilculturesand prefer tomaintainP. caprifoliacearum(inclusing caprifoliacearum dense orevencoremioidconidiophorefascicles,similarto of be variationwithinasinglespecies. The Chinesetypematerial between ChineseandIndiancollectionswereconsideredto Cercospora penicillata Synonyms P. viburni-cylindrici Crous, comb.nov. Rabenh., Herb. Italy [lectotype Viv. Mycol.,no.587(1857),withdescription onlabel Bot., : sér. 3,11 A. : : Passalorapenicillata ercellis, on Exosporium depazeoides Conidiophore fascicle.B. (designated here,MycoBank,MBT202789): , butnarrowerconidia,2–4.5 : 364(1849). ischaracterisedbyhavingloosetooften iv. Mycol.587(HAL)]. (Ces.) Fresen., Sambucus nigra Conidiophore tips. Ces., inRabenh., Beitr. Mykol .: 93(1863). Desm., , 1856, (K(M) IMI294111, Ann. Sci.Nat., μm wide.We V. deCesati, P. C. Conidia. khasiana Herb. Braun etal. P. P. )
Cercospora sambucina Cercospora depazeoides Cercospora ticinensis Cercospora depazeoides Cercospora depazeoides Cercospora sambuci Literature Pseudocercospora sambucigena Phaeoramularia penicillata Cercospora depazeoides 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, Sch Maurilaun, Fl.Exs. 1749. Kabát&Bubák,FungiImperf.Exs.97.Kernervon Everh., FungiColumb.599.Ellis&North Amer. Fungi Ess. 336.Cavara,FungiLongobard.Exs.245,984.Ellis & Exsiccatae: Briosi&Cav., FunghiParas.PianteColt. Util. 4), Crousetal.(2013:106, Crous (2005:402, g.6),Bakalova&Borisova(2010:136,pl. Util. Ess.336, 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), en. 280.D.Sacc., Mycoth.Ital.199,1587. S MBT202790): 17: 1166 (1883)[lecotype( 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 (designated here,MycoBank,MBT202792): Paras. Sacc., CUP 41160, NY 2343037]. 1883, nigra [Gagry], upperwaterfall Rast Pereira (Traz-os-Montes), onSambucusnigra,Dec.1902,A. Monogr 21 Aug. 1921, 21 Aug. Pennsylvania Schub., Phytopathol. Sin Stevens on Sambucus canadensis . , 9 Aug. 1912, : Saccardo(1886:468 W. A. Kellerman401 6: 108(1912)[ Syll. Fung.4:469(1886). ]. Piante Colt.Util.Ess . : Briosi&Cavara,FunghiParas.PianteColt. (ILL 15175); 11 Mycotaxon gs 1–3,Ellis(1976:246, : 59(1927)[ : Briosi&Cavara,FunghiParas.PianteColt. : DauphonCounty, onSambucuspubens et al.(2005:279–280),Bakalova&Borisova USA: O. E.Jennings, Acc. 6736(NY 142641)]. . 1:59(1903)[ Austr.-Hung. 788.Krieger, FungiSaxon. . 12:13(1982). et al.(2013:105–106). A. Elenkin(LE158291)]. Cavara, inBriosi& F. Stevens&King, Ohio, on isotypes
Ellis &Kellerm., 92: 400(2005)[ var. (Desm.) Sacc., var. var. type: g. 60). (Ces.) X.J.Liu& Y.L. Guo, holotype hoekvary River, on dl. PilzeKulturgew. 193,898. gagrensis amphigena designated here,MycoBank, [mexicana sambucina (NY 2343036); ., no.336(1900)[ Georgia –469), Lindau(1910:134), Sambucus canadensis : CUP 14660,NY 2343299]. Sambucus nigra U. Braun,Crous&K. type: assiljevsky &Karakulin : Costa Rica Elenkin &Ohl, g. 185D),Braun& Mycoth. Ven F. L. L. ], 7Jul.1923,F. : Sousa daCâmara, Portugal: Chaves (Ellis &Kellerm.) Abkhazia Amer. Naturalist holotype isolectotypes IMA FUNGUS Illinois Biol. vul., Herb. Sambucus , Briosi& : Cartago, lectotype : Gagra : ., Funghi Bolez. Cent USA: Italy , Jul. Acta . : : ,
Cercosporoid fungi 4 ARTICLE
Fig. 49. Pseudocercospora depazeoides (Rabenh., Herb. Viv. Mycol., no. 587, lectotype of Passalora penicillata, HAL). A. Conidiophore fascicles. B. Conidia. Bar = 10 μm.
Mycol. Rom. 195. Scheuer, Mycoth. Graec. 433. Seym. & μm diam, immersed to somewhat erumpent, medium to dark Earle, Econ. Fungi 476. Syd., Mycoth. March. 1290, 2177. brown, composed of swollen hyphal cells, 3–7 μm diam. Syd., Mycoth. Germ. 1045, 1774, Thüm., Mycoth. Univ. 984. Conidiophores in small to large fascicles, loose to very dense, ill, Fungi Bav. 996. ahlbruckner, 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 subcylindrical to exuous, somewhat geniculate-sinuous, angular-irregular, 1–10 mm diam, sometimes con uent unbranched, 20–200(–300) × 3–7 μm, short conidiophores and larger, at rst pale greenish or greyish green, later 0–1-septate, longer ones pluriseptate throughout, pale brownish, nally grey to greyish white, often zonate, 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 to conidiogenous cells, 10–30 μm long, sympodially or blackish, scattered. Mycelium internal; hyphae branched, occasionally percurrently proliferating, conidiogenous septate, 2–4 μm wide, subhyaline to pale olivaceous, 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 tips truncate or subtruncate, 1.5–2.5 μm wide, or slightly
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Culture characteristics darkened-refractive. almost so,notdarkenedtosomewhatrefractiveorslightly truncate, (1.5–)2–2.5(–3) apex obtuse,baseshorttooccasionallylongobconically thin-walled, smooth,occasionallysomewhatrough-walled, hyaline, subhyalinetopaleolivaceousorolivaceous-brown, 30–140(–155) ×(3.5–)4–6.5(–7.5) Conidia paracercosporoid, visibleasminutecircle,1.5–2 refractive todarkened-refractive,infrontviewoccasionally [ Host rangeanddistribution W. Crous(CBSH-20391). Netherlands: Milingerwaard,on 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: The genericaf West 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), New 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 , Paracercospora irginia, Wisconsin). : on ( Sambucus kamtschatica, (now solitary, obclavate-cylindrical,straighttocurved, designated here,MycoBank,MBT202794). (designated here,MycoBank,MBT202793): ], Sambucus , Desm.,Pl.Crypt.France1849(NY 2343300). : Desm.,Pl.Crypt.France1849,e.g.FH,G,PC. Pseudocercospora griseola Sambucus nigra ., andBraun&Crous(2005)introduced spp.Itemergedveryclearlythat asingle nigra : complex(Braun nity of Colonies sp.), Adox , Ex-epitype culture μ , mexicana pubens m wide,hilaunthickenedor C. depazeoides : On Cercospora depazeoideshas , autumn, Sambucus nigra on MEA reaching16mm aceae, Asia (China,Japan, μ forcollectionswith m, (1–)2–8(–9)-septate, Sambucus , ], chinensis racemosa Pseudocercospora et al ) andthespecies Phaeoisariopsis : CBS126000. ealand, Central ealand, Central J. B.H. . 2013,Crous as speciesof ( canadensis canadensis , williamsii williamsii P. , 2007,P. , μ ebulus m diam. The Braun etal. ,
to somewhattortuous, simpleorbranched, 50–190×4–5.5 stomata, erect,subcylindrical, curved,geniculate-sinuous internal hyphaeorsmallhyphal aggregations,through Conidiophores small, onlyafewsubstomatal swollenhyphalcells,brown. wide, septate,paleolivaceous. or partlyexternal;super brownish, formingirregularpatches. surface. later greyishwhite,margininde spots, 1–8mmdiam,palebrown,yellowishbrownto discolorations, yellowishtobrownish,orcircularirregular Description (1995: 49, g.46),Guoetal.(1998:60, Illustrations Braun (2003:168). & Hsieh(1995:47–48),Guo Literature Basionym (Fig. 50) Pseudocercospora ebulicola Ramularia sambucina HAL, hasbeenexaminedandturnedouttobeamixtureof being made. with undesirable andrequiresaproposaltoconserve older heterotypicsynonymof molecular sequenceanalyses,renders based onmorphologicalreassesmentsandresultsof the allocationof genus penicillata serious consequencesongeneticlevelsince reallocation of culture andsequencesobtainedfromthismaterialhelpto 187 (1876)”butitwas combination isusuallycitedas“ con introduction ofthecombination we followSaccardo’s treatmentontheoccasionofhis included intheverymeagreoriginaldescription.However, cercosporoid aswellthealternarioidhyphomycetewere also analternarioidfungus.Itcannotbeexcludedthatthe contains besidesthetypicalcercosporoidhyphomycete by Crousetal.(2013). the depazeoides sambucigena (Braun in thecurrentsenseandcircumscriptionofthisgenus x thephylogeneticpositionandaf eneta 280(1875).Thedesignatedepitype,theex-epitype Mycol. Pap.140:143(1976). Nat. Hist.Soc.13:139(1934). Eriksson, FungiParas.Scand.Exs.42,depositedat Lectotype materialof Pseudocercospora ne thisnametothecercosporoidelement.Saccardo’s C. apiiasconservedtypeunder Art. 14.9,whichis Cercospora et al.2013,Crousal.2013). : Chupp(1954:100),Hsieh&Goh(1990:50),Guo Caespituli : Cercosporaebulicola (i.e. : Hsieh&Goh(1990:50, : . The phylogeneticpositionofthisspeciesin mustbereducedtosynonymywith C. depazeoides Leaf spotsamphigenous,inde C. depazeoides in loosefascicles,3–15, arising from C. depazeoides (Braun 1995,Braun hypophyllous, effuse, olivaceousto and cladehasrecentlybeencon rst validlypublishedinMycotheca Cladosporium herbarum cial hyphaebranched,2–3 etal.(1998:60–61),Crous& ) beingthetypespeciesof Cercospora depazeoides Pseudocercospora toPseudocercosporahas Exosporium depazeoides W. Yamam., nite orbrownontheupper Nuovo Giorn.Bot.Ital Stromata (W. Yamam.)Deighton, nity ofthisspecies. The to g. 34),Guo&Hsieh g. 46). Pseudocercospora, et al Pseudocercospora Mycelium Cercospora lacking orvery Trans. Sapporo . 2013). Thus, IMA FUNGUS Cercospora Cercospora , whichis . internal rmed and nite . μ an 8 m 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. Conidia. Bar = 10 μm. U
Host range and distribution: On Sambucus (javanica [formosana]), Adoxaceae, Asia (China, Sichuan, hejiang; 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, super cial hypha. C. Conidiophore tips. D. Conidia. Bar = 10 μm. incorrect. Cercospora ebulicola is a species of the genus Pseudocercospora whereas C. catenospora is a Phaeoramularia-like species of Passalora. μm, 2–8-septate, olivaceous to olivaceous-brown, thin- 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 20–130 × 3–5 μm, 2–11-septate, subhyaline, yellowish 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). 1.5–2.5 μm wide, hila unthickened, not darkened. 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.
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Passalora viburni-sargentii Phaeoramularia penicillata Misapplied names Germany Lectotype not toslightlyrefractiveordarkened-refractive. short obconicallytruncate,2–3 short conicallytruncateincatenateconidia,base to of primaryandsolitaryconidiaobtuse,rounded,truncate to subhyaline, laterpaleolivaceous,thin-walled,smooth,apex occasionally somewhatconstrictedatthesepta,hyaline, curved, (20–)30–90(–145)×(2.5–)3–6(–7) obclavate-subcylindrical tocylindrical,straightsomewhat catenate, insimple,shortacropetalchainsordisarticulating, as minutecircles,about1.5–2.5 refractive, butunthickened,infrontviewsometimesvisible being truncateandsomewhatrefractiveordarkened- conidiogenous lociinconspicuoustosubconspicuousby reduced toconidiogenouscells,about10–40 conidiogenous cellsintegrated,terminalorconidiophores olivaceous-brown andwallssomewhatthickened,smooth; rst subhyalineandthin-walled,laterpaleolivaceousto aseptate orsparinglyseptate,usuallywith0–3septa,at with asinglebranchlet,15–70(–80)×(2.5–)3–7(–8) moderately geniculate-sinuous,unbranchedoroccasionally through stomataorerumpent,erect,straight,subcylindricalto fascicles, loosetodense,arisingfromstromata,emerging somewhat thickened.Conidiophores brown, composedofswollenhyphalcells,2–7 internal. punctiform, scattered,darkbrowntoblackish. white, withnarrowdarkmargin. greenish, ochraceoustobrown,centrelatergreyish con uent andlarger, occasionallysomewhatzonate,dull somewhat angular-irregular, 1–8mmdiam,sometimes Description Mycoth. Graec.382.Smarods,FungiLat.Exs.448. Mycoth. Gen.1312.S Fungi Imperf.Exs.445.Liro,Mycoth.Fenn.295.Petr., Exsiccatae: Fuckel,FungiRhen.Exs.118. Kabát&Bubák, g. 82),Guo(2012:161, Braun (1992:216, Illustrations (1954: 103),Ellis(1976:247),Braun&Mel’nik(1997:19,77). (1910: 136), Literature (FH, G,K(M),etc). 118] (HAL). Viburnum opulus 161 (2012)[ Inst. im.V.L. Komarova on (1886: 468)andChupp(1954:103). Phytopathol. Sin calvescens Viburnum opulus Stromata : Saccardo(1886:468),Lindau(1910:136),Ferraris : (designatedbyBraun& Mel’nik 1997:20): : : Chupp(1954:103),Ellis(1976:248, Rheinland-Pfalz Isolectotypes Leaf spots ], 5 Aug. 1984, assiljevsky &Karakulin(1937:236),Chupp , summer1863[Fuckel, holotype substomatal orimmersed,10–80 : g. 9),Braun&Mel’nik(1997:appendix, . 12(4):13(1982). Cercospora penicillatasensu vul., Herb.Mycol.Rom.432.Scheuer, g. 2). subsp. : Fuckel, : 20:19(1997). (Ces.) X.J.Liu& Y.L. Guo, : amphigenous, subcircularto China Y.L. Guo, Mycosystema Y. L.Guo266 “in sylvaHostrichiensi”,on μm diam.Conidia μm wide,hilaunthickened, calvescens Caespituli : Fungi Rhen.Exs Hubei in smalltofairlylarge μm, 1–10-septate, Fungi Rhen.Exs : Shennongjian, (HMAS47828)]. μm diam,walls amphigenous, [sargentii Saccardo g. 186A), solitary to μm long; Mycelium μm diam, . 118, Acta Braun etal. μm, var. 31: . Braun was reallocatedtothelattergenus(Crous&Braun2003, reduced tosynonymywith unthickened. Braun(in&Mel’nik1997)classi 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 opulus are homotypicsynonyms.Chinese materialon morphological reassessments, based onresultsofmolecularsequenceanalysesand genus Passalora loci andhilaofthisspeciesasintermediatebetween opulus on (syn. confusions andmisinterpretations. history ofthisspeciesiscomplicatedandcharacterisedby but burejaeticum unproven andmightratherpertainto the FarEastofRussia(Egorova2007)isuncertainand Notes: A recordofthisspecieson 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, depazeoides be indistinguishablefromEuropean materialof hosts (Crous &Braun2003)areadditionalunprovenanddoubtful Clement phylogenetic studies(Winkworth&Donoghue2005, which hasbeencon [opulus calvescens Host rangeanddistribution opulus Sambucus nigra Pseudocercospora viburni A collection ofthisspeciesfromthe Asian partof Turkey Sambucus nigra P. opuli , andFuckel(1870)treatedthefunguson of P. opuli subsp.calvescens et al.2013,Crous assolespeciesof Pseudophaeoramularia subsp. Cercospora penicillata var. et al.2014). andPseudocercospora C. penicillata , (Liu &Guo1982)wasalsobasedonChupp’s seems tobecon sargentii ) wasintroducedforacercosporoidfungus does notbelongto opulus americanum C. opulicanbeinterpretedasnewspecies not belongingtosect. . Fuckel(1863)added wasnotmentioned,whichseemsto var. , opulus . Chupp(1954)followedSaccardo’s Viburnum rmed aswellsupportedcladein calvescens et al.2013). hasbeenexaminedandfound to : On , trilobum C. penicillata Pseudocercospora subsp. and ned tospeciesofthissection, Viburnum , Pseudophaeoramularia for suchspecies.However,
Passalora viburni-sargentii now lantana Viburnum and proposedthenew Cercospora penicillata ], calvescens ]), opulus V. burejaeticumfrom Passalora penicillata Opuli. (edule P. varia . , i.e.“f. Pseudocercospora Adoxaceae andV. orientale Phaeoramularia C. penicillata
subsp. The taxonomic sect. [SDXFLÀRUXP andC.opuli P. opuli. IMA FUNGUS [opulus penicillata Viburnum Viburnum Viburnum trilobum Opulus , Asia was var. ed f. ], ” . ,
Cercosporoid fungi 4
has been examined (Kütahya, on Viburnum opulus, 9 Jul. ARTICLE 1953, ex herb. Petrak, G U). Pseudocercospora varia is morphologically very close to and confusable with P. opuli, but the conidia are formed singly. The latter species seems to be con ned to species 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: Hsieh & Goh (1990: 53, g. 36, as Stigmina tinea), Guo (1995: 125, g. 4), Guo et al. (1998: 64, g. 49).
Description: Leaf spots amphigenous, subcircular to angular- irregular and vein-limited, 0.5–7 mm diam, often con uent and along the leaf margin, brown to dark brown, margin inde nite 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 external; super cial hyphae hypophyllous, emerging through stomata or arising from the base of conidiophore fascicles, branched, septate, pale olivaceous, thin-walled, smooth, 1.5–2.5 μm wide. Stromata well-developed, substomatal, 15–75 μm diam, brown. Conidiophores in small to moderately large fascicles, divergent to usually dense, arising from Fig. 52. Pseudocercospora tinea (based on Guo 1995: 125, g. 4). A. stromata, through stomata or solitary, arising from super cial Conidiophore fascicles. B. Conidiophores. C. Solitary conidiophores hyphae, erect, straight to curved, subcylindrical, somewhat arising from super cial hyphae. D. Conidia. Bar = 10 μm. attenuated towards the tip or irregular in width, geniculate- sinuous, unbranched, relatively short, 5–35 × 2–5 μm, 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. μm long, conidiogenous loci inconspicuous or subdenticulate, 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 obclavate-cylindrical, 10–120 × 2–3.5(–4) μm, 1–11-septate, 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 to usually obconically truncate, 1–2.5 μm wide, hila Pseudocercospora viburnigena) is clearly distinct from the unthickened, not darkened. Chinese species by having non-geniculate, only percurrently proliferating conidiogenous cells with ne annellations. Holotype: China: Guangdong Province: Guangzhou, on Super cial hyphae are only rarely formed. Pseudocercospora 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 been examined and identi ed by C. Nakashima (TUA 40, 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, revised and con rmed as P. tinea by C. Nakashima (CNS567, Guangdong, Sichuan, hejiang; Japan, Taiwan). CNS976).
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brownish halo. raised, andsometimeswithochraceous, yellowishbrownto to moderaterlywide,darkbrown margin,sometimessomewhat white, marginindistinct,formed asmarginallineorwithnarrow to mediumdarkbrownorreddish brown,latergreytogreyish irregular, 1–8mmdiam,sometimescon Description 280. Ellis&Everh.,N. Amer. Fungi3190. Exsiccatae: Clements&Clements,Crypt.Format.Colorad. Hsieh (1995:51),Crous&Braun(2003:418). Literature Basionym (Fig. 53a) Pseudocercospora varia μm. A. Fig. 53a.Pseudocercosporavaria Conidiophore fascicles. State Mus.Nat.Hist var. varia Cheng, : Saccardo(1886:468),Chupp(1954:105),Guo& : Cercospora : Acta Mycol.Sin.11 Leaf spots Caespituli amphigenous,subcirculartoangular-
. 35:141(1884). varia B. amphigenous, punctiform, scattered, amphigenous, punctiform, scattered, Conidiophores. Peck,
var. (Peck) J.K.Bai&M.Y. : 123(1992). varia Rep. (Annual)New York uent andlarger, pale (BPI 442161,isotype). C. Conidia. Bar=10 Braun etal. Passalora antipus specimens, e.g.on lonicericola Collections on varia onLonicera rather infavourof (short petiolesdonotcoincidewith accordance with with solitaryconidiophoresfoundon“ nal identi (PDD 19826,19843,1985),butsuf unclear. The collectionsconcernedhavebeenexamined (Gagdil 2005)on to asP. opuli short chainsandlateralbranchlets. They arereferred opulus Notes: Collectionson Wisconsin, Wyoming). South Carolina, Texas, Massachusetts, Mississippi,New York, Oklahoma,Oregon, Alabama, Alaska, Florida,Idaho,Illinois,Iowa,Kansas, suspensum Holotype unthickened, notdarkened. thickened, to0.5 mostly palertowardsthetip,wallthintousuallysomewhat olivaceous-brown oryellowishbrown,aboveallbelow, at unbranched, (5–)10–80(–100)×2–9 occasionally somewhatgeniculate-sinuousintheupperhalf, or attenuatedtowardsthetip,mostlynotgeniculate-sinuous, erumpent, erect,straighttosomewhatcurved,subcylindrical rarely dense,arisingfromstromata,throughstomataor or inusualysmallfasciles,mostly2–15,usuallydivergent, μ cells globoseorsubglobosetosomewhatirregular, 3–10 subglobose, paletomediumbrownoryellowishbrown, or intraepidermal,smalltowell-developed,10–60 1.5–3 by conidialgermination);hyphaebranched,septate,pale, some collectionswithsuper brown, darkbrowntoalmostblackish. to shortorlongobconicallytruncate,1–2.5 smooth, apexobtuse,occasionallysubacute,basetruncate × 2–6.5μ subacicular, straighttosomewhatcurved,(15–)20–85(–90) circle. occasionally somewhatrefractiveorlocivisibleasminute or subdenticulate,butalwaysunthickenedandnotdarkened, conidiogenous lociinconspicuousorvisiblebybeingtruncate conidiophores reducedtoconidiogenouscells,15–40 thick, smooth;conidiogenouscellsintegrated,terminalor prunifolium alnifolium Host rangeanddistribution 442161. Viburnum acerifolium m diam,wallthickened,to1 rst very pale, hyaline to subhyaline, later pale olivaceous, rst verypale,hyalinetosubhyaline,laterpaleolivaceous, Conidia μ and m wide,thin-walled,smooth. : , cassinoides m, 1–7-septate, hyaline to subhyaline, thin-walled, m, 1–7-septate,hyalinetosubhyaline,thin-walled, USA: cation hasnotbeenfound.Supercialhyphae , pubescens , V. trilobum (material examined:BPI 442155), other Adoxaceae . RecordsofthisspeciesfromNew solitary, obclavate-cylindrical,cylindrical,rarely L. japonica New York P. varia. The hostidentityisalsouncertain spp.arealsobasedonmisidenti μ Viburnum lantata (Ellis &Holw.) U.Braun&Crous. m, aboveallbelow, sometimesto1 L. ciliata V. burejaeticum , Aug., , dilatatum Viburnum edule UD¿QHVTXLDQXP , have atendencytoformconidiain , North America (Canada;USA, : Albany County, EastBerne,on ermont, C. H.Peck(NYS). (BPI 442168–442170),refer to belong toPseudocercospora : OnViburnum cial hyphae probably formed cial hyphaeprobablyformed μ , lentago m. and cient fructicationfora μ V. lantana . OtherrecordsofC. m, usually 0–3-septate, m, usually0–3-septate, irginia, West Conidiophores Stromata V. lantana Mycelium SDXFLÀRUXP [9 V. carlesii var. , odoratissimum μ DI¿QH m wide, hila m wide,hila IMA FUNGUS but theyare Isotype: BPI (acerifolium substomatal substomatal ” arenotin internal (in internal (in arequite μ cations. μ m diam, m diam, ealand [ m long, m long, solitary solitary DI¿QH irginia, ], μ V. V. m ], , , Cercosporoid fungi 4
having much larger, dense fascicles of conidiophores. The ARTICLE taxonomic status of the Asian collections is not quite clear and in urgent need of molecular con rmation. It cannot be 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 having much longer, usually obclavate conidia, 25–125 × 4.5–8 μm, with 2–10 septa and somewhat thickened walls and subconspicuous conidiogenous loci. Pseudocercospora caprifoliacearum, known from India on Viburnum sp., has longer conidiophores, 60–155 μm, and the conidia are 3–6.5 μm wide and thin-walled.
Description: Leaf spots amphigenous, subcircular to angular- irregular, 1–8 mm diam, medium to dark brown, nally with pale centre, greyish brown to dingy greyish white, margin inde nite or narrow, darker, or surrounded by darker veins, occasionaly somewhat raised. Caespituli amphigenous, punctiform, scattered, blackish. Mycelium internal. Stromata lacking or substomatal to intraepidermal, 10–50 μm diam, brown, swollen hyphal cells 2–7 μm diam, walls slightly 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, unbranched, 20–100 × 3.5–8(–10) μm, (0–)1–4(–5)-septate, Fig. 53b. Pseudocercospora varia var. viburni-sargentii (KUS 14011, medium to dark brown throughout or somewhat paler holotype). A. Conidiophore fascicles. B. Conidia. Bar = 10 μm. towards the tip, wall somewhat thickened, to 0.8 μm, smooth; conidiogenous cells integrated, terminal, 10–30 μm long, 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) denticle-like, 1.5–3 μm diam, often somewhat refractive or 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 × 4.5–8 μm, 2–10-septate, occasionally subhyaline to pale Illustrations: Guo & Hsieh (1995: 51, g. 48), Guo et al. (1998: olivaceous, but usually medium olivaceous or olivaceous- 66: 50), Shin & Kim (2001: 227, g. 104). brown, wall thin to somewhat thickened (to 0.8 μm), smooth, apex obtuse to subacute, base obconically truncate, 2–3 μm 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; conidia, 35–145 × 3.5–6.5 μm, 2–15-septate. Holotype: USA: Wisconsin: Madison, Arboretum, on Viburnum sp. [?UD¿QHVTXLDQXP] (as Viburnum “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 irginia).
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
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(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.: 907(1979). hao, ActaMycol.Sin.12:198(1993). Ray, 1941. Adoxaceae et al.2013). : : Cercospora viburni-cylindrici Nova Hedwigia nom. illeg. 3815(BPI439059)]. This speciesiseasily Cercospora viburnicola byitsconsistentlysolitary, pigmented, , 6 Aug. 1909,B.Brooks (Art. 53.1),nonC.viburnicola Pseudocercospora , arequitedistinctbynarrower, 55:219(1992). (FL. Tai) Y.L. Guo&W.X. F.L. Tai, Viburnum F.L. Tai, Prathigada Pseudocercospora species onhosts Lloydia (F.L. Tai)U. spp.,e.g. Syll. Fung. , hb. species
11 W.W. W.W. : 54 Braun etal. J.
20–35 developed, subglobosetosomewhatirregular, substomatal, 25–75 ×2.5–4 solitary, obclavate-cylindrical, straightoronlyslightlycurved, loci inconspicuousunthickened andnotdarkened. cells integrated,terminal,about 10–30 to brown,wallsomewhatthickened, smooth;conidiogenous 50–150 ×3–4.5μm,3–7-septate,mediumolivaceous-brown straight, subcylindricaltogeniculate-sinuous,unbranched, fascicles, arisingfromstromata,throughstomata,erect, large, somewhatdivergenttodenseorevencoremioid dark, browntodarkbrown. halo. darker margin,brown,sometimeswithadiffuse olivaceous nally centrebrown,greyishbrownorwhitewith brown todarkorevenblackish,margininde irregular, sizevariable, 1–8mmdiam,sometimescon Description g. 51). g. 10),Guo&Hsieh(1995:53, Illustrations Crous &Braun(2003:421). Literature Caespituli μm diam,brown. : Guo&Hsieh(1995:52), : : Tai (1948:37, g.1),Braun(1992:216,plate2, Leaf spots μm, 2–5-septate, paleolivaceous,thin-walled, hypophyllous, punctiform,scattered,effuse, Bar =10μm. fascicles. (BPI 442174,holotype). Fig. 54.Pseudocercosporaviburnicola amphigenous, subcirculartoangular- Conidiophores Mycelium B. Conidiophoretips. g. 49),Guoetal. μm long,conidiogenous internal. in smalltomoderately etal. A. Stromata IMA FUNGUS Conidiophore (1998: 65); (1998: 67, C. nite or Conidia Conidia. uent, well- Cercosporoid fungi 4
smooth, apex obtuse to subacute, base short obconically Pseudocercospora viburni-erosi U. Braun & H.D. ARTICLE truncate, 1–2 μm wide, hila unthickened, not darkened. 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: Shin & Kim (2001: 229, g 105).
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 much longer, pluriseptate, cylindrical- liform conidia, length fascicles are smaller and not coremioid. Therefore, we prefer 50–130 μm, with truncate base. 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). Bar = 10 μm. A. Conidiophore fascicle. B. Conidia. Bar = 10 μm.
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wide, hilaunthickened,notdarkened. smooth, apexobtuse,basetruncateoralmostso,2–2.5 3–12-septate, subhyalinetoverypaleolivaceous,thin-walled, or almostsotomoderatelycurved,50–130×2.5–4 not darkened. terminal, conidiogenouslociinconspicuous,unthickened, the tip,thin-walled,smooth;conidiogenouscellsintegrated, μm, 3–8-septate,olivaceous-browntobrown,palertowards above allintheupperportion,unbranched,60–120×3–4 straight, subcylindricaloronlyslightlygeniculate-sinuous, coremioid, arisingfromstromata,throughstomata,erect, in well-developedfascicles,to30,denseverydense, μm diam,globoseorsubglobose,brown. Stromata internal; hyphaebranched,septate,hyaline,2–3 Caespituli centre turninggreyishwhite,withraisedbrownborderline. centre surroundedbyabrowntodarkborder, brown todarkbrown,laterwithgreyishgrey sometimes con circular oralmostsotoangular-irregular, 3–10mmdiam, Description substomatal, smalltowell-developed,15–30 : hypophyllous, brown,punctiform. Leaf spots uent andlarger, to15mmdiam,at Conidia amphigenous, scatteredtocon solitary, cylindrical- liform, straight Conidiophores Mycelium μm wide. uent, nally Braun etal. μm, rst μm separate species. better excludedfromP. viburni-cylindrici differences, theKoreanmaterialon 50–130 much longer, pluriseptate,cylindrical- dense, oftencoremoidconidiophorefasciclesandaboveall Korean materialon 2001). The twospeciesaremorphologicallyveryclose,but assigned to Notes: Type materialofthenewspecieswaspreviously collection. Host rangeanddistribution taquetii Holotype Basionym (Fig. 57) Pseudocercospora viburnigena Mycol. Progr 1994. non , 4Sep.1997, Pseudocercospora tinea μm, withtruncatebase.Basedontheseobvious : : Korea: Kwachon,onViburnum erosum Cercospora tinea Pseudocercospora viburni-cylindrici Conidia. Bars=10 fascicles. Mycoth. Ven. 1252,HAL,lectotype). Fig. 57. . 1:23(2002). Viburnum erosum H. D.Shin(KUS-F14126). Pseudocercospora viburnigena B. Stroma cells.C.Conidiophores. : Onlyknownfromthetype Sacc., μm. Y.L. Guo &W.H. Hsieh Michelia differs inhavinglarge,
liform conidia,length U. Braun&Crous, Viburnum erosum andtreatedas
1: 268(1878), A. IMA FUNGUS (Shin&Kim Conidiophore (Sacc., var. is D. Cercosporoid fungi 4
Synonyms: Cercoseptoria tinea (Sacc.) Deighton, Mycol. Mycoth. en. 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: Ellis (1976: 117, g. 83B), Kirk (1999: g., unnumbered), Crous et al. (2013: 109, g. 65). Notes: A lectotypi cation of Cercospora 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. en. 1252” is designated as lectotype. pale to darker brown or centre later greyish brown to dingy The phylogenetic af nity of this species, previously referred grey, margin inde nite or darker, sometimes with reddish 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 1.5–4 μm wide, subhyaline, thin-walled, smooth. Stromata from Cercospora tinea by having super cal hyphae with substomatal or immersed, 15–80 μm diam, occasionally solitary conidiophores and conidiogenous cells sympodially larger, to 120 μm diam (hypophyllous stromata smaller 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. composed of brown swollen hyphal cells, 3–6 μm diam, 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 μm. Conidiophores 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 super cial hyphae, straight to somewhat curved-sinuous, 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, con rmed (USA: Louisiana: Lafayette, on Viburnum plicatum, apex at rst rounded, later truncate or subtruncate, 5–30(– 21 Sep. 1886, A. B. Langlois, BPI 441939, 441940). 50) × 2–5(–6), 0(–2)-septate, pale brown or olivaceous- brown, thin-walled, smooth, occasionally slightly rough- Pseudocercospora viburni-nudi U. Braun, sp. nov. walled; conidiophores mostly reduced to conidiogenous MycoBank MB814562 cells, occasionally integrated, terminal, 5–20 μm long, with (Fig. 58) a single terminal conidiogenous locus, 2–3 μm wide, neither 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 to narrowly cylindrical- liform, occasionally somewhat Adoxaceae by having much shorter, 1–4(–5)-septate conidia, obclavate, straight to curved or somewhat sigmoid, (20–)30– 15–40 × 3–7 μm. 110(–120) × 2–5 μm, (1–)3–11-septate, subhyaline to very pale olivaceous or somewhat brownish, thin-walled, smooth, Description: Leaf spots amphigenous, subcircular to angular- guttulate when fresh, apex obtuse to subacute, base truncate irregular, sometimes vein-limited, 2–12 mm diam or con uent to somewhat obconically truncate, 1.5–3 μm wide, hila and larger, medium to dark brown, margin inde nite, later with 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 45 μm diam, dark brown. Conidiophores 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 unbranched, 40–105 × 3–5 μm, base occasionally somewhat [Sacc., Mycoth. en. 1252] (HAL, s.n.). Isolectotypes: Sacc., wider, to 7 μm, apex sometimes slightly swollen, to 8 μm wide,
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portulacastrum Conidiaobclavate-cylindricaltoacicular, to125 ...... 1 Key toCercosporaspecies on Aizoaceae Cercospora Aizoaceae 15–40 ×3–7 obclavate-cylindrical, fusiform,ellipsoid,straighttocurved, always unthickenedandnotdarkened. subdenticulate, denticle-likelociabout1.5–2 conidiogenous lociinconspicuoustoconspicuousbybeing integrated, terminal,15–40 a singleconspicuousannellation;conidiogenouscells occasionally withasingleenteroblasticrejuvenationleaving the tip,wallsomewhatthickened,to1 1–5-septate, mediumtodarkbrownthroughoutorpalertowards Conidiaacicular, to200 μ m, 1–4(–5)-septate, often slightly constricted m, 1–4(–5)-septate,oftenslightlyconstricted μ m long, sympodially proliferating, m long,sympodiallyproliferating, μm long,basetruncate; on Conidia μ m, conidiophores m, conidiophores μ m diam, but loci m diam,butloci solitary, short μm long,basetruncatetoobconically truncate;on Braun etal. Tetragonia tetragonoides nudum Holotype μ short obconicallytruncate,sometimesrounded,1.5–2.5(–3) or olivaceous-brown,thin-walled,smooth,apexobtuse,base at thesepta,subhyalinetomostlypalemediumolivaceous collection. Host rangeanddistribution m wide,hilaunthickened,notdarkened. , Adoxaceae : USASouthCarolina ...... , 15Oct.1943, : Onlyknownfromthetype ...... tips. D.Conidia.Bar=10 Conidiophore. A. nudi Fig. 58.Pseudocercosporaviburni- : MurrellsInlet,on Conidiophore fascicle. Carter 855 (BPI 442176,holotype). Trianthema C. trianthematis C. Conidiophore C. tetragoniae (BPI442176). IMA FUNGUS Viburnum μm. 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. substomatal, 10–25(–40) μm diam, brown. Conidiophores Nac. Hist. Nat. Buenos Aires 20: 429 (1911). solitary or fasciculate, 2–10, divergent, arising from internal Synonyms: Cercospora tetragoniae (Speg.) assiljevsky, hyphae or stromata, through stomata or occasionally in assiljevky & Karakulin, Fungi imperfecti parasitici 1. erumpent, erect, straight, subcylindrical to geniculate- Hyphomycetes: 221 (1937). sinuous, unbranched, 20–125 × 3–7 μm, 0–6-septate, pale Cercospora tetragoniae (Speg.) Chupp, in iégas, 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 & 10–30 μm long, conidiogenous loci conspicuous, thickened Pons (1980: 216), Braun & Mel’nik (1997: 97), Braun (2000b: and darkened, 2.5–4 μm diam. Conidia solitary, acicular, 78), Crous & Braun (2003: 400). straight to curved, 30–200 × 2.5–5 μm, 3- to pluriseptate, hyaline, thin-walled, smooth, apex pointed, base truncate, Illustration: Sutton & Pons (1980: 207, g. 1F). 2.5–4 μm wide, hila thickened and darkened.
Description: Leaf spots amphigenous, circular or subcircular, 2–10 mm diam, scattered to con uent and larger, up to 20
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). μm. A. Conidiophores fascicle. B. Conidia. Bar = 10 μm.
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Leafspotslarger, usually1–10mm diam; caespitulilessconspicuous,neverpustulate;conidiophorespaler, oftensomewhat raisedmargin;caespitulimainlyepiphyllous,conspicuous,punctiformtoalmostpustulate, Leafspotssmall, 0.5–4mmdiam,dingygreenishgrey, brown,grey, 2 (1) Withdistinctleafspots;myceliuminternalandoften alsoexternal;conidiophoresfasciculateandoften 1998) arebasedonmisidenti [expansa Host rangeanddistribution 241730 (slide). (Fig. 60) 1 Leaf Key tocercosporoidspecieson Altingiaceae spots Altingiaceae Synonym Cercospora 18 Nov. 1909, Holotype Records fromBrazilon Notes: This speciesbelongstothe (USA, Maryland,Indiana, Texas). America (Argentina,ElSalvador, Brazil),North America Israel, Japan),Caucasus(Georgia),CentralandSouth Sierra Leone, Tanzania, Uganda, Casepituli irregular, scattered,0.5–6mmdiam,darkbrowntoblackish. Description Bhartiya Illustrations Literature 25: 45“1997”(1998)[ Appl isotype Aizoaceae Gorakhpur, on onLiquidambar subhyalineorpaletomedium olivaceousorolivaceous-brown,hilaunthickened,notdarkened conidiogenouslociinconspicuous, neitherthickenednotdarkened;conidiashorter, to100 subhyalinetopaleolivaceousorolivaceous-brown,thin-walled, shorter, 5–35 somewhatdarkened-refractive;on conidiato150μmlong,subhyalineorverypaleolivaceous,hilaunthickened toslightlythickened, aroundtheouterrim,somewhatdarkened-refarctive,1–1.5 thickened(to1μm),(10–)25–90 darkbrowntoblackish;conidiophorespalemedium brown,palertowardsthetip,wallsomewhat conidiamuchnarrower, 2–4 alsosolitary, arisingfromsuper ...... unthickened,nordarkened;conidia20–80×4–7.5 super . 17:80(1962);as“ et al.(1998:45, : ]), Aizoaceae : Cercosporaaizoacearum : Crous&Braun(2003:409),Kamal(2010:14,94). Argentina: LaPlata,onTetragonia tetragonoides : GPU8073]. : amphigenous, mainly hypophyllous, punctiform, amphigenous, mainlyhypophyllous,punctiform, : Chiddarwar(1962:78,plateII, amphigenous, circular to oval or Leaf spotsamphigenous,circulartoovalor
, Sep.1998,H.D.Bhartiya trianthematis C. Spegazzini cial hyphaelacking;conidiophoresfasciculate,0–1-septate,3–7 absentoralmostso,sometimeswithslightdiscolorationsontheupperleafsurface;myceliuminternal; Trianthema portulacastrum , Africa (Cameroon,Kenya,Malawi, g. 2). Spinacia oleracea holotype formosana : On (LPS16153). cations. trianthemae Chidd., Tetragonia imbabwe), Asia (Brunei, imbabwe), Asia : Bhartiyaetal., C. apiis.lat India and/VW\UDFLÀXD, Asia andNorth America ...... Mycopathol. Mycol. μm 2 ...... : Isotype: K(M) cial hyphaeifpresent,aseptate,narrower, 2–5 ”. Uttar Paradesh μm long;conidiogenouslociconspicuous,slightlythickened, atleast (Mendes (tetragonoides (HCIO, s.n.); /LTXLGDPEDUVW\UDFLÀXD gs 12–13), [monogyna . complex. Kavaka et al. Braun etal.
IMI ], , : μm; on longer conidia,arangesimilarto shows cylindricaltosomewhatobclavateshorterandacicular conidia weredescribedtobecylindrical,buttheillustration undoubtedly conspeci C. aizoacearum complex byhavingobclavate-cylindricaltoacicularconidia. true Note: A Aizoaceae brown. Host rangeanddistribution 32 (K(M)IMI83193).Isolectotypes:BPI441988,HCIO. India Lectotype 2–2.5 subobtuse, basetruncatetoshortobconicallytruncate,about (1–)3–11-septate, hyaline,thin-walled,smooth,apexpointedto to curved,(10–)30–125×2.5–4 conidia obclavatetosubcylindrical,longeronesacicular, straight thickened anddarkened,2.5–3 to conidiogenouscells,10–30 cells integrated,terminal,intercalaryorconidiophoresreduced to brown,wallsomewhatthickened,smooth;conidiogenous unbranched, 15–100×3–5 straight, subcylindrical-conicaltogeniculate-sinuous,usually to divergent,arisingfromstromata,throughstomata,erect, large fascicles,2–20,occasionallysolitary, relativelydense diam, olivaceoustobrown. Trianthema portulacastrum , North America ...... /LTXLGDPEDUVW\UDFLÀXD : μm diam,infrontviewvisibleasminutecircle; μ Maharashtra m wide,hilathickenedanddarkened. Mycelium nally usuallygreyishwhite,withdark, , Asia (India,Maharashtra). (designated here,MycoBank,MBT202798): Cercospora s.str. , describedfromIndiaonthesamehost,is μm wide,conidiogenousloci internal. : c. Type materialwasnotavailable. The Pune, Wanowri, MilitaryHospital,on μm long; μm wide; . Conidiophores Pseudocercospora tuberculans Stromata , 12Oct.1956,P. P. Chiddarwar : On μ , North America m, 0–7-septate, dark olivaceous m, 0–7-septate,darkolivaceous μ m diam. distinct fromthe μ Cercospora liquidambaris Trianthema portulacastrum m long, conidiogenous loci m long,conidiogenousloci C. trianthematis μ m, occasionallybroader, μm, substomatal, 15–45 Conidia in small to moderately in smalltomoderately P. liquidambaricola solitary, shorter IMA FUNGUS C. apiis.lat. . μ m ,
Cercosporoid fungi 4 ARTICLE
Fig. 61. Cercospora liquidambaris (CUP- A-002227(AL), lectotype). A. Conidiophore fascicles. B. Solitary conidiophores arising from super cial hyphae. C. Conidiophores. D. Conidia. Bar = 10 μm.
Cercospora (s. lat.) substomatal or intraepidermal, 10–60 μm diam, brown. 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) from super cial hyphae, lateral, erect, straight, subcylindrical or attenuated towards the apex to usually geniculate-sinuous, Literature: Chupp (1954: 259). unbranched, (10–)25–90(–100) × 3–5 μm, (0–)1–8-septate, pale to medium brown, paler towards the tip, wall somewhat Description: Leaf spots amphigenous, subcircular to angular- thickened, at least below, to 1 μm, smooth or almost irregular, small, 0.5–4 mm diam, dingy greyish green, brown, so, proliferation sympodial and occasionally percurrent to nally usually greyish white or white, sometimes somewhat (conidiophores with enteroblastic rejuvenation leaving delicate raised, margin inde nite or usually darker, narrow. Caespituli annellations); conidiogenous cells integrated, terminal, 10–25 amphigenous, usually epiphyllous, very conspicuous, μm long, conidiogenous loci conspicuous, in front view visible punctiform to almost pustulate, scattered, dark brown to as minute circle, 1–1.5 μm diam, at least rim slightly thickened, blackish. Mycelium internal and external; super cial hyphae somewhat darkened or refractive. Conidia solitary, narrowly lacking or almost so to well-developed, branched, septate, obclavate-cylindrical, liform-subacicular, straight to curved, subhyaline to pale olivaceous or brownish, 1.5–4 μm wide. occasionally germinating with short to moderately long lateral Stromata almost lacking or small to usually well-developed, outgrowths, 40–150 × (2–)2.5–4(–4.5) μm, 3–12-septate,
VOLUME 6 · NO. 2 435 436 ARTICLE
Basionym Pseudocercospora liquidambaricola Pseudocercospora specieson Altingiaceae Pseudocercospora Pseudocercospora neoliquidambaris Pseudocercospora liquidambaris Cercoseptoria liquidambaricola Synonyms (Fig. 62) USA: Lectotype so toslightlythickened,somewhatdarkenedorrefractive. sometimes truncate,1–2 acute tosubobtuse,baseusuallyshortobconicallytruncate, subhyaline topaleolivaceous,thin-walled,smooth,apex lat. untilphylogeneticdatawillbeavailable. Therefore, thisspeciesistentativelyretainedin phylogenetically quiteheterogeneous(S.I.R. as inthePassalora darkened. The conidiaare cercospora-like,butpigmented The conidiogenouslociandconidialhilaarethickened The genericaf Atkinson’s herbariumat CUP isdesignatedaslectotype. collected in Alabama in1891. A correspondingsamplefrom clari been examined. The identity ofthetrue VW\UDFLÀXD. SeveralcollectionsfromMexicoandtheUSA have species isalsoknownfromNorth America on correctly assignedto these hostsin Asia belongsin which is,however, incorrectsincethecercosporoidfunguson to synonymywith material causingaleafspotdisease reduced liquidambaris Notes: The taxonomyandnomenclatureof Montana, Massachusetts,NorthCarolina, Texas). Delaware, Florida,Indiana,Louisiana,Maryland,Mississippi, Altingiaceae Host rangeanddistribution 1891, Braun, Kobay., 14 Oct.1928, (1990) [ & Goh,Cercosporaandsimilarfungi fromTaiwan Trimestriel Soc.Mycol.France PPE, hb.Sawada)]. Liquidambar formosana Cooke &EllisexG.F. Atk., 1892[ 39.1), and Agric. Res.Inst.Taiwan Trimestriel Soc.Mycol.France VXSHUÀ ed bylectotypi Alabama G. F. Atkinson [CUP-A-002227(AL)]. C. liquidambaris ( : Cercosporaliquidambaricola : Cercosporaliquidambaris ] (Art.52.1). type: Schlechtendalia Mycoscience , North America (Mexico;USA, Alabama, designated here,MycoBank,MBT202799): have been totally confused. Chupp (1954) havebeentotallyconfused.Chupp(1954) : Auburn, on : Auburn, nom. illeg Taiwan: Taipei,on nity of K. Sawada(NTU-PPE,hb.Sawada)]. C. liquidambaris cation. Atkinson (1892)citedaspecimen compex, which recently proved to be compex,whichrecentlyprovedtobe P. liquidambaricola μ
C. liquidambaris . (Art.53.1),nonC.liquidambaris 43: 224(2002), m wide, hila unthickened or almost m wide,hilaunthickenedoralmost Sawada,basedon Taiwanese , 14Oct.1928,
85: 112 (1943),nom.inval Liquidambar : OnLiquidambar 5:44(2000). Pseudocercospora
(J.M. Yen)J.M. Yen, Goh &W.H. Hsieh,inHsieh 97:92(1981). 94:52(1978). Cooke &EllisexG.F. Atk., of Liquibambarformosana Liquidambar formosana type: (seebelow). The later C. Nakash.& Tak. Sawada,
VW\UDFLÀXD, 14Oct. C. liquidambaris Taiwan: Taipei,on K. Sawada(NTU- nom. illeg.[ remains unclear.
J.M. Yen, (J.M. Yen)U. ideira, unpubl.). ideira, unpubl.). Cercospora s. Cercospora s. Liquibambar Liquibambar
and is now andisnow Cercospora Cercospora VW\UDFLÀXD, Rep. Gov. . (Art. : 150 Bull. Bull. Braun etal. is is , , from super 151, Fig. 62.Pseudocercosporaliquidambaricola external; super delicately punctiformtosubeffuse, brown. halo, occasionallysomewhatzonate. blackish, narrow, sometimesraised,withdiffuse yellowish greyish browntowhite, withdarkborder, brownto vein-limited, paleolivaceous,browntodarkorlater 1–10 mmdiamorcon Description 139, g.114), Kobayashi g. 113), Guo&Hsieh(1995:125, Illustrations Crous &Braun(2003:253). Guo &Hsieh(1995:123–124), Literature (mainly hypophyllous when developed), branched, 1.5–4 (mainly hypophyllous whendeveloped),branched, 1.5–4 g. 113). : Chupp(1954:259p.p.),Hsieh&Goh(1990:150), cial hyphae. : : Yen (1978a:53, A. Leaf spots Conidiophores fascicles. cial hyphaelackingoralmostso todeveloped C. uent andlarger, angularspotsoften Conidiophores. et al.(2002:224, subcircular toangular-irregular, g. 3),Hsieh&Goh(1990:151, g. 110), Guo Caespituli etal. B. D. Mycelium Conidiophores arising Conidia. Bar=10 (Hsieh &Goh1990: g. 6). (1998: 138–140), amphigenous, amphigenous, IMA FUNGUS etal. internal and internal and (1998: μm.
Cercosporoid fungi 4
μm wide, subhyaline to pale olivaceous-brown, thin-walled, Alabama in 1891, which can be used as lectotype. There is ARTICLE smooth. Stromata small to moderately large, 15–40 μm diam, 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. also solitary, arising from super cial hyphae if present, lateral, Braun, Schlechtendalia 2: 27 (1999). erect, straight, subcylindrical-conical to geniculate-sinuous, (Fig. 63) unbranched, 5–35 × 2–5 μm, 0–2-septate, subhyaline, pale Basionym: Cercospora tuberculans Ellis & Everh., J. Mycol. olivaceous to olivaceous-brown, thin-walled, smooth to 4: 115 (1888). somewhat rough; conidiophores reduced to conidiogenous cells or integrated, terminal, 5–25 μm long, conidiogenous 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., 20–100 × 2–4 μm, 2–10-septate, subhyaline to pale olivaceous 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 truncate, 1–2 μm wide, hila unthickened, not darkened 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). absent to well-developed, 10–80 μm diam, dark brown to blackish, substomatal to immersed, composed of swollen Host range and distribution: On Liquidambar (formosana, hyphal cells, 3–8 μm diam. Conidiophores 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 similar. If they are truly conspeci c or if two morphologically very close, but geographically and genetically distinct species are involved can only be clari ed on the base 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 be con rmed. Chupp (1954), who proposed the original 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) are probably topotypes. Guo & Hsieh (1995) con rmed the 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 can only be clari ed by lectotypi cation. Chupp (1954) interpreted this name in the 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. . 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 μm.
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(see Conidiaconsistentlyacicular, basetruncate,variousmorphologically barelydistinguishablespecies Atleastlongerconidia distinctlyacicularwithtruncatebaseandacuteorsubacuteapex;onother hosts ...... 7 (6) Conidianarrowlycylindricaltosubacicular, apexobtuse,notdistinctly pointed;on 6 (5) Leafspotsdeveloped,distinct;conidiaconsistently acicularand/ornarrower, 1.5–5 (see“Doubtful,excludedandinsuf Leafspotslackingor almostso;conidiaaciculartoobclavate-cylindrical,broad,35–195×(3–)4–6.5(–8) 5 (1) (see“Doubtful,excludedandinsuf Conidiophorespaleolivaceous-brownthroughoutortipspaler, hyalineorsubhyaline;conidianarrowlycylindrical Conidiophoreshyaline oronlypaleolivaceousatthebase;conidiaobclavate-cylindrical,20–80×3.5–7 4 (3) Conidiogenouslocinotcercospora-like,i.e.unthickenedtosomewhatthickened,butdarkened,atmost Conidiogenouslocicercospora-like,i.e.distinctlythickenedanddarkened,infront viewvisibleassmalldarkcircle 3 (2) Stromatadeveloped,10–60 Stromatalackingoralmostso;conidiophoresshort,5–30×3–5 2 (1) Conidiaconsistentlyacicularoratleastlongerconidiaandonlyshorteronessomewhatobclavate- Conidiacylindricalorobclavate-cylindrical...... 1 Key toCercosporaspecieson Amaranthaceae Cercospora 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, 27June1888, Mississippi Lectotype darkened. obconically truncate,2–3 subtruncate toshortobconicallytruncate,rarelylong to brownish,thin-walled,smooth,apexobtuse,base curved, 20–80×4–6.5 to somewhatobclavate-cylindrical,straightslightly unthickened andnotdarkened. loci inconspicuousorvisibleastruncatetips,always conidiogenous cells,10–25 cells integrated,terminalorconidiophoresreducedto olivaceous-brown, thin-walled,smooth;conidiogenous × 3–7μ straight toslightlygeniculate-sinuous,unbranched,10–35 ...... onAchyranthes tosubacicular, 15–90 ×2–4.5 onAchyranthes hyalineorsubhyaline ...... proliferation;conidiophoreshyaline,onlypaleolivaceousatthebaseorolivaceous-brown,tipsoften somewhatrefractive,formedoncharacteristic,bulging,convextipsandshoulderscausedbysympodial withminutecentralpore;conidiophoresdistinctlybrown;on 15–150 0–5-septate;on cylindrical ...... m, rarelylonger, 0–1-septate,paleolivaceousto ( : June1890–1893(BPI442031,442033,442034; : OktibbehaCounty, Starkville,on designated here,MycoBank,MBT202801): Tabular keytoCercospora specieson Acanthaceae accordingtohost genera μm, pluriseptate ...... μ S. M.Tracy (NYIsotypes: 2425377). m, 1–6-septate,paleolivaceous μ spp...... spp...... Celosia m wide,hilaunthickened,not Conidia μ m long,conidiogenous μm diam;conidiophoreslonger, 10–150 s...... sp...... solitary, cylindrical μm; onGomphrena ently knownspeciesof ently knownspeciesof Liquidambar USA: Braun etal. spp...... Nov., tuberculans undoubtedly doesnotpertainto of thiswood-inhabitingfungusisnotquiteclear, butit wood of Guo etal.(1998). They refertoChinesematerialondead unproven andnotincludedinGuo&Hsieh(1995) formosana Notes: RecordsofthisspeciesfromChinaon Missouri, Mississippi). Altingiaceae Host rangeanddistribution Chenopodiaceae in thelatterfamily( Amaranthaceae (The family Amaranthaceae s.str. Cercospora Cercospora μm, 0(–1)-septate;conidiashort,25–35×2.5–3.5 Celosia C. Keissler Liquidambar formosana μm, 0–8-septate;and/orconidialonger, (Keissler &Lohwag1937, Tai 1979)aredoubtful, inPetr., Crypt.Exs.3391[Hunan,Changcha, , North America (USA,Florida,Louisiana, Chenopodiaceae sp...... ”) ”) asseparatefamily, atleasttentatively.) and currentlyproposedtobeincluded &H.Handel-Mazzetti(W)]. The identity s. lat.),butweprefertomaintainthe Gomphrena ...... 6 μm ...... Cercosporella pseudachyranthis ...... 5 ...... LXGPDVW\UDFLÀXD : On/LTXLGDPEDU is phylogeneticallycloseto C. tuberculans ...... 4 ...... ) distributed asCercospora ...... 2 ...... spp., SouthAfrica ...... 3 ...... C. gorakhanathii . C. pretoriensis C. pretoriensis C. achyranthis μm; C. celosiigena μm; . IMA FUNGUS Liquidambar μm, . 7 ,
Cercosporoid fungi 4
Conidia acicular to obclavate-cylindrical, at least shorter conidia obclavate-cylindrical, with obconically truncate base ARTICLE ...... 8
8 (7) Stromata well-developed, large, 20–80 μm diam; conidiophores 10–80 μm long; conidial base 1–2 μm wide; on Alternanthera spp...... C. alternantherae Stromata lacking or 10–50 μm diam; conidiophores to 250 μm long; conidial base 1.5–4 μm wide; on Amaranthus 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 (see “Doubtful, excluded and insuf ciently known species of Cercospora”) Conidiophores brown, distinctly pigmented throughout; conidiogenous loci distinctly thickened and darkened, Cercospora type ...... 2
2 (1) Leaf spots lacking or almost so; conidiophores to 450 μm long; conidia acicular to obclavate-cylindrical, 35–195 × (3–)4–6.5(–8) μm ...... C. achyranthis Leaf spots distinct, well-developed; conidiophores to 250 μm long; conidia usually consistently acicular, much narrower, 25–240 × 2–5 μm ...... C. achyranthina
Aerva A single species ...... C. aervae-lanatae
Alternanthera A single species ...... C. alternantherae
Amaranthus A single species ...... C. brachiata
Celosia 1 Conidiophores short, 5–30 × 3–5 μm, 0(–1)-septate; conidia short, 25–35 × 2.5–3.5 μm, cylindrical to obclavate-cylindrical, 0–5-septate, hila 1–2 μm wide ...... C. celosiigena Conidiophores much longer, 20–220 μm, 0–8-septate; conidia much longer, 15–150 μm, (0–)1–14-septate, hila 1.5–3 μm wide ...... 2
2 (1) Stromata lacking or almost so; conidia acicular ...... C. celosiae Stromata developed, 10–40 μm diam; conidia cylindrical or obclavate-cylindrical ...... 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 1 Stromata lacking or small, 10–35 μm diam; conidia acicular, 30–450 μm long, apex pointed; mainly on Gomphrena globosa, Northeast Africa, Asia, North America, West Indies ...... C. gomphrenae Stromata 15–60 μm diam; conidiogenous loci not cercospora-like (neither distinctly thickened, nor darkened), conidia narrowly cylindrical to subacicular, apex obtuse; on Gomphrena spp., South Africa ...... C. pretoriensis (see “Doubtful, excluded and insuf ently known species of Cercospora”)
Pupalia A single species ...... C. achyranthina
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irregular, 0.5–6.5mmdiam,occasionallycon Description 27, g.9). 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) μm. A. Fig. 64.Cercosporaachyranthina Conidiophore fascicle. Mycologia : Chupp(1954:29), : : Ellis(1976:243, Leaf spotsamphigenous,circulartoangular- 40:352(1948). B. Conidiophore tip.C. asudeva (1963:31),Ibrahim& g. 183B),Guo Thirum. &Chupp, (BPI 4323844,holotype). et al.(2005:27),Kamal Conidia. Bar=10 uent andlarger, et al.(2005: Braun etal.
cells integrated,terminalandintercalary, 10–40 or brownish,wallsomewhatthickened,smooth;conidiogenous 15–250 ×3–8μ geniculate-sinuous, aboveallintheupperhalf,unbranched, stromata, throughstomataorerumpent,erect,straighttocurved, in fascicles,2–10,divergent,arisingfromswollenhyphalcellsor 25 dark. Achyranthes Host rangeanddistribution Topotype aspera Holotype truncate, 2–3.5 subobtuse, basetruncate,occasionallyslightlyobconically 3–10-septate, hyaline,thin-walled,smooth,apexacuteor obclavate-cylindrical, (25–)35–150(–240)×2–5 Conidia conidiogenous locithickenedanddarkened,2–3.5 purplish. grey orgreyishwhite,marginreddishbrowntopurple-violet, to 10mmdiam,browngreyishbrown, is maintainedanddepositedasCUP 37201. This specimen,whichcanbeconsideredtopotypematerial, mentioned acollectionfrom2Sep.1945astypematerial. collection from1944astype.Chupp(1954)erroneously complex. Notes: This speciesispartof theCercosporaapiis.lat. ( Bengal; Myanmar, Pakistan,Philippines),South America Pradesh, MadhyaMaharashtra,NewDelhi,West imbabwe), Asia (China;India,Bihar, Karnataka, Andra [atropurpurea so. septate, somewhatpigmented. dark greyorbrownish. hypophyllous, effuse, occose-velutinuous,dark,greyish to sometimes vein-limited,greyish, margininde only diffuse discolorations,greenishtobrownish, Description Exs. 546. Exsiccatae: Cif.,Myco 28, g.10). Illustrations Guo etal.(2005:28–29),Kamal(2010:12). (1965: 8),Shin&Kim(2001:24),BraunCrous(2003:42), (1937: 222),Chupp(1954:30), Literature (Fig. 65) Cercospora achyranthis nally very dark, sometimes with diffuse darker halo, often nally verydark,sometimeswithdiffuse darkerhalo,often enezuela). μ Conidiophores 7: 171(1909). m diam,substomataltoimmersed,brown. Mycelium , 20 Aug. 1944,M.J.Thirumalachar solitary, acicular, shorterconidiaoccasionallyslightly : Caespituli
: 2Sep.1945(CUP 37201). : Saccardo(1913:1429), Thirumalachar &Chupp(1948)citedasingle India : : Shin&Kim(2001:25, Leaf spotslackingoralmostso,indistinct ]. Amaranthaceae sp.), μ internal. m wide,hilathickenedanddarkened. : m, (0–)1–10-septate, pale olivaceous-brown m, (0–)1–10-septate,paleolivaceous-brown in small,loosefascicles, 2–12,arisingfrom Karnataka amphigenous, scattered, Digera muricata,Pupalialappacea . Doming.Exs.147.Syd.,FungiExot. Mycelium Stromata : Bangalore,on Syd. &P. Syd.,Ann.Mycol : OnAchyranthes , Africa (Sudan, Tanzania,(Sudan, , Africa Stromata asudeva (1963:31),Katsuki almost lackingorsmall,10– internal; hyphaebranched, g. 1),Guo assiljevsky &Karakulin nally sometimes dull nally sometimesdull lacking oralmost nite. nely punctiform, nely punctiform, (BPI 4323844). Conidiophores Conidiophores IMA FUNGUS Achyranthes et al.(2005: μ m, usually m, usually Caespituli μ μ (aspera m diam. m diam. m long, m long, nally . ,
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 truncate conidial bases are not con ned to shorter, young 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: Raghu Ram & Mallaiah (1996: 296, g. 2), Srivastava et al. (2001: 102, g. 1).
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 well-developed, gobose, 20–40 μm diam, brown to black. Conidiophores in loose fascicles, to 15, arising from stromata, through stomata, erect, subcylindrical to 1–4 times geniculate, unbranched, about 45–230 × 3.5–5.5 μm, 2–11-septate, pale olivaceous to brown, paler towards Fig. 65. Cercospora achyranthis (LEP, isolectotype). A. Conidiophore the tip, thin-walled, smooth; conidiogenous cells integrated, fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 μm. terminal and intercalary, conidiogenous loci thickened and darkened, 2–3.5 μm diam. Conidia solitary, acicular, straight to curved, 55–160 × 1.5–4 μm, 5–15-septate, hyaline, thin- substomatal swollen hyphal cells, through stomata, erect, walled, smooth, apex pointed, base truncate, 1.5–3 μm wide, straight to usually distinctly geniculate or geniculate-sinuous, hila thickened and darkened. often strongly geniculate, unbranched, 40–450 × 3.5–8 μm, 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). and intercalary, about 10–30 μm long, conidiogenous loci conspicuous, thickened and darkened, 1.5–2.5 μm diam. Host range and distribution: On Aerva (lanata, sanguinolenta Conidia solitary, acicular to obclavate-cylindrical, straight to [scandens]), Amaranthaceae, Asia (India, Andhra Pradesh, curved, 35–195 × (3–)4–6.5(–8) μm, 3–18-septate, hyaline Uttar Pradesh). or almost so, thin-walled, smooth, apex acute to subobtuse, base truncate to short obconically truncate, 1.5–4 μm wide, Note: This species is morphologically assignable to the hila thickened and darkened. Cercospora apii s. lat. complex.
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straight, subcylindrical-conical to oblong, geniculate-sinuous, straight, subcylindrical-conical tooblong,geniculate-sinuous, arising fromstromata,through stomataorerumpent,erect, developed, mostlyratherlarge fascicles,divergenttodense, immersed, large,20–80 to blackish. Caespituli 0.5–3 mmdiam,palegreenishtodingygrey, margin brownish. Description Illustration Crous &Braun(2003:53),Kamal(2010:15). Literature Cercospora alternantherae (Fig. 67) A. Fig. 66.Cercosporaaervae-lanatae 10 μm. Conidiophore fascicle. 6: 36(1890). : Saccardo(1892:637),Chupp(1954:30–31), : Chupp(1954:31, : amphigenous, punctiform, scattered, dark brown amphigenous, punctiform,scattered, darkbrown Leaf spotsamphigenous,circularorsubcircular, Mycelium μ B. m diam,brown. internal. Conidiophore tips.C.Conidia.Bar= g. 7).
Ellis &Langl., (K(M) IMI351224,holotype). Stromata Conidiophores substomatal to substomatal to J. Mycol. in well- Braun etal.
Jul. 1888, Louisiana Lectotype μ pointed orsubobtuse,basetruncatetoobconicallytruncate,1–2 125 ×2–4 solitary, obclavate-cylindrical,acicular, straighttocurved,20– thickened anddarkened,(1–)1.5–2(–2.5) intercalary, 10–30 smooth; conidiogenouscellsintegrated,terminal,occasionally pale olivaceoustobrown,palertowardsthetip,thin-walled, unbranched, 10–80×3–6 10 A. Fig. 67.Cercosporaalternantherae Host rangeand distribution BPI 432464. m wide,hilathickenedanddarkened. Conidiophore fascicles. μm. : St.Martinsville,on μ (designated here,MycoBank,MBT202806): A. B.Langlois1430 m, 0–10-septate, hyaline, thin-walled, smooth, apex m, 0–10-septate,hyaline,thin-walled,smooth,apex μ m long, conidiogenous loci conspicuous, m long,conidiogenouslociconspicuous, B. Conidiophore tips. : OnAlternanthera μ Alternanthera achyrantha m, 1–4-septate, subhyaline, m, 1–4-septate,subhyaline, (NY 830163). (NY 830163,lectotype). μ Conidia m diam.Conidia C. Conidia.Bar= IMA FUNGUS Isolectotype: (¿FRLGHD USA: , 18 ,
Cercosporoid fungi 4
halimifolia, crucis [portoricensis], pungens [achyrantha, ARTICLE repens]), Amaranthaceae, North America (USA, Louisiana, Texas), South America (Brazil, enezuela), West Indies (Cuba, Jamaica, Puerto Rico, irgin Islands).
Notes. This is a true Cercospora s. str. close to C. apii s. lat., but distinct by having large stromata (20–80 μm diam) with numerous, densely arranged conidiophores, smaller conidiogenous loci (1–2 μm wide), and obclavate-cylindrical to acicular conidia. Indian records of this species from Uttar Pradesh and West Bengal on A. sessilis (Kamal 2010) are doubtful and in need of revision and con rmation. They might rather pertain to the illegitimate species 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. μm. 1925, A. I. Lobik (not preserved)].
Literature: Saccardo (1892: 637), assiljevsky & 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 μm diam, olivaceous-brown to brown, cells 2–6 μm diam. et al. (2005: 29–30), Kamal (2010: 25). Conidiophores fasciculate, 2–12(–20), rarely solitary, arising from swollen hyphal cells or stromata, emerging Illustrations: Tai (1948: 37, g. 2), Guo et al. (2005: 29, g. through stomata or erumpent, erect, straight, subcylindrical 11), Pirnia et al. (2010: 185, g. 1). to usually distinctly geniculate or geniculate-sinuous, above all in the upper half, unbranched or tips occasionally Exsiccatae: Ellis & Everh., N. Amer. Fung. 2582. furcate, 20–250 × 3–6.5 μm, usually 1–8-septate, very long conidiophores sometimes with additional septa, pale Description: Leaf spots amphigenous, circular, subcircular to medium dark brown or olivaceous-brown throughout or to somewhat angular-irregular, 0.5–12 mm diam, at rst 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, nally with a tan, grey to greyish white centre surrounded 10–40(–70) μm long, conidiogenous loci conspicuous, by a brown, reddish to purplish brown or almost blackish thickened and darkened, 1.5–3.5 μm diam. Conidia margin. Caespituli amphigenous, mostly hypophyllous, solitary, acicular, shorter conidia may also be narrowly punctiform scattered, dark. Mycelium internal. Stromata obclavate-cylindrical, straight to curved, 25–250 × 1.5–5(–
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in 1890. material (formersyntypes)fromthetypelocality, butcollected G U, NY, PHandnumerousotherherbaria)isauthentic (Fig. 69) Cercospora celosiae (Pirnia & Braun2004),Indonesia(Shivas included inCrous&Braun(2003)refertoGermany(Jage former treated assubgenus,whichre genus The hostsof undoubtedly in 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 ofC.brachiata Type materialofthisspeciesisprobablynotmaintained. for typematerialof is morphologicallybarelydistinctfrom do withacomplexofcrypticspecies. if allcollectionspertaintoasinglespeciesorwehave different continentsareinvolved. Thus, itremainsunclear Many hostspeciesandawidedistributionrangecovering conidiophores aswelllengthandwidthofconidia. rather variable,aboveallwithregardtothelengthof lat. complex. Notes: This speciesbelongs to the Islands). Republic, Haiti,PuertoRico, Trinidad and Tobago, Texas, Wisconsin),West Indies(Barbados,Cuba,Dominican North America (USA,Delaware,Florida,Illinois,Nebraska, and South America (Brazil,Ecuador, Panama, Iran, Pakistan),Europe(Germany, Russia,Ukraine),Central Orissa, Maharashtra,UttarPradesh,West Bengal;Indonesia, Uganda), Asia (Brunei,China;India, Andhra Pradesh,Delhi, Amarathus polygamus 6) Host rangeanddistribution NY 24083120,2408321. Aug. 1887, USA: Lectotype hila thickenedanddarkened. or obconicallytruncateinshorterconidia,1.5–4 acute orsubacute,sometimessubobtuse,basetruncate [gangeticus [paniculatus blitoides japonica μ Ellis &Everh.,N. Amer. Fung.2582 (BPI 432395,FH, (1929). m, 1–20-septate,hyaline,thin-walled,smooth,apex Delaware et al.2010,Hesami Acnida Acnidia , cannabinus ] (Crous&Braun2003)aredoubtfulandbelong : Chupp(1954:32), sp.), UHWURÀH[XV , ], tuberculatus (designated here,MycoBank,MBT202808): A. Commons626 , C. achyranthina hybridus C. acnidae andAmaranthus , nowasynonymofAmaranthus Cercospora brachiata : Faulkland,on uenced byassumedhostrangedifferences. Amaranthaceae Cercospora amaranthi , subsp. , spinosus , viridis[blitum
werepreviouslyassignedtothe caudatus Syd., : On et al.2011). onAchyranthesbidentata . Somenewerrecordsnotyet (NY 2408319). cruentus asudeva (1963:76),Katsuki , Africa (Kenya,South Africa, PUQKV UHWURÀH[XV $PDUDQWKXV Amaranthus species. ects thecloseaf Ann. Mycol. , crassipes , tamariscinus et al Cercospora acnidae subsp. ], dubius Cercospora apiis. is morphologically . 1996)andIran A carefulsearch C. brachiata inLEfailed. (albus Isolectotypes: emarginatus ,
, hybridus enezuela), μ 27: 430 , cruentus usually m wide, , tricolor nity of blitum irgin , 18 Braun etal. et ], . , , integrated, terminal andintercalary, conidiogenousloci towards thetip,thin-walled, smooth;conidiogenouscells 0–6-septate, paletomedium brown,palerandnarrower usually unbranched,geniculate-sinuous, 20–220×3–6 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, occasionallysomewhatraised, subcircular, 1–12mmdiam,occasionallylarger, tanto Description 17, g.6),Guoetal.(2005:31, Illustrations al. (2005:30–31),Kamal(2010:31). fascicle. B.Conidiophoretip. Fig. 69.Cercosporacelosiae(BPI434404,holotype). : Ellis(1976:243, : Leaf spots C. Conidia.Bar=10μm. g. 183C),Hsieh&Goh(1990: amphigenous, circularor Mycelium g. 12). internal. Conidiophores Caespituli A. IMA FUNGUS Conidiophore Stromata usually μ m, in Cercosporoid fungi 4
conspicuous, thickened and darkened, 1.5–3 μm diam. ARTICLE Conidia solitary, acicular, straight to curved, 25–150 × 2–4(– 4.5) μm, hyaline, 2–12-septate, apex acute, base truncate to somewhat obconically truncate, 1.5–3 μm wide, hila 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, enezuela), 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 (con rmed by Ch. Nakashima).
Cercospora celosiigena U. Braun & Bagyan., sp. nov. MycoBank MB814563 (Fig. 70)
Literature: Bagyanarayana et al. (1991: 324).
Illustration: Bagyanarayana et al. (1991: 321, g. 6).
Diagnosis: Differs from C. celosiae in having much shorter conidiophores, 5–30 × 3–5 μm, 0(–1)-septate, and short, narrowly obclavate-cylindrical conidia, 25–35 × 2.5–3.5 μm, 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. Conidia. Bar = 10 μm. irregular, 1–3 mm diam, somewhat raised, greyish brown to greyish white, with a narrow purplish margin. Caespituli hypophyllous, scattered, ne, dark brown. Mycelium 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. or slightly geniculate-sinuous, unbranched, 5–30 × 3–5 μm, 0(–1)-septate, pale olivaceous to olivaceous-brown, thin- Notes: Bagyanarayana et al. (1991) identi ed the type 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. integrated, terminal, 5–25 μm long, conidiogenous loci However, C. celosiae belongs to the C. apii s. lat. complex. conspicuous, thickened and darkened, 1–2 μm diam. Conidia The conidia are consistently acicular, and the conidiophores solitary, narrowly obclavate-cylindrical, straight to slightly are much longer and septate. curved, 25–35 × 2.5–3.5 μm, 0–5-septate, hyaline, thin- 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). ones, 1–2 μm wide, hila slightly thickened and darkened. (Fig. 71)
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μm. Conidiophore fascicle.B. Fig. 71.Cercosporagomphrenae Conidiophore tips. (CUP 33132,holotype). C. Conidia.Bar=10 Braun etal. A. Iran, Nepaland Taiwan (Crous&Braun2003). W.W. Ray Holotype and darkened. acute tosubobtuse,basetruncate,2–4 × 2–5 Conidia conspicuously thickenedanddarkened,2–3.5 terminal andintercalary, 10–30 tip, thin-walled,smooth;conidiogenouscellsintergrated, 3–10-septate, paletomediumbrown,palertowardsthe geniculate-sinuous, unbranched,30–300×3–7 or erumpent,erect,straight,subcylindricaltodistinctly from internalswollenhyphaeorstromata,throughstomata Conidiophores diam, substomataltoimmersed,brown,cells2–5 pale, 2–3.5μ Mycelium Caespituli brown, yellowishreddishbrowntopurplishbrown. brown, laterdingygreytogreyishwhite,margindarker, angular-irregular, 1–10mmdiam,at Description 32, g.13). Illustrations et al.(2005:31),Kamal(2010:47). Crous &Braun(2003:201),ShinKim(2001:67–69),Guo Literature Synonym Cercospora gorakhanathii gomphrenaea complex. Duetoconfusionwiththename Notes: This speciesispartof theCercosporaapiis.lat. Oklahoma, Texas), West Indies (Cuba). India, Andhra Pradesh;Korea),North America (USA,Georgia, [decumbens Host rangeandditribution 39900). Isotype: K(M)IMI168993. Gomphrena globosa Illustrations Literature (Fig. 72) and larger, lightbrowntodingygrey. irregular, sometimesvein-linited,2–8mmdiam orcon Description (2003: 271, g.2). Brit. Mycol.Soc.89:124(1987). Phytopathol H. D.Bhartiya Pradesh μ m, 3–20-septate, hyaline, thin-walled, smooth, apex m, 3–20-septate,hyaline,thin-walled,smooth,apex solitary, acicular, straighttocurved,30–300(–450) : : Cercosporacelosiicola : Chupp(1954:34),Ibrahim&ElNurElamin(1974), : Kamal(2010:31). internal; hyphae branched, septate, subhyaline or internal; hyphaebranched,septate,subhyalineor are undoubtedlywrongordoubtful,e.g.thosefrom amphigenous, punctiform, dark brown, scattered. amphigenous, punctiform,darkbrown,scattered. : : : : Rai&Kamal(1987:125, g.1),Bhartiya et al. USA: : Basti,onCelosia ]), amphigenous, circular to somewhat Leaf spotsamphigenous,circulartosomewhat Shin &Kim(2001:68, g.25),Guoetal.(2005: Leaf spots m wide. Sawada,severalrecordsof Amaranthaceae . in small, divergent fascicles, 3–15, arising in small,divergentfascicles,3–15,arising 56: 271(2003)[ (HCIO 42689);isotype:GPU8026]. Oklahoma , 18 Aug. 1942,W. W. Ray Stromata amphigenous, circulartoangular- : OnGomphrena Topotype : sp. (as“coronata , Africa (Sudan), Asia (China; Stillwater, Ray’s yard,on A.N. Rai&Kamal, μ almost lackingor10–35 m long, conidiogenous loci m long,conidiogenousloci
Bhartiya : from8 Aug. 1942(CUP Caespituli holotype μ m wide, hila thickened m wide,hilathickened rst brownish, greyish rst brownish,greyish (globosa C. gomphrenae et al., : (CUP 33132). amphigenous, ”), Nov. 1997, IMA FUNGUS India Cercospora μ m, usually m, usually μ μ m diam. m diam. , serrata m diam. m diam. : Trans. Indian uent Uttar μ m
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. (1999: 63, g. 3).
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 to light olivaceous, about 2.5 μm wide. Stromata well- developed, substomatal, about 25 μm diam, pale olivaceous to olivaceous. Conidiophores solitary or in fascicles of 2–6, arising from stromata, through stomata, about 15–130 × 2–4.5 μm, cylindrical, straight to slightly 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, straight to curved, about 35–210 × 2–4 μm, 5–27-septate, 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. μm. Narayan (GPU 5023). Isotype: HCIO 41940.
Host range and distribution: On Alternanthera sessilis, nely punctiform, effuse, dark dingy olivaceous to brown. Amaranthaceae, Asia (India). Mycelium internal. Stromata developed, substomatal to immersed, 10–40 μm diam, medium to dark olivaceous. 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. unbranched, rarely branched, 20–150 × 3–6 μm, 1–8-septate, pale, subhyaline to pale olivaceous, thin-walled, smooth; Cercospora crassoides Davis, Trans. Wisconsin Acad. conidiogenous cells integrated, terminal or occasionally Sci. 21: 298 (1924). intercalary, conidiogenous loci conspicuous, 1.5–2 μm 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 curved, 15–150 × 1.5–4.5 μm, (0–)1–14-septate, hyaline, Literature: Chupp (1954: 33), Crous & Braun (2003: 143). thin-walled, smooth, apex subacute or subobtuse, base truncate to somewhat obconically truncate, 1.5–3 μm wide, Illustration: Simmons (1995: 147, g. 117). hila thickened and darkened.
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diagnosis ordescription. globosae clearly designated.Furthermore,thename from twoyears,1993and1994,butaholotypewasnot type materialwasnotavailable. The authorscitedcollections subcylindrical conidia,butavalidationisnotmadesince Notes: This speciesiswell-characterised byhaving Amaranthaceae Host rangeanddistribution 41978). 1993/1994, Pradesh Syntypes truncate, thickenedanddarkened. hyaline, thin-walled,smooth,apexsubacutetoobtuse,base curved, about13–205×2–3 (somewhat attenuatedtowardsthetip),straighttosomewhat whitish grey, marginred. Description Illustration Literature Cercospora nothosaervae and darkened.Conidia and intercalary, conidiogenouslociconspicuous,thickened walled, smooth;conidiogenouscellsintegrated,terminal × 3.5–5 unbranched, cylindrical,1–4timesgeniculate,about45–155 3–7, arisingfromstromata,erect,straightoralmostso, to brown.Conidiophores subepidermal, about10–30 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. , inRao : Gorakhpur, on , 23Jul.1921,J.Davis(WIS). Mycelium : isinvalidsinceitwaspublishedwithoutanyLatin μm, 2–7-septate,olivaceoustolightbrown,thin- : Crous&Braun(2003:292), Kamal (2010:69). : Crous&Braun(2003:2001),Kamal(2010:47). : Patil(1975:220, : Raoetal.(1999:70, g.6). : (holotype : USA: circular 1–2 mm diam, centre depressed, Leaf spotscircular1–2mmdiam,centredepressed, S. Narayan Leaf spots : 69(1999), , Asia (India,UttarPradesh). , North America (USA,Florida,Oklahoma, et al.,SugarcanePathology Wisconsin
internal; hyphaebranched,septate, Stromata not clearlyindicated): solitary, subcylindricaltosubacicular amphigenous, circularorsubcicular, (GPU5051,5061;HCIO41968, Caespituli solitary orinsmallfascicles, g. 9).
: LoneRock,on nom. inval M.S. Patil, Gomphrena globosa : On : On μm diam,lightolivaceous well-developed, compact, μm, usually7–18-septate, Caespituli amphigenous. 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 etal. et , , internal. Cercospora barely con Caespituli reddish tored-brownborder, onthelowersurfacepaler. the uppersurfacecentrepalebrowntodingygrey, with subcircular, 2–6mmdiam,sometimescon Description Crous &Braun(2003:335). Literature darkened, atmost somewhatrefractive. wide, hilaunthickenedoronly veryslightlythickened,not subtruncate orslightlyshortobconically truncate,1–2.5 walled, smooth,apexobtuse orsubobtuse,basetruncate, × 2–4.5μm,1-topluriseptate,hyaline orsubhyaline,thin- occasionally subacicular, straighttoslightlycurved, 15–90 somewhat darkerrim. refractive, infrontviewsometimesvisibleasminutecirclewith slightly thickened,nottodarkened,oftensomewhat sympodial proliferation(lightmicroscopy),unthickenedoronly lateral viewasconvextotruncatetiporshouldercausedby conspicuous orsubconspicuous,1.5–2 to conidiogenouscells,10–25 cells integrated,terminal,sometimesconidiophoresreduced pigmented throughout,thin-walled,smooth;conidiogenous paler towardsthetip,upperpartsubhyalineorsometime 0–3-septate, rarelypluriseptate,paleolivaceous-brown, often stronglygeniculate-sinuous,10–50(–100)×3–6 the tip,unbranched,straighttocurvedorusuallydistinctly, erumpent, erect,subcylindrical,somewhatattenuatedtowards densely fasciculate,fewtonumerous,arisingfromstromata, angular-subglobose, about3–9 olivaceous-brown, composedofswollenhyphalcells, Stromata Holotype prominent hilum,darkenedandthickened. brown, apexsubobtuse,baseobconicallytruncate,witha μ almost rostrateapex,straighttoslightlycurved,20–70×5–6.5 prominent. brown, palertowardsthetip;conidiogenouslociconspicuous, cells swollen,20–70×5–5.5 non-geniculate, unbranchedoroccasionallybranched,basal loose groups,2–3,erumpent,erect,straightto (Fig. 73) was notavailable. was notpossibletocon cercosporoid. Itseemstobeaspeciesof Notes: The generalcharactersofthisfungusarenot collections. Host rangeanddistribution 31692). brachiata m, 1–5-septate,constrictedatthesepta,paleolivaceous- 4: 890(1948). Stromata : , Amaranthaceae, : Chupp(1954:35),Crous&Braun(1996:298), hypophyllous,scatteredtodense,punctiform, immersed, 15–60 India Conidia : uent, darkolivaceous.Mycelium Leaf spots
pretoriensis : Maharashtra lacking. solitary, broadlyobclavatewithattenuated, Conidia rm thisassumption. Type material amphigenous, irregularlyscattered, Conidiophores 17 Dec.1973, μm diam,somewhaterumpent, : Onlyknownfromthetype Chupp &Doidge, : Kolhapur, on μ μm long,conidiogenousloci solitary, narrowlycylindrical, m, 1–2-septate,olivaceous- μm diam.Conidiophores μm diam,visiblein solitary or in small, solitary orinsmall, M. S.Patil(HCIO Alternaria Nothosaerva IMA FUNGUS uent, on Bothalia internal. exuous, exuous, , butit μm, μm
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 are required to elucidate the correct generic af nity of this 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: Castañeda & Braun (1989: 44, pl. 1, g. 1).
Description: Leaf spots amphigenous, subcircular to somewhat irregular, 0.5–4 mm diam, yellowish, ochraceous to brownish, margin inde nite or narrow and darker. Caespituli hypophyllous, punctiform-effuse, greyish white. Mycelium internal. Stromata well-developed, substomatal, subglobose, 20–40 μm diam, brownish, composed of swollen hyphal cells, 2–8 μm diam, subglobose-angular. 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. erect, straight to exuous, curved, in the upper part slightly Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 μm to mostly distinctly geniculate-sinuous, simple, rarely branched, (40–)50–200(–220) × (2–)3–5(–6) μm, septate, 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, cells integrated, terminal, 10–40 μm long, conidiogenous loci 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 somewhat refractive, 1.5–2.5 μm diam. Conidia solitary, Africa. obclavate-cylindrical, straight to slightly curved, 20–80 × 3.5–7 μm, 1–8-septate, hyaline or subhyaline, thin-walled, Notes: Cercospora pretoriensis is a cercosporoid species smooth, apex obtuse, base obconically truncate to rounded, that cannot be unequivocally assigned to any of the 1.5–2 μm wide, hila almost unthickened or only very slightly 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
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μm. aspera Conidiophore fascicle.B. Fig. 74.Cercosporellapseudachyranthis Host rangeanddistribution Isotype: HAL 1648F. Paratypes 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: The morphologicalcharacters ofthisspecieshave (Pakistan), West Indies(Cuba,PuertoRico, , 22Dec.1987, , asperavar. indica , Achryranthes Cercospora Passalora 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 irgin Islands). Cercosporella (aspera , ?Asia . Type Braun etal. var. A. ],
Holotype μm, septate,hyalinetoolivaceous. above. 96.6 ×3.7μm,darkerbrownbelow, palerorsubhyaline 4–10, arisingfromstromata,divergent,unbranched,18.5– 10–30 brown, marginbrowntoblack,mostlyepiphyllous. Description Illustration Literature Cercospora pupaliae on theupper surface becomingchlorotic, 4–8mmdiam, of typematerial):Leafspots amphigenous, greenishbrown, Description Illustration Literature Cercospora sessilis known insometropical-subtropicalspeciesof above allinthelowerhalf,butthiskindofpigmentationis 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). of revision. pupaliae shows themtobeobclavate.Hence,thegenericaf 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: Type materialofthisspecieswasnotavailable. The collection. Host rangeanddistribution Amaranthaceae ., i.e.basedontheoriginaldescriptionandillustration,this Sydowia non Mycol. Appl Chupp (1954)introducedthenewspeciesname μm diam,darkbrown. Conidia C. sessilis isnotquiteclearandthisspeciesinurgentneed : : Crous&Braun(2003:344),Kamal(2010:79). : Crous&Braun(2003:375),Kamal(2010:85). : Patwardhan&Pande(1970:99, : Pavgh&Singh(1964:192, India : (based onPavgh&Singh1964 andexamination C. centrostachydis 23:98“1969”(1970);as“ Leaf spots : , 23 Aug. 1966,G.P. Patwardhan(AMH294). . solitary, Maharashtra 23: 190(1964),nom.illeg. Sorokin, 1892. Pavgi &U.P. Singh,
obclavate, straight,51.8–70×3.7 circular toirregular, greytodirty C. achyranthis Patwardhan & A.K. Pande, : Onlyknownfromthetype : Pune,onPupaliaorbiculata with referencetoSolheim& Conidiophores fromPakistan(Ahmad C. pupaliae gs 10–12). ), butfailedtoadd pupalae . Heemphasized g. 6). Cercosporella in fasciclesof Mycopathol. Passalora s. IMA FUNGUS (Art. 53.1), C. apii nity ofC. ”. Stromata et al. , but ,
Cercosporoid fungi 4
con uent. Stromata substomatal, 10–25 μm diam, pale Notes: Status and generic af nity unclear, type material 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 for a nal conclusion. According to the original publication, from stromata, erect, straight, subcylindrical to somewhat syntype material has been deposited in HCIO, but an geniculate-sinuous, unbranched, 25–90 × 3–4 μm, continuous 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 × 1.5–3 μm, 3–11-septate, hyaline or subhyaline, thin-walled, conidium which seemed to have a slightly thickened and smooth, apex subacute, base short obconically truncate. darkened hilum.
Syntypes: India: Uttar Pradesh: aranasi, on Alternanthera sessilis, Amaranthaceae, 22 Oct. 1961, U. P. Singh (HCIO, K(M) IMI 113095).
Passalora
Key to Passalora species on Amaranthaceae 1 Conidia formed singly, obclavate-cylindrical, 30–85 × 3.5–6 μm, (1–)3–7-septate, pale olivaceous; mycelium internal; conidiophores fasciculate, arising from well-developed stromata, 25–75 μm diam; on 3IDI¿D sericea ...... 3SIDI¿DH Conidia at least partly catenate and/or super cial mycelium with solitary conidiophores formed and stromata lacking ...... 2
2 (1) Super cial hyphae with solitary conidiophores formed; stromata lacking (mycovellosiella-like species) ...... 3 Super cial hyphae and solitary conidiophores lacking; conidiophores fasciculate; stromata developed (phaeoramularia-like species) ...... 4
3 (2) Conidiophores relatively long, 10–200 × 3–7 μm, longer conidiophores pluriseptate; on Cyathula achyranthoides ...... P. cyathulae Conidiophores shorter, 5–30 × 3–5 μm, 0–1-septate; on Iresine spp...... P. gonoclada
4 (2) Conidia usually cylindrical or subcylindrical, hyaline or subhyaline, 20–60 × 2–5 μm, 0–5-septate; conidiophores narrow, 1.5–5 μm wide; on Iresine diffusa ...... P. gilbertii Conidial shape variable, cylindrical, fusiform to obclacate, olivaceous to olivaceous-brown, 3–8 μm wide; conidiophores broader, 2–8 μm ...... 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 1 Stromata lacking; conidiophores solitary, arising from super cial hyphae, 5–30 μm long, 0–1-septate ...... P. gonoclada Stromata developed; super cial hyphae and solitary conidiophores absent; conidiophores fasciculate, 20–60(–110) μm long, 0–5-septate ...... 2
2 (1) Conidia usually cylindrical or subcylindrical, hyaline or subhyaline, 20–60 × 3.5–6 μm; conidiophores narrow, 1.5–5 μm wide, unbranched ...... P. gilbertii Conidial shape variable, cylindrical, fusiform to obclacate, olivaceous to olivaceous-brown, 3–8 μm wide; conidiophores broader, 4–6(–8) μm, occasionally branched ...... P. iresines
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Conidiasolitary, obclavate-cylindrical,30–85×3.5–6 Illustration Literature Basionym Passalora cyathulae Passalora specieson Amaranthaceae Conidiainchains,ellipsoid,ovoid,broadlyfusiform,subcylindrical,(10–)15–60×3–8 1 3IDI¿D (Fig. 75) Holotype collection. Host rangeanddistribution Amaranthaceae super effuse, darkolivaceous. brown below, margininde irregular spots,0.5–2mmdiam,brownabove,olivaceous- discolorations orsmallcircular, subcirculartosomewhat Description walled, smooth. septate, 2–5 hila somewhatthickenedanddarkened. short obconicallytruncatetorounded,about2 subhyaline topalebrownish,thin-walled,smooth,ends ellipsoid tocylindrical,20–65×3–7.5 sometimes branchedchains,straighttosomewhatcurved, and darkened,(1.5–)2(–2.5) 10–30 sometimes conidiophoresreducedtoconidiogenouscells, conidiogenous cellsintegrated,terminalorintercalary, throughout, subhyalinetopalebrown,thin-walled,smooth; variable, 10–200×3–7 to geniculate-sinuous,unbranchedbranched,length decumbent conidiophoresdif conidiophores arisingfromsuper erect todecumbent(differentiation betweensolitary arising fromsuper Chupp’s (1954) descriptionof new speciesbasedon described anewIndianspecies butdidnotintroduceda which is,however, incorrect.Sydow(in cited “Cercosporacyathulae cyathulae confused and conidialhila,conidiaformedinchains.Chupp(1954) conidiophores, thickenedanddarkenedconidiogenousloci Passalora Notes: This speciesisatypicalmycovellosiella-like & Crous,Mycosphaerellaand Anam. Mycologia cial hyphaeemergingthroughstomata,branched, μ ...... m long,conidiogenouslociconspicuous,thickened : : Chupp(1954:33). Ragnhildiana cyathulae : Ragnhildianacyathulae specieswithsuper Guyana: Coverden,on : Solheim&Stevens(1931:403, described bySydow(1937). Furthermore,he : 23:403(1931). Leaf spotslackingorindistinct,yellowish μ , m wide,subhyalinetopalebrown,thin- 4 Aug. 1922, 4 Aug. Stromata cial hyphae,lateralorterminal,
(F. Stevens&Solheim)U.Braun Ragnhildiana cyatheae Mycelium lacking. μ nite. : Onlyknownfromthetype μ m, aseptatetopluriseptate (F. Stevens &Solheim)Syd.” F. L.Stevens743(ILL 11981). m diam.Conidia cult), straight,subcylindrical C. cyathulae cial hyphaewithsolitary and theIndian Cyathula achyranthoides Caespituli Conidiophores F. Stevens&Solheim, cial hyphaeandlong internal andexternal; μ m, 0–4-septate, g. 12). : 148(2003) was basedon hypophyllous, in simpleor . Moreover, Cercospora et al.1937) μ m wide, solitary, Braun etal. , μm, (1–)3–7-septate ...... C. Conidiophore. hyphae. Fig. 75. length andseptation. readily distinguishablebyobviousdifferences intheconidial characters ofbothspecies,althoughthetwospeciesare B. Solitaryconidiophoresarisingfrom super Passalora cyathulae D. Conidia.Bar=10 ...... (ILL 11981, holotype). μm, (0–)1–4(–5)-septate μm. P. gomphrenicola A. cial hyphae. IMA FUNGUS Super 3SIDI¿DH cial 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) developed, 20–60 μm diam, subglobose, immersed, brown. Basionym: Cercospora gilbertii Speg., Anales Soc. Ci. Argent. Conidiophores numerous, in loose to very dense fascicles, 10(1): 32 (1880). arising from stromata, erumpent, erect, liform, exuous, Synonym: Phaeoramularia gilbertii (Speg.) U. Braun, somewhat geniculate-sinuous, unbranched, 20–60 × Schlechtendalia 2: 11 (1999). 1.5–5 μm, aseptate to pluriseptate throughout, subhyaline to pale olivaceous, thin-walled, smooth; conidiogenous Literature: Saccardo (1886: 457), Chupp (1954: 33), Crous & cells integrated, terminal, 10–30 μm long, conidiogenous Braun (2003: 197). loci subconspicuous, minute, 0.75–1.5 μm diam, slightly thickened and darkened-refractive. Conidia solitary or Illustration: Braun (1999: 12, g. 14). catenate, in short chains, subcylindrical, occasionally narrowly obclavate to fusiform, 20–60 × 2–5 μm, 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 short to long obconically truncate, about 1 μm wide, hila 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 Pr ncipe), North America (Mexico), South America (Colombia, Ecuador, Uruguay), West Indies (Puerto Rico, irgin Islands).
Note: The orthographic variant “gibertii” sometimes used for this species is incorrect. The epithet was derived from the name of the collector, G. Gilbert. The phylogenetic af nity of 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 elucidate the true generic af nity of this species. For the 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: Muntañola (1960: 210–211, gs 16–17), Ellis (1971: 307, g. 213), Deighton (1979: 27, g. 13).
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 Bar = 10 μm. formation of caespituli, patches to 12 mm diam. Caespituli
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smooth; conidiogenous cellsintegrated,terminal, intercalary olivaceous toolivaceous-brown, darkerinmass,thin-walled, 1–5-septate, oftensomewhat constrictedatthesepta,pale conidiophores, 20–70(–90)× 2–7 confusable withsuper decumbent branchedconidiophores reminiscentofand somewhat geniculate-sinuous,unbranchedorbranched, decumbent, straight,subcylindrical-conicaltosinuous or or fromstromata,throughstomataerumpent,erect to or evenmoreconidiophores,arisingfrominternalhyphae loose todensefascicles,largerfasciclescomposedof50 in outline,2–6 swollen hyphalcells,circulartosomewhatangular-irregular 10–70 septate, 2–4 con uent. hypophyllous, effuse, olivaceous,formingsmallpatchesor μm diam,substomatal,paletobrown,composedof Mycelium μm wide. μm diam. internal, hyphaecolourless,branched, cial hyphaegivingrisetosolitary Stromata Conidiophores μm, oftenirregularinwidth, absent orsmall,about in smalltolarge, Braun etal. Holotype 2(–2.5) in catenateconidia,baseshortobconicallytruncate,(1.5–) smooth, occasionallyfaintlyrough,apexroundedtotruncate the septa,paleolivaceoustoolivaceous-brown,thin-walled, (0–)1–4(–5)-septate, occasionallysomewhatconstricted at straight toslightlycurved,(10–)15–60×3–8 chains, ellipsoid,ovoid,broadlyfusiform,subcylindrical, darkened, 1.5–2.5 conspicuous, sometimessubdenticulate,thickenedand sympodial, occasionallypercurrent,conidiogenousloci and occasionallypleurogenous,5–30 glomerata 436740, 722393;K(M)IMI7706 (slide),MICH15302. Speg., HongosSud-Amer. Dec.Mycol. Argent. 45,e.g.BPI μm wide,hilasomewhatthickenedanddarkened. : , Feb.1881,C.Spegazzini Argentina: Buenos Aires, Palermo,on μm wide.Conidia 914, holotype). Fig. 77.Passaloragomphrenicola Conidia. Bar=10 super B. Solitaryconidiophoresarisingfrom cial hypha.C. A. in simpleorbranched (LPS914). μm. μm long,proliferation Conidiophore fascicles. Conidiophores. IMA FUNGUS Isotypes: 3IDI¿D (LPS μm, 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, enezuela).
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: Muntañola (1960: 208, g. 15 A).
Description: Leaf spots indistinct, formed as yellowish discolorations on the upper leaf surface. Colonies hypophyllous, effuse, brownish. Mycelium internal and external; super cial hyphae solitary, occasionaly intertwined, branched, septate, subhyaline to olivaceous-brown, 2–8 μm wide (sterile hyphae paler and narrow, fertile hyphae with conidiophores broader and darker), thin-walled, smooth. Stromata lacking. Conidiophores solitary, arising from super cial hyphae, lateral or terminal, erect to decumbent, 5–30 × 3–10 μm, 0–1-septate (decumbent threats with terminal conidiophores may be much longer and pluriseptate), sometimes with intercalary cells giving rise to minute peg-like protuberances with a single terminal scar, only about 2–5 μm long and wide, subhyaline to pale olivaceous or olivaceous- brown, thin-walled, smooth; conidiophores mostly reduced to conidiogenous cells, conidiogenous loci conspicuous, somewhat thickened and darkened, 1.5–2 μm diam. Conidia Fig. 78. Passalora gonatoclada (Syd., Fungi Exot. Exs. 930, CUP, catenate, in simple or branched chains, ellipsoid-ovoid to isolectotype). A. Solitary conidiophores arising from super cial hyphae cylindrical, 12–60 × 3.5–8.5 μm, (0–)1–7-septate, subhyaline and through a stoma. B. Conidiophores. C. Conidia. Bar = 10 μm. 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) μm wide, hila thickened and darkened. (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: Muntañola (1960: 217, g. 20, 219, g. 21). 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. discolorations, scattered to con uent, circular, elliptical, Sydow, Fungi itin. Costaricensi Coll. 12]): E 417850, ILL diffuse, violaceous, nally ochraceous to brown. 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, irgin Islands). arising from stromata, through stomata, erect, exuous, geniculate-sinuous, simple or occasionally branched, 40–
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Holotype with 1–3hila. somewhat thickenedanddarkenedhilum,apexrounded or walled, smooth,baseobconicallytruncate,withasingle sigmoid, 20–60×3.5–6 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 base sometimesswollen,aseptatetosparinglyseptate,often 60(–110) ×4–6(–8) 219, Fig. 79.Passalorairesines Conidia. Bar=10 Iresine diffusa her timeinSouth America, sheworked inSerbia(University 2011. The fateofherherbariumcouldnotyet beclari herbarium (no.600)astype. Muntañola-Cvetkovi Note: Muntañola(1960)designated acollectioninherprivate Amaranthaceae Host rangeanddistribution g. 21). : Argentina: A. 1 Aug. 1959, , 1Aug. Conidiophore fascicles. , South America (Argentina). μm. μm, branchlets15–20(–50) Tucumán (based onMuntañola1960:217, μm, 0–3-septate,olivaceous,thin- : On M. Muntañola Iresine diffusa : ElCerroSanJavier, on B. Conidiophore tips.C. (nottraced). Conidia [polymorpha ed. After μm long, diedin catenate, g. 20, Braun etal. ], immersed, subglobose tooblong,25–75 to blackish.Mycelium Caespituli dingy grey, with narrowmarginalline,somewhatraised. angular-irregular, 1–6mmdiam,palegreyish brownto Description Literature Basionym (Fig. 80) SIDI¿DH 3DVVDORUD Fig. 80.3DVVDORUDSIDI¿DH fascicle. B.Conidiophores. curator ofBCN,Barcelona). South American collections arenotincluded(accordingtothe mainly fungicollectedinPortugal,arepreservedBCN,but Belgrad (J. of Biology),butMuntañola’s herbariumisnotpreservedin she movedtoPortugal(UniversityofBarcelona,Faculty of Belgrade,FacultyBiology),andafterherretirement Mycosphaerella and Anam Cercospora : Chupp(1954:35). HFVRD SIDI¿DH &HUFRVSRUD : amphigenous, punctiform,scattered, darkbrown : Leaf spotsamphigenous,circulartosomerwhat ukojevic, pers.comm.). A partofherherbarium, : 35(1954).
(CUP 40533,holotype). C. Conidia.Bar=10 internal. (Chupp) U.Braun&Crous, .: 318(2003). Stromata Chupp,Monograph μm. μm diam,dark substomatal to A. IMA FUNGUS Conidiophore
Cercosporoid fungi 4
brown, composed of subglobose cells, 2.5–5 μm diam. 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). sinuous or slightly geniculate, unbranched, 10–70 × 3.5–6 μm, 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. conidiophores reduced to conidiogenous cells, 10–30 μm long, proliferation sympodial, occasionally percurrent, with Note: Based on a combination of conspicuous conidiogenous ne annellations, conidiogenous loci conspicuous, small, loci, slightly thickened and darkened, 1–1.5 μm wide, and 1–1.5 μm diam, somewhat thickened and darkened. Conidia obclavate-cylindrical, pale olivaceous conidia, Crous & Braun solitary, obclavate-cylindrical, straight to somewhat curved, (2003) reallocated this species to Passalora. 30–85 × 3.5–6 μm, (1–)3–7-septate, pale olivaceous, thin- walled, smooth, apex obtuse, base short obconically truncate, 1.5–2 μm wide, hila somewhat thickened and darkened.
Pseudocercospora
Key to Pseudocercospora species on Amaranthaceae 1 Super cial hyphae with solitary conidiophores developed ...... 2 Super cial hyphae with solitary conidiophores lacking; conidiophores consistently fasciculate ...... 4
2 (1) Conidia broad, 4–8 μm; on Cyathula tomentosa ...... P. cyathulae Conidia narrower, 2–4 μm ...... 3
3 (2) Conidia obclavate-cylindrical, pale olivaceous-brown; on Alternanthra spp...... P. alternantherae Conidia cylindrical, subacicular to obclavate-cylindrical, subhyaline to pale olivaceous; on Celosia spp...... P. celosiarum
4 (1) Conidiophores 10–100 μm long; conidia narrowly liform-subcylindrical, subacicular or narrowly obclavate-cylindrical, (40–)60–130(–150) × 1.5–3.5 μm, 2–12-septate, hila 1–1.5 μm wide; on Gomphrena spp...... P. globosae Conidiophores much shorter, 10–35 μm long; conidia obclavate-cylindrical, 10–90 μm long, 0–10-septate or, if conidiophores longer, conidia 2.5–6 μm wide; on other hosts ...... 5
5 (4) Conidiophores 10–70 × 3–5.5 μm, 0–5-septate; conidia 25–135 × 2.5–6 μm, 3–16-septate, pale olivaceous-brown; on Alternanthera spp...... P. alternantherae-nodosae Conidiophores shorter, 5–35 μm long, only 0–1(–2)-septate; conidia shorter and narrower, 15–90 × 2–4 μm or subhyaline if broader; on other hosts ...... 6
6 (5) Stromata lacking; conidiophores in small fascicles; on Gomphrena spp...... P. gomphrenae Stromata developed, 10–100 μm diam; conidiophores mostly in larger fascicles arising from stromata ...... 7
7 (6) Stromata large, 10–100 μm diam; conidia (2–)3–5(–6) μm wide, hila 1.5–3 μm wide, subhyaline; on Gomphrena pulchella ...... P. gomphrenae-pulchellae Stromata smaller, 10–65 μm; conidia 2–3.5 μm wide, hila 1–2 μm wide, subhyaline to pale olivaceous-brown; on other hosts ...... 8
8 (7) Conidia mostly narrowly cylindrical with truncate base, pale olivaceous-brown; on Amaranthus 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
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narrowlyobclavate-cylindrical,narrower, 1.5–3.5 Stromata 10–40 unthickened, not darkened. conidiogenous cells, lociinconspicuous, cells integrated,terminalor conidiophoresreducedto pale olivaceous-brown,thin-walled, smooth;conidiogenous geniculate, unbranched,15–55 ×3–4.5 lateral, erect,straight,subcylindrical tosinuous,slightly stomata, andsolitary, arisingfromsuper 5–60, divergenttodense,arisingfromstromata,through Conidiophores globose, substomatal,20–40 2–5 brown, thin-walled,smooth,2–5 external, super hypophyllous, notveryconspicuous.Mycelium brown, marginindistinct.Caespituli to fusiform,3–12×2–4mm,oftencon Super (2–)3–5(–6) Stromata10–100 Stromatadeveloped,10–100 2 (1) Stromatalacking,conidiophoresinsmallfascicles ...... 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 1 Super Alternanthera Tabular keytoPseudocercosporaspecieson Amaranthaceae accordingtohostgenera (Fig. 81) Mycotaxon μ m wide,externalhyphae2–3.5 globosa : Kamal(2010:147). : Yen : cial hyphae and solitary conidiophores absent; conidiophores only in fascicles cial hyphaeandsolitaryconidiophoresabsent;onlyinfascicles ...... cial hyphaewithsolitaryconidiophoresdeveloped ...... Leaf spots in smalltowell-developedfascicles,about et al.(1982:40, cial; hyphaebranched,septate,olivaceous- 16: 39(1982)...... μm diam;conidiophores10–100×1.5–5.5 μm wide,subhyaline;on μm diam;conidiophores10–30×2.5–6 amphigenous, scattered,oval μ m diam,browntodarkbrown. Conidia g. 3). μ m wide,internalhyphae im oiipoe nsalt ag acce,aiigfo toaa ...... 2 μm diam;conidiophoresinsmalltolargefascicles,arisingfromstromata ...... amphigenous, mostly J.M. Yen etal., μ solitary, obclavate- m wide. μ m, 0–3-septate, uent, greyish cial hyphae, internal and Gomphrena pulchella Stromata Braun etal. μm, subhyalinetopaleolivaceous-brown;on μm; conidiaobclavate-cylindrical, only providedwithacollectionnumber). numerous unincorporated Yen collectionsinUC(unnamed, be traced.Itispossiblethatthismaterialstillamong at UC,butthetypematerialconcernedcouldcurrentlynot holotype. The mycologicalLAMcollectionsarenowhoused Notes: Yen collection. Host rangeanddistribution 10583] (nottraced). sp., Holotype unthickened, notdarkened. smooth, apexobtuse,baseshortobconicallytruncate,hila μ cylindrical, straighttoslightlycurved,30–90×2.5–4 Basionym (Fig. 82) Pseudocercospora Synonyms μm; conidia m, 3–10-septate,paleolivaceous-brown,thin-walled, China ada) Goh&W.H. Hsieh,Trans. Mycol.Soc.Republ. Mycopathol. Mycol. Appl Rep. Gov. Agric. Res.Inst.Taiwan 35:106(1928). Amaranthaceae ...... : HFVRD DOWHUQDQWKHUDHQRGLÀRUDH &HUFRVSRUD : 2:135(1987). : Cercosporaalternanthericola India et al.(1982)cited“LAM, Yen #10583”as liform-subcylindrical, subacicularto : West Bengal ...... , 7Jan.1981, DOWHUQDQWKHUDHQRGLÀRUDH . : Onlyknownfromthetype 27: 93(1965)[ : Purulea,on P. alternantherae-nodosae P. gomphrenae-pulchellae B. K.DasPcc4483[Yen ...... Gomphrena Pavgi &U.P. Singh, P. alternantherae P. amaranthicola P. chamissoana P. gomphrenae holotype P. celosiarum P. froelichiae Alternanthera P. cyathulae IMA FUNGUS P. globosae Sawada, : (Saw- India : Cercosporoid fungi 4 ARTICLE
Fig. 81. Pseudocercospora alternantherae (based on Yen et al. 1982: 40, g. 3). A. Conidiophore fascicles. B. Solitary conidiophores arising from super cial hyphae. C. Conidia. Bar = 10 μm.
Uttar Pradesh: aranasi, 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: on Hsieh & Goh (1990: 18, g. 7). A. Conidiophore fascicle. B. 53), Kamal (2010: 147). Conidiophores. C. Conidia. Bar = 10 μm.
Illustrations: Pavgi & Singh (1965: 94, pl. 1, gs 16–18), Hsieh & Goh (1990: 18, g. 7), Guo & Hsieh (1995: 9, g. 8), 3–5.5 μm, 0–5-septate, uniformly pale olivaceous-brown Guo et al. (1998: 19, g. 8). to brown, thin-walled, smooth; conidiogenous cells integrated, terminal, occasionally conidiophores reduced Description: Leaf spots at rst indistinct or visible as 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 discoloured, leaves nally dis gured, faded, necrotic. darkened. Conidia solitary obclavate-subcylindrical, Caespituli amphigenous, forming effuse, dark olivaceous straight to curved, 25–135 × 2.5–6 μm, 3–15-septate, pale patches, nally covering the whole leaf blade. Mycelium olivaceous, thin-walled, smooth, apex obtuse, rounded to internal. Stromata small, substomatal, 10–20 μm diam, subacute, base obconically truncate, 2–2.5 μm wide, hila 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): or distinctly geniculate-sinuous, unbranched, 10–70 × Taiwan: Taipei, on Alternanthera sessilis, 9 May 1924, K.
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Illustrations Guo etal. Literature Basionym Synonym reduced tosynonymy. 3VHXGRFHUFRVSRUDDOWHUQDQWKHUDHQRGLÀRUDH from Indiaon K(M) IMI31945). [QRGLÀRUD Host rangeanddistribution 05136, TNS-F-220608. Pseudocercospora amaranthicola sessilis from Taiwan havebeenexamined[Taipei, on Notes: Three duplicatesofanadditionalsyntypecollection India, UttarPradesh; Taiwan). Sawada latter herbarium. is stillamongthenumerous untreated Yen collectioninthe neither inPCnorUC.Itispossible thatthematerialconcerned Notes: Type material of thisspeciescouldnotbetraced, collection. Host rangeanddistribution 1971, Amaranthus tricolor Holotype truncate, hilaneitherthickenednordarkened. obtuse orsubobtuse,basetruncatetosomewhatobconically 80 ×2–3 or onlyslightlyobclavate-cylindrical,straighttocurved,15– sympodial proliferation. or visibleastruncatetipslateralshoulderscausedby conidiogenous cells,lociinconspicuous cells integrated,terminalorconidiophoresreducedto pale olivaceous-brown,thin-walled,smooth;conidiogenous or occasionallybranched,8–30×2.5–3.5 exuous, subcylindricaltogeniculate-sinuous,unbranched fascicles, arisingfromstromata,erumpent,erect,straight, μm diam,darkbrown. internal. amphigenous, punctiform,darkbrowntoblackish. diam, palebrown,marginbrown. Description 377, g.307). (Fig. 83) amaranticola Bull. Trimestriel Soc.Mycol.France Trimestriel Soc.Mycol.France (1978). Yen, J. M. Yen , 5 Aug. 1907,K.Kawakami (NTU-PPE,hb.Sawada). Stromata : ], : : Hsieh&Goh(1990:17),Guo(1995:357), Bull. Trimestriel Soc.Mycol.France : Taiwan: Peikuoshan, Yuanlin, ChanghuaHsien,on (1998: 376),Crous&Braun(2003:55). Cercoseptoria amaranthicola μm, 3–10-septate,paleolivaceous-brown,apex : : Yen (1977:146, Alternanthera Cercospora amaranthicola Leaf spots Alternanthera sessilis 71282. ”. immersed, globoseorsubglobose,25–65 Cercospora alternathericola [mangostanus Conidiophores Conidia circular orsubcircular, 0.5–4mm sp., Amaranthaceae : Onlyknownfromthetype : OnAlternantherasessilis g. 1A–C), Caespituli solitary, narrowlycylindrical 93:145(1977). ], Amaranthaceae in well-developed,dense (BPI 432466,432467; is barelydistinctfrom Isolectotypes (J.M. Yen)J.M. Yen,
J.M. Yen, Bull. 97: 91(1981);“as
(J.M. Yen)J.M. epiphyllous, rarely Guo etal. μm, 0–3-septate, and istherefore , Asia (China; , Asia Alternanthera , described
Mycelium 94: 385 : HMAS , 30Oct. (1998: Braun etal.
Bar =10μm. g. 1 A–C). A. Fig. 83. (Fig. 84) Description (1995: 10, g.9),Guo Illustrations Crous &Braun(2003:114), Kamal(2010:162). (1995: 9),Guoetal.(1998:20),Nakashima Literature Basionym Pseudocercospora celosiarum Mandal) Deighton, Trans. Brit.Mycol.Soc Pseudocercospora amaranthicola : Katsuki(1965:9,as“ : Cercosporacelosiarum : : Kar&Mandal(1970:423, Leaf spots Conidiophore fascicles. etal.(1998:20, amphigenous, subcircular toangular- Mycol. Pap.140:141(1976). . 54:423(1970). C. celosiae B. Conidiophores. A.K. Kar&M.Mandal, (based on Yen 1977:146, g. 9). g. 1),Guo&Hsieh (A.K. Kar&M. et al.(2002:98), ”), Guo&Hsieh IMA FUNGUS C. Conidia. Cercosporoid fungi 4
stromata, through stomata or solitary, arising from super cial ARTICLE hyphae, lateral, erect, straight, subcylindrical-conical, not to strongly geniculate-sinuous, unbranched, 5–75 × 2–4 μm, 0–5-septate, pale olivaceous to olivaceous-brown, thin- walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–20 μm long, conidiogenous loci inconspicuous, neither thickened nor darkened. Conidia solitary, cylindrical, subacicular, long acicular to obclavate-cylindrical, 10–105 × 2–4 μm, straight to curved, 0–10-septate, subhyaline to pale olivaceous, thin- walled, smooth, apex obtuse to subacute, base truncate to short obconically truncate, 1.5–2 μm wide, hila unthickened, 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 collections of this species were originally wrongly identi ed and published as Cercospora celosiae (Katsuki 1965), which was clari ed in Nakashima et al. (2002).
Pseudocercospora chamissoana R.F. Castañeda & U. Braun, Cryptog. Bot. 1: 51 (1989). (Fig. 85)
Illustration: Castañeda Ruiz & Braun (1989: 47, pl. 3, g. 18).
Description: Leaf spots amphigenous, subcircular to irregular, 1–10 mm diam, brownish, later pale, margin inde nite or narrow, light to dark brown, formed as mariginal line. Caespituli hypophyllous, punctiform, dark brown to blackish, scattered. Mycelium internal. Stromata substomatal, 20–45 μm diam, brown, composed of swollen hyphal cells, subglobose to angular in outline, 2–5 μm diam. Conidiophores in small to moderately large fascicles, divergent to dense, arising from stromata, through stomata, erect, straight, subcylindrical- conical to geniculate-sinuous, unbranched, 5–25 × 2–5 μm, 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 conidiogenous cells, 5–20 μm long, conidiogenous loci super cial hyphae. C. Conidiophores D. Conidia. Bar = 10 μm. inconspicuous to subdenticulate, but always unthickened and not darkened. Conidia solitary, obclavate-cylindrical to somewhat fusoid, straight to slightly curved, 40–65 × 2–3 μm, 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, truncate, 1–2 μm wide, hila neither thickened nor darkened. dark brown or more greyish by abundant conidiation. Mycelium internal and external; super cial hyphae, if present, Holotype: Cuba: Granma: Guisa, Los Corrales, on Chamissoa emerging through stomata, branched, septate, 2–3 μm diam, 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, 5–60 μm diam, pale olivaceous to brown. Conidiophores in Host range and distribution: Only known from the type small to large fascicles, divergent to very dense, arising from collection.
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patches caused bycoloniesofthefungus,margin inde diam orlarger, diffuse, belowvisibleassubcircularbrownish surface onlyformedasyellowish discolorations,5–10mm Description (2010: 115–116). Literature Basionym (Fig. 86) MycoBank MB814580 Pseudocercospora cyathulae(Syd.)U.Braun, Conidiophores fascicles. Fig. 85.Pseudocercosporachamissoana comb. nov. (1937). : Chupp(1954:33), : Cercosporacyathulae : Leaf spots B. Conidia.Bar=10 lacking oralmostso,ontheupper leaf asudeva (1963:95),Kamal Syd., (HAL 1651F, isotype). Ann. Mycol μm. . 35: 239 nite. Braun etal. A. Conidiophores constrictions atsepta,thin-walled, smooth. μm diam,olivaceoustomedium olivaceous-brown,oftenwith and darker, 3–8μmwide,swollencellsoccasionally to12 hyphae withconidiophoresor conidiogenouscellsbroader olivaceous-brown, mostlywithout constrictionsatsepta,fertile hyphae narrow, 1–3.5 cover, branched,sometimesanastomosing,septate,sterile plagiotropous orclimbingleafhairs,formingadensenet or brown. Caespituli hyphae. Super Fig. 86.Pseudocercosporacyathulae cial hyphae. C. Conidiophores. Mycelium hypophyllous, effuse, brownishordarkolivaceous- solitary, arisingfromsuper B. Solitaryconidiophoresarisingfromsuper internal andexternal;super μm, subhyalinetopaleolivaceousor D. Conidia.Bar=10 (CUP 39581,holotype). μm. Stromata cial hyphae, cial hyphae IMA FUNGUS lacking. cial A. Cercosporoid fungi 4
lateral or terminal, length inde nite (forming decumbent fertile ARTICLE threats, but differentiation between individual conidiophores and hyphal portions dif cult or even impossible), shorter lateral conidiophores arising from decumbent threats about 5–30 μm long, width about 3–8 μm, erect to decumbent, straight, subcylindrical to geniculate-sinuous, unbranched or branched, aseptate to pluriseptate, olivaceous, olivaceous- brown to light brown, thin-walled, smooth; conidiogenous cells integrated, terminal, 10–25 μm long, conidiogenous loci inconspicuous to subconspicuous by being slightly refractive, but unthickened, occasionally visible in front view as minute circle (paracercosporoid) or subdenticulate, 1.5–2 μm diam. Conidia solitary, rarely in short chains, cylindrical to vermiform, shorter conidia sometimes ovoid, obovoid, short cylindrical, broadly fusiform, straight to curved, occasionally sigmoid, 15–100 × (4–)5–8 μm, 0–7-septate, occasionally somewhat constricted at the septa, pale brown, thin-walled, smooth, apex broadly rounded, rarely subtruncate or somewhat attenuated, base rounded to short obconically truncate, 1.5–2 μm wide, hila unthickened, not darkened, 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. Conidia. Bar = 10 μm. that could be traced and examined. Chupp (1954) confused this species with Ragnhildiana cyathulae ( Passalora cyathulae) and used the wrong citation “Cercospora cyathulae (Stev. & Solh.) Sydow” although Sydow (1937) did not intend obclavate-cylindrical, 10–80 × 2–3.5 μm, 1–7-septate, to introduce a new combination based on Ragnhildiana pale olivaceous, thin-walled, smooth, apex obtuse or cyathulae. He undoubtedly published a new species without subacute, base short obconically truncate, 1–2 μm wide, hila any reference to R. cyathulae. The two species are neither unthickened, not darkened. conspeci c nor congeneric. 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: Braun & Freire (2004: 231, g. 7). Host range and distribution: On Froelichia sp., 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 fructi cation. Mycelium internal. Stromata immersed to P. chamissoana, described from Cuba on Chamissoa or somewhat erumpent, 10–50 μm diam, olivaceous- altissima (Amaranthaceae, Amaranthoideae), which differs, brown, composed of swollen hyphal cells, 2–7 μm diam, 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, slightly geniculate-sinuous, unbranched, 10–35 × 2–5 μm, 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: conidiophores reduced to conidiogenous cells, 10–25 μm 224 (1964). long, conidiogenous loci inconspicuous. Conidia solitary,
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10–40 subglobose tosomewhatirregular, substomataltoimmersed, to dense,velvety, greyishwhite. by abundantfungalcolonies. subcircular tosomewhatirregular, greyishwhitetodarkgrey Description g. 42). Illustrations 14), Crous&Braun(2003:199). Literature Conidiophores fascicles.B. Fig. 88.Pseudocercosporaglobosae and notdarkened. conical, apextruncate,1–2 loci inconspicuoustodistinctly denticle-like,subcylindrical- occasionally intercalary, 10–25 walled, smooth;conidiogenous cellsintegrated,terminal, subhyaline topaleyellowish orolivaceous-brown,thin- unbranched tobranched,10–100×1.5–5.5 and subcylindrical-conicaltodistinctlygeniculate-sinuous, arising fromstromata,throughstomataorerumpent,straight to moderatelylargefascicles,loosedense, μm diam,yellowishbrown. : Yen &Lim(1980:177),BraunSivapalan(1999: : : Yen227, (1964: Leaf spots Conidia Conidiophores. at Caespituli μm wide,alwaysunthickened solitary, g. 7), Yen &Lim(1980:239, Mycelium rst inconspicuous,later μm long,conidiogenous Conidiophores C.Conidia. liform-subcylindrical, amphigenous, effuse (PC, holotype).A. internal. μm, 0–6-septate,
Bar =10μm. Stromata in small Braun etal. Synonym Pseudocercospora gomphrenae Goh&W.H. Hsieh, 1990: 19, g.8). Fig. 89.Pseudocercosporagomphrenae (Fig. 89) Holotype darkened. obconically truncate,1–1.5 apex subacutetosubobtuse,basetruncateshortorlong pale olivaceousorolivaceous-brown,thin-walled,smooth, 130(-150) ×1.5–3.5 subacicular tonarrowlyobclavate-subcylindrical,(40–)60– Gomphrena Host rangeanddistribution Apr. 1964,S.H. Yen 19(PC). Trans. Mycol. Soc.Republ.China [type: seePseudocercospora gomphrenae Res. Inst.Formosa85:107(1943), : : Singapore Cercospora gomphrenae sp.), A. Conidiophorefascicle. Amaranthaceae : Katong,on μm, 2–12-septate,subhyalinetovery μm wide,hilaunthickened,not : OnGomphrena , Asia (Brunei,Singapore). Gomphrena B. Conidia.Bar=10 (based onHsieh&Goh nom. inval Sawada, 4(2–3): 8(1989). ].
IMA FUNGUS globosa . (Art.39.1) ( Rep. Gov. globosa , μm. 20 , Cercosporoid fungi 4 ARTICLE
Fig. 90. Pseudocercospora gomphrenae-pulchellae (HAL 1635 F, isotype). A. Conidiophores fascicles. B. Conidiophores. C. Conidia. Bar = 10 μm.
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: Hsieh & Goh (1990: 19, g. 8), Guo & Hsieh TNS-F-220432 (1995: 11, g. 10), Guo et al. (1998: 21, g. 10). 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). 30 × 3.5–4 μm, 0–2-septate, olivaceous, thin-walled, smooth; (Fig. 90) conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, conidiogenous loci Illustration: Braun et al. (2001: 29, g. 8). inconspicuous, unthickened, not darkened. Conidia solitary, cylindrical, obclavate-cylindrical to subacicular, straight Description: Leaf spots amphigenous, subcircular to irregular, to curved, 30–90 × 2.5–4 μm, 4–7-septate, subhyaline to 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, 1.5–2 μm wide, hila unthickened, not darkened. loose to dense, blackish, later greyish white by abundant
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and narrowerconidia,only1.5–3.5 has smallerstromata,muchlonger, septateconidiophores gomphrenae-pulchellae Note: collection. Host rangeanddistribution Delhey Gomphrena pulchella the hostsoffamiliestreatedin part4. during thecourseofmonographic studiesofcercosporoidsfungion WSP forloaningtypematerialandothercollectionsintheir keeping 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 Type Literature Holotype hila unthickened,notdarkened. rounded, truncatetoobconicallytruncate,1.5–3 tinge), smooth,apexobtuse,occasionallysubacute,base μm, (0–)1–8-septate,subhyaline(withaverypalegreenish Conidia darkened orthickened,infrontviewvisibleasminutecircle. subconspicuous, paracercosporoid,i.e.rimveryslightly μm long,conidiogenouslociinconspicuous,occasionally or conidiophoresreducedtoconidiogenouscells,10–25 walled, smooth;conidiogenouscellsintegrated,terminal 2.5–6 towards thetip,geniculate-sinuous,unbranched,10–30× or erumpent.erect,straight,subcylindricalattenuated moderately dense,arisingfromstromata,throughstomata Synonym olivaceous-brown. Stromata 1.5–6 conidial formation. Alternaria gomphrenae Kamal (2010)reduced with cicatrizedconidiogenouscellsandrostrateconidia, based ontheoriginaldescriptionandillustrationofaspecies Note: Type materialofthisspecieswasnotindicated, but P. Chiddarwar Agric. For. Morioka 22: 511 (1957). : India μm diam,septate,subhyalinetopalebrown,smooth. μm, 0–1-septate,subhyalinetopaleolivaceous,thin- 1340 Pseudocercospora gomphrenae solitary, obclavate-cylindrical,20–90×(2–)3–5(–6) : : : Kamal(2010:179). substomatal tointraepidermal,10–100 Argentina: Alternaria gomphrenae : (BB). Maharashtra (otherwisenotspeci Conidiophores Isotype: HAL 1635F. Mycelium , Prov. Buenos Aires . 9: 6(1926). P. gomphrenicola Amaranthaceae in lackingstromata,and : Pune,onGomphrenaglobosa : Onlyknownfromthetype internal; hyphaebranched, in smalltolargefascicles, ed). μm wide. Togashi,
Chidd., , : BahiaBlanca,on 17 Mar. 2000, to synonymywith differs from Bull. Imp.Coll. Sci. &Cult. P. globosa μm diam, μm wide, Braun etal. , R. P. P. P. P. Ahmad S,IqbalSH,Khalid AN (1997) REFERENCES Al eri JrSA,LangdonKR,Wehburg C,KimbroughJW(1984)Index Braun U(2001) T Braun, U.(1999) Taxonomic notesonsomespeciesofthe Braun U(1998) Braun U(1995) Braun U(1993) Taxonomic notesonsomespecies ofthe Braun U(1992) Taxonomic notesonsomespecies ofthe Boedijn KB(1961) The genus Bhartiya HD,Singh AN, KumariN,SinghPN(1998)[“1997”] Braun U,FreireFCO(2006)Somecercosporoidhyphomycetesfrom Braun U,FreireFCO(2004)Somecercosporoidhyphomycetesfrom Bhartiya HD,NarayanS,Singh AN, KumariN(2003) Additions Bakshi M, Arzanlou M,Babai-Ahari A (2012)Comprehensivecheck Bakalova GG,Borisova TI (2010)CercosporoidHyphomycetous Bagyanarayana G,JagadeeswarP, BraunU(1991)Miscellaneous Atkinson GF(1892)Some Braun U,Sivapalan A (1999)Cercosporoid hyphomycetesfrom Braun U,Mel’nik Braun U(2000b) Annotated listof Braun U(2000a)Miscellaneousnotesonsomemicromycetes. Braun U,HillCF(2002)SomenewmicromycetesfromNew Braun U,CrousPW(2005) Additions andcorrections tonames Braun U,CastañedaRuizRF(1991) Bulletin Agriculture andConsumerServices,DivisionofPlantIndustry, of PlantDiseasesonFlorida(Revised). Sultan Ahmad MycologicalSocietyofPakistan. Spegazzini. complex ( genera (phytopathogenichyphomycetes) genera (phytopathogenichyphomycetes). complex (III).Mycotaxon complex. NovaHedwigia Hedwigia Pradesh. Foliicolous fungiintheforest Phytopathology Brazil –I Brazil –III. 416. Cuba (II).CryptogamicBotany complex ( to 2: 44–55. list ofcercosporoidfungifromIran. Bulgarian Academy ofSciences. Fungi. [FungiofBulgariavol.7.]So a:InstituteBotany, notes onIndianCercosporae. the ElishaMitchelScienceSociety 1–130. Komarova, Rossijskaya Akademiya NaukSt.Petersburg adjacent countries. Schlechtendalia Mycological Progress published in erlag. erlag. Cercospora 11 3:411–436. Kavaka ). Schlechtendalia II). . CryptogamieMycologie : 1–389. Cryptogamie Mycologie A monographofCercosporella,Ramulariaandallied A monographofCercosporella,Ramulariaandallied Schlechtendalia axonomic notesonsomespeciesofthe Cercospora Fungal Diversity A (1997)Cercosporoid fungifromRussiaand 56: 270–275. 5: 31–56. from North-EasternUttarPradesh. 25: 43–51. Trudy BotanicheskogoInstituta ImeniV.L. 1: 19–30. Cercosporae 55:211–221. 48: 275–298. and 5: 57–79. 2:1–28. Mycotaxon Cercospora Cercospora 8:41–71. 2–3 Passalora Cercospora ora ofNorth-EasternUttar Plant Pathology&Quarantine : 289–297. 25:221–244. 27:231–248. 8: 33–76. Fungi ofPakistan from Alabama. . 42:319–326. . Florida Departmentof
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