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Cercosporoid fungi (Mycosphaerellaceae) 2. Species on monocots (Acoraceae to Xyridaceae, excluding Poaceae)

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Abstract

Cercosporoid fungi (formerly Cercospora s. lat.) represent one of the largest groups of hyphomycetes belonging to the Mycosphaerellaceae (Ascomycota). They include asexual morphs, asexual holomorphs, or species with mycosphaerella-like sexual morphs. Most of them are leaf-spotting plant pathogens with special phytopathological relevance. In the first part of a new monographic work, cercosporoid hyphomycetes occurring on other fungi (fungicolous species), on ferns (pteridophytes) and gymnosperms were treated. This second part deals with cercosporoid fungi on monocots (Liliopsida; Equisetopsida, Magnoliidae, Lilianae), which covers species occurring on host plants belonging to families arranged in alphabetical order from Acoraceae to Xyridaceae, excluding Poaceae (cereals and grasses) which requires a separate treatment. The species are described and illustrated in alphabetical order under the particular cercosporoid genera, supplemented by keys to the species concerned. A detailed introduction, a survey of currently recognised cercosporoid genera, a key to the genera concerned, and a discussion of taxonomically relevant characters were published in the first part of this series. Neopseudocercospora, an additional recently introduced cercosporoid genus, is briefly discussed. The following taxonomic novelties are introduced: Cercospora alpiniigena sp. nov., C. neomaricae sp. nov, Corynespora palmicola comb. nov, Exosporium miyakei comb. nov, E. petersii comb. nov., Neopseudocercospora zambiensis comb. nov., Passalora caladiicola comb. nov, P. streptopi comb. nov., P. togashiana comb. nov., P. tranzschelii var. chinensis var. nov, Pseudocercospora beaucarneae comb. nov, P. constrictoflexuosa comb. et stat. nov, P. curcumicola sp. nov, P. dispori comb. nov., P. smilacicola sp. nov, P. urariigena nom. nov, Zasmidium agavicola comb. nov, Z. cercestidis-afzelii comb. nov, Z. citri-griseum comb. nov., Z. cyrtopodii comb. nov, Z gahnae comb. nov, Z. indicum comb. nov., Z liriopes comb. nov., Z mycovellosielloides sp. nov, Z. scleriae comb. nov, Z. smilacicola comb. nov, and Z. thaliae comb. nov.

Introduction

Cercospora-like fungi represent one of the largest groups of mostly plant pathogenic, leaf-spotting hyphomycetes, causing numerous economically relevant diseases of cultivated plants. The only worldwide monograph of this fungal group, published by Chupp (1954), is badly out of date. A new “Chupp” is urgently required by plant pathologists. Therefore, Braun et al. (2013) initiated initiative series of papers to update the monograph of Cercospora and allied genera (Mycosphaerellaceae), which is being accomplished in a stepwise approach due to the huge size of this fungal group. Sexual morphs (teleomorphs) are included in the descriptions as far as known and proven. Mycosphaerella s. str., which is based on M. punctiformis, its type species, is a heterotypic synonym of Ramularia (Braun et al. 2013). Hence, the names of sexual morphs described as Mycosphaerella s. lat. species are cited as synonyms of the particular species of Cercospora, Passalora, Pseudocercospora, etc. A general introduction, covering all aspects of cercosporoid fungi, ranging from the history of genera to keys to the currently recognised genera, has been published in the first part of this series of monographic contributions together with detailed treatments of species on other fungi (mycophylic taxa), on ferns as well as gymnosperms. The present part is devoted to cercosporoid hyphomycetes on monocots, covering species on hosts belonging to the Acoraceae to the Xyridaceae, but excluding those occurring on true grasses and cereals (Poaceae), which necessitates a separate treatment owing to the numerous species involved. The treatment of cercosporoid fungi on monocots follows the principles outlined in part 1 (Braun et al. 2013).

Materials and Methods

The present work is a compilation based on papers and unpublished data of the authors as well as global literature. Details on methods are to be found in the papers cited. As far as new examinations are concerned, fungal structures have been examined on the base of standard methods of light microscopy, using an Olympus BX50 microscope, 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 magnification of x1000. All illustrations have been prepared by the first author. The following abbreviations are used: author names follow Brummit & Powell (1992), journals Bridson (2004a, b), and exsiccatae http://www.botanischestaatssammlung.de/DatabaseClient/IndExs/index.jsp (IndExs — Index of Exsiccatae). Taxonomy and nomenclature of plant families, genera and species are based on the “Angiosperm Phylogeny Website” (http://www.mobot.org/mobot/research/apweb/), Tropicos database (http://www.Tropicos.org/), and The Plant List (http://www.theplantlist.org).

Sexual Morphs and Currently Recognised Cercosporoid Genera — an Addition

Crous et al. (2013b) introduced the new genus Neopseudocercospora with its type species N. terminaliae, described from Zambia on Terminalia sp. This genus is morphologically intermediate between Pseudocercospora and Sporidesmium, but phylogenetically belongs in Mycosphaerellaceae and must thus be considered a true cercosporoid genus.

Neopseudocercospora Crous, Persoonia 31: 219 (2013).

Type species: Neopseudocercospora terminaliae Crous, 2013 (i.e. N. zambiensis (Deighton) Crous & U. Braun 2014).

Neopseudocercospora zambiensis (Deighton) Crous & U. Braun, comb. nov.

MycoBank MB809006

Basionym: Sporidesmium zambiense Deighton, Mycol. Pap. 117: 27 (1969).

Synonyms: Repetophragma zambiense (Deighton) Subram., Proc. Indian Acad. Sci., B, 58: 185 (1992). Pseudocercospora zambiensis (Deighton) B. Sutton, Mycopathologia 125: 61 (1994)

Neopseudocercospora terminaliae Crous, Persoonia 31: 219 (2013).

Description: Hyphomycetes (asexual morphs); Mycosphaerellaceae. Foliicolous, plant pathogenic. Mycelium superficial; hyphae branched, septate, pigmented, smooth-walled. Stromata lacking. Conidiophores solitary, arising from superficial hyphae, lateral, one-celled orseptate, pigmented; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, proliferation percurrent or sympodial and percurrent proliferations mixed, conidiogenous loci truncate, relatively broad, neither thickened nor darkened. Conidia solitary, rather sporidesmium/scolecostigmina-like, scolecosporous, pluriseptate, pigmented, rather dark in mass, wall somewhat thickened, hilum broadly truncate, neither thickened nor darkened, but with a minute frill.

Notes: Neopseudocercospora is morphologically reminiscent of the genus Sporidesmium (Sordariomycetes), but belongs to Mycosphaerellaceae where it clusters with Microcyclosporella and zasmidium-like species (clade 8 in Crous et al. 2013a). Due to the unthickened, non-pigmented conidiogenous loci and conidial hila, the type species of Neopseudocercospora would be morphologically assignable to Pseudocercospora, but phylogenetically it does not belong in the Pseudocercospora clade and warrants a genus of its own. Morphologically it differs from most Pseudocercospora species by its rather sporidesmium/scolecostigmina-like conidia. Deighton (in Cejp & Deighton 1969) introduced Sporidesmium zambiense, based on material on Terminalia mollis collected in Zambia, which Sutton (1994) reallocated to Pseudocercospora. This species is conspecific with Neopseudocercospora terminaliae. Subramanian (1992) assigned S. zambiense to Repetophragma, but that genus is not appropriate for the present species as it is characterised, based on its type species, by having consistently, conspicuously percurrent conidiogenous cells with numerous, dense annellations and didymo- to phragmosporous brown conidia with an almost colourless base. The phylogenetic position of this genus is not yet known, but probably does not lie within Mycosphaerellaceae.

Taxonomic Treatment

Cercosporoid species on monocots (Acoraceae to Xyridaceae)

Acoraceae

Passalora

On Acorus, see Passalora acori (Part 1, fungicolous species).

Alismataceae

Cercospora

Key to Cercospora species on Alismataceae
  1. 1

    Conidia acicular, base truncate ............................................................................ 2

    Conidia at least partly obclavate-cylindrical with obconically truncate base ............................................... 3

  2. 2

    (1) Conidiophores relatively short, 10–70 µm; conidia 40–125 µm long, hila about 2.5–5 µm wide; on Echinodorus spp. ....................................................................... C. echinodori

    Conidiophores longer, > 100 µm; conidia longer, 80–155 µm, hila about 2.5–3 µm wide; on Alisma orientale ......................................................................... C. alismaticola

    [If on Sagittaria montevidensis, Brazil, morphologically indistinguishable; see C. apii s. lat.]

  3. 3

    (1) Stromata lacking; conidiophores to about 320 µm long; on Echinodorus spp. ................................ C. osiridis

    Stromata at least partly developed, 10–40 µm diam ...................................................... 4

  4. 4

    (3) On Alisma spp. and Echinodorus spp. .......................................................... C. alismatis

    On Sagittaria spp. .................................................. C. sagittariae

Cercospora species on Alismataceae

Cercospora alismaticola Z.D. Jiang & P.K. Chi, in Chi, Fungal Diseases of Cultivated Medicall Plants in Guangdong Province: 97 (1994); also J. S. China Agric. Univ. 15: 17 (1994).

(Fig. 1)

Fig. 1
figure1

Cercospora alismaticola (based on Chi 1994: 97, fig. 88). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Zhuang (2001), Crous & Braun (2003: 51).

Illustration: Chi (1994: 97, fig. 88).

Description: Leaf spots amphigenous, circular, 2–5 mm diam, reddish brown, centre later becoming greyish white. Caespituli amphigenous. Mycelium internal. Stromata small, subglobose, about 10–40 µm diam, pigmented. Conidiophores fasciculate, 3–20, loose, arising from stromata, erect, straight, unbranched, 2–8 times geniculate, somewhat narrowed towards the apex, 100–177 × 4–6 µm, 3–5-septate, brown, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, with thickened and darkened conidiogenous loci. Conidia solitary, acicular, straight to curved, 80–155 × 3–4 µm, pluriseptate, hyaline, apex pointed, base truncate, about 2.5–3 µm wide, with thickened and darkened hilum.

Holotype: China: Guangdong: Guangzhou, on Alisma orientale [A. plantago-aquatica var. orientale], Alismataceae, Nov. 1988, Z. D. Jiang (hb. S. China Agric. Univ., Guangzhou).

Host range and distribution: Only known from the type collection.

Notes: This species belongs to the Cercospora apii s. lat. complex.

Cercospora alismatis Ellis & Holw., J. Mycol. 1: 63 (1885). (similar to Fig. 3)

Literature: Saccardo (1886: 478), Vassiljevsky & Karakulin (1937: 221), Chupp (1954: 28), El-Gholl et al. (1992: 265), Crous & Braun (2003: 51).

Exsiccatae: Ellis & Everh., Fungi Columb. 597. Ellis & Everh., N. Amer. Fungi 3191.

Description: Leaf spots amphigenous, subcircular to angularirregular, 2–15 mm diam, brown to greyish brown, margin indefinite or darker. Caespituli amphigenous. Mycelium internal. Stromata almost lacking or 10–40 µm diam, substomatal, brown. Conidiophores in small to moderately large fascicles, arising from internal hyphae or stromata, through stomata, erect, straight, subcylindrical to geniculate or somewhat sinuous, mostly unbranched, 15–180 × 4–7 µm, septate, pale yellowish green, olivaceous to medium olivaceous-brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, 10–50 µm long, sympodial, with thickened and darkened conidiogenous loci, 2–2.5 µm diam. Conidia solitary, obclavate(-subcylindrical), straight to curved, 50–160 × (3−)4–6 µm, 3- to pluriseptate, hyaline or subhyaline, thin-walled, smooth, apex obtuse to subacute, base obconically truncate, 2–3 µm wide, hila somewhat thickened and darkened.

Holotype: USA: Iowa: Decorah, on Alisma sp., 29 Jul. 1884, Holway (NY 00830162).

Host range and distribution: On Alisma (plantago-aquatica, subcordatum, triviale, Alisma sp.), Echinodorus (subalatus [intermedius], Echinodorus sp. [“alismatis”]), Alismataceae, Asia (Singapore), North America (USA, California, Delaware, Iowa, Indiana, Minnesota, Montana, Nebraska, New York, Wisconsin, Texas), West Indies (Dominican Republ., Haiti).

Notes: This is a true Cercospora s. str. close to C. apii s. lat., but distinct by having obclavate-cylindrical conidia, (3−)4–6 µm wide. Older records of C. alismatis on Sagittaria spp. belong to C. sagittariae. Cercospora alismatis and C. sagittariae are morphologically very close. Inoculation experiments, cultures and data of molecular sequence analyses are necessary to prove if a single or two genuine species are involved. The record of C. alismatis on Echinodorus intermedius from Singapore refers to a collection deposited at BPI on imported plants intercepted at California.

Cercospora apii Fresen. s. lat. (sensu Crous & Braun 2003: 35).

Literature: Soares et al. (2009: 404).

Host range and distribution: On Sagittaria montevidensis, Alismataceae, South America (Brazil, Rio de Janeiro).

Notes: Soares et al. (2009) found a collection of Cercospora on Sagittaria montevidensis characterised by having consistently acicular conidia that can be referred to as C. apii s. lat., at least tentatively.

Cercospora echinodori Chupp, Monograph of Cercospora: 28 (1954).

(Similar to Fig. 1)

Literature: Chupp (1954: 28), El-Gholl et al. (1992: 265), Crous & Braun (2003: 169).

Description: Leaf spots subcircular to somewhat angular-irregular, 2–12 mm diam or confluent and larger, centre grey, margin darker, brown to blackish, indistinctly zonate. Caespituli amphigenous, fine, dark. Mycelium internal. Stromata substomatal or immersed, 10–40 µm diam, subglobose to somewhat irregular, medium brown. Conidiophores in small to moderately large fascicles, loose to dense, arising from stromata, through stomata or erumpent, erect, straight, subcylindrical to somewhat geniculate-sinuous, unbranched, 10–70 µm long, (2.5−)4–6.5 µm wide below and 2–5 µm wide above, pale olivaceous to medium brown, septate, thin-walled, smooth; conidiogenous cells integrated, terminal, occasionally intercalary, 10–30 µm long, conidiogenous loci thickened and darkened, 2–2.5 µm diam. Conidia solitary, acicular, 40–125 × 2.5–5 µm, pluriseptate, hyaline, thin-walled, smooth, apex subobtuse to pointed, base truncate, 2.5–5 µm wide, hila somewhat thickened and darkened.

Holotype: Haiti: Etange, Saumatre, on Echinodorus cordifolius, 12 Apr. 1920, C. Leonard (CUP 39740). Isotype: BPI 4360474.

Host range and distribution: On Echinodorus (berteroi, cordifolius [ovalis]), Alismataceae, West Indies (Cuba, Haiti).

Notes: This species belongs to the Cercospora apii s. lat. complex.

Cercospora osiridis El-Gholl, Proffer & T.S. Schub., Mycotaxon 43: 266 (1992); as “osirisae”.

(Fig. 2)

Fig. 2
figure2

Cercospora osiridis (FLAS F55631). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 300)

Illustration: El-Gholl et al. (1992: 270, figs 1–4).

Description: Leaf spots amphigenous, circular to irregular, 1–14 mm diam, necrotic, brown, with chlorotic halo. Caespituli epiphyllous. Mycelium internal. Stromata lacking. Conidiophores in small to moderately large fascicles (6–22), loose to dense, erect, arising from immersed hyphae, simple or occasionally branched, subcylindrical or 1–6 times geniculate in the upper part, rather long, to 318 µm or even longer, width rather uniform, 3.5–6 µm, pluriseptate, medium brown, wall slightly thickened, smooth; conidiogenous cells integrated, mostly terminal, conidiogenous loci thickened and darkened. Conidia solitary, obclavate, (20−)30–80(−130) × (3−)4–6(−9) µm, 3–7(−12)-septate, hyaline or subhyaline, thin-walled, smooth, apex obtuse, base obconically truncate, hila somewhat thickened and darkened.

Holotype: USA: Florida: Ft. Lauderdale, on Echinodorus uruguayensis, June 1990 (FLAS F55631). Ex-type culture: ATCC 76111.

Host range and distribution: On Echinodorus (uruguayensis [osiris], Echinodorus sp.), Alismataceae, Asia (Singapore), North America (USA, California).

Notes: Material of this species on Echinodorus sp. from Singapore, intercepted in the Port of Los Angeles, is deposited as BPI 744439.

Cercospora sagittariae Ellis & Kellerm., J. Mycol. 2: 1 (1886).

(Fig. 3)

Fig. 3
figure3

Cercospora sagittariae (BPI 440960). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Synonym: Cercosporella macrospora Bres., Hedwigia 36: 201 (1896) [lectotype (designated by Braun 1995): Germany: Saxony: Königstein, on Sagittaria sagittifolia, 2 Sep. 1895, Krieger, Fungi Saxon. Exs. 1298 (HAL)].

Literature: Saccardo (1886: 479), Lindau (1910: 87), Vassiljevsky & Karakulin (1937: 221), Chupp (1954: 29), Katsuki (1965: 8), El-Gholl et al. (1992: 265), Braun (1995: 111), Braun & Mel’nik (1997: 91), Crous & Braun (2003: 362), Guo et al. (2005: 25), Soares et al. (2009: 404–406), Kamal (2010: 83).

Illustrations: Guo et al. (2005: 26, fig. 8), Soares et al. (2009: 405, fig. 3).

Exsiccatae: Ellis & Everh., Fungi Columb. 693. Ellis & Everh., N. Amer. Fungi 1502. Krieger, Fungi Saxon. Exs. 1298. Jaap, Fungi Sel. Exs. 50. Petrak, Crypt. Exs. 4102. Petrak, Mycoth. Gen. 1319. Poelt & Scheuer, Reliqu. Petrak. 2801.

Description: Leaf spots amphigenous, circular to somewhat elliptical-oval or irregular, 3–12 mm diam, occasionally confluent, sometimes vein-limited, straw-coloured, greyish brown to brown, margin brown or with yellowish halo. Caespituli amphigenous, mainly epiphyllous, punctiform, dark or greyish white by abundant sporulation. Mycelium internal. Stromata lacking or small, substomatal or immersed, 10–25 µm diam, brown, composed of swollen hyphal cells, rounded to angular in outline, 4–8 µm diam. Conidiophores in small to moderately large fascicles, mostly 2–15, loose to rather dense, arising from internal hyphae or stromata, through stomata or erumpent, erect, straight to slightly curved or geniculate-sinuous, unbranched, (20−)30–80(−150) × 4–8 µm, 0–4(−8)-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, about 10–30 µm long, conidiogenous loci 1–3(−4) per cell, somewhat thickened and darkened, 2–3 µm diam. Conidia solitary, acicular to obclavate-cylindrical, (15−)25–110(−150) × (2−)3–5.5(−6) µm, (0−)3–8-septate, hyaline, thin-walled, smooth, apex obtuse to pointed, base truncate to short obconically truncate, 2–3 µm wide, hila somewhat thickened and darkened.

In vitro (Soares et al. 2009): Colonies on PDA with compact aerial mycelium, felty, smoke-grey to grey-olivaceous with white sectors, periphery irregular, reaching 2–2.4 cm diam after 10 d, reverse dark green, with distinct pale sectors; on OA aerial mycelium sparse, cottony, white to smoke-grey, periphery irregular, 1.7–2 cm diam after 10 d; on MEA as on OA, reaching 2 cm diam after 10 d, reverse olivaceous-black; cultures not sporulating.

Lectotype (designated here, MycoBank MBT178175): USA: Kansas: Manhattan, on Sagittaria variablilis, Sep. 1884, W. A. Kellerman 681 (BPI 440960). Isolectotype: Ellis & Everh., N. Amer. Fungi 1502, e.g. CUP, OSC.

Host range and distribution: On Sagittaria (arifolia, cuneata, intermedia, lancifolia, latifolia, montevidensis subsp. montevidensis, montevidnesis subsp. calycina [Lophotocarpus calycinus], rigida, sagittifolia, trifolia, variabilis, Sagittaria sp.), Alismataceae, Asia (China, India, Japan, Korea), Europe (Czech Republic, Germany, Hungary, Latvia, Romania, Russia (European part), Ukraine), North America (Canada; USA, Delaware, Florida, Iowa, Illinois, Kansas, Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Nebraska, New York, Oklahoma, Pennsylvania, South Dakota, Texas, Vermont, Virginia, West Virginia, Wisconsin), Central and South America (Argentina, Costa Rica), Oceania (Hawaii), West Indies (Cuba, Puerto Rico, Virgin Islands).

Notes: A true Cercospora s. str. distinct from C. apii s. lat. by rather broad conidiophores and obclavate-cylindrical conidia.

Pseudocercospora

A single species.

Pseudocercospora arthrospora D.J. Soares, R.W. Barreto & U. Braun, Mycologia 101: 409 (2009).

(Fig. 4)

Fig. 4
figure4

Pseudocercospora arthrospora (VIC 30505). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Illustration: Soares et al. (2009: 410, fig. 6).

Description: Leaf spots amphigenous, circular to irregular, initially formed as raised, small, brown dots surrounded by a yellow margin, later forming larger necrotic areas, brown with small grey refracting dots in the centre corresponding to sporulating colonies. Caespituli epiphyllous, dense. Mycelium internal; hyphae branched, 2–3 µm wide, septate, smooth, hyaline to pale olivaceous. Stromata lacking or only with a few swollen hyphal cells. Conidiophores in well-developed, dense fascicles, erect, straight to somewhat curved, subcylindrical, not geniculate, unbranched, 20–40 × 2–4 µm, 1–3-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal, 10–20 µm long, subhyaline, conidiogenous loci indistinct to subdenticulate, 1–3 per cell, 1–2.5 µm diam, but wall not or barely thickened and not darkened. Conidia at first intact, later disarticulating in fragments (arthroconidia occasionally in vivo but often in vitro formed), also forming secondary conidia in branched chains, obclavate, cylindrical, straight to curved, 20–190 × 1.5–3.5 µm, hyaline or subhyaline, thin-walled, smooth, apex obtuse, base truncate to short obconically truncate, 1–2.5 µm wide, hila neither thickened nor darkened.

In vitro: Colonies on PDA slow-growing, reaching 2 cm after 10 d, sporulating abundantly, velvety to chamois-like, reverse black at centre, pale to dark grey towards the margin; on OA aerial mycelium sparse, flat, with irregular margin, light grey, reverse grey; on MEA like on OA.

Holotype: Brazil: Santa Catarina: Forquilhinha, SC 448, rice field at road side, on Sagittaria montevidensis, 21 Apr. 2005, D. J. Soares (VIC 30505).

Host range and distribution: On Sagittaria montevidensis, Alismataceae, South America (Brazil, Paraná, Rio de Janeiro, Santa Catarina).

Notes: This species is unusual due to its disarticulating hyaline or subhyaline conidia, which is a character rarely encountered in Pseudocercospora.

Amaryllidaceae (incl. Allioideae, syn. Alliaceae)

Cercospora

Key to Cercospora species on Amaryllidaceae
  1. 1

    Conidia obclavate-cylindrical, base truncate to somewhat obconically truncate, 20–220 × 4–8 µm; on Scardoxus spp., South Africa ....................................... C. haemanthi

    Conidia acicular or short cylindrical, base truncate ..................................... 2

  2. 2(1)

    Conidia small, 45–50 × 3.5–4 µm, usually 4-septate, cylindrical, ends rounded; on Allium nigrum, Europe ..................................... see Cercospora sancti-marini (doubtful, excluded and insufficiently known species)

    Conidia acicular, much longer ........................................... 3

  3. 3(2)

    Stromata very large, 64–112 µm diam; conidiophores very long, to 640 µm; on Amaryllis belladonna, India ............................................................... C. amaryllidicola

    Stromata smaller, to 70 µm diam; conidiophores usually much shorter ..................................... 4

  4. 4(3)

    Conidia narrow, 20–285 × 2–3.5 µm; on Crinum asiaticum, India ..................................... C. crinicola

    Conidia broader, (2−)3–7.5 µm; on other hosts .................................................. 5

  5. 5(4)

    Stromata lacking or small; on Allium spp. .............................................. C. duddiae

    Stromata well-developed, 25–70 µm diam ...................................................... 6

  6. 6(5)

    Conidiophores very long, 160–405 µm; conidia to 435 µm long; on Allium cepa, Brazil .......................... C. riofranciscana

    Conidiophores shorter, 30–90 µm long; conidia 60–120 µm long; on other hosts of the Amaryllidaceae ........... C. amaryllidis

Cercospora species on Amaryllidaceae

Cercospora amaryllidicola R.C. Rajak, Mycotaxon 10: 457 (1980); as “amaryllicola”.

(Fig. 5)

Fig. 5
figure5

Cercospora amaryllidicola (K(M) 225290). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 55), Kamal (2010: 16).

Illustration: Rajak (1980: 458, fig. 2).

Description: Leaf spots olivaceous-brown to greyish brown, halo indistinct, infections at first at the tip, developing downwards, die-back observed. Caespituli amphigenous, effuse, greyish black. Mycelium internal. Stromata well-developed, globose, 64–112 µm wide, dark brown to greyish black. Conidiophores fasciculate, 6–15, erect, straight to flexuous, subcylindrical to geniculate, unbranched, 225–640 × 5–7 µm, pluriseptate, olivaceous-brown to brown, paler towards the tip, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal or occasionally intercalary, with distinct conidiogenous loci, thickened and darkened. Conidia solitary, acicular-filiform, 70–450 × 4–6.5 µm, 6–25-septate, hyaline, thin-walled, smooth, apex pointed, base truncate or somewhat narrowed at the base, hila thickened and darkened.

Holotype: India: Madhya Pradesh: Jabalpur, Tagore garden, on Amaryllis belladonna, Jan. 1978, R. C. Rayak (K(M) IMI 225290).

Host range and distribution: On Amaryllis belladonna, Amaryllidaceae, Asia (India).

Notes: A true Cercospora s. str. distinct from C. apii s. lat. by having very long conidiophores, to 640 µm, and wider conidia, 72–450 × 4–6.5 µm.

Cercospora amaryllidis Ellis & Everh., J. Mycol. 3: 14 (1887).

(Similar to Fig. 1)

Literature: Saccardo (1892: 653), Lindau (1910: 91), Vassiljevsky & Karakulin (1937: 223), Chupp (1954: 36), Braun (1991: 295), Crous & Braun (2003: 55).

Illustration: Braun (1991: 295, plate 4, fig. 25 B).

Description: Leaf spots lacking or pale, diffuse or subcircular to irregular, 1–5 mm diam. Caespituli amphigenous, often epiphyllous, visible as minute black dots. Mycelium internal. Stromata well-developed, large, 30–70 µm diam, dark brown, composed of swollen hyphal cells, 5–12 µm diam, round to angular in outline, large stromata often rupturing the epidermis, erumpent. Conidiophores in moderately large to large fascicles, 10–35, loose to dense, arising from stromata, erumpent, erect, straight, subcylindrical to geniculate-sinuous, unbranched, 30–90 × 4–7 µm, septate, olivaceous-brown, yellowish brown to medium brown, thin-walled, smooth; conidiogenous cells integrated, terminal, rarely intercalary or conidiophores reduced to conidiogenous cells, 10–25 µm long, with thickened and darkened conidiogenous loci, about 1.5–3 µm diam. Conidia solitary, acicular, straight to curved, 60–120 × (2−)3–5 µm, indistinctly pluriseptate, hyaline, thin-walled, smooth, apex pointed, base truncate or only slightly narrowed at the very base, about 1.5–3 µm wide, hila somewhat thickened and darkened.

Holotype: USA: Louisiana: on Amaryllis sp., 26 Jul. 1886, Langlois 589 (NY 1840467). Isotype: BPI 432570. Topotype: BPI 432569 (collected 18 Nov. 1885).

Host range and distribution: On Agapanthus sp., Amaryllis (belladonna, Amaryllis sp.), Cephyranthes chloroselen (Cooperia drummondii), Cyrthanthus elatus (Amaryllis purpurea), Hippeastrum sp., Hymenocallis sp., Amaryllidaceae, Europe (Germany, Italy), North America (USA, Alabama, Florida, Louisiana, Texas), South America (Colombia), Central America and West Indies (Bermuda, Cuba, El Salvador, Panama, Puerto Rico, Virgin Islands).

Notes: The type specimen of C. amaryllidis is a mixed sample, containing C. amaryllidis (a true Cercospora), and a Pseudocercospora sp. Chupp (1954) confused the issue by describing characters of the Pseudocercospora sp. Braun (1991: 295) confined the name C. amaryllidis to the true Cercospora s. str. and published a redescription. Records of C. amaryllidis on Manfreda maculosa (Asparagaceae) are doubtful. Furthermore, it is not quite clear if all other records of this species are based on correct determinations and refer to true C. amaryllidis or rather to Pseudocercospora spp.

Cercospora crinicola R.K. Srivast., N. Srivast. & A.K. Srivast., Proc. Natl. Acad. Sci. India, B, 64: 107 (1994); as “criniicola”.

(Similar to Fig. 1)

Literature: Crous & Braun (2003: 143), Kamal (2010: 36).

Illustration: Srivastava et al. (1994: 106, fig. 1).

Description: Leaf spots amphigenous, circular to oblong, 2–20 mm diam, various shades of brown or red. Caespituli amphigenous, mainly hypophyllous. Mycelium internal. Stromata subglobose, 35–50 µm diam, substomatal, reddish brown to almost black. Conidiophores in dense fascles, arising from stromata, through stomata, erect, straight, curved to geniculate-sinuous, unbranched, about 50–180 × 3–6 µm, septate, light brown; conidiogenous cells integrated, terminal and intercalary, with thickened and darkened conidiogenous loci. Conidia solitary, acicular, shorter conidia sometimes more cylindrical, straight to somewhat curved, about 20–285 × 2–3.5 µm, 1–28-septate, hyaline, thin-walled, smooth, apex subacute to obtuse, base truncate with somewhat thickened and darkened hilum.

Holotype: India: Uttar Pradesh: Gorakhpur, Botanical Garden of the University, on Crinum asiaticum, Amaryllidaceae, Feb. 1992, R. K. Srivastava (GPU, Gorakhpur University, 1401). Isotype: HCIO 30933.

Host range and distribution: Only known from the type collection.

Note: A true Cercospora s. str. belonging to the C. apii s. lat. complex.

Cercospora duddiae Welles, Phytopathology 13: 362 (1923).

(Fig. 6)

Fig. 6
figure6

Cercospora duddiae (BPI 436011). A. Conidiophore fascicle. B. Conidiophore. C. Conidia. Bar = 10 µm.

Literature: Vassiljevsky & Karakulin (1937: 297), Chupp (1954: 346), Narain & Saksena (1971: 134), Ellis (1976: 271), Little (1987c), Chi (1994: 208), Braun & Mel’nik (1997: 57), Crous & Braun (2003: 167), Guo et al. (2005: 176–177), Kamal (2010: 42).

Illustrations: Narain & Saksena (1971: 135, figs 1–2), Ellis (1976: 271, fig. 205 B), Little (1987c: unnumbered figure), Chi (1994: 209, fig. 230), Guo et al. (2005: 177, fig. 124).

Description: Leaf spots varying on different hosts, but usually beginning as chlorotic spots, circular to oblong, 0.5–6 mm diam, pale brownish, brown, greyish brown or with dingy grey centre, margin indistinct, yellow or somewhat darker. Caespituli amphigenous, usually forming an olivaceous fruiting layer. Mycelium internal. Stromata lacking or small aggregations of swollen hyphal cells, brown. Conidiophores in small to moderately large fascicles, mostly 2–15, loose to moderately dense, arising from internal hyphae or small stromatic aggregations of hyphal cells, through stomata or erumpent, erect, straight, subcylindrical to somewhat geniculate-sinuous, unbranched, (15−)40–180(−475) × 4–8 µm, 0–8-septate, light brown to olivaceous-brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal or intercalary, short conidiophores sometimes reduced to conidiogenous cells, about 10–30 µm long, with a single or several thickened and darkened conidiogenous loci. Conidia solitary, acicular, short conidia sometimes subcylindrical, 25–150(−250) × (2.5−)3–5(−7.5) µm, usually 3–16-septate, hyaline, thin-walled, smooth, apex acute to subobtuse, base truncate or only slightly attenuated at the very base, mostly 2–4 µm wide, hila thickened and darkened.

Syntypes: Philippines: Laguna: Los Baños, on Allium cepa and A. sativum, C. B. Welles (not traced, probably not preserved). Neotype (designated here, MycoBank MBT178132): Philippines: Laguna: College, on Allium cepa, 6 Oct. 1959, A de Dios (BPI 436011).

Host range and distribution: On Allium (ascalonicum, cepa, fistulosum, porrum, sativum, schoenoprasum, tuberosum, Allium sp.), Amaryllidaceae (Allioideae, syn. Alliaceae), Africa (Mauritius, Nigeria, Sierra Leone, Somalia, Tanzania, Togo, Uganda), Asia (Brunei, Cambodia, China, India, Indonesia, Malaysia, Myanmar, Oman, Papua New Guinea, Philippines, Russia, Far East, Sabah, Thailand, Yemen), Australia, Caucasus (Azerbaijan, Georgia), North America (Mexico), Oceania (Palau, Samoa, Vanuatu), West Indies (Barbados, Cuba, Jamaica).

Note: A true Cercospora s. str. with relatively wide conidia (4–6 µm).

Cercospora haemanthi Kalchbr., Grevillea 9: 24 (1881).

(Fig. 7)

Fig. 7
figure7

Cercospora haemanthi (B, holotype). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Saccardo (1886: 477), Chupp (1954: 37), Crous & Braun (1996: 276; 2003: 209).

Description: Leaf spots amphigenous, circular to somewhat angular-irregular, 3–12 mm diam, pale tan to dark olivaceous, margin indefinite or narrow, reddish. Caespituli amphigenous, punctiform, pustulate, blackish, scattered. Mycelium internal. Stromata substomatal, well-developed, 20–80 µm diam, dark brown. Conidiophores in small to usually large fascicles, loose to mostly dense, sometimes almost coremioid, arising from stromata, through stomata, erect, straight to slightly geniculate, usually unbranched, 20–150 × 4–7 µm, aseptate to pluriseptate, pale to medium olivaceous-brown, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal, occasionally conidiophores reduced to conidiogenous cells, 15–50 µm long, conidiogenous loci conspicuous, thickened and darkened, 3–4 µm diam. Conidia solitary, cylindrical-obclavate, 20–220 × 4–8 µm, pluriseptate, mostly with 4–10 septa, hyaline, thin-walled, smooth, apex obtuse, base truncate to somewhat obconically truncate, 3–4 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank MBT178133): South Africa: Cape Province: on Scadoxus puniceus, Jan. 1875, MacOwan 1020 (B 700016004). Isolectotypes: B 700016005, 700016006.

Host range and distribution: On Scadoxus (membranaceus, multiflorus, puniceus [Haemanthus natalensis]), Amaryllidaceae, Africa (Ethiopia, South Africa).

Note: A true Cercospora s. str. distinct from C. apii s. lat. by having much wider obclavate-cylindrical conidia.

Cercospora riofranciscana Bezerra & Peres, Publ. Inst. Micol. Univ. Recife 450: 9 (1965).

(Fig. 8)

Fig. 8
figure8

Cercospora riofranciscana (IMUR 46193). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 356).

Illustration: Batista et al. (1965: 20, fig. 6).

Description: Leaf spots numerous, elliptical or oblong, at first brown, later dark, 0.5–4 mm diam. Mycelium internal; hyphae inter- and intracellular, hyphal cells 7.5–18.5 × 3.5–5.5 µm. Stromata intraepidermal, 25–65 µm diam, brown. Conidiophores fasciculate, subcylindrical-filiform, non-geniculate, unbranched, 160–405 × 3.5–7.5 µm, pluriseptate, brown; conidiogenous cells integrated, terminal, usually with terminal truncate, thickened and darkened conidiogenous locus. Conidia solitary, acicular, straight to curved, 120–435 × 3.5–4 µm, pluriseptate, hyaline, thin-walled, smooth, apex pointed, base truncate, hila thickened and darkened.

Holotype: Brazil: Pernambuco: Rio São Francisco, Belém de Maria, on Allium cepa, 26 May 1965, A. Lustosa Sobrinho (IMUR 46193).

Host range and distribution: On Allium cepa, Amaryllidaceae (Allioideae, syn. Alliaceae), South America (Brazil).

Doubtful, excluded and insufficiently known species

Cercospora sancti-marini (Sacc.) Vassiljevsky, in Vassiljevsky & Karakulin, Fungi Imperfecti Parasitici (Hyphomycetes) 1: 197 (1937).

Basionym: Cercosporina sancti-marini Sacc., Atti Mem. Accad. Sci. Lett. Arti Padova 33: 178 (1917).

Synonym: Cercospora sancti-marini (Sacc.) Chupp, Monograph of Cercospora: 353 (1954).

Literature: Saccardo (1931: 908), Chupp (1954: 353), Crous & Braun (2003: 364).

Description: Leaf spots indistinct. Colonies sparingly effuse, brown. Mycelium internal. Stromata present. Conidiophores mildly geniculate, unbranched, 60–70 × 4.5 µm, very pale olivaceous, paler towards the tip. Conidia cylindrical, 45–50 × 3.5–4 µm, usually 4-septate, hyaline, ends rounded.

Holotype: San Marino: Dogama, on Allium nigrum, Amaryllidaceae (Allioideae, syn. = Alliaceae) [not traced].

Host range and distribution: Only known from the type collection.

Notes: This species is insufficiently known. Type material of this species is not preserved in Saccardo’s herbarium at PAD.

The present description is based on the original publication and Chupp (1954). Without any material it is even unclear if this species represent a true Cercospora s. str.

Cercospora victorialis Thüm., Hedwigia 21: 172 (1882).

Synonyms: Cladosporium alliicola H.D. Shin & U. Braun, Korean J. Mycol. 23: 141 (1995) [holotype: Korea: Suwon, on Allium victorialis, 3 Sep. 1993, H. D. Shin (KUS 12597); isotype: HAL 1533 F.

Cladosporium victorialis (Thüm.) U. Braun & H.D. Shin, in Braun & Mel’nik, Trudy Bot. Inst. im. V.L. Komarova 20: 101 (1997).

Literature: Saccardo (1886: 477), Vassiljevsky & Karakulin (1937: 297), Chupp (1954: 355), Braun & Mel’nik (1997: 101), Crous & Braun (2003: 422), Schubert (2005: 148–149), Bensch et al. (2012: 288).

Illustrations: Shin & Braun (1995: 140–141, figs 1–2), Braun & Mel’nik (1997: fig. 71), Schubert (2005: 149, fig. 72, pl. 33, figs A–D), Bensch et al. (2012: 288–289, figs 338–339).

Lectotype (designated by Braun, in Braun & Mel’nik 1997): Russia: West Siberia, Mt Kerlygan, on Allium victorialis, Martjanov (LE 404551). Isolectotype: LE 404552.

Host range and distribution: On Allium (cepa, sativum, victorialis [ochotense]), Amaryllidaceae (Allioideae, syn. Alliaceae), Asia (Japan, Korea, Russia) South America (Brazil).

Pseudocercospora

Key to Pseudocercospora species on Amaryllidaceae
  1. 1

    Conidiophores narrow, 1.5–2.5 µm; conidia 9–39 × 1.5–3.5 µm; on Zephyranthes rosea, India ................................ P. indica

    Conidiophores wider, 2.5–6(−8) µm; conidia longer and broader, 20–125 × 2–6 µm; on various hosts of the Amaryllidaceae ........................................................ P. pancratii

Pseudocercospora species on Amaryllidaceae

Pseudocercospora indica D. Gupta, Padhi & Chowdhry, Sci. Cult. 48: 112 (1982).

(Fig. 9)

Fig. 9
figure9

Pseudocercospora indica (based on Gupta et al. 1982; 112, fig. 1–2). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Illustration: Gupta et al. (1982: 112, figs 1–2).

Description: Leaf spots amphigenous, linear, starting at the tip, proceeding downwards, extending to 10–65 mm in length, yellowish brown, margin black. Caespituli hypophyllous, punctiform, dark. Mycelium internal. Stromata well-developed, large, immersed, subglobose to globose, 45–75 µm diam, dark brown. Conidiophores numerous, in dense fascicles, almost sporodochial, arising from stromata, erect, straight, subcylindrical, barely geniculate, unbranched, 9–36 × 1.5–2.5 µm, deep olivaceous-brown, 0–2-septate, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, conidiogenous loci inconspicuous, unthickened, not darkened. Conidia solitary, obclavate-cylindrical, straight to curved, short and narrow, (9−)12–39 × 2–3.5 µm, 2–5-septate, pale to medium olivaceous-brown, thin-walled, smooth, apex obtuse, base short obconically truncate, hila unthickened, not darkened.

Holotype: India: Odisha: Bhubaneswar, botanical garden of the Utkal University, on Zephyranthes rosea, Amaryllidaceae, 26 Feb. 1978, D. Gupta (HCIO 32865).

Host range and distribution: Only known from the type collection.

Notes: Pseudocercospora indica Deighton (Deighton 1987), described from India on Uraria picta, is a homonym and needs a new name: Pseudocercospora urariigena U. Braun, nom. nov. [MycoBank MB809007. Basionym: Pseudocercospora indica Deighton, Trans. Brit. Mycol. Soc. 88: 371 (1987).]

Pseudocercospora pancratii (Ellis & Everh.) U. Braun & R.F. Castañeda, Cryptog. Bot. 2: 294 (1991).

(Fig. 10)

Fig. 10
figure10

Pseudocercospora pancratii (NY 00838178). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora pancratii Ellis & Everh., J. Mycol. 3: 15 (1887).

Synonyms: Cercospora hymenocallidis Pat., Bull. Trimestriel Soc. Mycol. France 28: 142 (1912) [holotype: Costa Rica: San José, museum garden, Jul. 1908, “Herb. Inst. Physio-geogr. Nat. costaricensis no. 11”, herb. Patouillard 7828 (FH)].

Cercosporidium hymenocallidis (Pat.) Sacc., Syll. Fung. 25: 895 (1931).

Missaplied name: Cercospora amaryllidis sensu Chupp (1954: 36) p.p.

Literature: Saccardo (1892: 654), Chupp (1954: 37), Vasudeva (1963: 157), Katsuki (1965: 9), Crous & Braun (2003: 306), Kamal (2010: 207), Braun & Urtiaga (2012: 318), Kirschner (2014: 487–488).

Illustration: Braun & Castañeda Ruiz (1991: 295, fig. 25 A), Kirschner (2014:488, fig.5).

Description: Leaf spots amphigenous, 1–20 mm diam or sometimes covering large leaf segments, subcircular to irregular, pale, greyish, brown, reddish, occasionally somewhat zonate, margin indefinite or surrounded by a diffuse darker to blackish, sometimes reddish margin. Caespituli amphigenous, punctiform, blackish to dingy grey by abundant conidial formation, when abundant in dark grey layers. Mycelium internal; hyphae branched, septate, 1–4 µm wide, hyaline to medium brown, thin-walled, smooth. Stromata well-developed, substomatal, 10–80(−125) µm diam, sometimes rupturing the stomata, often somewhat erumpent, brown to dark brown, visible as small black dots. Conidiophores in dense, rich fascicles, arising from, stromata, through stomata, straight and subcylindrical-lageniform to strongly flexuous, geniculate-sinuous, simple, rarely branched, 5–50 × 2.5–6(−8) µm, 0(−2)-septate, subhyaline to olivaceous-brown or medium brown, thin-walled, smooth; conidiophores mostly reduced to conidiogenous cells, occasionally with integrated, terminal conidiogenous cells, 5–25 µm long, conidiogenous loci inconspicuous, unthickened, not darkened. Conidia solitary, obclavate, subcylindrical, subacicular-filiform, 20–125 × 2–6 µm, 0–10-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex obtuse to subacute, base truncate to short obconically truncate, 1–3.5 µm wide, hila neither thickened nor darkened.

Holotype: USA: Louisiana: On Hymenocallis crassifolius, June 1886, Langlois 656 (NY 00838178). Isotype: BPI 457098.

Host range and distribution: On Agapanthus africanus [umbellatus], Ammocharis longifolia [Crinum longifolium], Crinum (americanum, asiaticum var. japonicum, zeylanicum, Crinum sp.), Hippeastrum (puniceum [equestre], Hippeastrum sp.), Hymenocallis (arenicola, caribaea [declinata], coronaria, crassifolia, latifolia, littoralis, tubiflora), Iris sp., Pancratium sp., Sprekelia formosissima, Amaryllidaceae, Africa (Kenya), Asia (India, Karnataka; Japan, Myanmar, Taiwan), Central America and West Indies (Barbados, Bermuda, Costa Rica, Cuba, Puerto Rico, Virgin Islands), South America (Brazil, Venezuela), North America (USA, Alabama, Florida, Louisiana, Mississippi, Texas).

Notes: Kirschner (2013) observed immature asci in stromata of a collection from Taiwan.

Zasmidium

A single species.

Zasmidium hymenocallidis (U. Braun & Crous) U. Braun & Crous, Schlechtendalia 20: 101 (2010).

(Fig. 11)

Fig. 11
figure11

Zasmidium hymenocallidis (NY 01043037). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Stenella hymenocallidis U. Braun & Crous, Mycotaxon 92: 403 (2005).

Illustration: Braun & Crous (2005: 411, fig. 8).

Exsiccatae: Ellis & Everh., North Amer. Fungi 1767.

Description: Lesions diffuse, shape and size variable, brown to reddish brown, margin indefinite. Caespituli amphigenous, punctiform, scattered, greyish brown. Mycelium internal; hyphae branched, septate, 2–5 µm wide, subhyaline, thin-walled, smooth. Stromata substomatal, 10–50 µm diam, brown, composed of swollen hyphal cells, 2–7 µm diam, walls somewhat thickened. Conidiophores in small to moderately large fascicles, loose to very dense, arising from stromata, emerging through stomata, erect, straight, subcylindricalconical to slightly geniculate-sinuous, unbranched, 5–30 × 3–5 µm, 0–2-septate, subhyaline to pale olivaceous-brown, thin-walled, smooth or almost so; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–20 µm long, conidiogenous loci conspicuous, somewhat thickened and darkened, 1–2 µm wide. Conidia solitary, short conidia ellipsoid-ovoid to short cylindrical, longer conidia cylindrical-filiform, subacicular, occasionally narrowly obclavate-cylindrical, (15−)20–110 × 2.5–4 µm, (0−)1–8-septate, without any constrictions at septa, subhyaline, pale olivaceous to olivaceous-brown, occasionally pale brown, thin-walled, verruculose, apex obtuse to subacute, base truncate to short obconically truncate, 1–1.5 µm wide, hila slightly thickened and darkened.

Holotype: USA: Louisiana: Point a la Hache, on living and senescing leaves of Hymenocallis crassifolia, June 1886, A. B. Langlois [Ellis & Everh., North Amer. Fungi 1767] (NY 01043037).

Host range and distribution: Only known from the type collection.

Notes: This is one of the few Zasmidium species without any superficial hyphae in vivo. However, molecular analyses of comparable species with lacking superficial mycelium revealed that such taxa belong in Zasmidium (see Braun et al. 2013, discussion under Zasmidium).

Aponogetonaceae

Cercospora

A single species.

Cercospora aponogetonicola M.S. Pavgi & L. Singh, Hydrobiologia 53: 89 (1977).

(Similar to Fig. 1)

Literature: Crous & Braun (2003: 62), Kamal (2010: 17).

Illustration: Pavgi & Singh (1977: 89, figs 5–8).

Description: Leaf spots amphigenous, on floating leaves, subcircular to somewhat angular-irregular, 1–4 mm diam, often confluent, pale olivaceous-brown, margin indistinct to darker. Caespituli amphigenous, dark, punctiform. Mycelium internal. Stromata substomatal, according to the original description large, about 70–110 µm diam. Conidiophores in loose to moderately large fascicles, arising from stromata, through stomata, erect, straight, subcylindrical to slightly geniculate, unbranched, 30–180 × 3.5–7.5 µm, pluriseptate, brown throughout or paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, about 10–30 µm long, conidiogenous loci thickened and darkened, 2.5–3 µm wide. Conidia solitary, acicular, young conidia occasionally somewhat obclavate-cylindrical, 60–280 × 3–5 µm, 5–17-septate, hyaline or subhyaline, very pale greenish olivaceous, thin-walled, smooth, apex obtuse to acute, base truncate, occasionally somewhat obconically truncate, 2–3 µm wide, hilum somewhat thickened and darkened.

Holotype: India: Uttar Pradesh: Varanasi, on Aponogeton crispus, Aponogetonaceae, 19 Sep. 1969, L. Singh (HCIO 31612). Isotype: K(M) IMI 174885.

Host range and distribution: Only known from the type collection.

Note: A true Cercospora s. str. close to C. apii s. lat.

Araceae

Cercospora

Tabular key to Cercospora species on Araceae according to host genera
Alocasia  
A single species ...................................................... C. alocasiae
Amorphophallus  
1   Conidia acicular, base truncate .......................................................... C. amorphophalli
1   Conidia obclavate-cylindrical, base obconically truncate ........................................ 2
2(1)   Conidia narrow, 20–100 × 2–3.5 µm, base 1–2 µm wide .................................... C. amorphophallicola Conidia broader, 30–140(−230) × 3–7 µm, base 2–3 µm wide .................................... C. chevalieri
Anchomanes  
1   Leaf spots 1–11 mm diam; conidiophores in dense fascicles, 35–75 µm long, 0–2-septate; conidia obclavate-subcylindrical, 35–90 × 5–6 µm .................................... C. anchomanes Leaf spots much larger, 10–30 mm diam; conidiophores in loose to moderately dense fascicles, much longer and pluriseptate, 50–300(−350) µm; conidia cylindrical, rarely obclavate-cylindrical, 30–165 × 4–5.5 µm ...................................................... C. anchomanicola
Anthurium  
A single species ............................................................... C. richardiicola
Arum  
1   Conidia cylindrical, 10–50 × 3–4 µm, 1–4-septate ......................................... C. ari Conidia obclavate, 40–70 × 4.5–5 µm, 8–12-septate .................................... C. aricola
Arisaema  
A single species ............................................... C. arisaematis
Caladium  
1   Conidia acicular, 25–260 × 2.5–5 µm, pluriseptate .................................... C. caladii Conidia broadly ellipsoid-ovoid, obovoid to fusiform-subcylindrical, 15–55(−70) × 5–9.5 µm, 2–7-septate .................................................................. C. verruculosa
Calla  
A single species .......................................................................... C. callae
Colocasia  
A single species .......................................................................... C. alocasiae
Cryptocoryne  
A single species .......................................................................... C. cryptocorynes
Epipremnum  
A single species .......................................................................... C. richardiicola
Monstera  
A single species .......................................................................... C. monsterae
Peltrandra  
A single species .......................................................................... C. callae
Philodendron  
A single species .......................................................................... C. apii s. lat.
Pistia  
1   Conidia narrowly acicular, short conidia sometimes obclavate-cylindrical, 30–300 × 2–4 µm ........................ C. alocasiae Conidia broadly acicular to obclavate-cylindrical, 30–150 × 3–6.5 µm ................................................... C. pistiae
Plesmonium  
A single species .......................................................................... C. chevalieri
Pothos  
A single species .......................................................................... C. bombycina
Symplocarpus  
A single species .......................................................................... C. symplocarpi
Syngonium  
A single species .......................................................................... C. syngoniicola
Typhonium  
A single species .......................................................................... C. typhoniae
Xanthosoma  
A single species .......................................................................... C. chevalieri
Zantedeschia  
1   Conidia acicular, 25–300 × 2–4 µm, 2–20-septate ............................................................ C. richardiicola Conidia broadly obclavate-cylindrical, 25–110 × 4–8 µm, 2–8-septate .................................................... C. callae

Cercospora species on Araceae

Cercospora alocasiae Goh & W.H. Hsieh, Trans. Mycol. Soc. Republ. China 2: 86 (1987).

(Similar to Fig. 1)

Synonyms: Cercospora alocasiae Sawada, Taiwan Agric. Rev. 38: 693 (1942), nom. inval. (ICN, Art. 39.1).

Cercospora colocasigena S. Narayan, Kharwar, R.K. Singh & Bhartiya, Kavaka 25: 88 (1998) [holotype: India: Uttar Pradesh: Gorakhpur, on Colocasia esculenta, Feb. 1992, S. Narayan (HCIO 419320); isotype: GPU 5003.

Literature: Chupp (1954: 55), Katsuki (1965: 11), Hsieh & Goh (1990: 28), Guo & Jiang (2000a), Crous & Braun (2003: 52), Guo et al. (2005: 34–35), Kamal (2010: 14), Phengsintham et al. (2013b: 46).

Illustrations: Hsieh & Goh (1990: 29, fig. 15), Narayan et al. (1997: 89, fig. 2), Guo et al. (2005: 35, fig. 15), Phengsintham et al. (2013b: 48, figs 4–5).

Description: Lesions indistinct or with distinct leaf spots, 1–25 mm diam, subcircular to somewhat irregular or oblong, brownish to grey or greyish white, sometimes zonate, margin indefinite to distinct, brown. Colonies amphigenous, effuse, dark. Mycelium internal; hyphae branched, septate, 2–4 µm diam. Stromata lacking or relatively small, about 10–30 µm diam, substomatal to immersed, olivaceous-brown. Conidiophores solitary or in small to moderately large fascicles, loose to dense, arising from internal hyphae or stromata, through stomata or erumpent, erect, straight, subcylindrical to geniculate, unbranched, 10–200 × 3.5–6 µm, continuous to pluriseptate, olivaceous to brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, 15–40 µm long, with conspicuous conidiogenous loci, thickened and darkened, about 1.5–2.5 µm diam. Conidia solitary, acicular, shorter conidia often somewhat obclavate-subcylindrical, straight to curved, 30–200 × 2–4 µm, about 3–24-septate, hyaline, thin-walled, smooth, apex subobtuse to pointed, base truncate to obconically truncate in shorter conidia, about 1.5–3 µm wide, hila thickened and darkened.

Holotype: Taiwan: Taipei, on Alocasia macrorrhizos, 27 Apr. 1921, K. Sawada (NTU-PPE [hb. Sawada]).

Host range and distribution: On Alocasia (macrorrhizos [indica, odora], Alocasia sp.), Arum sp., Colocasia esculenta, Pistia stratiotes, Araceae, Asia (China, India, Japan, Laos, Myanmar, Nepal, Taiwan), South America (Venezuela), West Indies (Cuba).

Notes: This species belongs to the Cercospora apii s. lat. complex. Kamal (2010) examined type material of C. colocasigena and reduced it to synonymy with C. alocasiae. Authentic collections for C. alocasiae Sawada are deposited at BPI (432460, 432461). Thaung (1984) recorded this species from Myanmar on Arum sp. Previous records of “C. caladii” on Colocasia esculenta from China and India belong to the present species.

Cercospora amorphophalli Henn., Hedwigia 41: 147 (1902).

(Fig. 12)

Fig. 12
figure12

Cercospora amorphophalli (B, holotype). A. Conidiophores. B. Conidia. Bar = 10 µm.

Synonym: Cercospora aracearum Firdousi, A.N. Rai & K.M. Vyas, Indian Phytopathol. 44: 225 (1991) [holotype: India: Madhyar Pradesh: Sagar, Gopalpura, South Forest Division, on Amorphophallus margaritifer, Aug. 1988, S. A. Firdousi (K(M) IMI 329641)].

Literature: Saccardo (1906: 611), Chupp (1954: 56), Boedijn (1961: 412), Katsuki (1965: 11), Guo & Jiang (2000a), Braun (2001a: 431), Crous & Braun (2003: 55, 62), Guo et al. (2005: 36), Kamal (2010: 16, 18).

Illustrations: Firdousi et al. (1991: 226, fig. 1), Braun (2001a: 433, fig. 16 B), Guo et al. (2005: 36, fig. 16).

Description: Leaf spots amphigenous, circular or subcircular, 1–8 mm diam, pale brownish, grey, greyish white, margin ochraceous to brown. Caespituli amphigenous, mostly hypophyllous, delicately punctiform, dark brown to blackish. Mycelium internal. Stromata lacking or almost so to well-developed, small to moderately large, 10–55 µm diam, substomatal to intraepidermal, yellowish to medium brown or olivaceous-brown. Conidiophores loosely fasciculate, about 2–20, arising from internal hyphae or stromata, through stomata or erumpent, erect, straight, subcylindrical to geniculate in the upper portion, unbranched, 8–250 × (2.5−)3–6(−7.5) µm, aseptate to pluriseptate, pale to medium brown or olivaceous-brown throughout or paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, about 10–30 µm long, with a single to several conspicuously thickened and darkened conidiogenous loci, (2−)3–4 µm diam. Conidia solitary, acicular, straight to somewhat curved, 10–110 × 2–5 µm, 3–24-septate, hyaline, thin-walled, smooth, apex pointed, base truncate, 3–4 µm wide, hila somewhat thickened and darkened.

Holotype: Indonesia: Java: Buitenzorg, on Amorphophallus sp., Dec. 1901, Zimmermann (B 700016010).

Host range and distribution: On Amorphophallus (paeoniifolius [campanulatus], konjac [rivieri], margaritifer, variabilis, Amorphophallus sp.), Araceae, Africa (Sudan), Asia (China, India, Indonesia, Japan, Philippines, Thailand).

Note: This species belongs to the Cercospora apii complex.

Cercospora amorphophallicola U. Braun, Nova Hedwigia 73: 430 (2001).

(Fig. 13)

Fig. 13
figure13

Cercospora amorphophallicola (L 53844). A. Conidiophores. B. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 56).

Illustration: Braun (2001a: 433, fig. 16 A).

Description: Leaf spots amphigenous, subcircular to angular-irregular, 1–15 mm diam, pale greenish, later pale to dark brown, blackish or reddish brown, finally greyish brown to dingy grey, margin indefinite or with a narrow, dark brown to blackish border or marginal line, sometimes somewhat raised or limited by veins, occasionally with a diffuse halo. Caespituli amphigenous, punctiform, dark, mostly scattered. Mycelium internal. Stromata lacking or small, 10–30 µm diam, dark brown, intraepidermal, rarely substomatal. Conidiophores in small to moderately large fascicles, loose to dense, arising from internal hyphae or stromata, mostly erumpent, occasionally emerging through stomata, erect, subcylindrical, in the upper half conspicuously geniculate-sinuous, unbranched, 20–120 × 3–7 µm, pluriseptate throughout, pale to medium dark brown or olivaceous-brown, paler towards the tip, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal, occasionally intercalary, 10–30 µm long, conidiogeous loci conspicuous, thickened and darkened, 1–2 µm diam. Conidia solitary, narrowly obclavate-subcylindrical, 25–100 × 2–3.5 µm, 1–8-septate, hyaline or subhyaline, smooth, apex subobtuse to subacute, base obconically truncate to subtruncate, 1–2 µm wide, hila thickened and darkened.

Holotype: Indonesia: Java: Bogor, Botanical Garden, on Amorphophallus sp., May 1950, K. B. Boedijn (L 53844).

Host range and distribution: On Amorphophallus sp., Araceae, Asia (Indonesia).

Note: This species differs from C. amorphophalli in having obclavate-subcylindrical conidia with obconically truncate base and much smaller conidiogenous loci.

Cercospora anchomanes J.M. Yen & Gilles, Cah. Maboké 8: 75 (1970).

(Fig. 14)

Fig. 14
figure14

Cercospora anchomanes (based on Yen & Gilles 1970: 75, fig. 1). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Literature: Kranz (1963, as C. chevalieri), Crous & Braun (2003: 57).

Illustration: Yen & Gilles (1970: 75, fig. 1).

Description: Leaf spots amphigenous, circular to irregular, 1–11 mm diam, mostly confluent, brown to whitish, border darker brown or yellowish. Caespituli amphigenous, usually hypophyllous. Mycelium internal. Stromata in the stomatal cavity, subglobose, 25–80 µm diam, pigmented. Conidiophores in dense fascicles, arising from stromata, through stomata, erect, straight to flexuous or somewhat geniculate, unbranched, 35–95 × 5 µm, 0–2-septate, pale brown, thin-walled, smooth; conidiogenous cells integrated, terminal, conidiogenous loci conspicuous, thickened and darkened. Conidia solitary, obclavate-subcylindrical, straight to mostly curved, 35–90 × 5–6 µm, hyaline, thin-walled, smooth, apex obtuse, base somewhat obconically truncate, hila thickened and darkened.

Holotype: Gabon: on Anchomanes difformis, 25 Dec. 1969, G. Gilles 30 (not traced).

Host range and distribution: On Anchomanes (difformis, giganteus), Araceae, Africa (Gabon, Guinea).

Note: Type material was expected to be deposited at PC, but could not be traced there.

Cercospora anchomanicola J.M. Yen & Gilles, in Yen, Bull. Trimestriel Soc. Mycol. France 90: 310 (1974).

(Fig. 15)

Fig. 15
figure15

Cercospora anchomanicola (PC, topotype). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 57).

Illustration: Yen (1974: 311, fig. 3).

Description: Leaf spots amphigenous, formed as greenish to greyish discolorations with indistinct margin and yellowish to ochraceous halo, subcircular to somewhat irregular, 10–30 mm diam. Caespituli amphigenous, scattered, effuse, punctiform, brown. Mycelium internal. Stromata substomatal, 20–75 µm diam, subglobose, brown. Conidiophores in small to moderately large fascicles, loose to moderately dense, arising from stromata, through stomata, erect, straight, subcylindrical to geniculate-sinuous, unbranched, 50–300(−350) × 4–6 µm, pluriseptate throughout, pale olivaceous-brown to brown, paler towards the apex, thin-walled, smooth; conidiogenous cells integrated, terminal or intercalary, 10–40 µm long, conidiogenous loci conspicuous, thickened and darkened, (2−)2.5–3(−3.5) µm diam. Conidia solitary, cylindrical (-obclavate-cylindrical), straight to somewhat curved, 30–165 × 4–5.5 µm, 3–17-septate, hyaline or subhyaline, thin-walled, smooth, apex obtuse, rounded, base truncate to slightly obconically truncate, 2.5–3.5 µm wide, hila thickened and darkened.

Holotype: Ivory Coast: Abidjan, on Anchomanes difformis, 9 Jul. 1972, G. Gilles (not traced).

Host range and distribution: On Anchomanes difformis, Araceae, Africa (Ivory Coast).

Notes: A true Cercospora s. str. distinct from C. apii s. lat. by having cylindrical to somewhat obclavate-cylindrical conidia. Type material could not be traced at PC, but topotype material, collected in 1973, is deposited in PC (Ivory Coast, Abidjan, on Anchomanes difformis, 10 Feb. 1973, G. Gilles, PC), and has been examined. An additional topotype from 1976 is deposited as K(M) IMI 212938.

Cercospora apii Fresen. s. lat. (sensu Crous & Braun 2003: 35).

Literature: Braun & Urtiaga (2013b).

Host range and distribution: On Philodendron sp., Araceae, South America (Venezuela).

Cercospora arisaematis F.L. Tai, Chinese Bot. Soc. Bull. 2: 47 (1936); as “arisemae”.

(Fig. 16)

Fig. 16
figure16

Cercospora arisaematis (HMAS 06993). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 56), Katsuki (1965: 11), Hsieh & Goh (1990: 29), Crous & Braun (2003: 65), Guo et al. (2005: 37), Kamal (2010: 18), Bhat & Pratibha (2010).

Illustrations: Hsieh & Goh (1990: 30, fig. 16), Guo et al. (2005: 37, fig. 17).

Description: Leaf spots circular or subcircular, 0.5–10 mm diam, at first water-soaked, later brown to dingy grey, margin darker brown. Caespituli amphigenous, punctiform, dark. Mycelium internal. Stromata lacking or small, 10–20 µm diam, substomatal, brown. Conidiophores in small fascicles, 2–9, loose, arising from internal hyphae or stromatic hyphal aggregations, through stomata, erect, straight, subcylindrical to geniculate-sinuous, unbranched, 10–140(−425) × 4–6.5(−8) µm, aseptate to pluriseptate, pale olivaceous to light brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, sometimes conidiophores reduced to conidiogenous cells, about 10–40 µm long, conidiogenous loci thickened and darkened, about 2–3 µm diam. Conidia solitary, acicular to obclavate-subcylindrical, straight to somewhat curved, 20–130 × 2.5–5 µm, 2–15-septate, hyaline or with a very pale olivaceous tinge, thin-walled, smooth, apex subobtuse to subacute, base truncate to obconically truncate, hila thickened and darkened.

Holotype: China: Jiangsu: Nanjing, on Arisaema heterophyllum, 27 Sep. 1925, T. F. Yu (HMAS 06993).

Host range and distribution: On Arisaema (heterophyllum [ambiguum], murrayi, Arisaema sp.), Araceae, Africa (Uganda), Asia (China, India, Japan, Taiwan).

Notes: Records of C. arisaematis on Typhonium roxburghii [divaricatum] from Mainland China and Taiwan (Hsieh & Goh 1990, Guo & Jiang 2000a) belong undoubtedly to C. typhonii.

Cercospora bombycina Chidd., Sydowia 13: 154 (1959).

(Fig. 17)

Fig. 17
figure17

Cercospora bombycina (BPI 433778). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Synonym: Cercospora aracearum S. Narayan, R.N. Kharwar, R.K. Singh & H.S.G. Rao, J. Indian Bot. Soc. 80: 213 (2001), nom. illeg. (ICN, Art. 53.1) [holotype: India: Uttar Pradesh: on Pothos scandens, R. K. Kharwar (HCIO 42013)].

Literature: Crous & Braun (2003: 85), Kamal (2010: 24).

Illustration: Chiddarwar (1959: plate 4, figs 8–9).

Description: Leaf spots amphigenous, angular-irregular, 3–6 mm diam, scattered, pale to medium brown, border distinct, darker brown. Caespituli amphigenous, finely punctiform, scattered, dark. Mycelium internal. Stromata substomatal, moderately large, 15–45 µm diam, compact, brown. Conidiophores in small to moderately large fascicles, usually 4–30, arising from stromata, emerging through stomata, loose, straight, subcylindrical to distinctly geniculate-sinuous, unbranched, 50–130 × 3.5–5 µm, pluriseptate, mostly with 2 to 7 septa, brown, thin-walled, smooth; conidiogenous cells integrated, terminal or intercalary, about 10–30 µm long, conidiogenous loci thickened and darkened, about 1.5–2 µm diam. Conidia solitary, obclavate, 50–130 × 3–4.5 µm, 4–12-septate, hyaline, thin-walled, smooth, apex subobtuse to subacute, base obconically truncate, about 1.5–2.5 µm wide, hila somewhat thickened and darkened.

Holotype: India: Maharashtra: Pune, on Pothos scandens, 22 Oct. 1954, P. P. Chiddarwar 3 (K(M) IMI 83164). Isotype: BPI 433778.

Host range and distribution: On Pothos (scandens, Pothos sp.), Araceae, Asia (India, Madhya Pradesh, Maharashtra, Uttar Pradesh).

Note: A true Cercospora s. str. distinct from C. apii s. lat. by its obclavate conidia.

Cercospora caladii Cooke, Grevillea 8: 95 (1880).

(Similar to Fig. 1)

Literature: Saccardo (1886: 478), Chupp (1954: 57), Vasudeva (1963: 58), Crous & Braun (2003: 93), Guo et al. (2005: 38), Kamal (2010: 26).

Illustration: Guo et al. (2005: 39, fig. 18).

Description: Leaf spots circular, 0.5–5 mm diam, dingy grey to brown, margin reddish brown. Caespituli amphigenous. Mycelium internal. Stromata lacking or small, brown. Conidiophores in small to moderately large fascicles, 2–12, arising from internal hyphae or small stromata, erect, straight to curved, subcylindrical to somewhat geniculate above, unbranched, 20–475 × 4–6.5 µm, continuous to pluriseptate, pale to medium brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, conidiogenous loci conspicuous, thickened and darkened, about 2.5–3 µm diam. Conidia solitary, acicular, straight to curved, 25–260 × 2.5–5 µm, pluriseptate, hyaline, apex subacute, base truncate, hila thickened and darkened.

Holotype: India: Karnataka: Belgaum, on Caladium sp., 1879, J. Hobson (K(M) 190717).

Host range and distribution: On Araceae sp., Caladium (bicolor, Caladium sp.), Araceae, Asia (India), South America (Brazil, Honduras), West Indies (Puerto Rico, Virgin Islands).

Notes: Records of “C. caladii” on Colocasia esculenta from Asia (China and India) do not belong to this species. They are excluded and referred to as C. alocasiae. Caladium (Caladieae) on the one hand and Alocasia/Colocasia (Pistia clade) on the other hand are not closely allied (Nauheimer et al. 2012).

Cercospora callae Peck & Clinton, Rep. (Annual) New York State Mus. Nat. Hist. 29: 52 (1876).

(Fig. 18)

Fig. 18
figure18

Cercospora callae (NYS, holotype). A. Conidiophore fascicle. B. Conidophore tips. C. Conidia. Bar = 10 µm.

Synonyms: Cercospora pachyspora Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia I, 43: 88 (1891) [lectotype (designated here, MycoBank MBT178134): USA: Delaware: Wilmington, on Peltandra virginica, 4 Oct. 1889, A. Commons, no. 1013 (NY 1840471)].

Cladosporium callae Peck & Clinton, in herb.

Literature: Saccardo (1886: 478–479; 1892: 654), Vassiljevsky & Karakulin (1937: 222), Chupp (1954: 57), El-Gholl et al. (1992: 265), Crous & Braun (2003: 94), Schubert & Braun (2007: 190).

Illustration: Schubert & Braun (2007: 191, fig. 1).

Exsiccatae: Ellis, N. Amer. Fungi 1253.

Description: Leaf spots mostly oblong, between veins, sometimes forming larger patches, extending from the mid-rib to the leaf margin, pale greenish, medium brown to blackish, later sometimes dingy grey. Caespituli amphigenous, mainly hypophyllous, scattered to dense, forming mouse-grey, downy layers. Mycelium internal. Stromata substomatal to intraepidermal, 10–60 µm diam, occasionally larger, pale, yellowish green to brownish. Conidiophores in small, loose to usually large and dense to very dense fascicles, arising from stromata, emerging through stomata or erumpent, erect, straight to mostly distinctly geniculate-sinuous unbranched, 50–150 × 4–9 µm, aseptate to pluriseptate throughout, pale yellowish to olivaceous-brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, occasionally conidiophores reduced to conidiogenous cells, conidiogenous loci conspicuous, thickened and darkened, about 2–3 µm wide. Conidia solitary, broadly obclavate-cylindrical, 25–110 × 4–8 µm, 2–9-septate, hyaline or subhyaline, thin-walled, smooth, apex obtuse, broadly rounded, base truncate to short obconically truncate, 2–4 µm wide, hila thickened and darkened.

Holotype: USA: New York: Buffalo, on Calla palustris, G. W. Clinton (NYS).

Host range and distribution: On Calla palustris, Peltandra (sagittifolia [alba, glauca], virginica), Zantedeschia aethiopica, Araceae, Europe (Belarus), North America (Canada, Ontario; USA, Alabama, Delaware, Florida, Maryland, Massachusetts, Minnesota, Mississippi, New Hampshire, New York, Wisconsin).

Notes: A true Cercospora s. str. distinct from C. apii s. lat. There are three syntypes of Cercospora pachyspora in NY [Commons 1013 (NY 1840471), Commons 1014 (NY 180472) and a third as 1013/1014 (NY 1840473)]. The host of Commons 1014 was indicated to be Alisma plantago-aquatica, which is, however, doubtful. The three collections have recently been checked by an expert for vascular plants in NY and it turned out that all of them, including Commons 1014 on “Alisma”, belong to Peltandra virginica. Syntype material ‘Commons 1013’ is designated as lectotype.

Cercospora chevalieri Sacc., Syll. Fung. 22: 1431 (1913).

(Fig. 19)

Fig. 19
figure19

Cercospora chevallieri (FH 7807). A. Conidiophore fascicle. B. Conidiophore. C. Conidia. Bar = 10 µm.

Synonyms: Cercospora amorphophalli Pat. & Har., Bull. Trimestriel Soc. Mycol. France 24: 15 (1909), nom. illeg. (ICN, Art. 53.1).

Cercosporina chevalieri (Sacc.) Sacc., Syll. Fung. 25: 896 (1931).

Literature: Saccardo (1913: 1431), Chupp (1954: 58), Ellis (1976: 244), Crous & Braun (2003: 119), Kamal (2010: 31).

Illustration: Ellis (1976: 245, fig. 184).

Description: Leaf spots amphigenous, subcircular to angular-irregular, small, yellowish to ochraceous, brown or greyish white, margin narrow, darker brown to blackish. Caespituli amphigenous, but mainly hypophyllous, punctiform to dense, dark. Mycelium internal. Stromata substomatal to immersed, subglobose, dark brown, 20–60 µm diam. Conidiophores in small, loose to large and dense fascicles, arising from stromata, through stomata or erumpent, erect, straight to mostly distinctly to strongly geniculate-sinuous, unbranched, 50–150 × 4–8 µm, pluriseptate throughout, pale olivaceous to mid pale brown throughout, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal and intercalary, about 10–35 µm long, conidiogenous loci thickened and darkened, about 2–3 µm diam. Conidia formed singly, broadly obclavate to obclavate-cylindrical, 30–140(− 230) × 3–7 µm, 3–14(−24)-septate, hyaline or subhyaline, thin-walled, smooth, apex obtuse, base obconically truncate, 2–3 µm wide, hila thickened and darkened.

Holotype: Sudan: Territoir de L’Oubangui, at Mission Chari, Lake Tchad, on Amorphophallus sp., 15 Oct. 1902, A. Chevalier, no. 5744 (FH 7807).

Host range and distribution: On Amorphophallus paeoniifolius [campanulatus],?Plesmonium margaritiferum, ?Xanthosoma sp., Araceae, Africa (Gabon, Ghana, Ivory Coast, Nigeria, Sierra Leone, Sudan, Togo, Uganda), Asia (India, Maharashtra, Madhya Pradesh), South America (Venezuela).

Notes: A true Cercospora s. str. distinct from C. apii s. lat. However, records of this species on hosts belonging to Anchomanes, Plesmonium and Xanthosoma spp. are unclear, unproven and possibly based on misidentifications. A record on Anchomes difformis from Guinea (Kranz 1963) belongs undoubtedly to C. anchomanes. Ellis (1976) described pale olivaceous conidia. However, conidia in the type material and other collections examined are hyaline or subhyaline, i.e. with a pale greenish tinge.

Cercospora cryptocorynes Chidd., Mycopathol. Mycol. Appl. 17: 75 (1962); as “cryptocoryneae”.

(Fig. 20)

Fig. 20
figure20

Cercospora cryptocorynes (BPI 435386). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 146), Kamal (2010: 37).

Illustrations: Chiddarwar (1962: 74, plate 1, figs 12–14).

Description: Leaf spots amphigenous, scattered, subcircular, oval to somewhat irregular, 2–25 mm diam, yellowish to ochraceous, pale brownish, finally greyish white, margin darker, brown. Caespituli amphigenous, punctiform, dark brown. Mycelium internal. Stromata substomatal, subglobose to irregular, 20–40 µm diam, dark brown. Conidiophores in small to moderately large fascicles, 4–40, loose to rather dense, arising from stromata, through stomata, erect, straight, subcylindrical or narrowed towards the apex, not or only slightly geniculate-sinuous, unbranched, 40–160 × 4–8 µm, 1–7-septate, pale to medium brown, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal and intercalary, about 10–40 µm long, with conspicuous conidiogenous loci, 2–3 µm diam, thickened and darkened. Conidia solitary, obclavate to obclavate-cylindrical, 25–110 × 3.5–5 µm, 3–14-septate, distance between septa 5–12 µm, hyaline, thin-walled, smooth, apex obtuse to subacute, base usually obconically truncate, occasionally truncate, 2–3 µm wide, hila thickened and darkened.

Holotype: India: Maharashtra: Pune, Mula (Mutha), river bed, on Cryptocoryne retrospiralis, 21 Nov. 1956, P. P. Chiddarwar, no. 26 (K(M) IMI 83187). Isotypes: BPI 435386, HCIO.

Host range and distribution: On Cryptocoryne (cognata, retrospiralis), Araceae, Asia (India, Maharashtra).

Note: A true Cercospora s. str. distinct from C. apii s. lat. by obclavate-cylindrical conidia.

Cercospora monsterae Narayan, Kharwar & R.K. Singh, Kavaka 28–29: 68 (2001).

(Similar to Fig. 1)

Literature: Kamal (2010: 67).

Illustration: Narayan et al. (2001:69, fig. 6).

Description: Leaf spots amphigenous, subcircular to irregular, 5–53 mm diam, later confluent, covering the whole leaf surface, dark olivaceous to brown on the upper side, brown to blackish below. Caespituli amphigenous, effuse. Mycelium internal and external; external hyphae branched, septate, subhyaline to olivaceous. Stromata well-developed, immersed, substomatal, compact, 25–40 µm diam, olivaceous to brown. Conidiophores fasciculate, 8–12, arising from stromata, through stomata, erect, straight to geniculate-flexuous, unbranched, about 45–90 × 2.5–4.5 µm, 2–5eptate, light olivaceous to olivaceous, thin-walled, smooth; conidiogenous cells integrated, terminal, subcylindrical, with a single terminal locus to sympodial, geniculate, conidiogenous loci conspicuous, thickened and darkened. Conidia solitary, acicular, straight to curved, about 55–167 × 2–5.5 µm, 7–19-septate, hyaline, thin-walled, smooth, apex subobtuse to pointed, base truncate, hila thickened and darkened.

Holotype: India: Uttar Pradesh: Gorakhpur, on Monstera deliciosa, Araceae, Mar. 1995, S. Narayan (HCIO 42014).

Host range and distribution: Only known from the type collection.

Note: This species belongs to the C. apii s. lat. complex.

Cercospora pistiae Nag Raj, Govindu & Thirum., Sydowia 24: 299 “1970” (1971).

(Fig. 21)

Fig. 21
figure21

Cercospora pistiae (BPI 439744). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Morris & Crous (1994: 327), Barreto et al. (1999), Crous & Braun (2003: 326), Guo et al. (2005: 39), Kamal (2010: 76).

Illustrations: Govindu et al. (1971: 300, fig. 7), Morris & Crous (1994: 327, fig. 4), Barreto et al. (1999: 82, fig. 1), Guo et al. (2005: 39, fig. 18).

Description: Leaf spots amphigenous, subcircular, elliptical to irregular, 2–15 mm diam, yellowish to brown, often within chlorotic streaks or areas to 40 mm diam, sometimes confluent, severely attacked leaves sometimes becoming necrotic or causing shot-hole symptoms. Caespituli amphigenous, mainly epiphyllous, punctiform, dark. Mycelium immersed; hyphae branched, septate, brownish, about 1.5–6 µm wide. Stromata lacking or only with small aggregations of swollen hyphal cells. Conidiophores solitary or in small to moderately large fascicles, about 2–15, loose to dense, arising from internal hyphae or small aggregations of swollen hyphal cells, through stomata or erumpent, erect, straight, subcylindrical to somewhat geniculate-sinuous, unbranched, 10–295 × 3–8.5 µm, conidiophores mostly 1–10-septate or occasionally with more septa, olivaceous-brown to brown, paler towards the apex, sometimes subhyaline at the very tip, wall thin to somewhat thickened, smooth; conidiogenous cells integrated, terminal or intercalary, about 10–80 µm long, conidiogenous loci conspicuous, thickened and darkened, about 2–4 µm diam. Conidia solitary, broadly acicular to obclavate (-subcylindrical), straight to curved, 30–150 × 3–6.5 µm, 1–14-septate, hyaline, thin-walled, smooth, apex obtuse to subacute, base truncate to somewhat obconically truncate, about 2.5–4 µm wide, hila thickened and darkened.

Holotype: India: Kerala: Ernakulam, on Pistia stratiotes, 11 Dec. 1964, T. R. Nag Raj (MYSP Herb. 803)

Host range and distribution: On Pistia stratiotes, Araceae, Africa (Ghana, South Africa), Asia (China; India, Kerala), Australia, South America (Brazil, Venezuela).

Notes: This species does not belong to the C. apii s. lat. complex. It differs in having subacicular to obclavate (-cylindrical), relatively broad conidia. Material deposited as BPI 439744 (India, “Emakulam”, 21 Jul. 1964, Nag Raj) is very probably topotype material.

Cercospora richardiicola G.F. Atk., J. Elisha Mitchell Sci. Soc. 8: 51 (1892); as “richardiaecola”.

(Similar to Fig. 1)

Literature: Saccardo (1892: 653), Chupp (1954: 60), Boedijn (1961: 413), Crous & Braun (1996: 305), Vieura & Barreto (2004), Nakashima et al. (2007: 266), Groenewald et al. (2013: 160).

Description: Leaf spots amphigenous, circular to somewhat irregular, about 2–8 mm diam, dark olivaceous, brown to reddish brown, sometimes with paler centre. Caespituli amphigenous. Mycelium internal. Stromata lacking or small, substomatal, 10–20 µm diam, brown. Conidiophores solitary or in small, loose fascicles, about 2–15, arising from internal hyphae or small hyphal aggregations, through stomata, erect, straight, subcylindrical to somewhat geniculate, unbranched or only rarely branched, about 20–400 × (2−)3–7 µm, aseptate to usually pluriseptate, pale olivaceous to brown throughout or paler towards the tip, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal and intercalary, conidiophores rarely reduced to conidiogenous cells, 10–40 µm long, conidiogenous loci thickened and darkened, 1.5–3.5 µm diam. Conidia solitary, acicular, 25–300 × 2–4 µm, 2–20-eptate, hyaline, thin-walled, smooth, apex pointed or subobtuse, base truncate, occasionally slightly obconically truncate, 2–3.5 µm wide, hila thickened and darkened.

Holotype: USA: Alabama: Lee County, Auburn, on Zantedeschia aethiopica, 7 Sep. 1891, G. F. Atkinson (CUP-A-2111).

Host range and distribution: On Anthurium sp., Epipremnum aureum, Zantedeschia (aethiopica [Richardia africana], albomaculata [angustiloba, melanoleuca], elliottiana, rehmannii, Zantedeschia sp.), Araceae, Africa (Ethiopia, Sierra Leone, South Africa, Zimbabwe), Asia (China [Hong Kong], Indonesia, Japan, Malaysia, Thailand), USA (Alabama, Florida, Gulf states, Mississippi), Central and South America, West Indies (Colombia, Guatemala, Puerto Rico, Virgin Islands).

Notes: This species belongs to the Cercospora apii s. lat. complex. Groenewald et al. (2013) included cultures based on Japanese material on Zantedeschia sp. (MAFF238210) in molecular studies of Cercospora s. str. species and found that this collection belongs to a clade of an obviously plurivorous Cercospora comprising other isolates from Tagetes erecta, Fuchsia × hybrida, Osteospermum sp. and Gerbera × hybrida. However, the application of the name C. richardiicola is only tentative as this species was described from North America, but American collections were not included in the phylogenetic studies.

Cercospora symplocarpi Peck, in Thüm., Mycoth. Univ., Cent. VII, no. 669 (1877).

(Fig. 22)

Fig. 22
figure22

Cercospora symplocarpi (HAL, Thüm., Mycoth. Univ. 669). A. Conidiophore fascicle. B. Conidia. C. Conidium with basal germ tube. Bar = 10 µm.

Literature: Saccardo (1886: 477), Vassiljevsky & Karakulin (1937: 225), Chupp (1954: 60), Crous & Braun (2003: 394).

Exsiccatae: Barthol., Fungi Columb. 4008. Ellis, N. Amer. Fungi 1295. Ellis & Everh., N. Amer. Fungi 95. Sydow, Fungi Exot. Exs. 97. Thüm., Mycoth. Univ. 669.

Description: Leaf spots amphigenous, subcircular to angular-irregular, 2–8 mm diam, dark olivaceous to reddish brown, centre sometimes paler. Caespituli amphigenous, punctiform, scattered, dark brown. Mycelium internal. Stromata almost lacking to well-developed, substomatal to intraepidermal, 10–60 µm diam, brown, composed of swollen hyphal cells, rounded to somewhat irregular in outline, 3–8 µm diam. Conidiophores in small to moderately large fascicles, loose to usually dense, arising from internal hyphae or stromata, through stomata or erumpent, erect, straight, subcylindrical, subclavate or slightly narrowed towards the apex, slightly geniculate-sinuous, unbranched, 10–60(−70) × 4–8 µm, 0–3-septate, pale to medium brown throughout, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–40 µm long, conidiogenous loci conspicuous, slightly thickened and somewhat darkened-refractive, usually several per cell, non-protuberant, more or less lying planely on the wall of the cell, about 2–3 µm wide. Conidia solitary, obclavate-subcylindrical, occasionally subacicular, 40–150 × 3.5–7 µm, 3–12-septate, distance between septa 5–20 µm, hyaline or subhyaline, i.e. with a pale greenish tinge, occasionally very pale olivaceous, thin-walled, smooth, apex obtuse to subacute, base short obconically truncate, occasionally subtruncate, 2–2.5 µm wide, hila somewhat thickened and darkened-refractive.

Lectotype (designated here, MycoBank MBT178176): USA: New York: West Albany, on Symplocarpus foetidus, Jul. 1877, C. H. Peck (NYS). Isolectotypes: Thüm., Mycoth. Univ. 669 (e.g. BPI 441828, CUP 041363, DAOM, HAL, MICH 15373).

Host range and distribution: On Symplocarpus foetidus, Araceae, North America (Canada; USA, Connecticut, Delaware, Indiana, Maryland, Massachusetts, Michigan, Minnesota, New Jersey, New Mexico, New York, Pennsylvania, Virginia, West Virginia, Wisconsin).

Notes: A true Cercospora s. str. distinct from C. apii s. lat. by it broadly obclavate-cylindrical, sometimes not quite colourless conidia and non-protuberant conidiogenous loci lying planely on the wall of the conidiogenous cells which are less thickened and darkened than in common Cercospora species. This species is morphologically somewhat intermediate between Cercospora s. str. and Passalora, but we prefer to maintain it in Cercospora, at least tentatively. The generic affinity of this species has to be proven by means of molecular sequence analyses.

Cercospora syngoniicola U. Braun & Urtiaga, Mycosphere 4: 593 (2013).

(Fig. 23)

Fig. 23
figure23

Cercospora syngoniicola (K(M) 180153). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Illustration: Braun & Urtiaga (2013b: 594, fig. 1).

Description: Leaf spots amphigenous, subcircular to angular-irregular, 2–18 µm diam, yellowish, ochraceous, brownish, olivaceous-brown, greyish to medium brown, later with paler centre, greyish brown to greyish white, with narrow darker border or diffuse brownish halo, occasionally somewhat zonate. Caespituli amphigenous, mostly epiphyllous, finely punctiform, dark. Mycelium internal. Stromata lacking or small, substomatal or immersed aggregations of swollen hyphal cells, 10–30 µm diam, olivaceous-brown, cells 2.5–11 µm diam. Conidiophores solitary or in small to large fascicles, 2–30, arising from internal hyphae or stromata, erumpent, loose to dense, erect, straight, subcylindrical or attenuated towards the tip to distinctly geniculate-sinuous, unbranched, 10–130 × 3–9 µm, 0–5-septate, pale to medium olivaceous-brown or yellowish brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–40 µm long, conidiogenous loci conspicuous, (1.5−)2–3.5(−4) µm diam, thickened and darkened, occasionally subdenticulate. Conidia solitary, acicular to distinctly obclavate or obclavate-subcylindrical, 25–120 × 3–6 µm, 2–10-septate, distance between septa 5–20 µm, hyaline, thin-walled, smooth, apex obtuse to subacute, base truncate to distinctly obconically truncate, 2–3.5 µm wide, hila thickened and darkened.

Holotype: Venezuela: Lara: Duaca, on living leaves of Syngonium sp., Nov. 1993, R. Urtiaga (K(M) 180153).

Host range and distribution: On Syngonium sp., Araceae, South America (Honduras, Venezuela).

Notes: Two collections from Honduras on “Nephthytis sp.” (rather Syngonium sp.) deposited as “Cercospora verruculosa” have been examined (BPI 442251, 442252). They agree well with C. syngoniicola.

Cercospora typhonii Munjal, Lal & Chona, Indian Phytopathol. 13: 148 (1960).

(Similar to Fig. 1)

Literature: Crous & Braun (2003: 414), Kamal (2010: 96).

Illustration: Munjal et al. (1960: 148, fig. 2).

Description: Leaf spots circular or subcircular, scattered to confluent, 2–9 mm diam, yellowsh green, centre verdigris, border yellowish, broard. Caespituli amphigenous, mostly epiphyllous. Mycelium internal. Stromata small, to about 25 µm diam, globose, dark brown. Conidiophores few, loose to numerous and dense, arising from stromata, erect, straight, subcylindrical to somewhat geniculate, 15–85 × 4–6 µm, width irregular, pluriseptate, pale to olivaceous-brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, usually terminal, conidiogenous loci conspicuous, about 2–3 µm diam, thickened and darkened. Conidia solitary, acicular, straight to somewhat curved, about 25–130 × 3–4 µm, pluriseptate, hyaline, thin-walled, smooth, apex acute, base truncate, about 2.5–3 µm wide, hila thickened and darkened.

Holotype: India: West Bengal: Calcutta, on Typhonium trilobatum, 2 Nov. 1939, A. K. Ghosh (HCIO 26616).

Host range and distribution: On Typhonium (flagelliformum, roxburghii [divarigatum], trilobatum), Araceae, Asia (China, India, Malaysia, Taiwan).

Notes: A true Cercospora s. str. close to C. apii s. lat., but distinct by having relatively short conidiophores (type material was not available, but Indian material from K(M) IMI examined).

Cercospora verruculosa F. Stevens & Solheim, Mycologia 23: 397 (1931).

(Fig. 24)

Fig. 24
figure24

Cercospora verruculosa (BPI 428776). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Synonyms: Helminthosporium caladii F. Stevens, Trans. Illinois Acad. Sci. 10: 209 (1917) [lectotype (designated here, MycoBank MBT178177): Puerto Rico: Mayaguez, on Caladium bicolor, 27 Oct. 1913, F. L. Stevens 3860 (BPI 428776). Isolectotypes: BPI 428777, 845005; K, PC], non Cercospora caladii Cooke, 1880.

Cercosporidium caladii (F. Stevens) Deighton, Mycol. Pap. 112: 32 (1967).

Passalora caladii (F. Stevens) Poonam Srivast., J. Living World 1: 113 (1994), comb. inval. (ICN, Art. 41.4).

Passalora caladii (F. Stevens) U. Braun & Sivap., Fungal Diversity 3: 10 (1999).

Literature: Chupp (1954: 60), Boedijn (1961: 413), Deighton (1967: 32), Saccardo (1972: 1386), Crous & Braun (2003: 93), Piepenbring (2006), Kamal (2010: 111).

Illustration: Deighton (1967: 33, fig. 17).

Description: Leaf spots amphigenous, circular to angular-irregular, 2–40 mm diam, straw-coloured, brown, dull brown, later greyish brown to grey with brown to dark brown border, sometimes zonate. Caespituli mostly hypophyllous, scattered to aggregated, punctiform, medium to dark brown. Mycelium internal; hyphae branched, septate, 2.5–4 µm wide, subhyaline or pale, thin-walled, smooth. Stromata substomatal, 10–50 µm diam, subglobose, brown. Conidiophores fasciculate, to 25, loose to mostly dense, arising from stromata, through stomata, erect, straight, subcylindrical to distinctly geniculate-sinuous, rhachis-like, unbranched, 30–135(−250) × 3.5–8 µm, sparingly septate, yellowish to pale olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal, occasionally intercalary, about 10–50 µm long, conidiogenous loci conspicuous, thickened and darkened, 2–3 µm diam. Conidia solitary, broadly ellipsoid-ovoid, obovoid, fusiform, subcylindrical, 15–55(−70) × 5–9.5 µm, 2–7-septate, hyaline or subhyaline, thin-walled, smooth, apex broadly rounded, base rounded to short obconically truncate, 2–3.5 µm wide, hila somewhat thickened and darkened.

Holotype: Trinidad: St. Augustine, on Caladium sp., 13 Aug. 1932, F. L. Stevens 829 (ILL 15793).

Host range and distribution: On Caladium (bicolor, Caladium sp.), Colocasia (esculenta [antiquorum]), Xanthosoma (sagittifolia, Xanthosoma sp.), Araceae, Asia (Brunei, India), Central and South America, West Indies (Panama, Puerto Rico, Trinidad and Tobago, Venezuela, Virgin Islands).

Notes: This species has hitherto been considered a member of Passalora. However, results of molecular sequence analyses have shown that passalora-like species with hyaline conidia rather pertain to Cercospora s. str. (Groenewald et al. 2013, Braun et al. 2013). Therefore, this species is now better treated under the latter genus. Since the name Cercospora caladii is preoccupied, a reallocation of Helminthosporium caladii to Cercospora is not possible and the name C. verruculosa (heterotypic synonym of H. caladii) has to be applied to this species.

Doubtful, excluded and insufficiently known species

Cercospora ari (Fautrey) Vasyag., in Shvartsman et al., Fl. Spor. Rast. Kazakhstana 8(2): 252 (1975).

Basionym: Ramularia ari Fautrey, Rev. Mycol. 17: 71 (1895).

Literature: Saccardo (1895: 605), Vassiljevsky & Karakulin (1937: 54), Braun (1995: 250), Braun & Mel’nik (1997: 40), Crous & Braun (2003: 65).

Illustration: Braun (1995: 247, fig. 230 b).

Exsiccatae: Roum., Fungi Sel. Exs. 6181.

Description: Leaf spots amphigenous, subcircular to irregular, 2–4 mm diam, membranous, brownish to grey. Caespituli epiphyllous. Mycelium internal. Stromata lacking or small. Conidiophores in small fascicles, subcylindrical, straight, 20–65 × 3–6 µm, 0–1-septate, brown; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, conidiogenous loci conspicuous, thickened and darkened. Conidia solitary, cylindrical, 10–50 × 3–4 µm, 1–4-septate, hyaline, apex obtuse to subacute, base truncate, about 2–3 µm wide, slightly thickened and darkened.

Lectotype (designated by Braun 1995): France: Côte-d’Or, on Arum maculatum, May 1892, Fautrey, Roum., Fungi Sel. Exs. 6181 (PC). Isolectotypes: Roum., Fungi Sel. Exs. 6181.

Host range and distribution: On Arum (maculatum, korolkowii), Araceae, Central Asia (Kazakhstan), Europe (France, Italy).

Notes: A true Cercospora s. str., but type material too poor for a final conclusion about the status of this species (see Braun 1995).

Cercospora aricola Sacc., Ann. Mycol. 11: 548 (1913). Synonym: Cercosporina aricola (Sacc.) Sacc., Syll. Fung. 25: 896 (1931).

Literature: Chupp (1954: 56), Crous & Braun (2003: 65), Guo et al. (2005: 278).

Description: Leaf spots conspicuous, on both sides, but more evident below, 2–4 mm diam, white, margin ochraceous to brown. Caespituli hypophyllous. Mycelium internal. Stromata present. Conidiophores 50–90 × 5–6 µm, more or less straight, mostly 1-septate, brown, slightly but closely denticulate above. Conidia solitary, obclavate, often curved, 40–70 × 4.5–5 µm, 8–12-septate, without constrictions, subhyaline, apex acute, base obconically truncate.

Holotype: Mexico: Vera Cruz, on unknown member of Araceae, S. Bonansea [Fungi Mexicani 14] (PAD).

Host range and distribution: On unknown member of Araceae, Typhonium giganteum, Araceae, Asia (China), North America (Mexico).

Notes: This species is insufficiently known. The present description is based on the orginal description. Type material was not available for re-examination. It is also unclear if Chinese material on Typhonium giganteum, assigned to C. aricola, is actually conspecific. Guo et al. (2005) described conidiophores of 44–92 × 4–6 µm and conidia of 35–80 × 3.5–4.5 µm.

Cercospora callae f. aethiopica Gonz. Frag., Bol. Soc. Esp. Hist. Nat. 233: 326 (1923).

Literature: Saccardo (1972: 1370).

Holotype: Spain: Madrid, Botanical Garden, on Zantedeschia aethiopica, Araceae, 4 Jan. 1922, F. Riofrio 6162 (MA).

Notes: Not belonging to C. callae and Cercospora at all, but rather a sporidesmium-like fungus.

Passalora

Key to Passalora species on Araceae
  1. 1

    Stromata present, 10–60 µm diam; conidiophores fasciculate; conidia solitary; on Colocasia ............. P. colocasiae

    Stromata absent; conidiophores solitary; conidia catenate; on Caladium ............................. P. caladiicola

Passalora species on Araceae

Passalora caladiicola (Chupp) U. Braun, comb. nov. Mycobank MB809008.

Basionym: Cercospora caladiicola Chupp Monograph of Cercospora: 57 (1954); as “caladicola”.

Synonym: Cercospora caladii Henn., Hedwigia 48: 17 (1908), nom. illeg. (ICN, Art. 53.1).

Literature: Saccardo (1931: 868), Chupp (1954: 57), Crous & Braun (2003: 93–94).

Description: Leaf spots minute, formed as dark brown to black specks, 0.5–1.5 mm diam. Caespituli hypophyllous. Stromata lacking. Conidiophores solitary, mostly formed as lateral branches of procumbent threads, or in groups of 2–3, irregular in outline, but not geniculate, 10–25 × 3–5 µm, or even as large as 85 × 7 µm, pale olivaceous-brown, when longer paler and narrower towards the tip; small scars at bluntly rounded tips. Conidia catenate, cylindrical, 10–50 × 3–6 µm, 1–5-septate, pale olivaceous, ends rounded.

Holotype: Brazil: São Paulo, on Caladium sp., Araceae, 14 Mar. 1902, A. Puttemans 476 (B, but currently lost).

Host range and distribution: Only known from the type collection.

Notes: This species is insufficiently known. Its type material, previously preserved at B, was sent as loan to Brazil in 1997 and has been lost. The present description is based on the original publication and Chupp (1954). However, the combination of superficial hyphae with solitary conidiophores, visible conidiogenous loci (scars) and cylindrical catenate conidia strongly suggests a mycovellosiella-like species that has to be assigned to Passalora.

Passalora colocasiae (Höhn.) U. Braun, in Braun et al., New Zealand J. Bot. 37: 308 (1999).

(Fig. 25)

Fig. 25
figure25

Passalora colocasiae (CUP 39261). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora caladii var. colocasiae Höhn., Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Cl., Abt. 1, 116: 150 (1907).

Synonyms: Cercospora colocasiae (Höhn.) Chupp, Monograph of Cercospora: 58 (1954).

Mycosphaerella alocasiae Syd. & P. Syd., Philipp. J. Sci. 8: 195 (1913) [syntypes: Philippines: Luzon, Manila, on Alocasia macrorhizon, Dec. 1911, Graff, Syd., Fungi Exot. Exs. 30 (e.g. BPI 607294, 607296; CUP; L)].

Sphaerella alocasiae (Syd. & P. Syd.) Trotter, Syll. Fung. 24: 850 (1928).

Literature: Saccardo (1913: 1431), Chupp (1954: 58), Vasudeva (1963: 87), Sivanesan (1984: 185), Crous & Braun (2003: 132), Aptroot (2006: 31–32), Kamal (2010: 113).

Illustrations: Sivanesan (1984: 186, fig. 94), Aptroot (2006: 10, fig.36).

Exsiccatae: Rehm, Ascomyc. 2060. Syd., Fungi Exot. Exs. 30.

Description: Leaf spots amphigenous, subcircular to somewhat angular-irregular, 2–8 mm diam, centre pale ochraceous to brownish, margin narrow to moderately wide, darker, dingy greyish brown to dark brown. Caespituli amphigenous, punctiform, scattered to dense, dark brown. Mycelium internal; hyphae sparingly branched, septate, pigmented. Stromata substomatal, small to moderately large, subglobose, 10–60 µm diam, pigmented, composed of swollen hyphal cells, 2–8 µm diam. Conidiophores in small to moderately large fascicles, arising from stromata, through stomata, loose to dense, erect, straight to curved, subcylindrical to attenuated towards the tip, usually not geniculate-sinuous, occasionally slightly so, unbranched, 5–35 × 2–5 µm, sometimes swollen at the very base, to 8 µm wide, 0–2(−3)-septate, pale olivaceous to light brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–25 µm long, conidiogenous loci conspicuous, slightly thickened and darkened, 1–2 µm diam. Conidia solitary, obclavate (-cylindrical), fusiform, (15−)25–60(−100) × 2–5 µm, (0−)1–5(−6)-septate, subhyaline to pale olivaceous or light brown, thin-walled, almost smooth to verruculose, apex obtuse to subacute, base obconically truncate, 1–2.5 µm wide, hila slightly thickened and darkened.

Sexual morph: Ascomata pseudothecial, in the centre of leaf spots, numerous, dense, immersed, globose, blackish, ostiolum 18–25 µm diam; asci fasciculate, saccate to cylindrical, 35–52 × 10–16 µm, aparaphysate, 8-spored, distichous to tristichous; ascospores narrowly pyriform, 16–22 × 3–5.5 µm, with a single median septum, hyaline, ends obtuse.

Holotype: Samoa: on living leaves of Colocasia esculenta, 1905, Rechinger 2297 (FH). Isotype: CUP 39261.

Host range and distribution: On Alocasia (macrorrhizos, Alocasia sp.), Colocasia (esculenta [antiquorum], Colocasia sp.), Araceae, Asia (Brunei, India, Indonesia, Malaysia, Nepal, Papua New Guinea, Philippines), Oceania (Fiji, Guam, Niue, Samoa, Solomon Islands, Tonga).

Pseudocercospora

Key to Pseudocercospora species on Araceae
  1. 1

    Conidia short clavate-turbinate, 20–35 × 10–14.5 µm, (1−)3(−5)-septate; on Colocasia ................................ P. colocasiae

    Conidia much narrower, < 10 µm; on other hosts ................................................... 2

  2. 2(1)

    Mycelium internal and external, superficial; stromata lacking; conidiophores fasciculate and solitary, arising from superficial hyphae, to 300 µm long; conidia cylindrical, 40–80 × 4–4.5 µm; on Amorphophallus ................................................... P. protensa

    Mycelium internal; superficial hyphae lacking; stromata developed; conidiophores much shorter, 5–80 µm; conidia obclavate-cylindrical, filiform or subacicular, if cylindrical then conidia wider, 6–9 µm; on other hosts ................................................... 3

  3. 3(2)

    Conidia mostly cylindrical, 20–50 × 6–9 µm, 1–3(−4)-septate; on Alocasia ......................................... P. alocasiicola

    Conidia narrower, 2–5.5 µm, (0−)1–12-septate; on other hosts ................................................... 4

  4. 4(3)

    Conidia narrowly obclavate to cylindrical-filiform, 40–110 × 2–4 µm, 4–12-septate; on Aglaonema .................. P. bruneiensis

    Conidia shorter and wider, to 75 × 5.5 µm, 0–8-septate; on other hosts ....................................... 5

  5. 5(4)

    Conidiophores 5–25 × 3–6 µm; conidia obclavate-cylindrical, hila 2–2.5 µm wide; on Colocasia ....................... P. aracearum

    Conidiophores narrower, 5–25 × 2–4 µm; conidia obclavate-cylindrical to subacicular, hila 1–2 µm wide; on Montrichardia .................................................................... P. montrichardiae

Pseudocercospora species on Araceae

Pseudocercospora alocasiicola U. Braun & McKenzie, New Zealand J. Bot. 37: 306 (1999).

(Fig. 26)

Fig. 26
figure26

Pseudocercospora alocasiicola (PDD 49041). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Illustration: Braun et al. (1999: 307, fig. 7).

Description: Leaf spots amphigenous, subcircular to irregular, 1–15 mm diam, at first brownish, dark greyish brown to blackish or violet-black, margin indefinite, later with pale greyish brown to dingy greyish centre and dark margin, dark brown to blackish or violet-black. Caespituli hypophyllous, punctiform, scattered to subgregarious, dark brown to blackish. Mycelium internal; hyphae septate, branched, pigmented. Stromata intraepidermal, occasionally substomatal, small to expanded, planate, composed of swollen hyphal cells, more or less angular, thick-walled, 3–10 µm diam. Conidiophores in small to moderately large fascicles, loose, occasionally solitary, arising from stromata, erumpent, erect, straight and subcylindrical to geniculate-sinuous, unbranched or occasionally branched, 15–80 × 3–8 µm, continuous to pluriseptate throughout, occasionally with constrictions at septa, olivaceous to medium dark brown, tips often paler, wall thin to somewhat thickened, smooth; conidiogenous cells integrated, terminal, occasionally intercalary, conidiogenous loci inconspicuous to subconspicuous by being frequently subdenticulate, but always unthickened and not darkened. Conidia solitary, usually cylindrical, some conidia subclavate, short conidia sometimes obclavate or broadly fusiform, 20–50 × 6–9 µm, 1–3(−4)-septate, not or only slightly constricted at septa, pale olivaceous to dingy olivaceous-brown, thin-walled, almost smooth to faintly rough-walled, apex obtuse, rounded, base rounded to somewhat obconically truncate or with denticle-like protuberance, hilum unthickened, not darkened.

Holotype: Vanuatu: Efate, Teouma, on living leaves of Alocasia macrorrhizos, Araceae, 21 Nov. 1983, E. H. C. McKenzie (PDD 49041).

Host range and distribution: Only known from the type collection.

Pseudocercospora aracearum U. Braun & McKenzie, in Braun et al. New Zealand J. Bot. 37: 308 (1999).

(Fig. 27)

Fig. 27
figure27

Pseudocercospora aracearum (PDD 60364). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Illustration: Braun et al. (1999: 307, fig. 9).

Description: Leaf spots subcircular, 4–6 mm diam, brownish, margin indefinite, but often with a greenish halo. Caespituli amphigenous, punctiform, dark brown. Mycelium internal; hyphae septate, branched, pigmented. Stromata well-developed, intraepidermal, erumpent, subglobose, 20–40 µm diam, brown, composed of swollen hyphal cells, 2.5–6 µm diam. Conidiophores usually in small fascicles, loose to dense, arising from stromata, erumpent, erect, straight, occasionally curved, subcylindrical or attenuated towards the tip, slightly geniculate-sinuous, 5–25 × 3–6 µm, 0–2-septate, subhyaline to pale olivaceous, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–20 µm long, conidiogenous loci inconspicuous, unthickened, not darkened. Conidia solitary, obclavate-subcylindrical, fusiform, 20–75 × (3−)4– 5(−5.5) µm, 1–8-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex usually obtuse, occasionally subacute, base obconically truncate, occasionally truncate in cylindrical conidia, mostly 2–2.5 µm wide, hila neither thickened nor darkened.

Holotype: Vanuatu: Santos, Sarete, on Colocasia esculenta, Araceae, 14 Feb. 1985, E. H. C. McKenzie (PDD 60364).

Host range and distribution: Only known from the type collection.

Pseudocercospora bruneiensis U. Braun & Sivap., Fungal Diversity 3: 11 (1999).

(Fig. 28)

Fig. 28
figure28

Pseudocercospora bruneiensis (HAL 1596 F). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Illustration: Braun & Sivapalan (1999: 12, fig. 7).

Description: Leaf spots amphigenous, subcircular to somewhat irregular, 3–15 mm diam, dingy greyish green, grey or greyish white, margin narrow, darker, dingy olivaceous to dark greyish brown. Caespituli amphigenous, mostly hypophyllous, punctiform to subeffuse, dingy greyish olivaceous to brownish. Mycelium internal; hyphae sparingly branched, pigmented, forming small to well-developed stromata, substomatal, 10–50 µm diam, brown, composed of swollen hyphal cells, 2–7 µm diam. Conidiophores in dense fascicles, usually numerous, arising from stromata, emerging through stomata, erect, straight, cylindrical, conical or flexuous, geniculate-sinuous, simple or occasionally branched, 5–40 × 2–5 µm, 0–1-septate, pale olivaceous to medium olivaceous-brown, tips often somewhat paler, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–30 µm long, conidiogenous loci inconspicuous, rounded to truncate, unthickened, not darkened. Conidia solitary, narrowly obclavate or cylindrical-filiform, 40–110 × 2–4 µm, pluriseptate, mostly with 4–12 septa, subhyaline to pale olivaceous, thin-walled, smooth, apex usually subacute, base obconically truncate or occasionally truncate, 1.5–3 µm wide, hila neither thickened nor darkened, conidia rarely with a subbasal lateral branchlet which possibly represents a germ tube.

Holotype: Brunei: Rimba, on Aglaonema sp., Araceae, 28 Aug. 1996, A. Sivapalan 7747 (HAL 1596 F).

Host range and distribution: Only known from the type collection.

Pseudocercospora colocasiae Deighton, Mycol. Pap. 140: 17 (1976).

(Fig. 29)

Fig. 29
figure29

Pseudocercospora colocasiae (K(M) IMI 77190). A. Conidiophore fascicle. B. Conidiophore tip with conidium. C. Conidia. Bar = 10 µm.

Literature: Matsushima (1985), Goh & Hsieh (1990: 30), Guo & Hsieh (1995: 22), Guo et al. (1998: 35–36).

Illustrations: Deighton (1976: 18–19, figs 5–6), Matsushima (1985: fig. 298), Goh & Hsieh (1990: 31, fig. 17), Guo & Hsieh (1995: 25, fig. 23), Guo et al. (1998: 35, fig. 23).

Description: Leaf spots indistinct, forming subcircular pale discolorations, to 10 mm diam, centre sometimes turning greyish with age, margin indefinite. Caespituli amphigenous, mainly hypophyllous, effuse, thin, medium to dark olivaceous. Mycelium internal; hyphae branched, septate, subhyaline, 1.5–4 µm wide. Stromata none. Conidiophores in loose fascicles, to 9, arising from internal hyphae, through stomata, erect, cylindrical-filiform, flexuous, somewhat sinuous, barely geniculate, unbranched, rarely with lateral branchlets, swollen at the very base, 50–425 × 4–7 µm, pluriseptate throughout, medium dark olivaceous, somewhat paler towards the tip, thin-walled, smooth or somewhat rough-walled above; conidiogenous cells integrated, terminal, proliferations sympodial and percurrent, annellations not very conspicuous, conidiogenous loci inconspicuous or visible as truncate apex, 2.5–4 µm wide, sometimes subdenticulate, but always unthickened and not darkened. Conidia solitary, short clavate-turbinate, mostly straight, rarely somewhat curved or asymmetric, 20–35 × 10–14.5 µm, (1−)3(−5)-septate, pale olivaceous, thin-walled, almost smooth to faintly rough-walled, apex broadly rounded, base obconically truncate, 2–3.5 µm wide, hila neither thickened nor darkened.

Holotype: Solomon Islands: Hauhui, Malaite, on Colocasia esculenta, 1 June 1959, A. Johnston (K(M) IMI 77190).

Host range and distribution: On Colocasia esculenta, Araceae, Asia (Bangladesh, Indonesia [Prov. Papua], Myanmar, Taiwan), Oceania (Samoa, Solomon Islands).

Pseudocercospora montrichardiae (Henn.) U. Braun & F.O. Freire, Cryptog. Mycol. 23: 313 (2002).

(Fig. 30)

Fig. 30
figure30

Pseudocercospora montrichardiae (S-F37470). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora montrichardiae Henn., Hedwigia 48: 115 (1909).

Literature: Saccardo (1931: 869), Chupp (1954: 59), Crous & Braun (2003: 281).

Illustration: Braun & Freire (2002: 314, fig. 20).

Description: Leaf spots amphigenous, subcircular to angular-irregular, 1–15 mm diam, yellowish to ochraceous, finally greyish brown to greyish white, margin indefinite or narrow and darker, often vein-limited. Caespituli amphigenous, punctiform, dark brown. Mycelium internal. Stromata variable, almost absent to well-developed, substomatal, 10–70 µm diam, olivaceous to yellowish brown or dark brown. Conidiophores in small to large fascicles, often almost sporodochial, loose to dense, arising from stromata, emerging through stomata, erect, straight, subcylindrical-conical to strongly geniculate-sinuous, unbranched or occasionally branched, 5–25 × 2–4 µm, 0–2-septate, subhyaline to pale olivaceous, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–20 µm long, conidiogenous loci inconspicuous to subdenticulate, but always unthickened, not darkened. Conidia solitary, narrowly obclavate-subcylindrical, subacicular, (8−)15–70 × 2.5–5.5 µm, (0−)1–7-septate, subhyaline to olivaceous, thin-walled, smooth, apex obtuse to subacute, base truncate to obconically truncate, 1–2 µm wide, hila unthickened, not darkened.

Lectotype (designated by Braun & Freire 2002): Brazil: Para: Ilha das Oncas, on Montrichardia arborescens, Oct. 1903, Huber 96 (S-F37470).

Host range and distribution: On Montrichardia (arborescens, linifera), Araceae, South America (Brazil).

Pseudocercospora protensa (Syd.) Deighton, Mycol. Pap. 140: 150 (1976).

(Fig. 31)

Fig. 31
figure31

Pseudocercospora protensa (B 70014865). A. Stroma. B. Conidiophore fascicle and superficial hypha with conidiophore emerging through a stoma. C. Conidiophore tips. D. Conidia. Bar = 10 µm.

Basionym: Cercospora protensa Syd., Ann. Mycol. 28: 446 (1930).

Literature: Chupp (1954: 59), Boedijn (1961: 413), Crous & Braun (2003: 336–337).

Description: Leaf spots formed as indistinct yellowish to brownish discolorations on the upper leaf surface, 3–10 mm diam, later expanded, larger, sometimes affecting almost entire leaves. Caespituli hypophyllous, punctiform to confluent, effuse, later visible as large patches or layers, loose to dense, dark olivaceous to brown. Mycelium internal, occasionally with a few procumbent threats growing like superficial hyphae, sometimes even with lateral conidiophores. Stromata lacking or only with small stromatic aggregations of swollen hyphal cells, substomatal, 10–25 µm diam, brown, individual cells subglobose, 2.5–6 µm diam, with somewhat thickened walls. Conidiophores in small to moderately large, loose fascicles, arising from internal hyphae or stromatic hyphal aggregations, emerging through stomata, occasionally solitary, arising from decumbent threads, erect, straight to flexuous, variously curved to somewhat geniculate-sinuous, simple or branched, branchings not rare, long, 60–300 × 2.5–6 µm, width uniform or somewhat attenuated towards the tip, plainly pluriseptate throughout, cells (5−)10–20(−30) µm long, pale to medium olivaceous-brown or yellowish brown, paler towards the apex, tips sometimes subhyaline, thin-walled, smooth; conidiogenous cells integrated, terminal, sometimes intercalary, 10–25 µm long, sympodial, conidiogenous loci inconspicuous or visible as truncate tips or lateral shoulders formed by sympodial proliferation, but always unthickened and not darkened. Conidia solitary, cylindrical to obclavate-cylindrical, straight to curved, often strongly curved, sigmoid or occasionally even uncinate to subspirally twisted, outline and width often somewhat irregular, 20–80 × 3–6.5 µm, 2–9-septate, pale yellowish, greenish, olivaceous to olivaceous-brown, thin-walled, smooth, apex subacute to obtuse, rounded, base subtruncate to usually short obconically truncate or gradually attenuated towards the base, 1.5–2.5 µm wide, hila unthickened, not darkened.

Lectotype (designated here, MycoBank MBT178135): Philippines: Province Laguna: Los Baños, on Amorphophallus paeoniifolius, 1 Oct. 1912, C. F. Baker 322 (S-F37711). Isolectotype: B 700014865.

Host range and distribution: On Amorphophallus (paeoniifolius [campanulatus], variabilis, Amorphophallus sp.), Araceae, Asia (Indonesia, Malaysia, Papua New Guinea, Philippines).

Notes: An additional collection deposited as C. protensa on Amorphophallus sp. has been examined and confirmed (North Borneo, 1 Sep. 1959, A. Johnston NB 28, K(M) IMI 78759).

Zasmidium

Key to Zasmidium species on Araceae
  1. 1

    Superficial hyphae not developed in vivo; conidiophores only fasciculate; conidia formed singly, obclavate, 30–60 × 3–4 µm; on Cercestis afzelii, Africa ....................................... Z. cercestidis-afzelii

    Superficial hyphae developed in vivo, with or without solitary conidiophores ....................................... 2

  2. 2(1)

    Conidia catenate, small, 4–20 × 2–3 µm, 0–1-septate; on Cercestis congoensis, Africa ................................ Z. deightoniana

    Conidia consistently solitary or solitary as well as catenate, much longer and pluriseptate ............................. 3

  3. 3(2)

    Superficial hyphae formed in vivo, but conidiophores only fasciculate, solitary conidiophores arising from superficial hyphae lacking; conidia to 350 µm long; on Homalomena spp., Asia ................................ Z. extremorum

    Solitary conidiophores arising from superficial hyphae formed in vivo; conidia much shorter, only to 140 µm; on other hosts ....................................... 4

  4. 4(3)

    Stromata lacking; on Anthurium or Cercestis ....................................... 5

    Stromata developed; on Alocasia or Colocasia ....................................... 6

  5. 5(4)

    Conidiophores solitary; conidia solitary, obclavate-cylindrical to filiform; on Anthurium sp., Asia ....................... Z. anthuriicola

    Conidiophores solitary as well as fasciculate; conidia catenate, cylindrical or subcylindrical; on Cercestis congoensis, Africa ....................................... Z. cercestidis

  6. 6(4)

    Hyphae regularly nodulose; conidia solitary as well as catenate; on Colocasia esculenta ............................ Z. colocasiae

    Hyphae not or only rarely with swellings; conidia mosty solitary; on Alocasia spp. ....................................... Z. alocasiae

Zasmidium species on Araceae

Zasmidium alocasiae (Sarbajna & Chattopadh.) Kamal, Cercosporoid Fungi of India: 237 (2010).

(Fig. 32)

Fig. 32
figure32

Zasmidium alocasiae (K(M) IMI 284569). A. Superficial hyphae. B. Superficial hyphae with conidiophores. C. Conidiophore fascicles. D. Conidiophores. E. Conidia. Bar = 10 µm.

Basionym: Stenella alocasiae Sarbajna & Chattopadh., J. Mycopathol. Res. 29: 33 (1991).

Literature: Braun & Sivapalan (1999: 23), Guo (2010).

Illustration: Sarbajna & Chattopadhyay (1991: 34, fig. 2).

Description: Leaf spots amphigenous, subcircular to somewhat irregular, scattered to confluent, 2–18 mm diam, at first pale greenish, later pale greyish brown to greyish white, sometimes zonate, margin mostly darker, narrow to moderately broad, dark brown, reddish brown or blackish brown, sometimes with yellowish halo. Caespituli amphigenous, scattered to confluent, punctiform, dark brown, blackish brown, later greyish brown by abundant sporulation. Mycelium internal and external; superficial hyphae emerging through stomata, sparingly branched, septate, occasionally with constrictions at septa and swellings, 1–3 µm wide, subhyaline to pale olivaceous, thin-walled, verruculose. Stromata substomatal, subglobose to somewhat irregular, 10–65 µm diam, yellowish brown, medium brown to dark brown, cells 2–8 µm diam. Conidiophores in small to moderately large fascicles, 3–30, loose to moderately dense, arising from stromata, through stomata, and solitary, arising from superficial hyphae, lateral, rarely terminal, erect, straight, subcylindrical or somewhat attenuated towards the tip, somewhat sinuous and geniculate near the tip, unbranched, very short conidiophores arising from superficial hyphae sometimes peg-like, aseptate, 5–200 × 2–5 µm, 0–15-septate, pale olivaceous to yellowish or olivaceous-brown to brown, wall thin to somewhat thickened, smooth or almost so; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, conidiogenous loci conspicuous, thickened and darkened, 1–1.5 µm diam. Conidia mostly solitary, occasionally catenate, cylindrical, fiiliform, narrowly obclavate, 10–140 × 2–4 µm, 1- to pluriseptate, subhyaline, pale olivaceous, olivaceous-brown or pale brownish, thin-walled, verruculose, apex subacute or obtuse, base subtruncate to short obconically truncate, 1–2.5 µm wide, hila somewhat thickened and darkened.

Holotype: India: West Bengal: Baruipur, on Alocasia macrorrhizos, 28 Dec. 1983, K. K. Sabajna PCC 3469 (K(M) IMI 284569).

Host range and distribution: On Alocasia (macrorrhizos [indica], Alocasia sp.), Araceae, Asia (Brunei, ?China, India, West Bengal).

Notes: Braun & Sivapalan (1999) discussed a collection on Colocasia sp. from Brunei, which was morphologically somewhat intermediate between Stenella alocasiae and S. colocasiae. The conidia are much longer in comparision to S. alocasiae (20–140 × 2–4 µm), pluriseptate and consistently formed singly. The identity of material on Alocasia sp. recorded and described from Hainan (Guo 2001c) is unclear. The conidiophores have been described to be 4–6.5 µm wide and the conidia are said to be catenate, cylindrical, to 195 µm long.

Zasmidium anthuriicola (U. Braun & C.F. Hill) Crous & U. Braun, Persoonia 23: 104 (2009).

(Fig. 33)

Fig. 33
figure33

Zasmidium anthuriicola (HAL 1870 F). A. Superficial hyphae. B. Superficial hyphae with conidiophores. C. Conidiophores. D. Conidia. Bar = 10 µm.

Basionym: Stenella anthuriicola U. Braun & C.F. Hill, in Braun et al. Fungal Diversity 22: 33 (2006).

Illustration: Braun et al. (2006: 29, fig. 9).

Description: Lesions formed as sharply delineated brown discolorations on leaves, to 5 mm wide, with abundant superficial mycelium. Colonies in vitro (prune extract agar) olivaceous-grey (top), and olivaceous-black (bottom), with regular to slightly irregular, smooth margin. Mycelium effuse; hyphae sparingly branched, mostly straight, occasionally anastomosing, 1–3(−5) µm wide, septate, subhyaline, pale to medium brown or olivaceous-brown, thin-walled, verruculose. Stromata lacking. Conidiophores solitary, arising from plagiotropous hyphae, lateral, occasionally terminal, erect, straight, neither geniculate nor sinuous, unbranched, subcylindrical or slighty attenuated towards the apex, occasionally swollen at the base, 10–60 × 2–4 µm, 0–3(−4)-septate, pale olivaceous to olivaceous-brown, often paler towards the tip, thin-walled, smooth to verruculose, especially in the lower half; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–30 µm long, conidiogenous loci conspicuous, 0.75–1.5 µm diam, thickened and darkened. Conidia solitary, narrowly obclavate-cylindrical, filiform, 10–90 × 2–3 µm, 0–6-septate, subhyaline to pale olivaceous, thin-walled, verruculose, apex obtuse to subacute, base short obconically truncate, 1 µm wide, hila somewhat thickened and darkened.

Holotype: Thailand: intercercepted at Auckland International Airport, New Zealand (imported from Thailand), on Anthurium sp., Araceae, 3 Aug. 2005, C. F. Hill (HAL 1870 F). Ex-type culture: CBS 118742.

Host range and distribution: Only known from the type collection.

Note: Crous et al. (2009c) analysed this species by means of molecular methods and confirmed its phylogenetic position.

Zasmidium cercestidis (J.M. Yen & Gilles) U. Braun, Schlechtendalia 20: 100 (2010).

(Fig. 34)

Fig. 34
figure34

Zasmidium cercestidis (K(M) IMI 212936). A. Superficial hypha with solitary conidiophores. B. Conidiophore fascicle. C. Conidiophore tip. D. Conidia. Bar = 10 µm.

Basionym: Cercospora cercestidis J.M. Yen & Gilles, in Yen, Bull. Trimestriel Soc. Mycol. France 91: 92 (1975).

Synonym: Stenella cercestidis (J.M. Yen & Gilles) Deighton, Mycol. Pap. 144: 53 (1979).

Literature: Crous & Braun (2003: 116).

Illustration: Yen (1975: 91, fig. 2).

Description: Leaf spots amphigenous, scattered, oval to irregular, 5–20 mm diam, brownish, surrounded by a yellowish halo, margin indistinct. Caespituli amphigenous. Mycelium internal and external; superficial hyphae emerging through stomata, branched, septate, 2–2.5 µm wide, pale brown to brown, wall thin and rough. Stromata lacking or only formed as small substomatal aggregations of swollen hyphal cells, brown. Conidiophores in small to moderately large, loose fascicles, emerging through stomata or solitary, arising from superficial hyphae, lateral, erect, straight, subcylindrical, barely geniculate, unbranched, 20–105 × 3–4 µm, 0–8-septate, brown, paler towards the tip, thin-walled, smooth or almost so; conidiogenous cells integrated, terminal, 10–30 µm long, sympodial, conidiogenous loci conspicuous, 1–2(−5) per cell, somewhat thickened and darkened. Conidia catenate, in simple or branched chains, cylindrical or subcylindrical, 8–75 × 2–3 µm, 0–7-septate, pale olivaceous-brown, wall thin, rough, ends rounded, truncate or short obconically truncate, 1–2 µm wide, hila thickened and darkened.

Holotype: Ivory Coast: Abidjan, Forêt de Yapo, on Cercestis afzelii, 10 Feb. 1974, G. Gilles PCI 93 (K(M) IMI 212936).

Host range and distribution: On Cercestis afzelii, Araceae, Africa (Ivory Coast).

Notes: The status of the type material is not quite clear, but the collection deposited at K is the only material that could be traced. Therefore, it is assumed that the material now in K represents the holotype.

Zasmidium cercestidis-afzelii (J.M. Yen & Gilles) U. Braun, comb. nov.

MycoBank MB809009

(Fig. 35)

Fig. 35
figure35

Zasmidium cercestidis-afzelii (K(M) IMI 212936). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Basionym: Cercospora cercestidis-afzelii J.M. Yen & Gilles, in Yen, Bull. Trimestriel Soc. Mycol. France 91: 93 (1975).

Literature: Crous & Braun (2003: 116).

Illustration: Yen (1975: 93, fig. 3).

Description: Leaf spots amphigenous, scattered, circular to somewhat irregular, 1–4 mm diam, brown to dark brown, with dark margin. Caespituli amphigenous. Mycelium internal. Stromata lacking or very small, only formed as aggregations of a few swollen hyphal cells, substomatal. Conidiophores in loose fascicles, 2–24, arising from internal hyphae or hyphal aggregations, emerging through stomata, erect, straight, subcylindrical, barely geniculate, unbranched, 35–85 × 3–4 µm, 1–7-septate, pale brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, 10–30 µm long, with 1–3 conspicuous conidiogenous loci, somewhat thickened and darkened, about 1–2 µm diam. Conidia solitary, obclavate, 30–60 × 3–4 µm, 1–6(−7)-septate, pale olivaceous-brown, wall thin and verruculose, apex obtuse to subacute, base obconically truncate, about 1–2 µm diam, hila somewhat thickened and darkened.

Holotype: Ivory Coast: Abidjan, Forêt de Yapo, on Cercestis afzelii, Araceae, 10 Feb. 1974, G. Gilles PCI 94 (K(M) IMI 212936).

Host range and distribution: Only known from the type collection.

Notes: Deighton (1979) supposed that Cercospora cercestidis-afzelii might be a synonym of Stenella cercestidis, although the leaf spots are quite distinct and superficial mycelium is not formed. We prefer to maintain two species and consider C. cercestidis-afzelii as one of the Zasmidium species without formation of superficial hyphae in vivo. With regard to the unclear status of the type collection see notes under Z. cercestidis.

Zasmidium colocasiae (Sarbajna & Chattopadh.) Kamal, Cercosporoid Fungi of India: 240 (2010).

(Fig. 36)

Fig. 36
figure36

Zasmidium colocasiae (K(M) IMI 311131). A. Conidiophore fascicle. B. Conidiophores. C. Solitary conidiophores arising from a superficial hypha. D. Conidia. Bar = 10 µm.

Basionym: Stenella colocasiae Sarbajna & Chattopadh., J. Mycopathol. Res. 29: 35 (1991).

Illustration: Sarbajna & Chattopadhyay (1991: 36, fig. 3).

Description: Leaf spots amphigenous, circular, 2–10 mm diam, at first small, with brown centre and light brown halo, later turning greyish white, surrounded by a somewhat raised dark brown border, to 1 mm wide, occasionally with shot-hole symptoms. Caespituli amphigenous, punctiform, blackish, unevenly scattered. Stromata epiphyllous, well-developed, substomatal, 10–50 µm diam, composed of swollen hyphal cells, brown. Mycelium internal and external; superficial hyphae profusely branched, septate, 2–4.5 µm wide, pale olivaceous to brownish, thin-walled, verruculose, nodulose, with conspicuous swellings formed in regular intervals. Conidiophores in small, loose fascicle, arising from stromata, through stomata, fascicles only formed on the upper leaf surface, and solitary, arising from superficial hyphae, lateral and terminal, erect, straight to somewhat curved, subcylindrical to somewhat geniculate-sinuous above, unbranched, occasionally with intercalary swellings, rarely with percurrent rejuvenation, 15–115 × 2.5–5 µm, 1–5-septate, brown, paler towards the tip, wall somewhat thickened, smooth or almost so; conidiogenous cells integrated, terminal, conidiogenous loci conspicuous, somewhat thickened and darkened, 0.8–1.5 µm diam. Conidia solitary, occasionally two in a short chain, ellipsoid-ovoid, subcylindrical-fusiform to narrowly obclavate, straight to curved, 5–100 × 2–4 µm, 0–5-septate, pale olivaceous, thin-walled, smooth to faintly rough, apex subacute or subobtuse, base subtruncate to short obconically truncate, 0.8–1.5 µm wide, hila almost unthickened to slightly thickened and somewhat darkened-refractive.

Holotype: India: West Bengal: Ichapur, on Colocasia esculenta [antiquorum], Araceae, 10 Sep. 1986, K. K. Sarbajna, PCC 4967 (K(M) IMI 311131).

Host range and distribution: Only known from the type collection.

Zasmidium deightonianum (U. Braun) U. Braun, Schlechtendalia 20: 100 (2010).

(Fig. 37)

Fig. 37
figure37

Zasmidium deightonianum (K(M) IMI 7735). A. Conidiophores and hyphae emerging through a stoma. B. Superficial hypha. C. Superficial hypha with solitary conidiophore. D. Conidia. Bar = 10 µm.

Basionym: Stenella deightoniana U. Braun, in Braun & Crous, Mycotaxon 92: 404 (2005).

Synonyms: Cladosporium cercestidis Deighton, Mycol. Res. 94: 570 (1990), non Zasmidium cercestidis (J.M. Yen & Gilles) U. Braun, 2010.

Stenella cercestidis (Deighton) U. Braun, Schlechtendalia 5: 54 (2000), nom. illeg. (ICN, Art. 53.1).

Illustration: Deighton (1990: 570, fig. 1).

Description: Leaf spots amphigenous, circular or subcircular, 1–3 mm diam, greyish, border darker. Caespituli amphigenous, thinly effuse, covered with numerous minute darker dots, medium brown. Mycelium internal and external; immersed hyphae branched, septate, subhyaline to very pale brownish, 1–2.5 µm wide; superficial hyphae emerging through stomata, sparingly branched, septate, subhyaline to medium brown, thin-walled, almost smooth to verruculose, 1–2.5 µm wide. Stromata substomatal, 10–25 µm diam, brown. Conidiophores in small to moderately large fascicles, about 10–30, arising from stromata, through stomata, loose to moderately dense, and conidiophores solitary, arising from superficial hyphae, lateral, erect, straight, subcylindrical or somewhat swollen at the tip to somewhat geniculate-sinuous, unbranched, 10–60 × 1.5–3 µm, 0–3-septate, pale to medium brown, thin-walled, smooth or almost so; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–25 µm long, conidiogenous loci conspicuous, about 0.5–1 µm diam, thickened and darkened, often numerous and aggregated near the apex. Conidia catenate, in simple or rarely branched chains, narrowly ellipsoid-ovoid, subcylindrical, fusiform, 4–20 × 2–3 µm, 0–1-septate, pale brownish, thin-walled, verruculose, ends somewhat pointed to short obconically truncate, 0.5–1 µm wide, hila slightly thickened and darkened.

Holotype: Sierra Leone: Njala (Kori), on Cercestis congoensis, 25 Apr. 1934, F. C. Deighton (K(M) IMI 7735).

Host range and distribution: On Cercestis congoensis, Araceae, Africa (Sierra Leone).

Zasmidium extremorum (Syd.) U. Braun, Schlechtendalia 20: 101 (2010).

(Fig. 38)

Fig. 38
figure38

Zasmidium extremorum (S-F14686). A. Superficial hypha. B. Conidiophore fascicles. C. Conidiophore tips. D. Conidia. Bar = 10 µm.

Basionym: Cercospora extremorum Syd., Ann. Mycol. 15: 264 (1917).

Synonym: Stenella extremorum (Syd.) U. Braun, Nova Hedwigia 73: 434 (2001).

Literature: Saccardo (1931: 868), Chupp (1954: 59), Boedijn (1961: 413), Crous & Braun (2003: 181).

Illustration: Braun (2001a: 435, fig. 19).

Description: Leaf spots circular, subcircular to elliptic, scattered, often zonate, 2–8 mm diam, centre pale brown, greyish brown to grey, surrounded by a brownish zone and dark margin or marginal line. Caespituli hypophyllous, effuse. Mycelium internal and external; superficial hyphae emerging through stomata, sparingly branched at right angles or almost so, 1–3 µm wide, subhyaline to pale olivaceous-brown, thin-walled, verruculose. Stromata lacking or almost so or developed and substomatal, small, 10–40 µm diam, brown. Conidiophores solitary or in small to moderately large fascicles, loose to moderately dense, arising from internal hyphae or stromata, through stomata (solitary conidiophores arising from superficial hyphae not formed or at least not yet observed), erect, straight to slightly curved or sinuous, non-geniculate or only slighty so near the apex, unbranched, 10–80 × 3–5 µm, 0–6-septate, pale olivaceous, olivaceous-brown, later pale to medium dark brown, paler towards the tip, wall thin to slightly thickened, smooth or almost so; conidiogenous cells integrated, terminal, occasionally conidiophores reduced to conidiogenous cells, about 10–40 µm long, sympodially proliferating, conidiogenous loci conspicuous, 1–1.5(−2) µm diam, barely to very slightly thickened, somewhat darkened or refractive. Conidia solitary, cylindrical-filiform, subacicular to almost obclavate-cylindrical, (15−)40–300(−350) × 2–5 µm, 1- to pluriseptate, pale olivaceous to very pale olivaceous-brown, thin-walled, verruculose to verrucose, apex obtuse to subacute, base truncate to slightly obconically truncate, 1.5–2 µm wide, hila slightly thickened and somewhat darkened-refractive.

Lectotype (designated by Braun 2001): Philippines: Prov. Laguna: Luzon, Los Baños, on Homalomena philippinensis, Feb. 1914, C. F. Baker, [Fungi Malayana 521] (S-F14686). Isolectotypes: B 700016016; BPI 436255, 436356; CUP 39793, S-F14687.

Host range and distribution: On Homalomena (pendula [rubra], philippinensis, Homalomena sp.), Araceae, Asia (Indonesia, Philippines).

Arecaceae (Palmae)

Cercospora

Key to Cercospora species on Arecaceae
  1. 1

    Conidiophores 6.5–8 µm wide; conidia obclavate-cylindrical, 50–155 × 5.5–8 µm, base obconically truncate; on Raphia farinifera ...................................... C. raphiae

    Conidiophores narrower, 4–6 µm; conidia acicular, base truncate ...................................... 2

  2. 2(1)

    Stromata lacking or poorly developed; conidia narrow, 25–120 × 2–4 µm; on Cocos nucifera .......................... C. nucifera

    Stromata well-developed, 10–100 µm diam; conidia wider, 40–320 × 4–5 µm; on Areca catechu ............. C. arecacearum

Cercospora species on Arecaceae

Cercospora arecacearum Hidayat & Meeboon, in To-anun et al., Mycol. Progr. 8: 116 (2009).

(Similar to Fig. 1)

Literature: To-anun et al. (2011: 37), Phengsintham et al. (2013a: 74).

Illustrations: To-anun et al. (2009: 117, fig. 1; 2011: 38, fig. 19), Phengsintham et al. (2013a: 75, figs 2–3).

Description: Leaf spots circular, 1–7 mm diam, pale to dark brown with medium to dark brown border. Caespituli amphigenous, scattered, dark brown. Mycelium internal. Stromata oval to ellipsoid, 10–100 µm diam, substomatal to intraepidermal, brown, composed of swollen hyphal cells, subglobose, rounded to angular, 6–13 µm diam. Conidiophores fasciculate, 2–22, divergent to dense, arising from stromata, through stomata or erumpent, erect, straight to curved, cylindrical, geniculate, unbranched, 45–310 × 4–6 µm, 3–8-septate, distance between septa 8–35 µm, yellowish to medium brown throughout or paler towards the tip, wall 0.5–0.8 µm wide, smooth; conidiogenous cells integrated, terminal, 10–65 µm long, conidiogenous loci thickened and darkened, 2–3 µm diam. Conidia solitary, acicular, straight to curved, 40–320 × 4–5 µm, 3- to pluriseptate, hyaline, thin-walled, smooth, apex pointed, base truncate to slightly obconically truncate, 1.5–3 µm wide, hila somewhat thickened and darkened.

Holotype: Thailand: Chiang Mai Province: Mae Taeng District, T. Pa Pae, Mushroom Research Centre, on Areca catechu, 17 Nov. 2006, I. Hidayat (CMU 27946).

Host range and distribution: On Areca catechu, Arecaceae, Asia (Thailand).

Notes: To-anun et al. (2009) sequenced the new species and analysed its phylogenetic position. Due to consistently acicular conidia it belongs to the C. apii complex.

Cercospora nucifera R.K. Srivast., S. Narayan & A.K. Srivast., Indian Phytopathol. 48: 106 (1995).

(Fig. 39)

Fig. 39
figure39

Cercospora nucifera (based on Srivastava et al. 1995: 106, fig. 1). A. Conidiophores. B. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 293), Kamal (2010: 69).

Illustration: Srivastava et al. (1995: 106, fig. 1).

Description: Leaf spots amphigenous, circular or subcircular, greyish black, 2–20 mm diam. Caespituli hypophyllous. Mycelium internal; hyphae branched, septate. Stromata immersed, poorly developed, pseudoparenchymatous, brown. Conidiophores fasciculate, arising from stromatic hyphal aggregations, straight to curved, geniculate-sinuous, rarely branched, about 40–165 × 4–5.5 µm, 2–7-septate, pale olivaceous to light brown, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci thickened and darkened. Conidia solitary, acicular, straight to curved, 25–120 × 2–4 µm, 2–8-septate, hyaline, thin-walled, smooth, apex pointed, base truncate, hila thickened and darkened.

Holotype: India: UttarPradesh: Gorakhpur, on Cocos nucifera, Mar. 1990, R. K. Srivastava (GPU 1311, Gorakhpur University, Dept. of Botany, India). Isotype: HCIO 30926.

Host range and distribution: On Cocos nucifera, Arecaceae, Asia (India, Uttar Pradesh).

Note: This species belongs to the Cercospora apii s. lat. complex.

Cercospora raphiae Deighton, Trans. Brit. Mycol. Soc. 85: 741 (1985).

(Fig. 40)

Fig. 40
figure40

Cercospora raphiae (K(M) IMI) 147583). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Literature: Leung et al. (1997a), Crous & Braun (2003: 348).

Illustrations: Deighton (1987: 741, fig. 2), Leung et al. (1997a: 5, fig. a–g).

Description: Leaf spots amphigenous, subcircular to elliptical, slighty swollen, 4–5 × 1–3 mm, sometimes confluent and larger, 4–40 × 4–5 mm, brown on the upper surface, paler below, centre later ash grey, with darker margin. Caespituli amphigenous, mostly epiphyllous, velutinous, dark brown, evenly distributed on leaf spots. Mycelium internal; hyphae branched, septate, almost colourless, about 2.5 µm wide. Stromata immersed to erumpent, 25–115 × 20–55 µm, dark brown. Conidiophores 2–10 in divergent fascicles, arising from stromata, erumpent, straight or almost so, cylindrical to subclavate, flexuous, but usually not geniculate, unbranched, to 210 µm long and 6.5–8 µm wide (tips often somewhat swollen, to 10 µm wide), pluriseptate, light brown; conidiogenous cells integrated, terminal, conidiogenous loci conspicuous, somewhat thickened and darkened, terminal and lateral, barely prominent, 1.5–2.5 µm diam. Conidia solitary, obclavate-cylindrical, straight, about 50–155 × 5.5–8 µm, 8–16-septate, colourless, smooth, apex obtuse, base obconically truncate, 1.5–2 µm wide, hilum thickened and darkened.

Holotype: Zimbabwe: Plam block, Mayo, on Raphia farinifera, 6 Feb. 1970, A. R. Rothwell (K(M) IMI 147583).

Host range and distribution: On Raphia farinifera, Arecaceae, Africa (Zimbabwe).

Doubtful, excluded and insufficiently known species

Cercospora acrocomiae J.A. Stev., Rep. (Annual) Insular Agric. Exp. Sta. Puerto Rico (Rio Piedras) 1916–1917: 89 (1917).

Synonym: Exosporium acrocomiae (J.A. Stev.) Chupp ex J.A. Stev., Contrib. Reed Herb. 23: 516 (1975).

Literature: Chupp (1954: 428), Crous & Braun (2003: 43), Guatimosim et al. (2013).

Illustration: Guatimosim et al. (2013: 63, plate 1).

Lectoype (designated by Guatimosim et al. 2013): Puerto Rico: Rio Pedras, on Acrocomia media [aculeata], 14 Feb. 1912, J. R. Johnston [Stevenson 4206] (BPI 432400).

Isolectotypes: BPI 432402, CUP 39018.

Host range and distribution: Acrocomia aculeata [media], Arecaceae, Puerto Rico.

Note: This is an Exosporium recently re-examined and redescribed by Guatimosim et al. (2013).

Cercospora calamicola Henn., Hedwigia (Beibl.) 42: 88 (1903).

Literature: Saccardo (1906: 611), Chupp (1954: 428).

Holotype: Australia: Queensland: Cairns, on Calamus caryotoides, May 1902, Pritzel 78 (B 700014861).

Host range and distribution: On Calamus caryotoides, Arecaceae, Australia.

Notes: Not a Cercospora according to Chupp (1954), but status still unclear. Type material has been examined, but conidiophores and conidia agreeing with the original description, which would allow a reassessment of the generic affinity of this species, have not been found.

Cercospora elaeidis Steyaert, Bull. Soc. Roy. Bot. Belgique 80: 35 (1948); as “elaedis”.

Synonym: Pseudospiropes elaeidis (Steyaert) Deighton, Trans. Brit. Mycol. Soc. 85: 739 (1985).

Literature: Chupp (1954: 428), Ellis (1976: 278), Mulder & Holliday (1975).

Illustrations: Ellis (1976: 277, fig. 211 B), Deighton (1985: 740, fig. 1), Mulder & Holliday (1975: fig., unnumbered).

Description: Leaf spots amphigenous, subcircular, elliptical to irregular, small, often slightly depressed, brown to dark brown, margin indefinite or with somewhat raised paler border or orange halo. Colonies hypophyllous, not very conspicuous. Mycelium internal; hyphae branched, about 1.5–3 µm wide, septate, thin-walled, subhyaline to brownish. Stromata almost lacking or small, 10–30 µm diam, immersed, brown. Conidiophores in small fascicles, mostly 2–7, occasionally solitary, arising from stromata, through stomata, erect, straight, subcylindrical to somewhat geniculate-sinuous, unbranched, about 100–350 × 5–9 µm, base often somewhat swollen, pluriseptate, medium to dark brown, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal, to 80 µm long, with a single to several conspicuous conidiogenous loci, circular, 2.5–4 µm diam, often somewhat prominent, with a conspicuous central pore. Conidia solitary, obclavate to broadly acicular, in longer conidia apical portion gradually attenuated towards the tip, rostrate, straight to curved, 40–185 × 5–9 µm, (1−)4–12-septate, pale golden brown, tips often paler, thin-walled, almost smooth to faintly verruculose, apex obtuse, base truncate to obconically truncate, 2–4 µm wide, hila somewhat thickened and darkened.

Holotype: Democratic Republic of Congo: Kodoro, on Elaeis guineensis, 19 Nov. 1942, Steyaert (INÉAC “Herbario Division de Phytopathologie de l’Institut National pour l’étude Agronomique du Congo Belge”, B.141[332]; not traced).

Host range and distribution: On Elaeis guineensis, Arecaceae, Africa (Angola, Benin, Cameroon, Congo, Ethiopia, Gabon, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, Nigeria, São Tomé e Príncipe, Senegal, Sierra Leone, Sudan, Tanzania, Togo, Zimbabwe), Australia, South America and West Indies (Cuba, Netherlands Antilles, Suriname).

Notes: This species seems to be confined to Elaeis guineensis (Deighton 1985). Records on Carpentaria and Raphia spp. are wrong and pertain to Pseudocercospora species. Other records on Phoenix sp. and unidentified palms are doubtful and unproven. The generic affinity of C. elaeidis is quite unclear. Due to pigmented conidia, this species has to be excluded from Cercospora. Deighton (1985) placed it in Pseudospiropes, but this genus, introduced for saprobic species on wood and bark and distoseptate conidia, is not suitable for this species. A formal inclusion in Passalora would be possible, but this genus is still polyphyletic, and verruculose conidia are also not in favour of Passalora. New collections, cultures and sequence data are necessary to point out the true generic affinity of this species. It is unclear what happened with the herbarium of the pytopathological division of INÉAC. This institute was closed in December 1962. Type material of C. elaeidis is not included in Steyaert’s herbarium at BR. Therefore, the designation of a neotype is needed. Collections of this species from Congo now deposited at K have been examined (K(M) IMI56702a and 59521), but they are too meagre and not suitable as neotypes.

Cercospora licualae Syd. & P. Syd., Philipp. J. Sci. (Bot.) 9: 188 (1914).

Synonym: Cercospora virens Sacc., Boll. Orto Bot. Regia Univ. Napoli 6: 62 (1921) [syntypes: Singapore: Straits Settlement, Licuala sp., Oct. 1917, C. F. Baker 407 (B 700014863; PAD)].

Literature: Saccardo (1931: 872, 885–886), Chupp (1954: 429).

Syntypes: Philippines: Palawan, Taytay, on Licuala spinosa, Apr. 1913, E. D. Merrill 8748 (S-F20464 [slide K(M) IMI 88990], W).

Host range and distribution: On Licuala (spinosa, Licuala sp.), Arecaceae, Asia (Philippines, Singapore).

Notes: Chupp (1954) excluded this species and stated that it does not belong to Cercospora, which could be confirmed by our re-examinations of the type collections. The true generic affinity is, however, quite unclear.

Cercospora palmae-amazonensis Bat. & Cavalc., Anais XII Congr. Soc. Bot. Brasil: 385 (1964).

Illustration: Batista & Cavalcanti (1964: 386, fig. 102).

Description: Leaf spots amphigenous, indeterminate, effuse, brown. Caespituli amphigenous, black. Stromata oblong, black, 22–25 µm diam. Conidiophores fasciculate, 2–10, erect, straight, simple or geniculate, unbranched, 112–250 × 4–5 µm, swollen at the base, 4–7.5 µm diam, 6–10-septate, cells 21–25 × 2.5–5 µm, brown, paler towards the tip, smooth; conidiogenous cells integrated, terminal and intercalary. Conidia solitary, obclavate, 40–90 × 2.5–7.5 µm, 6–9-septate, not constricted at the septa, brownish, apex obtuse, base obconically truncate.

Holotype: Brazil: Amazonas, Manaus, on an unidentified palm, 17 Feb. 1961, A. C. Batista (IMUR 21246). Isotypes: INPA, PC.

Host range and distribution: Unidentified palm, Arecaceae, South America (Brazil).

Notes: Type material of this species has been examined (Fungos do Brasil no. 21246, ex herb. IMUR, PC), but no sporing structure could be found. The identity of this fungus is unclear. It is even uncertain if this species is cercosporoid at all. According to the original drawing it could also be a helminthosporioid fungus.

Cercospora palmicola Speg., Anales Soc. Ci. Argent. 26: 72 (1888).

Synonyms: Drechslera palmicola (Speg.) F. Anderson, M.V. Bianchinotti & U. Braun, Schlechtendalia 5: 67 (2000).

Corynespora palmicola (Speg.) U. Braun, comb. nov.

MycoBank MB809010

Basionym: Cercospora palmicola Speg., Anales Soc. Ci. Argent. 26: 72 (1888).

Literature: Lindau (1910: 88); Vassiljevsky & Karakulin (1937: 317), Chupp (1954: 429), Crous & Braun (2003: 303).

Exsiccatae: Roum., Fungi Sel. Exs5188.

Description: Leaf spots amphigenous, elliptical to somewhat irregular, 10–20 mm long and 3–8 mm wide, brownish, confluent. Caespituli punctiform, scattered, dark. Mycelium internal. Stromata developed, composed of swollen hyphal cells, circular in outline, brown. Conidiophores in loose to dense fascicles, arising from stromata, erect, straight to curved, subcylindrical to clavate, unbranched, 15–50 × 3–6 µm, aseptate, wall thin to somewhat thickened, brown or olivaceous-brown, smooth; conidiophores reduced to conidiogenous cells, monotretic, with a single terminal minute porus, not darkened or slightly so around the porus, not proliferating. Conidia solitary, obclavate-subcylindrical, 40–70 × 6–9 µm, 5–7-distoseptate, appearing thick-walled by distoseptation and reduced lumina, smooth, apex obtuse, broadly rounded, base short obconically truncate, distinctly darkened.

Lectotype (designated here, MycoBank MBT178178): Paraguay: Guarapí, on Syagrum romanzoffianum [Cocos australis], 19 Oct. 1883, B. Balansa 4070 (LPS 925). Isolectotype: B 700016013. Topotypes: Roum., Fungi Sel. Exs. 5188.

Host range and distribution: On Syagrum romanzoffianum, Phoenix canariensis, Arecaceae, Europe (Germany, Russia), South America (Paraguay).

Notes: The generic affinity of this species is intricate and disputable. The reallocation to Drechslera (Anderson et al., in Braun 2000) is doubtful since this genus is characterized by having polytretic, sympodially proliferating conidiogenous cells. Due to monotretic conidiogenous cells and pluridistoseptate conidia with pigmented hila, C. palmicola is rather assignable to Corynespora, although aseptate, not proliferarting conidiophores and less conspicuous pores are unusual in this genus. However, some species with at least rather inconspicuous pores and not percurently proliferating conidiophores are known in Corynespora, e.g. C. cubensis

Hol.-Jech. and C. pseudolmediae (R.F. Castañeda) Hol.-Jech. (Mercado Sierra et al. 1997).

Cercospora palmicola f. stilbacea C. Moreau, Rev. Mycol. 12: 38 (1947).

Synonyms: Helminthosporium stilbaceum (C. Moreau) S. Hughes, Mycol. Pap. 48: 38 (1952).

Exosporium stilbaceum (C. Moreau) M.B. Ellis, Mycol. Pap. 82: 38 (1961).

Literature: Chupp (1954: 429), Ellis (1971: 403).

Illustrations: Moreau (1947: 39, figs 1–3), Ellis (1971: 401, fig. 274B).

Type: Republic of Congo: Etoumbi, Fort Rousset, on Elaeis guineensis, Mar. 1947, M. A. Bachy (probably not preserved).

Host range and distribution: On Elaeis (guineensis, Elaeis sp.), Arecaceae, Africa (Congo, Ghana, Guinea, São Tomé and Príncipe, Sierra Leone, Sudan, Tanzania, Zambia), Asia (Malaysia).

Cercospora virens — see Cercospora licualae

Distocercospora

A single species.

Distocercospora livistonae U. Braun & C.F. Hill, in Braun et al., Fungal Diversity 22: 23 (2006).

(Fig. 41)

Fig. 41
figure41

Distocercospora livistonae (HAL 1875 F). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Illustration: Braun et al. (2006: 24, fig. 3).

Description: Leaf spots amphigenous, subcircular to irregular, 2–15 mm diam, pale to dark brown, finally greyish brown to greyish white, margin indefinite or with a diffuse, irregular darker border. Caespituli hypophyllous, finely punctiform, dark brown. Mycelium internal. Stromata lacking or small, 10–30 µm diam, substomatal, brown. Conidiophores in small to moderately large, loose fascicles, arising from internal hyphae or stromata, through stomata, erect, straight, subcylindrical-filiform, usually distinctly geniculate-sinuous, above all in the upper half, unbranched, 40–280 × 3–6 µm, pluriseptate throughout, wall somewhat thickened below, thin-walled towards the apex, pale to medium dark brown or olivaceous-brown, smooth; conidiogenous cells integrated, terminal and intercalary, 10–30 µm long, proliferation sympodial or occasionally percurrent, conidiogenous loci conspicuous, thickened and darkened, 2–2.5 µm diam. Conidia solitary, obclavate, 20–85 × 4–7 µm, indistinctly 2–5-distoseptate, pale olivaceous, outer wall very thin, inner wall to 2 µm thick, almost smooth to distinctly verruculose, apex obtuse or subobtuse, base short obconically truncate, 2–3 µm wide, hila somewhat thickened and darkened.

Holotype: New Zealand: Auckland, Manurewa, Auckland Regional Botanic Gardens, Hill Road, on Livistona chinensis, 10 Sep. 2005, C. F. Hill 1247 (HAL 1875 F).

Host range and distribution: On Livistona chinensis [boninensis, chinensis var. boninensis], Arecaceae, Asia (Japan), New Zealand.

Passalora

Key to Passalora species on Arecaceae
  1. 1

    Stromata small, 10–30 µm diam; conidia 40–185 × 5–9 µm, (1−)4–12-septate, often rostrate, mostly verruculose, pale golden brown; on Elaeis guineensis ....................................... see Cercospora elaeidis (excluded and doubtful species)

    Stromata larger, 50–160 µm diam; conidia much shorter (to 70 µm) and only 1–3-septate, hyaline, subhyaline or brown to olivaceous-brown, smooth ....................................... 2

  2. 2(1)

    Stromata large, about 85–160 µm diam, textura intricata; conidiophores 90–105 µm long, 1–3-septate; conidia cylindrical to somewhat clavate, 8–13 µm wide, pale brown or olivaceous-brown; on Acrocomia ....................................... P. acrocomiae

    Stromata smaller, 50–70 µm diam, textura angularis; conidiophores much longer, 60–300 µm, (1−)3–5-septate; conidia obclavate, narrower, 2–7 µm wide, hyaline to subhyaline; on Syagrus ......................... P. eitenii

Passalora species on Arecaceae

Passalora acrocomiae Guatimosim & R.W. Barreto, Mycotaxon 122: 64 (2013).

(Fig. 42)

Fig. 42
figure42

Passalora acromiae (BPI 432398). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Illustration: Guatimosim et al. (2013: 65, plate 2).

Description: Leaf spots amphigenous, subcircular to elliptical, 0.9–2.3 × 0.3–0.6 cm, with pale centre surrounded by a pale brown border. Caespituli hypophyllous. Mycelium internal; hyphae branched, 2.5 µm wide, septate, subhyaline. Stromata subepidermal, erumpent, subglobose, 100–157 × 87–113 µm, composed of textura intricata, pale brown, smooth. Conidiophores numerous, to 20, in dense fascicles, arising from stromata, erumpent, erect, straight to somewhat curved-sinuous, subcylindrical or somewhat enlarged towards the tip, unbranched, 90–105 × 5–7 µm, 1–3-septate, pale brown, thin-walled, smooth; conidiogenous cells integrated, terminal, subcylindrical, 37–55 × 6–8 µm, conidiogenous loci conspicuous, one per cell, discoid, 1.5–4 µm diam. Conidia solitary, subcylindrical to slightly clavate, 35–68 × 8–13 µm, 1–3-septate, pale brown or olivaceous-brown, thin-walled, smooth, but apex distinctly roughened, apex rounded, base obconically truncate, 2–4 µm wide, hila somewhat thickened and darkened.

Types: Puerto Rico: Rio Piedras, on Acrocomia media, Arecaceae, 15 Jul. 1914, J. A. Stevenson 2090 (BPI 432398). Isotypes: BPI 432398, 432401.

Host range and distribution: Only known from the type collection.

Notes: Cercospora acrocomiae was a confused name. The original collections from Puerto Rico (Rio Pietras) contain two different fungal species, an Exosporium and a Passalora. Guatimosim et al. (2013) solved this problem by means of a lectotypification of the name C. acrocomiae in the sense of Chupp (1954) with material representing the Exosporium (see “Excluded and doubtful species”) and described the Passlora as new species.

Passalora eitenii R.B. Medeiros & Dianese, Mycotaxon 51: 509 (1994).

(Fig. 43)

Fig. 43
figure43

Passalora eitenii (UB, Coll. Mycol. 881). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Illustration: Medeiros & Dianese (1994: 511, plate 1, A–H).

Description: Leaf spots amphigenous, mainly hypophyllous, necrotic, dark brown to black, usually elliptical, rarely confluent, 10–15 × 2–2.5 mm. Caespituli hypophyllous, effuse, velutinous, brown. Mycelium immersed; hyphae septate, branched, 3–5 µm wide, brown, smooth. Stromata substomatal, 50–70 µm diam, brown, cells angular. Conidiophores fasciculate, 3–12, arising from stromata, through stomata, erect, cylindrical, distal end somewhat geniculate-sinuous, unbranched, 60–300 × 5–10 µm, (1−)3–5-septate, brown, smooth, wall somewhat thickened; conidiogenous cells integrated, terminal, sympodially and occasionally percurrently proliferating; conidiogenous loci conspicuous. Conidia solitary, obclavate, straight, 27–55 × 2–7 µm, 1–3-septate, mostly 1-septate, hyaline or subhyaline, wall thickened, smooth, apex obtuse, base obconically truncate, hilum conspicuous.

Holotype: Brazil: D.F.: Brasilia, on Syagrus comosa, Arecaceae, Nov. 1991, J. C. Dianese (UB, Coll. Mycol. 881).

Host range and distribution: Only known from the type collections.

Pseudocercospora

Key to Pseudocercospora species on Arecaceae
  1. 1

    Stromata large, 20–150 µm diam; conidiophores in sporodochial conidiomata, 10–30 × 4–7 µm, proliferation strictly percurrent, with 0–5 conspicuous annellations; conidia obclavate-cylindrical, 60–150 × 5–10 µm, verrucose; on various palms ................................................................... see Scolecostigmina palmicola

    Stromata usually 10–50 µm diam; conidiophores sympodially proliferating or occasionally a few percurrent proliferations mixed with sympodial ones; conidia smooth or almost so ................................... 2

  2. 2(1)

    Conidia narrow, 2–4 µm wide, hila 1–2.5 µm wide; on Copernicia or Rhapis ............................................... 3

    Conidia broader, 3.5–8 µm, hila wider, 2–5 µm; on other hosts ..................................................... 4

  3. 3(2)

    Conidiophores very short, 5–15 µm, usually aseptate; conidia (10−)15–50(−60) µm long, (0−)1–4(−6)-septate; on Copernicia ....................................................... P. coperniciae

    Conidiophores 10–30 µm long, 0–2-septate; conidia much longer, 30–160 µm, 0–10-septate; on Rhapis ................................................... P. rhapisicola

  4. 4(2)

    Conidiophores very long, to 170 µm, 5–7(−8) µm wide, pluriseptate; conidia obclavate, 40–90 × 5–9 µm; on Carpentaria ................................................................... P. carpentariae

    Conidiophores shorter, to 100 µm, 2.5–6 µm wide, aseptate or sparingly septate; conidia obclavate-cylindrical to fusiform, narrower, 2.5–6(−7) µm; on other hosts ...................................... 5

  5. 5(4)

    Leaf spots distinct, variable in shape and size, often oblong, covering large leaf segment or entire leaf blades discoloured; conidial shape very variable, broadly cylindrical, obclavate, fusiform to subacicular or subclavate; on Howea and Rhopalostylis, New Zealand .................... P. arecacearum

    Leaf spots lacking; conidia obclavate-cylindrical, rarely fusiform; on Roystonea, North America, USA ......... P. roystoneae

Pseudocercospora species on Arecaceae

Pseudocercospora arecacearum U. Braun & C.F. Hill, in Braun et al., Fungal Diversity 22: 25 (2006).

(Fig. 44)

Fig. 44
figure44

Pseudocercospora arecacearum (HAL 1876 F). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Illustration: Braun et al. (2006: 24, fig. 4).

Description: Leaf spots conspicuous, variable in shape and size, often oblong, covering large leaf segment or entire leaf blades discoloured, necrotic, straw-coloured, yellowish, ochraceous to dingy brown or greyish brown, margin indefinite. Mycelium internal, occasionally with superficial hyphae, emerging through stomata, branched, septate, 1.5–3 µm wide, pale olivaceous, thin-walled, smooth. Stromata substomatal, rarely intraepidermal, 10–50 µm diam, rarely confluent and larger, to 90 µm diam, olivaceous-brown, occasionally somewhat erumpent, composed of swollen hyphal cells, 2–6 µm diam. Conidiophores in small, loose to moderately large and dense fascicles, arising from stromata, emerging through stomata, occasionally erumpent, erect, straight, subcylindrical-conical to moderately geniculate-sinuous, usually unbranched, rarely branched, long conidiophores sometimes subclavate, 5–100 × 2.5–6 µm, aseptate or sparingly septate, pale to medium olivaceous or olivaceous-brown, wall thin or only slightly thickened, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–40 µm long, mostly with a single conidiogenous locus, determinate, occasionally sympodial, with 2–3 loci, truncate to convex, 2–4 µm wide, unthickened, not darkened. Conidia solitary, shape and size variable, broadly cylindrical, obclavate, fusiform to subacicular or subclavate, 20–130 × 3.5–7 µm, 2–20-septate, occasionally somewhat constricted at the septa, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth, fresh conidia with oil droplets, apex obtuse, base truncate or short to long obconically truncate, 2–5 µm wide, hila unthickened, not darkened.

Holotype: New Zealand: Auckland, St. Johns, Morrin Road, The Atrium, on Rhopalostylis sapida, 4 Jul. 2005, C. F. Hill 1209 (HAL 1876 F). Ex type culture: CBS 118406.

Host range and distribution: On Howea forsteriana, Rhopalostylis (baueri var. cheesemanii, sapida), Arecaceae, New Zealand.

Pseudocercospora carpentariae Deighton, Trans. Brit. Mycol. Soc. 89: 403 (1987).

(Fig. 45)

Fig. 45
figure45

Pseudocercospora carpentariae (K(M) IMI 305079). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Deighton (1985: 742, as Peudocercospora sp.), Leung et al. (1997b).

Illustrations: Deighton (1985: 742, fig. 3; 1987: 403, fig. 1), Leung et al. (1997b: 5, fig. a–f).

Description: Leaf spots amphigenous, circular, subcircular to elliptical, to 8 mm long and 2–4 mm wide, dark brown, surrounded by a yellowish halo, about 1 mm wide. Caespituli hypophyllous, dark brown, sparsely distributed at the centre. Mycelium internal; hyphae branched, septate, 1.5–2.5 µm wide, subhyaline. Stromata immersed, erumpent, compact, 15–45 µm diam, brown. Conidiophores in divergent fascicles, 10–20, arising from stromata, erumpent, erect, straight to slighty geniculate-sinuous, unbranched, to 170 × 5–7(−8) µm, pluriseptate, medium brown, wall thin to somewhat thickened, smooth; conidiogenous cells integrated, terminal, conidiogenous loci inconspicuous, sometimes subdenticulate, but always unthickened and not darkened. Conidia solitary, straight to slightly curved, obclavate, 40–90 × 5–9 µm, 4–10-septate, pale olivaceous-brown, thin-walled, smooth, apex obtuse to subacute, base short obconically truncate, hilum neither thickened nor darkened.

Holotype: Australia: Northern Territory: Darwin, on Carpentaria acuminata, 14 May 1986, J. Duff & B. Conde (K(M) IMI 305079).

Host range and distribution: On Carpentaria acuminata, Arecacae, Australia (Northern Territory).

Pseudocercospora coperniciae U. Braun & F.O. Freire, Cryptog. Mycol. 23: 308 “2002” (2003).

(Fig. 46)

Fig. 46
figure46

Pseudocercospora coperniciae (HAL 1709 F). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Illustration: Braun & Freire (2003: 310, fig. 12).

Description: Leaf spots amphigenous, subcircular to irregular, 3–15 mm diam or confluent and larger, pale to greyish brown, finally dingy grey, margin indefinite or with a diffuse darker border. Caespituli amphigenous, punctiform, loose to dense, dark brown. Mycelium internal. Stromata substomatal to intraepidermal, 10–30(−40) µm diam, olivaceous-brown. Conidiophores in small to moderately large fascicles, loose to moderately dense, arising from stromata, through stomata or erumpent, erect, straight, subcylindrical-conical to moderately geniculate-sinuous, unbranched, 5–15 × 2–5 µm, 0(−1)-septate, pale olivaceous, thin-walled, smooth; conidiophores usually reduced to conidiogenous cells, conidiogenous loci inconspicuous, occasionally subdenticulate, but always unthickened and not darkened. Conidia solitary, cylindrical to obclavate-cylindrical, occasionally fusiform, (10−)15–50(−60) × 2–3.5 µm, (0−)1–4(−6)-septate, subhyaline to pale olivaceous, apex obtuse to subacute, base truncate to obconically truncate, 1–2 µm wide, hila unthickened, not darkened.

Holotype: Brazil: State of Ceará: Cascaval County, Preaoca District, on Copernicia prunifera, 28 Feb. 2001, F. O. Freire (HAL 1709 F).

Host range and distribution: On Copernicia prunifera, Arecaceae, South America (Brazil, Ceará).

Pseudocercospora rhapisicola (Tominaga) Goh & W.H. Hsieh, Trans. Mycol. Soc. Republ. China 4: 39 (1989).

(Fig. 47)

Fig. 47
figure47

Pseudocercospora rhapisicola (NIAES 10300). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora rhapisicola Tominaga, Trans. Mycol. Soc. Japan 5: 57 (1965).

Literature: Hsieh & Goh (1990: 258), Guo & Hsieh (1995: 242), Guo et al. (1998: 253), Leung et al. (1997c).

Illustrations: Hsieh & Goh (1990: 258, fig. 198), Guo & Hsieh (1995: 243, fig. 205), Leung et al. (1997c: 17, fig. a–c), Guo et al. (1998:253, fig. 210).

Description: Leaf spots amphigenous, scattered, at first tiny spots, later small irregular patches, pale brown, margin indefinite, with yellowish halo. Caespituli amphigenous, punctiform, pale grey. Mycelium internal. Stromata substomatal, 10–40 µm diam, dark brown. Conidiophores in fascicles, 10–20, loose to dense, straight to curved, subcylindrical or attenuated towards the tip, geniculate-sinuous, unbranched, 10–30 × 2–4 µm, 0–2-septate, pale brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, conidiogenous loci inconspicuous, occasionally subdenticulate, unthickened and not darkened. Conidia solitary, straight, curved to somewhat geniculate-sinuous, narrowly obclavate to filiform, 30–160 × 2–4 µm, 0–10-septate, hyaline or subhyaline, thin-walled, smooth, apex subobtuse to subacute, base short obconically truncate, 1–2.5 µm wide, hila neither thickened nor darkened.

Holotype: Japan: Tokyo, Edogawa, on Rhapis excelsa, 12 Jul. 1960, M. Yokohama (NIAES 10300, Herb. National Institute of Agrico-Environmental Sciences, Tsukuba, Japan). Isotype: NIAES 10301. Possible ex-holotype isolates: MAFF 35042, NBRC 8433, CBS 282.66.

Host range and distribution: On Rhapis (excelsa [fiabelliformis], humilis), Arecaceae, Asia (Japan, Korea, Taiwan), North America (USA, California, Florida), Hawaii.

Pseudocercospora roystoneae U. Braun & Crous, Mycol. Progr. 2: 203 (2003).

(Fig. 48)

Fig. 48
figure48

Pseudocercospora roystoneae (CUP 41024). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Illustration: Braun et al. (2003a: 204, fig. 9).

Description: Leaf spots lacking, indefinite (probably on leaves from seedlings), or with large diffuse discolorations, forming necrotic patches, often large leaf segments or almost entire leaves discoloured, straw-coloured to brown or dingy greyish brown, margin indefinite. Caespituli amphigenous, punctiform, scattered, dark brown to black. Mycelium internal. Stromata substomatal, 10–50 µm diam, brown. Conidiophores in small to moderately large fascicles, loose to dense, arising from stromata, emerging through stomata, erect, straight, subcylindrical to strongly geniculate-sinuous, unbranched, orwith short lateral branchlets, 5–60 × 3–6 µm, 0–4-septate, pale olivaceous to medium olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–25 µm long, conidiogenous loci inconspicuous, unthickened, not darkened. Conidia solitary, obclavate-cylindrical to broadly fusiform, 40–100 × (4−)5–6(−7) µm, 4–10-septate, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth, apex obtuse, base obconically truncate, 2–3.5 µm wide, hila neither thickened nor darkened.

Holotype: USA: Florida: Gainesville, on Roystonea regia, Arecaceae, 16 May 1957, J. L. Smith (CUP 41024).

Host range and distribution: Only known from the type collection.

Note: This species resembles Pseudocercospora carpentariae, which differs, in having much smaller stromata, longer conidiophores, to 160 µm and wider conidia, 6.5–8 µm.

Scolecostigmina

A single species.

Scolecostigmina palmivora (Sacc.) Kamal, Cercosporoid Fungi of India: 260 (2010).

(Fig. 49)

Fig. 49
figure49

Scolecostigmina palmivora (HAL, Rabenh., Fungi Eur. Exs. 4392). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Exosporium palmivorum Sacc., Annual Rep. Missouri Bot. Gard. 9: 159 (1898).

Synonyms: Cercospora palmivora (Sacc.) Nannizzi, Atti Reale Accad. Fisiocrit. Siena X, 2: 491 (1928).

Stigmina palmivora (Sacc.) S. Hughes, Mycol. Pap. 49: 13 (1952).

Exosporium preisii Bubák, Ann. Mycol. 2: 400 (1904) [lectotype (designated here, MycoBank MBT178136): Czech Republic: Bohemia: Krc, near Prague, on Phoenix reclinata, 13 Sep. 1901, F. Petrák (BPI 454537); isolectotypes: Rabenh., Fungi Eur. 4392 (e.g. B, BPI 439978, LEP, and Vestergr., Micromyc. Rar. Sel. Praec. Scand. 471, e.g., BPI 439979, HBG].

Cercospora preisii Bubák, Ann. Mycol. 2: 400 (1904), valid alternative name (ICN, Art 36.2).

Literature: Saccardo (1892: 653; 1906, 687), Vassiljevsky & Karakulin (1937: 317), Chupp (1954: 429), Ellis (1971: 146), Crous & Braun (1996: 293), Colán et al. (2013).

Illustrations: Bubák (1904: 401, fig. 20–21), Ellis (1971: 145, fig. 97 D), Kamal (2010: 261, fig. 40), Colmán et al. (2014: 849, figs A–E).

Exsiccatae: Cif., Mycofl. Doming. Exs. 31. Kabát & Bubák, Fungi Imp. Exs. 398. Krypt. Exs. 2530. Petr., Fl. Bohem. Movar. Exs. 576. Petr., Mycoth. Gen. 631. Rabenh., Fungi Eur. 4392. Syd., Mycoth. Germ. 200. Vestergr., Micromyc. Rar. Sel. Praec. Scand. 471.

Description: Leaf spots amphigenous, subcircular to somewhat angular-irregular, about 1–10 mm diam, brown or pale, margin indefinite or darker. Caespituli amphigenous, punctiform, sometimes confluent, dark, scattered. Mycelium internal. Stromata well-developed, immersed to erumpent, 20–150 µm diam, dark olivaceous-brown or brown. Conidiophores numerous, in dense sporodochial conidiomata, arising from stromata, erect, straight to somewhat curved, cylindrical to subcylindrical, 10–30 × 4–7 µm, aseptate or only septate at the base, medium to dark brown, wall somewhat thickened, smooth to verrucose; conidiogenous cells integrated, terminal or conidiophores usually reduced to conidiogenous cells, 0–5 times percurrently proliferating, with distinct annellations, with a single terminal locus, broad and truncate, locus wall immediately after conidial secession not thickened, but annellations often conspicuous and somewhat darkened. Conidia solitary, obclavate-subcylindrical, straight to slightly curved, 60–150 × 5–10 µm, 4–16-septate, medium to dark brown, wall somewhat thickened, verrucose, apex obtuse, broadly rounded, base truncate to somewhat obconically truncate, hilum wall not thicker than lateral walls, with thin-walled somewhat bulging papilla-like projection.

Lectotype (designated here, MycoBank MBT178137): USA: Nebraska: Plattsmouth, on Phoenix canariensis, Oct. 1897, W. J. Hesser (BPI 454522).

Host range and distribution: On Acoelorrhaphe wrightii, Adonidia merrillii [Veitchia merrillii], Syagrum romanzoffianum [Arecastrum romanzoffianum], Borassus flabellifer, Brahea armata, Butia capitata, Caryota (mitis, urens), Chamaerops humilis, Cocos nucifera, Corypha sp., Howea forsteriana, Phoenix (acaulis, canariensis [tenuis], dactylifera, loureiroi [hanceana], reclinata, roebelenii, sylvestris), Rhapis excelsa, Roystonea (regia [elata]), Sabal palmetto, Thrinax sp., Washingtonia robusta, Arecaceae, Africa (Ghana, Kenya, South Africa, Zimbabwe), Asia (China, India, Japan, Myanmar, Taiwan), Caucasus (Armenia), Europe (Austria, Czech Republic, Germany), North America (Mexico; USA, Florida, Gulf states, Louisiana, Missouri, Mississippi, Nebraska, New York, Ohio, Texas), Central and South America (Brazil, El Salvador, Nicaragua), West Indies (Cuba, Dominican Republic).

Notes: Based on the phylogenetic position of its type species, Stigmina has been reduced to synonymy with Pseudocercospora. Due to its conidial shape and size, Stigmina palmivora is rather scolecostigmina-like, which is supported by its phylogenetic position. Brazilian material of this species has recently been cultivated and sequenced. Results of molecular sequence analyses revealed Pseudocercospora colombiensis and P. irregulariramosa as closest relatives with 93% ITS nucleotide homology and 98% LSU nucleotide homology. Pseudocercospora colombiensis does not cluster within the Pseudocercospora s. str. clade, but it belongs to a clade composed of Pallidocercospora, Scolecostigmina, Trochophora and some pseudocercosporoid species (Crous et al. 2013). Sequences based on material from Japan (unpublished data) are close to those from Brazil, suggesting that a single widespread species is involved. Type material of this species is not preserved in Saccardo’s herbarium at PAD. The designated lectotype, deposited at BPI and denominated as syntype, is the only type material that could be traced.

Zasmidium

A single species.

Zasmidium caryotae (X.J. Liu & Y.Z. Liao) Kamal, Cercosporoid Fungi of India: 239 (2010).

(Fig. 50)

Fig. 50
figure50

Zasmidium caryotae (HMAS 10262). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Basionym: Stenella caryotae X.J. Liu & Y.Z. Liao, Acta Microbiol. Sin. 20: 119 (1980).

Literature: Guo (1999: 358).

Illustration: Liu & Liao (1980: 120, fig. 4).

Description: Leaf spots subcircular, 1–7 mm diam, or oblong, 5–16 × 2–9 mm, dark brown to blackish, sometimes with yellowish border or halo. Caespituli amphigenous, mostly hypophyllous. Mycelium internal and external; superficial hyphae verruculose. Stromata hemispherical, 25–55 µm diam, dark brown. Conidiophores in divergent fascicles, arising from stromata, erect, straight to curved, subcylindrical, 0–4 times geniculate, apex truncate to somewhat swollen, unbranched, about 40–290 × 3–6.5 µm, 2–12-septate, olivaceous-brown to medium brown, wall thin to somewhat thickened, smooth, occasionally with percurrent rejuvenations; conidiogenous cells integrated, terminal, sympodial, with several conspicuous conidiogenous loci, thickened and darkened. Conidia solitary, obclavate, straight to curved, 30–290 × 5–8 µm, pluriseptate, pale olivaceous-brown, thin-walled, verruculose, apex obtuse to subacute, base truncate to obconically truncate, hila thickened and darkened.

Holotype: China: Guangdong: Gaoyang Xian, Dinghuashan, on Caryota mitis, 7 Feb. 1958 (HMAS 10262).

Host range and distribution: On Caryota (mitis, Caryota sp.), Arecaceae, Asia (China, Guangdong, Guangxi; India).

Doubtful, excluded and insufficiently known species

Stenella caryotae-urentis R.K. Chaudhary, Tripathi, P.N. Sing & S. Chaudhary, Indian Phytopathol. 54: 228 (2001); as “caryotae-urensis”.

Illustration: Chaudhary et al. (2001:230, fig. 3)

Description: Leaf spots amphigenous, circular to irregular, 1–6 mm diam, dark brown. Caespituli amphigenous, effuse, dark brown. Stromata lacking. Mycelium internal and external; superficial hyphae branched, 2.5–3.5 µm wide, septate, light brown, finely verruculose. Conidiophores solitary, arising from superficial hyphae, lateral, erect, about 25–75 × 3.5–4 µm, 2–4-septate, brown; conidiogenous cells integrated, terminal, cicatrized. Conidia solitary or catenate, in simple or branched chains, subcylindrical-vermiform, 10–100 × 3–5.5 µm, 1–11-septate, often constricted at the septa, pale olivaceous, verrucose, apex rounded to subobtuse, base truncate.

Holotype: India: Uttar Pradesh: Gorakhpur, on Caryota urens, Arecaceae, May 1997, R. K. Chaudhary (HCIO 42564).

Isotype: GPU Herb. 5/97.

Host range and distribution: Only known from the type collection.

Notes: The generic affinity of this species is unclear and has to be proven on the base of a re-examination of type material, which was not available for us. Stenella caryotae-urentis was neither cited nor reassessed in Kamal (2010). Details of the conidiogenous loci are not described, and the conidia are rather sirosporium-like.

Stenella novae-zelandiae Matsush., Matsush. Mycol. Mem. 4: 18 (1985).

Illustration: Matsushima (1985: figs 318, 319).

Holotype: New Zealand: near Hari hari, isolated from dead wood, Arecaceae, 21 May 1983 (MFC-12458).

Notes: This species has nothing to do with Stenella as well as Zasmidium and is not cercosporoid at all. The superficial mycelium is smooth, conidiogenous cells have a geniculate, characteristic zig-zag structure with numerous truncate, unthickened conidiogenous loci and catenate conidia with truncate base and unthickened hila. The generic affinity of this species is quite unclear, but this fungus reminds one of asexual morphs of the Venturiaceae, above all Pseudocladosporium.

Stenella palmicola Matsush., Matsush. Mycol.Mem. 4: 19 (1985).

Illustration: Matsushima (1985: fig. 320).

Holotype: New Zealand: Waipoua Forest, isolated from dead petioles of Rhopalostylis sapida, Arecaceae, 12 May 1983 (MFC-12291).

Notes: This species is quite distinct from Stenella and Zasmidium and rather pseudocladosporium-like. The mycelium, conidiophores and conidia are smooth-walled, the conidiogenous loci and conidial hila are truncate, neither thickened nor darkened, and the conidia are formed in simple or branched acropatal chains.

Stenella taiwanensis Matsush., Matsushima Mycol. Mem. 5: 30 (1987).

Illustration: Matsushima (1987: figs 447, 448).

Holotype: Taiwan: Nan-Jen-Shan, isolated from a dead rhachis of Areca catechu, Arecaceae, 27 Feb. 1986 (MFC-6T196).

Notes: This species is quite distinct from Stenella and Zasmidium. The conidiophores have a single terminal, broadly truncate, unthickened conidiogenous locus giving rise to sporidesmium-like conidia formed singly or in short chains. The general habit of this fungus suggests an affinity to the Sporidesmium complex.

Asparagaceae (including Agavaceae, Convallariaceae, and Hostaceae)

Cercospora

Key to Cercospora species on Asparagaceae
  1. 1

    Conidia consistently broadly cylindrical or subcylindrical .......................................................... 2

    Conidia acicular-filiform to obclavate-cylindrical .......................................................... 5

  2. 2(1)

    Conidia subcylindrical, apex sometimes distinctly swollen; on Agave .................................... C. agavicola

    Conidia never swollen at the apex; on other hosts .......................................................... 3

  3. 3(2)

    Leaf spots indistinct; conidiophores short, 10–35 µm; conidia 15–70 × 3–5.5 µm, (1−)3(−5)-septate; on Yucca .......................................................... C. floricola

    Leaf spots distinct; conidiophores much longer, to 350 µm; conidia pluriseptate (1–18); on other hosts ......................... 4

  4. 4(3)

    Conidiophores very long, to 350 µm; conidia consistently cylindrical, 50–80 × 4–5.5 µm, 4–7-septate; on Furcraea, South America ..................................................... C. fourcroyae

    Conidiophores shorter, to 160 µm; conidia cylindrical to obclavate-cylindrical, 20–150 × 4–8 µm, 1–18-septate; on Maianthemum, northern hemisphere ................................. C. maianthemi

  5. 5(1)

    Conidia cylindrical to obclavate-cylindrical, rather broad, 20–150 × 4–8 µm; on Maianthemum ............ C. maianthemi

    Conidia at least partly acicular, i.e. acicular, filiform to obclavate-cylindrical; on other hosts ...................................... 6

  6. 6(5)

    Stromata well-developed, 20–60 µm diam; conidia acicular-filiform to slightly obclavate; on Hosta .............................. C. hostae

    Stromata lacking or small, 10–30 µm diam, if larger conidia acicular to obclavate-cylindrical; on Asparagus or Polygonatum .......................................................... 7

  7. 7(6)

    Leaf spots circular to oblong, between veins, 1–7 mm diam or to 30 mm long, pale brown to dingy grey, with dark purple margin; stromata lacking or small, 10–30 µm diam; on Polygonatum .......................................................... C. chinensis

    On cladodes and branches, forming small subcircular, oval to elliptical spots, 0.5–4 mm diam, pale brownish, greyish brown to dingy grey, margin brown, reddish brown; stromata always present, 10–40 µm diam or oblong, to 100 µm; on Asparagus .................................. C. asparagi

Cercospora species on Asparagaceae

Cercospora agavicola Ayala-Escobar, Mycotaxon 93: 117 (2005).

(Fig. 51)

Fig. 51
figure51

Cercospora agavicola (HAL 1839 F). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Groenewald et al. (2013: 144).

Illustration: Ayala-Escobar et al. (2005: 119, figs 2–14).

Description: Leaf spots variable in size and shape, irregular necroses, dingy grey. Caespituli punctiform to pustulate, scattered to dense, dark to blackish brown, later grey-brown to greyish white by abundant sporulation. Mycelium internal; hyphae solitary or forming lax to dense ropes or planate aggregations of swollen hyphal cells of to 15 µm diam. Stromata well-developed, immersed, often somewhat erumpent, 20–150 µm diam or confluent and larger, composed of swollen hyphal cells, 3–8 µm diam, brown.

Conidiophores in small to moderately large lax fascicles, erect, divergent, subcylindrical-filiform to flexuous-sinuous, barely geniculate, unbranched, occasionally with constrictions and swellings, 20–100 × 3–6(−7) µm (to 200 µm long and strongly branched under high humidity in moist chambers), pluriseptate, at first subhyaline, later pale olivaceous, olivaceous-brown or pale brown, often paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, 10–40 µm long, 1–3 conidiogenous loci, thickened and darkened, 2–3 µm diam, terminal and lateral. Conidia solitary, subcylindrical, (35−)40–100(−120) × 3–5.5 µm, (0−)3–8-septate, hyaline, thin-walled, smooth, apex obtuse, sometimes with distinctly swollen tip, base truncate to somewhat obconically truncate, 1.5–2.5 µm wide, hila somewhat thickened and darkened.

In vitro: Colonies on all media green to greyish, reaching 20 mm diam within 3 weeks. Sporulation only observed on AA medium after 21 d. Conidiophores dense, 231–960 µm long, pluriseptate, with a single terminal conidiogenous locus or to five lateral slightly protruding loci. Conidia solitary, cylindrical, straight, 25–120 µm long, 2–8-septate, hyaline, apical cell often swollen, subglobose or clavate. Spermogonia formed on OA, exuding masses of hyaline, rod-shaped spermatia, 3–6 × 1–2 µm.

Holotype: Mexico: State of Guanajuto: Penjamo, on Agave tequilana, Asparagaceae, Jan. 2003, V. Ayala-Escobar & Ma. de Jesús Yáñez-Morales (CHAP 166). Isotype: HAL 1839 F. Ex-type culture: CBS 117292.

Host range and distribution: Only known from the type collection.

Notes: The position of C. agavicola as genetically distinct species has been proven by means of molecular sequence analyses (Groenewald et al. (2013).

Cercospora apii s. lat.

Notes: Two collections belonging to the C. apii complex have been examined on Dracaena spp. (Japan, intercepted at Hawaii, on Dracaena fragrans [deremenensis], 7 Aug. 1974, Kunisaki et al., BPI 435828; Mexico, Tamazunchale, on D. marginata, 11 June 1982, J. Okamura, BPI 435829). The sample on Dracaena fragrans is characterised by having loosely fasciculate, very long, straight, cylindrical, usually non-geniculate conidiophores, to 250 × 3–7 µm, pale to medium brown, paler towards the tip, pluriseptate, conidiogenous loci 2.5–4 µm diam. The conidia are formed singly, acicular, to 160 × 3–4.5 µm, hyaline pluriseptate. The Mexican collection on D. marginata differs in having shorter conidiophores, to 80 × 3–8 µm, pale to medium olivaceous-brown, with loci 2–3.5 µm diam, and narrower conidia, 2–4 µm.

Cercospora asparagi Sacc., Michelia 1: 88 (1877).

(Fig. 52)

Fig. 52
figure52

Cercospora asparagi (HAL, Sacc., Mycoth. Ven. 1052). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Synonyms: Cercospora caulicola G. Winter, J. Mycol. 1: 125 (1885) [syntypes: USA: Missouri: Perryville, on Asparagus officinaiis, Sep. 1883, C. H. Demetrio, Rabenh., Fungi Eur. Exs. no. 3591 (numerous herbaria, e.g. B, G, S)].

Cercosporina asparagicola Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 20: 424 (1910) [holotype: Argentina: La Plata, botanical garden, on Asparagus officinaiis, C. Spegazzini (LPS 4966); K(M) IMI 247001 (slide)].

Cercospora asparagicola (Speg.) Vassiljevsky, in Vassiljevsky & Karakulin, Fungi Imperfecti Parasitici (Hyphomycetes) 1: 296 (1937).

Literature: Saccardo (1886: 477), Lindau (1910: 89), Vassiljevsky & Karakulin (1937: 296), Chupp (1954: 343), Vasudeva (1963: 43), Katsuki (1965: 42), Ellis (1976: 270), Sutton & Pons (1980: 203), Hsieh & Goh (1990: 208), Braun & Mel’nik (1997: 40), Braun (2000: 74), Crous & Braun (2003: 68), Phengsintham et al. (2013b: 50), Guo et al. (2005: 174), Kamal (2010: 18–19), Han et al. (2013).

Illustrations: Ellis (1976: 271, fig. 205), Hsieh & Goh (1990: 209, fig. 161), Guo et al. (2005: 175, fig. 122), Phengsintham et al. (2013b:51, fig. 10–11).

Exsiccatae: Sacc., Mycoth. Ven. 1052. Rabenh., Fungi Eur. Exs. 3591.

Description: On cladodes and branches, forming small subcircular, oval to elliptical spots, 0.5–4 mm diam, pale brownish, greyish brown to dingy grey, margin brown, reddish brown. Caespituli amphigenous, punctiform, dark. Mycelium internal. Stromata small to well-developed, 10–40 µm diam, substomatal to immersed, sometimes oblong, to 100 µm in length, brown to dark brown, composed of swollen hyphal cells, 3–13 µm diam, rounded to angular-irregular in outline. Conidiophores in small to moderately large fascicles, loose to rather dense, arising from stromata, through stomata, erect, straight, subcylindrical to somewhat geniculate-sinuous above, unbranched, 10–170 × 4–8 µm, continuous to pluriseptate, pale to dark olivaceous-brown or medium brown, paler towards the tip, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal, occasionally conidiophores reduced to conidiogenous cells, 10–30 µm long, conidiogenous loci conspicuous, thickened and darkened, 1.5–3 µm diam. Conidia solitary, acicular to somewhat obclavate-subcylindrical, straight to somewhat curved, 35–150 × 2.5–5 µm, 3–12-septate, hyaline, thin-walled, smooth, apex subobtuse to subacute, base truncate to slightly attenuated, 2–3 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank MBT178138): Italy: Treviso, on Asparagus officinalis, Sep. 1875 (PAD). Isolectotypes: Sacc., Mycoth. Ven. 1052 (e.g. BPI 433642, HAL, HBG).

Host range and distribution: On Asparagus (densiflorus [Protasparagus densiflorus], officinalis, setaceus [plumosus, Protasparagus setaceus]), Asparagaceae, Africa (Ghana, Kenya, Malawi, South Africa, Zimbabwe, Zambia), Asia (Brunei, Cambodia, China, India, Israel, Japan, Korea, Laos, Malaysia, Nepal, Pakistan, Taiwan, Thailand), Europe (Italy, Russia, Ukraine), North America (USA, Arkansas, California, Delaware, Florida, Georgia, Illinois, Maryland, North Carolina, Nebraska, New Mexico, Oklahoma, South Carolina, Texas, Virginia), Oceania (Hawaii, Solomon Islands), Central and South America (Argentina, Brazil, Colombia, Honduras, Venezuela), West Indies (Cuba).

Notes: A true Cercospora s. str. pertaining to the C. apii s. lat. complex.

Cercospora chinensis F.L. Tai, Bull. Chin. Bot. Soc. 2: 49 (1936).

(Similar to Fig. 1)

Literature: Chupp (1954: 343), Crous & Braun (2003: 119), Guo et al. (2005: 175), Groenewald et al. (2013: 150).

Illustration: Guo et al. (2005: 176, fig. 123).

Description: Leaf spots amphigenous, circular to oblong, between veins, 1–7 mm diam or to 30 mm long, pale brown to dingy grey, with dark purple margin. Caespituli amphigenous, dark, punctiform. Mycelium internal. Stromata almost lacking to subglobose, 10–30 µm diam, substomatal or immersed, dark brown. Conidiophores in small to moderately large fascicles, 2–18, loose to dense, arising from internal hyhae or stromata, through stomata or erumpent, erect, straight, subcylindrical or attenuated towards the tip, geniculate, unbranched or rarely branched, 20–150 × 4–6 µm, pluriseptate, medium to dark brown, paler towards the tip, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal or intercalary, 10–30 µm long, conidiogenous loci thickened and darkened, 2.5–3 µm diam. Conidia solitary, acicular to somewhat obclavate-cylindrical, straight to slightly curved, (15−)30–220(−300) × 2–5 µm, 2–16-septate, hyaline, thin-walled, smooth, apex subacute, base truncate to obconically truncate, about 2–3.5 µm wide, hila somewhat thickened and darkened.

Holotype: China: Beijing, on Polygonatum officinale, Sep. 1933, L. Shih 1063 (HMAS 06992).

Host range and distribution: On Convallaria majalis, Polygonatum (humile, odoratum [oficinale], Polygonatum sp.), Asparagaceae, Asia (China, Korea).

Notes: ‘Sci. Rep. Natl. Tsing Hua Univ., Ser. B, Biol. Sci., 2: 428. 1937’ was given as the literature citation in Chupp (1954), but it actually appeared in a paper the previous year. This species belongs to the Cercospora apii complex. The phylogenetic position of this species has recently been examined by Groenewald et al. (2013).

Cercospora floricola Heald & F.A. Wolf, Mycologia 3: 17 (1911).

(Fig. 53)

Fig. 53
figure53

Cercospora floricola (BPI 436450). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Literature: Saccardo (1931: 883), Vassiljevsky & Karakulin (1937: 297), Chupp (1954: 346), Crous & Braun (2003: 187).

Description: Leaf spots indistinct. Caespituli formed on scapes, flower pedicels and outer portions of the perianth, effuse, olivaceous to blackish. Mycelium internal. Stromata formed as loose stromatic aggregations of swollen hyphal cells, 20–60 µm diam or confluent and larger, forming irregular layers, immersed, brown. Conidiophores in dense to very dense fascicles, arising from stromata, forming layers, erect, straight to slightly curved, subcylindrical, barely to slightly geniculate-sinuous, unbranched, 10–35 × 4–6.5 µm, often uniformly short and aseptate or somewhat longer and 0–2-septate, pale or very pale olivaceous-brown, thin-walled, smooth; conidiophores reduced to conidiogenous cells or conidiogenous cells integrated, terminal, 10–25 µm long, conidiogenous loci conspicuous, thickened and darkened, (1.5−)2(−2.5) µm diam. Conidia solitary, broadly ellipsoid-ovoid, cylindrical or somewhat obclavate, 15–70 × 3–6 µm, 1–4(−5)-septate, hyaline or subhyaline, thin-walled, smooth, apex obtuse, mostly broadly rounded, base truncate to short obconically truncate, (1.5−)2(−2.5) µm wide, hila somewhat thickened and darkened.

Lectotype (designated here, MycoBank MBT178151): USA: Texas: Austin, on Yucca rupicola, 19 June 1909, F. D. Heald &F. A. Wolf, no. 1438 (BPI 436450). Isolectotypes: BPI 436451, CUP 3915.

Host range and distribution: On Yucca rupicola, Asparagaceae, North America (USA, Texas).

Notes: This species is characterised by having passalora-like conidia, broadly ellipsoid-ovoid to cylindrical or somewhat obclavate and 1–4-septate, but the conidia are colourless or almost so and the conidiogenous loci agree with those of Cercospora s. str. species. Results of molecular sequence analyses showed that such species, as far as the conidia are hyaline, belong in Cercospora s. str.

Cercospora fourcroyae Obreg.-Bot., Caldasia 3: 49 (1941).

(Fig. 54)

Fig. 54
figure54

Cercospora fourcroyae (CUP 39835). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 37), Crous & Braun (2003: 188).

Description: Leaf spots amphigenous, subcircular to elliptical, 5–35 mm diam, brownish or zonate, with altering zones of green and brown, centre grey and often darkened by the fungal colonies. Caespituli amphigenous, punctiform, scattered to dense, dark brown or greyish brown by abundant conidiation. Mycelium internal. Stromata large, immersed, about 40–125 µm diam, dark brown, loosely woven masses to dense stromatic hyphal aggregations composed of swollen hyphal cells, 3–6 µm diam, circular to angular in outline, mostly formed as textura angulata. Conidiophores in large fascicles, arising from stromata, loose to moderately dense, erect, straight to strongly curved-sinuous, but not or only slighty geniculate, rarely forming an almost complete circle, unbranched, length variable, (15−)30–200(−350) × 3–9 µm, plainly pluriseptate, sometimes constricted at the septa, width irregular, pale to medium brown or olivaceous to yellowish brown, wall thin to slightly thickened, to 1 µm wide, smooth; conidiogenous loci conspicuous, at bluntly rounded tips or lateral, 2–3.5 µm diam, thickened and darkened. Conidia solitary, cylindrical-obclavate, straight to curved, 40–80 × 3.5–6 µm, 3–7-septate, hyaline or subhyaline, thin-walled, smooth, apex obtuse, base truncate to obconically truncate, 2–3 µm wide, hila thickened and darkened. Holotype: Colombia: Santander, San José de Suaita, on Furcraea foetida [gigantea], 5 Dec. 1937, R. Barrios-Ferrer (CUP 39835).

Host range and distribution: On Furcraea (foetida, Furcraea sp.), Asparagaceae, South America (Brazil, Colombia).

Notes: This species belongs in Cercospora s. str., but it is quite distinct from C. apii s. lat. by its very long conidiophores and obclavate-cylindrical conidia with blunt tips. This species was recorded on Alstroemeria sp., Alstroemeriaceae, in Brazil (Batista et al. 1965: 16, fig. 7; Mendes et al. 1998). The corresponding material has been re-examined (on Alstroemeria sp., Brazil, D.F., 10 May 1963, E.P. Heringer, IMUR 34448). This sample proved to belong to a species of Cercospora, but it was too poorly developed for a final conclusion and detailed description. As Alstroemeria pertains to another plant family, it remains doubtful if this collection has been correctly identified.

Cercospora hostae Katsuki, Ann. Phytopathol. Soc. Japan 20: 72 (1955).

(Fig. 55)

Fig. 55
figure55

Cercospora hostae (based on Shin & Kim 2001: 72, fig. 27). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Synonym: Cercospora hostae Hori, Ann. Phytopathol. Soc. Japan 1(4): 66 (1921); nom. inval. (nom. nud.).

Literature: Chupp (1954: 347), Katsuki (1965: 42), Kim & Shin (1999b), Shin & Kim (2001: 71), Guo & Xu (2002), Crous & Braun (2003: 219), Guo et al. (2005: 177).

Illustrations: Shin & Kim (2001: 72, fig. 27), Guo et al. (2005: 178, fig. 125).

Description: Leaf spots amphigenous, scattered to confluent, circular or subcircular to somewhat irregular, 1–10 mm diam, to 20 mm when confluent, at first brown to greyish brown on the upper leaf surface, later greyish brown with narrow reddish brown border, finally centre turning greyish white to white with reddish brown to dark brown border, and pale yellow halo on both sides of the leaf. Caespituli amphigenous. Mycelium internal; hyphae branched, 2.5–4 µm wide, septate, hyaline. Stromata small to large, subglobose to globose, immersed or substomatal, 20–60 µm diam, dark brown to blackish brown, composed of brown swollen hyphal cells. Conidiophores in loose to dense fascicles, 3–20, arising from stromata, through stomata or erumpent, erect, straight to somewhat curved, subcylindrical, barely to slightly geniculate, unbranched, 10–245 × 3.5–6.5 µm, width uniform, 0–8-septate, olivaceous-brown, paler towards the tip, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal, conidiogenous loci conspicuous, thickened and darkened, 2–3 µm diam. Conidia solitary, acicular-filiform to obclavate-subcylindrical, 25–250 × 2.5–5 µm, 5–14-septate, hyaline, thin-walled, smooth, apex subobtuse or subacute, base truncate to somewhat obconically truncate, about 2–3 µm wide, hila thickened and darkened.

Holotype: Japan: Tokyo, Murayama, on Hosta sp., 12 Oct. 1952, E. Kurosawa (not preserved).

Host range and distribution: On Hosta (plantaginea, ventricosa, Hosta sp.), Asparagaceae, Asia (China, Japan, Korea).

Notes: A true Cercospora s. str. close to or belonging to the C. apii s. lat. complex. Hori (1921) listed this fungus as new species from Mainland China, but without any descriptions or diagnosis (nom. nud.). The type collection cited in Katsuki (1955) is probably not maintained.

Cercospora maianthemi Fuckel, Fungi Rhen. Exs., Suppl., Fasc. 2, no. 1631 (1866); as “majanthemi”.

(Fig. 56)

Fig. 56
figure56

Cercospora maianthemi (HAL, Fuckel, Fungi Rhen Exs. Suppl. 1631). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Saccardo (1886: 476), Lindau (1910: 89), Vassiljevsky & Karakulin (1937: 298), Chupp (1954: 348), Katsuki (1965: 43), Ellis (1976: 272), Braun & Mel’nik (1997: 71), Crous & Braun (2003: 263).

Illustrations: Vassiljevsky & Karakulin (1937: 298, fig. 23), Ellis (1976: 272, fig.206 B).

Exsiccatae: Allesch. & Schnabl, Fungi Bav. 499. Barthol., Fungi Columb. 4110 (as “C. subsanguinea”). Bucholtz & Bondartsev, Fungi Ross Exs., Ser. B, 698. Erikss., Fungi Paras. Scand. Exs. 191. Fuckel, Fungi Rhen Exs. Suppl. 1631. Kabát & Bubák, Fungi Imperf. Exs. 50, 599. Kellerm., Ohio Fungi 123. Krieger, Fungi Saxon. Exs. 448. Petr., Fl. Bohem. Morav. Exs. Pilze 1213. Petr., Fungi Polon. Exs. 22. Petr., Mycoth. Carp. 7. Petr., Mycoth. Gen. 1718. Sacc., Mycoth. Ven. 1565. Poelt & Scheuer, Reliqu. Petrak. 2799. Rabenh., Fungi Eur. Exs. 2073, 3590. Scheuer, Mycoth. Graec. 251. Siemaszko, Fungi Bialowiez. Exs. 98. Syd., Mycoth. Germ. 1776, 1777, 2449. Vestergr., Micromyc. Rar. Sel. Praec. Scand. 237. Weese, Eumyc. Sel. Exs. 499. Wirtgen, Florae Rhenanae, Ed. Nov., II. Abt., 46.

Description: Leaf spots large, at first circular, later enlarged, covering large leaf segments or almost entire leaves, brownish with yellowish border. Caespituli amphigenous, mainly hypophyllous, punctiform to almost pustulate, scattered, effuse to dense, velvety, sooty, olivaceous-brown, dark brown to blackish. Mycelium internal. Stromata substomatal to immersed, 30–80 µm diam, medium to medium dark brown, sometimes almost blackish brown, composed of swollen hyphal cells, 3–10 µm diam. Conidiophores few to numerous in divergent to dense fascicles, straight, subcylindrical to distinctly sinuous or sinuous-geniculate, unbranched, 50–160 × 4.5–7 µm, aseptate to pluriseptate, pale olivaceous to olivaceous-brown, tips paler, darker in mass, wall thin to somewhat thickened, smooth; conidiogenous cells integrated, terminal and intercalary or conidiophores reduced to conidiogenous cells, 10–60 µm long, conidiogenous loci conspicuous, thickened and darkened, 2–4 µm diam. Conidia solitary, cylindrical to obclavate-cylindrical, short conidia sometimes broadly ellipsoid-obovoid to short cylindrical, 20–150 × 4–8 µm, 1–15-septa, distance between septa 5–20 µm, at first hyaline or subhyaline, later greenish to very pale olivaceous, olivaceous-brown or brownish with age, apex obtuse, rounded, base short obconically truncate, 2–3 µm wide, hila somewhat thickened and darkened.

Lectotype (designated here, MycoBank MBT178139): France: Alsace: Jura, Col Amic, on Maianthemum bifolium, summer 1866, Morthier [Fuckel, Fungi Rhen. Exs. Suppl. 1631] (BPI 438038). Isolectotypes: Fuckel, Fungi Rhen. Exs. Suppl. 1631 (e.g. FH, G, HAL).

Host range and distribution: On Maianthemum (bifolium, canadense), Asparagaceae, Asia (Japan, Russia), Europe (Austria, Belgium, Czech Republic, Denmark, Estonia, Finland, France, Germany, Hungary, Italy, Latvia, Lithuania, Netherlands, Poland, Romania, Russia, Sweden, Switzerland, Ukraine), North America (USA, Michigan, Ohio, Pennsylvania, New York, West Virginia, Wisconsin).

Notes: The identity of North American collections on Maianthemum canadense has been established. American collections coincide with European samples, at least morphologically. Braun & Mel’nik (1997) examined type material of this species and numerous other samples, and confirmed that the conidia are at first hyaline but become somewhat pigmented with age, i.e. they are rather passalora-like. However, the conidiogenous loci are large, thickened and rather cercospora-like. Therefore, they preferred to maintain this species in Cercospora s. str., at least tentatively. Results of molecular sequence analyses are urgently needed to determine the generic affinity of this species.

Doubtful, excluded and insufficiently known species

Cercospora montenegrina Bubák, Bot. Közlem. 14: 82

(1915).

Synonym: Cercosporina montenegrina (Bubak) Sacc., Syll. Fung. 25: 909 (1931).

Literature: Vassiljevsky & Karakulin (1937: 296), Chupp (1954: 350), Crous & Braun (2003: 281).

Description: Leaf spots linear, 2–3 mm long, dark to black. Caespituli amphigenous. Mycelium internal. Stromata large, pustulate, black. Conidiophores in very dense fascicles, arising from stromata, erect, straight to sinuous, narrower towards the tip, neither geniculate nor branched, 20–40 × 5.5–7 µm, septate, pale or very pale olivaceous, paler towards the tip. Conidia solitary, cylindrical-filiform or slightly attenuated towards the tip, straight to curved, 35–55 × 3–3.5 µm, 3–5-septate, hyaline, thin-walled, apex subobtuse, base obconic.

Holotype: Montenegro: Piva Monastery, on Anthericum ramosum, Asparagaceae, Jul. 1905, Rohlena (BPI 438578).

Host range and distribution: Only known from the type collection.

Notes: This collection is in poor condition and devoid of any mature conidiophores and conidia. The generic affinity of this species remains unclear. New collections are necessary.

Cercospora smilacinae Ellis & Everh., Bull. Torrey Bot. Club. 27: 577 (1900).

Literature: Saccardo (1902: 1073), Vassiljevsky & Karakulin (1937: 298), Chupp (1954: 353).

Holotype: USA: Idaho: Latah Co., on Maianthemum stellatum [Smilacina sessilifolia], Asparagaceae, 1 Jul. 1899, R. M. Horner 1293 (NY 838617).

Notes: This is a doubtful species. No sporulation has been found in the type collection, which has been annotated by C. G. Shaw (10 Apr. 1967) who stated that he only found Cylindrosporium smilacinae Ellis & Everh., and that “C. smilacinae is, therefore, probably best considered a synonym of the above”. This species was also reported on Smilax from Brazil by Mendes et al. (1998), which was undoubtedly based on a misidentification.

Passalora

Key to Passalora species on Asparagaceae
  1. 1

    Conidiophores verruculose; conidia medium brown and finely verruculose; on Cordyline ..................... P. cordylines

    Conidiophores and conidia smooth or almost so; conidia subhyaline to pale olivaceous or pale brown; on other hosts .................................................... 2

  2. 2(1)

    Stromata very large, to 180 µm diam; conidia olivaceous to pale brown; on Yucca ..................... P.yuccae-gloriosae

    Stromata lacking or smaller, 20–70 µm diam .................................................... 3

  3. 3(2)

    Stromata lacking or almost so; conidia 18–100 × 2.5–5 µm, brownish; on Ophiopogon japonica ....................... P. togashiana

    Stromata well-developed, 20–70 µm diam; conidia 4–8 µm wide, subhyaline to pale olivaceous; on Polygonatum ................................................................ 4

  4. 4(3)

    Conidia 50–85 × 4–6 µm wide, 4–7-septate; on Polygonatum spp., Europe, Caucasus, West Asia ................................................................ P. polygonati

    Conidia broader, 6–8 µm, and only 1–4-septate; on Polygonatum odoratum, Japan, endemic ................................................................ P. polygonati-maximoviczii

Passalora species on Asparagaceae

Passalora cordylines (Henn.) Crous & M.P.S. Câmara, Mycotaxon 68: 302 (1998).

(Fig. 57)

Fig. 57
figure57

Passalora cordylines (BPI 435225). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bars = 10 µm.

Basionym: Cercospora cordylines Henn., Hedwigia 41: 117 (1902).

Synonym: Cercospora cordylines Speg., Revista Mus. La Plata, Secc. Bot., 15: 45 (1908), nom. illeg. (ICN, Art. 53.1) [lectotype (designated here, MycoBank MBT178140): Brazil: São Paulo, on living leaves of Corydyline spectabilis [dracaenoides], Sep 1905, A. Usteri (CUP 39512)].

Literature: Saccardo (1906: 611), Chupp (1954: 344), Crous & Braun (2003: 138).

Illustration: Crous & Câmara (1998: 301, fig. 2).

Description: Leaf spots amphigenous, circular, 2–8 mm diam, brown to dark red-brown, with a dark brown to black border, frequently confluent, forming larger patches. Caespituli amphigenous, punctiform, dark. Mycelium internal; hyphae branched, septate, light brown, 3–4 µm wide, thin-walled, smooth to faintly rough. Stromata immersed, brown, large, to 100 µm diam. Conidiophores in dense, large fascicles, arising from stromata, straight, subcylindrical to geniculate-sinuous, unbranched, 30–75 × 3–5 µm, 1–7-septate, medium brown, smooth to verruculose, wall thin to somewhat thickened; conidiogenous cells integrated, terminal, 5–30 µm long, conidiogenous loci conspicuous, somewhat thickened and darkened, 1–2 µm wide. Conidia solitary, subcylindrical to obclavate, (25−)45–75 × (4−)5–6 µm, (1−)3–8(−10) µm, pale medium brown, thin-walled, finely verruculose, apex obtuse, base short obconically truncate, 1.5–2 µm wide, hila slighty thickened and darkened.

Lectotype (designated here, MycoBank MBT178141): Brazil: São Paulo, Botanical Garden, on Corydyline spectabilis, Mar. 1901, A. Puttemans [Fungi S. Paulensis 286] (BPI 435225).

Host range and distribution: On Cordyline (fruticosa [terminalis], spectabilis [dracaenoides], Cordyline sp.), Asparagaceae, South America (Brazil), North America (USA, Florida), Oceania (Hawaii).

Notes: Chupp (1954) described conidia to 175 µm long, which have not been found during the course of the re-examination of type material. Type material (Puttemans 286) was previously deposited at B as well, but in 1997 it was sent on loan to Brazil and has been lost.

Passalora polygonati (Rostr.) U. Braun & Crous, in Crous & Braun, Mycosphaerella and Anam. : 331 (2003).

(Fig. 58)

Fig. 58
figure58

Passalora polygonati (C, holotype). A. Conidiophore fascicle. B. Conidiophore. C. Conidiophore tips. D. Conidia. Bar = 10 µm.

Basionym: Cercospora polygonati Rostr., Bot. Tidsskr. 26: 314 (1905).

Literature: Saccardo (1913: 1413), Lindau (1910: 89), Vassiljevsky & Karakulin (1937: 299), Chupp (1954: 352), Braun & Mel’nik (1997: 81), Crous & Braun (2003: 331).

Description: Leaf spots amphigenous, subcircular, elliptical to somewhat irregular, 1–8 mm diam, centre yellowish, ochraceous or brownish, later becoming pale, greyish white with narrow to broad dark margin, reddish brown to purplish, finally very dark. Caespituli hypophyllous, punctiform, scattered to aggregated, dark. Mycelium internal. Stromata substomatal, about 20–50 µm diam, dark olivaceous-brown, composed of swollen hyphal cells, 3–8 µm diam. Conidiophores in small to usually large fascicles, divergent to dense, arising from stromata, through stomata, erect, straight to usually strongly geniculate-sinuous, unbranched, 25–140 × 4–8 µm, (0−)1–6-septate, brown, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal and intercalary, about 10–50 µm long, conidiogenous loci conspicuous, thickened and darkened, about 2–2.5 µm diam. Conidia solitary, subcylindrical, 50–85 × 4–6 µm, 4–7-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex obtuse, broadly rounded, base rounded to short obconically truncate, 2–3 µm wide, hila somewhat thickened and darkened.

Holotype: Denmark: Jütland: Vendsyssel-Thy, Baggesvogn, on Polygonatum multiflorum, 1 Sep. 1902, Rostrup (C).

Host range and distribution: On Polygonatum (humile, multiforum, orientale), Asparagaceae, Caucasus (Azerbaijan), Asia (Russia), Europe (Denmark, Russia).

Notes: Not a Cercospora as already emphasized by Chupp (1954). The conidiogenous loci are conspicuous and the conidia are subhyaline to pale olivaceous.

Passalora polygonati-maximoviczii Poonam Srivast., J. Living World 1: 117 (1994), nom. nov. (as “(Katsuki) Poonam Srivast.”).

(Fig. 59)

Fig. 59
figure59

Passalora polygonati-maximoviczii (TNS-F-60888). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Basionym: Cercosporidium polygonati-maximoviczii Katsuki, Trans. Mycol. Soc. Japan 16: 12 (1975), nom. superfl. (ICN, Art. 52.1).

Synonyms: Cercospora polygonati-maximowicziiTogashi, Jap. J. Bot. 2: 75 (1924) [syntypes: TNS-F-243801–243803].

Passalora polygonati-maximoviczii (Togashi) U. Braun & Crous, Mycosphaerella and Anam.: 331 (2003), nom. illeg. (ICN, Art. 53.1).

Literature: Vassiljevsky & Karakulin (1937: 298), Chupp (1954: 353), Katsuki (1965: 43), Braun & Crous (2007: 60).

Illustration: Katsuki & Kobayashi (1975: 12, fig. 8).

Description: Leaf spots amphigenous, at first formed as small brown specks, later almost elliptical to irregular, often angular and vein-limited, usually elongated, 5–25 mm long and 1–5 mm wide, on the upper leaf surface pale to greyish brown with purplish brown margin, below dull to greyish brown, sometimes uniformly blackish on both sides. Caespituli hypophyllous, punctiform, effuse to dense. Mycelium internal. Stromata substomatal to immersed, subepidermal, dark brown to blackish, subglobose, 30–70 µm diam. Conidiophores in large, divergent to dense fascicles, arising from stromata, through stomata or erumpent, erect, straight to curved, geniculate-sinuous, unbranched, 40–120 × 6–7 µm, 0–4-septate, brown, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, sympodially proliferating, conidiogenous loci conspicuous, about 2–2.5 µm diam, terminal and on shoulders caused by sympodial proliferation, somewhat thickened and darkened. Conidia solitary, cylindrical to somewhat obclavate-cylindrical, straight to somewhat curved, 45–90 × 6–8 µm, 1–4-septate, pale olivaceous, thin-walled, smooth, apex obtuse, base short obconically truncate to rounded, 2–3 µm wide, hila somewhat thickened and darkened.

Holotype: Japan: Niigata Pref.: Kitauonuma-gun, Horinouchi, on Polygonatum odotarum var. pluriflorum, 8 Jul. 1972, S. Katsuki SK 2635 (TNS-F-60888). Isotype: TNS-F-60889 (“SK 2636”).

Host range and distribution: On Polygonatum (odoratum var. maximoviczii [maximowiczii], odoratum var. pluriflorum, odoratum var. thunbergii [japonicum]) (Asparagaceae), Asia (Japan).

Notes: This species is morphologically close to P. polygonati, but differs in having 1–4-septate conidia. Katsuki & Kobayashi (1975) considered Cercospora polygonati-maximoviczii Togashi a “nom. seminud.” and cited it as a synonym of Cercosporidium polygonati-maximoviczii, but Togashi’s species was published before 1935 with a full English description and is valid. Katsuki’s species must be considered a superfluous name (nom. illeg.). Srivastava (1994) introduced the “new combitation” Passalora polygonati-maximoviczii with reference to the illegitimate name Cercosporidium polygonati-maximoviczii, published with a Latin description, as basionym. Hence, Srivastava’s name has to be considered a new name attributed to Srivastava as author.

Passalora togashiana (Katsuki & Urasawa) C. Nakash., comb. nov.

(Fig. 60)

Fig. 60
figure60

Passalora togashiana (TNS-F-243896). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

MycoBank MB809037

Basionym: Cercospora togashiana Katsuki & Urasawa, Ann. Phytopathol. Soc. Jap. 15: 144 (1951).

Literature: Chupp (1954: 355), Katsuki (1965: 43).

Description: Leaf spots indistinct or pale brown on the upper side. Caespituli amphigenous, mainly hyphophyllous, effuse, grey-olivaceous, vein-limited, covering an area of 4–6 × 0.3–0.5 mm. Stromata lacking or small, consisting of a few brown to reddish brown cells, substomatal to immersed. Conidiophores solitary or loosely fasciculate, arising from internal hyphae or the upper part of stromata, curved to sharply bent, sinuous, simple or branched, about 60–150 × 3.5–8 µm, pluriseptate, pale brown to reddish brown, darker in mass, wall thin to somewhat thickened, smooth; conidiogenous cells integrated, terminal and intercalary or conidiophores reduced to conidiogenous cells, proliferating sympodially, conidiogenous loci conspicuous, slightly thickened, 2–2.5 µm in diam. Conidia solitary, cylindrical to almost obclavate, straight to mostly curved, 18–100 × 2.5–5 µm, 4–8-septate, occasionally constricted at septa, pale to pale brown, smooth, apex obtuse, base rounded to or subtruncate, rounded, hila slightly thickened.

Holotype: Japan: Saitama Pref. : Minamisaitama-gun, Niiwa-mura, on Ophiopogon japonicus, 1 Jul. 1950, Y. Urasawa (TNS-F-243896).

Host range and distribution: On Ophiopogon japonicus, Asparagaceae, Asia (Japan).

Notes: Type material of Cercospora togashiana has been examined. Solitary conidiophores arising from internal hyphae and pigmented conidia suggest that this is not a member of Cercospora. Furthermore, this species has slightly thickened conidiogenous loci and conidial hila. Based on these morphological characters, the present fungus has to be assigned to the genus Passalora.

Passalora yuccae-gloriosae Crous, U. Braun & Alfenas, Mycotaxon 72: 190 (1999).

(Fig. 61)

Fig. 61
figure61

Passalora yuccae-gloriosae (IACM 3108). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Literature: Deighton (1973: 111–113, as Cercospora yuccae).

Illustrations: Deighton (1973: 112, fig. 3, as Cercospora yuccae), Crous et al. (1999: 189, fig. 13).

Description: Leaf spots amphigenous, subcircular to elliptical, 1–6 mm diam, pale brown, with a darker brown border, surrounded by a thin, red-brown margin. Caespituli amphigenous, dark brown, to 200 µm wide and 130 µm high. Mycelium internal; hyphae branched, septate, 2.5–4 µm wide, pale brown, smooth. Stromata immersed, large, to 180 µm wide and 60 µm high, dark brown. Conidiophores in dense, large fascicles, arising from stromata, straight, subcylindrical to geniculate-sinuous, unbranched, 15–65 × 3.5–6.5 µm, 2–5-septate, pale brown, thin-walled, verruculose; conidiogenous cells integrated, terminal, 15–22 µm long, sympodial, conidiogenous loci thickened and darkened, about 1.5–2.5 µm diam, usually non-protuberant. Conidia solitary, subcylindrical to obclavate-cylindrical, straight to somewhat curved, (20−)40–90(−160) × (3.5−)4–5.5(−6) µm, (1−)4–7(−13)-septate, olivaceous to pale brown, thin-walled, smooth or almost so (to very minutely rough-walled), apex obtuse, base obconically truncate, about 2 µm wide, hila somewhat thickened and darkened.

Holotype: Brazil: São Paulo, Estrada para Lindoia, Socorro, on Yucca gloriosa var. recurvifolia, 20 Sep. 1939, A. P. Viégas & O. Zagatto (IACM 3108).

Host range and distribution: On Yucca (gloriosa [recurvifolia], Yucca sp.), Asparagaceae, North America (USA, Georgia, North and South Carolina), South America (Brazil).

Notes: The name Cercospora yuccae has been confused and misinterpreted. A clarification of cercosporoid fungi on Yucca spp. was made in Crous et al. (1999).

Pseudocercospora

Key to Pseudocercospora species on Asparagaceae
  1. 1

    Conidiophores in sporodochial conidiomata, arising from large stromata, monopodial, percurrently proliferating, with conspicuous annellations; on Beaucarnea or Yucca ...................................... 2

    Conidiophores sympodial, annellations lacking; on Cordyline or Maianthemum ........................................ 4

  2. 2(1)

    Conidiophores with thin to somewhat thickened walls, verruculose to verrucose, 10–60 × 4–7 µm, 0–6-septate; conidia fainely verruculose, 25–125 × 4–7 µm; on Yucca .......................................................... see Scolecostigmina yuccae

    Conidiophores and conidia thin-walled and smooth; on Beaucarnea or Yucca (but then conidia only 2.5–5 µm wide) ........................................................... 3

  3. 3(2)

    Conidiophores rather broad, 5–25 × 4–6 µm; conidia 4–8 µm wide; on Beaucarnea .................... P. beaucarneae

    Conidiophores narrower, 10–50 × 2.5–5 µm; conidia 3–5(−5.5) µm wide; on Yucca.......................... P. concentrica

  4. 4(1)

    Stromata large, 50–80 µm diam; conidiophores 10–50 × 3–5.5 µm, 0–2-septate; on Cordyline ......................................................... P. cordylines

    Stromata smaller, 15–55 µm diam; conidiophores longer and wider, 95–235 × 4–7 µm, 7–16-septate; on Maianthemum ......................................................... P. maianthemi

Pseudocercospora species on Asparagaceae

Pseudocercospora beaucarneae (Pollack & D.F. Farr) U. Braun, comb. nov.

MycoBank MB809011

(Fig. 62)

Fig. 62
figure62

Pseudocercospora beaucarneae (BPI 71905). A. Conidioma. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Stigmina beaucarneae Pollack & D.F. Farr, Mycotaxon 8: 193 (1997).

Illustrations: Pollack & Farr (1979: 198, figs 2–7, 199, figs 8–13).

Description: Leaf spots oblong-ellipsoid, to 8 cm long and 6 mm wide, vein-limited, pale brown, margin reddish brown, centre later becoming straw-coloured. Conidiomata amphigenous, punctiform-pustulate to oblong, sporodochial to avervular, deeply immersed, at first circular or elliptical in outline, later elongated or confluent, forming oblong striae to 5 mm long, erumpent, rupturing the epidermis and cuticle. Mycelium internal. Stromata immersed, consisting of a prosenchymatous stratum of interwoven brown hyphae, surmounted by a saucer-shaped base of pseudoparenchyma consisting of thick-walled, dark brown cells. Conidiophores numerous, in large, dense conidiomata, arising from stromata, erumpent, simple or branched, erect, composed of to 11 cylindrical cells, 4–8 × 4 µm, forming columns, darker brown below and paler above, smooth or almost so; conidiogenous cells integrated, terminal, oblong cylindrical, ampulliform to subglobose, 5–25 × 4–8 µm, pale olivaceous, thin-walled, almost smooth to verruculose, percurrently proliferating, with visible annellations, with a single broad terminal locus, 2.5–5 µm wide, neither thickened nor darkened. Conidia solitary, obclavate-subcylindrical, straight to curved, 30–75 × 4–8 µm, 0–7-septate, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth or almost so, guttulate, apex obtuse, base truncate to short obconically truncate, 2.5–4 µm wide, hila neither thickened nor darkened.

Holotype: USA: Texas: Brownsville (plants from Mexico), on Beaucarnea recurvata, 2 June 1977, intercepted by J. Van Valkenburgh, no. 007449 (BPI 71905).

Host range and distribution: On Beaucarnea recurvata, Asparaceae, North America (Mexico; USA, Texas).

Notes: The genus Stigmina has been reduced to synonym with Pseudocercospora. Due to relatively thin-walled, smooth or almost smooth conidiophores and conidiophores with less conspicuous annellations, S. beaucarneae is rather close to the former genus Cercostigmina which is also synonymous with Pseudocercospora. On the other hand, the large, deeply immersed conidiomata are sporodochial to rather acervular. Among acervular genera, this species could easily be placed in Colletogloeum Petr. Stigmina beaucarneae resembles Colletogloeum sissoo (Syd.) B. Sutton (C. dalbergiae (S. Ahmad) Petr.), the type species of Colletogloeum which was demonstrated to cluster within the Mycosphaerellaceae near to the Pseudocercospora complex (Quaedvlieg et al. 2013).

Pseudocercospora concentrica (Cooke & Ellis) U. Braun & Crous, Mycol. Progr. 1: 22 (2002).

(Fig. 63)

Fig. 63
figure63

Pseudocercospora concentrica (NY 1102862). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora concentrica Cooke & Ellis, Grevillea 5: 90 (1877).

Synonyms: Stigmina concentrica (Cooke & Ellis) Deighton, Trans. Brit. Mycol. Soc. 61: 107 (1973).

Cercostigmina concentrica (Cooke & Ellis) U. Braun, Cryptog. Bot. 4: 108 (1993).

Cylindrosporium angustifoliae Ellis & Kellerm., J. Mycol. 2: 81 (1886) [holotype: USA: Kansas: Manhattan, on Yucca glauca, June 1886, Kellerman, no. 838 (NY 883634); K(M) IMI 119447 (slide), NY 883632, 883633 (slides)].

?Cylindrosporium yuccae Montemart., Rivista Patol. Veg. (Padua) 7: 231 (1915) [type: Italy: Milano, on Yucca sp. (probably not preserved)].

?Mycosphaerella deightonii M. Morelet, Bull. Soc. Sci. Nat. Archéol. Toulon & Var. 205: 9 (1973), sexual morph [holotype: USA: New Jersey: Gloucester Co., Newfield, on Yucca filamentosa, 1 June 1874, W. A. Kellerman 2150 (NY 838826); isotype: NY 1102861].

Literature: Saccardo (1886: 479), Vassiljevsky & Karakulin (1937: 297), Chupp (1954: 344), Sivanesan (1984: 248), Simay (1987), Kobayashi (2007).

Illustrations: Deighton (1973: 108, fig. 1, 110, fig. 2), Sivanesan (1984: 24, fig. 137).

Description: Leaf spots amphigenous, also at stems, elliptical, usually to 15 mm diam, occasionally to 40 × 10 mm, later often confluent, brownish to greyish brown, surrounded by a darker brown margin 1–2 mm wide, patches sometimes somewhat sunken with somewhat raised margin, spots less conspicuous on brown necrotic leaves. Caespituli amphigenous, punctiform to pustulate, scattered to dense, often in concentric zones, dark brown to blackish, often accompanied by spermogonia and mycosphaerella-like ascomata. Mycelium internal; hyphae branched, septate, subhyaline to olivaceous, occasionally darker, above all around stromata, 2–7 µm diam. Stromata deeply immersed, later erumpent, well-developed, large, 100–500 µm wide and 80–300 µm high, consisting of a basal prosenchymatous stratum, 25–40 µm thick, of interwoven hyphae surrounded by a palisade-like mass of dense, parallel, closely septate, branched hyphae, cells 6–14 µm long, pale olivaceous, thin-walled, smooth, giving rise to conidiophores at their distal ends. Conidiophores numerous, arising from stromata, loose to usually dense, forming sporodochial conidiomata, erect, straight to somewhat sinuous, but not geniculate, unbranched or only rarely branched, 10–50 × 2.5–5 µm, aseptate or only with few septa, pale olivaceous to very pale olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–30 µm long, with a single terminal locus, truncate to slighty convex, monoblastic, determinate to percurrent, with fine annellations. Conidia solitary, cylindrical, subcylindrical to obclavate-cylindrical, straight to curved, 20–75(−90) × 3–5(−5.5) µm, (0−)1–7-septate (in vitro conidia to 180 µm long and to 10−septate), pale olivaceous, thin-walled, smooth, apex obtuse to subacute, base subtruncate to short obconically truncate, 1.5–3 µm wide, hila unthickened, not darkened.

Lectotype (designated here, MycoBank MBT178142): USA: New Jersey: Newfield, on Yucca filamentosa, 1 June 1874, J. B. Ellis (K(M) 190710). Isolectotype: NY 1102862, K(M) IMI 119447 (slide).

Host range and distribution: On Yucca (aloifolia, brevifolia, filamentosa, glauca, gloriosa, guatemalensis, Yucca sp.), Asparagaceae, Caucasus (Georgia), Europe (Hungary, Italy, Russia, Switzerland), North America (Mexico; USA, Arizona, Connecticut, Georgia, Iowa, Illinois, Kansas, Mississippi, New Jersey, Oklahoma, Texas, Virginia), West Indies (Puerto Rico, Virgin Islands).

Notes: Deighton (1973) cited syntype material deposited at K as “holotype” and material deposited at NY as isotype, which was incorrect as in the original description a holotype designation is lacking and the cited collection comprises several duplicates. Morelet (1973) described Mycosphaerella deightonii as sexual morph of P. concentrica with the following brief description: ascomata 60–120 µm diam; asci 8-spored, 45–52 × 8.5–9 µm; ascospores 14.5–19 × 3.5–4 µm, hyaline, with a single medium septum. The genetic connection between M. deightonii and P. concentrica is, however, unproven and unclear. Aptroot (2006: 76 and 12, fig. 247) examined type material and emphasized that M. deightonii is morphologically indistinguishable from Planistromella acervata. Barr (1996) discussed M. deightonii in connection with P. acervata as well, but without final conclusion and with reference to smaller ascospores in M. deightonii. Crous (in Crous et al. 2011) described Mycosphaerella valgourgensis on Yucca sp. from France, which forms conidia of similar shape and size. However, this species is genetically distinct from Pseudocercospora concentrica (Mycosphaerella deightonii) and does not cluster within the Pseudocercospora s. str. clade and the Mycosphaerellaceae at all. Furthermore, its ascospores are wider and the conidia, only formed in vitro, are 45–150 µm long and hyaline, only becoming somewhat pigmented and verruculose with age. Records of “P. concentrica” from Japan have been established, also by means of molecular sequence analyses, and seem rather to belong to Mycosphaerella valgourgensis.

Pseudocercospora cordylines U. Braun, Schlechtendalia 1: 29 (1998).

(Fig. 64)

Fig. 64
figure64

Pseudocercospora cordylines (B 700014864). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Literature: Saccardo (1906: 611), Chupp (1954: 344).

Illustration: Braun (1998b: 27, fig. 6).

Description: Leaf spots amphigenous, subcircular, 2–5 mm diam, tan to dark reddish brown, margin narrow to moderately wide, dark brown to blackish. Caespituli amphigenous, punctiform, blackish. Mycelium internal. Stromata well-developed, 50–80 µm diam, dark brown to blackish, immersed to erumpent. Conidiophores numerous, in dense fascicles, arising from stromata, erect, straight, subcylindrical to somewhat flexuous, but barely geniculate-sinuous, unbranched, 10–50 × 3–5.5 µm, 0–2-septate, pale olivaceous to medium brown throughout, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–25 µm long, proliferation sympodial, conidiogenous loci inconspicuous or somewhat refractive, visible in front view as minute circle, but neither thickened nor darkened. Conidia solitary, obclavate-cylindrical, (15−)30–80(−100) × 4–6 µm, 2–8-septate, subhyaline to pale olivaceous-brown, thin-walled, smooth, apex usually obtuse, base obconically truncate to rounded, about 2–3 µm wide, hila neither thickened nor darkened, occasionally somewhat refractive.

Holotype: Brazil: São Paulo, Botanical Garden, on living leaves of Corydyline spectabilis [dracaenoides], 10 Aug. 1905, A. Usteri (B 700014864). Isotype: CUP 39513.

Host range and distribution: On Cordyline spectabilis, Asparagaceae, South America (Argentina, Brazil).

Notes: Type material of Cercospora cordylines Speg. (Argentina, Santa Catalina, Spegazzini and Brazil, São Paulo, Botanica Garden, Sep. 1905, Usteri 14) could not be traced at LPS. Therefore, Braun (1998) described Pseudocercospora cordylines as a new species with a collection from B as holotype, which is topotype material of C. cordylines, collected in Aug. 1905. However, syntype material of C. cordylines has since been found at CUP and proved to be Passalora cordylines.

Pseudocercospora maianthemi R. Kirschner, Mycol. Progr. 7: 24 (2008).

(Fig. 65)

Fig. 65
figure65

Pseudocercospora maianthemi (based on Kirschner & Piepenbring 2008: 26–27, figs 5–6). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Illustrations: Kirschner & Piepenbring (2008: 26–27, figs 5–6).

Description: Leaf spots amphigenous, pale to dark brown with irregular margin, confluent to streaks to 40 mm long and 5 mm wide. Caespituli hypophyllous. Mycelium internal, intercellular; hyphae branched, 2–5 µm wide, septate, pale brown, smooth. Stromata substomatal, irregularly shaped, 15–55 µm diam, pale to medium brown. Conidiophores formed singly or in fascicles of about 15, rarely to 30, arising from stromata, through stomata, erect, straight to somewhat geniculate-sinuous, unbranched, rather long, (95−)115–190(−235) × 4–7 µm, mostly about 5 µm wide, 7–16-septate, distance between septa 4–28 µm, dark brown, paler towards the tip, thin-walled, smooth, with percurrent rejuvenations, but not distinctly annellidic at the apex; conidiogenous cells integrated, terminal, pale to medium brown, proliferation sympodial, with 1–2 conidiogenous loci, inconspicuous to visible by being truncate, sometimes denticle-like, but neither thickened nor darkened, 2–3 µm diam. Conidia solitary, obclavate-cylindrical, straight to slightly curved, (20−)40–75(−90) × 4–5(−6) µm, tapering to 1.5–2 µm towards the tip, (2−)3–5(−7)-septate, very pale but distinctly brown, thin-walled, smooth, apex obtuse to subacute, base obconically truncate, 2–3 µm wide, neither thickened nor darkened.

Holotype: Panama: Chiriquí Province: Las Nubes, ca. 2.250 m alt., on Maianthemum gigas (or M. paniculatum), Asparagaceae, 25 Feb. 2003, R. Kirschner & J. A. Bernal Vega 1695 (FR). Isotype: PMA. Ex-type culture: CBS 103099.

Host range and distribution: Only known from the type collection.

Scolecostigmina

A single species.

Scolecostigmina yuccae (Cooke) Crous, U. Braun & Alfenas, Mycotaxon 72: 192 (1999).

(Fig. 66)

Fig. 66
figure66

Scolecostigmina yuccae (BPI 435122). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Basionym: Cercospora yuccae Cooke, Grevillea 7: 35 (1878).

Literature: Saccardo (1886: 479), Vassiljevsky & Karakulin (1937: 297), Chupp (1954: 355).

Illustration: Crous et al. (1999: 189, fig. 14).

Exsiccatae: Ravenel, Fungi Amer. Exs. 292

Description: Leaf spots amphigenous, oval to oblong, 8–30 × 2–6 mm, pale to medium brown on the upper surface, grey to medium brown below, margin lighter brown, somewhat raised, lesions occasionally bulging on upper surface. Caespituli amphigenous, punctiform, dark brown to blackish. Mycelium internal; hyphae branched, hyaline, thin-walled, 1.5–2.5 µm wide. Stromata immersed, well-developed, subglobose, 25–125 µm diam, dark brown. Conidiophores numerous, in dense fascicles, arising from stromata, erumpent, forming sporodochial conidiomata, erect, straight, subcylindrical-conical to somewhat curved, not geniculate, unbranched or occasionally branched at the very base, 10–60 × 4–7 µm, 0–6-septate, pale to dark brown, wall thin to somewhat thickened, fainly verruculose to verrucose; conidiogenous cells integrated, terminal or conidiophores aseptate, i.e. reduced to conidiogenous cells, 10–25 µm long, percurrently proliferating, with a single terminal locus, unthickened, not darkened. Conidia solitary, subcylindrical to obclavate, 25–125 × 4–7 µm, 1–6-septate, subhyaline, olivaceous to medium brown, thin-walled, fainly verruculose, apex obtuse, base short obconically truncate, hila unthickened, not darkened.

Lectotype (designated by Crous et al. 1999): USA: Georgia: Darien, on Yucca sp., H. W. Ravenel [Ravenel, Fungi Amer. Exs. 292] (BPI 435122). Isolectotypes: Ravenel, Fungi Amer. Exs. 292, e.g. CUP, K(M) IMI 119448.

Host range and distribution: On Yucca (gloriosa, Yucca sp.), Asparagaceae, South America (Brazil), North America (USA, Alabama, Georgia, Kansas).

Notes: The type species of Scolecostigmina has been sequenced and clusters distantly from the Pseudocercospora s. str. clade. On the other hand, various other former Scolecostigmina species cluster within this clade and belong to Pseudocercospora. Hence, the true generic affinity has to be proven on the basis of molecular sequence analyses. In the interim, we prefer to retain the present species in Scolecostigmina. “Cercospora yuccagena Cooke” is a typographical error for Cercospora yuccae Cooke (Chupp 1954: 356).

Zasmidium

Key to Zasmidium species on Asparagaceae
  1. 1

    Ramularia-like species; conidia catenate, cylindrical to fusiform, 10–55 × 2–6 µm, 0–3(−4)-septate, verruculose; conidiophores fasiculate as well as solitary, arising from superficial hyphae, 20–100 × 2–5 µm, subhyaline, greenish, yellowish, pale olivaceous to reddish brown, almost smooth; on Maianthemum .................... Z. subsanguinea

    Cercosporoid species; conidia formed singly, much longer, to 600 µm; on other hosts ....................... 2

  2. 2(1)

    Stromata lacking; conidiophores solitary, to 100 µm long, verruculose; conidia 170–600 × 7.5–10 µm, 5–55-septate, brown to dark brown,

    coarsely verrucose; on Agave .............................................. Z. agavicola

    Stromata developed, 10–35 µm diam; conidiophores fasciculate as well as solitary, 50–180 µm long, smooth; conidia much shorter and narrower, (8−)12–180 × 2–4 µm, 0–12-septate, subhyaline to pale olivaceous, verruculose; on Dracaena ...................................... Z. dracaenae

Zasmidium species on Asparagaceae

Zasmidium agavicola (R.F. Castañeda & L. Garcia) U. Braun & R.F. Castañeda, comb. nov. MycoBank MB809038

(Fig. 67)

Fig. 67
figure67

Zasmidium agavicola (INIFAT C87/431). A. Conidiophores. B. Conidia. Bar = 10 µm.

Basionym: Stenella agavicola R.F. Castañeda & L. García, Fungi Cubenses 3: 13 (1988).

Illustration: Castañeda (1988: fig. 25).

Description: Colonies amphigenous, forming circular, velutinous patches, olivaceous to dark brown. Mycelium internal and external; superficial hyphae branched, septate, 2–3 µm wide, brown, smooth to verruculose. Stroma lacking. Conidiophores solitary, arising from superficial hyphae, erect, straight to curved, geniculate, unbranched or branched, to 100 µm long and 3–4 µm wide at the base, septate, brown, verruculose, rarely smooth; conidiogenous cells integrated, usually terminal, with several conspicuous conidiogenous loci, truncate. Conidia solitary, cylindrical-filiform, very long, 170–600 × 7.5–10 µm, 5–55-septate, brown to dark brown, coarsely verrucose, apex obtuse, rounded, base short obconically truncate, hila somewhat thickened and darkened.

Holotype: Cuba: Prov. Pinar del Río: Soroa, on Agave sp., Asparagaceae, 24 Jan. 1987, M. Gonzáles Avila (INIFAT C87/431).

Host range and distribution: Only known from the type collection.

Zasmidium dracaenae (Hansf.) U. Braun & Crous, Schlechtendalia 20: 101 (2010).

(Fig. 68)

Fig. 68
figure68

Zasmidium dracaenae (K(M) IMI 53368). A. Hypha. B. hypha with conidiophore. C. Conidiophore fascicle. D. Conidiophore tips. E. Conidia. Bar = 10 µm.

Basionym: Cercospora dracaenae Hansf., Proc. Linn. Soc. London 1942–1943: 56 (1943).

Synonym: Stenella dracaenae (Hansf.) U. Braun & Crous, in Crous & Braun, Mycosphaerella and Anam.: 166 (2003).

Literature: Chupp (1954: 345).

Description: Leaf spots variable, irregularly shaped, small, 1–5 mm diam, to very large, to 80 mm diam, pale straw-coloured, yellowish, ochraceous, later brownish with white centre, margin narrow, darker, sometimes with yellowish halo and zonate. Caespituli amphigenous, mostly hypophyllous, punctiform to subeffuse, blackish. Mycelium internal and external; superficial hyphae branched, mostly at right angles, septate, 1–3 µm wide, hyaline to pale brown, thin-walled, verruculose. Stromata substomatal, 10–35 µm diam, occasionally erumpent, brown. Conidiophores in small to moderately, large, loose fascicles, 2–25, arising from stromata, through stomata, erect, cylindrical-filiform, setiform, only geniculate-sinuous near the apex, unbranched, rarely with short branchlets near the tip, about 50–180 × 2–4 µm, pluriseptate throughout, pale to medium brown or olivaceous, paler towards the tip, wall somewhat thickened below, thin above, smooth; conidiogenous cells integrated, terminal, about 10–30 µm long, conidiogenous loci conspicuous, thickened and darkened, about 1 µm diam. Conidia solitary, shape and size variable, short conidia narrowly ellipsoid-ovoid to obclavate-cylindrical, long conidia acicular, (8−)12–180 × 2–4 µm, 0–12-septate, hyaline to pale olivaceous, thin-walled, verruculose, apex obtuse to subacute, base truncate or short obconically truncate, 1–1.5 µm wide, hila slightly thickened and darkened.

Holotype: Uganda: Mukono, on Dracaena fragrans, Hansford 1982 (probably not preserved).

Host range and distribution: On Dracaena (arborea, fragrans [ugandensis], marginata hort., Dracaena sp.), Asparagaceae, Africa (Sierra Leone, Uganda), Asia (Japan), North America (Mexico; USA, Texas).

Notes: Type material has not been found. The present description is based on material on Dracaena arborea collected by Deighton in 1953 in Sierra Leone (K(M) IMI 53368). Material collected in Japan on D. fragrans is deposited as BPI 435828, and another collection from Mexico, intercepted at Brownsville, Texas, USA as BPI 435829. Braun & Sivapalan (1999) described and illustrated Stenella sp. on Dracaena sp. from Brunei (conidiophores solitary or emerging through stomata, 10–20 × 1.5–3 µm, 0–1-septate, subhyaline to pale olivaceous; conidia solitary, (5−)10–15(−25) × 1–2 µm, 0–1-septate, subhyaline to pale olivaceous, verruculose). Furthermore, Mall et al. (2013) recorded Stenella sp. on Dracaena marginata from India (Uttar Pradesh).

Zasmidium subsanguineum (Ellis & Everh.) U. Braun, Schlechtendalia 20: 103 (2010).

(Fig. 69)

Fig. 69
figure69

Zasmidium subsanguineum (HAL, Krieger, Fungi Saxon. Exs. 1293). A. Superficial hyphae. B. Conidiophore fascicles. C. Solitary conidiophores. D. Conidiophores. E. Conidia. Bar = 10 µm.

Basionym: Cercospora subsanguinea Ellis & Everh., J. Mycol. 4: 4 (1888).

Synonyms: Ramularia subsanguinea (Ellis & Everh.) Karak., in Vassiljevsky & Karakulin, Fungi Imperfecti Parasitici (Hyphomycetes) 1: 117 (1937).

Ramularia subsanguinea (Ellis & Everh.) Savile, Canad. J. Bot. 35: 205 (1957).

Stenella subsanguinea (Ellis & Everh.) U. Braun, Cryptog. Bot. 3: 242 (1993).

Ramularia rubicunda Bres., Hedwigia 36: 200 (1896) [lectotype (designated by Braun 1993): Germany: Saxony: Polenzthal, 13 June 1884, K. W. Krieger, Krieger, Fungi Saxon. Exs. 1293 (HAL); isolectotypes: Krieger, Fungi Saxon. Exs. 1293, e.g. BPI 419014, 4119023.

Literature: Saccardo (1892: 655), Vassiljevsky & Karakulin (1937: 217–118), Chupp (1954: 355), Braun (1998: 377).

Illustrations: Braun (1993a: 242, fig. 19; 1998: 375, fig. 636).

Exsiccatae: Kabát & Bubák, Fungi Imperf. Exs. 90. Krieger, Fungi Saxon. Exs. 1293. Krypt. Exs. 1495. Petr., Fl. Bohem. Morav. Exs. Pilze 1768. Syd., Mycoth. Germ. 2237.

Description: Leaf spots amphigenous, subcircular to somewhat angular-irregular, 0.5–6 mm diam, occasionally confluent and larger, at first yellowish or yellowish brown, later with pale centre and narrow to moderately wide, brown margin, finally often large leaf segments discoloured, brown, necrotic, sometimes with shot-hole symptoms. Caespituli usually hypophyllous, punctiform, greyish to pale reddish. Mycelium internal and external; superficial hyphae sparingly to well-developed, emerging through stomata or arising from stromata, branched, septate, 1–4 µm wide, yellowish to olivaceous, thin-walled, almost smooth to distinctly verruculose. Stromata substomatal, small, subhyaline to faintly pigmented. Conidiophores in small to moderately large, loose to dense fascicles, arising from stromatic hyphal aggregations, emerging through stomata, or solitary, arising from superficial hyphae, lateral, rarely terminal, erect, straight, flexuous, geniculate-sinuous, simple oroccasionally branched, 20–100 × 2–5 µm, continuous to pluriseptate, subhyaline, greenish, yellowish, olivaceous to reddish brown, thin-walled, almost smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, about 10–30 µm long, conidiogenous loci conspicuous, thickened and darkened, 1–1.5 µm diam. Conidia catenate, occasionally in branched chains, more or less cylindrical to fusiform, 10–55 × 2–6 µm, 0–3(−4)-septate, subhyaline, yellowish to pale olivaceous, thin-walled, verruculose, ends rounded to truncate, 1–2 µm diam, hila thickened and darkened.

Holotype: Canada: British Columbia: on Maianthemum canadense, 6 Oct. 1887, B.C. Macoun 12 (NY 838629). Isotypes: CUP 41349, NY 41348.

Host range and distribution: On Maianthemum (bifolium, canadense, dilatatum [kamtschaticum], racemosum subsp. amplexicaule), Asparagaceae, Asia (Kazakhstan; Russia, Siberia), Europe (Austria, Belarus, Czech Republic, Estonia, France, Finland, Germany, Hungary, Italy, Latvia, Lithuania, Poland, Russia, Slovakia, Sweden, Ukraine), North America (Canada, Ontario, Quebec; USA, Alaska, California, Michigan, New York, Ohio, Oregon, Pennsylvania, Vermont, Washington, Wisconsin).

Colchicaceae

Cercospora

Key to Cercospora species on Colchicaceae
  1. 1

    Conidiophores relatively short and narrow, 25–55 × 2–4 µm; conidia narrowly obclavate,

    30–80 × 2–3.5 µm, olivaceous; on Iphigenia .................................... C. iphigeniae

    Conidiophores longer, to 170 µm, and wider, 3–7 µm; conidia acicular to obclavate-cylindrical,

    hyaline, 2.5–5.5 µm wide; on other hosts ............................................... 2

  2. 2(1)

    On Disporum ................................................................. C. dispori

    On Gloriosa................................................................. C. gloriosae

Cercospora species on Colchicaceae

Cercospora dispori Togashi & Maki, Trans. Sapporo Nat. Hist. Soc. 17: 98 (1942).

(Fig. 70)

Fig. 70
figure70

Cercospora dispori (TNS-F-243895). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 345), Katsuki (1965: 42), Kim & Shin (1999a), Shin & Kim (2001: 62), Crous & Braun (2003: 164).

Illustration: Shin & Kim (2001: 63, fig. 22).

Description: Leaf spots amphigenous, scattered to confluent, usually vein-limited, subcircular, elliptical to angular, brown, ranging from yellowish brown, reddish brown to dark brown, 5–15 mm diam, confluent, later entire leaflets becoming necrotic, turning brown to blackish. Caespituli amphigenous, punctiform, scattered, dark brown to blackish. Mycelium internal; hyphae branched, septate, 1.5–3.5 µm wide, hyaline to pale olivaceous or brown, thin-walled, smooth. Stromata lacking to well-developed, 10–70 µm diam, substomatal or intraepidermal, subglobose, dark brown to blackish brown. Conidiophores in small to moderately large fascicles, 2–15, loose to moderately dense, occasionally solitary, arising from stromata, through stomata or erumpent, erect, straight, subcylindrical to geniculate in the upper half, unbranched, (10−)20–150(−200) × 3.5–7.5 µm, (0−)1–7(−9)-septate, pale olivaceous to brown, often paler towards the tip, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal and intercalary, about 10–30 µm long, proliferation sympodial, rarely percurrent, conidiogenous loci terminal and lateral, thickened and darkened, 1.5–2.5 µm diam. Conidia solitary, acicular or subacicular to cylindrical to almost filiform or somewhat obclavate-cylindrical, 25–130(−200) × 3.5–5.5 µm, 1–12-septate, hyaline, thin-walled, smooth, apex obtuse to subacute, base truncate to slightly obconically truncate, about 2–2.5 µm wide, hila somewhat thickened and darkened.

Holotype: Japan: Fukuoka Pref.: Mt Hiko, on Disporum smilacinum var. ramosum, 22 Sep. 1940, Y. Maki & T. Katsuki (TNS-F-243895).

Host range and distribution: On Disporum (smilacinum, viridescens), Colchicaceae, Asia (Japan, Korea).

Notes: Groenewald et al. (2013) included this species in molecular sequence analyses of the Cercospora s. str. complex and confirmed its position as species of its own. In a combined tree it clusters as sister to C. chinensis.

Cercospora gloriosae Syd., Ann. Cryptog. Exot. 2: 266 “1929” (1930).

(Fig. 71)

Fig. 71
figure71

Cercospora gloriosae (CUP 39896). A. Conidiophore fascicles. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Synonym: Cercospora gloriosicola J.M. Yen & Lim, Bull. Trimestriel Soc. Mycol. France 85: 467 “1969” (1970) [holotype: Singapore: on Gloriosa superba, 2 Aug. 1969, G. Lim (PC)].

Literature: Chupp (1954: 346), Vasudeva (1963: 114), Crous & Braun (2003: 199–200), Kamal (2010: 47).

Illustration: Yen & Lim (1980: 214, fig. 17).

Description: Leaf spots amphigenous, scattered, circular, subcircular to somewhat angular-irregular, 2–20 mm diam, at first greenish, yellowish, later dingy grey, with narrow darker border. Caespituli amphigenous, often more abundant on the lower leaf surface, punctiform, dark. Mycelium internal. Stromata substomatal, almost lacking or small, 10–40 µm diam, rarely larger, to 70 µm diam, medium to dark brown, composed of swollen hyphal cells, 3–7 µm diam. Conidiophores in small to large fascicles, loose to moderately dense, arising from stromata, through stomata, erect, straight, subcylindrical to distinctly geniculate or geniculate-sinuous towards the apex, unbranched or only rarely branched, 15–120(−170) × 3–7 µm, 1–8-septate, pale to medium brown or olivaceous thoughout or paler towards the tip, wall thin to somewhat thickened, smooth; conidiogenous cells integrated, terminal and intercalary, 10–30 µm long, conidiogenous loci conspicuous, thickened and darkened, 2.5–4 µm diam. Conidia solitary, shorter conidia obclavate-cylindrical, longer ones subacicular to distinctly acicular, 30–120 × 2.5–5 µm, 3–11-septate, hyaline, thin-walled, smooth, apex obtuse to acute, sometimes flagelliform, base truncate to short obconically truncate, 2.5–3.5 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank MBT178179): India: Bihar-: Pusa, on Gloriosa superba, 15 Sep. 1908, P. C. Kar, no. 2259 (CUP 39896). Isolectotype: HCIO 2259.

Host range and distribution: On Gloriosa (superba [virescens], Gloriosa sp.), Colchicaceae, Africa (Ethiopia, Uganda), Asia (India, Indonesia, Singapore).

Notes: A true Cercospora s. str. distinguished from C. apii s. lat., which possibly attacks Gloriosa spp. as well, by having distinct lesions and obclavate-cylindrical to acicular conidia with truncate to obconically truncate base and obtuse, rounded apices. The conidiogenous loci and conidial hila are 2.5–4 µm wide. Three well-developed Indian samples from Uttar Pradesh have been examined (K(M) 242532, 242946, 254732). The differentiation between C. gloriosa and C. gloriosicola is vague and could be proven by re-examinations of type material and other collections. Yen & Lim (1970) based the differentiation between the two species on Chupp’s (1954) description of C. gloriosae, who mentioned acicular conidia 2–4 µm wide. It is unclear on which specimens Chupp’s (1954) description and data were based. Yen & Lim (1970) described obclavate to obclavate-cylindrical conidia, 4–5 µm wide, and large stromata, 48–70 µm diam. Type material of C. gloriosicola, deposited at PC, has been re-examined, and acicular conidia 2.5–3.5 µm wide and stromata to 40 µm diam have been found. Thus, the size of the stromata and the conidial shape and size, obclavate-cylindrical to acicular, 2.5–5 µm wide, in C. gloriosa and C. gloriosicola are not genuinely different. In the type of C. gloriosicola the conidiophore fascicles are larger and denser, but such characters may be influenced by external conditions, and they are barely sufficient to warrant two separate species. The description of C. gloriosa in Chupp (1954) suggests that a second species with strictly acicular conidia, belonging to C. apii s. lat. may also occur on Gloriosa spp., but this question is still open and needs confirmation.

Cercospora iphigeniae Patwardhan & Sathe, Sydowia 19: 151 “1965”(1966).

(Fig. 72)

Fig. 72
figure72

Cercospora iphigeniae (based on Patwardhan & Sathe 1966: 152, fig. 2). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 228), Kamal (2010: 53).

Illustration: Patwardhan & Sathe (1966: 152, fig. 2).

Description: Leaf spots broadly fusiform, pale grey with black margin. Caespituli mainly epiphyllous. Stromata depressed globose to flattened, 10–20 µm diam, dark brown. Conidiophores fasciculate, slightly divergent, arising from stromata, erect, straight to curved, subcylindrical to geniculate-sinuous above, unbranched, 25–55 × 2–4 µm, 1–4-septate, brown, paler above, hyaline at the tip; conidiogenous cells integrated, terminal, with 1–3 thickened and darkened conidiogenous loci. Conidia solitary, narrowly obclavate, 30–80 × 2–3.5 µm, olivaceous, indistinctly 3–12-septate, thin-walled, apex pointed, base obconically truncate, hila thickened and darkened.

Holotype: India: Maharashtra: Pune, on Iphigenia indica, Colchicaceae, 23 Jul. 1964, P. G. Patwardhan & A. V. Sathe (AHMA 244).

Host range and distribution: Only known from the type collection.

Notes: Type material of this species was not available for re-examination. The generic affinity of this species is not quite clear as the conidia have been described to be olivaceous, so that it could also be a member of Passalora, but it seems rather to be a species of Cercospora s. str. A revision of type material is urgently necessary.

Commelinaceae

Cercospora

Key to Cercospora species on Commelinaceae
  1. 1

    Conidiophores 25–190 × 4–6.5 µm; conidia acicular or slightly obclavate, i.e. slightly obconically truncate at the base, about 1.5–3 µm wide, width variable 2–5 µm; on Pollia japonica ................................................................. C. polliae-japonicae

    Conidiophores shorter, usually 10–50 µm and/or conidia consistently acicular with truncate base or acicular to distinctly obclavate with smaller hila 1–2 µm wide; on other hosts ........................................................... 2

  2. 2(1)

    Conidia consistently acicular, 30–275 × 2–3 µm, base truncate; conidiophores 30–200 × 3–6 µm; conidiogenous loci 2.5–3 µm diam; on Commelina benghalensis ...................... C. benghalensis

    Conidia obclavate-cylindrical to acicular, at least partly with obconically truncate base; conidiogenous loci 1–2 µm wide ...................................................... 3

  3. 3(2

    Conidia 40–120 × (2−)3–5(−5.5) µm; conidiophores (20−)40–100 × 4–7 µm; on Amischotolype, Aneilema, Commelina and Murdannia spp., widespread ............................ C. commelinicola

    Conidia narrower, 1.5–3.5 µm; conidiophores shorter, usually 10–50 µm ................................ 4

  4. 4(3)

    With definite leafspots, subcircularto irregular, 1–4 mm diam;

    conidia occasionally in short chains; on Commelina longifolia ................................. C. commelinae-salicifoliae

    Entire leaflets turning brown, finally withering; conidia consistently solitary; on Murdannia nudiflora ....................................................... C. nudiflorae

Cercospora species on Commelinaceae

Cercospora benghalensis Chidd., Sydowia 13: 153

(1959).

(Similar to Fig. 1)

Literature: De (1991), Crous & Braun (2003: 78), Kamal (2010: 21–22).

Illustration: Chiddarwar (1959: plate V, figs 1–3).

Description: Leaf spots amphigenous, irregular-angular, 0.5–4 mm diam, scattered, dull to dark brown with paler brown centre. Caespituli amphigenous, scattered, punctiform, dark. Mycelium internal. Stromata substomatal, 10–25 µm diam, brown. Conidiophores in small to moderately large fasciles, 4–20, divergent, arising from stromata, through stomata, erect, straight to curved or geniculate-sinuous, unbranched, 25–235 × 3–4.5 µm, 0–11-septate, brown, thin-walled, smooth; conidiogenous cells integrated, terminal or intercalary, about 10–40 µm long, conidiogenous loci conspicuous, thickened and darkened, about 1.5–2 µm diam. Conidia solitary, acicular, sometimes almost filiform, straight to curved, 30–275 × 2–2.5 µm, 2–22-septate, hyaline, thin-walled, smooth, apex pointed, base truncate, about 1.5–2 µm wide, hila somewhat thickened and darkened.

Holotype: India: Maharashtra: Pune, Shivajinagar, Agricultural College, on Commelina benghalensis, 8 Sep. 1954, P. P. Chiddarwar 1 (K(M) IMI 83162). Isotype: BPI 433304.

Host range and distribution: On Commelina benghalensis, Commelinaceae, Asia (India, Maharashtra, West Bengal).

Notes: This species belongs to the Cercospora apii s. lat. complex.

Cercospora commelinae-salicifoliae A.K. Kar & M. Mandal, Indian Phytopathol. 26: 675 “1973”(1974).

(Fig. 73)

Fig. 73
figure73

Cercospora commelinae-salicifoliae (K(M) IMI 138302). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 133), Kamal (2010: 35).

Illustration: Kar & Mandal (1974: 676, fig. 2).

Description: Leaf spots amphigenous, scattered, subcircular to angular-irregular, 1–4 mm diam, pale brownish, later greyish brown to greyish white, margin darker, brown, reddish brown to purplish. Caespituli amphigenous, punctiform, dark. Mycelium internal. Stromata substomatal, subglobose to somewhat irregular, 10–40 µm diam, olivaceous-brown. Conidiophores in small to moderately large fascicles, loose to moderately dense, arising from stromata, through stomata, erect, straight, subcylindrical to geniculate-sinuous, unbranched, 10–45 × 2–5.5 µm, 0–4(−7)-septate, pale to medium olivaceous-brown or brown, thin-walled, smooth; conidiogenous cells integrated, terminal, occasionally intercalary or conidiophores reduced to conidiogenous cells, 10–25 µm long, conidiogenous loci thickened and darkened, 1–1.5 µm diam. Conidia solitary, narrowly obclavate-cylindrical, straight to slightly curved, 20–70 × 1.5–3.5 µm, 1–6-septate, hyaline, thin-walled, smooth, apex obtuse to acute, base subtruncate to short obconically truncate, 1–2 µm wide, hila thickened and darkened.

Holotype: India: West Bengal: Dasnapur, Midnapur, on Commelina longifolia [salicifolia], 13 Jan. 1969, M. Mandal (K(M) IMI 138302).

Host range and distribution: On Commelina longifolia, Commelinaceae, Asia (India, West Bengal).

Note: A true Cercospora s. str. distinct from C. apii s. lat. by its narrowly obclavate-cylindrical conidia with obconically truncate base and small conidiogenous loci.

Cercospora commelinicola Chupp ex U. Braun, Cryptog. Mycol. 20: 156 (1999).

(Fig. 74)

Fig. 74
figure74

Cercospora commelinicola (HBG, Syd., Fungi Exot. Exs. 1238). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Synonym: Cercospora commelinicola Chupp, Monograph of Cercospora: 116 (1954), nom. inval. (ICN, Art. 39.1).

Literature: Chupp (1954: 116), Katsuki (1965: 19), Crous & Braun (2003: 133), Guo et al. (2005: 65), Braun & Urtiaga (2008).

Illustrations: Braun (1999: 157, fig. 1), Guo et al. (2005: 65, fig. 40).

Exsiccatae: Sydow, Fungi Exot. Exs. 1238.

Description: Leaf spots amphigenous, subcircular to irregular, 1–15 mm diam or confluent and larger, forming large oblong blotches, to 30 mm diam, ochraceous to medium brown, later dull greyish brown to grey, margin narrow, dark brown to reddish brown. Caespituli usually epiphyllous, punctiform, dark brown. Mycelium internal. Stromata 10–45 µm diam, substomatal or intraepidermal, brown. Conidiophores in small to moderately large fascicles, 2–20, loose to moderately dense, arising from stromata, emerging through stomata or erumpent, erect, straight, subcylindrical and mostly barely or not geniculate to slightly geniculate-sinuous, unbranched, apex rarely swollen, (10−)30–100(−155) × 4–7 µm, 1- to pluriseptate, pale to medium dark brown or olivaceous-brown, paler towards the tip, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal, 10–40 µm long, conidiogenous loci conspicuous, often aggregated, not or barely prominent, 1.5–2 µm diam, thickened and darkened. Conidia solitary, narrowly obclavate-subcylindrical (-acicular), 40–150(−200) × (2−)3–5.5(−6) µm, pluriseptate, distance between septa 5–10 µm, rarely constricted at septa, hyaline, thin-walled, smooth, apex subacute, base short to long obconically truncate, 1.5–2.5 µm diam, hila somewhat thickened and darkened.

Holotype: Ecuador: Prov. Pichincha: Mindo, on Commelina virginica, 9 Nov. 1937, H. Sydow [Fungi Exot. Exs. 1238] (HBG). Isotypes: Syd., Fungi Exot. Exs. 1238, e.g., B, CUP 39483.

Host range and distribution: On Amischotolype hookeri, Aneilema aequinoctiales, Commelina (benghalensis, communis, diffusa, erecta [elegans], imberbis [kotschyi], paludosa, virginica), Commelinaceae, Africa (Kenya, Malawi, Sudan, Tanzania), Asia (China, India, Japan), South America (Ecuador, Venezuela), West Indies (Barbados, Cuba).

Notes: Chupp proposed this name as “new combination” with reference to “C. commelynae Kalchbr. & Cooke” as described in Sydow (Ann. Mycol. 37: 425, 1939), but there was no valid basionym, as Sydow did not provide any Latin description or diagnosis. This species is a true Cercospora s. str. distinct from C. apii s. lat.

Cercospora nudiflorae Chupp, Monograph of Cercospora: 117 (1954).

(Fig. 75)

Fig. 75
figure75

Cercospora nudiflorae (CUP 35213). A. Conidiophore fascicles. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 293).

Description: Leaf spots lacking, entire attacked leaves turning brown and withering. Caespituli amphigenous, punctiform, dark. Mycelium internal. Stromata substomatal, small, 10–30 µm diam, brown. Conidiophores in small to moderately large fascicles, loose to dense, arising from stromata, through stomata, erect, straight, barely or only slightly geniculate-sinuous, unbranched, (10−)15–50(−80) × 3–6 µm, usually 1–3-septate, pale to medium olivaceous-brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, occasionally conidiophores reduced to conidiogenous cells, 10–25 µm long, with a single or several conidiogenous loci, sometimes aggregated, barely prominent, 1–2 µm diam, thickened and darkened. Conidia solitary, narrowly obclavate to acicular, 20–80 × 1.5–3.5 µm, indistinctly pluriseptate, hyaline, thin-walled, smooth, apex acute or subobtuse, base truncate to obconically truncate, 1–2 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank MBT178143): Bermuda: Devonshire Marsh, On Murdannia nudiflora [Commelina nudiflora], 12 Jan. 1926, H. H. Whetzel (CUP 35213). Isolectotype: CUP 40395.

Host range and distribution: On Murdannia nudiflora, Commelinaceae; South America (Venezuela), West Indies (Bermuda).

Notes: Records of Cercospora commelinicola from Venezuela and West Indies (Minter et al. 2001, Urtiaga 1966) undoubtedly refer to C. nudiflorae. The two species have rather similar conidiophores and conidia. A record on Commelina sp. from Florida (Crous & Braun 2003) is doubtful and appears to belong to C. commelinicola.

Cercospora polliae-japonicae Y.L. Guo & Y. Jiang, Mycosystema 19: 302 (2000).

(Fig. 76)

Fig. 76
figure76

Cercospora polliae-japonicae (HMAS 77628). A. Conidiophores. B. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 331), Guo et al. (2005: 66).

Illustrations: Guo & Jiang (2000b: 303, fig. 1), Guo et al. (2005: 657, fig.41).

Description: Leaf spots amphigenous, subcircular, 2–12 mm diam, pale brown to brown. Caespituli amphigenous. Mycelium internal. Stromata lacking or small, 10–45 µm diam, brown. Conidiophores solitary or in small to moderately large, divergent fascicles, 2–28, erect, straight to curved, unbranched, somewhat geniculate, 25–190 × 4–6.5 µm, 0–7-septate, brown; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci thickened and darkened, 2–3.5 µm diam. Conidia solitary, acicular to somewhat obclavate, straight to somewhat curved, 40–200 × 2–5 µm, pluriseptate, hyaline, thin-walled, smooth, apex pointed, base truncate to slightly obconically truncate, about 1.5–3 µm wide, hila thickened and darkened.

Holotype: China: Zhejiang Province: Hangzhou, on Pollia japonica, Commelinaceae, 26 May 1961, Q. M. Ma & X. J. Liu (HMAS 77628).

Host range and distribution: Only known from the type collection.

Notes: This species belongs to the Cercospora apii s. lat. complex.

Doubtful, excluded and insufficiently known species

Cercospora commelinae Kalchbr. & Cooke, Grevillea 9: 24 (1881); as “commelynae”.

Literature: Chupp (1954: 116), Crous & Braun (1996: 260; 2003: 133).

Syntypes: South Africa: Cape, on Commelina benghalensis, MacOwan 1346 (B 700016011, K(M) 193949, 193950).

Host range and distribution: On Commelina benghalensis, Commelinaceae, Africa (South Africa, Zimbabwe).

Notes: Chupp (1954) examined the type specimen lodged at Kew (MacOwen 1346), and concluded that C. commelinae was in fact a species of Septoria. An examination of a South African collection (PREM 32777) identified by Chupp & Doidge (1948) to be C. commelinae and the type from B confirmed that the fungus would be better placed in Septoria.

Pseudocercospora

Key to Pseudocercospora species on Commelinaceae
  1. 1

    In vivo with external mycelium; stromata lacking or almost so; with solitary conidiophores

    arising from superficial hyphae; on Pollia japonica, Taiwan ............................................... P. polliae

    Superficial mycelium in vivo lacking; conidiophores fasciculate, arising from small

    to well-developed stromata; on other hosts ........................................................ 2

  2. 2(1)

    Conidiophores only 0–3-septate; on Amischotolype hispida, Taiwan ............................................... P. forrestiae

    Conidiophores pluriseptate; on Commelina erecta, Trinidad ............................................. P. maracasensis

Pseudocercospora species on Commelinaceae

Pseudocercospora forrestiae Goh & W.H. Hsieh, Trans. Mycol. Soc. Republ. China 2: 130 (1987).

(Fig. 77)

Fig. 77
figure77

Pseudocercospora forrestiae (NTU-PPE, holotype). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Synonym: Cercospora forrestiae Sawada, Taiwan Agric. Res. Inst. Rept. 85: 107 (1943), nom. inval. (ICN, Art. 39.1).

Literature: Chupp (1954: 117), Hsieh & Goh (1990: 59), Guo & Hsieh (1995: 58), Guo et al. (1998: 74), Crous & Braun (2003: 187).

Illustrations: Hsieh & Goh (1990: 59, fig. 41), Guo & Hsieh (1995: 61, fig. 56), Guo et al. (1998: 75, fig. 58).

Description: Leaf spots amphigenous, elliptical, 5–12 × 3–5.5 mm, greyish brown, with dark brown margin. Caespituli amphigenous. Mycelium internal. Stromata dark brown, to 50 µm diam. Conidiophores in well-developed, dense fascicles, 10–25, arising from stromata, erect, straight to curved, cylindrical to geniculate-sinuous, unbranched, 20–80 × 3.5–5 µm, 0–3-septate, pale olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or occasionally conidiophores reduced to conidiogenous cells, conidiogenous loci neither thickened nor darkened, inconsociuous or visible as truncate tips or shoulders. Conidia solitary, obclavate-cylindrical, straight to curved, 35–145 × 3–5 µm, 2–13-septate, pale olivaceous, thin-walled, smooth, apex obtuse to pointed, base obconically truncate, hila unthickened, not darkened.

Holotype: Taiwan: Taipei, Shihting, on Amischotolype hispida [chinensis], Commelinaceae, 6 Nov. 1927, K. Sawada (NTU-PPE [herb. Sawada]). Isotype: TNS-F-220428.

Host range and distribution: Only known from the type collection.

Pseudoercospora maracasensis (R.E.D. Baker & W.T. Dale) Deighton, Mycol. Pap. 140: 147 (1976). (Fig. 78)

Fig. 78
figure78

Pseudocercospora maracasensis (CUP 35326). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercospora maracasensis R.E.D. Baker & W.T. Dale, Mycol. Pap. 33: 103 (1951).

Literature: Chupp (1954: 117), Crous & Braun (2003: 266–267).

Description: Leaf spots amphigenous, subcircular to irregular or somewhat angular-irregular, 4–30 mm diam, at first dull greenish, greyish green or greenish brown, later gradually changing to yellowish brown or brown, finally grey or greyish white, with distinct margin or marginal line, narrow, dark olivaceous-brown to dark reddish brown or blackish. Caespituli amphigenous, mainly hypophyllous, punctiform, scattered, blackish brown or dull greyish brown by abundant conidiation. Mycelium internal. Stromata small to well-developed, substomatal, subglobose to flattened above, 10–60 µm diam, medium to dark olivaceous-brown, cells 2–5 µm diam, rounded to angular in outline. Conidiophores in small to large fascicles, divergent to dense, sometimes almost coremioid, arising from stromata, through stomata, erect, subcylindrical-filiform, straight to curved, sinuous or somewhat geniculate, unbranched, 30–200 × 3–6 µm, pluriseptate, pale to medium olivaceous-brown, darker in mass, paler towards the tip, ultimate tip sometimes subhyaline, thin-walled, smooth; conidiogenous cells integrated, terminal, about 10–30 µm long, conidiogenous loci inconspicuous, unthickened and not darkened, occasionally somewhat refractive. Conidia solitary, obclavate-cylindrical, straight to curved, 35–110 × 3–6 µm, 3–9-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex obtuse or subobtuse, base short to long obconically truncate, (1.5−)2(−2.5) µm wide, hila unthickened, not darkened.

Holotype: Trinidad: Maracas Valley, on Commelina erecta, 4 Oct. 1944, R. E. D. Baker 226 (K(M) IMI 24457). Isotypes: CUP 35326, TRIN.

Host range and distribution: On Commelina erecta [elegans, virginica], Commelinaceae, West Indies (Trinidad and Tobago).

Pseudocercospora polliae Goh & W.H. Hsieh, Trans. Mycol. Soc. Republ. China 2: 134 (1987).

(Fig. 79)

Fig. 79
figure79

Pseudocercospora polliae (NTU-PPE, holotype). A. Conidiophore fascicle. B. Conidiophores and superficial hypha with solitary conidiophore emerging through a stoma. C. Conidia. Bar = 10 µm.

Synonym: Cercospora polliae Sawada, Rep. Gov. Agric. Res. Inst. Taiwan 87: 86 (1944), nom. inval. (ICN, Art. 39.1).

Literature: Chupp (1954: 117, 118), Katsuki (1965: 19), Hsieh & Goh (1990: 60), Guo & Hsieh (1995: 59), Guo et al. (1998: 75), Crous & Braun (2003: 330).

Illustrations: Hsieh & Goh (1990: 60, fig. 42), Guo & Hsieh (1995: 63, fig. 57), Guo et al. (1998: 76, fig. 59).

Description: Leaf spots indistinct to circular or elliptical, 10–15 mm diam, centre greyish brown, margin dark brown.

Caespituli amphigenous, mainly hypophyllous. Mycelium internal and external; superficial hyphae emerging through stomata, sparingly branched, septate, 1.5–3.8 µm wide, pale olivaceous, thin-walled, smooth. Stromata lacking or consisting of a few substomatal swollen hyphal cells to medium-seized, to 50 µm diam. Conidiophores in small fascicles, 2–10, emerging through stomata, or solitary, arising from superficial hyphae, lateral, erect, straight to somewhat curved, subcylindrical to somewhat geniculate-sinuous, unbranched, (15−)20–60 × 4–5 µm, 0–3-septate, brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, conidiogenous loci inconspicuous. Conidia solitary, obclavate-filiform, subacicular, straight to somewhat curved, 30–130 × 3–4.5 µm, 3–13-septate, subhyaline to very pale olivaceous, thin-walled, smooth, apex pointed to subobtuse, base subtruncate to short obconically truncate, 2–2.5 µm wide, hilum neither thickened nor darkened.

Holotype: Taiwan: Taipei, on Pollia japonica, 23 Mar. 1919, K. Sawada (NTU-PPE [hb. Sawada]). Isotype: TNS-F-220503.

Host range and distribution: On Pollia japonica, Commelinaceae, Asia (China, Japan, Taiwan).

Costaceae

Pseudocercospora

Key to Pseudocercospora species on Costaceae
  1. 1

    Conidiophores short, in dense fascicles, 5–30 × 2–5 µm, usually aseptate; conidia straight to slightly curved, 25–110 × 2–4 µm; on Costus spicatus and Costus sp.,

    Central and South America ................................................. P. costi

    Conidiophores very long, in loose to mostly dense, often even coremioid fascicles, 70–650 × 3–7 µm, pluriseptate; conidia straight to mostly distinctly curved, often strongly curved, sometimes apex even uncinate to circinate, much wider,

    20–90 ×(3.5−)5–7(−8) µm; on Cheilocostus speciosus, Asia and Oceania ................................................. P. costina

Pseudocercospora species on Costaceae

Pseudocercospora costi (F.L. Stevens) U. Braun & Crous, in Crous & Braun, Mycosphaerella and Anam.: 140 (2003).

(Fig. 80)

Fig. 80
figure80

Pseudocercospora costi (ILL 15148). A. Superficial hypha. B. Superficial hypha with solitary conidiophore. C. Conidiophore fascicles. D. Conidiophores. E. Conidia. Bar = 10 µm.

Basionym: Cercospora costi F.L. Stevens, Illinois Biol. Monogr. 11: 57 (1927).

Literature: Chupp (1954: 607), Urtiaga (1986), Braun & Urtiaga (2012, 2013a).

Description: Lesions variable, irregular, small to large blotches, to 50 mm diam, pale brown, with narrow yellowish border. Caespituli amphigenous, scattered, punctiform, dark. Mycelium internal, occasionally also external with a few superficial hyphae, unbranched or sparingly branched, 1–3 µm wide, subhyaline to pale olivaceous, septate, thin-walled, smooth. Stromata 10–40 µm diam, substomatal, globose, yellowish brown to brown. Conidiophores in small to moderately large fascicles, arising from stromata, emerging through stomata, dense, occasionally solitary, arising from superficial hyphae, erect, straight, subcylindrical-conical to somewhat geniculate-sinuous, unbranched, 5–30 × 2–5 µm, 0(−1)-septate, subhyaline to pale olivaceous or yellowish brown, thin-walled, smooth; conidiophores usually reduced to conidiogenous cells, conidiogenous loci inconspicuous, occasionally subdenticulate, but always unthickened and not darkened. Conidia solitary, narrowly obclavate-cylindrical to almost filiform, 25–110 × 2–4 µm, 2–10-septate, subhyaline to very pale olivaceous, thin-walled, smooth, apex subacute or subobtuse, base short to long obconically truncate, 1–2 µm wide, hila unthickened, not darkened.

Holotype: Panama: Gatun, on Costus sp., 24 Aug. 1923, F. L.

Stevens (ILL 15148).

Host range and distribution: On Costus (spicatus, Costus sp.), Costaceae, Central and South America (Panama, Venezuela).

Notes: The conidiophores are very short, arranged in dense, often almost sporodochial conidiomata. The conidiogenous loci are inconspicuous. Some superficial hyphae with solitary conidiophores have been observed.

Pseudocercospora costina (Syd. & P. Syd.) Deighton, Mycol. Pap. 140: 135 (1976). (Fig. 81)

Fig. 81
figure81

Pseudocercospora costina (K(M) IMI 82089). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora costina Syd. & P. Syd., Ann. Mycol. 14: 372 (1916).

Synonyms: Ancylospora costi Sawada, Rep. Gov. Res. Inst. Formosa 87: 78 (1944), nom. inval. (ICN, Art. 39.1) [syntypes: Taiwan: Kaohsiung, on Cheilocostus speciosus [Costus speciosus], 6 Oct. 1908, R. Suzuki (BPI 4428256; K(M) IMI 31902, NTU-PPE, hb. Sawada].

Helicomina costi M.A. Salam & P.N. Rao, Indian Phytopathol. 11: 123 (1958) [holotype: India: Andhra Pradesh: Hyderabad, Pakhal forest, on Cheilocostus speciosus, 28 Sep. 1956, M. A. Salam (K(M) IMI 90212a)].

Helicomina dracaenae Hasija, Indian Phytopathol. 19: 373 (1967) [holotype: India: Madhya Pradesh, Jabalpur, on Costus sp. (not Dracaena sp.), Sep. 1964, S. K. Hasija (K(M) IMI 109484)].

Phaeoisariopsis costi A.K. Singh, S.K. Singh & Kamal (“costusae”), Curr. Sci. 53: 876 (1984) [holotype: India: Uttar Pradesh, Gorakhpur, on Cheilocostus speciosus, Jan. 1980, A. K. Singh, KA-25 (K(M) IMI 244879)].

Passalora costi (A.K. Singh, S.K. Singh & Kamal) U. Braun & Crous, in Crous & Braun, Mycosphaerella and Anam.: 449 (2003).

Literature: Saccardo (1931: 895), Chupp (1954: 607), Deighton (1976: 135), Hsieh & Goh (1990: 356), Guo & Hsieh (1995: 355), Guo et al. (1998: 374), Braun (2001b: 62), Crous & Braun (2003: 141), Kamal (2010: 115, 167).

Illustrations: Deighton (1976: 136, fig. 81, 137, fig. 82), Singh et al. (1984: 876, fig. 1), Hsieh & Goh (1990: 356, fig. 273), Guo & Hsieh (1995: 356, fig. 300), Guo et al. (1998: 375, fig. 306).

Description: Leaf spots almost lacking, yellowish discolorations on the upper leaf surface, becoming dark when dry or with large subcircular to irregular patches, 5–50 mm diam, dingy grey to greyish white, margin indefinite. Caespituli hypophyllous, punctiform, scattered to dense, dark olivaceous, brown to blackish. Mycelium internal; hyphae sparingly branched, 2–5 µm diam, septate, pale olivaceous, thin-walled, smooth. Stromata substomatal, subglobose to somewhat irregular, 20–70 µm diam, brown or olivaceous-brown. Conidiophores in small to moderately large fascicles, to 20 conidiophores, loose to usually dense, often almost coremioid, arising from stromata, through stomata, erect, straight, subcylindrical-filiform, usually barely geniculate-sinuous, unbranched, 70–650 × 3–7 µm, pluriseptate throughout, pale to medium dark olivaceous or brown, paler towards the tip, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal or intercalary, 10–40 µm long, sympodially and occasionally percurrently proliferating, conidiogenous loci inconspicuous to conspicuous by being subdenticulate, but always unthickened and not darkened. Conidia solitary, subcylindrical, cylindrical-obclavate to somewhat clavate, short conidia sometimes broadly ellipsoid-ovoid, straight to mostly distinctly curved, often strongly curved, sometimes apex even uncinate to circinate, 20–90 × (3.5−)5–7(−8) µm, (1−)2–8(−9)-septate, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth, apex obtuse, rounded, base short obconically truncate, gradually attenuated towards the base or sometimes abruptly attenuated at the very base, peg-like, 1.5–2.5(−3) µm wide, hila neither thickened nor darkened.

Type: Philippines: Prov. Laguna: Los Baños, on Cheilocostus speciosus [Costus speciosus], Jan. 1916, C. F. Baker 4149 p.p. (K(M) IMI 82089).

Host range and distribution: On Cheilocostus speciosus [Costus speciosus], Costus sp., Costaceae, Asia (China, India, Nepal, Philippines, Taiwan), Oceania (Fiji, Micronesia, Solomon Islands, Vanuatu).

Notes: Type material of this species is not preserved in Sydow’s herbarium at S. K(M) IMI 82089 is the only fragment of type material that could be traced. It is unclear if additional duplicates exist. Therefore, we prefer to postpone a final typification. Records of this species on Hedychium sp. (Zingiberaceae) are doubtful (Braun 2001, Crous & Braun 2003). A collection from India on “Hedychium” sp. (Jabalpur, 19 Oct. 1977, R. C. Rayak, K(M) IMI 217586) has been examined and undoubtedly represents P. costina, but the identity of the host plant is unclear. It cannot be excluded that the leaves belong to Costus s. lat.

Cyperaceae

Cercospora

Key to Cercospora species on Cyperaceae
  1. 1

    Conidia strictly acicular, base of the conidia truncate ..................................................... 2

    Conidia not strictly acicular, conidia at least partly obclavate-cylindrical or filiform and base of the conidia at least partly obconically truncate ..................................................... 4

  2. 2(1)

    With distinct leaf spots; conidiophores short, 20–45 µm; on Cyperus fuscus, Europe

    ..................................................... C. cyperi-fusci (see “doubtful, excluded and insufficiently known species”)

    Leaf spots lacking; conidiophores much longer, 30–120 µm; on Kyllinga erecta, Africa ...................................................... 3

  3. 3(2)

    Conidiophores often branched; conidia 20–120 × 1.5–3 µm, 1–12-septate ....................................... C. kyllingae

    Conidiophores unbranched; conidia longerand wider, 60–250 × 3–4 µm, 8–21-septate ................................... C. kyllingicola

  4. 4(1)

    Conidiophores very long, 30–200 µm, pluriseptate; conidia short, cylindrical-fusiform,

    (12−)15–35 × 2–4 µm, 1–3-septate; on Bulbostylis, Rhynchospora and Scleria spp ........................ C. glauciana

    Conidiophores shorter, to 90 µm; conidia longer, 20–120 µm, often more than 3 septa ........................ 5

  5. 5(4)

    Conidiophores (10−)20–90 µm long; conidia (30−)40–120(−150) × (2−)2.5–5(−5.5) µm; on Cyperus spp., Africa and Asia ................................... C. cyperi

    Conidiophores shorter, usually 5–35 µm; conidia narrower, 1.5–3 µm .................................... 6

  6. 6(5)

    Conidiophores very short, 5–20 µm, 0–1-septate ........................................ 7

    Conidiophores longer, 10–35(−60) µm, 0–4-septate ........................................8

  7. 7(6)

    Conidiophores 2–5 µm wide, hila 1–2 µm wide; conidia narrowly cylindrical-filiform to acicular; on Cyperus spp .................................................... C. cyperigena

    Conidiophores narrower, 1.5–3 µm, hila 1 µm diam; conidia narrowly obclavate-filiform; on Eleocharis spp ........................................................ C. eleocharidis

  8. 8(6)

    Conidiophores very pale, subhyaline to pale olivaceous; conidiogenous loci very small, 1–1.5(−2) µm diam; on Carex, Cyperus, Eleocharis, Fuirena and Kobresia spp. ........................................ C. caricis

    Conidiophores distinctly pigmented, olivaceous-brown; conidiogenous loci somewhat broader, 1.5–2 µm; on species of Scirpus s. lat. ........................................ C. uredinophila

Cercospora species on Cyperaceae

Cercospora caricis Oudem., Nederl. Kruidk. Arch. II, 6: 59 (1892).

(Fig. 82)

Fig. 82
figure82

Cercospora caricis (LEP, Ellis & Everh., Fungi Columb. 1170). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Synonyms: Cercospora caricina Ellis & Dearn., Proc. Canad. Inst., N.S., Part 3, 1: 91 (1897) [lectotype (designated here, MycoBank MBT178144): Canada: Ontario: London, on Carex rosea, Aug. 1896, J. Dearness 2390 (DAOM); isotypes: NY and Ellis & Everh., Fungi Columb. 1170, e.g., BPI 434250–434253, HBG, LEP].

Cercospora microstigma Sacc., Ann. Mycol. 10: 315 (1912) [holotype: Canada: Ontario: London, on Carex laxiflora, 6 Aug. 1910, J. Dearness (PAD); isotypes: Barthol., Fungi Columb. 4904, e.g. BPI 438409].

Cercospora caricis Dearn. & House, New York State Mus. Bull. 188: 29 (1916), nom. illeg. (ICN, Art. 53.1) [type: USA: New York: Old Forge, on Carex folliculata, Aug., C. H. Peck (NYS)].

Cercosporina caricis Sacc., Syll. Fung. 25: 900 (1931), nom. nov., as “(Dearn. & House) Sacc., comb. nov.”.

Literature: Saccardo (1899: 1105; 1913: 1431), Lindau (1910: 88), Chupp (1954: 191), Vasudeva (1963: 70), Blaney et al. (1988), Crous & Braun (1996: 254; 2003: 105), Braun et al. (1999: 298), Inglis et al. (2001), Kamal (2010: 29), Pirnia et al. (2012).

Illustration: Vasudeva (1963: 71, fig. 36).

Exsiccatae: Barthol., Fungi Columb. 4904. Ellis & Everh., Fungi Columb. 1170. Ellis & Everh., N. Amer. Fungi 3489.

Description: Leaf spots amphigenous, elliptical to oblong or somewhat irregular, about 2–10 × 1–3 mm, pale brown to brown, sometimes paler with age, margin indefinite or narrow and darker brown or lesions expanded, or the whole leaf or larger leaf segments becoming necrotic. Caespituli hypophyllous, punctiform, dark, often linear between veins. Mycelium internal. Stromata substomatal, 10–60 µm diam, brown to dark brown. Conidiophores in small to moderately large fascicles, divergent to dense, sometimes very dense, almost coremioid, arising from stromata, through stomata, erect, straight, subcylindrical to moderately geniculate-sinuous, unbranched, 10–45 × 3–6 µm, 0–3(−4)-septate, olivaceous-brown throughout or with paler tips, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, about 10–30 µm long, conidiogenous loci conspicuous, thickened and darkened, 1–1.5(−2) µm diam. Conidia solitary, narrowly obclavate-cylindrical to acicular, (20−)25–80(−120) × 1.5–3 µm, usually 2–8-septate, hyaline, thin-walled, smooth, apex pointed to subobtuse, base truncate to short obconically truncate, 1–2 µm wide, hila somewhat thickened and darkened.

Holotype: The Netherlands: The Hague, on Carex sp., Sep. 1891, C. E. Destrée (not traced).

Host range and distribution: On Carex (albursina, alopecoidea, arctata, bebbii, bicknellii, bromoides, castanea, cephalophora, crawfordii, cristatella, davisii, deflexa, folliculata, granularis, gravida, interior, intumescens, laxiflora, lupulina, microptera, normalis, orbicularis, pensylvanica, pilulifera [oederi], plantaginea, projecta, pubescens, retrorsa, rosea [convoluta], sartwelliana, stipata, stricta, subcapitatus, tenera, vulpinoidea [muhlenbergii], Carex sp.), Cyperus (articulatus [fistulosus], esculentus, filiculmis, mitis [subcapitatus], rotundus, Cyperus sp.), Eleocharis acutangula [fstulosa], Fuirena pubescens [Carex pubescens], Kobresia simpliciuscula [Carex mirabilis], Cyperaceae, Africa (South Africa), Asia (India, Iran), Australia, Caucasus (Georgia), Europe (Great Britain, Ireland, The Netherlands), South America (Brazil), Oceania (New Caledonia), North America (Canada; USA, Arizona, Iowa, Illinois, Kansas, Maine, North Carolina, New York, Washington, Wisconsin).

Notes: A true Cercospora s. str. distinct from C. apii s. lat. by having much smaller conidiogenous loci, 1–1.5(−2) µm wide, and obclavate-subcylindrical conidia with obconically truncate bases. Further studies including pathogenicity tests on Carex and Cyperus are required to prove the conspecificity of species occurring on these two host genera. Material from the Netherlands appropriate for a neotypification was not found. Therefore, we preferto postpone a neotypification.

Cercospora cyperi Sawada, Trans. Nat. Hist. Soc. Taiwan 17: 179 (1914).

(Fig. 83)

Fig. 83
figure83

Cercospora cyperi (NTU-PPE, holotype). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Synonyms: Cercospora cypericola Chupp & H.C. Greene, Amer. Midl. Naturalist 50: 508 (1953) [lectotype (designated here, MycoBank MBT178145): USA: Wisconsin: Dane County, Madison, University of Wisconsin Arboretum, on Cyperus strigosus, 12 Aug. 1952, H. C. Greene (CUP 40865); isotypes: BPI 435441, CUP 40873, WIS].

Cercospora ugandensis Hansf., Proc. Linn. Soc. London 1942–1943: 59 (1943) [type: Uganda: Kampala Swamp, on Cyperus sp. (Mariscus sp.), Jul. 1930, Hansford 1311 (not traced)].

Cercospora cyperi-rotundi Thirum. & Govindu, Sydowia 7: 312 (1953) [type: India: Bihar: Patna, on Cyperus rotundus, 3 Dec. 1952, M. J. Thirumalachar (probably not preserved)].

Literature: Chupp (1954: 191), Vasudeva (1963: 95), Hsieh & Goh (1990: 103), Crous & Braun (2003: 150), Guo et al. (2005: 105), Kamal (2010: 38).

Illustrations: Thirumalachar & Govindu (1953: plate 7, figs 19–20), Hsieh & Goh (1990: 104, fig. 78), Guo et al. (2005: 106, fig.71).

Description: Leaf spots amphigenous, irregular to oblong, 1–25 × 0.5–5 mm, brown, margin indefinite. Caespituli hypophyllous, punctiform, fine, brown to dark brown. Mycelium internal. Stromata lacking or small, substomatal, 10–25 µm diam, brown, composed of thick-walled cell, 2–6 µm diam. Conidiophores in small to moderately large fascicles, loose to rather dense, rarely solitary, arising from internal hyphae or stromata, through stomata, erect, straight, subcylindrical to somewhat geniculate-sinuous, unbranched, (10−)20–90 × 3–6(−7) µm, 0–5-septate, pale to medium olivaceous-brown or brown throughout or paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, 10–30 µm long, conidiogenous loci small, (1−)1.5–2(−2.5) µm wide, thickened and darkened. Conidia solitary, narrowly obclavate-cylindrical, subacicular, (30−)40–120(−150) × (2−) 2.5–5(−5.5) µm, 3–10-septate, hyaline, thin-walled, smooth, apex pointed or subobtuse, base short to long obconically truncate to subtruncate, 1.5–2 µm wide, hila thickened and darkened.

[Holotype: Taiwan: Taipei, on Cyperus pilosus, 8 Nov. 1913, K. Sawada (not preserved)]. Neotype (designated here, MycoBank MBT178174): Taiwan: Taipei, on Cyperus sp., 25 Oct. 1924, K. Sawada (NTU-PPE [hb. Sawada]).

Host range and distribution: On Cyperus (esculentus, filiculmis, houghtonii, pilosus, rotundus, schweinitzii, strigosus, Cyperus spp.), Cyperaceae, Africa (Libya, Morocco, Nigeria, Uganda), Asia (China, India, Indonesia, Taiwan), ?North America (USA, Wisconsin), ?South America (Brazil).

Notes: Numerous collections on Cyperus spp., determined as C. cyperi, C. cypericola, C. cyperi-rotundi as well as C. ugandensis, have been examined and proved to represent a single species, characterised by having similar leaf spots, small conidiogenous loci, 1.5–2 µm diam, and obclavate-cylindrical conidia, (2−)2.5–5(−6) µm wide, with small hila, 1–2 µm diam. The status of C. ugandensis, described from Cyperus sp. [Mariscus sp.], is uncertain. Type material could not be traced, but the original description of this species agrees well with C. cyperi. Therefore, C. ugandensis is tentatively reduced to synonym with the latter species. Description and illustration of “C. ugandensis” from India (see Vasudeva, 1963: 205, fig. 151) do not agree with C. cyperi. The Indian collection is characterised by having large stromata, 30–70 µm diam, very numerous densely fasciculate conidiophores, and narrow conidia, 14–71 × 2–3 µm. African records of C. ugandensis on Kyllinga elatior and Kyllinga sp. (Crous & Braun 2003) are unclear. A specimen from Zimbabwe has been examined (on Kyllinga sp., Uruma, 29 Mar. 1957, J. O. Whiteside, K(M) IMI 69047a). In any case, this collection does neither belong to C. kyllingae nor to C. kyllingicola. It is closer to C. cyperi, but the conidiophores are uniformly short, about 10–30 µm, and the conidia are very narrow, about 1.5–2.5 µm.

Cercospora cyperi is morphologically very close to C. caricis, but differs in having somewhat wider conidia. Inoculation experiments and molecular examinations are necessary to find out if these differences are sufficient to maintain two distinct species. Cercospora cyperi seems to be confined to Africa and Asia. Records from North America are doubtful and seem to refer to C. caricis. Another record on Cyperus sp. from Brazil (Mendes et al. 1998) is also uncertain. Type material of C. cyperi, collected up to 1919, is not preserved in Sawada’s herbarium at the National Taiwan University. Therefore, authentic material from 1924 is designated as neotype.

Cercospora cyperigena U. Braun & Crous, in Crous & Braun Mycosphaerella and Anam.: 151 (2003).

(Fig. 84)

Fig. 84
figure84

Cercospora cyperigena (K(M) IMI 259773). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bars = 10 µm.

Literature: Braun & Urtiaga (2012: 302–303).

Illustration: Crous & Braun (2003: 151, fig. 8).

Description: Leaf spots amphigenous, subcircular to elliptical-fusiform, 1–8 mm diam, occasionally confluent, centre yellowish olivaceous to light brown, with a narrow brown margin. Caespituli hypophyllous, finely punctiform, often in lines, blackish brown, later greyish white by rich conidial formation. Mycelium internal. Stromata substomatal, 10–40 µm diam, olivaceous-brown. Conidiophores numerous, in dense fascicles, arising from stromata, emerging through stomata, erect, straight, subcylindrical-conical, slightly geniculate-sinuous, unbranched, 5–20 × 2–5 µm, 0–1-septate, pale olivaceous or olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–15 µm long, conidiogenous loci conspicuous, 1–2 µm wide, thickened and darkened. Conidia solitary, narrowly cylindrical-filiform to subacicular, 40–160 × 1–2.5 µm, pluriseptate, mostly 6–14-septate, hyaline, thin-walled, smooth, apex subacute, base truncate, hila 1–2 µm wide, slightly thickened and darkened.

Holotype: Tanzania: on Cyperus sp., 28 May 1981, C. L. Keswani (K(M) IMI 259573).

Host range and distribution: On Cyperus (rotundifolius, Cyperus sp.), Cyperaceae, Africa (Tanzania), South America (Venezuela).

Note: This species is quite distinct from Cercospora cyperi (incl. C. cypericola and C. cyperi-rotundi) by very short, 0–1-septate conidiophores and very long and narrowly cylindrical-filiform to subacicular conidia.

Cercospora eleocharidis Davis, Trans. Wisconsin Acad. Sci. 24: 300 (1929). (Fig. 85)

Fig. 85
figure85

Cercospora eleocharidis (BPI 436100). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 192), Crous & Braun (2003: 172), Braun & Crous (2005: 409).

Description: Leaf spots formed as small, oblong, brown discolorations, finally the whole leaf blade sometimes turning brown. Caespituli amphigenous, finely punctiform, dark. Mycelium internal. Stromata lacking or formed as small aggregations of swollen hyphal cells in the substomatal cavity, 10–20 µm diam, brown. Conidiophores in small to moderately large fascicles, mostly dense, arising from stromata, emerging through stomata, erect, straight, subcylindrical-conical, not or only slightly geniculate, unbranched, 5–15 × 1.5–3 µm, aseptate, pale olivaceous or brownish, thin-walled, smooth; conidiophores reduced to conidiogenous cells, conidiogenous loci minute but conspicuous, 1 µm diam, somewhat thickened and darkened. Conidia solitary, narrowly obclavate to almost linear, straight to somewhat curved, (20−)30–70 × 1.5–2.5(−3) µm, indistinctly 2–6-septate, hyaline, thin-walled, smooth, apex pointed, gradually narrowed towards the obconically truncate base, 1 µm wide, hila slightly thickened and darkened.

Lectotype (designated here, MycoBank MBT178146): USA: Wisconsin: Barron County, Brill, on Eleocharis palustris, 23 Jul. 1928, J. J. Davis (BPI 436100). Isolectotypes: CUP 39751, WIS.

Host range and distribution: On Eleocharis (acicularis, compressa, elliptica, kuroguwai, obtusa, palustris, Eleocharis sp.), Cyperaceae, Asia (Japan), North America (USA, Idaho, Louisiana, Wisconsin).

Notes: A true Cercospora s. str. quite distinct from C. apii s. lat. by having very short and narrow conidiophores, 5–15 × 2–3 µm, minute but distinct conidiogenous loci, 1 µm diam. Conidia are narrowly obclavate-fusiform to filiform, 20–70 × 1.5–2.5 µm.

Cercospora glauciana Viégas, Bol. Soc. Bras. Agron. 8: 27 (1945).

(Fig. 86)

Fig. 86
figure86

Cercospora glauciana (PDD 46385). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 192), Braun & Sivapalan (1999: 5), Crous & Braun (2003: 198), Kirschner & Piepenbring (2006: 214).

Illustrations: Braun & Sivapalan (1999: 4, fig. 2), Kirschner & Piepenbring (2006: 214, fig. 7).

Description: Leaf spots amphigenous, almost indistinct, diffuse discolorations to subcircular, elliptical or fusiform, oblong, 1–10 × 0.5–5 mm, sometimes larger, to 20 mm diam, dull brown, greyish brown to blackish brown, later with pale centre, greyish white, surrounded by a darker margin. Caespituli hypophyllous, fine, not very conspicuous. Mycelium internal; hyphae sparingly branched, septate, pale brown, 1–3 µm wide, thin-walled, smooth. Stromata almost lacking or small, substomatal aggregations of swollen hyphal cells in the substomatal cavity, 10–35 µm diam, brown, cells 2–5 µm diam. Conidiophores in small fascicles, 1–12, divergent to moderately dense, arising from internal hyphae or stromata, through stomata, erect, cylindrical-filiform, straight to somewhat curved, not or only slightly geniculate-sinuous near the apex, unbranched, occasionally torulose, with swellings, 30–200 × 3–5 µm, swellings occasionally to 8 µm wide, 1–10-septate, medium to dark olivaceous-brown or medium brown, paler towards the tip, wall somewhat thickened below and thin towards the tip, smooth or almost so; conidiogenous cells integrated, terminal and intercalary, 10–35 µm long, sympodially and sometimes also percurrently proliferating, conidiogenous loci solitary or several, sometimes aggregated, thickened and darkened, minute, about 1 µm diam. Conidia solitary, cylindrical-fusiform, (12−)15–35 × 2–4 µm, 1–3-septate, hyaline or subhyaline, thin-walled, smooth, apex obtuse to attenuated, base short obconically truncate, 1–1.5 µm wide, hila somewhat thickened and darkened.

Holotype: Brazil: São Paulo: Campinas, Faz. Staz. Elisa, on Bulbostylis major, 18 Feb. 1943, A. P. Viégas (IACM 4141).

Host range and distribution: On Bulbostylis major, Rhynchospora sp., Scleria (ciliaris, lithosperma, Scleria sp.), unidentified Cyperaceae, Cyperaceae, Asia (Brunei), Oceania (Fiji, Micronesia, Palau, Solomon Islands, Vanuatu), Central and South America (Brazil, Panama).

Note: A true Cercospora s. str. quite distinct from C. apii s. lat. by its short and narrowly subcylindrical-fusiform conidia.

Cercospora kyllingae J.M. Yen & Gilles, Cah. Maboké 8: 81 (1970).

(Fig. 87)

Fig. 87
figure87

Cercospora kyllingae (based on Yen & Gilles 1970: 80, fig. 4). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 240).

Illustration: Yen & Gilles (1970: 80, fig. 4).

Description: Leaf spots lacking. Caespituli hypophyllous, indistinct. Mycelium internal. Stromata lacking. Conidiophores solitary or in small divergent or somewhat denser fascicles, 2–5, arising from internal hyphae, emerging through stomata, erect, straight to flexuous, subcylindrical or narrowed towards the apex, fertile part distinctly geniculate-sinuous, simple or often branched, 30–105 × 4–5 µm, 3–7-septate, pale brown to brown, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci conspicuous, thickened and darkened. Conidia solitary, acicular, 20–120 × 1.5–3 µm, 1–12-septate, hyaline, thin-walled, smooth, apex acute, base truncate or subtruncate, hila thickened and darkened.

Holotype: Gabon: Libreville, on Kyllinga erecta, Cyperaceae, 22 Jan. 1970, G. Gilles (“Par.Gab. no. 41”) (not traced).

Host range and distribution: Only known from the type collection, which we have not traced.

Notes: This species with its acicular conidia undoubtedly belongs to the C. apii s. lat. complex.

Cercospora kyllingicola J.M. Yen & Gilles, Cah. Maboké 8: 83. 1970.

(Fig. 88)

Fig. 88
figure88

Cercospora kyllingicola (based on Yen & Gilles 1970: 82, fig. 5). A. Conidiophore fascicle and conidiophore. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 240).

Illustration: Yen & Gilles (1970: 82, fig. 5).

Description: Leaf spots lacking. Caespituli hypophyllous, indistinct. Mycelium internal. Stromata lacking. Conidiophores solitary or in small divergent fascicles, arising from internal hyphae, emerging through stomata, erect, straight to slightly flexuous, 0–2 times geniculate, unbranched, 35–120 × 4–5 µm, 1–6-septate, pale brown to brown, paler towards the tip, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci conspicuous, thickened and darkened.

Conidia solitary, acicular, straight to somewhat curved, 60–250 × 3–4 µm, 8–21-septate, hyaline, thin-walled, smooth, apex pointed, base truncate, hila thickened and darkened.

Holotype: Gabon: Libreville, on Kyllinga erecta, Cyperaceae, 22 Jan. 1970, G. Gilles (“Par.Gab. no. 41”) (not traced).

Host range and distribution: Only known from the type collection which we have not traced.

Notes: This species with its acicular conidia does undoubtedly belong to the C. apii s. lat. complex. It differs from C. kyllingae by its unbranched conidiophores and longer, broader conidia. However, C. kyllingae and C. kyllingicola occur both on Kyllinga erecta, are from the same locality and belong to the C. apii complex. If the described morphological differences between the two species are sufficient to keep two separate species is uncertain, somewhat doubtful and requires further collections and examinations.

Cercospora uredinophila (Sacc.) Deighton, Mycol. Pap. 118: 40 (1969).

(Fig. 89)

Fig. 89
figure89

Cercospora uredinophila (PAD, holotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercosporella uredinophila Sacc., Ann. Mycol. 12: 312 (1914).

Synonym: Cercosporella scirpina Davis, Trans. Wisconsin Acad. Sci. 18: 266 (1915) [lectotype (designated here, MycoBank MBT178147: USA: Wisconsin: St. Croix Falls, on Scirpus pedicellatus, 25 Aug. 1914, J. J. Davis (BPI 420940); isolectotype: WIS].

Literature: Saccardo (1931: 744), Braun (1993a: 236; 1995: 122), Crous & Braun (2003: 416).

Illustrations: Deighton (1969: 40, fig. 23), Braun (1993a: 237, fig. 3; 1995: 123, fig. 107).

Exsiccatae: Syd., Fungi Exot. Exs. 444.

Description: Leaf spots amphigenous, subcircular, elliptical, oblong, irregular, 1–4 mm diam, scattered to confluent, pale brown, margin indefinite or darker. Caespituli amphigenous, punctiform, whitish, pale pink to brownish, not very conspicuous. Mycelium internal; hyphae branched, septate, 1–3 µm wide, hyaline or subhyaline. Stromata small, substomatal, 10–30 µm diam, subhyaline, yellowish to pale olivaceous, composed of subhyaline to yellowish swollen hyphal cells, 2–5 µm diam. Conidiophores in small to usually moderately large fascicles, about 8–20, divergent to dense, arising from stromata, through stomata, erect, straight, subcylindrical, unbranched, somewhat attenuated towards the tip to moderately geniculate-sinuous, unbranched, 20–35(−60) × 2–6(−7) µm, aseptate, almost hyaline to pale yellowish olivaceous, thin-walled, smooth; conidiophores reduced to conidiogenous cells, conidiogenous loci conspicuously thickened and darkened, planate to subdenticulate, about 1.5–2 µm diam. Conidia formed singly, acicular, filiform, narrowly obclavate, straight to somewhat curved or sinuous, (35−)40–80(−140) × 2–3 µm, mostly (2−)3–7(−12)-septate, hyaline, thin-walled, smooth, apex acute to subobtuse, base truncate or short obconically truncate, 1.5–2.5 µm wide, hilum thickened and darkened.

Holotype: Philippines: Manila, on Scirpus grossus, 29 Mar. 1913, P. W. Graff 170 (PAD). Isotypes: Syd., Fungi Exot. Exs. 444, e.g. BPI 420980, CUP, K(M) IMI 16432a, MICH 15397.

Host range and distribution: On Actinoscirpus grossus [Scirpus grossus], Scirpus (cyperinus, pedicellatus, Scirpus sp.), Cyperaceae, Asia (Malaysia, Philippines), North America (USA, Wisconsin).

Doubtful, excluded and insufficiently known species

Cercospora crinospora G.F. Atk., J. Elisha Mitchell Sci. Soc. 8: 58 (1892).

Synonym: Dactylaria crinospora (G.F. Atk.) U. Braun & Crous, in Crous & Braun, Mycosphaerella and Anam.: 143 (2003).

Literature: Saccardo (1892: 655), Chupp (1954: 191).

Holotype: USA: Alabama: Auburn, on Rhynchospora glomerata, Cyperaceae, 27 Aug. 1891, B. M. Duggar (CUP-A 2034).

Host range and distribution: Only known from the type collection.

Notes: This species was placed in Dactylaria sect. Mirandina (sensu de Hoog 1985). The conidiophores are formed in small fascicles, 15–60 × 2–3.5 µm, pale to medium olivaceous-brown, septate, wall somewhat thickened; conidiogenous cells integrated, terminal, denticulate. The conidia are solitary, cylindrical-filiform to subfusiform, 15–60 × 1–2.5 µm, obscurely 1–4-septate, hyaline, smooth. This species is morphologically close to D. congesta (conidiophores subhyaline) and D. irregularis (isolated from bark, conidiophores shorter and wider).

Cercospora cyperi-fusci Sandu, Lucr. Şti. Inst. Agron. ‘Prof. Ion Ionescu de la Brad’ 1960: 384 (1960).

Literature: Crous & Braun (2003: 151).

Description: Lesions necrotic, brown, indeterminate, covering almost entire leaves. Conidiophores fasciculate, 4–12, rarely solitary, emerging through stomata, erect, geniculate-sinuous (“nodulose”), unbranched, 20–45 × 3.5–4 µm, aseptate or with a single septum, brown, paler towards the tip. Conidia solitary, acicular, 90–110 × 4 µm, pluriseptate, hyaline, apex acute, base truncate.

Holotype: Romania: Cîrjoaia, Zbereni forest near Cotnari, on Cyperus fuscus, 30 Jul. 1957, C. Sandu-Ville (not traced).

Host range and distribution: On Cyperus fuscus, Cyperaceae, Europe (Romania).

Notes: It was not possible to get type material or any other collections of this species. It is unclear and unknown to us if and where type material has been deposited. According to the original description and illustration (Sandu-Ville et al. 1960), the conidia of C. cyperi-fusci are strictly acicular placing this species in the C. apii s. lat. complex, i.e. it does not belong to C. cyperi which is common and widespread on Cyperus spp.

Cercospora scirpi Zaprom., Bolezni Rast. 12: 92 (1923). Synonyms: Cercosporella scirpi (Zaprom.) Karak., in Vassiljevsky & Karakulin, Fungi Imperfecti Parasitici (Hyphomycetes) 1: 173 (1937).

Cercosporella scirpi Moesz, Magyar Biol. Kutatóint. Munkái 1: 114 (1930) [holotype: Hungary: Hévizfürdö, on Schoenoplectus litoralis, 10 Sep. 1927, Moesz (BP); isotype: B].

Pseudocercosporella scirpi (Moesz) Deighton, Mycol. Pap. 133: 55 (1973).

Literature: Chupp (1954: 192), Braun (1995: 160; 1998a: 404), Braun & Mel’nik (1997: 91).

Holotype: Uzbekistan: Tashkent Gub., on Schoenoplectus lacustris, 18 Jul. 1918, Zaprometov (LE 159401).

Host range and distribution: On Eleocharis palustris, Schoenoplectus (lacustris, litoralis), Scirpus acutus, Cyperaceae, Asia (Uzbekistan), Europe (Germany, Great Britain, Hungary), North America (USA, Wisconsin).

Notes: Based on the original description, Braun (1995: 122) interpreted the name Cercospora scirpi as synonym of C. uredinicola, but later Braun (1998: 404) traced and examined type material of C. scirpi and reduced it to synonymy with Pseudocercosporella scirpi.

Cercospora scirpicola (Fuckel) Zind.-Bakker, Rev. Mycol. 5: 64 (1940).

Basionym: Sporidesmium scirpicola Fuckel, Fungi Rhen. Exs., no. 78 (1863).

Synonyms: Clasterosporium scirpicola (Fuckel) Sacc., Syll. Fung. 4: 393 (1886).

Alternaria scirpicola (Fuckel) Sivan., Bitunicate Ascomycetes and their Anamorphs: 526 (1984).

Nimbya scirpicola (Fuckel) E. Simmons, Sydowia 41: 318 (1989).

Sphaeria scirpicola DC., Fl. franç., edn. 3, 2: 300 (1805) : Fr., Syst. Mycol. 2: 510 (1823).

Macrospora scirpicola (DC. : Fr.) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 140 “1869” (1870). Pleospora scirpicola (DC. : Fr.) P. Karst., Bidrag Kännedom Finlands Natur Folk 19: 72 (1871).

Clathrospora scirpicola (DC. : Fr.) Höhn., Ann. Mycol. 18: 77 (1920).

Pyrenophora scirpicola (DC. : Fr.) E. Müll., Sydowia 5: 256 (1951).

Sphaeria scirpi Rabenh., Deutschl. Krypt.-Fl. 1 (Pilze): 170 (1844), nom. superfl.

Pleospora scirpi (Rabenh.) Ces. & De Not., Comment. Soc.

Crittog. Ital. 1: 217 (1863).

Pyrenophora scirpi (Rabenh.) Wehmeyer, World Monograph of the Genus Pleospora and its Segregates: 287 (1961).

Pleospora eleocharidis Plowr., Fungi of Norfolk: 18 (1884).

Pleospora palustris Berl., Nuovo Giorn. Bot. Ital. 20: 67 (1888).

Literature: Chupp (1954: 193), Ellis (1976: 421), Crous & Braun (2003: 369), Woudenberg et al. (2013: 198, current taxonomy and phylogeny).

Syntypes: Germany: Hattenheim, Fuckel, Fungi Rhen. Exs. 78 (e.g., FH, G, HAL).

Host range and distribution: On Eleocharis, Eriophorum, Festuca, Juncus, Scirpus (s. lat.) spp., Cyperaceae, Juncaceae, Poaceae, Europe and North America.

Notes: Sphaeria scirpi is a superfluous name since Rabenhorst (l. c.) cited in his original description the valid name Sphaeria scirpicola DC. as synonym. Rabenhorst also referred to ‘Fries, Elench. II: 108’ and ‘Syst. II: 510’, but in these works Fries only cited S. scirpicola DC.

Passalora

A single species.

Passalora cyperi (Durgas Gupta, Padhi & Chowdhry) U. Braun & Crous, in Crous & Braun, Mycosphaerella and Anam.: 450 (2003).

Basionym: Phaeoramularia cyperi Durgas Gupta, Padhy & Chowdhry, Curr. Sci. 50: 140 (1981).

Literature: Singh (2003: 527), Kamal (2010: 116).

Illustration: Gupta et al. (1981: 140, fig. 1).

Description: Leaf spots amphigenous, circular to elongated, 0.5–2 mm diam, deep brown. Caespituli hypophyllous, effuse, brown. Mycelium internal. Stromata pseudoparenchymatous, globose, small, 15–18 µm diam, medium brown. Conidiophores fasciculate, arising from stromata, about 7–9 per fascicle, erect, straight, subcylindrical or occasionally geniculate, rarely branched, 45–90 × 3.5–6(−7) µm, 3–10-septate, olivaceous-brown, smooth; conidiogenous cells integrated, terminal, conidiogenous loci conspicuous, thickened and darkened. Conidia solitary to catenate, in branched chains, cylindrical to ellipsoid-fusiform, 4–22 × 2–5 µm, 0–1-septate, pale olivaceous-brown, ends rounded, with thickened and darkened hila.

Holotype: India: Odisha: Bhubaneswar, on Cyperus alternifolius, Cyperaceae, 22 Nov. 1977, D. Gupta (HCIO 32895).

Host range and distribution: Only known from the type collection.

Notes: This species is only tentatively maintained in Passalora (including Phaeoramularia). The conidia are rather cladosporioid, but details of the structure of the conidiogenous loci and conidial hila are unknown. Type material was not available for re-examination.

Zasmidium

Key to Zasmidium species on Cyperaceae
  1. 1

    Stromata present, 10–30 µm diam; conidiogenous loci and conidial hila 2–3 µm wide; conidia 20–300 × 3–5 µm, pluriseptate (with to 30 septa); on Gahnia ........................................ Z. gahniae

    Stromata lacking; conidiogenous loci and conidial hila narrower, 1–1.5 µm diam; conidia shorter and above all narrower, 20–50(−70) × 2.5–3.5(−4) µm, only (0−)1–3-septate; on Scleria ........................................ Z. scleriae

Zasmidium species on Cyperaceae

Zasmidium gahniae (McKenzie) U. Braun & McKenzie, comb. nov.

MycoBank MB809039

(Fig. 90)

Fig. 90
figure90

Zasmidium gahniae (PDD 38658). A. Solitary conidiophore arising from superficial hypha. B. Conidiophore. C. Conidia. Bar = 10 µm.

Basionym: Stenella gahniae McKenzie, New Zealand J. Bot. 20: 248 (1982).

Illustration: McKenzie (1982: 249–250, figs 4–5).

Description: Colonies epiphyllous. Mycelium internal and external; superficial hyphae branched, septate, 1.5–2.5 µm wide, pale straw-coloured, thin-walled, minutely verruculose. Stromata absent. Conidiophores solitary, arising from superficial hyphae, lateral, originating from brown, smooth, sometimes swollen hyphal cells, 10–30 µm wide, erect, straight or flexuous, occasionally 1–2 times geniculate, unbranched, to 150 µm long and 3–4.5 µm wide, pluriseptate, brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, sympodial, conidiogenous loci conspicuous, numerous, slightly prominent, thickened and darkened, 2–3 µm wide. Conidia solitary, straight or almost so, cylindrical to narrowly obclavate, 20–300 × 3–5 µm, pluriseptate (to 30 septa), straw-coloured, thin-walled, minutely verruculose, apex obtuse, rounded, base truncate to short obconically truncate, 2–3 µm wide, hila thickened and darkened.

Holotype: New Zealand: Auckland Domain, on Gahnia setifolia, Cyperaceae, 28 Sep. 1978, W. S. M. Versluys & E. H. C. McKenzie (PDD 38658).

Host range and distribution: On Gahnia (lacera, setifolia), Cyperaceae, New Zealand.

Notes: This species is characterised by having scolecosporous (cercosporoid), pluriseptate conidia formed singly, with planate conidiogenous loci and hila. Such cercosporoid former Stenella species belong to the Mycosphaerellaceae and have to be reallocated to Zasmidium.

Zasmidium scleriae (McKenzie) U. Braun & McKenzie, comb. nov.

MycoBank MB809040

(Fig. 91)

Fig. 91
figure91

Zasmidium scleriae (PDD 40911). A. Solitary conidiophores arising from superficial hypha. B. Conidiophore. C. Conidia. Bar = 10 µm.

Basionym: Stenella scleriae McKenzie, New Zealand J. Bot. 20: 250 (1982).

Illustration: McKenzie (1982: 251, fig. 6).

Description: Colonies amphigenous, mainly epiphyllous. Mycelium internal and external; superficial hyphae, branched, septate, 1.5–5 µm wide, pale straw-coloured, thin-walled, minutely verruculose. Stromata absent. Conidiophores solitary, arising from superficial hyphae, lateral, erect, originating from brown, smooth, sometimes swollen cells, 5–10 µm diam, straight to flexuous or somewhat geniculate, unbranched, to 210 µm long and 2.5–4(−4.5) µm wide, pluriseptate, brown, paler towards the tip; conidiogenous cells integrated, terminal or intercalary, sympodial, conidiogenous loci thickened and darkened, slightly prominent, 1–1.5 µm wide. Conidia solitary, straight or curved to sinuous, cylindrical to narrowly obclavate, 20–50(−70) × 2.5–3.5(−4) µm, indistinctly (0−)1–3-septate, straw-coloured, thin-walled, minutely verruculose, apex obtuse, rounded, base truncate to short obconically truncate, 1–1.5 µm wide, hila somewhat thickened and darkened.

Holotype: Solomon Islands: Kolombangara, on Scleria polycarpa, Cyperaceae, 12 Jul. 1980, E. H. C. McKenzie (PDD 40911).

Host range and distribution: Only known from the type collection.

Notes: See Z. gahniae.

Dioscoreaceae

Cercospora

Key to Cercospora species on Dioscoreaceae
  1. 1

    Conidiophores very short, 7–45 × 3–7 µm, 0–1-septate; on Dioscorea pyrenaica, Europe ............................. C. aragonensis

    Conidiophores much larger, (10−)40–320 µm, pluriseptate; on Dioscorea spp., Africa, Asia ............................. 2

  2. 2(1)

    Stromata well-developed, large, about 30–65 µm diam; conidia strictly acicular; on Dioscorea oppositifolia, China ........................................ C. cantoniensis

    Stromata lacking or small, 10–40 µm diam; conidia acicular, shorter ones obclavate-cylindrical; on Dioscorea spp. ........................................ C. dioscoreae-pyrifoliae

Cercospora species on Dioscoreaceae

Cercospora aragonensis Durrieu, Bull. Trimestriel Soc. Mycol. France 80: 169 (1964).

(Fig. 92)

Fig. 92
figure92

Cercospora aragonensis (K(M) IMI 255661). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Pons & Sutton (1988: 15), Crous & Braun (2003: 63).

Illustration: Pons & Sutton (1988: 14, fig. 5).

Description: Leaf spots yellowish to brownish, necrotic, at the leaf edge, extending to the petioles, to 10 mm diam. Caespituli amphigenous. Mycelium internal. Stromata substomatal, 20–60 µm diam, pseudoparenchymatic, brown. Conidiophores fasciculate, to 20, arising from stromata, through stomata, short, subcylindrical-conical or only slighty geniculate-sinuous, unbranched, 7–45 × 3–7 µm, 0–1-septate, pale olivaceous, thin-walled, smooth; conidiophores reduced to conidiogenous cell or occasionally integrated, terminal, sympodial and occasionally percurrent, conidiogenous loci conspicuous, somewhat thickened and darkened, about 2.5–3 µm diam. Conidia solitary, acicular to narrowly obclavate, straight to curved, 65–155 × 3–4 µm, 2–12-septate, hyaline, thin-walled, smooth, apex pointed, base truncate to somewhat obconically truncate (gradually attenuated towards the base), about 2–3 µm wide, hila somewhat thickened and darkened.

Holotype: Spain: Prov. Hueasca: Pyrenees, Mt Tourbón, on Dioscorea pyrenaica, 25 Jul. 1963, G. Durrieu (TLA 1828).

Isotype: K(M) IMI 255661 (slide).

Host range and distribution: On Dioscorea pyrenaica Dioscoreaceae, Europe (Spain).

Notes: A true Cercospora s. str. distinct from C. apii s. lat. by its very short conidiophores and narrowly obclavate to acicular conidia. However, this species is only known from the type collection. New collections are necessary to establish the morphological variability of conidiophores and conidia. As already mentioned by Sutton & Pons (1988), it cannot be excluded that C. aragonensis was based on immature material.

Cercospora cantonensis P.K. Chi, Fungal Diseases of Cultivated Medical Plants in GuangdongPrivince: 84 (1994); also in J. S. China Agric. Univ. 15: 14 (1994)

(Similar to Fig. 1)

Literature: Guo et al. (2005: 107).

Illustrations: Chi (1994: 84, fig. 72), Guo et al. (2005: 107, fig. 72).

Description: Leaf spots amphigenous, circular, elliptical to irregular, greyish white with brown border. Caespituli amphigenous. Mycelium internal. Stromata well-developed, large, about 30–65 µm diam, globose, olivaceous-brown. Conidiophores fasciculate, 5–18, divergent to dense, arising from stromata, erect, straight, subcylindrical to distinctly geniculate, to six times, unbranched, about 40–325 × 3–6.5 µm, usually 3–13-septate, olivaceous-brown; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci distinct, thickened and darkened. Conidia solitary, acicular, about 30–255 × 2–4 µm, 5–30-septate, hyaline, thin-walled, smooth, apex pointed, base truncate, with thickened and darkened hilum.

Holotype: China: Prov. Guangdong: Luoding, on Dioscorea oppositifolia [opposita], Jul. 1986, S. Q. Chen 341 (Hb. South China Agric. Univ., Guangzhou).

Host range and distribution: On Dioscorea (gracillima, oppositifolia), Dioscoreaceae, Asia (China).

Notes: Crous & Braun (2003) reduced C. cantoniensis to synonymy with C. dioscoreae-pyrifoliae, but we prefer to maintain it tentatively as a separate species based on differences in the formation of stromata. Cercospora cantoniensis was described to have well-developed stromata, about 30–65 µm diam, whereas Cercospora collections on other Dioscorea spp., referred to as C. dioscoreae-pyrifoliae, are characterised by having small or no stromata. Guo et al. (2005) maintained C. cantoniensis as a separate species based on a comparison with the original description of C. dioscoreae-pyrifoliae which is, however, misleading with regard to the given conidial size (see notes under the latter species).

Cercospora dioscoreae-pyrifoliae J.M. Yen, Bull. Trimestriel Soc. Mycol. France 84: 5 (1968); as “dioscorae-pyrifoliae”.

(Fig. 93)

Fig. 93
figure93

Cercospora dioscoreae-pyrifoliae (PC, holotype). A. Conidiophore fascicle. B. Conidiophore. C. Conidiophore tip. D. Conidia. Bar = 10 µm.

Synonyms: Cercospora pachyderma var. indica Munjal, Lall & Chona, Indian Phytopathol. 14: 187 (1961) [holotype: India: Uttarakhand: Jeolikote, Kumaon, on Dioscorea sp., 22 Oct. 1959, J. N. Kapoor (HCIO 26873); isotype: K(M) IMI 256769].

? Mycosphaerella papuana Sivan., Trans. Brit. Mycol. Soc. 85: 743 (1985) [holotype: Papua New Guinea: Nembi Valley, on Dioscorea sp., 17 Sep. 1984, J. M. Waller 2144 (K(M) IMI 293049)].

Literature: Yen & Lim (1980: 159), Pons & Sutton (1988: 7–15, as C. apii), Crous & Braun (2003: 162), Aptroot (2006: 149), Kamal (2010: 41), Nakashima et al. (2011).

Illustrations: Yen (1968: 7, fig. 1), Yen & Lim (1980: 212, fig. 15), Sivanesan (1985: 744–746, figs 1–3), Pons & Sutton (1988: 8–11, figs 1–3), Nakashima et al. (2011: 256, fig. 1c,d).

Description: Leaf spots amphigenous, subcircular, elliptical to angular-irregular, 0.5–10 mm diam, at first brown, later with pale centre, finally greyish white with dark border, brown, and sometimes yellowish halo. Caespituli amphigenous, punctiform, dark, scattered. Mycelium internal. Stromata lacking or almost so to small, 10–40 µm diam, substomatal to intraepidermal, brown. Conidiophores in small to moderately large fascicles (usually 2–30), divergent to moderately dense, arising from internal hyphae or stromata, emerging through stomata or erumpent, straight to distinctly geniculate-sinuous, unbranched, (10−)40–200 × 3–7 µm, usually 0–8-septate, pale to dark olivaceous-brown or brown, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal and intercalary, rarely conidiophores reduced to conidiogenous cells, 10–40 µm long, sympodial, rarely percurrent, conidiogenous loci 2–4 µm diam. Conidia solitary, acicular, shorter conidia sometimes obclavate-subcylindrical, 30–185 × 2–4 µm, rarely longer, 3–18-septate, hyaline, thin-walled, smooth, apex pointed, base truncate or only slightly attenuated at the very base, 1.5–3.5 µm wide, hila thickened and darkened.

Holotype: Singapore: Jurong, on Dioscorea pyrifolia, 17 Dec. 1965, J. M. Yen 738 (PC). Isotype: K(M) IMI 255659.

Host range and distribution: On Dioscorea (bulbifera, cayennensis subsp. rotundata [rotundata], deltoidea, esculenta, pyrifolia, sylvatica, tokoro, villosa [glauca], Dioscorea sp.), Dioscoreaceae, Africa (Ghana, Nigeria, Tanzania), Asia (Borneo, India, Japan, Malaysia, Papua New Guinea, Singapore).

Notes: Cercospora dioscoreae-pyrifoliae can currently only be circumscribed and used as a heterogeneous morpho-taxon. This species is morphologically part of the C. apii s. lat. complex, but phylogenetically all examined isolates derived from Dioscorea spp. are distinct from C. apii s. str. However, in comprehensive phylogenetic studies on Cercospora s. str., sequences retrieved from Dioscorea isolates clustered in four clades, which represent different plurivorous Cercospora species. Currently it is quite unclear to which group the name C. dioscoreae-pyrifoliae pertains. Yen (1968) described relatively broad conidia, 3.5–6 µm wide, which is, however, not in agreement with the conidial size found in type material, which is narrower, about 2–4 µm (see also Pons & Sutton 1988).

The cited asexual/sexual connection has not been experimentally proven, i.e. it is unclear if Mycosphaerella papuana represents the genuine sexual morph of C. dioscoreae-pyrifoliae. Mycosphaerella papuana was described as follows (Sivanesan 1985): “Ascomata often mixed with the anamorph, amphigenous, mainly epiphyllous, solitary, rarely aggregated, immersed to somewhat erumpent, globose, sometimes immersed in stromata giving rise to conidiophores, 85–110 µm diam, peridium to 20 µm thick, composed of 4–6 layers of brown, polygonal, pseudoparenchymatous cells. Asci saccate-cylindrical, apex rounded, 34–46 × 7–10 µm, colourless, aparaphysate, 8-spored; ascospores narrowly obovate, 10–16 × 2–4.5 µm, to 18 µm long and subhyaline when discharged, with a single median septum, hyaline, ends acute, guttulate when young”.

Doubtful, excluded and insufficiently known species

Cercospora golaghati Saikia & A. Sarbhoy, Curr. Sci. 49: 830 (1980).

Literature: Pons & Sutton (1988: 70), Crous & Braun (2003: 201), Kamal (2010: 47).

Illustration: Saikia & Sarbhoy (1980: 830, fig. 1).

Description: Leaf spots apical and marginal, yellowish to brown. Caespituli hypophyllous, effuse. Mycelium internal. Stromata small, to 25 µm diam, composed of pale to dark brown cells. Conidiophores fasciculate, 2–7, arising from stromata, divergent, erect, straight or flexuous, geniculate, unbranched, (60−)80–100(−147) × 4–5.5 µm, 2–6-septate, mid golden brown, olivaceous or greyish brown; conidiogenous cells integrated, terminal, with thickened and darkened conidiogenous loci. Conidia solitary, obclavate-cylindrical, straight to slightly curved, about (35−)50–60(−80) × 4–6 µm, 3–7-septate, pale orange brown, thin-walled, smooth, apex conical, base obconically truncate with thickened and darkened hilum.

Holotype: India: Assam: Golaghat, on Dioscorea alata, Dioscoreaceae, 21 Nov. 1977, U. N. Saikia (HCIO 32660).

Host range and distribution: On Dioscorea alata, Dioscoreaceae, Asia (India).

Notes: An attempt to obtain type material of this species on loan from HCIO was not successful. The generic affinity of this species is unclear. Conidia were described as “pale orange brown” which would be in favour of Passalora, above all in combination with obclavate conidia.

Distocercospora

Key to Distocercospora species on Dioscoreaceae
  1. 1

    Conidiophores very long and usually multibranched and pluriseptate, 50–490(−625) × 3–6.5 µm; conidia with thickened inner wall layer, usually finely verruculose; on numerous Dioscorea spp. ........................................ D. pachyderma

    Conidiophores much shorter, 15–80 × 3–10 µm, shorterones aseptate, longer ones only 1–2(−3)-septate; conidia thin-walled; on Dioscorea sp., South Africa ........................... D. africana

Distocercospora species on Dioscoreaceae

Distocercospora africana Crous & U. Braun, Sydowia 46: 208 (1994).

(Fig. 94)

Fig. 94
figure94

Distocercospora africana (PREM 10125). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Literature: Crous & Braun (1996: 247).

Illustration: Crous & Braun (1994: 209, fig. 3).

Description: Leaf spots amphigenous, scattered, subcircular to angular, vein-limited, 8–20 mm diam. Caespituli hypophyllous, medium brown, punctiform, 40–80 × 30–40 µm. Mycelium internal; hyphae branched, septate, 1–2.5 µm diam. Stromata well-developed, substomatal, erumpent, large stromata rupturing the stomata, 15–50 µm wide and 10–40 µm high, medium brown. Conidiophores in small to moderately large fascicles, loose to dense, arising from stromata, through stomata, straight and subcylindrical to moderately geniculate-sinuous, unbranched or occasionally branched, 15–80 × 3–10 µm, shorter ones aseptate, longer ones 1–2(−3)-septate, olivaceous to brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, about 15–30 µm long, sympodial, conidiogenous loci conspicuous, somewhat thickened and darkened. Conidia solitary, subacicular to obclavate (-subcylindrical), straight to curved, 30–110 × 3–5 µm, 1–5(−6)-distoseptate, subhyaline to olivaceous, thin-walled, smooth, apex obtuse to subacute, base short obconically truncate, hila slightly thickened and darkened.

Holotype: South Africa: Cape Province: Kentani, on living leaves of Dioscorea sp., Dioscoreaceae, 26 Apr. 1917, A. Pegleu (PREM 10125).

Host range and distribution: On Dioscorea (sylvatica, Dioscorea sp.), Dioscoreaceae, South Africa.

Distocercospora pachyderma (Syd. & P. Syd.) Pons & B. Sutton, Mycol. Pap. 160: 60 (1988).

(Fig. 95)

Fig. 95
figure95

Distocercospora pachyderma (S-F37683). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercospora pachyderma Syd. & P. Syd., Ann. Mycol. 12: 203 (1914).

Synonyms: Cercosporina pachyderma (Syd. & P. Syd.) Sacc., Syll. Fung. 25: 900 (1931).

Cercospora dioscoreae-bulbiferae J.M. Yen & Gilles, Cah. Maboké 9: 105 “1971” (1973) [holotype: Gabon: Libreville, 28 km on Kango Rd, on Dioscorea bulbifera, 21 Feb. 1971, G. Gilles PC 86 (K(M) IMI 216331)].

Literature: Saccardo (1931: 900), Chupp (1954: 197), Katsuki (1965: 28), Pons & Sutton (1988: 60), Guo (1999), Shin & Kim (2001: 119–121), Kirschner et al. (2004: 61–63), Kamal (2010: 277).

Illustrations: Yen & Gilles (1973: 104, fig. 2), Pons & Sutton (1988: 61–64, figs 28–31), Shin & Kim (2001: 120, fig. 51), Kirschner et al. (2004: 62, figs 10–13), Kamal (2010: 276, fig. 47).

Description: Leaf spots lacking or almost so or formed as indistinct to distinct yellowish to brown discolorations or spots, 2–10 mm diam, with diffuse to distinct margin, sometimes much darker, blackish grey to blackish. Caespituli amphigenous, effuse to discrete, velutinous, greyish, grey-olivaceous to darker olivaceous, brown or blackish brown. Mycelium internal; hyphae branched, straight to strongly sinuous, tortuose, 1.5–5 µm wide, subhyaline to medium brown, thin-walled, smooth. Stromata lacking or almost so to small, 10–30 µm diam, substomatal, subglobose to irregularly shaped, brown, composed of swollen hyphal cells, 3–6 µm diam, more or less rounded to mostly angular-irregular in outline. Conidiophores in mostly lax to moderately dense fascicles, about 5–30, arising from internal hyphae or small stromata, through stomata, erect, subcylindrical-filiform, flexuous to strongly geniculate-sinuous, sometimes subdenticulate in the upper fertile part, simple or branched, ranging from slightly branched to multibranched, 50–490((−625) × 3–6.5 µm, pluriseptate, individual cells about 15–45 µm long, pale brown, brown or olivaceous-brown, somewhat paler towards the apex, wall thin to somewhat thickened, to 0.75 µm, smooth to somewhat rough; conidiogenous cells integrated, terminal and intercalary, 15–55 µm long, conidiogenous loci conspicuous, 1–2 µm diam, unthickend or almost so to slighty thickened, not to somewhat darkened-refractive, not to somewhat prominent or even subdenticulate. Conidia solitary, rarely in short chains, obclavate-cylindrical, short conidia sometimes ellipsoid-ovoid to cylindrical-fusiform, straight to strongly curved orsigmoid, (15−)25–95(−140) × (1.5(−)3–6.5((−7) µm, (0(−)1–5–distoseptate, subhyaline to pale olivaceous or brownish, inner wall layer somewhat thickened, smooth or almost so to finely verruculose, apex obtuse, base subtruncate to short obconically truncate, 1–2 µm wide, hila unthickened to slightly thickened and not to slightly darkened-refractive.

[Holotype: Philippines: Prov. Laguna: Luzon, Los Baños, on Dioscorea alata, 10 Nov. 1913, M. B. Raimundo, C. F. Baker 2053 (not preserved)]. Neotype (designated here, MycoBank MBT178148): Philippines: Prov. Laguna: Luzon, Morong Valley, on Dioscorea alata, 9 Nov. 1913, M. B. Raimundo, C. F. Baker 2051 (S-F37683). Topotype material: Philippines: Prov. Laguna: Luzon, Los Baños, on Dioscorea alata, Nov. 1913, C. F. Baker 522 (B; BPI 439183–439184; K(M) IMI 256649, S-F37682). Epitype (designated here, MycoBank MBT178149): Japan: Iwate Pref.: Morioka, Koma, on Dioscorea sp., 10 Sep. 2013, C. Nakashima (MUMH11476). Duplicate: CBS H-21733; ex-epitype cultures: MUCC1716, CBS 138247.

Host range and distribution: On Dioscorea (alata, bulbifera, esculenta, glabra, japonica, oppositifolia, pentaphylla, polystachya [batatas], quartiniana [beccariana], sagittata, subcalva, tokoro, yunnanensis, Dioscoreaceae, Africa (Gabon, Sierra Leone, Uganda), Asia (China, India, Japan, Korea, Malaysia, Myanmar, Philippines), Oceania (American Samoa, Micronesia, Solomon Islands), West Indies (Barbardos, Trinidad and Tobago).

Notes: The structure of the conidiogenous loci and conidial hila is intermediate between corresponding structures in Passalora and Pseudocercospora and above all confusable with some Pseudocercospora species with more conspicuous, subdenticulate, refractive scars and hila. The abundant hypophyllous sporulation, and long, flexuous conidiophores with dispersed sympodial conidiogenous loci, and the distoseptate conidia (also prominent in culture), are definitive characters for this species.

Passalora

Key to Passalora species on Dioscoreaceae
  1. 1

    Superficial hyphae with solitary conidiophores in vivo present (mycovellosiella-like) ........................................ 2

    Superficial hyphae and solitary conidiophores in vivo lacking, conidiophores only fasciculate ........................................ 4

  2. 2(1)

    Conidiophores long, 40–120 µm; on Dioscorea alata, China ........................................ P. dioscoreigena

    Conidiophores shorter, 5–50 µm; on Dioscorea nipponica ........................................ 3

  3. 3(2)

    Conidiogenous loci 1–2 µm diam; conidia narrow, 15–90 × 2–5 µm, 0–3-septate;

    Far East of Russia and Japan ........................................ P. tranzschelii var. tranzschelii

    Conidiogenous loci 1.5–3 µm diam; conidia broader, 30–125 × 4–7 µm,

    3–10-septate; China ........................................ P. tranzschelii var. chinensis

  4. 4(1)

    Conidia catenate; on Dioscorea spp., widespread ........................................ P. dioscoreae

    Conidia formed singly; on Dioscorea spp., China ...................................................... 5

  5. 5(4)

    Conidiophores long, 35–175 µm, often branched; conidia very broad, 10–55 × 7.5–10 µm, 1–2-septate ........................................ P. dioscoreicola

    Conidiophores shorter, 25–65(−115) µm, usually unbranched; conidia much narrower, 30–85(−105) × 4.5–6.5 µm ........................................ P. dioscoreae-subcalvae

Passalora species on Dioscoreaceae

Passalora dioscoreae (Ellis & G. Martin) U. Braun & Crous, in Crous & Braun, Mycosphaerella and Anam.: 162 (2003).

(Fig. 96)

Fig. 96
figure96

Passalora dioscoreae (NY 838293). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercospora dioscoreae Ellis & G. Martin, Amer.

Naturalist 16: 1003 (1882).

Synonyms: Phaeoramularia dioscoreae (Ellis & G. Martin) Deighton, in Ellis, More Dematiaceous Hyphomycetes: 319 (1976).

Cercospora nubilosa Ellis & Everh., J. Mycol. 4: 115 (1888) [holotype: USA: Ohio: Cleveland, on Dioscorea sp., as “Smilax sp.”, 20 Aug. 1888, S. M. Tracy (NY 838195); isotypes: K(M) 255658, NY 838194].

Cercospora tokoroi Togashi, Bull. Imp. Coll. Agric. Forest. (Morioka) 22: 46 (1936) [lectotype (designated here, MycoBank MBT178150): Japan: Iwate Pref.: Mt Iwate, on Dioscorea tokoro, 15 Sep. 1934, K. Togashi (TNS-F-243948); isolectotype: K(M) IMI 166671].

Literature: Saccardo (1886: 479; 1892: 654), Vasudeva (1963: 98), Chupp (1954: 197), Katsuki (1965: 28), Ellis (1976: 319), Pons & Sutton (1988: 33), Guo et al. (2003: 140–141), Kamal (2010: 117).

Illustrations: Ellis (1976: 320, fig. 241 C), Pons & Sutton (1988: 34–36, figs 14–16), Guo et al. (2003: 141, fig. 88).

Exsiccatae: Ellis & Everh., North Amer. Fungi 2471.

Description: Leaf spots amphigenous, variable, subcircular, angular-irregular to diffuse, sometimes vein-limited, 2–45 mm diam, pale to dark brown or olivaceous to olivaceous-brown, purplish brown, border diffuse, yellowish to cream, sometimes with distinct darker margin. Caespituli amphigenous, mainly hypophyllous, punctiform, dark. Mycelium internal. Stromata substomatal, 15–40 µm diam, brown, composed of subhyaline to brown swollen hyphal cells. Conidiophores in small to moderately large fascicles, about 5–30, arising from stromata, through stomata, erect, straight, subcylindrical to somewhat geniculate-sinuous, unbranched, 8–60 × 3–6 µm, rarely longer, 0–4-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores sometimes reduced to conidiogenous cells, 10–35 µm long, conidiogenous loci conspicuous, thickened and darkened, 1.5–2.5 µm diam. Conidia solitary or catenate, in simple or occasionally branched chains, subcylindrical to obclavate-cylindrical, rarely subclavate, straight to curved, 15–120 × (2−)3–5(−5.5) µm, 1–8-septate, pale olivaceous, thin-walled, smooth to somewhat rough, apex obtuse, subobtuse to truncate, base short obconically truncate, about 2–2.5 µm wide, hila somewhat thickened and darkened.

Holotype: USA: Pennsylvania: Delaware Co., on Dioscorea villosa, 1 Aug. 1882, W. Trimble (NY 838293). Isotype: K(M) IMI 256891.

Host range and distribution: On Dioscorea (alata, bulbifera, cayennensis, composita, deltoidea, esculenta, floribunda, hispida, japonica, nipponica, polystachya [batatas], quinquelobata, sativa, subcalva, tokoro, ubi, villosa), Dioscoreaceae, Africa (Togo, Uganda), Asia (China, India, Indonesia, Japan, Korea, Philippines, Sri Lanka, Taiwan), Europe (Italy), North America (Canada; USA, Delaware, Iowa, Illinois, Indiana, Maryland, Michigan, North Carolina, Ohio, Pennsylvania, Washington, Wisconsin), Central and South America (Brazil, Guatemala, Panama, Venezuela), West Indies (Cuba, Trinidad and Tobago).

Passalora dioscoreae-subcalvae Y.L. Guo, Mycosystema 20: 301 (2001). (Fig. 97)

Fig. 97
figure97

Passalora dioscoreae-subcalvae (HMAS 78803). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Literature: Guo et al. (2003: 81), Crous & Braun (2003: 451).

Illustrations: Guo (2001b: 301, fig. 1), Guo et al. (2003: 82, fig. 50).

Description: Leaf spots amphigenous, subcircular to irregular, 3–12 mm diam, reddish brown or centre grey to yellowish brown, with dark brown border line, halo yellowish brown above and pale yellowish to greyish brown below. Caespituli hypophyllous. Mycelium internal. Stromata lacking or small, substomatal. Conidiophores in loose fascicles, 3–14, through stomata, erect, straight, subcylindrical to geniculate above, usually unbranched, 25–65(−115) × 4.5–6.5(−8.5) µm, 0–4-septate, pale olivaceous-brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, sympodial, conidiogenous loci conspicuous, thickened and darkened, about 2–3 µm diam. Conidia solitary, obclavate to obclavate-cylindrical, straight to curved, 30–85(−105) × 4.5–6.5 µm, 2–6-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex obtuse, base short obconically truncate, 2–3 µm wide, hila thickened and darkened.

Holotype: China: Guangxi: Shangsi, on Dioscorea subcalva, 27 Oct. 1957, L. W. Xu, no. 532 (HMAS 78803).

Host range and distribution: On Dioscorea (alata, subcalva), Dioscoreaceae, Asia (China, Guangxi).

Passalora dioscoreicola Y.L. Guo, Mycosystema 20: 302 (2001); as “dioscoreiicola”. (Fig. 98)

Fig. 98
figure98

Passalora dioscoreicola (HMAS 79141). A. Conidiophores. B. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 451).

Illustrations: Guo (2001b: 302, fig. 2), Guo et al. (2003: 83, fig.51.

Description: Leaf spots amphigenous, subcircular to irregular, 4–10 mm diam, greyish brown to brown, halo pale yellowish brown to greyish brown. Caespituli mainly hypophyllous. Mycelium internal. Stromata lacking or small, substomatal. Conidiophores in loose to dense fascicles, through stomata, erect, straight, subcylindrical to usually distinctly or strongly geniculate above, unbranched or branched, 35–175 × 4–6 µm, width often irregular, 1–6-septate, olivaceous to medium olivaceous-brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, sympodial, conidiogenous loci conspicuous, thickened and darkened, about 2–3 µm diam. Conidia solitary, obclavate, clavate, fusiform-ellipsoid, straight to slightly, curved, 10–55 × 7.5–10 µm, 1–2-septate, olivaceous, thin-walled, smooth, apex obtuse, broadly rounded, base rounded to short obconically truncate, about 2–3 µm wide, hila thickened and darkened.

Holotype: China: Yunnan: Menglun, on Dioscorea sp., 22 Oct. 1973, Y. C. Zong & X. J. Liu 133 (HMAS 79141).

Host range and distribution: On Dioscorea sp., Dioscoreaceae, Asia (China, Yunnan).

Passalora dioscoreigena U. Braun & Crous, in Crous & Braun, Mycosphaerella and Anam.: 451 (2003). (Fig. 99)

Fig. 99
figure99

Passalora dioscoreigena (HMAS 79139). A. Superficial hypha with solitary conidiophore. B. Conidiophore fascicle. C. Conidia. Bar = 10 µm.

Basionym: Mycovellosiella dioscoreicola Y.L. Guo, Mycosystema 21 : 21 (2002), non Passalora dioscoreicola Y.L. Guo, 2001.

Literature: Guo et al. (2003: 27–28).

Illustrations: Guo (2002: 18, fig. 2), Guo et al. (2003: 27, fig. 13).

Description: Leaf spots amphigenous, circular to irregular, 1–5 mm diam, often confluent, yellowish brown to reddish brown, margin indefinite, with pale olivaceous-brown halo on the upper leaf surface, effuse, grey, greyish brown to dark brown below. Caespituli hypophyllous. Mycelium internal and external; superficial hyphae emerging through stomata, branched, septate, 2.5–4 µm wide, thin-walled, pale olivaceous. Stromata lacking or small, substomatal, brown. Conidiophores loosely fasciculate, emerging through stomata, or solitary, arising from superficial hyphae, lateral, erect, straight to curved, subcylindrical to somewhat geniculate, simple, rarely branched, 40–120 × 5–8.5 µm, 1–5-septate, often with constrictions at septa, pale olivaceous-brown or olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, sympodial, conidiogenous loci conspicuous, 2–3 µm diam, thickened and darkened. Conidia catenate, in simple or branched chains, obclavate-cylindrical, 25–110 × 4.5–6.5 µm, 1–6-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth, apex obtuse to truncate, base short obconically truncate, hila thickened and darkened.

Holotype: China: Sichuan: Emeishan, on Dioscorea alata, 6 Oct. 1956, X. J. Liu (HMAS 79139).

Host range and distribution: On Dioscorea (alata, Dioscorea sp.), Dioscoreaceae, Asia (China, Sichuan).

Passalora tranzschelii (Vassiljevsky) U. Braun & Crous, in Crous & Braun, Mycosphaerella and Anam.: 473 (2003). var. tranzschelii

(Fig. 100)

Fig. 100
figure100

Passalora tranzschelii var. tranzschelii (LE 505). A. Superficial hypha with solitary conidiophore. B. Conidiophore fascicle. C. Conidiophore. D. Conidia. Bar = 10 µm.

Basionym: Ragnhildiana tranzschelii Vassiljevsky, in Vassiljewsky & Karakulin, Fungi Imperfecti Parasitici (Hyphomycetes) 1: 379 (1937).

Synonyms: Ragnhildiana dioscoreae Vassiljevsky, in Vassiljewsky & Karakulin, Fungi Imperfecti Parasitici (Hyphomycetes) 1 : 379 (1937), non Passalora dioscoreae (Ellis & G. Martin) U. Braun & Crous, 2003 [probable type: Russia: Far East, near Vladivostok, on Dioscorea nipponica [giraldii], 15 Aug. 1929, W. Tranzschel (LE 347)].

Mycovellosiella dioscoreae (Vassiljevsky) N. Pons & B. Sutton, Mycol. Pap. 160: 49 (1988).

Mycovellosiella deightonii Katsuki & Y. Harada, Rep. Tottori Mycol. Inst. 10: 566 (1973) [holotype: Japan: Aomori Pref.: Hirosaki, on Dioscorea nipponica, 15 Aug. 1971, Y.

Harada (HHUF4130); isotype: K(M) 165059].

Passalora dioscoreae-nipponicae Y.L. Guo, Mycosystema 30: 868 (2011).

Illustration: Pons & Sutton (1988: 46–48, figs 20–22).

Description: Leaf spots amphigenous, circular to angular-irregular, scattered to confluent, 3–35 mm diam, at first yellowish, later pale to darker brown, margin narrow, darker. Caespituli hypophyllous, effuse, greyish to grey-brown. Mycelium internal and external; superficial hyphae emerging through stomata, branched, 2–4 µm wide, hyaline to pale brown, thin-walled, smooth. Stromata absent or almost so, only with a few substomatal swollen hyphal cells. Conidiophores occasionally in small, loose groups or fascicles, to six, through stomata, but usually solitary, arising from superficial hyphae, lateral, erect to occasionally decumbent, straight, subcylindrical-conical to somewhat geniculate-sinuous, unbranched or rarely branched, 5–50 × 3–7 µm, 0–2-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores often reduced to conidiogenous cells, 5–40 µm long, sympodial, conidiogenous loci conspicuous, 1–2 µm diam, somewhat thickened and darkened. Conidia solitary as well as catenate in simple or sometimes branched chains, obclavate-cylindrical, short conidia sometimes short cylindrical to ellipsoid-ovoid, 15–90 × 2–5 µm, 0–3-septate, subhyaline to pale olivaceous, apex obtuse to truncate, base subtruncate to short obconically truncate, 1–2 µm wide, hila slightly thickened and darkened.

Holotype: Russia: Ussuria: Voroshilov, on Dioscorea nipponica [giraldii], 30 Jul. 1927, W. Tranzschel (LE 505).

Host range and distribution: On Dioscorea nipponica, Dioscoreaceae, Asia (Japan; Russia, Far East).

var. chinensis Y.L. Guo, var. nov. MycoBank MB809022

(Fig. 101)

Fig. 101
figure101

Passalora tranzschelii var. chinensis (HMAS 65914). A. Superficial hyphae with solitary conidiophores. B. Conidia. Bar = 10 µm.

Illustration: Guo (2011: 869, fig. 4).

Diagnosis: Distinguished from var. tranzschelii by its larger conidiogenous loci, 1.5–3 µm diam, as well as longer and above all broader conidia, 30–125 × 4–7 µm, with 3–10 septa.

Holotype: China: Hebei Province: Yu County, Xiaowutai Mountains, on Dioscorea nipponica, 29 Aug. 1990, Y. L. Guo 1293-a (HMAS 65914). Paratype: China: Hebei Province: Zhuolu County, Yangjiaping, 31 Aug. 1990, Y. L. Guo 1332 (HMAS 65915).

Note: Guo (2011) assigned the material of the fungus now described as var. chinensis to Passalora dioscoreae-nipponicae, a new name introduced for Ragnhildiana dioscoreae. However, type material of the latter species agrees with Passalora tranzschelii s. str. (now var. tranzschelii).

Doubtful, excluded and insufficiently known species

Cercosporidium dioscoreae S. Singh, Plant Parasitic Fungi of Gorakhpur [University of Gorakhpur, PhD thesis (1994)], not effectively published (ICN, Art. 30.8).

Synonym: Passalora dioscoreae Poonam Srivast., J. Living World 1: 115 (1994); as “(S. Singh) Poonam Srivast.” nom. inval. (ICN, Art. 39.1 and 40.1).

Pseudocercospora

Key to Pseudocercospora species on Dioscoreaceae
  1. 1

    Conidiophores very long, 50–490(−625) × 3–6.5 µm, pluriseptate, often branched; conidia (15−)25–95(−140) × (1.5−)3–6.5(−7) µm, (0−)1–5-distoseptate, subhyaline to pale olivaceous or brownish ........................................ see Distocercospora pachyderma

    Conidiophores much shorter, mostly unbranched; conidia euseptate ........................................ 2

  2. 2(1)

    Conidia solitary or in short chains, small, above all short, (10−)15–30(−35) × 2.5–6 µm, 0–3-septate, very pale, subhyaline to pale olivaceous; on Dioscorea spp., North America .................................... P. subrufa

    Conidia much longer, about 25–130 µm, pluriseptate ........................................ 3

  3. 3(2)

    Stromata lacking; conidiophores very long, 40–270 µm, pluriseptate; conidia mostly distinctly to strongly curved or sigmoid ........................................ P. cylindrata

    Stromata developed, if lacking conidiophores much shorter and only 0–3-septate; conidia straight to somewhat curved, but not strongly so ...................................... 4

  4. 4(3)

    Conidia narrow, 1–3 µm ..................................................... 5

    Conidia much broader, 2.5–8 µm ................................................. 6

  5. 5(4)

    Stromata lacking; mycelium internal and external; superficial hyphae with solitary conidiophores abundant; conidiophores short, 4–30 µm; conidia subhyaline and very narrow, 1–2.5 µm ........................................ P. ubicola

    Stromata well-developed, 20–65 µm diam; mycelium consistently internal; superficial hyphae with solitary conidiophores lacking ........................................ P. hiratsukana

  6. 6(4)

    Stromata lacking; conidiophores long, 20–150 µm; conidia smooth ........................................ P. ubi

    Stromata developed, 10–90 µm diam; conidiophores much shorter, 5–65 µm, if longer conidia verruculose ................................................... 7

  7. 7(6)

    Conidiophores long, 15–180 µm; conidia cylindrical to subcylindrical, faintly verruculose ...................... P. carbonacea

    Conidiophores shorter, 5–65 µm; conidia not consistently cylindrical-subcylindrical, either consistently obclavate and smooth to somewhat rough or variable in shape, cylindrical, obclavate to clavate and smooth. ........................................ 8

  8. 8(7)

    Conidia cylindrical to obclavate-cylindrical, base often peg-like (abruptly attenuated), pale to moderately deep olivaceous-brown, smooth. ........................................ P. contraria

    Conidia consistently obclavate, base obconically truncate, subhyaline to pale olivaceous, smooth to somewhat rough-walled ........................................ P. dioscoreae

Pseudocercospora species on Dioscoreaceae

Pseudocercospora carbonacea (L.E. Miles) N. Pons & B. Sutton, Mycol. Pap. 160: 26 (1988).

(Fig. 102)

Fig. 102
figure102

Pseudocercospora carbonacea (NY 936951). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercospora carbonacea L.E. Miles, Trans. Illinois Acad. Sci. 10: 255 (1917).

Synonym: Stenella dioscoreicola J.M. Yen, A.K. Kar & B.K. Das, Mycotaxon 16: 53 (1982) [holotype: India: West Bengal: Cooch Behar, Forest of Chelapata, on Dioscorea sp., 17 Oct. 1980, B. K. Das Pcc4452 (LAM, Yen #10578)].

Literature: Saccardo (1931: 874), Chupp (1954: 196), Vasudeva (1963: 68), Ellis (1976: 257), Little (1987a), Pons & Sutton (1988: 26), Crous & Braun (2003: 104), Phengsintham et al. (2003a: 107), Kamal (2010: 159).

Illustrations: Ellis (1976: 257, fig. 194 A), Yen et al. (1982: 52, fig. 8), Little (1987a: fig., unnumbered), Pons & Sutton (1988: 29, fig. 12), Phengsintham et al. (2003a: 108, figs 58–59).

Description: Leaf spots amphigenous, subcircular to angular-irregular, 3–20 mm diam, brown, margin indistinct, sometimes with brownish halo. Caespituli hypophyllous. Mycelium internal; rarely with a few superficial hyphae, branched, 2–3 µm wide, septate, pale olivaceous, thin-walled, fainly verruculose. Stromata substomatal, 10–90 µm diam. Conidiophores in small to large fascicles, about 5–40, loose to dense, arising from stromata, through stomata (rarely with solitary conidiophores arising from superficial hyphae when present), erect, straight, subcylindrical to somewhat geniculate-sinuous, unbranched, 10–180 × 3–5 µm, (0−)1–3-septate, pale to medium olivaceous-brown, wall smooth or almost so to somewhat rough; conidiogenous cells integrated, terminal, 10–60 µm long, sympodial, rarely percurrent, conidiogenous loci inconspicuous to subdenticulate, but always unthickened and not darkened. Conidia solitary, cylindrical or subcylindrical, straight to strongly curved or sigmoid, 40–130 × 3.5–6 µm, 3–9-septate, subhyaline to pale olivaceous, thin-walled, smooth to somewhat verruculose, apex obtuse, rounded, base short obconically truncate, occasionally somewhat peg-like, about 2–2.5 µm wide, hila unthickened, not darkened.

Lectotype (designated here, MycoBank MBT178152): Puerto Rico: Vega Alta, on Dioscorea alata, 1913, F. L. Stevens, 4178 (NY 936951). Isolectotypes: BPI 434203, 424305, 845236; MICH 15267.

Host range and distribution: On Dioscorea (alata, bulbifera, caucasica, cayennensis, deltoides, dumetorum, glabra, microbotrya [gilbertii], nipponica, oppositifolia, pentaphylla [spinosa], trifida, Dioscorea spp.), Dioscoreaceae, Africa (Ethiopia, Ghana, Guinea, Nigeria, Sierra Leone, Tanzania, Togo), Asia (India, Indonesia, Myanmar, Thailand), Australia, North America (Canada), Central and South America (Brazil, Panama, Venezuela), West Indies (Barbados, Cuba, Dominican Republ., French Antilles, Grenada, Haiti, Jamaica, Puerto Rico, Saint Lucia, Saint Vincent and the Grenadines, Trinidad and Tobago, Virgin Islands).

Notes: Due to inconspicuous conidiogenous loci, Stenella dioscoreicola has to be excluded from Stenella (Zasmidium). It agrees well with Pseudocercosora carbonacea, which is known from India, and is undoubtedly a synonym. P. carbonacea is one of few Pseudocercospora species with rough-walled conidiophores and conidia. The type collection of St. dioscoreicola differs from other collections of P. carbonacea by the formation of a few superficial hyphae with solitary conidiophores, which is, however, not unusual in Pseudocercospora species.

Pseudocercospora contraria (Syd. & P. Syd.) Deighton, Mycol. Pap. 140: 30 (1976).

(Fig. 103)

Fig. 103
figure103

Pseudocercospora contraria (B, holotype). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercospora contraria Syd. & P. Syd., Ann. Mus. Congo, Bot., ser. V, 3: 21 (1909).

Synonyms: Cercospora wildemanii Syd. & P. Syd., Ann. Mus. Congo, Bot., ser. V, 3: 21 (1909) [lectotype (designated here, MycoBank MBT178154: Republic of Congo: Kiduma, on Dioscorea sp. (as Dolichos sp.), 28 Feb. 1907, H. Vanderyst (BR-MYC 039969,05); isolectotypes: B 700016014, K(M) IMI 90862].

Mycosphaerella contraria Hansf., Proc. Linn. Soc. London 153: 22 (1941) [type: Uganda: Kampala plantation, on Dioscorea sp., June 1937, Chandler (K(M) IMI 7956)].

Literature: Saccardo (1913: 1419, 1430), Chupp (1954: 196, 340), Katsuki (1965: 28), Deighton (1976: 30), Ellis (1976: 257), Sivanesan (1984: 207), Little (1987b), Pons & Sutton (1988: 17), Guo & Hsieh (1995: 84), Guo et al. (1998: 101), Shin & Kim (2001: 178), Aptroot (2006: 68), Meeboon et al. (2008).

Illustrations: Deighton (1976: 31–32, figs 16–17), Sivanesan (1984: 208, fig. 109), Little (1987b: fig., unnumbered), Pons & Sutton (1988: 19–21, figs 6–8), Guo & Hsieh (1995: 86, fig. 78), Guo et al. (1998: 102, fig. 82), Shin & Kim (2001: 179, fig. 78).

Exsiccatae: Poelt & Scheuer, Reliqu. Petrak. 2797.

Description: Leaf spots subcircular, elliptical to irregular, 2–10 mm diam or confluent and larger, dingy grey or whitish, greyish brown, reddish to blackish brown, sometimes surrounded by a light brown border of 1–5 mm, less distinct on the lower side, sometimes with brownish or reddish halo. Caespituli amphigenous, punctiform, at first greenish, later deeper brown. Mycelium internal; hyphae branched, septate, hyaline, 1–3.5 µm wide. Stromata substomatal or occasionally immersed, about 15–60 µm diam, subglobose, brown to dark brown. Conidiophores in small and loose to usually large and dense fascicles, to 30 or even more, arising from stromata, through stomata or occasionally erumpent, erect, straight, subcylindrical to somewhat geniculate-sinuous, unbranched, 10–65 × 2–6.5 µm, 0–3-septate, pale to medium olivaceous or olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–25 µm long, sympodial or occasionally percurrent, conidiogenous loci inconspicuous to subdenticulate, 1–2 µm diam, but always unthickened, not darkened. Conidia solitary, cylindrical to obclavate-cylindrical, straight to curved, (15−) 30–115 × 2.5–8 µm, 2–22-septate, pale to moderately deep olivaceous-brown, wall thin to somewhat thickened, smooth, apex obtuse, rounded, base obconically truncate, sometimes peg-like, 1.5–2 µm wide, hila unthickened, not darkened. Sexual morph: Ascomata immersed, to 120 µm diam; asci 28–40 × 8–10 µm, 8-spored; ascospores fusiform to ellipsoid-ovoid, 8–12 × 2–4 µm, with a single median septum, not or barely constricted, colourless, ends rounded.

Lectotype (designated here, MycoBank MBT178153): Republic of Congo: Lazard, Kisantu, on Dioscorea sp., 9 Feb. 1908, H. Vanderyst (BR-MYC 039968,04). Isolectotypes: B 700016012, K(M) IMI 91059.

Host range and distribution: On Dioscorea (alata, asteriscus, bulbifera, cochleari-apiculata, dumetorum, hirtiflora, oppositifolia, pentaphylla [triphylla], quartiniana, quinquelobata, schimperiana, trifida, villosa [sativa], Dioscorea spp.), Dioscoreaceae, Africa (Cameroon, Congo, Ghana, Guinea, Nigeria, Sierra Leone, Sudan, Tanzania, Togo, Uganda), Asia (China, India, Indonesia, Japan, Java, Korea, Thailand), South America (Brazil), Oceania (Solomon Islands).

Pseudocercospora cylindrata (Chupp & Linder) N. Pons & B. Sutton, Mycol. Pap. 160: 27 (1988).

(Fig. 104)

Fig. 104
figure104

Pseudocercospora cylindrata (FH 2515). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercospora cylindrata Chupp & Linder, Mycologia 29: 29 (1937).

Synonym: Helicomina cylindrata (Chupp & Linder) Chupp, Monograph of Cercospora: 197 (1954).

Literature: Chupp (1954: 196), Pons & Sutton (1988: 27), Goh & Hsieh (1995: 357), Guo et al. (1998: 379), Crous & Braun (2003: 149–150), Kamal (2010: 169).

Illustrations: Pons & Sutton (1988: 32, fig. 13), Guo et al. (1998:380, fig. 311).

Description: Leaf spots indistinct or above all evident on the lower leaf surface, angular, vein-limited, 1–5 mm diam or confluent and larger, to 20 mm, mid dark brown, less evident on the upper leaf surface, only visible as diffuse pale brown angular discolorations. Caespituli hypophyllous, dark olivaceous to almost blackish. Mycelium internal and external, with some superficial threads. Stromata lacking. Conidiophores in small fascicles, 2–10, arising from internal hyphae, through stomata or solitary, arising from procumbent hyphae, lateral, erect, straight to flexuous, sinuous, but mostly not distinctly geniculate, unbranched or occasionally branched near the apex, 40–270 × 4–5 µm, width often irregular, plainly pluriseptate, olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci inconspicuous to subdenticulate, but always unthickened and not darkened. Conidia solitary, cylindrical to somewhat obclavate-cylindrical, mostly distinctly to strongly curved to sigmoid, 15–80 × 3–6.5 µm, 1–8-septate, pale to medium brown or olivaceous-brown, thin-walled, smooth, apex obtuse, base short obconically truncate, sometimes almost peg-like, about 1.5–2.5 µm wide, hila unthickened, not darkened.

Holotype: China: Kwangsi: Yung Hsien, Ta Tseh Tsuen, on Dioscorea sp., 18 Aug. 1933, S. Y. Cheo, no. 2515 (FH 2515). Isotypes: CUP 39586, K(M) IMI 255655 (slide), VIA 4733.

Host range and distribution: On Dioscorea (japonica, Dioscorea sp.), Dioscoreaceae, Asia (China, Japan; India, West Bengal).

Notes: Chupp (1954) recorded this species from Japan on Dioscorea septemloba, which is, however, unconfirmed. This species is not listed in Katsuki (1965). However, a Japanese collection on Dioscorea japonica, deposited by R. Kurata as Cercospora ubi at IUM-FY682, has been re-examined and proved to be Pseudocercospora cylindrata.

Pseudocercospora dioscoreae U. Braun, Mouch. & McKenzie, New Zealand J. Bot. 37: 313 (1999).

(Fig. 105)

Fig. 105
figure105

Pseudocercospora dioscoreae (PC, holotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Illustration: Braun et al. (1999: 311, fig. 16).

Description: Leaf spots amphigenous, angular-irregular, 2–25 mm diam, dingy greyish brown to greyish white, margin narrow, dark brown to blackish. Caespituli amphigenous, mostly epiphyllous, punctiform, scattered, dark brown. Mycelium internal; hyphae septate, branched, brown, smooth. Stromata substomatal, 10–60 µm diam, brown. Conidiophores numerous, in dense fascicles, arising from stromata, through stomata, erect, straight or often curved throughout or only at the tip, subcylindrical, barely or only slightly geniculate, unbranched, 5–50 × 3–8 µm, 0–1-septate, pale to medium brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–30 µm long, conidiogenous loci inconspicuous, occasionally subconspicuous (in front view visible as minute slightly darkened circle). Conidia solitary, almost consistently obclavate, rarely fusiform or subcylindrical, 30–85 × (3−)4–6(−7) µm, 3–7-septate, subhyaline to pale olivaceous, thin-walled, smooth to somewhat rough, apex subacute, base obconically truncate, 2–3 µm wide, hila unthickened, not darkened.

Holotype: New Caledonia: Monts Koghis (Lavoix), on Dioscorea bulbifera, 6 May 1964, Huguenin, NC 64.256 (PC).

Host range and distribution: On Dioscorea (alata, bulbifera), Dioscoreaceae, Oceania (New Caledonia, Vanuatu).

Notes: Braun et al. (1999) cited two collections from Vanuatu on Dioscorea alata and D. bulbifera, respectively, as paratypes (PDD 46864 and 57186).

Pseudocercospora hiratsukana (Togashi & Katsuki) Deighton, Mycol. Pap. 140: 34 (1976).

(Fig. 106)

Fig. 106
figure106

Pseudocercospora hiratsukana (CUP 40760). A. Conidiophore fascicle and solitary conidiophore. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora hiratsukana Togashi & Katsuki, J. Jap. Bot. 28: 286 (1953).

Literature: Katsuki (1965: 28), Deighton (1976: 34–36), Yen & Lim (1980: 177), Pons & Sutton (1988: 26), Crous & Braun (2003: 218).

Illustrations: Deighton (1976: 35, fig. 18), Pons & Sutton (1988:28, fig. 11).

Description: Leaf spots amphigenous, subcircular, elliptical to irregular-angular, somewhat vein-limited, 2–8 mm diam, brown, later greyish brown or grey, margin indefinite or darker, brown. Caespituli amphigenous, mainly hypophyllous, punctiform, dark. Mycelium internal, rarely with repent hyphae, 1.5–3 µm wide, very pale olivaceous. Stromata substomatal, subglobose, 20–65 µm diam, olivaceous-brown. Conidiophores in dense, mostly large fascicles, to 30 or even more, arising from stromata, through stomata, sometimes rupturing the stomata, erect, straight to slightly flexuous-sinuous, barely geniculate, subcylindrical or somewhat attenuated towards the tip, unbranched, 5–40 × 2–4 µm, 0–2(−4)-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores often reduced to conidiogenous cells, 5–25 µm long, conidiogenous loci inconspicuous or visible as truncate tips, unthickened, not darkened. Conidia solitary, obclavate, obclavate-subcylindrical, subcylindrical, fusiform, straight to somewhat curved, 25–60 × 1.5–3 µm, 2–7-septate, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth, apex obtuse to subacute, base obconically truncate, 1–2 µm wide, hila unthickened, not darkened.

Spermogonia sometimes developed, amphigenous, mainly epiphyllous, scattered, immersed, globose, 50–55 µm diam, sometimes apparently developed in old conidiophore stromata; spermatia rod-shaped, 3–4 × 1 µm, hyaline.

[Type: Japan: Kagoshima Pref.: Kusukawa, Yaku Island, on Dioscorea quinqueloba, 14 Oct. 1949, S. Katsuki (not preserved)]. Neotype (designated here, MycoBank MBT178155): Japan: Kagoshima, Yaku Island, on Dioscorea quinqueloba, 5 Aug. 1951, S. Katsuki (CUP 40760). Epitype (designated here, MycoBank MBT178156): Japan : Tokyo, Inagi, Kurihira, on Dioscorea tokoro, 23 Oct. 1999, E. Imaizumi (TNS-F-61275). Ex-epitype culture: MAFF238300.

Host range and distribution: On Dioscorea (quinqueloba, tokoro, Dioscorea sp.), Dioscoreaceae, Asia (Japan, Singapore).

Notes: Type material of this species could not be traced but other authentic collections were examined. The collection designated as neotype represents topotype material, and there is an additional specimen collected by Katsuki elsewhere [Fukuoka, Shikano-shima, 27 Nov. 1949, S. Katsuki (CUP 39641 and YNU 24495; slide at K(M) IMI 92182a)].

Pseudocercospora subrufa (Ellis & Holw.) U. Braun, Cryptog. Bot. 3: 241 (1993).

(Fig. 107)

Fig. 107
figure107

Pseudocercospora subrufa (NY 830555). A. Conidiophore fascicle. B. Solitary conidiophores arising from superficial hyphae. C. Conidia. Bar = 10 µm.

Basionym: Ramularia subrufa Ellis & Holw., J. Mycol. 4: 2 (1888).

Synonyms: Mycovellosiella subrufa (Ellis & Holw.) U. Braun, Mycotaxon 48: 288 (1993).

Ramularia dioscoreae Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 43: 85 (1891) [lectotype (designated by Braun 1993): USA: Wisconsin: Racine, on Dioscorea villosa, 17 Aug. 1889, J. J. Davis (NY 1042940); isolectotype: NY 1042938].

Didymaria fulva Ellis & Everh., in herb. [authentic material: USA: Indiana: Crowforsville, on Dioscorea villosa, 7 June 1894, Olive (BPI 415505, NY 928029, 928030, 928031)].

Literature: Saccardo (1892: 562), Sutton & Pons (1988: 37), Braun (1998: 373).

Illustrations: Sutton & Pons (1988: 38, fig. 17), Braun (1993a: 242, fig. 18; 1998:372, fig.633).

Description: Leaf spots amphigenous, small, 1–4 mm diam, angular-irregular, vein-limited, yellowish, ochraceous to pale brown, finally sometimes greyish white, margin indefinite or sometimes with darker marginal line. Caespituli hypophyllous, greyish white to faintly pigmented, yellowish ochraceous. Mycelium internal and external; superficial hyphae climbing leaf hairs, branched, 1–4 µm wide, septate, subhyaline to pale olivaceous, thin-walled, smooth. Stromata small to well-developed, substomatal to erumpent, pigmented. Conidiophores solitary, arising from superficial hyphae, lateral, as well as in small to moderately large, loose to dense fascicles, arising from stromata, through stomata or erumpent, erect, straight and subcylindrical to geniculate-sinuous, unbranched or rarely branched, 5–60 × 2–5.5 µm, aseptate or sparingly septate, subhyaline, yellowish, greenish to olivaceous, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–30 µm long, conidiogenous loci often somewhat protuberant, subdenticulate, but unthickened, sometimes conspicuous by being slightly darkened or refractive. Conidia solitary, occasionally catenate, rarely in branched chains, ellipsoid-ovoid, fusiform, straight to curved, (10−)15–30(−35) × 2.5–6 µm, (0−)1–2(−3)-septate, subhyaline to faintly yellowish green or pale olivaceous, thin-walled, smooth, apex rounded to subacute, base obconically truncate, 1–1.5 µm wide, hila unthickened, not or barely darkened.

Holotype: USA: Iowa: Decorah, on Dioscorea villosa (as “Smilax sp.”), 28 June 1885, Holway (NY 830555). Isotype: B 700016015.

Host range and distribution: On Dioscorea villosa, Dioscoreaceae, North America (USA, Iowa, Indiana, Wisconsin).

Notes: The conidiogenous loci of this unusual species are subconspicuous, i.e., unthickened, but occasionally somewhat darkened, protuberant, subdenticulate. Taxa with subconspicuous (above all unthickened) scars have to be placed in Pseudocercospora, which has recently been confirmed by molecular examinations (Crous et al. 2000, 2013).

Pseudocercospora ubi (Racib.) Deighton, Mycol. Pap. 140: 36 (1976).

(Fig. 108)

Fig. 108
figure108

Pseu