IMA FuNguS · 5(2): 203–390 (2014)
doi:10.5598/imafungus.2014.05.02.04
Uwe Braun1, Pedro W. Crous2, and Chiharu Nakashima3
ART I CLE
Cercosporoid fungi (Mycosphaerellaceae) 2. Species on monocots
(Acoraceae to Xyridaceae, excluding Poaceae)
1
Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik und Botanischer Garten, Herbarium, Neuwerk 21, 06099 Halle (Saale),
Germany; corresponding author e-mail: uwe.braun@botanik.uni-halle.de
2
3
CBS-KNAW, Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands
Graduate School of Bioresources, Mie University, 1577 Kurima-machiya, Tsu, Mie 514-8507, Japan
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 irst 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 irst part of this series. Neopseudocercospora, an
additional recently introduced cercosporoid genus, is briely 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. constrictolexuosa 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.
Key words:
Ascomycota
Cercospora s. lat.
Liliopsida
hyphomycetes
Article info: Submitted: 2 June 2014; Accepted: 31 October 2014; Published: 25 November 2014.
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 irst 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).
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VOLUME 5 · NO. 2
203
Braun et al.
ART I CLE
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 magniication of ×1000. All illustrations
have been prepared by the irst 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
Crous & U. Braun, comb. nov.
MycoBank MB809006
(Deighton)
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
supericial; hyphae branched, septate, pigmented, smoothwalled. Stromata lacking. Conidiophores solitary, arising from
supericial hyphae, lateral, one-celled or septate, 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/scolecostigminalike, 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 conspeciic 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
204
Conidia acicular, base truncate ....................................................................................................................................... 2
Conidia at least partly obclavate-cylindrical with obconically truncate base ................................................................... 3
IMA FUNGUS
Cercosporoid fungi 2
2 (1)
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 (3)
On Alisma spp. and Echinodorus spp. .......................................................................................................... C. alismatis
On Sagittaria spp. ....................................................................................................................................... C. sagittariae
Cercospora species on Alismataceae
Literature: Zhuang (2001), Crous & Braun (2003: 51).
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)
Illustration: Chi (1994: 97, ig. 88).
ART I CLE
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.]
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.
Fig. 1. Cercospora alismaticola (based on Chi 1994: 97, ig. 88). A.
Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
VOLUME 5 · NO. 2
Description: Leaf spots amphigenous, subcircular to angularirregular, 2–15 mm diam, brown to greyish brown, margin
indeinite 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
205
ART I CLE
Braun et al.
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.
wide above, pale olivaceous to medium brown, septate, thinwalled, 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, thinwalled, 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)
Literature: Crous & Braun (2003: 300)
Illustration: El-Gholl et al. (1992: 270, igs 1–4).
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
Cercospora: 28 (1954).
(Similar to Fig. 1)
Chupp,
Monograph
of
Literature: Chupp (1954: 28), El-Gholl et al. (1992: 265),
Crous & Braun (2003: 169).
Description: Leaf spots subcircular to somewhat angularirregular, 2–12 mm diam or conluent and larger, centre grey,
margin darker, brown to blackish, indistinctly zonate. Caespituli
amphigenous, ine, 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
206
Fig. 2. Cercospora osiridis (FLAS F55631). A. Conidiophore fascicle.
B. Conidiophore tips. C. Conidia. Bar = 10 µm.
IMA FUNGUS
Cercosporoid fungi 2
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.
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 olivaceousbrown, 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.
ART I CLE
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.
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.
Cercospora sagittariae Ellis & Kellerm., J. Mycol. 2:
1 (1886).
(Fig. 3)
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, ig. 8), Soares et al. (2009:
405, ig. 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
conluent, 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
VOLUME 5 · NO. 2
Fig. 3. Cercospora sagittariae (BPI 440960). A. Conidiophore
fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.
207
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Braun et al.
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).
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.
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)
Illustration: Soares et al. (2009: 410, ig. 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 welldeveloped, dense fascicles, erect, straight to somewhat
curved, subcylindrical, not geniculate, unbranched, 20–40
× 2–4 µm, 1–3-septate, pale olivaceous to olivaceousbrown, 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 irst 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, lat, with irregular margin, light
grey, reverse grey; on MEA like on OA.
Holotype: Brazil: Santa Catarina: Forquilhinha, SC 448, rice
ield at road side, on Sagittaria montevidensis, 21 Apr. 2005,
D. J. Soares (VIC 30505).
208
Fig. 4. Pseudocercospora arthrospora (VIC 30505). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
IMA FUNGUS
Cercosporoid fungi 2
Amaryllidaceae (incl. Allioideae, syn. Alliaceae)
ART I CLE
Cercospora
Key to Cercospora species on Amaryllidaceae
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 (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 insuficiently known species)
Conidia acicular, much longer ......................................................................................................................................... 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 (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 (4)
Stromata lacking or small; on Allium spp. ....................................................................................................... C. duddiae
Stromata well-developed, 25–70 µm diam ...................................................................................................................... 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)
Literature: Crous & Braun (2003: 55), Kamal (2010: 16).
Illustration: Rajak (1980: 458, ig. 2).
Description: Leaf spots olivaceous-brown to greyish brown,
halo indistinct, infections at irst at the tip, developing
downwards, die-back observed. Caespituli amphigenous,
effuse, greyish black. Mycelium internal. Stromata welldeveloped, globose, 64–112 µm wide, dark brown to greyish
black. Conidiophores fasciculate, 6–15, erect, straight to
lexuous, 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-iliform, 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).
VOLUME 5 · NO. 2
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 x 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, ig. 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
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Braun et al.
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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) conined 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, ig. 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.
Fig. 5. Cercospora amaryllidicola (K(M) 225290). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
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.,
210
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)
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).
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
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,
istulosum, 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)
Literature: Saccardo (1886: 477), Chupp (1954: 37), Crous &
Braun (1996: 276; 2003: 209).
Fig. 6. Cercospora duddiae (BPI 436011). A. Conidiophore fascicle.
B. Conidiophore. C. Conidia. Bar = 10 µm.
Illustrations: Narain & Saksena (1971: 135, igs 1–2), Ellis
(1976: 271, ig. 205 B), Little (1987c: unnumbered igure), Chi
(1994: 209, ig. 230), Guo et al. (2005: 177, ig. 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
VOLUME 5 · NO. 2
Description: Leaf spots amphigenous, circular to somewhat
angular-irregular, 3–12 mm diam, pale tan to dark olivaceous,
margin indeinite 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,
multilorus,
puniceus
[Haemanthus
natalensis]),
Amaryllidaceae, Africa (Ethiopia, South Africa).
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Braun et al.
Fig. 7. Cercospora haemanthi (B, holotype). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
Fig. 8. Cercospora riofranciscana (IMUR 46193). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
Note: A true Cercospora s. str. distinct from C. apii s. lat. by
having much wider obclavate-cylindrical conidia.
Illustration: Batista et al. (1965: 20, ig. 6).
5.5 µm. Stromata intraepidermal, 25–65 µm diam, brown.
Conidiophores fasciculate, subcylindrical-iliform, nongeniculate, 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.
Description: Leaf spots numerous, elliptical or oblong, at
irst brown, later dark, 0.5–4 mm diam. Mycelium internal;
hyphae inter- and intracellular, hyphal cells 7.5–18.5 × 3.5–
Holotype: Brazil: Pernambuco: Rio São Francisco, Belém de
Maria, on Allium cepa, 26 May 1965, A. Lustosa Sobrinho
(IMUR 46193).
Cercospora riofranciscana Bezerra & Peres, Publ.
Inst. Micol. Univ. Recife 450: 9 (1965).
(Fig. 8)
Literature: Crous & Braun (2003: 356).
212
IMA FUNGUS
Cercosporoid fungi 2
Doubtful,
species
excluded
and
insuficiently
known
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 insuficiently 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).
ART I CLE
Host range and distribution: On Allium cepa, Amaryllidaceae
(Allioideae, syn. Alliaceae), South America (Brazil).
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, igs 1–2), Braun
& Mel’nik (1997: ig. 71), Schubert (2005: 149, ig. 72, pl. 33,
igs A–D), Bensch et al. (2012: 288–289, igs 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
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)
Illustration: Gupta et al. (1982: 112, igs 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 olivaceousbrown, 0–2-septate, thin-walled, smooth; conidiogenous
VOLUME 5 · NO. 2
cells integrated, terminal or conidiophores reduced to
conidiogenous cells, conidiogenous loci inconspicuous,
unthickened, not darkened. Conidia solitary, obclavatecylindrical, 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.
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Braun et al.
Fig. 9. Pseudocercospora indica (based on Gupta et al. 1982; 112,
ig 1–2). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.
Fig. 10. Pseudocercospora pancratii (NY 00838178). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
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)
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.
214
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, ig. 25 A),
Kirschner (2014: 488, ig. 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 indeinite 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.
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Mycelium internal; hyphae branched, septate, 1–4 µm wide,
hyaline to medium brown, thin-walled, smooth. Stromata welldeveloped, 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 lexuous,
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-iliform, 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, tubilora), 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)
Basionym: Stenella hymenocallidis U. Braun & Crous,
Mycotaxon 92: 403 (2005).
Illustration: Braun & Crous (2005: 411, ig. 8).
Fig. 11. Zasmidium hymenocallidis (NY 01043037). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
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 olivaceousbrown, 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-iliform, 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.
Exsiccatae: Ellis & Everh., North Amer. Fungi 1767.
Description: Lesions diffuse, shape and size variable, brown
to reddish brown, margin indeinite. Caespituli amphigenous,
punctiform, scattered, greyish brown. Mycelium internal;
hyphae branched, septate, 2–5 µm wide, subhyaline, thinwalled, 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
VOLUME 5 · NO. 2
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.
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Notes: This is one of the few Zasmidium species without
any supericial hyphae in vivo. However, molecular analyses
of comparable species with lacking supericial 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, igs 5–8).
Description: Leaf spots amphigenous, on loating leaves,
subcircular to somewhat angular-irregular, 1–4 mm diam,
often conluent, 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
1
Conidia acicular, base truncate ............................................................................................................ C. amorphophalli
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 obclavatesubcylindrical, 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
216
Conidia acicular, 25–260 × 2.5–5 µm, pluriseptate ............................................................................................ C. caladii
IMA FUNGUS
Cercosporoid fungi 2
Calla
A single species ............................................................................................................................................................... C. callae
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Conidia broadly ellipsoid-ovoid, obovoid to fusiform-subcylindrical, 15–55(–70) × 5–9.5 µm, 2–7-septate
........................................................................................................................................................... C. verruculosa
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:
VOLUME 5 · NO. 2
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).
217
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Braun et al.
Illustrations: Hsieh & Goh (1990: 29, ig. 15), Narayan et al.
(1997: 89, ig. 2), Guo et al. (2005: 35, ig. 15), Phengsintham
et al. (2013b: 48, igs 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 indeinite 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, olivaceousbrown. 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, thinwalled, 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)
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, ig. 1), Braun (2001a:
433, ig. 16 B), Guo et al. (2005: 36, ig. 16).
218
Fig. 12. Cercospora amorphophalli (B, holotype). A. Conidiophores.
B. Conidia. Bar = 10 µm.
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 welldeveloped, 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
IMA FUNGUS
Cercosporoid fungi 2
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Fig. 13. Cercospora amorphophallicola (L 53844). A. Conidiophores. B. Conidia. Bar = 10 µm.
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).
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.
Note: This species belongs to the Cercospora apii complex.
Cercospora amorphophallicola U. Braun, Nova
Hedwigia 73: 430 (2001).
(Fig. 13)
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).
Literature: Crous & Braun (2003: 56).
Illustration: Braun (2001a: 433, ig. 16 A).
Description: Leaf spots amphigenous, subcircular to angularirregular, 1–15 mm diam, pale greenish, later pale to dark
brown, blackish or reddish brown, inally greyish brown to
dingy grey, margin indeinite 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
VOLUME 5 · NO. 2
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)
Literature: Kranz (1963, as C. chevalieri), Crous & Braun
(2003: 57).
Illustration: Yen & Gilles (1970: 75, ig. 1).
219
Braun et al.
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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)
Literature: Crous & Braun (2003: 57).
Illustration: Yen (1974: 311, ig. 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 (-obclavatecylindrical), 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).
Fig. 14. Cercospora anchomanes (based on Yen & Gilles 1970: 75,
ig. 1). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar
= 10 µm.
Description: Leaf spots amphigenous, circular to irregular,
1–11 mm diam, mostly conluent, 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 lexuous 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,
220
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).
IMA FUNGUS
Cercosporoid fungi 2
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Fig. 15. Cercospora anchomanicola (PC, topotype). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
Cercospora arisaematis F.L. Tai, Chinese Bot. Soc.
Bull. 2: 47 (1936); as “arisemae”.
(Fig. 16)
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, ig. 16), Guo et al. (2005:
37, ig. 17).
Description: Leaf spots circular or subcircular, 0.5–10
mm diam, at irst 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,
VOLUME 5 · NO. 2
Fig. 16. Cercospora arisaematis (HMAS 06993). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
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, thinwalled, 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.
221
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Braun et al.
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)
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, igs 8–9).
Description: Leaf spots amphigenous, angular-irregular, 3–6
mm diam, scattered, pale to medium brown, border distinct,
darker brown. Caespituli amphigenous, inely 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, ig. 18).
222
Fig. 17. Cercospora bombycina (BPI 433778). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
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.
IMA FUNGUS
Cercosporoid fungi 2
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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)
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), ElGholl et al. (1992: 265), Crous & Braun (2003: 94), Schubert
& Braun (2007: 190).
Illustration: Schubert & Braun (2007: 191, ig. 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, thinwalled, 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 obclavatecylindrical, 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,
VOLUME 5 · NO. 2
Fig. 18. Cercospora callae (NYS, holotype). A. Conidiophore
fascicle. B. Conidophore tips. C. Conidia. Bar = 10 µm.
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 plantagoaquatica, 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.
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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 misidentiications. 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. 19. Cercospora chevallieri (FH 7807). A. Conidiophore fascicle.
B. Conidiophore. C. Conidia. Bar = 10 µm.
Literature: Crous & Braun (2003: 146), Kamal (2010: 37).
Illustrations: Chiddarwar (1962: 74, plate 1, igs 12–14).
Cercospora chevalieri Sacc., Syll. Fung. 22: 1431
(1913).
(Fig. 19)
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, ig. 184).
Description: Leaf spots amphigenous, subcircular to angularirregular, 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.
224
Description: Leaf spots amphigenous, scattered, subcircular,
oval to somewhat irregular, 2–25 mm diam, yellowish to
ochraceous, pale brownish, inally 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.
IMA FUNGUS
Cercosporoid fungi 2
Literature: Kamal (2010: 67).
Description: Leaf spots amphigenous, subcircular to
irregular, 5–53 mm diam, later conluent, 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-lexuous, unbranched, about 45–90 × 2.5–
4.5 µm, 2–5-septate, light olivaceous to olivaceous, thinwalled, 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.
ART I CLE
Illustration: Narayan et al. (2001: 69, ig. 6).
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)
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, ig. 7), Morris & Crous
(1994: 327, ig. 4), Barreto et al. (1999: 82, ig. 1), Guo et al.
(2005: 39, ig. 18).
Fig. 20. Cercospora cryptocorynes (BPI 435386). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
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)
VOLUME 5 · NO. 2
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
conluent, 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
225
Braun et al.
ART I CLE
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-septate, 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).
Fig. 21. Cercospora pistiae (BPI 439744). A. Conidiophore fascicle.
B. Conidia. Bar = 10 µm.
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.
Host range and distribution: On Pistia stratiotes, Araceae,
Africa (Ghana, South Africa), Asia (China; India, Kerala),
Australia, South America (Brazil, Venezuela).
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.
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
Cercospora symplocarpi Peck, in Thüm., Mycoth.
Univ., Cent. VII, no. 669 (1877).
(Fig. 22)
Holotype: India: Kerala: Ernakulam, on Pistia stratiotes, 11
Dec. 1964, T. R. Nag Raj (MYSP Herb. 803)
226
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).
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
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 darkenedrefractive, 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 afinity of this
species has to be proven by means of molecular sequence
analyses.
Fig. 22. 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
VOLUME 5 · NO. 2
Cercospora syngoniicola U. Braun & Urtiaga,
Mycosphere 4: 593 (2013).
(Fig. 23)
Illustration: Braun & Urtiaga (2013b: 594, ig. 1).
Description: Leaf spots amphigenous, subcircular to angularirregular, 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, inely
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 geniculatesinuous, unbranched, 10–130 × 3–9 µm, 0–5-septate,
pale to medium olivaceous-brown or yellowish brown, thinwalled, smooth; conidiogenous cells integrated, terminal or
227
Braun et al.
ART I CLE
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, ig. 2).
Description: Leaf spots circular or subcircular, scattered to
conluent, 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 (lagelliformum,
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).
Fig. 23. Cercospora syngoniicola (K(M) 180153). A. Conidiophore
fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
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 obclavatesubcylindrical, 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).
228
Cercospora verruculosa F. Stevens & Solheim,
Mycologia 23: 397 (1931).
(Fig. 24)
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).
IMA FUNGUS
Cercosporoid fungi 2
Holotype: trinidad: St. Augustine, on Caladium sp., 13 Aug.
1932, F. L. Stevens 829 (ILL 15793).
ART I CLE
base rounded to short obconically truncate, 2–3.5 µm wide,
hila somewhat thickened and darkened.
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,
species
excluded
and
insuficiently
known
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).
Fig. 24. Cercospora verruculosa (BPI 428776). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
Literature: Saccardo (1895: 605), Vassiljevsky & Karakulin
(1937: 54), Braun (1995: 250), Braun & Mel'nik (1997: 40),
Crous & Braun (2003: 65).
Illustration: Braun (1995: 247, ig. 230 b).
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, ig. 17).
Description: Leaf spots amphigenous, circular to angularirregular, 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 geniculatesinuous, 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,
VOLUME 5 · NO. 2
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 inal conclusion about the status of this species (see
Braun 1995).
229
Braun et al.
ART I CLE
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).
Notes: This species is insuficiently 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 conspeciic. 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.
Host range and distribution: On unknown member of Araceae,
Typhonium giganteum, Araceae, Asia (China), North America
(Mexico).
Passalora
Key to Passalora species on Araceae
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).
passalora colocasiae (Höhn.) U. Braun, in Braun et
al., New Zealand J. Bot. 37: 308 (1999).
(Fig. 25)
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.
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).
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.
230
Notes: This species is insuficiently 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 supericial 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.
Literature: Saccardo (1913: 1431), Chupp (1954: 58),
Vasudeva (1963: 87), Sivanesan (1984: 185), Crous & Braun
(2003: 132), Aptroot (2006: 31–32), Kamal (2010: 113).
IMA FUNGUS
Cercosporoid fungi 2
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.
ART I CLE
Exsiccatae: Rehm, Ascomyc. 2060. Syd., Fungi Exot. Exs.
30.
Holotype: Samoa: on living leaves of Colocasia esculenta,
1905, Rechinger 2297 (FH). Isotype: CUP 39261.
Fig. 25. Passalora colocasiae (CUP 39261). A. Conidiophore
fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
Illustrations: Sivanesan (1984: 186, ig. 94), Aptroot (2006:
10, ig. 36).
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
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 (1)
Mycelium internal and external, supericial; stromata lacking; conidiophores fasciculate and solitary,
arising from supericial hyphae, to 300 µm long; conidia cylindrical, 40–80 × 4–4.5 µm;
on Amorphophallus ................................................................................................................................. p. protensa
Mycelium internal; supericial hyphae lacking; stromata developed; conidiophores much shorter, 5–80 µm;
conidia obclavate-cylindrical, iliform or subacicular, if cylindrical then conidia wider, 6–9 µm;
on other hosts .......................................................................................................................................................... 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
VOLUME 5 · NO. 2
231
ART I CLE
Braun et al.
4 (3)
Conidia narrowly obclavate to cylindrical-iliform, 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 (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
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. 26. Pseudocercospora alocasiicola (PDD 49041). A.
Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
Pseudocercospora species on Araceae
pseudocercospora alocasiicola U. Braun &
McKenzie, New Zealand J. Bot. 37: 306 (1999).
(Fig. 26)
Illustration: Braun et al. (1999: 307, ig. 7).
Description: Leaf spots amphigenous, subcircular to irregular,
1–15 mm diam, at irst brownish, dark greyish brown to blackish
or violet-black, margin indeinite, 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
232
Illustration: Braun et al. (1999: 307, ig. 9).
Description: Leaf spots subcircular, 4–6 mm diam, brownish,
margin indeinite, but often with a greenish halo. Caespituli
amphigenous, punctiform, dark brown. Mycelium internal;
hyphae septate, branched, pigmented. Stromata welldeveloped, 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, thinwalled, smooth, apex usually obtuse, occasionally subacute,
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 27. Pseudocercospora aracearum (PDD 60364). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
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)
Illustration: Braun & Sivapalan (1999: 12, ig. 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,
VOLUME 5 · NO. 2
Fig. 28. Pseudocercospora bruneiensis (HAL 1596 F). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
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 lexuous, geniculate-sinuous, simple or occasionally
branched, 5–40 × 2–5 µm, 0–1-septate, pale olivaceous to
medium olivaceous-brown, tips often somewhat paler, thinwalled, 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-iliform, 40–110 × 2–4 µm,
233
ART I CLE
Braun et al.
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)
Literature: Matsushima (1985), Goh & Hsieh (1990: 30), Guo
& Hsieh (1995: 22), Guo et al. (1998: 35–36).
Illustrations: Deighton (1976: 18–19, igs 5–6), Matsushima
(1985: ig. 298), Goh & Hsieh (1990: 31, ig. 17), Guo & Hsieh
(1995: 25, ig. 23), Guo et al. (1998: 35, ig. 23).
Description: Leaf spots indistinct, forming subcircular pale
discolorations, to 10 mm diam, centre sometimes turning
greyish with age, margin indeinite. 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-iliform, lexuous, 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 roughwalled 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 roughwalled, 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).
Fig. 29. Pseudocercospora colocasiae (K(M) IMI 77190). A.
Conidiophore fascicle. B. Conidiophore tip with conidium. C. Conidia.
Bar = 10 µm.
Literature: Saccardo (1931: 869), Chupp (1954: 59), Crous &
Braun (2003: 281).
Illustration: Braun & Freire (2002: 314, ig. 20).
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)
Basionym: Cercospora montrichardiae Henn., Hedwigia 48:
115 (1909).
234
Description: Leaf spots amphigenous, subcircular to angularirregular, 1–15 mm diam, yellowish to ochraceous, inally
greyish brown to greyish white, margin indeinite 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,
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 30. Pseudocercospora montrichardiae (S-F37470). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
erect, straight, subcylindrical-conical to strongly geniculatesinuous, unbranched or occasionally branched, 5–25 ×
2–4 µm, 0–2-septate, subhyaline to pale olivaceous, thinwalled, 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).
VOLUME 5 · NO. 2
Fig. 31. Pseudocercospora protensa (B 70014865). A. Stroma.
B. Conidiophore fascicle and supericial hypha with conidiophore
emerging through a stoma. C. Conidiophore tips. d. Conidia. Bar =
10 µm.
pseudocercospora protensa (Syd.) Deighton, Mycol.
Pap. 140: 150 (1976).
(Fig. 31)
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 conluent, effuse, later visible as large patches or layers,
loose to dense, dark olivaceous to brown. Mycelium
235
ART I CLE
Braun et al.
internal, occasionally with a few procumbent threats
growing like supericial 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 lexuous, 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 olivaceousbrown 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 conirmed
(North Borneo, 1 Sep. 1959, A. Johnston NB 28, K(M) IMI
78759).
Zasmidium
Key to Zasmidium species on Araceae
1
Supericial hyphae not developed in vivo; conidiophores only fasciculate; conidia formed singly,
obclavate, 30–60 × 3–4 µm; on Cercestis afzelii, Africa .......................................................... Z. cercestidis-afzelii
Supericial hyphae developed in vivo, with or without solitary conidiophores ................................................................. 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 (2)
Supericial hyphae formed in vivo, but conidiophores only fasciculate, solitary conidiophores arising
from supericial hyphae lacking; conidia to 350 µm long; on Homalomena spp., Asia ..................... Z. extremorum
Solitary conidiophores arising from supericial hyphae formed in vivo; conidia much shorter,
only to 140 µm; on other hosts ................................................................................................................................ 4
4 (3)
Stromata lacking; on Anthurium or Cercestis .................................................................................................................. 5
Stromata developed; on Alocasia or Colocasia .............................................................................................................. 6
5 (4)
Conidiophores solitary; conidia solitary, obclavate-cylindrical to iliform; on Anthurium sp., Asia ............. Z. anthuriicola
Conidiophores solitary as well as fasciculate; conidia catenate, cylindrical or subcylindrical;
on Cercestis congoensis, Africa .......................................................................................................... Z. cercestidis
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
Illustration: Sarbajna & Chattopadhyay (1991: 34, ig. 2).
Zasmidium alocasiae (Sarbajna & Chattopadh.)
Kamal, Cercosporoid Fungi of India: 237 (2010).
(Fig. 32)
Description: Leaf spots amphigenous, subcircular to somewhat
irregular, scattered to conluent, 2–18 mm diam, at irst 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
conluent, punctiform, dark brown, blackish brown, later
Basionym: Stenella alocasiae Sarbajna & Chattopadh., J.
Mycopathol. Res. 29: 33 (1991).
Literature: Braun & Sivapalan (1999: 23), Guo (2010).
236
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
cylindrical, iliform, 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)
Basionym: Stenella anthuriicola U. Braun & C.F. Hill, in Braun
et al. Fungal Diversity 22: 33 (2006).
Illustration: Braun et al. (2006: 29, ig. 9).
Fig. 32. Zasmidium alocasiae (K(M) IMI 284569). A. Supericial
hyphae. B. Supericial hyphae with conidiophores. C. Conidiophore
fascicles. d. Conidiophores. e. Conidia. Bar = 10 µm.
greyish brown by abundant sporulation. Mycelium internal
and external; supericial 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 supericial 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 supericial 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,
VOLUME 5 · NO. 2
Description: Lesions formed as sharply delineated brown
discolorations on leaves, to 5 mm wide, with abundant
supericial 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, iliform, 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.
237
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Braun et al.
Fig. 33. Zasmidium anthuriicola (HAL 1870 F). A. Supericial hyphae.
B. Supericial hyphae with conidiophores. C. Conidiophores. d.
Conidia. Bar = 10 µm.
Fig. 34. Zasmidium cercestidis (K(M) IMI 212936). A. Supericial
hypha with solitary conidiophores. B. Conidiophore fascicle. C.
Conidiophore tip. d. Conidia. Bar = 10 µm.
Host range and distribution: Only known from the type
collection.
Note: Crous et al. (2009c) analysed this species by means of
molecular methods and conirmed its phylogenetic position.
Zasmidium cercestidis (J.M. Yen & Gilles) U. Braun,
Schlechtendalia 20: 100 (2010).
(Fig. 34)
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, ig. 2).
238
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; supericial 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 supericial 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
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
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)
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, ig. 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, thinwalled, 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 supericial
mycelium is not formed. We prefer to maintain two species
and consider C. cercestidis-afzelii as one of the Zasmidium
species without formation of supericial hyphae in vivo. With
regard to the unclear status of the type collection see notes
under Z. cercestidis.
VOLUME 5 · NO. 2
Fig. 35. Zasmidium cercestidis-afzelii (K(M) IMI 212936). A.
Conidiophore fascicle. B. Conidia. Bar = 10 µm.
Zasmidium colocasiae (Sarbajna & Chattopadh.)
Kamal, Cercosporoid Fungi of India: 240 (2010).
(Fig. 36)
Basionym: Stenella colocasiae Sarbajna & Chattopadh., J.
Mycopathol. Res. 29: 35 (1991).
Illustration: Sarbajna & Chattopadhyay (1991: 36, ig. 3).
Description: Leaf spots amphigenous, circular, 2–10 mm
diam, at irst 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,
239
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Braun et al.
Fig. 36. Zasmidium colocasiae (K(M) IMI 311131). A. Conidiophore
fascicle. B. Conidiophores. C. Solitary conidiophores arising from a
supericial hypha. d. Conidia. Bar = 10 µm.
substomatal, 10–50 µm diam, composed of swollen hyphal
cells, brown. Mycelium internal and external; supericial
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 supericial 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
240
Fig. 37. Zasmidium deightonianum (K(M) IMI 7735). A. Conidiophores
and hyphae emerging through a stoma. B. Supericial hypha. C.
Supericial hypha with solitary conidiophore. d. Conidia. Bar = 10 µm.
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)
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.
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 38. Zasmidium extremorum
(S-F14686). A. Supericial hypha.
B. Conidiophore fascicles. C.
Conidiophore tips. d. Conidia. Bar
= 10 µm.
Stenella cercestidis (Deighton) U. Braun, Schlechtendalia 5:
54 (2000), nom. illeg. (ICN, Art. 53.1).
somewhat pointed to short obconically truncate, 0.5–1 µm
wide, hila slightly thickened and darkened.
Illustration: Deighton (1990: 570, ig. 1).
Holotype: Sierra leone: Njala (Kori), on Cercestis
congoensis, 25 Apr. 1934, F. C. Deighton (K(M) IMI 7735).
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; supericial 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
supericial 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
VOLUME 5 · NO. 2
Host range and distribution: On Cercestis congoensis,
Araceae, Africa (Sierra Leone).
Zasmidium
extremorum
(Syd.)
Schlechtendalia 20: 101 (2010).
(Fig. 38)
U.
Braun,
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, ig. 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; supericial hyphae emerging
through stomata, sparingly branched at right angles or
241
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Braun et al.
almost so, 1–3 µm wide, subhyaline to pale olivaceousbrown, 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 supericial hyphae not formed or at least not yet
observed), erect, straight to slightly curved or sinuous, nongeniculate 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-iliform, subacicular to
almost obclavate-cylindrical, (15–)40–300(–350) × 2–5 µm,
1- to pluriseptate, pale olivaceous to very pale olivaceousbrown, 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 darkenedrefractive.
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
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 (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 Toanun 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, ig. 1; 2011: 38, ig.
19), Phengsintham et al. (2013a: 75, igs 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,
242
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)
Literature: Crous & Braun (2003: 293), Kamal (2010: 69).
Illustration: Srivastava et al. (1995: 106, ig. 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,
IMA FUNGUS
Cercosporoid fungi 2
Note: This species belongs to the Cercospora apii s. lat.
complex.
ART I CLE
Host range and distribution: On Cocos nucifera, Arecaceae,
Asia (India, Uttar Pradesh).
Cercospora raphiae Deighton, Trans. Brit. Mycol.
Soc. 85: 741 (1985).
(Fig. 40)
Literature: Leung et al. (1997a), Crous & Braun (2003: 348).
Illustrations: Deighton (1987: 741, ig. 2), Leung et al. (1997a:
5, ig. a–g).
Description: Leaf spots amphigenous, subcircular to elliptical,
slighty swollen, 4–5 × 1–3 mm, sometimes conluent 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, lexuous, 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).
Fig. 39. Cercospora nucifera (based on Srivastava et al. 1995: 106,
ig. 1). A. Conidiophores. B. Conidia. Bar = 10 µm.
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: Uttar Pradesh: Gorakhpur, on Cocos
nucifera, Mar. 1990, R. K. Srivastava (GPU 1311, Gorakhpur
University, Dept. of Botany, India). Isotype: HCIO 30926.
VOLUME 5 · NO. 2
Doubtful,
species
excluded
and
insuficiently
known
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.
243
Braun et al.
ART I CLE
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
afinity 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, ig. 211 B), Deighton (1985:
740, ig. 1), Mulder & Holliday (1975: ig., unnumbered).
Fig. 40. Cercospora raphiae (K(M) IMI) 147583). A. Conidiophore
fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.
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).
Description: Leaf spots amphigenous, subcircular, elliptical to
irregular, small, often slightly depressed, brown to dark brown,
margin indeinite 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).
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.
244
Notes: This species seems to be conined to Elaeis
guineensis (Deighton 1985). Records on Carpentaria and
Raphia spp. are wrong and pertain to Pseudocercospora
species. Other records on Phoenix sp. and unidentiied palms
are doubtful and unproven. The generic afinity of C. elaeidis
is quite unclear. Due to pigmented conidia, this species has
to be excluded from Cercospora. Deighton (1985) placed it
IMA FUNGUS
Cercosporoid fungi 2
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 licualae Syd. & P. Syd., Philipp. J. Sci.
(Bot.) 9: 188 (1914).
Basionym: Cercospora palmicola Speg., Anales Soc. Ci.
Argent. 26: 72 (1888).
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: Lindau (1910: 88); Vassiljevsky & Karakulin (1937:
317), Chupp (1954: 429), Crous & Braun (2003: 303).
ART I CLE
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 afinity 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 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
Exsiccatae: Roum., Fungi Sel. Exs5188.
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 conirmed by
our re-examinations of the type collections. The true generic
afinity is, however, quite unclear.
Cercospora palmae-amazonensis Bat. & Cavalc.,
Anais XII Congr. Soc. Bot. Brasil: 385 (1964).
Illustration: Batista & Cavalcanti (1964: 386, ig. 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 unidentiied
palm, 17 Feb. 1961, A. C. Batista (IMUR 21246). Isotypes:
INPA, PC.
Host range and distribution: Unidentiied palm, Arecaceae,
South America (Brazil).
VOLUME 5 · NO. 2
Description: Leaf spots amphigenous, elliptical to somewhat
irregular, 10–20 mm long and 3–8 mm wide, brownish,
conluent. 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 olivaceousbrown, 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 romanzofianum [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 romanzofianum,
Phoenix canariensis, Arecaceae, Europe (Germany, Russia),
South America (Paraguay).
Notes: The generic afinity 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
245
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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, igs 1–3), Ellis (1971: 401,
ig. 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)
Illustration: Braun et al. (2006: 24, ig. 3).
Description: Leaf spots amphigenous, subcircular to irregular,
2–15 mm diam, pale to dark brown, inally greyish brown to
greyish white, margin indeinite or with a diffuse, irregular
darker border. Caespituli hypophyllous, inely 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-iliform, 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,
246
Fig. 41. Distocercospora livistonae (HAL 1875 F). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
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.
IMA FUNGUS
Cercosporoid fungi 2
Passalora
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 (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
ART I CLE
Key to Passalora species on Arecaceae
Passalora species on Arecaceae
passalora acrocomiae Guatimosim & R.W. Barreto,
Mycotaxon 122: 64 (2013).
(Fig. 42)
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.
Fig. 42. Passalora acromiae (BPI 432398). A. Conidiophore fascicle.
B. Conidia. Bar = 10 µm.
passalora eitenii R.B. Medeiros & Dianese, Mycotaxon
51: 509 (1994).
(Fig. 43)
Illustration: Medeiros & Dianese (1994: 511, plate 1, A–H).
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 lectotypiication 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.
VOLUME 5 · NO. 2
Description: Leaf spots amphigenous, mainly hypophyllous,
necrotic, dark brown to black, usually elliptical, rarely conluent,
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,
247
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Braun et al.
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.: Brasília, 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
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 (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 (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 (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 (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)
Illustration: Braun et al. (2006: 24, ig. 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 indeinite. Mycelium internal, occasionally with
supericial 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 conluent 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
248
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 olivaceousbrown, 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.
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 44. Pseudocercospora arecacearum (HAL 1876 F). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
Fig. 43. Passalora eitenii (UB, Coll. Mycol. 881). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
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)
Literature: Deighton (1985: 742, as Peudocercospora sp.),
Leung et al. (1997b).
Illustrations: Deighton (1985: 742, ig. 3; 1987: 403, ig. 1),
Leung et al. (1997b: 5, ig. a–f).
VOLUME 5 · NO. 2
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 olivaceousbrown, thin-walled, smooth, apex obtuse to subacute, base
short obconically truncate, hilum neither thickened nor
darkened.
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Fig. 46. Pseudocercospora coperniciae (HAL 1709 F). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
Illustration: Braun & Freire (2003: 310, ig. 12).
Fig. 45. Pseudocercospora carpentariae (K(M) IMI 305079). A.
Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
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)
250
Description: Leaf spots amphigenous, subcircular to
irregular, 3–15 mm diam or conluent and larger, pale to
greyish brown, inally dingy grey, margin indeinite 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 obclavatecylindrical, 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).
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Description: Leaf spots amphigenous, scattered, at irst
tiny spots, later small irregular patches, pale brown, margin
indeinite, 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, geniculatesinuous, 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 iliform, 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
[labelliformis], humilis), Arecaceae, Asia (Japan, Korea,
Taiwan), North America (USA, California, Florida), Hawaii.
pseudocercospora roystoneae U. Braun & Crous,
Mycol. Progr. 2: 203 (2003).
(Fig. 48)
Illustration: Braun et al. (2003a: 204, ig. 9).
Fig. 47. Pseudocercospora rhapisicola (NIAES 10300). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
pseudocercospora rhapisicola (Tominaga) Goh &
W.H. Hsieh, Trans. Mycol. Soc. Republ. China 4: 39
(1989).
(Fig. 47)
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, ig. 198), Guo & Hsieh
(1995: 243, ig. 205), Leung et al. (1997c: 17, ig. a–c), Guo
et al. (1998: 253, ig. 210).
VOLUME 5 · NO. 2
Description: Leaf spots lacking, indeinite (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 indeinite. 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, or with 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.
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Fig. 48. Pseudocercospora roystoneae (CUP 41024). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
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.)
Cercosporoid Fungi of India: 260 (2010).
(Fig. 49)
252
Kamal,
Fig. 49. 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].
IMA FUNGUS
Cercosporoid fungi 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, ig. 20–21), Ellis (1971: 145,
ig. 97 D), Kamal (2010: 261, ig. 40), Colmán et al. (2014:
849, igs A–E).
Exsiccatae: Cif., Mycol. 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
indeinite or darker. Caespituli amphigenous, punctiform,
sometimes conluent, 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 thinwalled 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 romanzofianum
[Arecastrum romanzofianum], Borassus labellifer, 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).
VOLUME 5 · NO. 2
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.
ART I CLE
Cercospora preisii Bubák, Ann. Mycol. 2: 400 (1904), valid
alternative name (ICN, Art 36.2).
Zasmidium
A single species.
Zasmidium caryotae (X.J. Liu & Y.Z. Liao) Kamal,
Cercosporoid Fungi of India: 239 (2010).
(Fig. 50)
Basionym: Stenella caryotae X.J. Liu & Y.Z. Liao, Acta
Microbiol. Sin. 20: 119 (1980).
Literature: Guo (1999: 358).
Illustration: Liu & Liao (1980: 120, ig. 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;
supericial 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 olivaceousbrown, 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).
253
Braun et al.
ART I CLE
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 afinity 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: igs 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 supericial
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 afinity
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).
Fig. 50. Zasmidium caryotae (HMAS 10262). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
Doubtful,
species
excluded
and
insuficiently
known
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, ig. 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;
supericial hyphae branched, 2.5–3.5 µm wide, septate, light
brown, inely verruculose. Conidiophores solitary, arising from
supericial 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
254
Illustration: Matsushima (1985: ig. 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: igs 447, 448).
Holotype: taiwan: Nan-Jen-Shan, isolated from a dead
rhachis of Areca catechu, Arecaceae, 27 Feb. 1986 (MFC6T196).
IMA FUNGUS
Cercosporoid fungi 2
rise to sporidesmium-like conidia formed singly or in short
chains. The general habit of this fungus suggests an afinity
to the Sporidesmium complex.
Asparagaceae (including Agavaceae, Convallariaceae, and Hostaceae)
ART I CLE
Notes: This species is quite distinct from Stenella and
Zasmidium. The conidiophores have a single terminal,
broadly truncate, unthickened conidiogenous locus giving
Cercospora
Key to Cercospora species on Asparagaceae
1
Conidia consistently broadly cylindrical or subcylindrical ................................................................................................ 2
Conidia acicular-iliform to obclavate-cylindrical ............................................................................................................. 5
2 (1)
Conidia subcylindrical, apex sometimes distinctly swollen; on Agave ......................................................... C. agavicola
Conidia never swollen at the apex; on other hosts ......................................................................................................... 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. loricola
Leaf spots distinct; conidiophores much longer, to 350 µm; conidia pluriseptate (1–18); on other hosts ....................... 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 (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, iliform to obclavate-cylindrical; on other hosts .......................................... 6
6 (5)
Stromata well-developed, 20–60 µm diam; conidia acicular-iliform 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 (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)
Literature: Groenewald et al. (2013: 144).
Illustration: Ayala-Escobar et al. (2005: 119, igs 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 greybrown 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 conluent and larger,
composed of swollen hyphal cells, 3–8 µm diam, brown.
VOLUME 5 · NO. 2
Conidiophores in small to moderately large lax fascicles,
erect, divergent, subcylindrical-iliform to lexuoussinuous, 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 irst 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
255
Braun et al.
ART I CLE
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 nongeniculate 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)
Synonyms: Cercospora caulicola G. Winter, J. Mycol. 1: 125
(1885) [syntypes: uSA: Missouri: Perryville, on Asparagus
oficinalis, 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 oficinalis, C.
Spegazzini (LPS 4966); K(M) IMI 247001 (slide)].
Cercospora asparagicola (Speg.) Vassiljevsky, in Vassiljevsky
& Karakulin, Fungi Imperfecti Parasitici (Hyphomycetes)
1: 296 (1937).
Fig. 51. Cercospora agavicola (HAL 1839 F). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
ive 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).
256
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, ig. 205), Hsieh & Goh (1990:
209, ig. 161), Guo et al. (2005: 175, ig. 122), Phengsintham
et al. (2013b: 51, ig. 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 geniculatesinuous 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,
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Host range and distribution: On Asparagus (densilorus
[Protasparagus
densilorus],
oficinalis,
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, ig. 123).
Fig. 52. Cercospora asparagi (HAL, Sacc., Mycoth. Ven. 1052). A.
Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
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 oficinalis, Sep. 1875 (PAD).
Isolectotypes: Sacc., Mycoth. Ven. 1052 (e.g. BPI 433642,
HAL, HBG).
VOLUME 5 · NO. 2
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, thinwalled, smooth, apex subacute, base truncate to obconically
truncate, about 2–3.5 µm wide, hila somewhat thickened and
darkened.
Holotype: China: Beijing, on Polygonatum oficinale, 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).
257
Braun et al.
ART I CLE
conidiogenous cells integrated, terminal, 10–25 µm long,
conidiogenous loci conspicuous, thickened and darkened,
(1.5–)2(–2.5) µm diam. Conidia solitary, broadly ellipsoidovoid, 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)
Literature: Chupp (1954: 37), Crous & Braun (2003: 188).
Fig. 53. Cercospora loricola (BPI 436450). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
Cercospora loricola Heald & F.A. Wolf, Mycologia 3:
17 (1911).
(Fig. 53)
Literature: Saccardo (1931: 883), Vassiljevsky & Karakulin
(1937: 297), Chupp (1954: 346), Crous & Braun (2003: 187).
Description: Leaf spots indistinct. Caespituli formed on
scapes, lower 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 conluent 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
258
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).
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 54. Cercospora fourcroyae (CUP 39835). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
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, ig. 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 inal conclusion and detailed
description. As Alstroemeria pertains to another plant family, it
remains doubtful if this collection has been correctly identiied.
Cercospora hostae Katsuki, Ann. Phytopathol. Soc.
Japan 20: 72 (1955).
(Fig. 55)
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, ig. 27), Guo et al. (2005:
178, ig. 125).
Description: Leaf spots amphigenous, scattered to conluent,
circular or subcircular to somewhat irregular, 1–10 mm diam,
VOLUME 5 · NO. 2
Fig. 55. Cercospora hostae (based on Shin & Kim 2001: 72, ig. 27).
A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10
µm.
to 20 mm when conluent, at irst brown to greyish brown
on the upper leaf surface, later greyish brown with narrow
reddish brown border, inally 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
259
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Braun et al.
darkened, 2–3 µm diam. Conidia solitary, acicular-iliform 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)
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, ig. 23),
Ellis (1976: 272, ig. 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 irst 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, olivaceousbrown, 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 olivaceousbrown, tips paler, darker in mass, wall thin to somewhat
260
Fig. 56. Cercospora maianthemi (HAL, Fuckel, Fungi Rhen Exs.
Suppl. 1631). A. Conidiophore fascicles. B. Conidiophore tips. C.
Conidia. Bar = 10 µm.
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 irst 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,
IMA FUNGUS
Cercosporoid fungi 2
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
conirmed that the conidia are at irst hyaline but become
somewhat pigmented with age, i.e. they are rather passaloralike. 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 afinity of this species.
Doubtful, excluded and insuficiently known
species
Cercospora montenegrina Bubák, Bot. Közlem. 14: 82
(1915).
Synonym: Cercosporina montenegrina (Bubák) 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-iliform 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).
ART I CLE
Switzerland, Ukraine), North America (USA, Michigan, Ohio,
Pennsylvania, New York, West Virginia, Wisconsin).
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 afinity 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 misidentiication.
Passalora
Key to Passalora species on Asparagaceae
1
Conidiophores verruculose; conidia medium brown and inely 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 (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 (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 (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
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Braun et al.
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Description: Leaf spots amphigenous, circular, 2–8 mm
diam, brown to dark red-brown, with a dark brown to
black border, frequently conluent, 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, inely 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 reexamination 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.
Fig. 57. Passalora cordylines (BPI 435225). A. Conidiophore
fascicle. B. Conidiophores. C. Conidia. Bars = 10 µm.
Passalora species on Asparagaceae
passalora cordylines (Henn.) Crous & M.P.S.
Câmara, Mycotaxon 68: 302 (1998).
(Fig. 57)
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, ig. 2).
262
passalora polygonati (Rostr.) U. Braun & Crous, in
Crous & Braun, Mycosphaerella and Anam.: 331
(2003).
(Fig. 58)
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,
inally very dark. Caespituli hypophyllous, punctiform,
scattered to aggregated, dark. Mycelium internal. Stromata
substomatal, about 20–50 µm diam, dark olivaceousbrown, 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.
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 58. Passalora polygonati (C, holotype). A. Conidiophore fascicle.
B. Conidiophore. C. Conidiophore tips. d. Conidia. Bar = 10 µm.
Fig. 59. Passalora polygonati-maximoviczii (TNS-F-60888). A.
Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.
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 multilorum, 1 Sep. 1902, Rostrup (C).
Host range and distribution: On Polygonatum (humile,
multilorum,
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.
VOLUME 5 · NO. 2
passalora
polygonati-maximoviczii
Poonam
Srivast., J. Living World 1: 117 (1994), nom. nov.
(as “(Katsuki) Poonam Srivast.”).
(Fig. 59)
Basionym: Cercosporidium polygonati-maximoviczii Katsuki,
Trans. Mycol. Soc. Japan 16: 12 (1975), nom. superl.
(ICN, Art. 52.1).
Synonyms: Cercospora polygonati-maximowiczii Togashi, Jap.
J. Bot. 2: 75 (1924) [syntypes: TNS-F-243801–243803].
Passalora polygonati-maximoviczii (Togashi) U. Braun &
Crous, Mycosphaerella andAnam.: 331 (2003), nom.
illeg. (ICN, Art. 53.1).
Literature: Vassiljevsky & Karakulin (1937: 298), Chupp
(1954: 353), Katsuki (1965: 43), Braun & Crous (2007: 60).
263
Braun et al.
ART I CLE
Illustration: Katsuki & Kobayashi (1975: 12, ig. 8).
Description: Leaf spots amphigenous, at irst 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, geniculatesinuous, 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 obclavatecylindrical, 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. plurilorum, 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. plurilorum,
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 polygonatimaximoviczii 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 superluous 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)
MycoBank MB809037
Basionym: Cercospora togashiana Katsuki & Urasawa, Ann.
Phytopathol. Soc. Jap. 15: 144 (1951).
Fig. 60. Passalora togashiana (TNS-F-243896). A. Conidiophore
fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.
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, Niiwamura, on Ophiopogon japonicus, 1 Jul. 1950, Y. Urasawa
(TNS-F-243896).
Literature: Chupp (1954: 355), Katsuki (1965: 43).
Description: Leaf spots indistinct or pale brown on the upper
side. Caespituli amphigenous, mainly hyphophyllous, effuse,
264
Host range and distribution: On Ophiopogon japonicus,
Asparagaceae, Asia (Japan).
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
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)
Literature: Deighton (1973: 111–113, as Cercospora yuccae).
Illustrations: Deighton (1973: 112, ig. 3, as Cercospora
yuccae), Crous et al. (1999: 189, ig. 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 geniculatesinuous, 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 roughwalled), 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).
Fig. 61. Passalora yuccae-gloriosae (IACM 3108). A. Conidiophore
fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
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 clariication of cercosporoid fungi on
Yucca spp. was made in Crous et al. (1999).
Pseudocercospora
Key to Pseudocercospora species on Asparagaceae
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 (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
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Braun et al.
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 (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
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 irst circular or elliptical in outline, later
elongated or conluent, 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 saucershaped 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).
Fig. 62. Pseudocercospora beaucarneae (BPI 71905). A. Conidioma.
B. Conidiophore tips. C. Conidia. Bar = 10 µm.
Pseudocercospora species on Asparagaceae
pseudocercospora beaucarneae (Pollack & D.F.
Farr) U. Braun, comb. nov.
MycoBank MB809011
(Fig. 62)
Basionym: Stigmina beaucarneae Pollack & D.F. Farr,
Mycotaxon 8: 193 (1997).
Illustrations: Pollack & Farr (1979: 198, igs 2–7, 199, igs
8–13).
266
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)
Basionym: Cercospora concentrica Cooke & Ellis, Grevillea
5: 90 (1877).
IMA FUNGUS
Cercosporoid fungi 2
Illustrations: Deighton (1973: 108, ig. 1, 110, ig. 2),
Sivanesan (1984: 24, ig. 137).
Fig. 63. Pseudocercospora concentrica (NY 1102862). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
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., Newield,
on Yucca ilamentosa, 1 June 1874, W. A. Kellerman
2150 (NY 838826); isotype: NY 1102861].
VOLUME 5 · NO. 2
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Literature: Saccardo (1886: 479), Vassiljevsky & Karakulin
(1937: 297), Chupp (1954: 344), Sivanesan (1984: 248),
Simay (1987), Kobayashi (2007).
Description: Leaf spots amphigenous, also at stems, elliptical,
usually to 15 mm diam, occasionally to 40 × 10 mm, later
often conluent, 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, thinwalled, 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 olivaceousbrown, 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
ine 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: Newield, on Yucca ilamentosa, 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,
ilamentosa, 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
267
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Braun et al.
Fig. 64. Pseudocercospora cordylines
(B 700014864). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia.
Bar = 10 µm.
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, ig. 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 inal 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
Schlechtendalia 1: 29 (1998).
(Fig. 64)
U.
Literature: Saccardo (1906: 611), Chupp (1954: 344).
268
Braun,
Illustration: Braun (1998b: 27, ig. 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 lexuous,
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, obclavatecylindrical, (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).
IMA FUNGUS
Cercosporoid fungi 2
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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. 65. Pseudocercospora maianthemi (based on Kirschner &
Piepenbring 2008: 26–27, igs 5–6). A. Conidiophore fascicle. B.
Conidiophores. C. Conidia. Bar = 10 µm.
Basionym: Cercospora yuccae Cooke, Grevillea 7: 35 (1878).
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.
Illustration: Crous et al. (1999: 189, ig. 14).
pseudocercospora maianthemi R. Kirschner, Mycol.
Progr. 7: 24 (2008).
(Fig. 65)
Illustrations: Kirschner & Piepenbring (2008: 26–27, igs 5–6).
Description: Leaf spots amphigenous, pale to dark brown
with irregular margin, conluent to streaks to 40 mm long
and 5 mm wide. Caespituli hypophyllous. Mycelium internal,
intercellular; hyphae branched, 2–5 µm wide, septate, pale
VOLUME 5 · NO. 2
Literature: Saccardo (1886: 479), Vassiljevsky & Karakulin
(1937: 297), Chupp (1954: 355).
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, welldeveloped, 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
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Braun et al.
Fig. 66. Scolecostigmina yuccae (BPI 435122). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
Fig. 67. Zasmidium agavicola (INIFAT C87/431). A. Conidiophores.
B. Conidia. Bar = 10 µm.
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.
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 afinity 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).
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).
270
IMA FUNGUS
Cercosporoid fungi 2
Zasmidium
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 supericial 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 (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. García)
U. Braun & R.F. Castañeda, comb. nov.
MycoBank MB809038
(Fig. 67)
Basionym: Stenella agavicola R.F. Castañeda & L. García,
Fungi Cubenses 3: 13 (1988).
Illustration: Castañeda (1988: ig. 25).
Description: Colonies amphigenous, forming circular,
velutinous patches, olivaceous to dark brown. Mycelium
internal and external; supericial hyphae branched, septate,
2–3 µm wide, brown, smooth to verruculose. Stroma lacking.
Conidiophores solitary, arising from supericial 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-iliform, 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)
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).
VOLUME 5 · NO. 2
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Key to Zasmidium species on Asparagaceae
Description: Leaf spots variable, irregularly shaped, small,
1–5 mm diam, to very large, to 80 mm diam, pale strawcoloured, 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; supericial hyphae branched, mostly at right
angles, septate, 1–3 µm wide, hyaline to pale brown, thinwalled, 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-iliform, 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 ellipsoidovoid to obclavate-cylindrical, long conidia acicular, (8–)12–
180 × 2–4 µm, 0–12-septate, hyaline to pale olivaceous, thinwalled, 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
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Braun et al.
Fig. 68. Zasmidium dracaenae (K(M)
IMI 53368). A. Hypha. B. hypha with
conidiophore. C. Conidiophore fascicle. d.
Conidiophore tips. e. Conidia. Bar = 10 µm.
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)
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.
272
Literature: Saccardo (1892: 655), Vassiljevsky & Karakulin
(1937: 217–118), Chupp (1954: 355), Braun (1998: 377).
Illustrations: Braun (1993a: 242, ig. 19; 1998: 375, ig. 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
conluent and larger, at irst yellowish or yellowish brown,
later with pale centre and narrow to moderately wide, brown
margin, inally often large leaf segments discoloured, brown,
necrotic, sometimes with shot-hole symptoms. Caespituli
usually hypophyllous, punctiform, greyish to pale reddish.
Mycelium internal and external; supericial 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
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 69. Zasmidium subsanguineum
(HAL, Krieger, Fungi Saxon. Exs.
1293). A. Supericial hyphae. B.
Conidiophore fascicles. C. Solitary
conidiophores. d. Conidiophores. e.
Conidia. Bar = 10 µm.
from supericial hyphae, lateral, rarely terminal, erect,
straight, lexuous, geniculate-sinuous, simple or occasionally
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
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 (1)
On Disporum ..................................................................................................................................................... C. dispori
On Gloriosa ................................................................................................................................................... C. gloriosae
Cercospora species on Colchicaceae
Illustration: Shin & Kim (2001: 63, ig. 22).
Cercospora dispori Togashi & Maki, Trans. Sapporo
Nat. Hist. Soc. 17: 98 (1942).
(Fig. 70)
Description: Leaf spots amphigenous, scattered to conluent,
usually vein-limited, subcircular, elliptical to angular, brown,
ranging from yellowish brown, reddish brown to dark brown,
5–15 mm diam, conluent, later entire lealets becoming
necrotic, turning brown to blackish. Caespituli amphigenous,
punctiform, scattered, dark brown to blackish. Mycelium
Literature: Chupp (1954: 345), Katsuki (1965: 42), Kim & Shin
(1999a), Shin & Kim (2001: 62), Crous & Braun (2003: 164).
VOLUME 5 · NO. 2
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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 conimed 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)
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, ig. 17).
Fig. 70. Cercospora dispori (TNS-F-243895). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
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 iliform 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.
274
Description: Leaf spots amphigenous, scattered, circular,
subcircular to somewhat angular-irregular, 2–20 mm diam, at
irst 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 geniculatesinuous 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 obclavatecylindrical, longer ones subacicular to distinctly acicular, 30–
120 × 2.5–5 µm, 3–11-septate, hyaline, thin-walled, smooth,
apex obtuse to acute, sometimes lagelliform, 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
IMA FUNGUS
Cercosporoid fungi 2
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Fig. 71. Cercospora gloriosae (CUP 39896). A. Conidiophore
fascicles. B. Conidiophore tip. C. Conidia. Bar = 10 µm.
Fig. 72. Cercospora iphigeniae (based on Patwardhan & Sathe
1966: 152, ig. 2). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.
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 inluenced by external conditions, and
they are barely suficient 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 conirmation.
VOLUME 5 · NO. 2
Cercospora iphigeniae Patwardhan & Sathe, Sydowia
19: 151 “1965”(1966).
(Fig. 72)
Literature: Crous & Braun (2003: 228), Kamal (2010: 53).
Illustration: Patwardhan & Sathe (1966: 152, ig. 2).
275
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Braun et al.
Description: Leaf spots broadly fusiform, pale grey with
black margin. Caespituli mainly epiphyllous. Stromata
depressed globose to lattened, 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 reexamination. The generic afinity 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
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 (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 (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 (3)
With deinite leaf spots, subcircular to irregular, 1–4 mm diam;
conidia occasionally in short chains; on Commelina longifolia ..................................... C. commelinae-salicifoliae
Entire lealets turning brown, inally withering; conidia consistently solitary;
on Murdannia nudilora ........................................................................................................................ C. nudilorae
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, igs 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,
276
unbranched, 25–235 × 3–4.5 µm, 0–11-septate, brown, thinwalled, 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 iliform, straight
to curved, 30–275 × 2–2.5 µm, 2–22-septate, hyaline, thinwalled, 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.
IMA FUNGUS
Cercosporoid fungi 2
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Cercospora commelinae-salicifoliae A.K. Kar & M.
Mandal, Indian Phytopathol. 26: 675 “1973”(1974).
(Fig. 73)
Literature: Crous & Braun (2003: 133), Kamal (2010: 35).
Illustration: Kar & Mandal (1974: 676, ig. 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 obclavatecylindrical, 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.
Fig. 73. Cercospora commelinae-salicifoliae (K(M) IMI 138302). A.
Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.
Cercospora commelinicola Chupp ex U. Braun,
Cryptog. Mycol. 20: 156 (1999).
(Fig. 74)
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, ig. 1), Guo et al. (2005: 65,
ig. 40).
Exsiccatae: Sydow, Fungi Exot. Exs. 1238.
Description: Leaf spots amphigenous, subcircular to irregular,
1–15 mm diam or conluent 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,
VOLUME 5 · NO. 2
Fig. 74. Cercospora commelinicola (HBG, Syd., Fungi Exot. Exs.
1238). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia.
Bar = 10 µm.
277
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Braun et al.
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 olivaceousbrown, 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 nudilorae Chupp,
Cercospora: 117 (1954).
(Fig. 75)
Monograph
of
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 geniculatesinuous, 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.
278
Fig. 75. Cercospora nudilorae (CUP 35213). A. Conidiophore
fascicles. B. Conidiophore tip. C. Conidia. Bar = 10 µm.
Lectotype (designated here, MycoBank MBT178143):
Bermuda: Devonshire Marsh, On Murdannia nudilora
[Commelina nudilora], 12 Jan. 1926, H. H. Whetzel (CUP
35213). Isolectotype: CUP 40395.
Host range and distribution: On Murdannia nudilora,
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. nudilorae. 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.
IMA FUNGUS
Cercosporoid fungi 2
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Cercospora polliae-japonicae Y.L. Guo & Y. Jiang,
Mycosystema 19: 302 (2000).
(Fig. 76)
Literature: Crous & Braun (2003: 331), Guo et al. (2005: 66).
Illustrations: Guo & Jiang (2000b: 303, ig. 1), Guo et al.
(2005: 657, ig. 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 insuficiently 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
Fig. 76. Cercospora polliae-japonicae
Conidiophores. B. Conidia. Bar = 10 µm.
(HMAS
77628).
A.
was in fact a species of Septoria. An examination of a
South African collection (PREM 32777) identiied by Chupp
& Doidge (1948) to be C. commelinae and the type from B
conirmed that the fungus would be better placed in Septoria.
Pseudocercospora
Key to Pseudocercospora species on Commelinaceae
1
In vivo with external mycelium; stromata lacking or almost so; with solitary conidiophores
arising from supericial hyphae; on Pollia japonica, Taiwan ........................................................................ p. polliae
Supericial mycelium in vivo lacking; conidiophores fasciculate, arising from small
to well-developed stromata; on other hosts ............................................................................................................. 2
VOLUME 5 · NO. 2
279
Braun et al.
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2 (1)
Conidiophores only 0–3-septate; on Amischotolype hispida, Taiwan ........................................................... p. forrestiae
Conidiophores pluriseptate; on Commelina erecta, Trinidad ................................................................. p. maracasensis
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, thinwalled, 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 (NTUPPE [ 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)
Basionym: Cercospora maracasensis R.E.D. Baker & W.T.
Dale, Mycol. Pap. 33: 103 (1951).
Literature: Chupp (1954: 117), Crous & Braun (2003: 266–
267).
Fig. 77. Pseudocercospora forrestiae (NTU-PPE, holotype). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
Pseudocercospora species on Commelinaceae
pseudocercospora forrestiae Goh & W.H. Hsieh,
Trans. Mycol. Soc. Republ. China 2: 130 (1987).
(Fig. 77)
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, ig. 41), Guo & Hsieh
(1995: 61, ig. 56), Guo et al. (1998: 75, ig. 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
280
Description: Leaf spots amphigenous, subcircular to
irregular or somewhat angular-irregular, 4–30 mm diam, at
irst dull greenish, greyish green or greenish brown, later
gradually changing to yellowish brown or brown, inally
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 lattened
above, 10–60 µm diam, medium to dark olivaceousbrown, 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-iliform, 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, thinwalled, smooth, apex obtuse or subobtuse, base short
to long obconically truncate, (1.5–)2(–2.5) µm wide, hila
unthickened, not darkened.
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 78. Pseudocercospora maracasensis (CUP 35326). A.
Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
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)
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, ig. 42), Guo & Hsieh
(1995: 63, ig. 57), Guo et al. (1998: 76, ig. 59).
Description: Leaf spots indistinct to circular or elliptical,
10–15 mm diam, centre greyish brown, margin dark brown.
VOLUME 5 · NO. 2
Fig. 79. Pseudocercospora polliae (NTU-PPE, holotype). A.
Conidiophore fascicle. B. Conidiophores and supericial hypha with
solitary conidiophore emerging through a stoma. C. Conidia. Bar =
10 µm.
Caespituli amphigenous, mainly hypophyllous. Mycelium
internal and external; supericial 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 supericial hyphae, lateral, erect, straight to
somewhat curved, subcylindrical to somewhat geniculatesinuous, 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-iliform, 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
Commelinaceae, Asia (China, Japan, Taiwan).
japonica,
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Costaceae
Pseudocercospora
Key to Pseudocercospora species on Costaceae
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)
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 supericial hyphae, unbranched or sparingly branched,
1–3 µm wide, subhyaline to pale olivaceous, septate, thinwalled, 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
supericial 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 iliform, 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 supericial hyphae with solitary
conidiophores have been observed.
282
Fig. 80. Pseudocercospora costi (ILL 15148). A. Supericial hypha.
B. Supericial hypha with solitary conidiophore. C. Conidiophore
fascicles. d. Conidiophores. e. Conidia. Bar = 10 µm.
IMA FUNGUS
Cercosporoid fungi 2
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pseudocercospora costina (Syd. & P. Syd.) Deighton,
Mycol. Pap. 140: 135 (1976).
(Fig. 81)
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, ig. 81, 137, ig. 82), Singh
et al. (1984: 876, ig. 1), Hsieh & Goh (1990: 356, ig. 273),
Guo & Hsieh (1995: 356, ig. 300), Guo et al. (1998: 375, ig.
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 indeinite.
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-iliform,
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
VOLUME 5 · NO. 2
Fig. 81. Pseudocercospora costina (K(M) IMI 82089). A. Conidiophore
fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
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 inal typiication. 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.
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Cyperaceae
Cercospora
Key to Cercospora species on Cyperaceae
1
Conidia strictly acicular, base of the conidia truncate ..................................................................................................... 2
Conidia not strictly acicular, conidia at least partly obclavate-cylindrical or iliform
and base of the conidia at least partly obconically truncate ..................................................................................... 4
2 (1)
With distinct leaf spots; conidiophores short, 20–45 µm; on Cyperus fuscus, Europe
.............................................................. C. cyperi-fusci (see “doubtful, excluded and insuficiently known species”)
Leaf spots lacking; conidiophores much longer, 30–120 µm; on Kyllinga erecta, Africa ................................................. 3
3 (2)
Conidiophores often branched; conidia 20–120 × 1.5–3 µm, 1–12-septate ................................................. C. kyllingae
Conidiophores unbranched; conidia longer and wider, 60–250 × 3–4 µm, 8–21-septate ......................... C. kyllingicola
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 (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 (5)
Conidiophores very short, 5–20 µm, 0–1-septate ........................................................................................................... 7
Conidiophores longer, 10–35(–60) µm, 0–4-septate ....................................................................................................... 8
7 (6)
Conidiophores 2–5 µm wide, hila 1–2 µm wide; conidia narrowly cylindrical-iliform
to acicular; on Cyperus spp. ............................................................................................................... C. cyperigena
Conidiophores narrower, 1.5–3 µm, hila 1 µm diam; conidia narrowly obclavate-iliform;
on Eleocharis spp. ............................................................................................................................ C. eleocharidis
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)
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 laxilora, 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.”.
284
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, ig. 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 indeinite 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,
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Host range and distribution: On Carex (albursina,
alopecoidea, arctata, bebbii, bicknellii, bromoides, castanea,
cephalophora, crawfordii, cristatella, davisii, delexa,
folliculata, granularis, gravida, interior, intumescens, laxilora,
lupulina, microptera, normalis, orbicularis, pensylvanica,
pilulifera [oederi], plantaginea, projecta, pubescens,
retrorsa, rosea [convoluta], sartwelliana, stipata, stricta,
subcapitatus, tenera, vulpinoidea [muhlenbergii], Carex sp.),
Cyperus (articulatus [istulosus], esculentus, iliculmis, mitis
[subcapitatus], rotundus, Cyperus sp.), Eleocharis acutangula
[istulosa], 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 conspeciicity
of species occurring on these two host genera. Material from
the Netherlands appropriate for a neotypiication was not
found. Therefore, we prefer to postpone a neotypiication.
Cercospora cyperi Sawada, Trans. Nat. Hist. Soc.
Taiwan 17: 179 (1914).
(Fig. 83)
Fig. 82. Cercospora caricis (LEP, Ellis & Everh., Fungi Columb.
1170). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia.
Bar = 10 µm.
erect, straight, subcylindrical to moderately geniculate-sinuous,
unbranched, 10–45 × 3–6 µm, 0–3(–4)-septate, olivaceousbrown 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 obclavatecylindrical 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).
VOLUME 5 · NO. 2
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, igs
19–20), Hsieh & Goh (1990: 104, ig. 78), Guo et al. (2005:
106, ig. 71).
Description: Leaf spots amphigenous, irregular to oblong,
1–25 × 0.5–5 mm, brown, margin indeinite. Caespituli
hypophyllous, punctiform, ine, 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
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[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, iliculmis,
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).
Fig. 83. Cercospora cyperi (NTU-PPE, holotype). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
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 olivaceousbrown or brown throughout or paler towards the tip, thinwalled, 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.
286
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 obclavatecylindrical 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, ig. 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 ind out if these differences are suficient to
maintain two distinct species. Cercospora cyperi seems to
be conined 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)
Literature: Braun & Urtiaga (2012: 302–303).
Illustration: Crous & Braun (2003: 151, ig. 8).
Description: Leaf spots amphigenous, subcircular to ellipticalfusiform, 1–8 mm diam, occasionally conluent, centre
yellowish olivaceous to light brown, with a narrow brown
margin. Caespituli hypophyllous, inely punctiform, often
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 85. Cercospora eleocharidis (BPI 436100). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
conidiogenous loci conspicuous, 1–2 µm wide, thickened
and darkened. Conidia solitary, narrowly cylindrical-iliform
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).
Fig. 84. Cercospora cyperigena (K(M) IMI 259773). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bars = 10 µm.
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, thinwalled, smooth; conidiogenous cells integrated, terminal or
conidiophores reduced to conidiogenous cells, 5–15 µm long,
VOLUME 5 · NO. 2
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-iliform to subacicular conidia.
Cercospora eleocharidis Davis, Trans. Wisconsin
Acad. Sci. 24: 300 (1929).
(Fig. 85)
287
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Braun et al.
Fig. 86. Cercospora glauciana (PDD 46385). A. Conidiophore fascicle. 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, inally the whole leaf blade sometimes turning
brown. Caespituli amphigenous, inely 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.
288
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 iliform, 20–70 ×
1.5–2.5 µm.
Cercospora glauciana Viégas, Bol. Soc. Bras. Agron.
8: 27 (1945).
(Fig. 86)
Literature: Chupp (1954: 192), Braun & Sivapalan (1999: 5),
Crous & Braun (2003: 198), Kirschner & Piepenbring (2006:
214).
Illustrations: Braun & Sivapalan (1999: 4, ig. 2), Kirschner &
Piepenbring (2006: 214, ig. 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
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
with pale centre, greyish white, surrounded by a darker
margin. Caespituli hypophyllous, ine, 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-iliform, 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.), unidentiied 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)
Literature: Crous & Braun (2003: 240).
Illustration: Yen & Gilles (1970: 80, ig. 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 lexuous, 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).
VOLUME 5 · NO. 2
Fig. 87. Cercospora kyllingae (based on Yen & Gilles 1970: 80, ig.
4). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar =
10 µm.
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)
Literature: Crous & Braun (2003: 240).
289
Braun et al.
ART I CLE
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 suficient 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)
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, ig. 23), Braun (1993a: 237,
ig. 3; 1995: 123, ig. 107).
Exsiccatae: Syd., Fungi Exot. Exs. 444.
Fig. 88. Cercospora kyllingicola (based on Yen & Gilles 1970: 82, ig.
5). A. Conidiophore fascicle and conidiophore. B. Conidiophore tips.
C. Conidia. Bar = 10 µm.
Illustration: Yen & Gilles (1970: 82, ig. 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
lexuous, 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.
290
Description: Leaf spots amphigenous, subcircular, elliptical,
oblong, irregular, 1–4 mm diam, scattered to conluent,
pale brown, margin indeinite 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, iliform,
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
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 89. Cercospora uredinophila (PAD, holotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
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 insuficiently known
species
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-iliform 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).
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).
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: uSA: Alabama: Auburn, on Rhynchospora
glomerata, Cyperaceae, 27 Aug. 1891, B. M. Duggar (CUP-A
2034).
Holotype: romania: Cîrjoaia, Zbereni forest near Cotnari, on
Cyperus fuscus, 30 Jul. 1957, C. Sandu-Ville (not traced).
Host range and distribution: Only known from the type
collection.
Host range and distribution: On Cyperus fuscus, Cyperaceae,
Europe (Romania).
VOLUME 5 · NO. 2
291
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Braun et al.
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).
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 superluous 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
Host range and distribution: On Eleocharis palustris,
Schoenoplectus (lacustris, litoralis), Scirpus acutus,
Cyperaceae, Asia (Uzbekistan), Europe (Germany, Great
Britain, Hungary), North America (USA, Wisconsin).
A single species.
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.
Basionym: Phaeoramularia cyperi Durgas Gupta, Padhy &
Chowdhry, Curr. Sci. 50: 140 (1981).
passalora cyperi (Durgas Gupta, Padhi & Chowdhry)
U. Braun & Crous, in Crous & Braun, Mycosphaerella
and Anam.: 450 (2003).
Literature: Singh (2003: 527), Kamal (2010: 116).
Illustration: Gupta et al. (1981: 140, ig. 1).
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. superl.
Pleospora scirpi (Rabenh.) Ces. & De Not., Comment. Soc.
292
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, olivaceousbrown, 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.
IMA FUNGUS
Cercosporoid fungi 2
loci and conidial hila are unknown. Type material was not
available for re-examination.
Zasmidium
ART I CLE
Notes: This species is only tentatively maintained in
Passalora (including Phaeoramularia). The conidia are rather
cladosporioid, but details of the structure of the conidiogenous
Key to Zasmidium species on Cyperaceae
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)
Basionym: Stenella gahniae McKenzie, New Zealand J. Bot.
20: 248 (1982).
Illustration: McKenzie (1982: 249–250, igs 4–5).
Description: Colonies epiphyllous. Mycelium internal and
external; supericial hyphae branched, septate, 1.5–2.5
µm wide, pale straw-coloured, thin-walled, minutely
verruculose. Stromata absent. Conidiophores solitary,
arising from supericial hyphae, lateral, originating from
brown, smooth, sometimes swollen hyphal cells, 10–
30 µm wide, erect, straight or lexuous, 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.
VOLUME 5 · NO. 2
Fig. 90. Zasmidium gahniae (PDD 38658). A. Solitary conidiophore
arising from supericial hypha. B. Conidiophore. C. Conidia. Bar =
10 µm.
293
ART I CLE
Braun et al.
Zasmidium scleriae (McKenzie) U. Braun & McKenzie,
comb. nov.
MycoBank MB809040
(Fig. 91)
Basionym: Stenella scleriae McKenzie, New Zealand J. Bot.
20: 250 (1982).
Illustration: McKenzie (1982: 251, ig. 6).
Description: Colonies amphigenous, mainly epiphyllous.
Mycelium internal and external; supericial hyphae,
branched, septate, 1.5–5 µm wide, pale straw-coloured,
thin-walled, minutely verruculose. Stromata absent.
Conidiophores solitary, arising from supericial hyphae,
lateral, erect, originating from brown, smooth, sometimes
swollen cells, 5–10 µm diam, straight to lexuous 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.
Fig. 91. Zasmidium scleriae (PDD 40911). A. Solitary conidiophores
arising from supericial hypha. B. Conidiophore. C. Conidia. Bar =
10 µm.
Dioscoreaceae
Cercospora
Key to Cercospora species on Dioscoreaceae
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 (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
Illustration: Pons & Sutton (1988: 14, ig. 5).
Cercospora aragonensis Durrieu, Bull. Trimestriel
Soc. Mycol. France 80: 169 (1964).
(Fig. 92)
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,
Literature: Pons & Sutton (1988: 15), Crous & Braun (2003: 63).
294
IMA FUNGUS
Cercosporoid fungi 2
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.
ART I CLE
Host range and distribution: On Dioscorea pyrenaica
Dioscoreaceae, Europe (Spain).
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, ig. 72), Guo et al. (2005: 107,
ig. 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).
Fig. 92. Cercospora aragonensis (K(M) IMI 255661). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
short, subcylindrical-conical or only slighty geniculatesinuous, 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).
VOLUME 5 · NO. 2
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)
295
Braun et al.
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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, ig. 1), Yen & Lim (1980: 212, ig.
15), Sivanesan (1985: 744–746, igs 1–3), Pons & Sutton
(1988: 8–11, igs 1–3), Nakashima et al. (2011: 256, ig. 1c,d).
Description: Leaf spots amphigenous, subcircular, elliptical to
angular-irregular, 0.5–10 mm diam, at irst brown, later with
pale centre, inally 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 obclavatesubcylindrical, 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).
Fig. 93. 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)].
296
Notes: Cercospora dioscoreae-pyrifoliae can currently only
be circumscribed and used as a heterogeneous morphotaxon. 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,
IMA FUNGUS
Cercosporoid fungi 2
Doubtful, excluded and insuficiently 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, ig. 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 lexuous, 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.
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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”.
Holotype: India: Assam: Golaghat, on Dioscorea alata,
Dioscoreaceae, 21 Nov. 1977, U. N. Saikia (HCIO 32660).
Host range and distribution:
Dioscoreaceae, Asia (India).
On
Dioscorea
alata,
Notes: An attempt to obtain type material of this species
on loan from HCIO was not successful. The generic afinity
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
Conidiophores very long and usually multibranched and pluriseptate, 50–490(–625) × 3–6.5 µm;
conidia with thickened inner wall layer, usually inely verruculose;
on numerous Dioscorea spp. ............................................................................................................ d. pachyderma
Conidiophores much shorter, 15–80 × 3–10 µm, shorter ones 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)
Literature: Crous & Braun (1996: 247).
Illustration: Crous & Braun (1994: 209, ig. 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, thinwalled, smooth; conidiogenous cells integrated, terminal
or conidiophores reduced to conidiogenous cells, about
VOLUME 5 · NO. 2
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)
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,
297
Braun et al.
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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, greyolivaceous 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 angularirregular in outline. Conidiophores in mostly lax to moderately
dense fascicles, about 5–30, arising from internal hyphae
or small stromata, through stomata, erect, subcylindricaliliform, lexuous 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 or sigmoid, (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 inely verruculose, apex obtuse, base subtruncate
to short obconically truncate, 1–2 µm wide, hila unthickened to
slightly thickened and not to slightly darkened-refractive.
Fig. 94. Distocercospora africana (PREM 10125). A. Conidiophore
fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
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, ig. 2), Pons & Sutton
(1988: 61–64, igs 28–31), Shin & Kim (2001: 120, ig. 51),
Kirschner et al. (2004: 62, igs 10–13), Kamal (2010: 276,
ig. 47).
298
[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
IMA FUNGUS
Cercosporoid fungi 2
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Fig. 95. Distocercospora
pachyderma (S-F37683).
A. Conidiophore fascicles.
B. Conidiophore tips. C.
Conidia. Bar = 10 µm.
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, lexuous
conidiophores with dispersed sympodial conidiogenous loci,
and the distoseptate conidia (also prominent in culture), are
deinitive characters for this species.
Passalora
Key to Passalora species on Dioscoreaceae
1
Supericial hyphae with solitary conidiophores in vivo present (mycovellosiella-like) ..................................................... 2
Supericial hyphae and solitary conidiophores in vivo lacking, conidiophores only fasciculate ...................................... 4
2 (1)
Conidiophores long, 40–120 µm; on Dioscorea alata, China ................................................................ p. dioscoreigena
Conidiophores shorter, 5–50 µm; on Dioscorea nipponica ............................................................................................. 3
VOLUME 5 · NO. 2
299
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Braun et al.
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 (1)
Conidia catenate; on Dioscorea spp., widespread ..................................................................................... p. dioscoreae
Conidia formed singly; on Dioscorea spp., China ........................................................................................................... 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)
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, ig. 241 C), Pons & Sutton
(1988: 34–36, igs 14–16), Guo et al. (2003: 141, ig. 88).
Exsiccatae: Ellis & Everh., North Amer. Fungi 2471.
Fig. 96. Passalora dioscoreae (NY 838293). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
300
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
IMA FUNGUS
Cercosporoid fungi 2
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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, loribunda,
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
Mycosystema 20: 301 (2001).
(Fig. 97)
Y.L.
Guo,
Literature: Guo et al. (2003: 81), Crous & Braun (2003: 451).
Illustrations: Guo (2001b: 301, ig. 1), Guo et al. (2003: 82,
ig. 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)
VOLUME 5 · NO. 2
Fig. 97. Passalora dioscoreae-subcalvae (HMAS 78803). A.
Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.
Literature: Crous & Braun (2003: 451).
Illustrations: Guo (2001b: 302, ig. 2), Guo et al. (2003: 83,
ig. 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
301
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Braun et al.
Fig. 98. Passalora dioscoreicola (HMAS 79141). A. Conidiophores.
B. Conidia. Bar = 10 µm.
× 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)
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, ig. 2), Guo et al. (2003: 27, ig.
13).
302
Fig. 99. Passalora dioscoreigena (HMAS 79139). A. Supericial
hypha with solitary conidiophore. B. Conidiophore fascicle. C.
Conidia. Bar = 10 µm.
Description: Leaf spots amphigenous, circular to irregular, 1–5
mm diam, often conluent, yellowish brown to reddish brown,
margin indeinite, with pale olivaceous-brown halo on the
upper leaf surface, effuse, grey, greyish brown to dark brown
below. Caespituli hypophyllous. Mycelium internal and external;
supericial 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 supericial 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, thinwalled, smooth; conidiogenous cells integrated, terminal and
intercalary, sympodial, conidiogenous loci conspicuous, 2–3
µm diam, thickened and darkened. Conidia catenate, in simple
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
or branched chains, obclavate-cylindrical, 25–110 × 4.5–6.5
µm, 1–6-septate, pale olivaceous to olivaceous-brown, thinwalled, 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)
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, igs 20–22).
Description: Leaf spots amphigenous, circular to angularirregular, scattered to conluent, 3–35 mm diam, at irst
yellowish, later pale to darker brown, margin narrow, darker.
Caespituli hypophyllous, effuse, greyish to grey-brown.
Mycelium internal and external; supericial 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 supericial
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.
VOLUME 5 · NO. 2
Fig. 100. Passalora tranzschelii var. tranzschelii (LE 505). A.
Supericial hypha with solitary conidiophore. B. Conidiophore
fascicle. C. Conidiophore. d. Conidia. Bar = 10 µm.
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)
303
Braun et al.
ART I CLE
Illustration: Guo (2011: 869, ig. 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 dioscoreaenipponicae, 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 insuficiently known
species
Cercosporidium dioscoreae S. Singh, Plant Parasitic
Fungi of Gorakhpur [University of Gorakhpur, PhD
thesis (1994)], not effectively published (ICN, Art.
30.8).
Fig. 101. Passalora tranzschelii var. chinensis (HMAS 65914). A.
Supericial hyphae with solitary conidiophores. B. Conidia. Bar = 10
µm.
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
304
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 (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 (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 (3)
Conidia narrow, 1–3 µm .................................................................................................................................................. 5
Conidia much broader, 2.5–8 µm .................................................................................................................................... 6
5 (4)
Stromata lacking; mycelium internal and external; supericial 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;
supericial hyphae with solitary conidiophores lacking ....................................................................... p. hiratsukana
6 (4)
Stromata lacking; conidiophores long, 20–150 µm; conidia smooth ........................................................................ p. ubi
IMA FUNGUS
Cercosporoid fungi 2
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 (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
ART I CLE
Stromata developed, 10–90 µm diam; conidiophores much shorter, 5–65 µm,
if longer conidia verruculose .................................................................................................................................... 7
Pseudocercospora species on Dioscoreaceae
pseudocercospora carbonacea (L.E. Miles) N. Pons
& B. Sutton, Mycol. Pap. 160: 26 (1988).
(Fig. 102)
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, ig. 194 A), Yen et al. (1982: 52,
ig. 8), Little (1987a: ig., unnumbered), Pons & Sutton (1988:
29, ig. 12), Phengsintham et al. (2003a: 108, igs 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 supericial hyphae,
branched, 2–3 µm wide, septate, pale olivaceous, thinwalled, 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 supericial
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.
VOLUME 5 · NO. 2
Fig. 102. Pseudocercospora carbonacea (NY 936951). A.
Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
305
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Braun et al.
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], triida, 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 supericial 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)
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, igs 16–17), Sivanesan
(1984: 208, ig. 109), Little (1987b: ig., unnumbered), Pons
& Sutton (1988: 19–21, igs 6–8), Guo & Hsieh (1995: 86, ig.
78), Guo et al. (1998: 102, ig. 82), Shin & Kim (2001: 179,
ig. 78).
Exsiccatae: Poelt & Scheuer, Reliqu. Petrak. 2797.
Description: Leaf spots subcircular, elliptical to irregular,
2–10 mm diam or conluent 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
306
Fig. 103. Pseudocercospora contraria (B, holotype). A. Conidiophore
fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
on the lower side, sometimes with brownish or reddish halo.
Caespituli amphigenous, punctiform, at irst 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
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
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 ellipsoidovoid, 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,
hirtilora, oppositifolia, pentaphylla [triphylla], quartiniana,
quinquelobata, schimperiana, triida, 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)
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, ig. 13), Guo et al.
(1998: 380, ig. 311).
Description: Leaf spots indistinct or above all evident on
the lower leaf surface, angular, vein-limited, 1–5 mm diam
or conluent 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 supericial 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 lexuous,
sinuous, but mostly not distinctly geniculate, unbranched
or occasionally branched near the apex, 40–270 × 4–5
µm, width often irregular, plainly pluriseptate, olivaceousbrown, 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.
VOLUME 5 · NO. 2
Fig. 104. Pseudocercospora cylindrata (FH 2515). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
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, unconirmed. 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.
307
Braun et al.
ART I CLE
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)
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, ig. 18), Pons & Sutton
(1988: 28, ig. 11).
Fig. 105. Pseudocercospora dioscoreae (PC, holotype). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
pseudocercospora dioscoreae U. Braun, Mouch. &
McKenzie, New Zealand J. Bot. 37: 313 (1999).
(Fig. 105)
Illustration: Braun et al. (1999: 311, ig. 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,
308
Description: Leaf spots amphigenous, subcircular, elliptical
to irregular-angular, somewhat vein-limited, 2–8 mm diam,
brown, later greyish brown or grey, margin indeinite 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, olivaceousbrown. Conidiophores in dense, mostly large fascicles, to
30 or even more, arising from stromata, through stomata,
sometimes rupturing the stomata, erect, straight to slightly
lexuous-sinuous, barely geniculate, subcylindrical or
somewhat attenuated towards the tip, unbranched, 5–40
× 2–4 µm, 0–2(–4)-septate, pale olivaceous to olivaceousbrown, 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 olivaceousbrown, 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.
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 106. Pseudocercospora hiratsukana (CUP 40760). A.
Conidiophore fascicle and solitary conidiophore. B. Conidiophores.
C. Conidia. Bar = 10 µm.
[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
VOLUME 5 · NO. 2
Fig. 107. Pseudocercospora subrufa (NY 830555). A. Conidiophore
fascicle. B. Solitary conidiophores arising from supericial hyphae. C.
Conidia. Bar = 10 µm.
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)
Basionym: Ramularia subrufa Ellis & Holw., J. Mycol. 4: 2
(1888).
Synonyms: Mycovellosiella subrufa (Ellis & Holw.) U. Braun,
Mycotaxon 48: 288 (1993).
309
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Braun et al.
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, ig. 17), Braun (1993a:
242, ig. 18; 1998: 372, ig. 633).
Description: Leaf spots amphigenous, small, 1–4 mm
diam, angular-irregular, vein-limited, yellowish, ochraceous
to pale brown, inally sometimes greyish white, margin
indeinite or sometimes with darker marginal line. Caespituli
hypophyllous, greyish white to faintly pigmented, yellowish
ochraceous. Mycelium internal and external; supericial
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 supericial
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 geniculatesinuous, 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, thinwalled, 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
conirmed by molecular examinations (Crous et al. 2000,
2013).
310
Fig. 108. Pseudocercospora ubi (K(M) IMI 166111). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
pseudocercospora ubi (Racib.) Deighton, Mycol.
Pap. 140: 36 (1976).
(Fig. 108)
Basionym: Cercospora ubi Racib., Paras. Algen Pilze Javas
3: 39 (1900).
Synonym: Cercospora brasiliensis Av.-Saccá, Bol. Agric.
São Paulo 18: 580 (1917) [type: Brazil: São Paulo, on
Dioscorea sp. (not traced)].
Literature: Saccardo (1906: 1073), Chupp (1954: 198),
Batista et al. (1965: 12), Katsuki (1965: 28), Deighton (1976:
36), Pons & Sutton (1988: 24), Hsieh & Goh (1990: 104), Guo
& Hsieh (1995: 86), Guo et al. (1998: 102), Crous & Braun
(2003: 414), Kamal (2010: 229).
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Illustrations: Batista et al. (1965: 22, ig. 8), Deighton (1976:
37, ig. 19), Pons & Sutton (1988: 25, ig. 10), Hsieh & Goh
(1990: 105, ig. 79), Guo & Hsieh (1995: 87, ig. 79), Guo et
al. (1998: 103, ig. 83).
Description: Leaf spots subcircular to angular-irregular,
4–15 mm diam, yellowish to brown, margin indeinite.
Caespituli amphigenous, mainly hypophyllous, effuse,
greyish brown. Mycelium internal. Stromata lacking or only
small substomatal aggregations of swollen, pigmented
hyphal cells. Conidiophores in small, loose fascicles,
2–12, occasionally solitary, arising from internal hyphae or
stromatic hyphal aggregations, emerging through stomata,
erect or occasionally decumbent, straight, subcylindrical
or somewhat attenuated towards the tip, lexuous, slightly
geniculate-sinuous, unbranched, 20–120(–150) × 3.5–6
µm, 0–4-septate, pale to medium olivaceous-brown or
brown, thin-walled, smooth; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells,
mosty 15–30 µm long, conidiogenous loci inconspicuous
or only visible as truncate tips, unthickened, not darkened.
Conidia solitary, obclavate-cylindrical, subclavate, straight
to somewhat curved, (20–)25–70(–120) × 3–6(–7.5)
µm, (0–)1–6(–9)-septate, subhyaline to pale olivaceous
or olivaceous-brown, thin-walled, smooth, apex obtuse,
base short obconically truncate, about 1.5–3 µm wide, hila
unthickened, not darkened.
Holotype: Indonesia: Java: Buitenzorg, on Dioscorea alata
(KRA). Isotype: K(M) IMI 166111 (slide).
Host range and distribution: On Dioscorea (alata, bulbifera,
caucasica, esculenta, microbotrya [gibertii], nipponica,
pentaphylla [spinosa], polystachya [batatas], villosa [sativa]),
Dioscoreaceae, Africa (Togo), Asia (China, India, Indonesia,
Japan, Malaysia, Papua New Guinea, Philippines, Russia,
Taiwan), Oceania (Samoa, Solomon Islands, Tonga,
Vanuatu), Central and South America (Brazil, Panama,
Venezuela), West Indies (Dominican Republ.).
pseudocercospora ubicola (J.M. Yen) Deighton,
Mycol. Pap. 140: 155 (1976).
(Fig. 109)
Basionym: Cercospora ubicola J.M. Yen, Rev. Mycol. 30: 200
(1965).
Literature: Yen & Lim (1980: 189), Shaw (1984), Sutton &
Pons (1988: 22), Guo & Jiang (2000a), Crous & Braun (2003:
414).
Illustrations: Yen (1965: 201, ig. 14), Yen & Lim (1980: 262,
ig. 64), Sutton & Pons (1988: 23, ig. 9).
Description: Leaf spots amphigenous, irregular-angular,
yellowish to brown, vein-limited, margin indeinite, conluent.
Caespituli hypophyllous, indistinct. Mycelium internal and
external; supericial hyphae emerging through stomata,
branched, 1–2.5 µm wide, septate, pale olivaceous to
olivaceous-brown, thin-walled, smooth. Stromata lacking
or small stromatic hyphal aggregations, 8–15 µm diam,
VOLUME 5 · NO. 2
Fig. 109. Pseudocercospora ubicola (PC, holotype). A. Conidiophore
fascicle and conidiophores. B. Solitary conidiophores arising from
supericial hyphae. C. Conidia. Bar = 10 µm.
substomatal, somewhat pigmented. Conidiophores in small,
loose fascicles, 2–5, emerging through stomata or solitary,
arising from supericial hyphae, erect, straight to curved,
subcylindrical-conical,
somewhat
geniculate-sinuous,
unbranched, 4–30 × 2–5 µm, 0–3-septate, subhyaline to
pale olivaceous or olivaceous-brown, thin-walled, smooth;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, 5–20 µm long, conidiogenous
loci inconspicuous to subdenticulate, but unthickened and
not darkened. Conidia solitary, acicular-iliform to narrowly
obclavate-cylindrical, straight to curved, (12–)25–100 × 1–2.5
µm, (0–)2–8-septate, subhyaline, thin-walled, smooth, apex
subacute to subobtuse, base truncate to short obconically
truncate, 1–2 µm wide, hila unthickened, not darkened.
311
ART I CLE
Braun et al.
Holotype: Singapore: Botanic Garden, on Dioscorea
pyrifolia, 7 June 1964, S. H. Sun (PC). Isotype: K(M) IMI
120988 (slide).
Host range and distribution: On Dioscorea (pyrifolia,
subcalva), Dioscoreaceae, Asia (China, Papua New Guinea,
Singapore).
Zasmidium
A single species.
Zasmidium dioscorinum Archana Singh, R. Singh,
Sham. Kumar, Upadhyaya & R.F. Castañeda, Nova
Hedwigia 98: 258 (2014).
(Fig. 110)
Illustration: Singh et al. (2014: 259, ig. 1).
Description: Leaf spots amphigenous, small to large, 1.5–
15 mm diam, subcircular to angular, vein-limited, scattered,
blackish brown. Colonies hypophyllous, effuse, blackish
brown. Mycelium internal and external; supericial hyphae
branched, septate, 1.5–2.5 µm wide, thin-walled, verruculose.
Stromata supericial, prosenchymatous, 20–30 × 20–25
µm, dark brown. Conidiophores in loose fascicles, 8–19,
arising from stromata, or solitary, arising from supericial
hyphae, lateral, erect, straight to curved-sinuous, lexuous
to 0–3 times geniculate, unbranched, 6–130 × 3–4 µm,
2–7-septate, brown, paler towards the apex, wall thickened,
smooth; conidiogenous cells integrated, terminal, 8.5–28.5
µm long, sympodial, conidiogenous loci conspicuous,
thickened and darkened, 0.5–1.5 µm diam. Conidia solitary,
short obclavate-subcylindrical, ellipsoid-fusiform, straight to
slightly curved, 12–40 × 4–6 µm, 1–3-septate, light brown,
wall slightly thickened, verruculose, obtuse, rounded to
somewhat attenuated, base rounded to short obconically
truncate, 0.5–1.5 µm wide, hila somewhat thickened and
darkened.
Holotype: India: Uttar Pradesh: Siddharthnagar, Bansi, on
Dioscorea oppositifolia, Dioscoreaceae, Mar. 2008, A. Singh
(HCIO 42396). Isotype: GPU–KSR 398.
Host range and distribution. Only known from the type
collection.
Flagellariaceae
Pseudocercospora
A single species.
Pseudocercospora lagellariae Goh & W.H. Hsieh,
Trans. Mycol. Soc. Republ. China 2: 130 (1987).
(Fig. 111)
Synonym: Cercospora lagellariae Sawada, Taiwan Agric.
Rev. 38: 696 (1942), nom. inval. (ICN, Art. 39.1).
Literature: Chupp (1954: 237), Goh & Hsieh (1990: 132), Guo
312
Fig. 110. Zasmidium dioscorinum (based on Singh et al. 2014: 259,
ig. 1). A. Solitary conidiophore arising from supericial hypha. B.
Conidiophore fascicle. C. Conidia. Bars = 10 µm.
& Hsieh (1995: 121), Guo et al. (1998: 137), Crous & Braun
(2003: 186).
Illustrations: Goh & Hsieh (1990: 133, ig. 101), Guo & Hsieh
(1995: 124, ig. 109), Guo et al. (1998: 138, ig. 113).
Description: Leaf spots elliptical, 3–5 mm diam, pale to
dark brown. Caespituli amphigenous. Mycelium internal.
Stromata hemispherical, 50–70 µm wide and to 40
µm high, dark brown. Conidiophores in dense almost
sporodochial fascicles, arising from stromata, erect,
straight, subcylindrical, not or barely geniculate, apex
mostly truncate, unbranched, short, 5–20 × 4–6 µm,
0–2-septate, brown, thin-walled, smooth; conidiogenous
cells integrated, terminal or conidiophores often reduced
to conidiogenous cells, conidiogenous loci inconspicuous
or visible as truncate tip, but always unthickened and not
darkened. Conidia solitary, cylindrical, obclavate-cylindrical
or slightly clavate, almost straight to curved, 55–70 × 4–5
µm, 3–5-septate, colourless, thin-walled, smooth, apex
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 111. Pseudocercospora lagellariae (NTU-PPE, holotype). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
obtuse, rounded, base truncate to obconically truncate, hila
neither thickened nor darkened.
Holotype: taiwan: Hengchun, Pingtung Hsieh, on Flagellaria
indica, 26 Apr. 1931, K. Sawada (NTU-PPE, herb. Sawada).
Host range and distribution:
Flagellariaceae, Asia (Taiwan).
On
Flagellaria
indica,
Heliconiaceae
Cercospora
A single species.
Cercospora heliconiae Chowdhry, D. Gupta & Padhi,
Indian Phytopathol. 36: 625 (1983).
(Fig. 112)
Literature: Crous & Braun (2003: 214), Kamal (2010: 50).
Illustration: Chowdry et al. (1983: 625, ig. 2).
VOLUME 5 · NO. 2
Fig. 112. Cercospora heliconiae (based on Chowdhry et al. 1983:
625, ig. 2). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.
Description: Leaf spots amphigenous, brown, at irst
marginal, then extending to the midrib, 16–80 mm diam.
Caespituli amphigenous. Stromata globose, 30–50 µm
diam, dark brown. Conidiophores in well-developed, dense
fascicles, 11–40, arising from stromata, erect, subcylindrical,
313
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Braun et al.
straight to sinuous, 2–7 times geniculate, unbranched,
95–180 × 3–4.5 µm, 2–8-septate, deep olivaceous-brown;
conidiogenous cells integrated, terminal and intercalary,
conidiogeous loci conspicuous, thickened and darkened.
Conidia solitary, acicular, straight to curved, 35–180 × 1.5–3
µm, 8–23-septate, hyaline, thin-walled, smooth, apex acute,
base truncate, hila thickened and darkened.
Holotype: India: Odisha: Bhubaneswar, on Heliconia caribaea,
Heliconiaceae, 28 Nov. 1977, D. Gupta (HCIO 32855).
Host range and distribution: Only known from the type
collection.
Note: A true Cercospora s. str. belonging to the C. apii s. lat.
complex.
Pseudocercospora
Key to Pseudocercospora species on Heliconiaceae
1
Conidiophores (20–)30–90 × 4.5–9 µm; conidiogenous loci unthickened,
not darkened; conidia cylindrical, subcylindrical to somewhat fusiform,
50–120 × 4.5–6 µm, 0–5-septate, pale brown; on Heliconia psittacorum ............................................. p. heliconiae
Conidiophores shorter and narrower, 10–65 × 3–6(–7) µm; conidiogenous loci subconspicuous,
paracercospora-like, i.e. only ultimate rim slightly thickened and darkened, in front view
visible as minute circle; conidia obclavate-cylindrical, somewhat narrower,
30–130 × 2.5–5 µm, 1–10-septate; on Heliconia psittacorum ............................. P. ijiensis (see under Musaceae)
pseudocercospora heliconiae Meiriele Silva & O.L.
Pereira, Mycotaxon 113: 366 (2010).
(Fig. 113)
Illustration: Silva & Pereira (2010: 367–369, igs 1–6).
Description: Leaf spots amphigenous, variable, subcircular
to subangular-irregular, brownish to dingy grey or greyish
white, border darker, at irst small, 2–10 mm diam, later
spreading or conluent, covering large leaf areas. Caespituli
amphigenous, mainly hypophyllous, punctiform, dark
olivaceous-brown to blackish brown. Mycelium internal.
Stromata absent or small, 10–40 µm diam, brown to dark
brown, substomatal. Conidiophores in small to moderately
large, loose to dense fascicles, arising from internal hyphal
cells or small stromata, erect, almost straight to curved,
subcylindrical to geniculate-sinuous, unbranched, (20–)
30–90 × 4.5–9 µm, 0–4(–5)-septate, medium brown, paler
towards the tip, wall thin to somewhat thickened, to 1 µm, at
the very base sometimes to 2 µm, smooth; conidiogenous
cells integrated, terminal or conidiophores reduced to
conidiogenous cells, 15–50 µm long, conidiogenous loci
inconspicuous or visible as truncate tips, 2–3 µm wide, but
always unthickened and not darkened. Conidia solitary,
cylindrical, subcylindrical to somewhat fusiform, i.e.
gradually attenuated to base and apex, 50–120 × 4.5–6 µm,
0–5-septate, pale brown, thin-walled, smooth, apex obtuse,
base obconically truncate, 2–3.5 µm wide, hila unthickened,
not darkened.
Holotype: Brazil: Minas Gerais: Viçosa, on Heliconia
psittacorum, Heliconiaceae, 12 May 2008, O. L. Pereira (VIC
31221. Isotype: HAL 2356 F.
Host range and distribution: Only known from the type
collection.
314
Fig. 113. Pseudocercospora heliconiae (HAL 2356 F). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Note: A record of Pseudocercospora ijiensis on Heliconia
psittacorum was published from Brazil (Gasparotto et al.
2005).
Hydrocharitaceae
Cercospora
A single species.
Cercospora limnobii Conway, Trans. Brit. Mycol. Soc.
71: 523 “1978” (1979).
(Similar to Fig. 1)
Literature: Crous & Braun (2003: 252).
Illustration: Conway (1979b: 522, igs 1–4).
Description: Leaf spots irregularly shaped, 10–15 mm
diam, later coalescing along the margin and tip, medium
to dark brown. Stromata lacking or small, 10–30 µm diam,
substomatal, mostly only a few swollen hyphal cells, brown,
wall thickened. Mycelium internal; hyphae branched, straight
to sinuous-torulose, pale olivaceous to medium brown, 2–6
µm wide. Conidiophores solitary or in small fascicles, 2–6,
arising from internal hyphae or stromata, through stomata,
erect, straight, subcylindrical to geniculate-sinuous in the
upper fertile part, 75–130(–260) × 4–8 µm, septate, brown
throughout or with paler tips; conidiogenous cells integrated,
terminal and intercalary, about 10–35 µm long, conidiogenous
loci conspicuous, thickened and darkened, 2–4 µm diam.
Conidia solitary, acicular, straight or almost so, 75–150(–
250) × 3–4.5 µm, pluriseptate, hyaline, thin-walled, smooth,
apex acute, base truncate, 2–3 µm wide, hila thickened and
darkened.
Holotype: uSA: Florida: Deep Creek, Rodman Reservoir,
Orange Springs, on Limnobium spongia, Hydrocharitaceae,
12 Sep. 1975, K. E. Conway & D. Bowman (BPI 71882).
Host range and distribution: Only known from the type
collection.
Note: This species belongs to the C. apii complex.
Hypoxidaceae
Fig. 114. Pseudocercospora curculiginis (HMAS 62713). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
Pseudocercospora
A single species.
pseudocercospora curculiginis Y.L. Guo & X.J. Liu,
Mycosystema 5: 101 (1992).
(Fig. 114)
Literature: Guo & Hsieh (1995: 10–11), Guo et al. (1998: 22).
Illustrations: Guo & Liu (1992: 102, ig. 2), Guo & Hsieh
(1995: 12, ig. Fig.11), Guo et al. (1998: 22, ig. 11).
VOLUME 5 · NO. 2
Description: Leaf spots amphigenous, elliptical to oblongelliptical, 0.5–10 × 0.3–10 mm, often conluent, pale grey
to pale brown, surrounded by a dark greyish marginal line
and yellowish brown to pale greyish brown halo on the
upper side and grey to pale greyish brown below. Caespituli
amphigenous. Mycelium internal. Stromata none or small,
10–30 µm diam, globose, greyish brown. Conidiophores
fasciculate, 2–15, arising from internal hyphal cells or stromata,
through stomata, erect, straight to curved, subcylindrical to
somewhat geniculate-sinuous, mostly unbranched, 6.5–
96.5 × 3–4.5 µm, 1–7-septate, uniformly pale to medium
olivaceous-brown, thin-walled, smooth; conidiogenous cells
315
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Braun et al.
integrated, terminal and intercalary, conidiogenous loci
inconspicuous, unthickened, not darkened. Conidia solitary,
acicular to somewhat obclavate-subcylindrical, 40–120 ×
3–4 µm, 4–12-septate, olivaceous, thin-walled, smooth, apex
obtuse to pointed, base truncate or occasionally somewhat
obconically truncate, hila unthickened, not darkened.
Holotype: China: Guangdong: Dinghushan, on Molinearia
capitulata [Curculigo capitulata], Hypoxidaceae, 21 Oct.
1981, Y. L. Guo & X. J. Liu 17 (HMAS 62713)
Host range and distribution: Only known from the type
collection.
Iridaceae
Cercospora
Key to Cercospora species on Iridaceae
1
Conidiophores short, 15–50 × 3–5.5 µm; conidia acicular, 25–70 × 2–4 µm; on Iris spp. ................................... C. iridis
Conidiophores very long, about 60–340 × 3–5 µm; conidia obclavate-cylindrical,
57–133 × 4–7 µm; on Neomarica caerulea ........................................................................................ C. neomaricae
Cercospora species on Iridaceae
Cercospora iridis Chupp, Monograph of Cercospora:
260 (1954).
Literature: Crous & Braun (2003: 229).
Description: Leaf spots amphigenous, elliptical to oblong,
5–20 mm in length, straw-coloured to brownish, darkened
when fruiting is abundant. Caespituli amphigenous, dark.
Mycelium internal. Stromata globose to lattened, 20–40
µm diam, dark brown. Conidiophores fasciculate, 2–20,
divergent, straight to slightly curved, 0–2 times geniculate,
unbranched, 15–50 × 3–5.5 µm, sparingly septate, pale to
medium brown, paler towards the tip; conidiogenous loci
conspicuous, thickened and darkened. Conidia solitary,
acicular, straight to somewhat curved, 25–70 × 2–4 µm,
indistinctly pluriseptate, thin-walled, smooth, tip subacute or
subobtuse, base truncate, hila thickened and darkened.
Holotype: Mexico: Matamoros, on Iris sp., 12 May 1944, C. J.
Hansen 57462 (not traced).
Host range and distribution: On Iris spp., Iridaceae, Asia
(Nepal), North America (Mexico), South America (Brazil).
Notes: A true Cercospora s. str. close to C. apii s. lat. but
with short conidiophores and obclavate-acicular conidia.
According to Chupp (1954), type material was said to be
deposited at “U.S.D.A. Herbarium” (now BPI), but this
collection is not preserved in this herbarium.
Cercospora neomaricae Macedo & R.W. Barreto, sp.
nov.
MycoBank MB809023
(Fig. 115)
Synonym: Cercospora neomaricae Macedo & R.W. Barreto,
Australas. Pl. Pathol. 37: 581 (2008); nom. ival. (ICN, Art.
40.6).
Illustration: Macedo & Barreto (2008: 582, Fig. 2).
316
Fig. 115. Cercospora neomaricae (VIC 305338). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
IMA FUNGUS
Cercosporoid fungi 2
obconically truncate, hila thickened and darkened. In vitro:
Colonies slow-growing (3–6 cm in 15 d), white to pale
pinkish cottony aerial mycelium which may raise centrally
with irregular greyish sectioning (in MEA), rays of mycelial
threads immersed in the medium formed at the periphery (in
MEA), colony in reverse white to pink, greenish brown, dark
grey or almost black, not sporulating.
ART I CLE
Description: Leaf spots amphigenous, subcircular,
about 3–10 mm diam, brown with yellow halo, later
centre greyish with dark brown margin (eye-spots) and
yellow halo. Caespituli amphigenous. Mycelium internal.
Stromata lacking to well-developed. Conidiophores in
loose fascicles, arising from internal hyphae or stromata,
erect, straight to slightly curved, geniculate, unbranched,
about 60–340 × 3–5 µm, 2–6-septate, dark brown, thinwalled, smooth; conidiogenous cells integrated, terminal
or intercalary, sympodial, with conspicuous conidiogenous
loci, thickened and darkened. Conidia solitary, obclavatesubcylindrical, straight to somewhat curved, 57–133 ×
4–7 µm, 6–16-septate, hyaline, thin-walled, smooth, apex
obtuse to somewhat pointed, base truncate to usually
Holotype: Brazil: Rio de Janeiro, municipality Petrópolis,
Haipava, on Neomarica caerulea, Aug. 2007, D. M. Macedo
& R. W. Barreto (VIC 305338).
Host range and distribution: On Neomarica caerulea,
Iridaceae, South America (Brazil).
Pseudocercospora (including Stigmina)
Key to Pseudocercospora species on Iridaceae
1
Conidiogenous cells sympodially proliferating, without annellations;
conidia acicular-iliform, 70–135 × 2–3 µm, 5–14-euseptate, smooth;
on Libertia, New Zealand .......................................................................................................................... p. libertiae
Conidiophores percurrently proliferating, with annellations; conidia subcylindrical,
18–25 × 7–8 µm, 1–3-distoseptate, verruculose; on Dierama, South Africa .............................. Stigmina dieramae
Pseudocercospora species on Iridaceae
pseudocercospora libertiae U. Braun & C.F. Hill, in
Braun et al., Australas. Pl. Pathol. 32: 89 (2003).
(Fig. 116)
Holotype: New Zealand: Auckland, Auckland University
Campus, on Libertia ixioides, Iridaceae, 8 Apr. 2001, C. F.
Hill 398 (HAL 1728 F).
Illustration: Braun et al. (2003b: 90, ig. 4).
Description: Leaf spots amphigenous, oblong, spread along
the leaf margin, inally very long, expanded, covering large
parts of the leaves, pale to dark brown, reddish brown, later
pale straw-coloured to dull greyish white, with brown to
reddish brown border. Caespituli amphigenous, punctiform,
dark brown, loose to dense, pale dingy olivaceous to greyish
white by abundant conidial formation. Mycelium internal.
Stromata lacking or small, substomatal, 10–30 µm diam,
olivaceous-brown. Conidiophores in small to moderately
large fascicles, loose to dense, arising from internal hyphae
or stromata, through stomata, erect, straight, subcylindricalconical to geniculate-sinuous, unbranched, 5–25 × 2–4 µm,
occasionally to 6 µm wide at the very base, conidiophores
often appearing to be longer by persistent conidia,
0–1-septate, subhyaline to pale olivaceous or olivaceousbrown, thin-walled, smooth; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous
cells, 5–20 µm long, sympodial, conidiogenous loci
inconspicuous. Conidia solitary, acicular-iliform, 70–135
× 2–3 µm, obscurely 5–14-septate, often with minute oil
droplets when fresh, subhyaline to pale olivaceous, thinwalled, smooth, apex subacute, base truncate or gradually
attenuated towards the base (i.e. only slightly obconically
truncate), (1.5–)2(–2.5) µm wide, hila unthickened, not
darkened.
VOLUME 5 · NO. 2
Fig. 116. Pseudocercospora libertiae (HAL 1728 F). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
317
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Braun et al.
Host range and distribution: Only known from the type
collection.
Stigmina dieramae Crous & B. Sutton, S. Afr. J. Bot.
63: 284 (1997).
(Fig. 117)
Illustration: Crous & Sutton (1997: 284, ig. 7).
Description: Leaf spots amphigenous, black, diffuse, 1–12 × 1
mm. Caespituli amphigenous, scattered, punctiform, blackish.
Mycelium internal and external, supericial; hyphae light brown,
branched, septate, 4–6 µm wide, smooth to verruculose,
forming small substomatal stromatic aggregations.
Conidiophores fasciculate, 2–8, arising from stromatic hyphal
aggregations, forming small conidiomata, 13–30 µm diam,
emerging through stomata, erect, subcylindrical, straight or
once geniculate, unbranched or only branched at the base,
8–16 × 6–8 µm, aseptate or 1-septate, brown, thin-walled,
verruculose; conidiogenous cells integrated, terminal or
conidiophores aseptate, i.e. reduced to conidiogenous cells,
tapering to a rounded end, brown, verruculose, percurrently
proliferating, with 1–3 irregular annellations, conidiogenous
locus neither thickened nor darkened. Conidia solitary,
holoblastic, subcylindrical, straight to somewhat curved,
18–25 × 7–8 µm, 1–3-distoseptate, brown, verruculose,
with longitudinal striations, apex obtuse, broadly rounded,
base truncate to subtruncate, with marginal frill, but neither
thickened nor darkened.
Holotype: lesotho: Thaba Putsoa, Maseru district, on
Dierama sp., Iridaceae, 23 Feb. 1967, A. J. Guillarmod
(PREM 44333).
Host range and distribution: Only known from the type
collection.
Notes: Due to didymo- to phragmosporous, distoseptate
conidia, this species is morphologically close to the type
species of Stigmina, which proved to be a species of
Pseudocercospora based on its phylogenetic position.
Therefore, Stigmina was reduced to synonymy with
Pseudocercospora. On the other hand, S. diramae is
characterised by having verruculose conidiophores and
conidia which is reminiscent of Pseudoasperisporium.
However, the latter genus differs by its euseptate conidia.
Fig. 117. Stigmina dieramae (PREM 44333). A. Conidiophore
fascicles. B. Conidia. Bar = 10 µm. P.W. Crous del.
Stigmina dieramae has not yet been cultured and its
phylogenetic position is still unknown. Therefore, we hesitate
to reallocate this species to Pseudocercospora and retain it
tentatively in Stigmina awaiting conirmation by cultures and
phylogenetic data.
Doubtful, excluded and insuficiently known
species
Cercospora iridicola Tracy & Earle, nom. nud.
uSA: Mississippi: Starkville, on Iris pabularia, Iridaceae, Jan
1894, S. M. Tracy (not traced).
Notes: A heterosporium-like fungus (Cladosporium),
according to Chupp (1954: 261), probably Cladosporium
iridis.
Juncaceae
Cercospora
Key to Cercospora species on Juncaceae
1
318
Conidiophores (5–)10–60(–80) µm long, pale to dark brown or olivaceous-brown;
conidia obclavate-cylindrical, 2.5–5(–6) µm wide, hyaline, subhyaline to pale olivaceous;
on numerous Juncus spp. ......................................................................................................................... C. juncina
Conidiophores 5–30 µm long, paler, subhyaline to pale olivaceous to medium brown;
conidia acicular, rarely obclavate, narrower, 1.5–3 µm, hyaline;
on Juncus marginatus, USA ......................................................................................................... C. junci-marginati
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
subcylindrical to geniculate-sinuous, unbranched, 5–30 × 2–5
µm, aseptate, subhyaline, pale olivaceous to medium brown,
thin-walled, smooth; conidiophores reduced to conidiogenous
cells, conidiogenous loci minute, 1–2 µm diam, somewhat
thickened and darkened. Conidia solitary, acicular, rarely
narrowly obclavate, 25–65 × 1.5–3 µm, 1–5-septate, hyaline,
thin-walled, smooth, apex pointed, base truncate, 1–2.5 µm
wide, hila somewhat thickened and darkened.
Holotype: uSA: New York: Selkirk, on Juncus marginatus,
Peck (NYS).
Host range and distribution: On Juncus marginatus,
Juncaceae, North America (USA, NY).
Cercospora juncina Sacc., Ann. Mycol. 11: 552
(1913).
(Fig. 119)
Synonyms: Cercosporina juncicola Hori & Kasai, Jap. J.
Bot. 2: 105 (1923) [type: Japan: Okayama Pref.: Kanaeura, on Juncus effusus, 28 June 1921, M. Kasai (not
preserved)].
Cercospora junci Davis, Trans. Wisconsin Acad. Sci. 24:
300 (1929) [holotype: uSA: Wisconsin: Brill, on Juncus
brevicaudatus, 23 Jul. 1928, J. J. Davis (WIS)].
Cercospora juncicola (Hori & Kasai) Vassiljevsky, in
Vassiljevsky & Karakulin, Fungi Imperfectii Parasitici
(Hyphomycetes) 1: 275 (1937).
Cercospora juncicola (Hori & Kasai) Chupp, Monograph of
Cercospora: 263 (1954), comb. superl.
Cercospora junci-iliformis Mel'nik, Novosti Sist. Nizsh. Rast.
1966: 214 (1966) [holotype: russia: St. Petersburg area,
Tosnensky Rayon, Lisino-Korpus, 15 Jul. 1963, V. A.
Mel’nik (LE 40408)].
Literature: Saccardo (1931: 877), Chupp (1954: 263), Braun
& Mel'nik (1997: 65), Crous & Braun (2003: 233).
Fig. 118. Cercospora junci-marginati (NYS, holotype). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
Illustration: Braun & Mel'nik (1997: ig. 33).
Exsiccatae: Barthol., Fungi Columb. 4902.
Cercospora species on Juncaceae
Cercospora junci-marginati U. Braun, Mycotaxon 48:
277 (1993).
(Fig. 118)
Basionym: Ramularia junci Peck, Rep. (Annual) New York
State Mus. Nat. Hist. 44: 26 (1891), non Cercospora junci
Davis, 1929.
Literature: Saccardo (1892: 563), Braun (1998: 300), Crous
& Braun (2003: 233).
Illustration: Braun (1993b: 280, ig. 3).
Description: Leaf spots amphigenous, oblong, 2–10 × 1–2
mm, brown, margin indeinite. Caespituli amphigenous.
Mycelium internal, forming substomatal stromata, 10–35 µm
diam, yellowish brown. Conidiophores in dense fascicles,
arising from stromata, emerging through stomata, straight,
VOLUME 5 · NO. 2
Description: Lesions at irst small, subcircular, elliptical-oval,
oblong, 1–10 × 1–3 mm, sometimes diffuse, brown with diffuse
yellowish halo, later larger segments of the leaf sheath or
stem turning dark brown. Caespituli punctiform, minute, dark
brown to blackish. Mycelium internal. Stromata substomatal,
10–40 µm diam, rarely larger, to 75 µm, brown to dark brown,
composed of swollen hyphal cells, 2–7 µm diam, rounded to
angular in outline. Conidiophores in small to moderately large
fascicles, about 3–20, loose to usually dense or very dense,
arising from stromata, through stomata, erect, straight,
subcylindrical to distinctly geniculate-sinuous, unbranched,
(5–)10–60(–80) × 2–6 µm, rarely swollen at the base, to 8 µm
wide, 0–2-septate, pale to dark brown or olivaceous-brown,
wall thin or slightly thickened; conidiogenous cells integrated,
terminal or conidiophores often reduced to conidiogenous
cells, sympodial, 5–30 µm long, conidiogenous loci
conspicuous, about 1.5–2 µm diam, thickened and darkened.
Conidia solitary, obclavate-cylindrical, straight to curved, 25–
319
Braun et al.
ART I CLE
Notes: Type material of Cercosporina juncicola is not preserved,
and even Katsuki (1965) failed to trace type material or any
other collections of this species. However, based on the data
given in the original description, this species seems to be a
synonym of C. juncina, above all due to short conidiophores
and obclavate-cylindrical, hyaline to olivaceous conidia. The
fungus described and illustrated by Ellis (1976: 262–263,
ig. 199C) does not agree with the original description of C.
juncicola (conidiophores 50–90 × 4–6 µm, pale to medium
brown; conidia solitary, obclavate-subcylindrical to subacicular,
straight to curved, 40–140 × 3–5 µm, 2–7-septate, pale strawcoloured, often minutely verruculose). The identity of the
described material, probably from UK, is unclear. The source
of Chupp’s (1954) description is also unclear and not in
accordance with the original data of C. juncicola.
Liliaceae (s. str.)
Passalora
A single species.
passalora streptopi (Dearn. & Barthol.) U. Braun,
comb. nov.
MycoBank MB809041
(Fig. 120)
Basionym: Cercospora streptopi Dearn. & Barthol., Mycologia
9: 363 (1917).
Literature: Saccardo (1931: 883), Chupp (1954: 354).
Exsiccatae: Barthol., Fungi Columb. 5004.
Fig. 119. Cercospora juncina (BPI 437515). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
75 × 2.5–5(–6) µm, 2–6-septate, hyaline, subhyaline to pale
olivaceous, thin-walled, smooth, apex obtuse to pointed, base
short obconically truncate, 1.5–2.5 µm wide, hila thickened
and darkened.
Holotype: Canada: Ontario, London, on Juncus canadensis,
12 Aug. 1910, J. Dearness, 3502 [Barthol., Fungi Columb.
4902] (PAD). Isotypes: Barthol., Fungi Columb. 4902, e.g.
BPI 437515, 437516, DAOM, K(M) IMI 8038.
Host range and distribution: On Juncus (balticus,
brachycephalus, brevicaudatus, canadensis, confusus,
dudleyi, effusus, ensifolius, iliformis, greenei, pygmaeus,
Juncaceae, Africa (Morocco), Asia (Japan), Europe
(Romania, Russia), North America (Canada; USA, Utah,
Washington, Wisconsin).
320
Description: Leaf spots amphigenous, subcircular to angularirregular, 1–6 mm diam or conluent and larger, yellowish,
olivaceous to brownish, inally sometimes greyish white, margin
indeinite or darker, brown, sometimes with diffuse yellowish,
straw-coloured to pale brownish halo. Caespituli amphigenous,
punctiform to effuse, brown, scattered to aggregated, loose
to dense. Mycelium internal. Stromata almost lacking, only
with small substomatal aggregations of swollen hyphal cells,
globose to subglobose, brown, substomatal, 10–50 µm
diam. Conidiophores in small to moderately large, loose to
dense fascicles, arising from internal hyphae or stromata,
emerging through stomata, erect, straight and subcylindrical to
subclavate, i.e. width somewhat increasing from base to top,
barely or only slightly geniculate-sinuous, unbranched, 10–70
× 2.5–5 µm, 0–4-septate, pale olivaceous to brown below, paler
towards the apex, tips often subhyaline, darker brown in mass,
thin-walled, smooth; conidiogenous cells integrated, terminal,
occasionally conidiophores reduced to conidiogenous cells,
10–35 µm long, often with a single, sometimes with several
conspicuous conidiogenous loci, 1–1.5 µm diam, somewhat
thickened and darkened-refractive. Conidia solitary as well
as catenate, usually in simple chains, broadly ellipsoid-ovoid,
cylindrical, subclavate to somewhat obclavate, straight to
slightly curved, 18–70 × (3–)4–7(–9) µm, (0–)1–7-septate,
sometimes constricted at septa, subhyaline to yellowish, pale
olivaceous or olivaceous-brownish to medium brown, thinwalled, smooth, older conidia occasionally rough-walled, apex
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 120. Passalora streptopi (BPI 441721). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
Fig. 121. Pseudocercospora dispori (WSP 35442). A. Conidiophore
fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
obtuse, rounded to obconically truncate in catenate conidia,
base subtruncate to obconically truncate, 1–2 µm wide, hila
slighty thickened and darkened-refractive.
Notes: This species is a typical phaeoramularioid Passalora
species characterised by having pigmented conidiophores
and conidia, conspicuous (thickened and darkened)
conidiogenous loci and conidial hila and catenate conidia.
Lectotype (designated here, MycoBank MBT178157): uSA:
Washington: Jefferson, Duckabush River, Olympic National
Park, on Streptopus amplexifolius, Aug. 1912, E. Bartholomew
[Fungi Columb. 5004] (BPI 441721). Isolectotypes: Barthol.,
Fungi Columb. 5004, BPI 441722, 845248, DAOM, MICH
5601.
Host range and distribution: On Prosartes (hookeri,
trachycarpum), Streptopus amplexifolius, Liliaceae s. str.,
North America (Canada; USA, Montana, Oregon, Tennessee,
Washington).
VOLUME 5 · NO. 2
Pseudocercospora
A single species.
pseudocercospora dispori (U. Braun) U. Braun,
comb. nov.
MycoBank MB809012
(Fig. 121)
Basionym: Stigmina dispori U. Braun, Mycotaxon 51: 66
(1994).
321
Braun et al.
ART I CLE
Illustration: Braun (1994: 64, ig. 33).
Description: Leaf spots amphigenous, angular-irregular, 1–15
mm diam or conluent and larger, often large leaf segments
or entire leaves discoloured, yellowish ochraceous, later
brownish, margin indeinite. Caespituli amphigenous,
mostly hypophyllous, punctiform, yellowish, brownish,
inally almost black. Mycelium internal, pigmented, forming
well-developed, substomatal stromata. Conidiophores
fasciculate, sporodochial, arising from stromata, emerging
through stomata, fascicles rich and dense, erect, straight to
somewhat lexuous, unbranched or occasionally branched
near the apex, subcylindrical to subclavate, 10–40 × 2–6
µm, aseptate or septate, at irst subhyaline, later yellowish to
brown, smooth, thin-walled; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells,
monoblastic, determinate or percurrently proliferating, with
delicate annellations, conidiogenous loci broad and truncate,
unthickened, not darkened. Conidia solitary, subcylindrical,
subclavate or obclavate, 25–60 × 4–9 µm, (0–)2–5-septate,
occasionally constricted at the septa, subhyaline to faintly
yellowish green or olivaceous, thin-walled, smooth, apex
obtuse, base truncate, hila neither thickened nor darkened.
Holotype: uSA: Washington: Mason County, Lilliwaup, on
Prosartes hookeri [Disporum hookeri], Liliaceae, 27 Jul.
1937, L. K. Jones (WSP 35442).
Host range and distribution: Only known from the type
collection.
Notes: This species is a cercostigmina-like hyphomycete.
However, Cercostigmina has been reduced to synonym with
Pseudocercospora.
Zasmidium
A single species.
Zasmidium liriopes (F.L. Tai) U. Braun, Y.L. Guo &
H.D. Shin, comb. nov.
MycoBank MB809013
(Fig. 122)
Basionym: Cercospora liriopes F.L. Tai, Bull. Chinese Bot.
Soc. 2: 55 (1936).
Synonyms: Stenellopsis liriopes (F.L. Tai) H.D. Shin & U.
Braun, Mycotaxon 74: 116 (2000).
Passalora liriopes (F.L. Tai) Y.L. Guo, Mycosystema 20: 303
(2001).
Literature: Chupp (1954: 348), Shin & Kim (2001: 276–277),
Guo et al. (2003: 103).
Illustration: Shin & Braun (2000: 117, ig. 5), Shin & Kim
(2001: 277, ig. 127), Guo et al. (2003: 103, ig. 65).
Exsiccatae: Triebel, Microfungi Exs. 637.
Description: Leaf spots amphigenous, scattered, circular,
1–5 mm diam, centre tan to dingy grey with reddish brown,
322
Fig. 122. Zasmidium liriopes (HAL, Triebel, Microfungi Exs. 637). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
slightly raised margin and occasionally yellowish brown halo
on the upper leaf surface, sometimes with dots in the centre
of spots on the upper side when viewed with a hand lense.
Caespituli amphigenous, punctiform, blackish. Mycelium
internal; hyphae branched, septate, 2–4 µm wide, hyaline.
Stromata well-developed, globular, substomatal, 25–70 µm
diam, brown to blackish brown. Conidiophores in loose to
dense fascicles, (5–)10–40, arising from stromata, through
stomata, erect, almost straight to curved, sometimes sinuous,
0–2 times geniculate, but mostly not distinctly geniculate,
unbranched, 5–55 × 2–5.5 µm, 0–3-septate, olivaceous-brown
or pale brown throughout, thin-walled, smooth or almost so;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, 5–25 µm long, conidiogenous
IMA FUNGUS
Cercosporoid fungi 2
Holotype: China: Jiangsu: Wuxi, on Liriope spicata, 4 Oct.
1930, C. T. Wei 1061 (HMAS 06991). Paratype: China:
Jiangsu: Wuxi, on “Ophiopogon” sp. [Liriope sp.?], 19 Sep.
1934, C. T. Wei 1030 (HMAS 06990).
Cylindrosporium inconspicuum var. candidum Sacc. &
Fautrey, in Saccardo, Syll. Fung. 16: 1010 (1902).
Cercosporella inconspicua (G. Winter) Höhn., Ann. Mycol. 1:
413 (1903) and Hedwigia 42: (178) (1903).
Cercosporella lilii Dearn., Mycologia 21: 327 (1929) [holotype:
uSA: N.Y.: Washington Co., north of Devines woods,
Vaughns, north of Hudson Falls, on Lilium canadense, 1
Jul. 1919, Burnham (DAOM 5975)].
Pseudocercosporella hungarica (Bäumler) Sivan., Bitunicate
Ascomycetes and their Anamorphs (Vaduz): 202 (1984).
Pseudocercosporella inconspicua (G. Winter) U. Braun,
Nova Hedwigia 47: 343 (1988).
ART I CLE
loci conspicuous, 1–2.5 µm diam, truncate, almost unthickened
to somewhat thickened and somewhat darkened-refractive.
Conidia solitary, subcylindrical-obclavate, straight to somewhat
curved, (10–)20–105 × 2.5–4.5 µm, mostly (0–)2–7-septate,
subhyaline to pale olivaceous or olivaceous-brown, thin-walled,
inely but distinctly verruculose, apex obtuse, base obconically
truncate or subtruncate, 1–2 µm wide, hila unthickened or only
slightly so, not or only slightly darkened-refractive.
Marantaceae
Host range and distribution: On Liriope (muscari [platyphylla],
spicata, Liriope sp.), Liliaceae, Asia (China, Japan, Korea),
North America (USA, Florida).
Cercospora
A single species.
Notes: Cercospora liriopes is an unusual species. Due to
distinct conidiogenous loci and verruculose conidia, Shin
& Braun (2000) assigned this species to Stenellopsis. This
reallocation was based on material collected from Korea, and
type material had not been examined. Guo (2001b) introduced
the combination Passalora liriopes, and Guo et al. (2003) added
a Chinese description and illustration. Details of the structure
of the conidial surface were not provided. Korean material
agrees very well with the Chinese type material of C. liriopes.
Y.L. Guo recently re-examined this material and found that the
conidia are slightly verruculose. Thus, there is no doubt that the
Korean collections pertain to C. liriopes. However, the generic
afinity of this species is not quite clear and confused. Narrow
long, pluriseptate, verruculose conidia are not in favour of
Passalora. They are rather reminiscent of former Stenellopsis
species, which are stenella- and zasmidium-like, but without
formation of verruculose supericial hyphae. However, it has
been proven, also based on molecular methods, that such
species rather belong to Zasmidium (see Braun et al. 2013).
Furthermore, C. liriopes was included in molecular sequence
analyses and proved to cluster together with Zasmidium citrigrisea and Z. musicola (Arzanlou et al. 2008) in a groups with
100 % bootstrap support, which fully substantiates the need to
reallocate C. liriopes to Zasmidium.
Doubtful, excluded and insuficiently known
species
Cercospora liliicola Richon, Cat. Champ. Marne: No.
2032 (1889) [type: France: Marne, St. Amand, on
Lilium candidum (not preserved)].
Synonyms: Cylindrosporium inconspicuum G. Winter, in
Rabenh., Fungi Eur. Exs., Cent. 12 (resp. Cent. 32),
no. 3178, Dresden 1884 and Jahresber. Naturf. Ges.
Graubündens, N.S., 34: 69 (1890) [lectotype (designated
by Braun 1995a): Switzerland: Zürich, on Lilium
martagon, Jul. 1883, G. Winter, Rabenh., Fungi Eur. Exs.
3178 (HAL)].
Cercosporella hungarica Bäumler, Verh. Zool.-Bot. Ges.
Wien 38: 707 (1888) [lectotype (designated by Braun
1995a): Slovakia: Banská Štiavnica, Prenčov, on Lilium
martagon, Kmet (HBG)].
VOLUME 5 · NO. 2
Cercospora calatheae Viégas & Chupp, Bol. Soc.
Brasil. Agron. 8: 13 (1945).
(Fig. 123)
Literature: Chupp (1954: 378), Crous & Braun (2003: 94).
Description: Leaf spots amphigenous, circular or subcircular,
0.5–3 mm diam, pale brown to dingy grey, margin dark
brown. Caespituli amphigenous, mainly hypophyllous, ine,
not very conspicuous. Mycelium internal. Stromata lacking
or only with a few substomatal swollen, brown hyphal cells.
Conidiophores solitary or in small to moderately large
fascicles, 2–15, arising from internal hyphae or hyphal
aggregations, through stomata, erect, straight, subcylindrical
to slightly geniculate-sinuous near the apex, unbranched,
40–250 × 4–8 µm, pluriseptate throughout, pale to medium
olivaceous-brown, wall thin to slightly thickened, smooth;
conidiogenous cells integrated, terminal, about 10–25 µm
long, with a single or several conspicuous conidiogenous
loci, 2–3 µm diam, thickened and darkened. Conidia solitary,
broadly acicular to subcylindrical, 30–200 × 4–6.5 µm, 3- to
pluriseptate, hyaline thin-walled, smooth, apex obtuse or
subobtuse, base truncate or subtruncate, hila 2.5–3 µm wide,
thickened and darkened.
Lectotype (designated here, MycoBank MBT178158):
Brazil: São Paulo: Cunha, on Calathea sp., 13 Apr. 1939, A.
P. Viégas & J. Kiehl 3029 (CUP 39264). Isolectotype: IACM.
Host range and distribution: On Goeppertia (allouia [Calathea
allouia], macrosepala [Calathea macrosepala], Goeppertia
[Calathea] sp.), Marantaceae, North America (USA, Florida),
Central and South America (Brazil, Dominican Republ.,
Panama).
Notes: A true Cercospora s. str. distinct from C.apii s. lat. by
wider conidia.
Pseudocercospora
A single species.
323
ART I CLE
Braun et al.
Fig. 124. Pseudocercospora pruinosivora (based on Rao & Yadav
1991: 188, ig. 1). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.
Fig. 123. Cercospora calatheae (CUP 39264). A. Conidiophores. B.
Conidia. Bar = 10 µm.
pseudocercospora pruinosivora (V.G. Rao & B.R.D.
Yadav) Kamal, Cercosporoid Fungi of India: 211
(2010); as “priunosivora”.
(Fig. 124)
Basionym: Cercospora pruinosivora V.G. Rao & B.R.D.
Yadav, Cryptog. Bot. 2: 188 (1991); as “priunosivora”.
Stromata well-developed, immersed to erumpent, olivaceous
to dark brown. Conidiophores numerous in dense fascicles,
arising from stromata, erect, straight to curved-sinuous,
short, to about 30 µm, apically geniculate, septate, brown;
conidiogenous loci inconspicuous. Conidia solitary, obclavatesubacicular, 56–142 × 5–6.5 µm, 3–6-septate, hyaline or
subhyaline, thin-walled, smooth, apex acute to subobtuse, base
short obconically truncate, hila unthickened, not darkened.
Holotype: India: Maharashtra: Poona, on Pleiostachya
pruinosa, Marantaceae, Dec. 1977, B. R. D. Yadav (AMH
4129).
Host range and distribution: Only known from the type
collection.
Literature: Braun & Crous (2003: 337).
Illustration: Rao & Yadav (1991: 188, ig. 1).
Description: Leaf spots amphigenous, marginal, irregularly
shaped, pale buff with cinnamon-coloured margin. Caespituli
amphigenous, mainly hypophyllous. Mycelium internal.
324
Notes: This species is insuficiently known, details of stromata
and conidiophores were not described, and the type material
was not available. Kamal’s (2010) reallocation of this species
suggests that the conidiogenous loci and conidial hila are
neither thickened nor darkened, which is supported by the
original illustration.
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Description: Leaf spots amphigenous, oblong, 1–5 × 1–2 mm
or conluent and larger, vein-limited, medium to medium dark
brown, margin indeinite. Caespituli mainly hypophyllous,
punctiform, scattered to aggregated, dark brown. Mycelium
internal. Stromata lacking or almost so. Conidiophores solitary
or in small fascicles, 2–15, arising from internal hyphae,
emerging through stomata, erect, straight to somewhat
curved, subcylindrical or somewhat attenuated towards the
tip, neither geniculate nor sinuous, unbranched, about 30–60
× 6–12 µm, sparingly septate, brown, paler towards the apex,
wall somewhat thickened, but thin near the apex, verruculose;
conidiogenous cells integrated, terminal, about 10–30 µm
long, conidiogenous loci conspicuous, cercosporoid, i.e.
planate, somewhat thickened and darkened, 3–4 µm diam.
Conidia solitary, cylindrical or subcylindrical, 35–100 × 6–9
µm, 1–5-septate, brown, wall slightly thickened, verruculose,
apex obtuse, broadly rounded, base subtruncate, truncate
to short obconically truncate, 3–4.5 µm wide, hila somewhat
thickened and darkened.
Lectotype (designated here, MycoBank MBT178159): uSA:
Louisiana: St. Martinsville, on Thalia dealbata, 6 Oct. 1889, A. B.
Langlois 2130 (NY 945725). Isolectotype: BPI 441902. Syntype:
NY 945726 (from 1 Oct. 1889). Topotypes (from Nov. 1889):
CUP 41402, NY 838631–838635; Ellis & Everh., N. Amer. Fungi
2476, e.g. BPI 441901, 441903, CUP, OSC 53168.
Host range and distribution: On Thalia dealbata, Marantaceae,
North America (USA, Louisiana).
Notes: Chupp (1954) exluded this species from Cercospora
and suggested an afinity to Coryneum and Heterosporium.
This species resembles, indeed, heterosporium-like
Cladosporium species, but the conidiogenous loci are
cercosporoid, i.e. planate, thickened and darkened and is
morphologically closer to Stenellopsis and Verrucisporota,
two genera which are now tentatively considered to be
synonyms of Zasmidium.
Melanthiaceae
Cercospora
Fig. 125. Zasmidium thaliae (BPI 441902). A. Conidiophore fascicle
and conidiophore. B. Conidia. Bar = 10 µm.
A single species.
Cercospora paridis Erikss., Hedwigia 22: 158 (1883).
(Fig. 126)
Zasmidium
Basionym: Cercospora thaliae Ellis & Langl., J. Mycol. 6: 36
(1890).
Synonyms: Cercosporidium paridis (Erikss.) X.J. Liu & Y.L.
Guo, Acta Mycol. Sin. 1: 99 (1982).
Passalora paridis (Erikss.) Poonam Srivast., J. Living World
1: 117 (1994), comb. inval. (ICN, Art. 41.5).
Passalora paridis (Erikss.) Y.L. Guo, Mycosystema 20: 157
(2001).
Cercospora majanthemi var. paridis Bäumler, Verh. Zool.Bot. Ges. Wien. 38: 717 (1888) [type: Slovakia:
Banská Štiavnica (Schemnitz), on Paris quadrifolia (not
preserved)].
Literature: Saccardo (1892: 654), Chupp (1954: 378), Crous
& Braun (2003: 401).
Literature: Saccardo (1886: 476; 1892: 654), Lindau (1910:
90, 799), Chupp (1954: 351), Braun & Mel'nik (1997: 78),
A single species.
Zasmidium thaliae (Ellis & Langl.) U. Braun, comb.
nov.
MycoBank MB809014
(Fig. 125)
VOLUME 5 · NO. 2
325
Braun et al.
ART I CLE
geniculate, unbranched, 30–160 × 4–6 µm, (0–)1–4-septate,
pale to medium olivaceous-brown or brownish, thin-walled,
smooth; conidiogenous cells integrated, terminal, occasionally
intercalary, 15–50 µm long, conidiogenous loci conspicuous,
thickened and darkened, 2–3 µm diam. Conidia solitary,
cylindrical-obclavate, rarely subacicular, short conidia
occasionally broadly fusiform, 25–80 × 4–7 µm, 1–7-septate,
hyaline or subhyaline, apex obtuse or subobtuse, base short
to long obconically truncate, occasionally subtruncate, 2–3 µm
wide, hila thickened and darkened.
Lectotype (designated here, MycoBank MBT178160):
Sweden: Stockholm, experimental ield, on Paris quadrifolia,
12 Jul. 1882, Erikss., Fungi Paras. Scand. Exs. 85 (SF37688). Isolectotypes: Erikss., Fungi Paras. Scand. Exs.
85, e.g., B, BPI 439237, FH, HAL.
Host range and distribution: On Paris (quadrifolia, verticillata),
Trillium apetalon, Melanthiaceae, Asia (China, Japan,
Kazakhstan, Russia), Europe (Czech Republic, Denmark,
Estonia, Finland, Germany, Hungary, Italy, Latvia, Norway,
Poland, Romania, Russia, Slovakia, Slovenia, Sweden,
Ukraine).
Fig. 126. Cercospora paridis (HAL, Erikss., Fungi Paras. Scand. Exs.
85). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar
= 10 µm.
Guo (2001a: 157), Crous & Braun (2003: 308), Guo et al.
(2003: 103–104).
Notes: The conidial shape of this species is rather passaloralike, but on account of colourless conidia and cercosporalike conidiogenous loci, it is maintained in Cercospora s.
str. Molecular sequence analyses of morphologically similar
species proved that such species belong in Cercospora as
far as the conidia are colourless. A collection from Japan
on Trillium apetalon (unpublished record) is morphologically
indistinguishable from specimens on Paris spp. and at least
tentatively assigned to C. paridis. Inoculation tests and/or
molecular sequence analyses are necessary to prove the
conspeciicity of the fungi concerned.
Musaceae
Illustration: Guo et al. (2003: 104, ig. 66).
Cercospora
Exsiccatae: Bucholtz & Bondatsev, Fungi Ross. Exs. 699.
Erikss., Fungi Paras. Scand. Exs. 85. Jaczewski, Komarov &
Tranzschel, Fungi Ross. Exs. 149. Petr., Fl. Bohem. Morav.
Exs., Pilze 1910. Smarords, Fungi Lat. Exs. 199. Syd., Mycoth.
Germ. 291, 1779. Tranzschel & Sereb., Mycoth. Ross. 147,
199. Vestergr., Micromyc. Rar. Sel. Praec. Scand. 1444.
Description: Leaf spots amphigenous, at irst not very
conspicuous, only diffuse pale greenish discolorations,
later subcircular, oval to irregular, 2–15 mm diam, greyish
greenish, greenish brown, yellowish, pale straw-coloured,
margin indeinite or somewhat darker, sometimes limited by
darker veins, without or with diffuse halo, yellowish or greenish
grey, occasionally subzonate. Caespituli mainly hypophyllous,
punctiform to pustulate, dark olivaceous-brown to blackish
brown, scattered to conluent. Stromata globose or subglobose,
well-developed, substomatal to immersed, 20–110 µm diam,
dark brown, composed of swollen hyphal cells, 2–8 µm wide,
brown, thick-walled. Conidiophores in small, loose to mostly
large and dense fascicles, arising from stromata, through
stomata or erumpent, erect, subcylindrical, usually distinctly
326
A single species.
Cercospora apii Fresen. s. lat. (Crous & Braun 2003:
35)
Synonym: Cercospora hayi Calp., Studies on the Sigatoka
Disease of Bananas and its Fungus Pathogens, Atkins
Garden and Research Laboratory: 63 (1955) [type: Cuba:
on Musa ×paradisiaca, 1955, Calpouzos (FH); ex type
culture: ATCC 12234].
Literature: Kaiser & Lukezic (1965), Crous & Braun (2003:
212).
Illustration: Calpozos (1955: 61, igs 17–18).
Host range and distribution: On Musa (acuminata [?nana],
balbisiana, ×paradisiaca [×sapientum], Musa sp.), Musaceae,
Asia (Bangladesh, Philippines, Thailand, Yemen), Australia,
Central and South America (Brazil, Panama), West Indies
(Cuba), Oceania (Fiji).
IMA FUNGUS
Cercosporoid fungi 2
Doubtful, excluded and insuficiently known
species
Cercospora musae var. paradisiaca Bat. & Garnier,
Revista Agric. (Recife) 3: 54 (1963).
Notes: Type material of this variety has been found at PC
(Brazil: Pernambuco: on Musa paradisiaca, 28 Apr. 1962,
L. de Geos Vieira [Fongos do Brasil 30658], ex herb.
URM), but no sporophore of a cercosporoid fungus could
be traced. Thus, its taxonomic status and value remain
unclear.
ART I CLE
Notes: Cercospora hayi was described as follows (Calpozos
1955): “Probably not forming its own spots but found on
necrotic leaf areas or on spots of other fungi. Conidiophores
loosely to dense fasciculate, arising through the stomata,
unbranched, sparsely 2–4-septate, slightly lexuous to
tortuous, pale olivaceous to olivaceous-brown, 30–80 ×
3.5–7 µm, apex truncate, marked with one or two scars,
lateral conidial scars present. Conidia hyaline, slenderly
obclavate, straight to curved, 6–14-septate, 54–154 µm
long, maximum with 3–4.5 µm, truncate base 2.2–3.5
µm wide, tip acute, 0.7–0.9 µm wide”. The conidia were
described to be “slenderly obclavate, but they are obviously
acicular. Calpozos (1955) recorded this species from Cuba
on Musa ×paradisiaca, M. nana and M. balbisiana. Due to
similar conidiophores and colourless acicular conidia, Crous
& Braun (2003) reduced the name C. hayi to synonymy with
C. apii s. lat. (see description in Crous & Braun 2003: 35),
and Arzanlou et al. (2003: 24) conirmed that this treatment
appears to be correct based on a comparision of rDNA
sequence data. However, C. apii s. lat. (C. hayi) collections
on banana are genetically not uniform and seem to belong
to several species. Based on ITS rDNA data, isolates of C.
hayi from banana clustered with C. apii and C. nicotiana
as well as separately together with C. kikuchii (Goodwin et
al. 2011). Groenewald et al. (2013) found sequence data of
Cercospora isolates from banana in three different clades,
viz. in the Cercospora cf. citrullina clade (material from
Bangladesh), in clade N (from Bangladesh) and clade O
(from Thailand). Cercospora hayi has to be epitypyied with
material from Cuba, and this collection should be cultured
and sequenced to reveal the genuine identity of this species.
Based purely on morphology, Cercospora collections on
banana, which morphologically agree with the concept of C.
apii s. lat. as outlined in Crous & Braun (2003), can only be
refered to as the latter species in this sense.
Cercospora musarum S.F. Ashby, Bull. Dept. Agric.
(Kingston) 2: 109 (1913).
Synonyms: Brachysporium torulosum Syd., Hedwigia 49:
83 (1909) [holotype: Brazil: Pará: on Musa acuminata
[cavendishii], 24 Jan. 1908, C. F. Baker (S-F21932);
isotype: K(M) 9810].
Helminthosporium torulosum (Syd.) S.F. Ashby, Trop. Agric.
(Trinidad) 10: 6 (1932).
Deightoniella torulosa (Syd.) M.B. Ellis, Mycol. Pap. 66: 7
(1957).
Corynespora torulosa (Syd.) Crous, Persoonia 31: 211
(2013).
Literature: Chupp (1954: 403), Crous & Braun (2003: 406),
Crous et al. (2013b).
Neotype (designated here, MycoBank MBT178161):
Jamaica: on Musa ×paradisiaca, 11 Oct. 1926, Maxwell
[annotated by S. F. Ashby as Helminthosporium torulosum]
(BPI 438705).
Notes: Crous et al. (2013b) elucidated the phylogenetic
position of this species, which clusters very close to other
Corynespora species. Details of the type material were
not given in the original description, but collections made
before 1913 could not be traced so that a neotypiication
is needed.
“Cercospora stachytarphetae Ellis & Everh.”
Notes: Records of this species as a “hyperparasite” on
Macrophoma musae on Musa ×paradisiaca in India (see
Kamal 2010) are doubtful.
Pseudocercospora
Key to Pseudocercospora species on Musaceae
1
Conidia broadly obclavate-cylindrical (-fusiform), (20–)40–75 × (4–)5–8 µm ................................................ p. musicola
Conidia narrower, 2–5(–6) µm ........................................................................................................................................ 2
2 (1)
Supericial hyphae with solitary conidiophores present in vivo; fasciculate conidiophores very long,
30–120 µm and pluriseptate .................................................................................................... p. musae-sapientum
Supericial hyphae and solitary conidiophores lacking in vivo or conidiophores much shorter,
to 30 µm ................................................................................................................................................................... 3
3 (2)
Caespituli predominantly epiphyllous, in vivo sporodochial, to 100 µm wide, composed of
large stromata, to 70 µm diam, and numerous densely arranged short conidiophores,
10–25 µm long, sympodially or to 4 times percurrently proliferating; conidia subcylindrical,
mostly 30–50 × 2.5–3 µm; ascospores often germinating with 3 or 4 germ tubes,
polar and lateral, 5–6 µm wide, often distorted during germination ........................................................ p. eumusae
VOLUME 5 · NO. 2
327
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Braun et al.
Conidiophores in vivo fasciculate, large sporodochia not formed, stromata smaller,
percurrent proliferations lacking; conidia longer if cylindrical; ascospores (if sexual morph formed)
with two polar germ tubes, parallel to the long axis of the spore, 4–5 µm wide, not distorted ................................. 4
4 (3)
Conidia obclavate-cylindrical, 2–5(–6) µm wide .............................................................................................................. 5
Conidia consistently cylindrical or subcylindrical, narrow, 2–4 µm .................................................................................. 8
5 (4)
Conidiogenous loci visible as minute circles, rim very slightly thickened and darkened,
hila with similar structure; conidia subhyaline to pale olivaceous-brown;
ascospores without mucoid sheath .............................................................. P. ijiensis [Paracercospora ijiensis]
Conidiogenous loci and hila neither thickened nor darkened; conidia pale olivaceous
to brown; ascospores [in the sexual morph of P. musae] with or without mucoid sheath ........................................ 6
6 (5)
Conidiophores 10–55 µm long, 1–5-septate, usually pluriseptate throughout and geniculate-sinuous
....................................................................................................................................................... p. fengshanensis
Conidiophores 5–30 µm long, 0–2-septate, mostly 0–1-septate, barely geniculate ....................................................... 7
7 (6)
Conidia (10–)20–80(–110) × (2–)2.5–5(–6) µm; ascospores 14–18 × 3–4 µm,
with mucoid sheath ..................................................................................................................................... p. musae
Conidia shorter and above all narrower, (25–)30–45(–60) × 2–2.5(–3) µm; ascospores shorter
and narrower, (9–)10–11(–12) × (2–)2.5–3 µm, without mucoid sheath; mainly on Acacia mangium,
but also reported from Musa sp. .......................................................................................................... p. thailandica
8 (4)
Conidia very long, 80–120 × 2.5–4 µm, mostly gradually tapering toward the tip and
sometimes also base, widest in the middle .......................................................................................... p. longispora
Conidia shorter, usually < 100 µm, and narrower, 2–3 µm, cylindrical-subcylindrical, not tapering ................................ 8
9 (8)
Conidia long, (40–)78–95(–120) µm ........................................................................................................ p. indonesiana
Conidia shorter, (25–)30–70(–83) µm ........................................................................................................................... 10
10 (9) Conidia (30–)60–70(–83) µm long, cylindrical .......................................................................................... p. assamensis
Conidia shorter, (25–)30–45(–60) µm long, cylindrical to obclavate-cylindrical,
mainly on Acacia mangium, but also reported from Musa sp. ............................................................. p. thailandica
Pseudocercospora species on Musaceae
pseudocercospora assamensis Arzanlou & Crous,
Persoonia 20: 28 (2008).
(Fig. 127)
Illustration: Arzanlou et al. (2008: 28–29, igs 7–8).
Description: In vitro (on MEA): Mycelium submerged and
supericial; submerged hyphae 2.5–4 µm wide, branched,
septate, medium brown, thin-walled, smooth; aerial hyphae
medium brown, thin-walled, smooth. Conidiophores solitary,
arising from supericial hyphae, lateral, subcylindrical,
unbranched or branched, to 20 µm long and 2–3 µm
wide, 0–1-septate, medium brown, thin-walled, smooth;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, sympodial, conidiogenous
loci inconspicuous. Conidia solitary, subcylindrical, straight to
curved, older conidia sometimes with irregular swellings, (30–
)60–70(–83) × 2–3 µm, pluriseptate, pale brown, thin-walled,
smooth, apex rounded, base truncate to slightly obconically
truncate, about 1 µm wide, hila unthickened, not darkened.
aerial mycelium and entire, smooth margin, surface pale
mouse-grey to mouse-grey, olivaceous in reverse. Colonies
on OA reaching 35 mm diam after 30 d at 24 °C, effuse, with
moderate velvety aerial mycelium and entire, smooth margin,
surface pale mouse-grey and iron-grey in reverse.
Holotype: India: Assam: Naojan, on Musa cv. Nanderan
(Plantain), Musaceae, 2005, I. Buddenhagen (CBS H-20044).
Ex-type culture: CBS 122467.
Host range and distribution: Only known from the type
collection.
pseudocercospora eumusae Crous & Mour., Sydowia
54: 36 (2002).
(Fig. 128)
Synonym: Mycosphaerella eumusae Crous & Mour., Sydowia
54: 36 (2002) [holotype: réunion: on Musa sp., 2001, J.
Carlier (PREM 57314)].
Literature: Arzanlou et al. (2008: 26).
Illustration: Crous & Mourichon (2002: 37–38, igs 1–12).
Culture characteristics: Colonies on MEA reaching 47 mm
diam after 30 d at 24 °C, elevated in the centre, with abundant
328
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 127. Pseudocercospora assamensis (CBS H-20044). A. Hyphae
with conidiophores. B. Conidia. Bar = 10 µm. P.W. Crous del.
Description: Leaf spots amphigenous, initially visible as faint
brown streaks, later oval to elliptical light brown lesions with grey
centre and dark brown border, inally conluent, forming larger,
brown, necrotic areas under favourable conditions, grey spots
and patches visible in necrotic areas, leasion surrounded by a
diffuse chlorotic yellowish halo. Caespituli mainly epiphyllous,
punctiform, grey. Mycelium internal; hyphae branched, septate,
1.5–2.5 µm wide, pale brown, thin-walled, smooth. Stromata
well-developed, to 70 µm diam, subepidermal or substomatal,
brown. Conidiophores numerous, in dense fascicles, arising
from stromata, forming sporodochial conidiomata, through
stomata or erumpent, erect, straight, subcylindrical or somewhat
attenuated towards the tip to geniculate-sinuous, unbranched
or branched, 10–25 × 3–5 µm, 0–3-septate, hyaline to pale
brown, thin-walled, smooth; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells, 10–
20 µm long, sympodial to percurrent, with 1–4 ine annellations,
conidiogenous loci inconspicuous or visible as truncate tip,
but always unthickened and not darkened. Conidia solitary,
subcylindrical, straight to curved, (18–)30–50(-65) × (2–)2.5–3
µm, 3–8-septate, subhyaline to pale olivaceous, thin-walled,
smooth, apex obtuse, base subtruncate to short obconically
truncate, hila unthickened, not darkened.
Spermogonia mainly hypophyllous, subepidermal,
substomatal, to 75 µm diam, dark brown; spermatia rodshaped, 3–6 × 1 µm, hyaline. Mycosphaerella-like sexual
morph: Ascomata amphigenous, mainly hypophyllous,
subepidermal, somewhat erumpent, black, to 80 µm diam,
VOLUME 5 · NO. 2
Fig. 128. Pseudocercospora eumusae (PREM 57315). A–B.
Conidiophore fascicles. C–d. Conidia. e. Spermatia. F. Asci. g.
Ascospores. h. Germination ascospore. Bar = 10 µm. P.W. Crous
del.
apical ostiole 10–15 µm diam, wall composed of 2–3 layers
of medium brown textura angularis, asci aparaphysate,
fasciculate, bitunicate, subsessile, obovoid, straight to slightly
curved, 30–50 × 9–15 µm, 8-spored, ascospores multiseriate,
overlapping, obovoid, tapering towards both ends, but with
more pronounced taper towards the base, widest in the middle
of the upper cell, (11–)12–13(–16.5) × (3–)3.5–4(–4.5) µm,
medianly 1-septate, hyaline, guttulate, ends rounded.
Holotype: réunion: on Musa sp., 2001, J. Carlier (PREM
57315).
Host range and distribution: On Musa spp., Musaceae,
Africa (Mauritius, Nigeria, Réunion), Asia (India, Sri Lanka,
Thailand, Malaysia, Vietnam).
pseudocercospora fengshanensis (T.Y. Lin & J.M.
Yen) J.M. Yen & S.K. Sun, Cryptog. Mycol. 4: 197
(1983).
(Fig. 129)
Basionym: Cercospora fengshanensis T.Y. Lin & J.M. Yen,
Rev. Mycol. 35: 320 “1970” (1971).
Literature: Hsieh & Goh (1990: 243), Guo & Hsieh (1995:
217), Guo et al. (1998: 229–230), Crous & Braun (2003: 183).
329
Braun et al.
ART I CLE
paler towards the tip, thin-walled, smooth; conidiogenous
cells integrated, terminal, conidiogenous loci inconspicuous
or subdenticuate, but always unthickened and not darkened.
Conidia solitary, obclavate-cylindrical, straight to somewhat
curved, 15–95 × 3–5 µm, 2–9-septate, occasionally
constricted at septa, pale olivaceous or brown, thin-walled,
smooth, apex obtuse, base short obconically truncate, hila
neither thickened nor darkened.
Holotype: taiwan: Feng-Shan, on Musa sp., Apr. 1970, T.
Whang F.S. 4 (not traced; probably not preserved).
Host range and distribution: On Musa (acuminata [nana],
Musa sp.), Musaceae, Asia (China, Taiwan).
Note: This species is morphologically close to P. musae,
but differs in having to 5-septate, mostly geniculate-sinuous
conidiophores.
Pseudocercospora ijiensis (M. Morelet) Deighton,
Mycol. Pap. 140: 144 (1976).
(Fig. 130)
Basionym: Cercospora ijiensis M. Morelet, Ann. Soc. Sci.
Nat. Archéol. Toulon Var. 21: 105 (1969).
Synonyms: Mycosphaerella ijiensis M. Morelet, Ann. Soc.
Sci. Nat. Archéol. Toulon Var. 21: 105 (1969) [holotype:
see P. ijiensis].
Cercospora pingtungensis T.Y. Lin & J.M. Yen, Bull. Trimestriel
Soc. Mycol. France 87: 427 (1971) [type: taiwan:
Pingtung, on Musa acuminata [canvendishii], 5 and 27
Oct. 1970, B. Shiu, P.T. no. 52 (probably not preserved)].
Cercospora ijiensis var. difformis J.L. Mulder & Stover, Trans.
Brit. Mycol. Soc. 67: 82 (1976) [holotype: honduras: La
Lima, on Musa sp., Mar. 1974, R. H. Stover (K(M) IMI
183747)].
Mycosphaerella ijiensis var. difformis J.L. Mulder & Stover,
Trans. Brit. Mycol. Soc. 67: 82 (1976).
Paracercospora ijiensis (M. Morelet) Deighton, Mycol. Pap.
144: 51 (1979).
Paracercospora ijiensis var. difformis (J.L. Mulder & Stover)
Deighton, Mycol. Pap. 144: 52 (1979).
Fig. 129. Pseudocercospora fengshanensis (based on Guo & Hsieh
1995: 219, ig. 185). A. Conidiophore fascicles. B. Conidiophore. C.
Conidia. Bar = 10 µm.
Illustrations: Lin & Yen (1971b: 319, ig. 1), Guo & Hsieh
(1995: 219, ig. 185), Guo et al. (1998: 230, ig. 190).
Description: Leaf spots amphigenous, distinct, oblongelliptical, scattered to conluent, 6–45 × 4–12 mm, centre
greyish brown to greyish white, with narrow, dark brown to
blackish margin and pale grey to yellowish brown halo on
the upper leaf surface. Caespituli amphigenous. Mycelium
internal. Stromata lacking or small, 15–40 µm diam,
substomatal, globose, dark brown. Conidiophores in small,
loose to dense fascicles, 2–7, rarely solitary, arising from
internal hyphae or stromata, through stomata, erect, straight
to lexuous, somewhat geniculate-sinuous, unbranched,
10–55 × 4–6 µm, 1–5-septate, brown or olivaceous-brown,
330
Literature: Meredith & Lawrence (1970b), Mulder & Stover
(1976), Deighton (1979: 51–52), Sivanesan (1984: 216–
218), Mulder & Holliday (1974a), Johanson & Jeger (1993),
Johanson et al. (1994), Crous et al. (2002: 51), Crous &
Braun (2003: 185), Gasparotto et al. (2005), Aptroot (2006:
91), Arzanlou et al. (2008: 26).
Illustrations: Meredith & Lawrence (1970b: 465–471, igs 1–8),
Lin & Yen (1971a: 426, ig. 1), Mulder & Stover (1976: 79–82,
igs 2–11), Mulder & Holliday (1974a: ig., unnumbered),
Deighton (1979: 51, ig. 27 A–C), Sivanesan (1984: 217, ig.
116), Guo et al. (2005: 293, ig. 215), Aptroot (2006: 13, ig.
320), Arzanlou et al. (2008: 35, igs 18 b, e, 19).
Description: Leaf spots amphigenous, symptoms variable,
at irst formed as minute, faint, reddish brown specks, later
elongated and wider, visible as narrow reddish brown streaks,
to 20 × 2 mm, parallel to leaf veins, conluent, later turning
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
straight to curved or somewhat lexuous, subcylindrical or
somewhat attenuated towards the tip, barely to moderately
geniculate-sinuous, unbranched, 10–65 × 3–6(–7) µm,
0–3(–5)-septate, pale to medium olivaceous-brown or
brown, thin-walled, smooth; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells,
about 10–30 µm long, conidiogenous loci inconspicuous
to subconspicuous by being ring-like, i.e. with slighty
thickened and darkened rim (paracercospora-like), about
1.5–2.5 µm wide. Conidia solitary, obclavate to cylindrical
or subcylindrical, size rather variable, 30–130 × 2.5–5 µm,
1–10-septate, subhyaline to pale olivaceous-brown, thinwalled, smooth, apex obtuse to subacute, base subtruncate to
short obconically truncate, 1.5–3 µm wide, hila unthickened,
not darkened or only ultimate rim slighty so.
Spermogonia often associted with conidiophores,
amphigenous,
immersed,
hour-glass-shaped,
oval,
obpyriform to subglobose, about 30–90 µm diam, pale to
medium brown; spermatia short cylindrical-bacilliform, 2.5–5
× 1–2.5 µm.
Mycosphaerella-like
sexual
morph:
Ascomata
amphigenous, scattered, immersed, globose, 40–85 µm
diam, with narrow to moderately thick papillata ostiole,
somewhat erumpent, aparaphysate, wall of three or more
layers of polygonal, brown cells; asci numerous, cylindricalobclavate, about 28–35 × 6.5–8 µm, 8-spored; ascospores
biseriate, fusiform-clavate, 11.5–20 × 2.5–6 µm, with a single
septum, median or somewhat supramedian, not or slightly
constricted at the septum, colourless, without mucoid sheath,
germinating with two germ tubes both from the polar ends,
not distorted, growth parallel to the long axis of the spore,
4–5 µm wide.
Holotype: hawaii: on Musa sp., 12 Dec. 1968, D. S. Meredith
& J. S. Lawrence (K(M) IMI 136696). Epitype (designated by
Arzanlou et al. 2008: 26): Cameroon: on Musa sp., date and
collector unknown (CBS H-20037), ex epitype culture CIRAD
86 = CBS 120258.
Fig. 130. Pseudocercospora ijiensis (K(M) IMI 136696). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
dark brown to blackish, occasionally with a dark purplish
tinge, older leaf spots elliptical-fusiform, with light brown
water-soaked border, inally with dark brown to blackish
centre, slightly depressed, at the end drying, turning to pale
grey or buff-coloured, with distinct dark brown to blackish
margin. Caespituli amphigenous, distinct on the upper leaf
surface, punctiform, dark brown, less conspicuous on the
lower leaf surface. Mycelium internal. Stromata lacking or
almost so on the lower leaf surface, developed on the upper
side, subglobose, substomatal to immersed, erumpent, about
10–50 µm diam, composed of pigmented cells, circular to
polygonal in outline. Conidiophores solitary or in small (2–8)
to moderately large fascicles, loose to moderately dense,
arising from internal hyphae or stromata, through stomata
or erumpent, more numerous and denser when arising from
stromata on the upper leaf surface, fascicles occasionally
arising from immature spermogonia or ascoma initials, erect,
VOLUME 5 · NO. 2
Host range and distribution: On Musa (acuminata,
×paradisiaca, Musa spp.), Musaceae, ?Heliconia psittacorum,
Heliconiaceae; Africa (Benin, Burundi, Cameroon, Central
African Republic, Congo, Comoros, Gabon, Ghana, GuineaBissau, Ivory Coast, Kenya, Malawi, Mayotte, Niger, Nigeria,
Rwanda, São Tomé e Príncipe, Somalia, Tanzania, Togo,
Tonga, Uganda, Zambia), Asia (Bhutan, Brunei, China,
Indonesia, Japan, Malaysia, Papua New Guinea, Peru,
Philippines, Singapore, Taiwan, Thailand, Vietnam), Australia,
Oceania (American Samoa, Cook Islands, Fiji, French
Polynesia, Hawaii, Micronesia, New Caledonia, Niue, Norfolk
Islands, Northern Mariana Islands, Samoa, Solomon Islands,
Tahiti, Tonga, Vanuatu, Wallis and Futuna Islands), Central
and South America (Belize, Bolivia, Brazil, Colombia, Costa
Rica, Dominican Republ., Ecuador, El Salvador, Guatemala,
Guyana, Honduras, Nicaragua, Panama, Venezuela), North
America (Mexico; USA, Florida), West Indies (Cuba, Jamaica,
Martinique, Netherlands Antilles, Haiti).
Notes: As the conidiogenous loci and conidial hila are similar
to those of Paracercospora egenula, the type species of
331
ART I CLE
Braun et al.
Paracercospora, Pseudocercospora ijiensis, was previously
considered to be a species of the latter genus. However,
P. ijiensis clusters within the large Pseudocercospora
clade (Arzanlou et al. 2008, Crous et al. 2013a), and
must thus be retained in this genus, although in molecular
analyses Paracercospora egenula and thus also the genus
Paracercospora proved to represent a separate genus of its
own (Crous et al. 2013a), i.e. the special paracercosporalike scar type is not suficient to distinguish Paracercospora
and Pseudocercospora (see Braun et al. 2013a). The status
of Cercospora pingtungensis (Lin & Yen 1971a; Hsieh &
Goh 1990: 242–243; Guo et al. 2005: 292–293, ig. 215) is
unclear, as type material could not be traced. According to
the original description and illustration, this species cannot
be maintained in Cercospora s. str. The more or less
cylindrical conidia are pigmented and the conidiogenous loci
range from being inconspicuous, truncate, unthickened, not
darkened to conspicuous, prominent. These characters and
the dimensions of conidiophores and conidia agree well with
those of Pseudocercospora (Paracercospora) ijiensis. The
described stromata in C. pingtungensis are larger than in
common collections of P. ijiensis, but the description of Lin &
Yen (1971a) was mainly based on epiphyllous colonies which
differ from hyphophyllous ones in having larger stromata,
corresponding to the description of Cercospora ijiensis
var. difformis (Mulder & Stover 1976). Furthermore, very
large stromata seem to refer to immature spermogonia, as
suggested by Lin & Yen (1971a: 426, ig. 1A), who classiied
these structures as “pycnidiostromata”. It is not unusual in
cercosporoid fungi that conidiophore fascicles may arise from
ascomata and that epiphyllous and hypophyllous colonies
are morphologically differentiated. Cercospora pingtungensis
is tentatively reduced to synonymy with P. ijiensis as
this species is very probably identical, but it is, of course,
necessary to conirm this treatment by a re-examination of
type material or a neotypiication with material from Taiwan.
The differentiation between P. ijiensis var. ijiensis and var.
difformis is vague and barely justiied. Pseudocercospora
ijiensis is morphologically very variable, and the occurrence
of morphological differences between epi- and hypophyllous
colonies in Pseudocercospora spp. is not unusual and a
common phenomenon in numerous species of this genus.
Gasparotto et al. (2005) recorded Heliconia psittacorum
as host of P. ijiensis, based on positive results of inoculation
tests. Morphological details of the fungus on Heliconia
psittacorum were not given. As this host belongs to another
family, cultures and molecular sequence analyses would be
useful to conirm that P. ijiensis is able attack Heliconia.
pseudocercospora indonesiana Arzanlou & Crous,
Persoonia 20: 29 (2008).
(Fig. 131)
Illustration: Arzanlou et al. (2008: 29, ig. 9, 30, ig. 10).
Description: In vitro (on MEA): Mycelium submerged and
supericial; submerged hyphae 2.5–4 µm wide, branched,
septate, medium brown, thin-walled, smooth; aerial hyphae
medium brown, thin-walled, smooth. Conidiophores solitary,
arising from supericial hyphae, lateral, subcylindrical,
332
Fig. 131. Pseudocercospora indonesiana (CBS H-20045). A.
Hyphae with conidiophores. B. Conidia. Bar = 10 µm. P.W. Crous del.
unbranched, to 30 µm long and 2–2.5 µm wide, 0–2-septate,
medium brown, thin-walled, smooth; conidiogenous cells
integrated, terminal or conidiophores reduced to conidiogenous
cells, sympodial, conidiogenous loci inconspicuous. Conidia
solitary, subcylindrical, straight to curved, (40–)78–95(–120)
× 2–3 µm, 3–7-septate, guttulate, pale brown, thin-walled,
smooth, apex rounded, base truncate to slightly obconically
truncate, hila unthickened, not darkened.
Culture characteristics: Colonies on MEA reaching 27 mm
diam after 30 d at 24 °C, low convex, with abundant aerial
mycelium and entire, smooth margin, surface pale mousegrey to mouse-grey, dark mouse-grey in reverse. Colonies
on OA reaching 35 mm diam after 47 d at 24 °C, effuse, with
moderate aerial mycelium and entire, smooth margin, surface
pale mouse-grey and olivaceous-black in reverse.
Holotype: Indonesia: Western Sumatra: Kumango, on Musa
cv. Buai, Musaceae, 2004, I. Buddenhagen (CBS H-20045).
Ex-type culture: CBS 122473.
Host range and distribution: Only known from the type
collection.
pseudocercospora longispora Arzanlou & Crous,
Persoonia 20: 30 (2008).
(Fig. 132)
Illustration: Arzanlou et al. (2008: 30–31, igs 11–12).
IMA FUNGUS
Cercosporoid fungi 2
Host range and distribution: Only known from the type
collection.
ART I CLE
Holotype: Indonesia: Western Sumatra: Kumango, on Musa
cv. Buai, Musaceae, 2004, I. Buddenhagen (CBS H-20043).
Ex-type culture: CBS 122470.
pseudocercospora musae (Zimm.) Deighton, Mycol.
Pap. 140: 148 (1976).
(Fig. 133)
Basionym: Cercospora musae Zimm., Centralbl. Bakteriol.,
Abt. 2, 8: 219 (1902).
Synonyms: Cercospora musae Massee, Bull. Misc. Inform.
Kew 28: 159 (1914), nom. illeg. (ICN, Art. 53.1) [type: Fiji:
on Musa sp., C. H. Knowles (K)].
Mycosphaerella musicola R. Leach, Trop. Agric. (Trinidad)
18: 92 (1941), nom. inval. (ICN, Art. 39.1).
Mycosphaerella musicola R. Leach, J.L. Mulder & R.H. Stover,
Trans. Brit. Mycol. Soc. 67: 77 (1976) [holotype: Jamaica:
on Musa ×paradisiaca [×sapientum], Jan. 1959, R. Leach
(K(M) IMI 75804a); epitype (designated in Arzanlou et al.
2008: 28): Cuba: Bayamo, on Musa ×paradisiaca, 1 Dec.
1966, R. Urtiaga (CBS H-20038), ex epitype culture K(M)
IMI 123823 = CBS 116634].
Fig. 132. Pseudocercospora longispora (CBS H-20043). A. Hyphae
with conidiophores. B. Conidia. Bar = 10 µm. P.W. Crous del.
Description: In vitro: Mycelium on MEA submerged and
supericial; submerged hyphae 2–3 µm wide, branched,
septate, medium brown, thin-walled, smooth; aerial
hyphae medium brown, thin-walled, smooth, later hyphal
cells becoming thick-walled, swollen, forming dark brown,
monilioid, muriform cells, 5–17 × 7–12 µm. Conidiophores
solitary, arising from supericial hyphae, lateral, straight,
subcylindrical, unbranched or branched, to 30 µm long
and 2–3 µm wide, 0–2-septate, medium brown, thinwalled, smooth; conidiogenous cells integrated, terminal
or conidiophores reduced to conidiogenous cells,
sympodial, conidiogenous loci inconspicuous. Conidia
solitary, cylindrical to subcylindrical, widest in the middle,
gradually tapering towards the apex and sometimes also
base, straight to curved, 80–120 × 2.5–4 µm, pluriseptate,
pale brown, thin-walled, smooth, apex subobtuse, base
truncate to slightly obconically truncate, hila unthickened,
not darkened.
Culture characteristics: Colonies on MEA reaching 15 mm
diam after 30 d at 24 °C, erumpent, with moderately developed
aerial mycelium and entire, smooth margin, surface buff
to rosy-buff, mouse-grey to dark grey, dark mouse-grey in
reverse. Colonies on OA reaching 15 mm diam after 30 d
at 24 °C, effuse, with abundant aerial mycelium and entire,
smooth margin, surface pale mouse-grey and dark mousegrey in reverse.
VOLUME 5 · NO. 2
Literature: Saccardo (1906: 610; 1931: 885), Chupp (1954:
402–403), Batista et al. (1965: 7), Meredith & Lawrence
(1970a), Mulder & Stover (1976: 77), Mulder & Holliday
(1974b), Sivanesan (1984: 209), Hsieh & Goh (1990: 243),
Guo & Hsieh (1995: 218), Crous & Braun (1996b: 284), Guo
et al. (1998: 230–231), Crous & Braun (2003: 286), Aptroot
(2006: 139), Arzanlou et al. (2008: 28), Kamal (2010: 200),
Phengsintham et al. (2013b: 116), Braun & Urtiaga (2013a:
195).
Illustrations: Batista et al. (1965: 18, ig. 4), Meredith &
Lawrence (1970a: 270, ig. 2), Mulder & Holliday (1974b, ig.,
unnumbered), Guo & Hsieh (1995: 220, ig. 186), Guo et al.
(1998: 231, ig. 191), Aptroot (2006: 16, ig. 579), Arzanlou et
al. (2008: 35, ig. 18 c, f), Phengsintham et al. (2013b: 11, ig.
116, 118, ig. 117).
Description: Leaf spots amphigenous, subcircular, elliptical,
oblong or somewhat irregular, 2–50(–75) mm diam, scattered
to conluent, brownish, reddish brown, dingy greyish brown
or grey, margin brown or purplish to reddish. Caespituli
amphigenous, punctiform, dark. Mycelium internal. Stromata
substomatal to immersed, subglobose, 15–40 µm diam, brown
to blackish brown. Conidiophores in small to moderately
large, usually dense fascicles, mostly 5–20, arising from
stromata, sporodochium-like, emerging through stomata
or erumpent, erect, straight to slightly curved or sinuous,
subcylindrical to conical, usually none or at most slightly
geniculate, unbranched or only rarely branched, 5–30 × 2–6
µm, 0–2-septate, very pale olivaceous to olivaceous-brown,
thin-walled, smooth; conidiogenous cells integrated, terminal
or conidiophores reduced to conidiogenous cells, 5–20 µm
long, conidiogenous loci inconspicuous or visible as truncate
tip, but always unthickened and not darkened. Conidia
solitary, cylindrical to obclavate-cylindrical, straight to curved
333
Braun et al.
ART I CLE
diam, aparaphysate; asci oblong-clavate, 29–36 × 8–11 µm;
ascospores 14–18 × 3–4 µm, short subcylindrical, upper cells
a little wider, medianly 1-septate, hyaline, with mucoid sheath,
germinating from both polar ends, 4–5 µm wide, germ tubes
parallel to the long axis of the spore, not becoming distorted.
Culture characteristics: Colonies on PDA reaching 8.5
mm diam after 38 d at 26 °C (average, maximum 12 mm),
at irst pale to medium grey (after 2–3 d), later dark green to
almost black (on V-8 juice agar pale to medium grey with more
abundant aerial mycelium compared to PDA), aerial mycelium
developed in clamps made of contorted, branched hyphae,
older colonies somewhat raised, almost hemispherical in side
view, margin lobed and irregular in surface view, centre inally
becoming hallow, outer shell consisting of a thick, almost
black rind made of dumb-bell-shaped hyphal cells; stromata
not developed in vitro, but colonies sporulating on PDA after
2–3 d.
[Type: Java: Buitenzorg, on Musa ×paradisiaca [×sapientum]
(not preserved)]. Neotype (designated here, MycoBank
MBT178162): Cuba: Bayamo, on Musa ×paradisiaca, 1 Dec.
1966, R. Urtiaga (CBS H-20038). Ex-neotype culture: K(M)
IMI 123823 = CBS 116634.
Fig. 133. Pseudocercospora musae (CBS H-20038). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
or somewhat sinuous, (10–)20–80(–110) × (2–)2.5–5(–6) µm
in vivo (in vitro with a tendency to be somewhat longer and
narrower, 2–3.5 µm), (0–)2–7(–9)-septate, subhyaline to pale
olivaceous, thin-walled, smooth, apex obtuse, rounded to
somewhat narrowed, base rounded to obconically truncate,
1.5–2.5 µm wide, hila unthickened, not darkened.
Spermogonia amphigenous, mainly hypophyllous,
immersed, hour-glass-shaped to lask-shaped, (30–)45–
90(–110) µm diam, wall composed of 2–3 layers of pale to
medium brown rectangular-polygonal cells; spermatia rodshaped, 2–6 × 0.8–1.4 µm, hyaline.
Ascomata
amphigenous,
immersed,
somewhat
erumpent, dark brown to blackish, ostiole small, 45–70 µm
334
Host range and distribution: On Musa (acuminata [cavendishii,
?nana], balbisiana var. liukiuensis [liukiuensis], banksii,
basjoo, ×paradisiaca [×sapientum], textilis, ventricosa,
Musaceae (widely distributed, wherever Musa spp. are
cultivated or grow naturally), Africa (Angola, Cameroon,
Cape Verde, Congo, Egypt, Ethiopia, Gabon, Ghana,
Guinea, Guinea-Bissau, Ivory Coast, Kenya, Madagascar,
Malawi, Mauritius, Mozambique, Nigeria, São Tomé e
Príncipe, Sierra Leone, Somalia, South Africa, Tanzania,
Togo, Uganda, Zambia, Zimbabwe), Asia (Brunei, Bhutan,
Cambodia, China, India, Indonesia, Japan, Laos, Malaysia,
Nepal, Papua New Guinea, Philippines, Sri Lanka, Taiwan,
Thailand, Vietnam, Yemen), Australia, Oceania (American
Samoa, Fiji, Hawaii, Palau, French Polynesia, Guam,
Kiribati, Micronesia, New Caledonia, Niue, Norfolk Island,
Samoa, Solomon Islands, Tonga, Tuvalu, Vanuatu, Wallis
and Futuna Islands), Central and South America (Argentina,
Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, El
Salvador, French Guiana, Guatemala, Guyana, Honduras,
Nicaragua, Panama, Peru, Suriname, Venezuela), North
America (Mexico; USA, Florida, Texas), West Indies
(Antigua and Barbuda, Bahamas, Barbados, Cayman
Islands, Cook Islands, Cuba, Dominican Republ., Grenada,
Guadeloupe, Haiti, Jamaica, Martinique, Montserrat, Puerto
Rico, Saint Kitts and Nevis, Saint Lucia, Saint Vincent and
the Grenadines, Trinidad and Tobago).
pseudocercospora musae-sapientum (A.K. Kar &
M. Mandal) U. Braun & Mouch., New Zealand J.
Bot. 37: 317 (1999); as “musae-sapienti”.
(Fig. 134)
Basionym: Cercospora musae-sapientum A.K. Kar & M.
Mandal, Norw. J. Bot. 22: 105 (1975); as “musae-sapienti”.
Literature: Crous & Braun (2003: 287), Kamal (2010: 200).
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 134. Pseudocercospora musaesapientum (K(M) IMI 147922). A. Solitary
conidiophores arising from supericial
hyphae. B. Conidiophore fascicles.
C. Conidiophores and supericial
hyphae emerging through stomata. d.
Conidiophores. e. Conidia. Bar = 10 µm.
Illustrations: Kar & Mandal (1975: 105, ig. 1), Braun et al.
(1999: 315, ig. 18).
Description: Leaf spots amphigenous, numerous,
scattered to conluent, oblong, limited by veins, about
2–10 × 0.2–4 mm, brown to deep brownish grey. Caespituli
hypophyllous, punctiform to subeffuse, dark. Mycelium
internal and external; supericial hyphae emerging through
stomata, 1.5–3 µm wide, branched, septate, olivaceous
to pale olivaceous-brown, thin-walled, smooth. Stromata
almost lacking or small, 10–35 µm diam, substomatal,
olivaceous-brown to brown. Conidiophores in loose, mostly
small fascicles, emerging through stomata and solitary,
arising from supericial hyphae, lateral, erect, straight,
subcylindrical to geniculate, unbranched or occasionally
branched, 15–120 × 3–7 µm, aseptate to pluriseptate
throughout, olivaceous-brown to medium dark brown,
wall thin to somewhat thickened, smooth; conidiogenous
cells integrated, terminal and intercalary or conidiophores
occasionally reduced to conidiogenous cells, about 10–30
µm long, conidiogenous loci inconspicuous to denticle-like,
but always unthickened, not darkened. Conidia solitary,
obclavate-cylindrical, 30–100 × 3–5 µm, 3–8-septate,
apex subacute to obtuse, base obconically truncate, hila
unthickened, not darkened. Conidia solitary, obclavate
VOLUME 5 · NO. 2
to cylindrical, straight to curved, 30–100 × 3–5 µm,
2–8-septate, pale olivaceous to olivaceous-brown, thinwalled, smooth, apex obtuse to subacute, base almost
truncate or short to long obconically truncate, about 2–2.5
µm wide, hila unthickened, not darkened.
Holotype: India: West Bengal: Midnapur, Daspur, on Musa
×paradisiaca [×sapientum], 16 Jan. 1970 (K(M) IMI 147922).
Host range and distribution: On Musa (×paradisiaca, Musa
sp.), Musaceae, Asia (India, West Bengal), Oceania (New
Caledonia, Vanuatu, Wallis and Futuna).
Notes: Beside type material, several collections of this
species from Oceania, deposited at PC, have been
examined. Pseudocercospora musae-sapientium is the
only Pseudocercospora on banana which forms supericial
hyphae with solitary conidiophores in vivo.
pseudocercospora musicola U. Braun, New Zealand
J. Bot. 37: 317 (1999).
(Fig. 135)
Synonyms: Cercospora musicola Sawada, Agric. Res. Inst.
Formosa Rep. 85: 116 (1943); as “musaecola; nom. inval.
(ICN, Art. 36.1).
335
Braun et al.
ART I CLE
Conidiophores solitary or in small to moderately large,
usually loose fascicles, arising from internal hyphae or
hyphal aggregations, emerging through stomata, geniculatesinuous, unbranched, 50–180 × 3–9 µm, pluriseptate
throughout, pale to medium brown, wall somewhat thickened,
smooth or almost so; conidiogenous cells integrated, terminal
and intercalary, about 10–30 µm long, conidiogenous loci
subdenticulate, but neither thickened nor darkened. Conidia
solitary, obclavate-cylindrical (-fusiform), (20–)40–75 × (4–)
5–8 µm, 1–7-septate, occasionally constricted at the septa,
pale yellowish or olivaceous to medium brown, wall thin
to somewhat thickened, smooth, apex obtuse, base short
obconically truncate, about 2–3 µm wide, hila unthickened,
not darkened.
Holotype: taiwan: Taipei, on Musa acuminata [cavendishii],
1 Jul. 1909, K. Sawada (BPI 438701). Isotype: BPI 438702.
Paratype: taiwan: Taipei, on Musa balbisiana var. liukiuensis,
18 Jul. 1911, Y. Fujikuro, [deposited as Cercospora musaeliukiuensis] (BPI 43873).
Host range and distribution: On Musa (acuminata, balbisiana
var. liukiuensis [liukiuensis], basjoo, ×paradisiaca),
Musaceae, Asia (Taiwan).
Note: The holotype of this species is syntype material of the
invalid designation Cercospora musicola .
pseudocercospora thailandica Crous, Himaman &
M.J. Wingf., Stud. Mycol. 50: 465 (2004).
(Fig. 136)
Synonym: Mycosphaerella thailandica Crous, Himaman &
M.J. Wingf., Stud. Mycol. 50: 465 (2004) [holotype: see
type of P. thailandica].
Illustration: Crous et al. (2004: 466, igs 11–15).
Fig. 135. Pseudocercospora musicola (BPI 438701). A. Conidiophore
fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
Cercospora musicola Sawada ex Goh & W.H. Hsieh,
Cercospora and Similar Fungi from Taiwan: 242 (1990);
nom. inval. (ICN, Art. 37.1).
Cercospora musae-liukiuensis Sawada, Coll. Agric. Natl.
Taiwan Univ., Spec. Publ. 8: 221 (1959), nom. nud. [type:
taiwan: on Musa balbisiana var. liukiuensis [liukiuensis]
(NTU-PPE, hb. Sawada)].
Literature: Chupp (1954: 403), Katsuki (1965: 48), Lin & Yen
(1971: 318), Tai (1979: 891), Hsieh & Goh (1990: 202), Crous
& Braun (2003: 287).
Illustrations: Lin & Yen (1971b: 319, ig. 1 D–G), Braun et al.
(1999: 315, ig. 19).
Description: Leaf spots lacking or diffuse. Caespituli
hypophyllous, arrangement linear, 2–15 × 0.2–2 mm, reddish
to dark brown, mostly dense. Mycelium internal; hyphae
septate, branched, brown. Stromata lacking or small, only
composed of a few swollen hyphal cells, substomatal.
336
Description: Leaf spots amphigenous, irregular blotches
covering large parts of the leaf lamina, associated with
symptoms including tip blight or lesions all along the margin
of the leaf, frequently extending to the middle of the leaf
lamina, medium brown, margin dark brown, raised. Caespituli
amphigenous, punctiform, brown. Mycelium internal.
Stromata well-developed, immersed, to 25 µm wide and 30
µm high, brown. Conidiophores in dense fascicles, arising
from stromata, erect, straight to curved, subcylindrical,
unbranched, 10–20 × 5–5 µm, 0–2-septate, pale brown, thinwalled, smooth; conidiogenous cells integrated, terminal or
conidiophores reduced to conidiogenous cells, 10–15 µm long,
conidiogenous loci inconspicuous, truncate, unthickened, not
darkened. Conidia solitary, narrowly obclavate-cylindrical,
straight to somewhat curved, (25–)30–45(–60) × 2–2.5(–3)
µm, 3–6-septate, pale brown, thin-walled, smooth, apex
subobtuse, base short obconically truncate.
Spermogonia intermixed with and similar to ascomata
in general morphology; spermatia rod-shaped, 3–5 × 1 µm,
hyaline.
Sexual morph: Ascomata amphigenous, subepidermal,
becoming erumpent, to 80 µm diam, black, ostiole 5–10 µm
diam, wall composed of 2–3 layers, medium brown, textura
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
angularis; asci fasciculate, obovoid to narrowly ellipsoid, straight
to slightly curved, subsessile, 30–40 × 6–8 µm, bitunicate,
8-spored; ascospores tri- to multiseriate, overlapping, fusoidellipsoid, straight to slightly curved, (9–) 10–11(–12) × (2–)
2.5–3 µm, medianly 1-septate, slightly constricted at the
septum, widest in the middle of the upper cell, hyaline, ends
obtuse, attenuated towards both ends, but more prominently
to the base, germination (on MEA after 24 h) with germ tubes
parallel to the long axis of the spore, developing several lateral
branches, constricted at the septum.
In culture: Colonies slightly erumpent, with smooth regular
margin, fast growing, covering the dish after 60 d, aerial
mycelium luffy, surface grey-olivaceous, reverse olivaceousblack.
Holotype:
thailand:
Chachoengsao
Province:
Sanamchaikhet, on Acacia mangium, 28 May 2003, K.
Pongpanich (CBS H-9875). Ex-type culture: CBS 116367
Host range and distribution: On Acacia mangium, Fabaceae,
and Musa sp., Musaceae, Asia (Thailand), West Indies
(Windward Islands).
Fig. 136. Pseudocercospora thailandica (CBS H-9875). A.
Conidiophore fascicle. B. Conidia. Bars = 10 µm.
Notes: Pseudocercospora thailandica is a primary pathogen
of Acacia mangium in Thailand, although Crous et al. (2004)
assigned Musa sp. to the host range of this species based on
morphology and DNA similarity.
Zasmidium
Key to Zasmidium species on Musaceae
(morphological differentiation dificult, sequence analyses more reliable)
1
Conidia rather long, 55–200 × 2.6–3.2 µm .................................................. Zasmidium sp. (see notes under Z. musae)
Conidia shorter, 6–83(–120) µm ..................................................................................................................................... 2
2 (1)
Conidia rather long and narrow, 50–83 × 2–2.5 µm;
conidiogenous loci and hila 0.5–1 µm diam ............................................................................... Z. queenslandicum
Conidia shorter, 6–65(–120) × 1.5–4.5 µm; conidiogenous loci and hila 1–1.5 µm diam ............................................... 3
3 (2)
Conidia solitary, small, (7–)27–40(–70) × 1.5–3 µm .......................................................................................... Z. musae
Conidia solitary to catenate, on average somewat longer and slightly wider,
usually (6–)35–65(–120) × 2–4 µm .......................................................................................................................... 4
4 (3)
Conidia solitary as well as catenate, at least in vivo; conidiophores solitary and in vivo
also fasciculate, arising from stromata, to 80 µm long; greasy spot disease of citrus,
rarely on banana .............................................................................................................................. Z. citri-griseum
Only known in vitro; conidia inely verruculose, genetically distinct but morphologically
barely distinguishable in vitro .................................................................................................................. Z. musicola
Zasmidium species on Musaceae
Zasmidium citri-griseum (F.E. Fisher) U. Braun &
Crous, comb. nov.
MycoBank MB809015
(Fig. 137)
Basionym: Cercospora citri-grisea F.E. Fisher, Phytopathology
51: 300 (1961).
VOLUME 5 · NO. 2
Synonyms: Stenella citri-grisea (F.E. Fisher) Sivan., Bitunicate
Ascomycetes and their Anamorphs: 226 (1984).
?Mycosphaerella citri Whiteside, Phytopathology 62: 263
(1972) [isotype: uSA: Florida: University of Florida, Citrus
Experimental Station, on Citrus paradisi, 1969, J. O.
Whiteside, no. 108 (K(M) 57614), dried culture; topotype
material: Jul. 1996 (K(M) IMI 141543)].
Zasmidium citri (Whiteside) Crous, Persoonia 23: 105 (2009).
337
Braun et al.
ART I CLE
mid rib, deep olivaceous, medium brown to dark brown or
even blackish, tar-like, margin diffuse, yellowish green to
distinct, raised, dark brown, on fruits mainly formed as
blackish specks. Colonies amphigenous. Mycelium internal
and external; internal hyphae hyaline to pale, supericial
hyphae emerging through stomata, branched, septate, 2–3
µm wide, pale olivaceous to red-brown or medium brown,
thin-walled, verruculose. Stromata lacking or small, external,
dark brown or with small stromatic hyphal aggregations in
the substomatal cavities. Conidiophores solitary, arising
from supericial hyphae, lateral, or in loose fascicles, arising
from stromatic hyphal aggregations, erect, subcylindrical,
straight to somewhat lexuous or geniculate-sinuous in the
fertile portion, unbranched, 5–80 × 2.5–6 µm, 0–6-septate,
deep olivaceous to red-brown or medium brown, thin-walled,
smooth to verruculose; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous
cells, about 5–30 µm long, sympodial, conidiogenous loci
conspicuous, lat, non-protuberant to subdenticulate, lattipped, somewhat thickened and darkened, about 1–1.5 µm
wide. Conidia solitary or catenate, in simple or occasionally
branched chains, subcylindrical to narrowly obclavatecylindrical (0–1-septate conidia sometimes short cylindrical
to ellipsoid-fusoid), straight to somewhat curved, 6–70(–120)
× 2–4.5 µm, (0–)1–6(–10)-septate, pale olivaceous to pale
brown, thin-walled, verruculose, apex obtuse or subobtuse to
truncate in catenate conidia, base obconically truncate, 1–2
µm wide, hila thickened and darkened.
[Holotype: uSA: Florida: Polk County, Babson Park, on Citrus
limon, 15 Jan. 1958, F. E. Fisher (probably not preserved)].
Neotype (designated here, MycoBank MBT178163): uSA:
Florida: Lake Alfred & Haines City, on Citrus sp., May 1970,
F. E. Fisher (K(M) IMI 148810).
Fig. 137. Zasmidium citri-griseum (K(M) IMI 148810). A. Supericial
hypha. B. Supericial hypha with solitary conidiophore. C. Conidiophore
fascicles. d. Conidiophore. e. Conidia. Bar = 10 µm.
Literature: Sivanesan & Holliday (1976), Sivanesan (1984:
226), Hsieh & Goh (1990: 295), Crous & Braun (2003: 125),
Pretorius et al. (2003), Crous et al. (2004: 464–465), Maxwell
et al. (2005), Aptroot (2006: 63), Mondal & Timmer (2006),
Arzanlou et al. (2008), Crous et al. (2009).
Illustrations: Sivanesan & Holliday (1976: ig., unnumbered),
Sivanesan (1984: 227, ig. 122), Pretorius et al. (2003: 300,
ig. 4), Crous et al. (2004: 465, ig. 10), Aptroot (2006: 12, ig.
194).
Description: On leaves, fruits and occasionally twigs, mainly
causing citrus greasy spot, leaf spots amphigenous, shape
and size variable, circular to angular-irregular, small to
large, covering large leaf segments between margin and
338
Host range and distribution: Acacia mangium, Fabaceae;
Aeglopsis chevalieri, Citrus (aurantiifolia, ×jambhiri, limon,
×microcarpa [×Citrofortunella mitis], paradisi, reticulata
[tangerina], sinensis, trifoliata [Poncirus trifoliata], Citrus
spp.), Fortunella sp., Murraya paniculata, Rutaceae; Musa
sp., Musaceae; Africa (Gabon), Asia (China [Hong Kong],
Japan, Thailand, Taiwan), Australia, North America (Mexico;
USA, Florida, New Jersey, Texas), South and Central America
(Argentina, Bolivia, Costa Rica, El Salvador, Paraguay,
Suriname, Venezuela), Oceania (Haiwaii, Tonga), West
Indies (Cuba, Dominican Republ., Haiti, Puerto Rico, Trinidad
and Tobago, Virgin Islands).
Notes: According to the current ICN, Cercospora citri-grisea
is the oldest valid name for this species and has priority.
Furthermore, it is still unclear and unproven if C. citri-grisea
and Mycosphaerella citri are actually conspeciic. This
species was regarded as a species complex by Pretorius
et al. (2003), but maintained as a single species with wider
host range in Crous et al. (2004), who proved by means of
molecular sequence analyses that Zasmidium citri-griseum
is able to infect Acacia mangii (in Thailand) and Musa sp. (in
North America, USA, Florida). Arzanlou et al. (2008) listed
Musa sp. from Tonga as host, Maxwell et al. (2005) revealed
infections of this species on Eucalyptus camaldulensis in
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Vietnam, and Crous et al. (2009) added Eucalyptus sp. from
Thailand as proven host.
The genetic connexion between Zasmidium citrigriseum and Mycosphaerella citri is unclear, not deinitively
proven and may be questioned. Aptroot (2006) examined
iso- and topotype material of M. citri and found the
isotype morphologically indistinguishable from Davidiella
ammophilae. Topotype material was re-identiied as saprobic
Mycosphaerella punctiformis. However, the Davidiella
morphology of M. citri is in contradiction with the phylogenetic
position of Z. citri-griseum, which clusters with other
Zasmidium species in Mycosphaerellaceae (Pretorius et al.
2003, Crous et al. 2004, Arzanlou et al. 2008, Crous et al.
2009) and not in Cladosporiaceae (Cladosporium/Davidiella
clade). Mycosphaerella citri is morphologically characterised
as follows: Spermogonia formed prior to ascomata, with rodshaped, colourless spermatia, 2–3.5 × 1 µm; ascomata on
decomposing leaves, amphigenous, immersed, to 90 µm
diam; asci broadly ellipsoid, 25–35 × 8–10 µm, 8-spored;
ascospores slightly fusiform-ellipsoid to narrowly obovoid,
with upper cells slightly wider, 6–12 × 2–3 µm, ± medianly
1-septate, hyaline, with minute oil droplets when fresh.
Further research on the true identity of M. citri, and allied
species in this complex is needed.
Zasmidium musae (Arzanlou & Crous) Crous & U.
Braun, Schlechtendalia 20: 102 (2010).
(Fig. 138)
Basionym: Stenella musae Arzanlou & Crous, Persoonia 20:
31 (2008).
Illustration: Arzanlou et al. (2008: 31–32, igs 13, 14a).
Description: In vitro (on MEA): Mycelium submerged and
supericial; submerged hyphae 2–3 µm wide, thin-walled,
smooth to verrucose, subhyaline to medium brown, with thin
septa; aerial hyphae 2–2.5 µm wide, coarsely verrucose,
olivaceous-brown to medium brown, rather thick-walled,
with thin septa. Conidiophores arising from supericial
hyphae, erect, straight subcylindrical to geniculatesinuous, unbranched, to 30 µm long and 2–2.5 µm wide,
0–3-septate, medium brown, wall rather thick, verruculose;
conidiogenous cells integrated, terminal, sometimes
intercalary or conidiophores reduced to conidiogenous
cells, sympodial, subcylindrical, tapering towards lat-tipped
subdenticulate apical conidiogenous loci, 1–1.5 µm diam,
thickened and darkened. Conidia solitary, subcylindrical to
obclavate, straight to somewhat curved, (7–)27–40(–70) ×
1.5–3 µm, 0–7-septate, pale brown, thin-walled, verruculose,
apex subobtuse, base short to long obconically truncate or
subtruncate, 1–1.5 µm wide, hila thickened and darkened.
Culture characteristics: Colonies on MEA reaching 30
mm after 30 d at 24 °C, erumpent, unevenly folded, with
moderate aerial mycelium and entire smooth margin, surface
pale mouse-grey to mouse-grey, in reverse dark mousegrey. Colonies on OA reaching 48 mm after 30 d at 24 °C,
effuse, with moderately developed aerial mycelium and entire
margin, pale mouse-grey to mouse-grey, in reverse dark
mouse-grey.
VOLUME 5 · NO. 2
Figs 138–140. 138. Zasmidium musae (CBS H-2047). 139. Z.
musicola (CBS H-2046), 140. Z. queenslandicum (CBS H-2050). A.
Hyphae with conidiophores. B. Conidia. Bar = 10 µm. P.W. Crous del.
Holotype: tonga: ACIAR Plot, Tongatapu, on Musa cv. TU8
AAAA, Mar. 1990, R. A. Fullerton (CBS H-20047). Ex-type
culture: CBS 122476.
Host range and distribution: On Musa sp., Musaceae,
Oceania (Tonga), West Indies (St. Lucia).
Notes: This and all other Zasmidium species on banana are
only known in vitro. All of them are morphologically similar
and only gradually differentiated, but, on the other hand,
genetically clearly distinct and form separate clusters, even
in pure ITS analyses (Arzanlou et al. 2008). There is a fourth
unnamed Zasmidium species discussed and illustrated in
Stover (1994) who coined the name “Cercospora nonvirolentum” (not formally published) which was considered
the asexual morph of Mycosphaerella musae (Speg.) Syd.
& P. Syd. and a prevalent co-inhabitant with Black Leaf
Streak and Sigatoka. However, the conspeciicity of this
Zasmidium and M. musae is uncertain and has to be proven
by new collections and epitypiications of M. musae and
other confusable Mycospharella spp. that have in the past
been isolated from Mycosphaerella speckle symptoms on
banana.
339
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Braun et al.
Zasmidium musicola (Arzanlou & Crous) Crous & U.
Braun, Schlechtendalia 20: 102 (2010).
(Fig. 139)
Basionym: Stenella musicola Arzanlou & Crous, Persoonia
20: 33 (2008).
Illustration: Arzanlou et al. (2008: 32–33, igs 14c, 16).
Description: In vitro (on MEA): Mycelium submerged and
supericial; submerged hyphae 2–3 µm wide, thin-walled,
smooth to verrucose, subhyaline to olivaceous-brown, with
thin septa; aerial hyphae 2–2.5 µm wide, coarsely verrucose,
olivaceous-brown, rather thick-walled, with thin septa.
Conidiophores arising from supericial hyphae, erect, straight,
subcylindrical to geniculate-sinuous, unbranched, (18–)
30–36(–45) × (2–)2.5–3(–4) µm, 0–2-septate, sometimes
continuous with its supporting hypha, pale brown, rather thickwalled, inely verruculose; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells,
smooth to verruculose, sometimes with swollen apex,
sympodial, conidiogenous loci truncate, lat-tipped, 1–1.5
µm diam, thickened and darkened. Conidia solitary, rarely
in short unbranched chains, subcylindrical to obclavate, (7–)
37–57(–120) × 2–4 µm, aseptate to pluriseptate, medium
brown, thin-walled, inely verruculose, apex obtuse, base
subtruncate or short to long obconically truncate, 1–1.5 µm
wide, hila thickened and darkened.
Culture characteristics: Colonies on MEA reaching 28
mm after 30 d at 24 °C, effuse, slightly raised at the centre,
with moderate, velvety to hairy aerial mycelium, folded, with
entire smooth margin, surface pale mouse-grey to mousegrey, in reverse dark mouse-grey. Colonies on OA reaching
39 mm after 30 d at 24 °C, effuse, with moderately velvety or
hairy aerial mycelium and entire margin, pale mouse-grey to
mouse-grey, in reverse olivaceous.
Holotype: Australia: Queensland: Mount Lewis, Mount
Lewis Rd., 16° 34’ 47.2’’ S, 145° 19’ 7’’ E, 538 m alt, on Musa
banksii, Musaceae, Aug. 2006, P. W. Crous, W. Gams & B.
Summerell (CBS H-20050). Ex-type culture: CBS 122475.
Host range and distribution: Only known from the type
collection.
Nartheciaceae
Cercospora
Doubtful, excluded and insuficiently known
species
Cercospora narthecii F.L. Balf.-Browne, Trans. & Proc.
Bot. Soc. Edinburgh 35: 434 (1951).
Basionym: Stenella queenslandica Arzanlou & Crous,
Persoonia 20: 34 (2008).
Synonyms: Napicladium ossifragi Rostr., Bot. Færöes 1: 316
(1901) [neotype (designated by Schubert et al. 2007:
135): denmark: Undallslund, on Narthecium ossifragum,
13 Sep. 1885, E. Rostrup (CP)].
Heterosporium magnusianum Jaap, Schriften Naturwiss.
Vereins Schleswig-Holstein 12: 346 (1902) [lectotype
(designated by Schubert et al. 2007: 135): germany:
Hamburg, Eppendorfer Moor, on Narthecium ossifragum,
12 Sep. 1887, O. Jaap (HBG)].
Heterosporium ossifragi (Rostr.) Lind, Danish Fungi: 531
(1913).
Cladosporium magnusianum (Jaap) M.B. Ellis, More
Dematiaceous Hyphomycetes: 337 (1976).
Cladosporium ossifragi (Rostr.) U. Braun & K. Schub.,
Stud. Mycol. 58: 133 (2007).
Illustration: Arzanlou et al. (2008: 32, ig. 14b, 34, ig.17).
Literature: Crous & Braun (2003: 289).
Description: In vitro (on MEA): Mycelium submerged
and supericial; submerged hyphae 2–3 µm wide, thinwalled, smooth, subhyaline to olivaceous-brown, with thin
septa; aerial hyphae 2–2.5 µm wide, coarsely verrucose,
olivaceous-brown, rather thick-walled, with thin septa.
Conidiophores arising from supericial hyphae, erect, straight
subcylindrical to geniculate-sinuous, unbranched, to 40 µm
long, and 2–3 µm wide, 0–4-septate, pale brown, thin-walled,
verruculose; conidiogenous cells integrated, terminal or
Holotype: uK: Scotland: Southerland, Inchnadamph forest,
on Narthecium ossifragum, June 1948, F. L. Balfour-Browne
(K(M) 191244).
Holotype: India: Tamila Nadu: Tiruchirapally, on Musa cv.
Grand Nain AAA (Cav.), Musaceae, 2005, I. Buddenhagen
(CBS H-20046). Ex-type culture: CBS 122479.
Host range and distribution: Only known from the type
collection.
Zasmidium queenslandicum (Arzanlou & Crous)
Crous & U. Braun, Schlechtendalia 20: 103 (2010).
(Fig. 140)
340
conidiophores reduced to conidiogenous cells, sympodial,
smooth to inely verruculose, conidiogenous loci lat-tipped,
1 µm diam, thickened and darkened-refractive. Conidia
solitary, subcylindrical to obclavate, straight to curved, 50–83
× 2–2.5 µm, aseptate to pluriseptate, medium brown, thinwalled, verruculose, apex obtuse or subobtuse, base short to
long obconically truncate or subtruncate, 0.5–1 µm wide, hila
thickened and darkened-refractive.
Culture characteristics: Colonies on MEA reaching 24 mm
after 30 d at 24 °C, effuse, slightly elevated at the centre, with
abundant aerial mycelium and entire smooth margin, surface
mouse-grey to dark mouse-grey, in reverse dark mouse-grey.
Colonies on OA reaching 41 mm after 30 d at 24 °C, effuse,
with moderately developed aerial mycelium and entire,
smooth margin, surface olivaceous-grey, in reverse iron-grey.
Notes: Cercospora narthecii was described to have long
conidiophores, 100–150 × 5–6 µm, and ellipsoid, aseptate
conidia, 12–15 × 4–5 µm, occasionally formed in chains,
hyaline to olivaceous (Balfour-Browne 1951). Type material
of this species has been examined and proved to belong to
IMA FUNGUS
Cercosporoid fungi 2
conidiophores this species is probably a synonym of
Cladosporium ossifragi, which is common on Narthecium
ossifragum.
Orchidaceae
ART I CLE
Cladosporium (conidiogenous loci and hila coronate). The
fructiication is rather immature and the conidia resemble
those of Cladosporium herbarum, but due to non-nodulose
Cercospora
Key to Cercospora species on Orchidaceae
1
Stromata lacking or small; conidiophores very short, 10–25 µm; conidia narrow,
40–100 × 2–3 µm; on Cephalanthera, Europe
......................................................... [C. cephalantherae, see Doubtful, excluded and insuficiently known species]
Stromata developed, 20–75 µm diam; conidiophores mostly much longer;
conidia broader, 3–5 µm .......................................................................................................................................... 2
2 (1)
Conidia subhyaline to brown; on Eulophia sp., India
................................................................... [C. eulophiae, see Doubtful, excluded and insuficiently known species]
Conidia hyaline ................................................................................................................................................................ 3
3 (2)
Conidiophores relatively short, 10–50 µm, 0–3-septate; on Cypripedium and Epipactis spp.,
Europe and North America .................................................................................................................. C. epipactidis
Conidiophores much longer, usually 50–285 µm, pluriseptate;
on Habenaria and Pecteilis spp. ..................................................................................................... C. habenariicola
Cercospora species on Orchidaceae
Cercospora epipactidis C. Massal., Ann. Mycol. 9:
256 (1911).
(Fig. 141)
Literature: Saccardo (1913: 1430), Chupp (1954: 425), Braun
& Mel'nik (1997: 58), Crous & Braun (2003: 175).
Exsiccatae: Syd., Mycoth. Germ. 2035.
Description: Leaf spots amphigenous, oblong or streak-like,
between veins, dark purplish violet to reddish dark brown,
5–40 × 1–2 mm or conluent and larger, margin indeinite.
Caespituli hypophyllous, punctiform to minutely pustulate,
dark brown to blackish. Mycelium internal. Stromata
substomatal to immersed, 20–60 µm diam, subglobose to
irregular, dark brown. Conidiophores in small to moderately
large fascicles, loose to dense, pale to medium brown, paler
towards the tip, arising from stromata, through stomata or
erumpent, erect, straight to curved-lexuous, geniculatesinuous, unbranched, 10–50 × 3–6 µm, 0–3-septate, thinwalled, smooth; conidiogenous cells integrated, terminal
or conidiophores reduced to conidiogenous cells, 10–25
µm long, conidiogenous loci conspicuous, thickened and
darkened, about 2–2.5 µm diam. Conidia solitary, cylindrical to
obclavate-subcylindrical, occasionally subacicular, 25–120(–
150) × 3–5 µm, 3–10-septate, hyaline, thin-walled, smooth,
Fig. 141. Cercospora epipactidis (LEP, Russia, Tersk, 2 Jul. 1914,
Woronichin). A. Conidiophore fascicle. B. Conidiophores. C. Conidia.
Bar = 10 µm.
VOLUME 5 · NO. 2
341
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Braun et al.
apex obtuse to subacute, base truncate to short obconically
truncate, 1.5–3 µm wide, hila thickened and darkened.
Holotype: Italy: S. Michele near Verona, on Epipactis
palustris, Aug. 1910, C. Massalongo (VER).
Host range and distribution: On Cypripedium guttatum,
Epipactis (atrorubens [rubiginosa], helleborine [latifolia],
palustris), Orchidaceae, Asia (Russia, West Siberia), Europe
(Czech Republic, Italy, Germany, Romania, Russia), North
America (USA, Florida).
Notes: A true Cercospora s. str. distinct from C. apii s. lat. by
having obclavate-cylindrical conidia. A single Asian collection
of this species from West Siberia has been examined
(Novosibirsk, Tersk, on Epipactis latifolia, 2 Jul. 1914,
Woronichin, LEP).
Cercospora habenariicola Meeboon, Hidayat & C.
Nakash., Mycotaxon 99: 118 (2007).
(Fig. 142)
Literature: To-anun et al. (2011: 72), Patil et al. (2012).
Illustrations: Meeboon et al. (2007: 119, igs 1–2), To-anun et
al. (2011: 72, ig. 55), Patil et al. (2012; 113, igs 1–6).
Description: Leaf spots amphigenous, circular, subcircular to
somewhat angular-irregular, about 3–30 mm diam, at irst pale
greenish to ochraceous, later brown to dark brown, inally with
greyish brown centre and darker margin, surrounded by a
brownish halo, sometimes zonate. Caespituli amphigenous,
mainly hypophyllous, punctiform, ochre-yellow, velvety. Mycelium
internal. Stromata well-developed, substomatal to intraepidermal,
25–75 µm diam, subglobose, brown to blackish brown.
Conidiophores numerous, in loose to dense fascicles, arising
from stromata, through stomata or erumpent, erect to decumbent,
straight, subcylindrical, geniculate-sinuous, unbranched or
rarely branched, 50–285 × 3–7.5 µm, rarely to 950 µm long,
2–9-septate, pale yellow to pale brown, wall thin or slightly
thickened, smooth; conidiogenous cells integrated, terminal,
sympodial, conidiogenous loci conspicuous, about 2.5–3.5 µm
diam, thickened and darkened. Conidia solitary, subacicular
to narrowly obclavate, 35–110 × 3–5 µm, mosty 3–10-septate,
hyaline, thin-walled, smooth, apex pointed, base obconically
truncate, 1.2–3 µm wide, hila thickened and darkened.
Holotype: thailand: Chiang Mai Province: Queen Sirikit
Botanic Garden, on Habenaria susannae, 14 Jul. 2006, J.
Meeboon, JM 155 (CMU 27883).
Host range and distribution: On Habenaria (heyneana,
longicorniculata, roxburghii), Pecteilis susannae [Habenaria
susannae],
Orchidaceae,
Asia
(India,
Karnataka,
Maharashtra; Thailand).
Notes: Cercospora habenariicola is characterised by having
obclavate conidia, about 3–5 µm wide, with obconically
truncate bases. It is not quite clear if Indian collections on all
hosts recorded by Patil et al. (2012) are truly conspeciic with
342
Fig. 142. Cercospora habenariicola (CMU 27883). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
C. habenariicola. Indian specimens on Habenaria furcifera
[ovalifolia], Peristylus densus, P. goodyeroides as well as
Pecteilis gigantea are described as having much narrower
conidia, only 2–2.5 µm wide, which are, according to Patil et
al. (2012; 113, ig. 6) rather acicular as far as longer conidia
are concerned. It is possible that two different species are
involved, viz. C. habenariicola and a Cercospora belonging
the the C. apii s. lat. complex.
Doubtful, excluded and insuficiently known
species
Cercospora cephalantherae Ondrěj & Zavřel, Čas.
Slez. Mus. Opavě, ser. A., Hist. Nat., 20: 22 (1971);
as "cephalenterae".
Illustration: Ondrěj & Zavřel (1971: 23, ig. 2).
IMA FUNGUS
Cercosporoid fungi 2
Holotype: Czech republic: Moravia: Štramberk, Kotouč, on
Cephalanthera damasonium [alba], Orchidaceae, 20 Aug.
1950, H. Zavřel (not preserved).
Host range and distribution: Only known from the type
collection.
Literature: Crous & Braun (2003: 178), Kamal (2010: 43).
Illustration: Patil (1978: 328, ig. 3).
Description: Leaf spots amphigenous, circular to angular, 3–5
mm diam, brown with pale centre. Caespituli hypophyllous,
olivaceous-brown. Mycelium internal. Stromata 30–70 µm
diam, globose, brown. Conidiophores in small to moderately
large fascicles, 5–12, arising from stromata, erect, straight
to lexuous, non-geniculate, unbranched, 50–195 × 5
µm, pluriseptate, olivaceous-brown, paler towards the tip;
conidiogenous loci distinct, prominent. Conidia solitary,
acicular, obclavate, straight to curved, 75–250 × 3.2–4 µm,
3–5-septate, subhyaline to brown, apex acute or subobtuse,
base truncate to obconically truncate.
Notes: Ondrěj & Zavřel (1971) mentioned that the type of this
species is deposited at BRA, but according to the curator
of this herbarium type material could not be traced. Based
on the original description and illustration, this species is
probably a true Cercospora s. str.
Holotype: India: Maharashtra: Kolhapur, on Eulophia sp.,
Orchidaceae, 18 Aug. 1977, M. S. Patil (HCIO 32545).
Cercospora epidendronis Bolick, nom. nud.
Notes: Type material of this species was not available for a
re-examination. Details of the conidiogenous loci and conidial
hila are not given in the original description and subhyaline
to brown conidia are not in favour of Cercospora s. str., i.e.
it cannot be excluded that C. eulophiae belongs to another
cercosporoid genus, e.g. Pseudocercospora. A revision of
the type material is necessary.
Notes: According to Alieri et al. (1984) and Farr et al. (1989:
609) [on Epidendrum sp., USA, Florida].
Cercospora eulophiae M.S. Patil, Indian Phytopathol.
31: 328 “1978” (1979).
ART I CLE
Description: Leaf spots amphigenous, brown or black, veinlimited, conluent. Mycelium internal. Stromata probably not
developed. Conidiophores fasciculate, straight, curved or
lexuous, short, unbranched, 10–25 × 3.5 µm, pale brown,
septate at the base. Conidia solitary, obclavate-cylindrical,
iliform, 40–80(–100) × 2–3 µm, hyaline, 4–8-septate, apex
subacute or subobtuse, base almost truncate to short
obconically truncate.
Host range and distribution: Only known from the type
collection.
Pseudocercospora
Key to Pseudocercospora species on Orchidaceae
1
Mycelium in vivo internal and external; supericial hyphae with solitary conidiophores developed ................................ 2
Mycelium in vivo internal; supericial hyphae with solitary conidiophores lacking ........................................................... 4
2 (1)
Stromata large, 10–100 µm diam; conidia narrow, 1.5–3 µm; on Cymbidium, New Zealand .................. p. cymbidiicola
Stromata smaller, 10–50 µm diam; conidia broader, 2–5 µm; on other hosts ................................................................. 3
3 (2)
On Dendrobium spp. ..................................................................................................................................... p. dendrobii
On Peristeria spp. ....................................................................................................................................... p. peristeriae
4 (1)
Stromata lacking or small, 10–25 µm diam; conidiophores loosely fasciculate and long,
50–250 × 3–6 µm, pluriseptate; on hosts of various genera ............................................................ p. odontoglossi
Stromata always developed, 10–200 µm diam; conidiophores much shorter,
10–60 µm long, 0–3-septate .................................................................................................................................... 5
5 (4)
Conidia narrow, 10–85 × 1–3 µm; on Epidendrum, Brazil ............................................................................. p. epidendri
Conidia broader, 2.5–5 µm .............................................................................................................................................. 6
6 (5)
Conidia subcylindrical to attenuated towards the apex, broadly acicular, base truncate,
2.5–4 µm wide; on Lycaste sp., Colombia .................................................................................... p. orchidacearum
Conidia obclavate-cylindrical, base usually obconically truncate, (1–)1.5–2.5(–3) µm wide .......................................... 7
7 (6)
Stromata small, 10–40 µm diam; conidiophores 5–15 per fascicle; conidiogenous loci
paracercosporoid, i.e. ranging from inconspicuous to subconspicuous by being
slighty thickened and darkened around the ultimate rim (ring-like in front view);
on Cypripedium spp., North America .................................................................................................... p. cypripedii
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Braun et al.
Stromata larger, 20–90 µm diam; conidiophores usually numerous in dense fascicles;
conidiogenous loci inconspicuous, unthickened, not darkened, not paracercosporoid;
on Cattleya, Jumenella, Odontoglossum and Oncidium spp. .................................................................. p. angraeci
Pseudocercospora species on Orchidaceae
pseudocercospora angraeci (Feuilleb. & Roum.) U.
Braun & Urtiaga, Mycosphere 4: 596 (2013)
(Fig. 143)
Basionym: Cercospora angraeci Feuilleb. & Roum., Rev.
Mycol. 5: 177 (1883); as “angreci”.
Literature: Saccardo (1886: 478), Chupp (1954: 424), Crous
& Braun (2003: 58).
Illustration: Braun & Urtiaga (2013b: 597, ig. 3).
Exsiccatae: Roum., Fungi Sel. Exs. 2522.
Description: Leaf spots small to large, 1–25 mm diam or
conluent and larger, subcircular to somewhat irregular,
dingy grey to blackish, mainly due to abundant colonies,
margin indistinct or darker. Caespituli amphigenous,
punctiform, scattered to gregarious, dark brown to blackish.
Mycelium internal. Stromata substomatal to intraepidermal
or deeply immersed, large, 20–100 µm diam, medium to
dark olivaceous-brown or brown. Conidiophores in dense
fascicles, usually numerous, arising from stromata, erumpent,
straight to somewhat sinuous, but not or barely geniculate,
unbranched, subcylindrical to somewhat attenuated towards
the tip, apex usually obtuse, (5–)10–40(–60) × 2.5–5 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 rarely percurrent, conidiogenous
loci inconspicuous or visible as truncate tip, 1.5–3 µm
wide, but always unthickened and not darkened. Conidia
solitary, obclavate-cylindrical, occasionally subacicular,
straight to somewhat curved, 20–85(–100) × 2.5–5 µm,
(0–)2–8-septate, pale olivaceous to olivaceous-brown, thinwalled, smooth, apex obtuse to subacute, base truncate to
mostly short obconically truncate, (1–)1.5–2.5(–3) µm wide,
hila unthickened, not darkened.
Syntypes: réunion: on Jumenella fragrans [Angraecum
fragrans], Oct. 1882 [Roum., Fungi Sel. Exs. 2522] (B, LEP,
PC).
Host range and distribution: On Cattleya (mossiae, Cattleya
sp.), Jumenella fragrans, Laelia sp., Oncidium (alexandrae
[Odontoglossum crispum], Oncidium sp.), Sobralia
(xantholeuca, Sobralia sp.), Orchidaceae, Africa (Réunion
[Island of Bourbon]), Europe (France; UK, England), North
America (Mexico; USA, Florida), South America (Guatemala,
Venezuela).
Notes: All syntype collections examined were in poor
condition, and not suitable to serve as a lectotype. Braun
& Urtiaga (2013b) examined a collection from Venezuela,
344
Fig. 143. Pseudocercospora angraeci (K(M) IMI 180170). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
in better agreement with the description, on which the
reallocation to Pseudocercospora, and the present description
and illustration are based. Furthermore, some collections on
Sobralia spp. desposited at BPI have been examined and
proved to represent P. angraeci.
pseudocercospora cymbidiicola U. Braun & C.F.
Hill, Mycol. Progr. 1: 23 (2002).
(Fig. 144)
Literature: Nakashima et al. (2006), Han et al. (2007).
Illustrations: Braun & Hill (2002: 24, ig. 5), Han et al. (2007:
127–128, igs 1–2).
Description: Leaf spots lacking or amphigenous, subcircular
to irregular, later effuse, large, 5–20 mm diam or even larger,
covering large leaf segments or almost entire leaves, dingy
greyish brown to dark brown, margin indeinite. Caespituli
amphigenous, variable, punctiform to subeffuse, greyish to
dark brown or even blackish. Mycelium internal and external;
supericial hyphae sparingly branched, septate, 1.5–3 µm
wide, pale olivaceous, thin-walled, smooth. Stromata lacking,
small to well-developed, 10–100 µm diam, substomatal to
intraepidermal, brown. Conidiophores in small, loose to large,
dense fascicles, arising from internal hyphae or stromata,
often with numerous short conidiophores arising from
IMA FUNGUS
Cercosporoid fungi 2
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thin-walled, smooth; conidiogenous cells integrated, terminal
or conidiophores reduced to conidiogenous cells, 5–30 µm
long, conidiogenous loci inconspicuous, neither thickened nor
darkened. Conidia solitary, acicular to cylindrical-iliform, 50–
110 × (1.5–)2–2.5(–3) µm, indistinctly 3–12-septate, distance
between septa 5–15 µm, subhyaline to pale olivaceous, thinwalled, smooth, apex subacute, base truncate, 1.5–2 µm
wide, hila neither thickened nor darkened.
Holotype: New Zealand: Auckland, Mt Albert, on Cymbidium
sp., 24 Sep. 2000, C. F. Hill 303 (HAL 1585 F). Isotype:
PDD 71965. Paratype: New Zealand: Auckland, Mt Eden,
on Cymbidium sp., 19 Apr. 1979, F. J. Newhook (K(M) IMI
239798), deposited as Pseudocercosporella sp.
Host range and distribution: On Cymbidium sp., Orchidaceae,
Asia (Japan, Korea), New Zealand.
pseudocercospora cypripedii (Ellis & Dearn.) U.
Braun & Crous, Mycosphaerella and Anam.: 152
(2003).
(Fig. 145)
Basionym: Cercospora cypripedii Ellis & Dearn., Canad. Inst.
Trans. 6: 637 (1899).
Literature: Saccardo (1902: 1073), Chupp (1954: 425).
Exsiccatae: Rogerson et al., Fungi Bor.-Amer. Exs. 317.
Fig. 144. Pseudocercospora cymbidiicola (HAL 1585 F). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
large stromata, forming sporodochial conidiomata, solitary
conidiophores arising from supericial hyphae also formed,
solitary, lateral, conidiophores erect, long conidiophores
sometimes decumbent (differentiation between decumbent
conidiophores and supericial hyphae often dificult),
subcylindrical-iliform to somewhat geniculate-sinuous,
unbranched, 5–100 × (1.5–)2–5(–5.5) µm, aseptate to
pluriseptate throughout, pale olivaceous to olivaceous-brown,
VOLUME 5 · NO. 2
Description: Leaf spots oblong, spread between veins, 1–3
mm wide, dark brown to blackish, sometimes in long streaks.
Caespituli amphigenous, often epiphyllous, punctiform,
dark. Mycelium internal. Stromata substomatal to immersed,
10–40 µm diam, dark brown, composed of brown swollen
hyphal cells, 2–5 µm wide. Conidiophores in small to
moderately large fascicles, mostly 5–15, loose to mostly
moderately dense, arising from stromata, through stomata or
erumpent, erect, straight to curved or somewhat geniculatesinuous, unbranched, 10–40 × 3–5 µm, 0–2-septate, pale
to medium dark brown, wall thin, smooth; conidiogenous
cells integrated, terminal or conidiophores reduced to
conidiogenous cells, 10–25 µm long, conidiogenous loci
inconspicuous to subconspicuous by being paracercosporalike, i.e. with minute thickened and darkened outer rim, visible
in front view as minute circle, 1–2 µm diam. Conidia solitary,
obclavate to obclavate-cylindrical, straight to curved, 30–150
× 3–5 µm, 3–8-septate, pale olivaceous, thin-walled, smooth,
apex acute to subobtuse, base subtruncate or short to long
obconically truncate, 1.5–2 µm wide, hila unthickened, not
darkened or at most somewhat refractive or ultimate rim
slightly darker.
Lectotype (designated here, MycoBank MBT178164):
Canada: Ontario: shore of Lake Huron near Southampton
and Komoka, on Cypripedium sp., 20 Aug. 1898, J. Dearness
2883 (NY 838300). Isolectotype: DAOM.
Host range and distribution: On Cypripedium (acaule,
calceolus, candidum, parvilorum var. pubescens [pubescens],
reginae [spectabile], Cypripedium sp.), Orchidaceae, North
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Fig. 146. Pseudocercospora dendrobii (CUP 41041). A. Solitary
conidiophores arising from supericial hyphae. B. Supericial hyphae
emerging through a stoma. C. Conidiophore fascicle. d. Conidia. Bar
= 10 µm.
Fig. 145. Cercospora cypripedii (NY 838300). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
America (Canada; USA, Florida, Iowa, Massachusetts, New
York, Wisconsin).
Notes: Based on conidiogenous loci ranging from being
inconspicuous to subconspicuous, i.e., ultimate rim
slightly thickened and darkened, this species seems to be
intermediate between Cercospora s. str. and Passalora.
However, the loci are rather Paracercospora-like and the
conidia are subhyaline to very pale olivaceous, so that this
species is better placed in Pseudocercospora.
pseudocercospora dendrobii Goh & W.H. Hsieh, in
Hsieh & Goh, Cercospora and Similar Fungi from
Taiwan: 255 (1990).
(Fig. 146)
Synomyms: Cercospora dendrobii H.C. Burnett, Proc. Florida
State Hort. Soc. 77: 465 “1964” (1965) [holotype: uSA:
Florida: Winter Haven, on Dendrobium sp., 17 Jan. 1962,
H. C. Burnett (CUP 41041)].
346
Pseudocercospora dendrobii (H.C. Burnett) U. Braun &
Crous, in Crous & Braun, Mycosphaerella and Anam: 452
(2003), nom. illeg. (ICN, Art. 53.1).
Literature: Guo & Hsieh (1995: 239), Guo et al. (1998: 250),
Braun & Crous (2005: 408; 2007: 61).
Illustrations: Guo & Hsieh (1995: 240, ig. 202), Hsieh & Goh
(1990: 256, ig. 196), Guo et al. (1998: 250, ig. 207), Braun &
Crous (2007: 57, ig. 4).
Description: Leaf spots amphigenous, at irst small, later
forming large patches, 2–30 mm diam or conluent and
larger, covering large leaf segments or almost entire leaves.
Caespituli hypophyllous, punctiform, dense, greyish brown
to medium dark brown. Mycelium internal and external;
supericial hyphae branched, septate, 1.5–4.5 µm wide, pale
olivaceous to medium olivaceous-brown, thin-walled, smooth.
Stromata 10–40 µm diam, substomatal, olivaceous-brown.
Conidiophores in small to moderately large fascicles, loose to
dense, arising from stromata or solitary, arising from supericial
IMA FUNGUS
Cercosporoid fungi 2
Illustration: Silva & Pereira (2011: 96–99, plates 1–6).
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pseudocercospora epidendri Meiriele Silva & O.L.
Pereira, Mycotaxon 11: 898 (2011).
(Fig. 147)
Description: Leaf spots amphigenous, distinct, scattered,
irregular, 3–12 mm diam, pale brown, margin black. Caespituli
hypophyllous, punctiform, brown. Mycelium internal.
Stromata well-developed, immersed, later erumpent, 47.5–
166 µm wide and 57–213 µm high, brown. Conidiophores in
loose to dense fascicles, arising from stromata, erect, straight
to curved, subcylindrical to geniculate-sinuous, unbranched,
10–57 × 2–6 µm, 0–2-septate, brown, paler towards the tip,
thin-walled, smooth; conidiogenous cells integrated, terminal,
pale brown or conidiophores reduced to conidiogenous
cells, conidiogenous loci inconspicuous to subdenticulate,
but always unthickened, not darkened. Conidia solitary,
obclavate-subcylindrical to subacicular, straight to curved,
10–85 × 1–3 µm, 1–9-septate, pale brown, thin-walled,
smooth, apex pointed or subobtuse, base rounded or short
to long obconically truncate, about 1–1.5 µm wide, hila
unthickened, not darkened.
Holotype: Brazil: Minas Gerais: Araponga, Parque Estadual
da Serra do Brigadeiro, on Epidendrum secundum,
Orchidaceae, 8 Jan. 2008, O. L. Pereira (VIC 30553). Extype culture: OLP 30553.
Host range and distribution: Only known from the type
collection.
pseudocercospora odontoglossi (Prill. & Delacr.) U.
Braun, Mycol. Progr. 1: 23 (2002).
(Fig. 148)
Basionym: Cercospora odontoglossi Prill. & Delacr., Bull.
Soc. Mycol. France. 9: 271 (1893).
Fig. 147. Pseudocercospora epidendri (VIC 30553). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
hyphae, lateral, rarely terminal, erect to decumbent, straight,
subcylindrical-conical to geniculate-sinuous, unbranched
or occasionally branched, 10–100 × 2–5 µm, continuous
to pluriseptate, olivaceous to olivaceous-brown, thinwalled, smooth; conidiogenous cells integrated, terminal or
conidiophores reduced to conidiogenous cells, 10–30 µm
long, conidiogenous loci inconspicuous to subdenticulate, but
always unthickened, not darkened. Conidia solitary, narrowly
obclavate to subacicular, 15–80 × 2–4.5 µm, 3–8-septate, pale
olivaceous to olivaceous-brown, thin-walled, smooth, apex
subacute, base truncate to obconically truncate, (1–)1.5–2.5(–
3) µm wide, hila neither thickened nor darkened.
Holotype: taiwan: Taichung Hsien, Tali, on Dendrobium sp.,
27 Nov. 1986, T. K. Goh (NCHUPP-233).
Host range and distribution: On Dendrobium spp.,
Orchidaceae, Asia (Japan, Taiwan), North America (USA,
Florida), Haiwaii.
VOLUME 5 · NO. 2
Literature: Saccardo (1895: 629), Lindau (1910: 91), Chupp
(1954: 426), Ellis (1976: 278), Crous & Braun (2003: 296).
Illustration: Ellis (1976: 277, ig. 211 A).
Description: Leaf spots lacking or indistinct, forming yellowish
to brownish discolorations with indistinct margin, turning to
dark olivaceous patches by abundant fructiication. Caespituli
hypophyllous, effuse, thin, velvety, dark olivaceous. Mycelium
internal. Stromata lacking or only with small, substomatal
stromatic hyphal aggregations, 10–25 µm diam, brown.
Conidiophores solitary or in small, divergent fascicles,
2–7, arising from internal hyphae or hyphal aggregations,
emerging through stomata, erect, occasionally decumbent,
usually distinctly to strongly geniculate-sinuous, with
constrictions, unbranched or branched, long, 50–250 × 3–6
µm, pluriseptate throughout, pale to medium dark brown, wall
somewhat thickened, smooth; conidiogenous cells integrated,
terminal and intercalary, 10–40 µm long, conidiogenous loci
inconspicuous to mostly conspicuous by being distinctly
subdenticulate to denticulate, but wall of the loci always
unthickened and not darkened. Conidia solitary, obclavate,
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Fig. 148. Pseudocercospora odontoglossi (K(M) IMI 95083). A.
Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
Fig. 149. Pseudocercospora orchidacearum (BPI 435447). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
cylindrical-obclavate, subacicular, short conidia sometimes
fusiform, 30–105 × 3–5 µm, 3–8-septate, pale olivaceous to
olivaceous-brown, thin-walled, smooth, apex obtuse to acute,
base truncate to obconically truncate, 1.5–3 µm wide, hila
unthickened, not darkened.
[Type: France: Versailles, greenhouse, on Oncidium
alexandrae [Odontoglossum crispum], Ed. André (not
preserved)]. Neotype (designated here, MycoBank
MBT178165): New Zealand: Auckland, Mt Eden, on
Odontoglossum sp., Apr. 1954, F. J. Newhook (K(M) IMI
95083).
Host range and distribution: Brassia sp., Cattleya sp.,
Cymbidium sp., Dendrobium sp., Epidendrum sp., Laelia sp.,
Laeliocattleya sp., Odontoglossum sp., Oncidium alexandrae,
348
Orchidaceae, Europe (France, Great Britain), New Zealand,
North America (Mexico; USA, Florida, Massachussetts,
Texas).
pseudocercospora orchidacearum U. Braun, sp.
nov.
MycoBank MB809024
(Fig. 149)
Diagnosis: Distinguished from Pseudocercospora angraeci
and P. cypripedii by its cylindrical to broadly acicular conidia
with truncate base, 2.5–4 µm wide, and from P. cypripedii
in addition in having inconspicuous, non-paracercosporoid
conidiogenous loci.
IMA FUNGUS
Cercosporoid fungi 2
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Description: Leaf spots amphigenous, variable, formed as
oblong discolorations between veins, to 20 × 5 mm or even
longer, pale to darker by abundant colonies. Caespituli
amphigenous, punctiform, scattered, dark brown to blackish.
Mycelium internal. Stromata substomatal to intraepidermal,
immersed, 10–60 µm diam, medium brown, composed of
swollen hyphal cells, circular to somewhat irregular in outline,
2.5–6 µm diam, wall somewhat thickened. Conidiophores in
small to moderately large, loose to dense fascicles, arising
from stromata, emerging through stomata or erumpent, erect,
straight to curved, subcylindrical or somewhat attenuated
towards the tip, not or barely geniculate, unbranched, apex
rounded to truncate, 10–55 × 3–5 µm, with attached conidia
occasionally to 70 µm long, 0–2-septate, pale olivaceous
to medium olivaceous-brown, thin-walled, smooth;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, 10–25 µm long, mostly
unilocal, conidiogenous loci inconspicuous or visible as
truncate apex, 2–4 µm wide, but always unthickened and
not darkened. Conidia solitary, subcylindrical to attenuated
towards the apex, i.e. broadly acicular, 25–65 × 3–4.5
µm, (2–)3–6(–7)-septate, subhyaline to pale olivaceous
to olivaceous-brown, thin-walled, smooth, apex obtuse
to somewhat pointed, base truncate, 2.5–4 µm wide, hila
unthickened, not darkened.
Holotype: Colombia: intercepted at Miami, Florida, USA, on
Lycaste sp., Orchidaceae, 3 Dec. 1951, A. S. Mills, 2275 (BPI
435447).
Host range and distribution: Only known from the type
collection.
Notes: This species resembles Pseudocercospora
cypripedii and P. angraeci, but differs in having cylindrical
to broadly acicular conidia with broad truncate base, 2.5–
4 µm. Furthermore, P. cypripedii has paracercosporoid
conidiogenous loci.
pseudocercospora peristeriae (H.C. Burnett) U.
Braun & Crous, Mycosphaerella and Anam.: 316
(2003).
(Fig. 150)
Basionym: Cercospora peristeriae H.C. Burnett, Proc. Florida
State Hort. Soc. 77: 465 “1964” (1965).
Synonym: Cercospora peristeriae H.C. Burnett, Bull. State Pl.
Board Florida 12: 16 (1958), nom. inval. (ICN, Art. 39.1).
Fig. 150. Pseudocercospora peristeriae (CUP 41003). A. Supericial
hyphae. B. Supericial hyphae with solitary conidiophores. C.
Conidiophore fascicles. d. Conidiophores. e. Conidia. Bar = 10 µm.
Description: Leaf spots amphigenous, large, oblong to
irregular, 5–50 mm diam, brown, margin indeinite. Caespituli
hypophyllous, punctiform to conluent and dense or subeffuse,
dingy greyish brown to medium brown. Mycelium internal
and external; supericial hyphae branched, occasionally
anastomosing, septate, subhyaline to pale olivaceous, thinwalled, smooth, 1.5–4.5 µm wide. Stromata substomatal, 10–
50 µm diam, immersed to erumpent, brown. Conidiophores in
small to moderately large fascicles, loose to moderately dense,
arising from stromata, through stomata or solitary, arising
from supericial hyphae, lateral or occasionally terminal,
erect to decumbent, subcylindrical, conical, straight, curved,
lexuous to geniculate-sinuous, simple or branched, with
swellings and constrictions, 10–60 × 2.5–6 µm, 0–3-septate,
subhyaline, pale olivaceous to olivaceous-brown, thinwalled, smooth; conidiogenous cells integrated, terminal or
conidiophores reduced to conidiogenous cells, 10–25 µm
long, conidiogenous loci inconspicuous, unthickened, not
darkened. Conidia solitary, narrowly obclavate to obclavatecylindrical, rarely subacicular, 40–100 × 2–5 µm, 3–8-septate,
subhyaline to pale olivaceous, apex subacute to subobtuse,
base subtruncate to short obconically truncate, 1–2 µm wide,
hila unthickened, not darkened.
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Braun et al.
[Type: USA: Florida: on Peristeria elata (CUP ‘3580’), not
traced]. Neotype (designated here, MycoBank MBT178166):
uSA: Florida: Gainesville, on Peristeria elata, Apr. 1957,
H. C. Burnett (CUP 41003).
Host range and distribution: On Peristeria elata and an
unnamed orchid, Orchidaceae, North America (USA, Florida).
Notes: Burnett (1965) cited “CUP 3580” without any other
details of this collection as type. This collection could not
be traced at CUP, but another authentic sample marked as
“Cercospora sp. nov.” (on Peristeria elata, USA, Florida,
Gainesville, Apr. 1957, H.C. Burnett, CUP 41003) was
found, which is designated as neotype. Based on the
examination of this collection, it could be demonstrated that
this fungus belongs in Pseudocercospora (conidiogenous
loci inconspicuous). Some supericial hyphae with solitary
conidiophores, not described by Burnett (1965), were also
observed. Pseudocercospora peristeriae and P. dendrobii
are morphologically very similar.
Zasmidium
Key to Zasmidium species on Orchidaceae
1
Conidiophores consistently solitary, arising from supericial hyphae, 3–80 × 1.5–5 µm,
fasciculate conidiophores not developed; conidia 4–140 × 1–4 µm; on Vanda, Asia .................. Z. orchidacearum
Conidiophores solitary as well as fasciculate, longer and broader, 60–230 × 5–7 µm;
conidia shorter and broader, 17–55 × 5–6 µm; on Cyrtopodium, South America ................................. Z. cyrtopodii
Zasmidium species on Orchidaceae
Zasmidium cyrtopodii (Dorn.-Silva, Pereira-Cavalho
& Dianese)) U. Braun & Dianese, comb. nov.
MycoBank MB809029
(Fig. 151)
Basionym: Stenella cyrtopodii Dorn.-Silva, Pereira-Cavalho
& Dianese, Mycologia 99: 756 (2008).
Illustration: Dornelo-Silva et al. (2008: 758, igs 22–30).
Description: Leaf spots amphigenous, elliptical, 5–25 ×
3–10 mm, sometimes conluent, dark brown. Colonies
amphigenous, effuse, velutinous, downy, olivaceous-brown
to brown. Mycelium internal and external; internal hyphae
intercellular, branched, septate, 5–7 µm wide, sometimes
monilioid, dark brown; supericial hyphae branched, septate,
3–4 µm wide, light brown to brown, verruculose. Stromata welldeveloped, subcuticular, immersed to somewhat erumpent,
about 25–50 µm diam, textura angularis, cells 3–9 µm diam,
brown. Conidiophores fasciculate, arising from stromata,
60–230 × 5–7 µm, or solitary, arising from supericial hyphae,
lateral or terminal, 9–28 µm long, conidiophores erect,
almost straight to somewhat curved, geniculate-sinuous,
unbranched, pigmented, wall smooth; conidiogenous cells
integrated, terminal and intercalary, with 1–8 conspicuous
conidiogenous loci, thickened and darkened. Conidia solitary
or in short chains, cylindrical or subcylindrical, straight, 17–55
× 5–6 µm, 2–7-septate, pigmented, verruculose, apex obtuse,
rounded, base short obconically truncate, hila thickened and
darkened.
Holotype: Brazil: Distrito Federal: Planaltina, Estação
Ecológia de Águas Emendadas, on Cyrtopodium eugenii, 23
Jan. 1998, D. Dornelo-Silva 9 (UB, Mycol. Coll.15854).
Host range and distribution: On Cyrtopodium eugenii,
Orchidacearum, South America (Brazil).
350
Fig. 151. Zasmidium cyrtopodii (UB, Myc. Coll. 15854). A. Supericial
hypha. B. Conidiophore fascicle. C. Conidiophores. d. Conidia. Bar
= 10 µm.
IMA FUNGUS
Cercosporoid fungi 2
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Fig. 152. Zasmidium orchidacearum (HAL 1595 F). A. Supericial hyphae. B. Conidiophores. C. Conidia. Bar = 10 µm.
Zasmidium orchidacearum (U. Braun & Sivap.) U.
Braun, Schlechtendalia 20: 102 (2010).
(Fig. 152)
Basionym: Stenella orchidacearum U. Braun & Sivap., Fungal
Diversity 3: 23 (1999).
Illustration: Braun & Sivapalan (1999: 25, ig. 15).
Description: Leaf spots lacking, diffuse or subcircular to
irregular and large, to 4 × 1.5 cm, dingy greyish brown,
margin indeinite. Colonies hypophyllous, effuse, thin and
inconspicuous to velvety, dull greyish brown. Mycelium
internal and external; supericial hyphae branched, septate,
1–3.5 µm wide, subhyaline, pale yellowish, olivaceous-brown
or pale brown, thin-walled, verruculose. Stromata lacking to
well-developed, substomatal or intraepidermal, 10–50 µm
diam, brown. Conidiophores solitary, arising from supericial
hyphae, lateral and occasionally terminal, rarely in small,
loose fascicles, arising from stromata or erumpent through
the cuticle, erect, straight, unbranched, subcylindrical to
strongly geniculate-sinuous, nodulose, 3–80 × 1.5–5 µm,
0–5-septate, pale olivaceous, olivaceous-brown to medium
brown, thin-walled, smooth or almost so; conidiogenous
cells integrated, terminal or intercalary, 3–30 × 1.5–4 µm,
proliferation mostly sympodial, occasionally percurrent;
conidiogenous loci often aggregated, minute, 0.5–1.5 µm
diam, planate, somewhat thickened and darkened. Conidia
solitary, cylindrical-iliform, occasionally narrowly obclavate,
small conidia fusiform or narrowly ellipsoid-ovoid, 4–140
× 1–4 µm, 0–10-septate, subhyaline, pale olivaceous or
olivaceous-brown, thin-walled, verruculose, apex obtuse to
subacute, base truncate or somewhat obconically truncate,
0.5–3 µm wide, hila slighty thickened and darkened.
Holotype: Malaysia: Sabah: Jesselton, on Vanda sp., 25 Feb.
1966, Tay Eng Bok, PP 630/60 (K(M) IMI 119139). Paratypes:
Brunei: Tutong, on leaves of an unidentiied orchid, 26 Aug.
1996, H. Fuziah 7733 (HAL 1595 F, Hb. Brunei Agricultural
Reseach Centre, Kilanas, Brunei).
Host range and distribution: Only known from the type
collections.
Notes: Stenella sp. on Vanda sp. was reported by Peregrine &
Ahmad (1982). This material could not be traced, but another
collection on Vanda sp. from Sabah has been examined.
Pontederiaceae
Cercospora
Key to Cercospora species on Pontederiaceae
1
Conidiophores uniformly short and aseptate, 5–25 × 1.5–4 µm, hyaline to only faintly pigmented,
pale yellowish to very pale brownish; conidia acicular to narrowly subcylindrical,
short and narrow, 15–45 × 1–2.5 µm, only with 1–2(–3) septa; on Pontederia cordata ................... C. pontederiae
Conidiophores much longer and pluriseptate, to 375 µm; conidia also much longer,
wider and pluriseptate, 25–350 × 2–5 µm; on Eichhornia crassipes ....................................................................... 2
VOLUME 5 · NO. 2
351
Braun et al.
ART I CLE
2 (1)
Conidia subacicular-iliform to narrowly obclavate, base of most conidia short to
long obconically truncate, narrow, 1.5–2 µm wide ..................................................................................... C. piaropi
Conidia acicular, base truncate, 2.5–4 µm wide ............................................................................................ C. rodmanii
Cercospora species on Pontederiaceae
Cercospora piaropi Tharp, Mycologia 9: 113 (1917).
(Fig. 153)
Synonym: Cercosporina piaropi (Tharp) Sacc., Syll. Fung.
25: 914 (1931).
Literature: Saccardo (1931: 914), Chupp (1954: 457),
Vasudeva (1963: 165), Crous & Braun (2003: 324), Kamal
(2010: 75), Conway (1976a), Groenewald et al. (2010, 2013),
Montenegro-Calderón et al. (2011).
Illustration: Conway (1976a: 1081, igs 1 right, 4–5).
Description: Leaf spots amphigenous, subcircular to
oval, 0.5–5 mm diam, dark brown, later with paler brown
or greyish brown centre and darker margin. Caespituli
amphigenous, often epiphyllous. Mycelium internal.
Stromata lacking or small, formed as small, substomatal
aggregations of swollen hyphal cells, brown, 10–20 µm diam.
Conidiophores solitary or in small fascicles, 2–10, arising
from internal hyphae or stromatic hyphal aggregations,
through stomata, erect, straight, subcylindrical to curved,
moderately geniculate-sinuous, unbranched, 10–200 ×
2.5–5 µm, continuous to pluriseptate, brown to medium
brown throughout or paler towards the tip, wall thin
or almost so, smooth; conidiogenous cells integrated,
terminal, occasionally intercalary, about 10–40 µm long,
conidiogenous loci conspicuous, 1.5–2.5 µm diam,
thickened and darkened. Conidia solitary, subaciculariliform to narrowly obclavate, 25–220 × 2–4(–5) µm, 2- to
pluriseptate, hyaline, thin-walled, smooth, apex pointed,
base almost truncate to usually short to long obconically
truncate, 1.5–2 µm wide, hila thickened and darkened.
Holotype: uSA: Texas: Palestine, on Eichhornia crassipes,
30 Oct. 1914, Lewis & Tharp,183 (BPI 439685).
Host range and distribution: On Eichhornia crassipes,
Pontederiaceae, Africa (South Africa, Nigeria, Zambia), Asia
(India, Bihar, Karnataka; Sri Lanka), North America (Mexico;
USA, Florida, Louisiana, Texas), South America (Brazil,
Venezuela).
Notes: A true Cercospora s. str. distinct from C. apii s. lat.
and C. rodmanii by having narrowly obclavate-iliform conidia
with obconically truncate base, only 1.5–2 µm wide. Several
collections from Brazil, India, South Africa and Sri Lanka have
been examined, but it is not clear if all records of C. piaropi
pertain to this species. It is possible that some of them refer
to C. rodmanii. Cercospora piaropi and C. rodmanii are two
very similar, confusable species on Eichhornia. Tessmann et
al. (2001) reduced C. rodmanii to synonym with C. piaropi,
but based on results of molecular sequence analyses and
morphology, Braun & Crous (2001), Groenewald et al. (2010,
352
Fig. 153. Cercospora piaropi (BPI 439685). A. Conidiophore
fascicles. B. Conidiophore tip. C. Conidia. Bar = 10 µm. U. Braun del.
2013) and Montenegro-Calderón et al. (2011) conirmed
that two separate species are involved. Cercospora piaropi,
characterised by its mostly narrowly obclavate-iliform conidia
with obconically truncate base, belongs to a clade treated as
“Cercospora cf. lagellaris” in Groenewald et al. (2010) and
was proven to be biologically conined to Eichhornia as host
(Montenegro-Calderón et al. 2011), whereas C. rodmanii, a
C. apii-like species with acicular conidia and truncate base,
forms a clade of its own, but differs biologically by being
plurivorous, which is not surprising for a species of the C.
apii complex.
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 154. Cercospora pontederiae (DAOM, Dearness 1800). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
Cercospora pontederiae Ellis & Dearn., Canad. Rec.
Sci. 5: 270 (1893).
(Fig. 154)
Synonym: Cercosporella pontederiae (Ellis & Dearn.) Davis,
Trans. Wisconsin Acad. Sci. 22: 182 (1926).
Literature: Saccardo (1895: 629), Chupp (1954: 458), Braun
(1995a: 126), Crous & Braun (2003: 333).
Illustration: Braun (1995: 125, ig. 111).
Description: Leaf spots amphigenous, irregular, indeinite,
brownish to reddish brown discolorations. Caespituli usually
epiphyllous, inconspicuous. Mycelium internal; hyphae
narrow, branched, septate, subhyaline. Stromata formed as
small substomatal hyphal aggregations, hyaline to faintly
pigmented, composed of a few slighty swollen hyphal cells.
Conidiophores in small, loose to dense fascicles, arising from
internal hyphae or hyphal aggregations, emerging through
stomata, straight, subcylindrical to geniculate-sinuous,
unbranched, 5–25 × 1.5–4 µm, aseptate, hyaline to faintly
pigmented, pale yellowish to very pale brownish, thin-walled,
smooth; conidiophores reduced to conidiogenous cells,
conidiogenous loci conspicuous but minute, 1–1.5 µm diam,
thickened and darkened. Conidia solitary, acicular to narrowly
subcylindrical, 15–45 × 1–2.5 µm, 1–2(–3)-septate, hyaline,
thin-walled, smooth, apex subacute, base ± truncate, 0.8–2
µm wide, thickened and darkened.
Lectotype (degignated by Braun 1995: 126): Canada:
Ontario: Niagara-on-the-Lake, on Pontederia cordata, 8 Aug.
1891, J. Dearness, no. 1800 (DAOM).
VOLUME 5 · NO. 2
Fig. 155. Cercospora rodmanii (BPI 440487). A. Conidiophores. B.
Conidia. Bar = 10 µm.
Host range and distribution: On Pontederia cordata,
Pontederiaceae, North America (Canada; USA, Texas,
Wisconsin).
Cercospora rodmanii Conway, Canad. J. Bot. 54:
1082 (1976).
(Fig. 155)
Literature: Crous & Braun (2003: 356), Groenewald et al.
(2010, 2013), Montenegro-Calderón et al. (2011).
353
Braun et al.
ART I CLE
Illustration: Conway (1976a: 1081, ig. 1 left, 2–3).
Description: Leaf spots punctiform to circular or subcircular,
1–3 mm diam, sometimes conluent and larger, blackish,
causing chlorosis of leaves and petioles. Caespituli
amphigenous. Mycelium internal. Stromata small, but mostly
developed, substomatal to immersed, 10–30 µm diam, brown.
Conidiophores in fascicles, 3–12, arising from stromata,
through stomata or erumpent, erect, straight, subcylindrical
to somewhat geniculate-sinuous, unbranched, 65–375 ×
4–5 µm, pluriseptate, brown, wall thin or almost so, smooth;
conidiogenous cells integrated, terminal, conidiogenous
loci conspicuous, about 2.5–3.5 µm diam, thickened and
darkened. Conidia solitary, acicular, straight to curved, 60–
350 × 4–5 µm, pluriseptate, hyaline, thin-walled, smooth,
apex pointed, base truncate, 2.5–4 µm, hila thickened and
darkened.
Holotype: uSA: Florida: Lake Alice, Gainesville, University of
Florida Campus, on Eichhornia crassipes, 19 Nov. 1974, K.
E. Conway (BPI 440487).
Host range and distribution: On Eichhornia crassipes,
Pontederiaceae, Africa (South Africa, Zambia), North
America (Mexico; USA, Florida, Texas), South America
(Brazil, Venezuela).
Notes: A true, plurivorous species of Cercospora s. str.
belonging to the C. apii s. lat. complex, distinct from C. piaropi
by having acicular conidia with truncate base. Tessmann et
al. (2001) considered C. rodmanii a synonym of C piaropi, but
the two species are genetically and morphologically distinct
(see C. piaropi).
Cercospora
Host range and distribution: Only known from the type
collection.
A single species.
Braun
&
Crous,
Illustration: Braun & Crous (2005: 397, ig. 3).
Description: Leaf spots amphigenous, subcircular to irregular,
2–10 mm wide, centre pale greenish, whitish, yellowish to
ochraceous, later becoming darker by abundant fructiication,
with narrow margin or halo, brownish, reddish, purple-violet,
occasionally somewhat raised. Caespituli amphigenous,
punctiform, dark brown, scattered to dense. Mycelium internal.
Stromata substomatal to intraepidermal, erumpent, 10–150
µm diam, medium brown, composed of swollen hyphal cells,
2–8 µm diam. Conidiophores in small to mostly large fascicles,
mostly more or less loose, arising from stromata, erect,
straight, subcylindrical to somewhat geniculate-sinuous,
unbranched, 20–110 × 3–8 µm, pluriseptate, subhyaline to
paler olivaceous or olivaceous-brown, smooth, wall thin to
slightly thickened; conidiogenous cells integrated, terminal,
354
occasionally intercalary, 10–60 µm long, conidiogenous
loci conspicuous, thickened and darkened, 2–3 µm diam.
Conidia solitary, obclavate-cylindrical, 25–100 × 4–7 µm,
(1–)3–8(–12)-septate, colourless, smooth, thin-walled, apex
obtuse, base truncate to usually obconically truncate, 2–3 µm
wide, hila thickened and darkened.
Holotype: Colombia: Bonda, on leaves of Smilax sp.,
Smilacaceae, 16 Nov. 1898, C. F. Baker 98 (NY 945706).
Smilacaceae
Cercospora smilacigena U.
Mycotaxon 92: 398 (2005).
(Fig. 156)
Fig. 156. Cercospora smilacigena (NY 945706). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
Doubtful, excluded and insuficiently known
species
Cercospora miyakei Henn., Bot. Jahrb. Syst. 37: 166
(1906).
(Fig. 157)
Synonym: Exosporium miyakei (Henn.) U. Braun & C.
Nakash., comb. nov.
MycoBank MB809016
Basionym: Cercospora miyakei Henn., Bot. Jahrb. Syst. 37:
166 (1906).
Literature: Saccardo (1913: 1429), Chupp (1954: 350),
Katsuki (1965: 43).
Illustration: Chupp (1954: 349, ig. 143).
Description: Leaf spots amphigenous, circular to angularirregular, 2–6 mm diam, often vein-limited, conluent, pale to
medium dark brown or with paler centre, surrounded by a
dark margin, dark brown to blackish, occasionally entire spots
IMA FUNGUS
Cercosporoid fungi 2
Host range and distribution: On Heterosmilax japonica,
Smilax nipponica, Smilacaceae, Asia (China, Japan).
ART I CLE
Lectotype (designated here, MycoBank MBT178167):
Japan: Tokyo, Meguro, on Smilax nipponica, 9 Oct. 1904,
J. Miyake 9 (B 700014862). Isolectotypes: CUP 40319,
TNS-F-15260.
Notes: Chupp (1954) excluded this species from Cercospora
and classiied it as helminthosporioid, which was conirmed
by Crous & Braun (2003). Lectotype and isolectotype
material has recently been re-examined. Due to fasciculate
conidiophores arising from well-developed stromata,
thickened and darkened conidiogenous loci and hila with
distinct central pore and scolecosporous, thick-walled,
distoseptate conidia, this species is better reallocated to
Exosporium.
Cercospora petersii (Berk. & M.A. Curtis) G.F. Atk., J.
Elisha Mitchell Sci. Soc. 8: 57 (1892).
(Fig. 158)
Fig. 157. Exosporium miyakei (B 700014862). A. Conidiophore
fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.
blackish. Caespituli amphigenous, mainly hypophyllous,
punctiform, scattered to aggregated. Mycelium internal.
Stromata substomatal to immersed, globose, 10–70 µm
diam, brown to dark brown, cells 2–8 µm diam, brown, wall
somewhat thickened. Conidiophores in loose, occasionally
denser fascicles, arising from stromata, through stomata
or erumpent, erect, straight to curved, lexuous, iliform to
strongly geniculate-sinuous throughout or only in the upper
half, simple or occasionally branched, irregular in width, (30–
)60–200(–250) × 3–6 µm, uniformly medium brown or tips
slightly paler, 1- to pluriseptate, cells 10–45 µm long, smooth,
wall thickened, to 0.8 µm; conidiogenous cells integrated,
terminal and intercalary, 10–35 µm long, conidiogenous loci
conspicuous, often on shoulders caused by geniculation
due to sympodial proliferation, 1.5–2 µm diam, in front view
circular, somewhat thickened and above all darkened, with
small central pore, conidiogenesis tretic. Conidia solitary,
occasionally catenulate, obclavate-cylindrical, straight to
somewhat curved, 30–150 × 4–9 µm, 5–12-distoseptate,
pale olivaceous to medium olivaceous-brown, outer wall thin,
but appearing thick-walled due to the inner wall structure
responsible for distoseptation, smooth or almost so, apex
obtuse, base gradually or short obconically truncate, 1.5–2
µm wide, with darkened or darkened-refractive hila.
VOLUME 5 · NO. 2
Basionym: Helminthosporium petersii Berk. & M.A. Curtis,
Grevillea 3: 102 (1875).
Synonyms: Exosporium petersii (Berk. & M.A. Curtis) U.
Braun, comb. nov.
MycoBank MB809016
Basionym: Helminthosporium petersii Berk. & M.A. Curtis,
Grevillea 3: 102 (1875).
Cercospora mississippiensis Tracy & Earle, Bull. Torrey
Bot. Club 22: 179 (1895) [lectotype (designated here,
MycoBank MBT178168): uSA: Mississippi: Starkville, on
Smilax glauca, 28 Oct. 1893, S. M. Tracy (BPI 438529);
isolectotypes: BPI 438534; NY 937094, 937095; OSU
9903; syntypes: NY 937094, 937096].
Pseudocercospora mississippiensis (Tracy & Earle) R.F.
Castañeda & U. Braun, Cryptog. Bot. 1: 52 (1989).
?Cercospora confusa G.F. Atk., undescribed ide Chupp
(1954: 352).
Literature: Saccardo (1895: xlviii), Chupp (1954: 349, 352),
Crous & Braun (2003: 277-278).
Illustration: Chupp (1954: 352, ig. 145).
Exsiccatae: Nash, Pl. Florida 2125. Ravenel, Fungi Amer.
Exs. 166, 616. Seym. & Earle, Econ. Fungi 199.
Description: Leaf spots amphigenous, circular, subcircular
to somewhat angular-irregular, 1–8 mm diam, occasionally
conluent and larger, at irst dark purple to blackish, later with
paler centre, pale to medium dark brown, inally greyish brown
do dingy grey, margin darker, often somewhat raised, narrow
to moderately wide, brown to blackish, sometimes reddish
brown to violet, or leaf spots sometimes somewhat zonate,
with brown centre, followed by a dark brown to blackish ringshaped zone surrounded by a paler narrow border. Caespituli
amphigenous, mainly hypophyllous, conspicuous, inely to
conspicuously punctiform or even coremioid, dark brown
to blackish. Mycelium internal; occasionally with supericial
hyphae, branched, septate, 2–5 µm wide, pale brown.
355
Braun et al.
ART I CLE
4942 (K(M) 190719). Isolectotype: BPI 864486 (ex herb.
Curtis 313). Syntype: K(M) 190718.
Host range and distribution: On Smilax (glauca, pseudochina
[tamnifolia], rotundifolia, tamnoides [hispida], Smilax sp.),
Smilacaceae, North America (Mexico; USA, Alabama,
Arkansas, Delaware, Florida, Georgia, Illinois, Maryland,
Montana, New York, North Carolina, Pennsylvania, South
Carolina, Tennessee, Texas, Virginia, Washington DC, West
Virginia, Wisconsin).
Fig. 158. Exosporium petersii (K(M) 190719). A. Conidiophore
fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
Stromata small, a few aggregated swollen hyphal cells
or larger, 10–40 µm diam, substomatal or intraepidermal,
medium to dark brown, cells 2–10 µm diam. Conidiophores in
loose to dense, sometimes coremioid fascicles, 2–25, arising
from stromatic cells or stromata, emerging through stomata
or erumpent, erect, lower part straight, subcylindrical, upper
fertile part geniculate-sinuous, usually strongly so, not or only
rarely branched, 30–250 × 3–6 µm, pluriseptate throughout,
medium to dark brown, wall thin to slightly thickened, to
1 µm, smooth; conidiogenous cells integrated, terminal
and intercalary, 10–40 µm long, proliferation sympodial
and occasionally also percurrent, enteroblastic, leaving a
conspicuous fringe; conidiogenous loci conspicuous, terminal
and laterial or often on shoulders caused by sympodial
proliferation, sometimes subdenticulate, slightly thickened,
darkened, in front view visible as small dark circles with
minute centre pore, 1–2 µm diam. Conidia solitary, distinctly
obclavate, often with broad dark base and narrow, pale
beak (almost alternarioid), the beak may be very long in
some conidia, small conidia occasionally broadly fusiform,
(20–)30–120(–130) × 4–8 µm, 2–9(–11)-distoseptate, pale
to medium olivaceous-brown, wall thin, but often appearing
somewhat thicker by a second inner wall layer, compound
width to 1.5 µm, smooth, apex obtuse or subobtuse, base
short obconically truncate, 1.5–2 µm wide, hila slightly
thickened, darkened-refractive.
Lectotype (designated here, MycoBank MB178169): uSA:
South Carolina: on Smilax glauca, Peters, herb. Curtis no.
356
Notes: The examination of type material and numerous
other collections of Cercospora petersii revealed that this
species has to be excluded from the cercosporoid fungi
and is better assigned to Exosporium. It seems to be close
to the Asian E. miyakei, which differs, however, in having
much larger stromata and obclavate-cylindrical conidia. E.
petersii has been recorded from different Smilax species.
It was not possible to re-examine and check material
on all recorded hosts. Smilax glauca seems to be the
most common host (numerous collections on this host,
deposited under different names, e.g. Cercospora smilacis,
C. smilacina, etc., have been examined and all of them
are true E. petersii, e.g. BPI 441298–441302, 441303,
441305, 441307, 441349–441355, 868949, and 864660).
A single collection deposited as C. smilacis on Smilax
tamnoides [hispida] from Illinois (BPI 441361) proved
to be E. petersii, but the host identiication could not be
checked. Cercospora s. lat. species on Smilax spp. are
often confused and identiications are often not reliable,
for instance, all examined collections on Smilax laurifolia
proved to be misidentiied and turned out to belong to
Pseudocercospora smilacicola.
Chupp (1954) treated Cercospora mississippiensis
and C. petersii, both described on Smilax glauca from
the USA, as two distinct species and confused several
cercosporoid fungi on Smilax species. Castañeda & Braun
(1989) introduced the combination Pseudocercospora
mississippiensis based on Cuban collections on Smilax
laurifolia and Smilax sp. and Chupp’s (1954) description.
Type material had not been re-examined. However, the
re-examination of type material of C. mississippiensis and
numerous additional samples showed that this species
is conspeciic with C. petersii. Material on Smilax glauca
from Cuba and North America previously assigned to C.
mississippiensis and C. petersii, respectively, proved to
represent a true Pseudocercospora, supericially confusable
with Exosporium petersii but readily distinguishable by
having unthickened, not darkened conidiogenous loci and
euseptate conidia, which is described as Pseudocercospora
smilacicola.
Cercospora mississippiensis has been recorded on
Smilax auriculata, S. hispida, S. pseudochina [S. tamnifolia],
S. riparia and S. tamnoides. It is unclear if all records on these
hosts belong to E. petersii. Re-examinations are necessary.
Material recorded from Laos as C. petersii on Smilax
chinensis (Phensingtham et al. 2013b: 154) has been reexamined (duplicate deposited as HAL 2660 F) and proved
to be Passalora smilacis.
IMA FUNGUS
Cercosporoid fungi 2
Passalora
1
Conidia obclavate-cylindrical, thin-walled, euseptate; conidiogenous cells with
thickened and darkened conidiogenous loci ............................................................................................................ 2
Conidia distoseptate; conidiogenous loci thickened and darkened, with conspicuous central pore ............................... 3
2 (1)
Conidiophores long, 10–130 × 2.5–7 µm, 0–12-septate, olivaceous to dark olivaceous-brown or
medium brown throughout or tips paler, wall to 1 µm thick ...................................................................... p. smilacis
Conidiophores uniformly short, 5–30 × 2–5 µm, 0–1(–2)-septate, subhyaline to pale olivaceous-brown,
paler towards the tip, thin-walled ....................................................................................................... p. pallidissima
3 (1)
Stromata lacking or small, 10–40 µm diam; conidia distinctly obclavate, often with long,
narrow, paler beak; on Smilax spp. in North America ....................................................... see exosporium petersii
Stromata well-developed, 10–70 µm diam; conidia obclavate-cylindrical;
on Smilax spp. in Asia ....................................................................................................... see exosporium miyakei
ART I CLE
Key to Passalora species on Smilacaceae
Passalora species on Smilacaceae
passalora pallidissima (Chupp) U. Braun, comb.
nov.
MycoBank MB809018
(Fig. 159)
Basionym: Cercospora pallidissima Chupp, Monograph of
Cercospora: 350 (1954).
Synonym: Pseudocercospora pallidissima (Chupp) Deighton,
Mycol. Pap. 140: 149 (1976).
Literature: Chupp (1954: 350), Crous & Braun (2003: 304),
Braun & Crous (2005: 413).
Illustration: Chupp (1954: 349, ig. 144).
Description: Leaf spots amphigenous, 2–15 mm diam,
circular to angular-irregular, sometimes vein-limited, brown,
later dull grey, with darker margin or marginal line, brown
to reddish brown, narrow. Caespituli amphigenous, mainly
hypophyllous, punctiform, scattered to somewhat aggregated,
medium to dark brown. Mycelium internal. Stromata welldeveloped, substomatal to immersed, globose, 20–70 µm
diam, medium brown. Conidiophores fasciculate, arising
from stromata, in small to moderately large fascicles, mostly
2–15, divergent to dense, erect, straight to slighty curved,
subcylindrical to conical, somewhat narrowed towards the tip,
barely or only slightly geniculate, unbranched, 5–30 × 2–5
µm, 0–1(–2)-septate, subhyaline to pale olivaceous-brown,
paler towards the tip, thin-walled, smooth; conidiophores
integrated, terminal or conidiophores often reduced to
conidiogenous cells, conidiogenous loci conspicuous, slightly
thickened and somewhat darkened-refractive, 0.8–1.5 µm
diam. Conidia solitary, obclavate-cylindrical, straight to
somewhat curved, 25–70 × 2–5 µm, 1–7-septate, subhyaline
to very pale yellowish or olivaceous, thin-walled, smooth
or almost so, apex obtuse, subobtuse or subacute, base
subtruncate to short obconically truncate, 1–1.5 µm wide,
hila almost unthickened to slightly thickened, somewhat
darkened-refractive.
VOLUME 5 · NO. 2
Fig. 159. Passalora pallidissima (CUP 40454). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
357
ART I CLE
Braun et al.
Holotype: Brazil: São Paulo, Campinas, Bosque dos
Jequitibas, on Smilax sp., Smilacaceae, 5 Sep. 1943, A. P.
Viégas & A. R. Teixeira 4276 (CUP 40454).
Host range and distribution: Only known frome the type
collection.
Notes: Deighton (1976) introduced the combination
Pseudocercospora pallidissima, but it is quite unclear on which
collections his reallocation was based. It seems that type
material of this species was not examined by Deighton. The
type collection does not contain any corresponding annotations.
Due to the conspicuous conidiogenous loci and conidial hila,
Cercospora pallidissima is better placed in Passalora. This
species is probably only known from the type collection in
Brazil. Indian records of “Pseudocercospora pallidissima” on
Smilax spp., including P. pallidissima var. constrictolexuosa,
are distinguished by having strongly geniculate-sinuous
conidiophores, inconspicuous conidiogenous loci (unthickened,
not darkened) and belong to separate species of the genus
Pseudocercospora (see P. constrictolexuosa and P. smilacis).
A record of P. pallidissima on S. anceps [kraussiana] from South
Africa (Crous & Braun 1996b) refers to a true Pseudocercospora
(see Pseudocercospora pycnidioides). Other records on
S. aspera (Crous & Braun 2003) and S. laurifolia from North
America (Braun & Crous 2005) are unclear, have to be proven
and seem to belong to Pseudocercospora species as well.
passalora smilacis (Thüm.) U. Braun, Arnoldia 14: 30
(1997).
(Fig. 160)
Basionym: Cercospora smilacis Thüm., Contrib. Fl. Mycol.
Lusit. 2: 14 (1879).
Synonyms: Cercospora smilacina Sacc., Michelia 2: 364
(1881) [holotype: France: Collioure, on Smilax aspera, O.
Debeaux (PAD); isotypes: Roum., Fungi Gall. Exs. 723].
Cercospora smilacis var. asperae Gonz. Frag.,Trab. Mus.
Nac. Ci. Nat., Madrid, ser. Bot., 9: 66 (1916) [holotype:
Spain: Barcelona, on Smilax aspera, 20 Sep. 1915,
Sennen (MA)].
Literature: Saccardo (1886: 476), Lindau (1910: 799), Gonzáles
Fragoso (1927: 230), Chupp (1954: 354), Ellis (1976: 271),
Braun & Mel'nik (1997: 93), Crous & Braun (2003: 378–379),
Phensingtham et al. (2013b: 154, as Cercospora petersii).
Illustration: Ellis (1976: 272, ig. 206A).
Exsiccatae: Kabát & Bubák, Fungi Imperf. Exs. 198. Herb.
Crypt. Ind. Orient. Exs. 27 (as Cercospora mississippiensis).
Krypt. Exs. 728a,b. Migula, Krypt. Germ. Austr. Helv. Exs.
179. Rabenh., Fungi Eur. Exs. 2975. Roum., Fungi Gall. Exs.
723. D. Sacc., Mycoth. Ital. 396. Thüm., Mycoth. Univ. 1670.
Triebel, Microf. Exs. 245.
Description: Leaf spots amphigenous, circular to somewhat
angular-irregular, 1–12 mm diam, sometimes conluent, pale
to dark brown or reddish brown with narrow to moderately
wide, darker border, sometimes somewhat raised and with
yellowish halo, occasionally entire spots almost black. Caespituli
358
Fig. 160. Passalora smilacis (HAL, Thüm., Mycoth. Univ. 1670). A.
Supericial hyphae. B. Supericial hypha with solitary conidiophore. C.
Conidiophore fascicles. d. Conidiophores. e. Conidia. Bar = 10 µm.
amphigenous, punctiform, scattered to dense, dark brown to
blackish. Mycelium internal, sometime also external; internal
hyphae 1.5–5 µm wide, branched, septate, subhyaline to medium
olivaceous-brown; supericial hyphae when present sparingly
branched, 1.5–3.5 µm wide, septate, occasionally constricted
at the septa, subhyaline, pale greenish, yellowish to olivaceousbrown, thin-walled, smooth, but solitary conidiophores arising from
supericial hyphae rare. Stromata well-developed, substomatal
to intraepidermal, immersed, 10–80 µm diam, brown to dark
brown. Conidiophores in small to large fascicles, rarely solitary,
arising from stromata, rather variable, loose to dense, sometimes
very dense, almost coremioid, uniformly short conidiophores
when dense forming almost sporodochial conidiomata, long
conidiophores when dense sometimes almost coremioid,
occasionally with solitary conidiophores arising from supericial
hyphae, lateral, conidiophores erect, straight, subcylindrical to
strongly geniculate-sinuous, simple or occasionally branched,
10–130 × 2.5–7 µm, 0–12-septate, olivaceous to dark olivaceousbrown or medium brown throughout or tips paler, wall to 1 µm
IMA FUNGUS
Cercosporoid fungi 2
Lectotype (designated here, MycoBank MBT178170):
portugal: Coimbra, on Smilax aspera [mauritanica], May
1879, F. Moller (BPI 441368). Topotypes (Jul. 1879): Thüm.,
Mycoth. Univ. 1670, e.g. BPI 441367, 441368, CUP 41239,
HAL, LEP.
Host range and distribution: On Smilax (aspera [goetzeana,
mauritanica, nigra], bona-nox, chinensis, excelsa, guianensis
[macrophylla], havanensis, herbacea, Smilacaceae, Africa
(Algeria, Canary Islands, Kenya, Libya, Morocco), Asia
(India, Himachal Pradesh; Iran, Israel, Jordan, Laos, Nepal,
Turkey), Caucasus (Georgia), Europe (Cyprus, France,
Italy, Greece, Malta, Montenegro, Portugal, Russia, Spain,
including Balearic Islands, Turkey, former Yugoslavia), North
America (USA, Colombia, Connecticut, Delaware, District
of Columbia, Florida, Georgia, Indiana, Iowa, Kansas,
Louisiana, Maryland, Minnesota, Mississippi, Missouri,
North Carolina, Nebraska, New Jersey, New York, North
Carolina, Oklahoma, South Carolina, Tennessee, Texas,
Virginia, Washington DC, West Virginia), South America
(Brazil, Uruguay, Venezuela).
Notes: The generic afinity of Cercospora smilacis is uncertain
and needs to be established by means of molecular sequence
analyses. Due to the pigmented conidia and conspicuous
conidiogenous loci, this species has been assigned to
Passalora, which is the best solution for now, but the loci are
in front view visible as minute circles and hence somewhat
paracercosporoid. Moreover, the loci resemble those of
Exosporium miyakei and E. petersii. Relations between these
species are also unclear. Collections of Cercospora s. lat. on
Smilax spp. in herbaria are often misidentiied. Records of
C. smilacis and C. smilacina on Smilax brasiliensis belong to
Pseudocercospora pycnidioides, other samples on Smilax
auriculata and S. laurifolia to Pseudocercospora smilacicola and
all collections on S. glauca have to be assigned to Exosporium
petersii, but North American specimens on S. bona-nox and
S. herbacea (e.g. BPI 441356, 441357) represent genuine
Passalora smilacis. Several collections of P. smilacis from Asia
have been examined, e.g. from India on S. aspera, from Laos
on S. chinensis and from Turkey on S. excelsa (BPI 441296).
Records of C. smilacis from Jamaican and Mexico are unproven
and doubtful. True Passalora smilacis is distributed in North
America, but it is unclear if all records from all states of the
USA being correct. It was not possible to conirm the identity of
material from all recorded areas.
ART I CLE
thick, above all below, smooth; conidiogenous cells integrated,
terminal, sometimes conidiophores reduced to conidiogenous
cells, 10–35 µm long, proliferation sympodial, occasionally
percurrent, conidiogenous loci minute but conspicuous, 1–1.5(–2)
µm diam, almost unthickened to slightly thickened and somewhat
darkened-refractive, in front view visible as minute dark circle.
Conidia solitary, obclavate-cylindrical, 20–90(–120) × (2–)3–6
µm, 1–12-septate, subhyaline to pale olivaceous or brownish,
thin-walled, smooth, apex obtuse or subobtuse, base rounded to
short obconically truncate, 1–2 µm wide, hila slightly thickened
and somewhat darkened-refractive.
Pseudocercospora
Key to Pseudocercospora species on Smilacaceae
1
Conidiophores very long, 20–350 µm, thick-walled, subcylindrical, straight to curved,
only terminal fertile part of long conidiophores geniculate-sinuous, unbranched;
stromata lacking or small, 10–40 µm diam ......................................................................................... p. smilacicola
Conidiophores much shorter, about 10–80 µm, thin-walled or almost so;
either non-geniculate or strongly geniculate-sinuous throughout;
stromata well-developed, 20–100 µm ...................................................................................................................... 2
2 (1)
Conidiophores straight, subcylindrical-conical, neither geniculate-sinuous nor denticulate,
usually unilocal, occasionally percurrent with ine annellations, unbranched;
Africa and South America ................................................................................................................. p. pycnidioides
Conidiophores strongly geniculate-sinuous, often with constrictions and subdenticulate, sympodial,
multilocal, unbranched or often branched; Asia (India) ............................................................................................ 3
3 (2)
Stromata 20–50 µm diam; conidiophores 10–60 × 3–6 µm, unbranched; conidia subhyaline;
on Smilax perfoliata ............................................................................................................... P. constrictolexuosa
Stromata larger, 40–80 µm diam; conidiophores simple or often branched, longer,
60–75 × 3–4 µm; conidia pale brown; on Smilax aspera ......................................................................... p. smilacis
Pseudocercospora species on Smilacaceae
Pseudocercospora constrictolexuosa (U. Braun)
U. Braun, comb. stat. nov.
MycoBank MB809019
(Fig. 161)
VOLUME 5 · NO. 2
Basionym:
Pseudocercospora
pallidissima
var.
constrictolexuosa U. Braun, Monogr. Cercosporella,
Ramularia Allied Genera (Phytopath. Hyphom.). 1: 198
(1995).
Illustration: Braun (1995a: 197, ig. 193).
359
Braun et al.
ART I CLE
10–30 µm long, conidiogenous loci inconspicuous or
subdenticulate, but always unthickened and not darkened.
Conidia solitary, obclavate-subcylindrical, rarely subacicular,
30–70 × (2.5–)3–5 µm, 2–6-septate, subhyaline or only with
a faintly chlorine tinge, thin-walled, smooth, apex subacute or
subobtuse, base short obconically truncate, 1.5–2.5 µm wide,
hila unthickened, not darkened.
Holotype: India: Uttar Pradesh: Gorakhpur, on Smilax
perfoliata [prolifera], Smilacaceae, 28 Feb. 1979, B. Rai
(K(M) IMI 235973).
Host range and distribution: Only known from the type
collection.
Notes: Due to its pale conidia, this Indian fungus was
described as variety of Pseudocercospora pallidissima.
However, the basionym, Cercospora pallidissima, proved to
be a species of Passalora, and owing to strongly geniculatesinuous conidiophores with numerous constrictions, this
species is closer to the Indian P. smilacis, from which
it differs in having quite distinct leaf spots, much smaller
stromata and shorter conidiophores as well as subhyaline
conidia. Type material of P. constrictolexuosa contains
traces of Zasmidium smilacis.
pseudocercospora pycnidioides (Chupp) U. Braun
& Crous, Mycosphaerella and Anam. 1: 345 (2003).
(Fig. 162)
Basionym: Cercospora pycnidioides Chupp, Monograph of
Cercospora: 353 (1954).
Synonym: Cercospora smilacina Speg., Rev. Mus. La Plata
15: 46 (1908), nom. illeg. (ICN, Art. 53.1), non C. smilacina
Sacc. 1881.
Literature: Chupp (1954: 353), Crous & Braun (1996b: 293,
as Pseudocercospora pallidissima).
Illustration: Chupp (1954: 353, ig. 146).
Exsiccatae: Herter, Pl. Uruguayenses Exs. 1486, as
Cercospora smilacis.
Fig. 161. Pseudocercospora constrictolexuosa (K(M) IMI 235973).
A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
Description: Leaf spots amphigenous, angular-irregular, 2–8
mm diam, pale, greenish white, surrounded by a narrow dark
brown to blackish brown margin or marginal line. Caespituli
amphigenous, punctiform, scattered, blackish brown.
Mycelium internal. Stromata substomatal to intraepidermal,
20–50 µm diam, brown. Conidiophores in moderately large
fascicles, arising from stromata, through stomata or erumpent,
erect, straight to curved or lexuous, subcylindrical, mostly
strongly geniculate-sinuous, with constrictions, unbranched,
10–60 × 3–6 µm, 0–5-septate, olivaceous to medium brown,
paler above, pale olivaceous or subhyaline, wall thin to
slightly thickened, smooth; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells,
360
Description: Leaf spots amphigenous, circular, 1–20 mm
diam, pale brown to brown with darker brown border.
Caespituli amphigenous, punctiform, dark. Mycelium
internal; occasionally with a few supericial hyphae. Stromata
substomatal to immersed, well-developed, 20–150 µm diam,
pycnidioid, brown to dark brown, composed of swollen
hyphal cells, 2–12 µm diam, thick-walled. Conidiophores in
dense, compact fascicles, arising from stromata, forming
sporodochial conidiomata, occasionally solitary, arising from
supericial threads between stromata when formed, erect,
straight to curved, subcylindrical-conical, not geniculate,
unbranched, short, 5–60 × 3–6 µm, 0–2-septate, pale to
medium olivaceous-brown, darker in mass, wall thin to
slightly thickened, smooth to inely roughened; conidiogenous
cells integrated, terminal, but conidiophores often reduced
to conidiogenous cells, 5–25 µm long, without distinct
proliferation or percurrent, with 1–2 irregular ine annellations,
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Notes: Cercosporoid species on Smilax have often been
confused. Many collections in herbaria are misidentiied.
Numerous collections are deposited under C. smilacina, but
almost all of them are based on wrong identiications as already
mentioned in Chupp (1954). Records from India on S. guianensis
[macrophylla] (Crous & Braun 2003) are also doubtful, unveriied
and not included in Kamal (2010). A collection from Zambia [near
Victoria Falls, on Smilax sp., 13 Mar. 2006, Ch. Hahn (GZU 212007)] is an additional sample that agrees with type material
of this species, at least morphologically. Material previously
recorded from South Africa as “Pseudocercospora pallidissima”
(Crous & Braun 1996b: 293) has been re-examined and proved
to be a collection belonging to P. pycnidioides as well [South
Africa: Mpumalanga, Nelspruit district, Schagen, on living leaves
of Smilax anceps [kraussiana], June 1939, L. C. C. Liebenberg
(PREM 32922)]. Type material of Cercospora pallidissima has
been re-examined and proved to be a species of Passalora due
to thickened and darkened conidiogenous loci and conidial hila.
pseudocercospora smilacicola U. Braun, sp. nov.
MycoBank MB809025
(Fig. 163)
Etymology: “smilacicola”, derived from the host genus
(inhabitant of Smilax).
Synonym: Cercospora petersii f. smilacis-beyrichii Elllis, in
Nash, Pl. Florida 2073 (1895).
Missapplied names: Cercospora mississippiensis auct. non
Tracy & Earle and Pseudocercospora mississippiensis auct.
Literature: Castañeda Ruiz & Braun (1989: 52), Kim & Shin
(1999b), Shin & Kim (2001: 206), all as Pseudocercospora
mississippiensis.
Illustrations: Castañeda Ruiz & Braun (1989: 48, plate 4, ig.
24), Shin & Kim (2001: 207, ig. 93), all as Pseudocercospora
mississippiensis.
Fig. 162. Pseudocercospora pycnidioides (LPS 953). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
rarely sympodial, conidiogenous loci inconspicuous, neither
thickened nor darkened. Conidia solitary, cylindrical to
obclavate-cylindrical, 20–85(–100) × 3.5–6 µm, 1–7-septate,
subhyaline, pale olivaceous to olivaceous-brown or brown,
wall thin to slightly thickened, smooth to faintly roughened,
apex obtuse, base short obconically truncate, 2–2.5 µm wide,
hila unthickened, not darkened.
Lectotype (designated here, MycoBank MBT178171):
Brazil: São Paulo, on Smilax sp., A. Usteri 953 (LPS 953).
Isolectotype: CUP 41237.
Host range and distribution: On Smilax (anceps [kraussiana],
brasiliensis, Smilax sp.), Smilacaceae, Africa (South Africa,
Zambia), South America (Brazil, Uruguay).
VOLUME 5 · NO. 2
Exsiccatae: Nash, Pl. Florida 1788, 1893, 2073 (as
Cercospora petersii).
Diagnosis: Morphologically resembling Exosporium petersii
but conidiogenous loci and conidial hila unthickened and
conidia euseptate.
Description: Leaf spots amphigenous, subcircular to angularirregular, 1–10 mm diam, at irst pale, later very dark brown,
inally with paler centre, brownish to greyish brown, occasionally
somewhat zonate, with darker brown to blackish border
or marginal line, sometimes somewhat raised, sometimes
surrouned by a diffuse lighter halo. Caespituli amphigenous,
mainly hypophyllous, very dark, brown to black, punctiform,
scattered. Mycelium internal; hyphae branched, septate, 1.5–3
µm wide, pale. Stromata lacking or 10–60 µm diam, substomatal,
brown to dark brown, composed of swollen hyphal cells,
rounded to somewhat angular-irregular in outline, 2–7 µm diam.
Conidiophores in small to moderately rich fascicles, mostly 5–20,
divergent to dense, occasionally almost coremioid, arising from
361
ART I CLE
Braun et al.
Fig. 163. Pseudocercospora smilacicola (HAL 2657 F). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
internal hyphae or stromata, emerging through stomata, erect,
straight to curved, subcylindrical, upper fertile part distinctly, often
strongly geniculate-sinuous, at least in older long conidiophores,
but not denticulate, unbranched, 20–250 × 3–6 µm, pluriseptate
throughout, pale to medium olivaceous-brown when young, later
medium to darker olivaceous-brown or brown, but tips usually
paler, sometimes much paler, wall thin to somewhat thickened,
to 0.8 µm, smooth; conidiogenous cells integrated, terminal and
intercalary, 10–40 µm long, conidiogenous loci inconspicuous
or visible as “shoulders” caused by sympodial proliferation,
but always unthickened and not darkened, at most somewhat
refractive. Conidia solitary, obclavate to obclavate-cylindrical,
shorter conidia often subcylindrical, straight to somewhat curved,
20–120 × 3–7 µm, 1–11-septate, subhyaline to pale olivaceous,
thin-walled, smooth, apex obtuse to somewhat pointed, base short
or long obconically truncate, 1.5–3 µm wide, hila unthickened, not
darkened.
Holotype: Cuba: Matanzas: San Miguel de los Baños, on
Smilax laurifolia [lanceolata], 11 Mar. 1987, R. F. Castañeda
(HAL 2657 F ). Isotype: INIFAT C87/14.
Host range and distribution: On Smilax (auriculata [beyrichii],
laurifolia, pseudochina, riparia, rotundifolia, Smilax sp.),
Smilacaceae, Asia (Japan, Korea), North America (USA,
Georgia, Florida, Louisiana, Mississippi, Pennsylvania), West
Indies (Cuba).
362
Fig. 164. Pseudocercospora smilacis (K(M) IMI 291864). A.
Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.
Notes: Pseudocercospora smilacicola comprises several
collections of cercosporoid fungi previously referred to
as Cercospora petersii and C. (or Pseudocercospora)
mississippiensis. Although supericially similar and confusable,
the new species is easily distinguishable from Exosporium
petersii (= C. mississippiensis) by having unthickened, not
darkened conidiogenous loci and hila as well as euseptate
conidia. “Pseudocercospora mississippiensis” on Smilax
riparia from Korea (Shin & Kim 2001) is morphologically
indistinguishable from Cuban and North American collections
of P. smilacicola (all examined North American collections
on Smilax laurifolia, e.g. BPI 441364 from Florida, belong
to this species). Some North American samples on Smilax
auriculata (BPI 438526, 439606–439607, 441343–441344)
and single samples on Smilax pseudochina (BPI 441369)
and S. rotundifolia (BPI 441314), deposited as C. smilacis,
proved to be P. smilacicola. Furthermore, an unpublished
collection on Smilax sp. from Japan deposited as MUMH
11273 agrees morphologically well with P. smilacicola.
IMA FUNGUS
Cercosporoid fungi 2
Literature: Kamal (2010: 206, as P. pallidissima).
Illustration: Budathoki & Singh (1995: 231, ig. 3).
Description: Leaf spots amphigenous, circular, 1–6 mm diam,
dark brown. Caespituli hypophyllous. Mycelium internal;
hyphae branched, septate, subhyaline, thin-walled, smooth.
Stromata substomatal, subglobose, pseudoparenchymatic,
40–80 µm diam, brown. Conidiophores in loose fascicles,
arising from stromata, erect, straight to lexuous,
subcylindrical to strongly geniculate-sinuous, simple
or branched, 60–75 × 3–4 µm, septate, brown to
dark brown, thin-walled, smooth; conidiogenous cells
integrated, terminal and intercalary, about 10–25 µm long,
conidiogenous loci inconspicuous to subdenticulate, but
always unthickened and not darkened. Conidia solitary
obclavate(-subcylindrical), straight to curved, 30–80 × 3–6.5
µm, 1–8-septate, pale brown, thin-walled, smooth, apex
acute to obtuse, base short obconically truncate, 1.5–3 µm
wide, hila unthickened, not darkened.
ART I CLE
pseudocercospora smilacis Budathoki & S.K. Singh,
Mycol. Res. 99: 232 (1995).
(Fig. 164)
Holotype: Nepal: Kathmandu Valley, Godawari, on Smilax
aspera, Nov. 1985, U. Budathoki KU 59 (K(M) IMI 291864).
Host range and distribution: On Smilax aspera, Smilacaceae,
Asia (India, Himachal Pradesh; Nepal).
Zasmidium
Key to Zasmidium species on Smilacaceae
1
In vivo supericial hyphae lacking; conidiophores solitary or fasciculate, arising from stromata;
conidia long and broad, 15–110 × 3–9 µm, pluriseptate (1–14); on Smilax perfoliata ............................. Z. indicum
In vivo at least partly with supericial mycelium, hyphae verruculose; solitary conidiophores
arising from supericial hyphae present ................................................................................................................... 2
2 (1)
Conidiophores consistently short, 10–30 µm, 0–1-septate; conidia solitary, smooth,
at most some conidia faintly rough-walled with age, 15–60 × (2.5–)3–5(–5.5) µm;
on Smilax sp., India .............................................................................................................. Z. mycovellosielloides
Conidiophores much longer, 50–165 µm, 1–12-septate and conidia verruculose or 12–85 µm
and 1–4-septate, but then conidia 4–6.5 µm wide, 1–6-septate and verrucose ...................................................... 3
3 (2)
Conidiophores consistently solitary, arising from supericial hyphae; conidia solitary as well as catenate,
narrow, 2–4 µm; on Smilax spp. .............................................................................................................. Z. smilacis
Conidiophores emerging through stomata as well as solitary, arising from supericial hyphae;
conidia solitary, much broader, 4–6.5 µm; on Smilax ovalifolia ....................................... Z. smilacis-macrophyllae
Zasmidium species on Smilacaceae
Zasmidium indicum (Kamal & R.P. Singh) U. Braun,
comb. nov.
MycoBank MB809020
(Fig. 165)
Basionym: Verrucispora indica Kamal & R.P. Singh, Canad.
J. Bot. 56: 2785 (1978).
Synonym: Sirosporium indicum (Kamal & R.P. Singh) D.E.
Shaw, Alcorn & B. Sutton, Austral. Syst. Bot. 6: 274
(1993).
Literature: Kamal (2010: 274).
Illustrations: Kamal & Singh (1978: 2786, ig. 1), Kamal
(2010: 275, ig. 46).
Description: Leaf spots amphigenous, circular, 2.5–12
mm diam, centre dark brown, inally almost blackish,
margin pale. Caespituli punctiform, blackish. Mycelium
internal; hyphae branched, septate, colourless, smooth.
Stromata well-developed, substomatal to intraepidermal,
VOLUME 5 · NO. 2
pseudoparenchymatic, 20–85 µm diam, medium to dark brown
or reddish brown, cells rounded to angular, 2.5–7 µm diam.
Conidiophores in loose to moderately dense fascicles, arising
from stromata, through stomata or erumpent, subcylindrical,
straight to somewhat curved-lexuous, apical part often
somewhat swollen or somewhat geniculate, 70–210 µm long
and 3–5.5 µm wide, brown or reddish brown, wall thickened,
smooth or almost so; conidiogenous cells integrated,
terminal, 5–20 µm long, sympodial, somewhat paler, with a
single or several distinct conidiogenous loci, thickened and
darkened, about 2–2.5 µm diam. Conidia solitary, cylindrical
or somewhat obclavate-cylindrical, 15–110 × 3–9 µm,
0–10-septate, wall thickened, pale brown or reddish brown,
verrucose, apex obtuse, base short obconically truncate,
2–2.5 µm wide, hila somewhat thickened and darkened.
Holotype: India: Uttar Pradesh: Gorakhpur, on Smilax
perfoliata [prolifera], Feb. 1976, R. P. Singh 190 (K(M) IMI
210821).
Host range and distribution: On Smilax perfoliata [prolifera],
Smilacaceae, Asia (India, Uttar Pradesh).
363
ART I CLE
Braun et al.
364
Fig. 165. Zasmidium indicum (K(M) IMI 210821). A. Conidiophore
fascicle. B. Conidiophores. C. Conidiophore tip. d. Conidia. Bar =
10 µm.
Fig. 166. Zasmidium mycovellosielloides (BPI 438522). A. Supericial
hypha. B. Solitary conidiophores arising from supericial hyphae. C.
Conidia. Bar = 10 µm.
Notes: This species belongs to a group of Zasmidium species
without any supericial mycelium in vivo. It is a typical member
of the former genus Verrucisporota, which has been reduced
to synonymy with Zasmidium (see Braun et al. 2013).
McKenzie (1982) discussed Verrucispora indica, compared
it with Biharia smilacis and considered the two species
conspeciic. Kamal (2010) followed McKenzie’s (1982)
treatment, but Shaw & Alcorn (1993) maintained two different
species, introduced the new name Stenella smilacicola for
Biharia smilacis under Stenella and reallocated Verrucispora
indica to Sirosporium. Conidiophores and conidia of these
species are, indeed, very similar, but they occur on two
different Smilax spp. and supericial mycelium with solitary
conidiophores is lacking in V. indica. Therefore, they are
maintained as two different species.
Zasmidium mycovellosielloides U. Braun, sp. nov.
MycoBank MB809026
(Fig. 166)
Etymology: mycovellosielloides = resembling species of the
former genus Mycovellosiella.
Diagnosis: Resembling Zasmidium smilacis-macrophyllae,
but conidiophores shorter, 10–30 µm, 0–1-septate, and
narrower, paler, and above all smooth conidia.
Description: Leaf spots amphigenous, circular, subcircular to
irregular, 2–25 mm diam or conluent and larger, brownish,
later pale, surrounded by a darker margin, medium to dark
brown. Colonies amphigenous, inconspicuous (surrounded
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
by distinct punctiform immature fruiting bodies which might
be immature spermogonia or ascomata). Mycelium internal
and external; supericial hyphae branched, 1–5 µm wide,
thin-walled, subhyaline, pale olivaceous to pale brown
or olivaceous-brown, smooth to verruculose, wider fertile
hyphae usually smooth and darker, paler hyphae without
conidiophores often somewhat verruculose. Stromata lacking
or small, substomatal to epidermal, brown, composed of
some swollen hyphal cells, 3–7 µm wide, wall somewhat
thickened. Conidiophores solitary, arising from supericial
hyphae, occasionally arising from stromatic hyphae cells,
solitary or in small groups, erect, straight, subcylindricalconical, barely geniculate-sinuous, unbranched, 10–30 × 2–5
µm, 0–1-septate, pale to medium brown or olivaceous-brown,
thin-walled, smooth or faintly rough with age; conidiophores
usually reduced to conidiogenous cells, occasionally
integrated, terminal, 10–25 µm long, with a single to several
conspicuous conidiogenous loci near the apex, thickened
and darkened, 1–1.5 µm diam. Conidia solitary, obclavatesubcylindrical, short conidia ellipsoid-ovoid, straight to slightly
curved, 15–60 × (2.5–)3–5(–5.5) µm, (0–)1–3(–4)-septate,
subhyaline to pale olivaceous or olivaceous-brown, thinwalled, smooth, some conidia rough-walled with age, apex
obtuse, base short obconically truncate, 1–1.5 µm wide, hila
slightly thickened and darkened.
Holotype: India: Karnataka: Bangalore, Yercaud, on Smilax
sp., Smilacaceae, 25 Feb. 1965, R. Swaminarhan & V.
Raghunath (BPI 438522).
Host range and distribution: Only known from the type
collection.
Notes: The host leaves are only identiied as Smilax sp. They
are very large, to about 20 × 15 mm, broadly oval-elliptical
and remind one of leaves of Smilax macrophylla. Zasmidium
smilacis-macrophyllae is a comparable species but differs
in having much longer, septate conidiophores and broader,
darker, verrucose conidia.
Zasmidium smilacicola (D.E. Shaw & Alcorn) U.
Braun, comb. nov.
MycoBank MB809021
(Fig. 167)
Basionym: Stenella smilacicola D.E. Shaw & Alcorn,
Australian Systematic Botany 6: 274 (1993).
Synonyms: Biharia smilacis Agarwal, J. Indian Bot. Soc. 39:
354 (1960), non Zasmidium smilacis (P. Kumar, D.N.
Shukla & Kamal) Kamal 2010.
Verrucispora smilacis (Agarwal) McKenzie, New Zealand J.
Bot. 20: 252 (1982).
Stenella liliacearum N. Sharma, Soni & R.K. Verma, Indian
J. Trop. Biodiv. 14: 36 (2006) [holotype: India: Madhya
Pradesh: Balaghat, Supkhar, on Smilax ovalifolia
[macrophylla], 19 Dec. 2004, R. K. Verma (Hb. Tropical
Research Institute Jabalpur, TF213); isotype: HCIO
45901].
Literature: Subramanian (1971: 225), McKenzie (1982),
Shaw & Alcorn (1993).
VOLUME 5 · NO. 2
Fig. 167. Zasmidium smilacicola (K(M) IMI 75641). A. Supericial
hyphae. B. Solitary conidiophores arising from supericial hypha. C.
Conidiophore fascicle. d. Conidiophore tips. e. Conidia. Bar = 10 µm.
Illustrations: Agarwal & Beliram (1960: 355, igs 1–2), Sharma
et al. (2006: 36, ig. 2).
Description: Leaf spots amphigenous, circular to irregular,
to 20 mm diam, medium to dark brown, margin indeinite or
narrow and somewhat darker. Caespituli amphigenous, rather
inconspicuous to delicately punctiform, dark brown to blackish
or greyish black. Mycelium internal and external; supericial
hyphae branched, septate, 1.5–4(–6) µm wide, thin-walled,
subhyaline to brownish, verruculose. Stromata immersed, to
50 µm diam, brown. Conidiophores in small to moderately
large, loose fascicles, arising from stromata or solitary, arising
from supericial hyphae, erect, straight, subcylindrical-iliform
to slightly sinuous or subgeniculate, unbranched, (12.5–)60–
250 × 2.5–6 µm, occasionally with intercalary swellings, to
8 µm wide, often with percurrent rejuvenations giving raise
to new conidiogenous cells (visible as ine annellations),
pluriseptate, septa thin and not very conspicuous, wall thin
to somewhat thickened, brown, tips often paler, smooth;
conidiogenous cells integrated, terminal, 10–40 µm long,
apex of conidiogenous cells often somewhat swollen, with a
single to several distinct conidiogenous loci, often numerous
and aggregated, about 2 µm diam, slightly thickened and
darkened. Conidia solitary, ellipsoid-ovoid, cylindrical to
obclavate, 15–75(–105) × 3–9 µm, 0–7(–15)-septate, brown,
365
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Braun et al.
wall somewhat thickened, verrucose, apex obtuse, rounded,
base rounded to short obconically truncate, 1.5–2 µm wide,
hila almost unthickened or slightly so, somewhat darkenedrefractive.
Lectotype (designated here, MycoBank MBT178172):
India: Madhya Pradesh: Jabalpur, on Smilax ovalifolia
[macrophylla], 26 Dec. 1958, G. P. Agarwal 3 (K(M) IMI 75641).
Isolectotype: Mahakoshal Mahavidyalaya, Jabalpur, India,
Botany Department, Herbarium [now Gov. Model Science
College, Rani Durgawati University, Jabalpur] (unclear if still
maintained).
Host range and distribution: On Smilax
Smilacaceae, Asia (India, Madhya Pradesh).
ovalifolia,
Notes: See comments under Zasmidium indicum. Supericial
hyphae with solitary conidiophores were neither mentioned
in the original description nor depicted in the illustration, but
they are present in type material and abundant.
Zasmidium smilacis (P. Kumar, D.N. Shukla & Kamal)
Kamal, Cercosporoid Fungi of India: 248 (2010).
(Fig. 168)
Basionym: Stenella smilacis P. Kumar, D.N. Shukla & Kamal,
Curr. Sci. 49: 234 (1980).
Synonym: Veronaea smilacis R.P. Singh, Kamal & Abassi,
Curr. Sci. 50: 237 (1981) [holotype: India: U.P.:
Gorakhpur, Gorakhpur Forest Division, Madhaulia range,
on Smilax macrophylla, Feb.-Mar. 1978, R. P. Singh 310
(not preserved)].
Literature: de Hoog et al. (1983: 487), Braun & Castañeda
Ruiz (1991: 296).
Illustrations: Kumar et al. (1980: 235, ig. 1), Braun & Castañeda
Ruiz (1991: 295, ig. 31), Kamal (2010: 249, ig. 35).
Description: Leaf spots lacking or amphigenous, only formed
as pale yellowish discolorations, diffuse, or forming yellowish
to brown, irregularly shaped patches. Colonies hypophyllous,
olivaceous-brown. Mycelium internal and external, supericial
hyphae straight to lexuous, branched, 2–5 µm wide, septate,
pigmented, ± dimorphic, somewhat broader hyphae darker,
wall slightly thickened and smooth or almost so or hyphae
narrower, paler and verruculose. Conidiophores solitary,
arising from supericial hyphae, lateral and sometimes
terminal, erect, straight, subcylindrical-iliform to slightly
geniculate-sinuous, unbranched, long, 50–165 × 3–5.5
µm, 1–12-septate, brown, often somewhat paler towards
the tip, wall thin to somewhat thickened, smooth or almost
so; conidiogenous cells integrated, terminal, about 10–30
µm long, sympodial, with conspicuous conidiogenous loci,
often numerous and aggregated, somewhat thickened and
darkened, about 1 µm diam. Conidia solitary as well as
catenate, shape variable, ellipsoid-ovoid, subcylindrical,
fusiform, somewhat clavate or obclavate, 5–75 × 2–4 µm,
0–6-septate, hyaline, subhyaline to pale olivaceous, smooth
or almost so, ends rounded to truncate, 1–1.5 µm wide, hila
somewhat thickened and darkened.
366
Fig. 168. Zasmidium smilacis (K(M) IMI 227071). A. Supericial
hypha. B. Solitary conidiophores arising from supericial hyphae. C.
Conidia. Bar = 10 µm.
Holotype: India: U.P.: Gorakhpur, on Smilax ovalifolia
[macrophylla], Mar. 1978, P. Kumar 103 (K(M) IMI 227071).
Host range and distribution: On Smilax (ovalifolia, mollis,
perfoliata [prolifera]), Smilacaceae, Asia (India, Uttar
Pradesh), West Indies (Cuba).
Notes: Singh et al. (1981: 237, ig. 1) described and illustrated
Veronaea smilacis. Type material was said to be deposited
at IMI as “IMI 212616”, but this material was apparently not
retained (B. Aguirre-Hudson, K, in litt.). In the IMI accession
books, IMI 212616 is listed as discarded and the substrate
(host plant) is recorded as Coffea. It is quite unclear if the
authors deposited type material of V. smilacis at all, but that
there could be some error in the accession number cannot
be discounted. In any case, the description of this species
is close to Zasmidium smilacis (conidia 14–32.5 × 2–4 µm,
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
(1–)3–6(–12)-septate). This species is probably conspeciic
with Z. smilacis as already supposed by de Hoog et al. (1983).
Kamal (2010), one of the authors of V. smilacis, reduced this
name to synonym with Z. smilacis.
Zasmidium smilacis-macrophyllae (S. Chaudhary
& R.K. Chaudhary) Kamal, Cercosporoid Fungi of
India: 250 (2010).
(Fig. 169)
Basionym: Stenella smilacis-macrophyllae S. Chaudhary
& R.K. Chaudhary, in Rao et al., Frontiers of Fungal
Diversity in India (Prof. Kamal Festschrift): 601 (2003).
Illustration: Chaudhary & Chaudhary (2003: 602, ig. 7).
Description: Leaf spots amphigenous, circular, rarely
subcircular, centre black, margin dark brown. Colonies
hypophyllous, dark to blackish brown. Mycelium internal
and external; supericial hyphae branched, septate, 2–3
µm wide, olivaceous-brown, verruculose. Stromata present,
but only reduced to a few swollen substomatal cells.
Conidiophores emerging through stomata, in small, loose
fascicles and solitary, arising from supericial hyphae, lateral
or sometimes terminal, erect, straight to curved, lexuous,
subcylindrical, unbranched or rarely branched, 12–85 ×
3–4.5 µm, 1–4-septate, brown to dark brown, smooth;
conidiogenous cells integrated, terminal and intercalary,
with distinct conidiogenous loci. Conidia solitary, cylindrical
or subcylindrical, straight to somewhat curved, 15–45 ×
4–6.5 µm, 1–6-septate, occasionally somewhat constricted
at the septa, dark brown, wall thickened, verruculose, apex
rounded, base rounded to somewhat attenuated, with a
distinct hilum.
Holotype: India: Uttar Pradesh: Nichlaul Forest, on Smilax
ovalifolia [macrophylla], Smilacaceae, Sep. 1997, S.
Chaudhary (HCIO 43689). Isotype: GPU 9021.
Host range and distribution: Only known from the type
collection.
Typhaceae
Doubtful, excluded and insuficiently known
species
Cercospora typhae N.P. Golovina, Novosti Sist. Nizsh.
Rast. 1: 212 (1964).
Literature: Braun & Mel'nik (1997: 98), Crous & Braun (2003:
414).
Description: Leaf spots indistinct, oblong, brown or olivaceousbrown. Conidiophores 70–110 × 5–6 µm, unbranched,
olivaceous-brown, septate below. Conidia cylindrical to
obclavate, at irst one-celled, later 3–5-septate.
Fig. 169. Zasmidium smilacis-macrophyllae (based on Chaudhary &
Chaudhary 2003: 602, ig. 7). A. Supericial hypha. B. Conidiophores
and supericial hyphae emerging through a stoma. C. Conidiophores.
d. Conidia. Bar = 10 µm.
Host range and distribution: Only known from the type
collection.
Notes: This species is insuficiently known. The original
description is very meagre. Golovina (1964) cited that type
material of this species was deposited at the herbarium of
the Tashkent University. However, collections of TAK have
been transferred to TASH, but this does not refer to fungal
collections, which are lost according to Y. Gafforov, curator
of TASM.
Holotype: uzbekistan: Kurama District, on Typha angustifolia,
Typhaceae, N. P. Golovina (lost).
VOLUME 5 · NO. 2
367
Braun et al.
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Stromata immersed, intra- to subepidermal, often situated
in deep concave interveinal depressions of the leaves, to 70
µm diam and 30–60 µm deep, medium to dark brown, textura
angularis. Conidiophores numerous in dense to divergent
sporodochial fascicles, erumpent, erect, straight, cylindrical
to somewhat curved or geniculate-sinuous, unbranched,
20–45 × 3–4.5 µm, 0–2-septate, olivaceous-brown, paler
towards the tip, thin-walled, smooth; conidiogenous
cells integrated, terminal or conidiophores reduced to
conidiogenous cells, 10–20 µm long, proliferation sympodial
and percurrent, with to ive ine annellations, conidiogenous
loci truncate, 2.5–4.5 µm wide, neither thickened nor
darkened. Conidia solitary, cylindrical, subcylindrical or
somewhat tapering towards the tip, straight to curved or
slightly sinuous, about 65–90 × 4.5–5 µm, 3–5-septate, pale
olivaceous-brown, thin-walled, verruculose, apex obtuse,
base truncate to slighty obconically truncate, 3–4 µm wide,
hila unthickened, not darkened.
Holotype: Malawi: Mt Mulanje, Chambe to Lichenya track,
S1580, E03540, 2200 m alt, on Xerophyta splendens,
Velloziaceae, 16 Apr. 1991, B. C. Sutton MM156 (K(M) IMI
347226).
Host range and distribution: Only known from the type
collection.
Xanthorrhoeaceae
Cercospora
A single species.
Cercospora hemerocallidis Tehon, Mycologia 16:
139 (1924); as “hemerocallis”.
(Fig. 171)
Fig. 170. Pseudocercospora xerophytae (K(M) IMI 347226). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
Velloziaceae
Pseudocercospora
A single species.
pseudocercospora xerophytae B. Sutton, Mycol.
Pap. 167: 51 (1993).
(Fig. 170)
Illustration: Sutton (1993: 50–51, igs 12–13).
Description: Leaf spots at irst formed as small medium
brown lecks, 1 mm long, parallel to the leaf veins, later
enlarging to 5 × 1 mm, darker with diffuse lighter brown
border, oblong elliptical to navicular, sometimes conluent.
Caespituli amphigenous, not very conspicuous. Mycelium
immersed, intercellular; hyphae branched, septate, 2.5–4
µm wide, medium to dark brown, thin-walled, smooth.
368
Literature: Chupp (1954: 347), Saccardo (1972: 1375), Crous
& Braun (2003: 215).
Description: Leaf spots amphigenous, circular to subcircular
or somewhat irregular, 1–8 mm diam or even larger, at irst
pale greenish, later pale brownish to dingy grey in the
centre with darker margin or somewhat raised marginal
line, reddish to reddish brown, sometimes lower surface
more or less uniformly reddish to reddish brown. Caespituli
amphigenous, indistinct to inely punctiform, dark brown
to blackish. Mycelium internal. Stromata lacking or small,
substomatal, olivaceous-brown. Conidiophores solitary
to densely fasciculate, arising from stromata, emerging
through stomata, erect, straight, subcylindrical or somewhat
attenuated towards the tip, somewhat geniculatesinuous, unbranched, 10–80 × 2–6 µm, pale olivaceousbrown, yellowish brown, paler above, tip sometimes
subhyaline, continuous to pluriseptate, wall thin to slightly
thickened, smooth; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells,
conidiogenous loci distinct, thickened and darkened, about
1–2.5 µm diam. Conidia solitary, acicular to narrowly
obclavate, straight to somewhat curved or sinuous, 30–120
IMA FUNGUS
Cercosporoid fungi 2
Holotype: uSA: Illinois: Boomield, on Hemerocallis fulva, 25
Jul. 1922, P. A. Young 2897 (ILLS 2897).
Host range and distribution: On Hemerocallis (fulva,
Hemerocallis sp.), Xanthorrhoeaceae, North America (USA,
Florida, Illinois), West Indies (Bermuda).
Notes: The material examined from ILLS was in poor
condition, only with traces of Cercospora, and was dominated
by Alternaria and Cladosporium species. Records from China
on Hosta plantaginea (Hostaceae) are very doubtful.
ART I CLE
× 2–4 µm, indistinctly pluriseptate, hyaline or subhyaline,
thin-walled, smooth, apex subacute, base truncate to
obconically truncate, about 2–3 µm wide, hila somewhat
thickened and darkened.
Pseudocercospora
Key to Pseudocercospora species on Xanthorrhoeaceae
1
Mycelium internal; supericial hyphae with solitary conidiophores lacking; stromata well-developed,
25–80 µm diam; conidia (15–)20–50(–60) × 2–4 µm, 0–4-septate;
on Asphodelus spp. .......................................................................................................................... p. asphodelina
Mycelium internal and external; supericial hyphae with solitary conidiophores present; stromata smaller,
10–40 µm diam; conidia longer and pluriseptate, (20–)30–85 × 2–4 µm, (1–)3–12-septate;
on Dianella nigra ..................................................................................................................................... p. dianellae
Pseudocercospora species on Xanthorrhoeaceae
pseudocercospora asphodelina (Sacc.) U. Braun,
Mycotaxon 48: 293 (1993).
(Fig. 172)
Basionym: Septoria asphodelina Sacc., Syll. Fung. 3: 571
(1884).
Synonyms: Septoria aphodeli Westend., Bull. Acad. Roy.
Belg., Ser. II, 12: 573 (1857), non S. asphodeli Mont.,
1849.
Cylindrosporium asphodeli Kuhnh.-Lord. & J.P. Barry, Bull.
Trimestriel Soc. Mycol. France 65: 126 (1949) [holotype:
France: Hérault, Garrigue de la Gardiole, près Fabrègues,
on Asphodelus cerasifer, 11 May 1939, Kuhnholtz-Lordat
(not seen)].
Illustration: Braun (1993b: 295, ig. 22).
Fig. 171. Cercospora hemerocallidis (ILLS 2897). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
VOLUME 5 · NO. 2
Description: Leaf spots amphigenous, subcircular to irregular,
2–15 mm diam or conluent and larger, at irst yellowish,
ochraceous to brownish, later greyish brown, inally
greyish white, with darker margin. Caespituli amphigenous,
punctiform, dark brown to almost black. Mycelium internal;
hyphae septate, branched, 1–4 µm wide, subhyaline to
faintly pigmented. Stromata well-developed, 25–80 µm diam,
brown, substomatal. Conidiophores numerous, in dense
fascicles, arising from stromata, most of the conidiophores
immersed in the substomatal cavity, barely emerging through
stomata, straight, subcylindrical to lexuous, unbranched,
10–25 × 2–4 µm, 0(–1)-septate, pale, subhyaline to
greenish or olivaceous, thin-walled, smooth; conidiophores
mostly reduced to conidiogenous cells, conidiogenous loci
inconspicuous, unthickened and not darkened. Conidia
solitary, subcylindrical-acicular, obclavate, (15–)20–50(–
60) × 2–4 µm, 0–4-septate, subhyaline to greenish, thinwalled, smooth, apex subacute, base truncate to somewhat
obconically truncate, 1–2 µm wide, hila neither thickened not
darkened.
369
ART I CLE
Braun et al.
Fig. 172. Pseudocercospora asphodelina (BR, holotype). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
Holotype: Belgium: on Asphodelus albus, without date, R. P.
Cl. Dumont (BR).
Host range and distribution: On Asphodelus (albus, cerasifer,
microcarpus), Xanthorrhoeaceae, Europe (Belgium, France,
Spain).
Note: This speces is rather pseudocercosporella-like, but the
conidiophores are not quite colourless.
pseudocercospora dianellae U. Braun & C.F. Hill, in
Braun et al., Australas. Pl. Pathol. 32: 89 (2003).
(Fig. 173)
Illustration: Braun et al. (2003b: 90, ig. 3).
Exsiccatae: U. Braun, Fungi Sel. Exs. 15 (topotype material).
Description: Leaf spots lacking or almost so, or welldeveloped, amphigenous, shape and size variable, elliptical
to irregular, oblong, to about 40 × 10 mm, brown, sometimes
reddish brown, margin indeinite, old patches becoming
pale brown to greyish white, with an irregular dark border.
Caespituli hypophyllous, punctiform, scattered, dark brown
to blackish, later greyish brown. Mycelium internal and
external; supericial hyphae sparingly branched, septate,
1.5–4 µm wide, subhyaline to pale olivaceous-brown,
thin-walled, smooth. Stromata lacking to well-developed,
substomatal to intraepidermal, 10–40 µm diam, olivaceousbrown. Conidiophores in small to moderately large fascicles,
loose to moderately dense, arising from stromata, through
stomata or erumpent, also solitary, arising from supericial
hyphae, lateral, erect, occasionally decumbent, straight,
subcylindrical-conical to strongly geniculate-sinuous,
unbranched, 5–40 × 2–4 µm, 0–3-septate, subhyaline to
pale olivaceous or olivaceous-brown, thin-walled, smooth;
conidiogenous cells integrated, terminal or conidiophores
370
Fig. 173. Pseudocercospora dianellae (HAL 1729 F). A. Conidiophore
fascicles. B. Solitary conidiophores arising from supericial hyphae.
C. Conidiophores. d. Conidia. Bar = 10 µm.
reduced to conidiogenous cells, 5–25 µm long, conidiogenous
loci inconspicuous. Conidia solitary, obclavate-cylindrical,
subacicular, (20–)30–85 × 2–4 µm, (1–)3–12-septate,
subhyaline to pale olivaceous, thin-walled, smooth, apex
subacute to obtuse, base truncate to obconically truncate,
(1–)1.5–2.5(–3) µm wide, hila unthickened, not darkened.
Holotype: New Zealand: Auckland, Grey Lynn, Western
Springs Park, on Dianella nigra, 31 Mar. 2001, C. F. Hill 379
(HAL 1729 F). Isotype: PDD 75122.
Host range and distribution:
Xanthorrhoeaceae, New Zealand.
On
Dianella
nigra,
Zasmidium
A single species.
Zasmidium dianellae (Sawada & Katsuki) U. Braun,
Stud. Mycol. 72: 336 (2012).
(Fig. 174)
Basionym: Cercospora dianellae Sawada & Katsuki, Spec.
Publ. Coll. Agric. Natl. Taiwan Univ. 8: 216 (1959).
Synonyms: Heterosporium dianellae Sawada, Rep. Gov. Res.
Inst. Formosa 87: 76 (1944), nom. inval. (ICN, Art. 39.1)
[holotype: taiwan: Taipei, on Dianella ensifolia, 19 Nov.
1925, K. Sawda (NTU-PPE, National Taiwan University,
herb. Sawada)].
Stenella dianellae (Sawada & Katsuki) Goh & W.H. Hsieh,
Trans. Mycol. Soc. Republ. China 2: 137 (1987).
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
none to well-developed, globose, 40–125 µm diam, dark
brown. Conidiophores in small to moderately large fascicles,
to 20, loose, arising from stromata, or solitary, arising from
supericial hyphae, lateral, rarely terminal, erect, straight,
subcylindrical, slightly geniculate, unbranched, 20–75 × 2.5–4
µm, 0–3-septate, uniformly medium to dark reddish brown or
paler towards the apex, thin-walled, smooth; conidiogenous
cells integrated, terminal, conidiogenous loci thickened
and darkened, about 1.5–2.5 µm diam. Conidia solitary,
subcylindrical to obclavate-cylindrical, straight to somewhat
curved, 10–60 × 3–4.5 µm, 0–6-septate, subhyaline to
pale brownish, thin-walled, inely verruculose, ends bluntly
rounded, hila somewhat thickened and darkened.
Holotype: taiwan: Taipei, on Dianella ensifolia, 19 Nov. 1925,
K. Sawada (NTU-PPE, National Taiwan University, herb.
Sawada). Isotype: TNS-F-220610.
Host range and distribution: On Dianella
Xanthorrhoeaceae, Asia (Taiwan), New Zealand.
ensifolia,
Notes: Records of this species on Phormium sp., Phormiaceae
(Crous & Braun 2003) are unveriied.
Doubtful, excluded and insuficiently known
species
Cercospora phormii Bolick, nom. nud., ide Alieri et al.
(1984) and Farr et al. (1989: 615), on Phormium tenax,
USA, Florida.
Xyridaceae
Cercospora
Doubtful, excluded and insuficiently known
species
Cercospora xyridis Miles, Mycologia 18: 168 (1926).
Fig. 174. Zasmidium dianellae (NTU-PPE, holotype). A. Solitary
conidiophores arising from supericial hyphae. B. Conidiophore
fascicle. C. Conidia. Bar = 10 µm.
Literature: David (1997: 108–110), Goh & Hsieh (1990: 209),
Crous & Braun (2003: 159).
Illustrations: David (1997: 110, ig. 31), Goh & Hsieh (1990:
210, ig. 162).
Description: Leaf spots amphigenous, elliptical to fusiform,
scattered, 0.5–6 mm long, sometimes conluent and larger,
coalescing into elongated patches, at irst purplish red, later
dingy grey. Caespituli epiphyllous, punctiform-pustulate,
blackish, scattered to aggregated. Mycelium internal and
external; supericial hyphae sparingly branched, septate,
very pale olivaceous to brownish, 1.5–3.5 µm wide, thinwalled, pale and narrower hyphae verruculose. Stromata
VOLUME 5 · NO. 2
Literature: Chupp (1954: 606), Saccardo (1972: 1388), Crous
& Braun (2003: 432).
Description: Leaf spots oval, 2–4 mm diam, brown, margin
purplish brown. Caespituli amphigenous. Mycelium internal.
Stromata lacking or only small aggregations of a few swollen
hyphal cells, substomatal, dark brown, wall thickened.
Conidiophores solitary or fasciculate, 2–8, straight to slightly
sinuous, rarely geniculate, unbranched, 50–150 × 4–5.5 µm,
plainly pluriseptate, uniformly dark reddish brown. Conidia
solitary, obclavate, 40–90 × 2.5–4 µm, indistinctly septate,
subhyaline to very pale olivaceous, thin-walled, smooth, apex
subobtuse, base obconically truncate.
Syntypes: uSA: Mississippi: Wiggans, on Xyris difformis
[elata], 15 Sep. 1920, L. E. Miles 455 (BPI 442562, CUP
41582, NY 945737).
Host range and distribution: On Xyris difformis, Xyridaceae,
North America (USA, Mississippi).
371
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Braun et al.
Notes: Status unclear; type material very meagre, only dark
brown substomatal stromata and conidiophore fragments
found. The conidiophores seem to be cercospora-like, but
conidiogenous cells and loci as well conidia not seem. New
collections are necessary to prove the generic afinity of this
species.
Zingiberaceae
Cercospora
Key to Cercospora species on Zingiberaceae
1
Conidia acicular, base truncate or at least long acicular conidia mixed with
obclavate-subcylindrical conidia .............................................................................................................................. 2
Conidia obclavate-cylindrical, base short obconically truncate, acicular conidia not formed .......................................... 6
2 (1)
Conidia obclavate-cylindrical to acicular, rather broad, 40–160 × 4.5–6.5 µm; on Alpinia ........................... p. alpinigena
Conidia narrower, 2–4 µm ............................................................................................................................................... 3
3 (2)
Conidia acicular; on Alpinia or Curcuma spp. ................................................................................................................. 4
Conidia acicular, shorter conidia obclavate-cylindrical; on Stahlianthus or Zingiber spp. ............................................... 5
4 (3)
Conidiophores very long, 100–265 µm; conidia 165–335 µm long; on Alpinia ......................... C. alpiniae-katsumadae
Conidiophores shorter, 50–190 µm; conidia 40–130 µm long; on Curcuma ............................................... C. hitcheniae
5 (3)
Conidiophores 20–260 µm long; on Stahlianthus spp. ............................................................................... C. stahlianthi
Conidiophores shorter, 15–90 µm; on Zingiber spp. ............................................................................... C. zingibericola
6 (1)
Conidia rather broad, 40–160 × 3.5–6.5 µm, obclavate-cylindrical to acicular; on Alpinia spp. .................. C. alpinigena
Conidia obclavate-cylindrical, acicular conidia lacking, much narrower, 1.5–4 µm, if broader,
3.5–5 µm, pale, not quite colourless ........................................................................................................................ 7
7 (6)
Stromata lacking or only with a few substomatal swollen hyphal cells; conidiophores longer,
40–100 µm; conidia broader, 25–90 × 3.5–5 µm, 3–5-septate, subhyaline; on Zingiber spp. ................ C. zingiberi
Stromata well-developed, about 20–65 µm diam; conidia narrower, 1.5–4 µm; on other hosts ..................................... 8
8 (7)
Conidia quite colourless, 3–4 µm wide; on Curcuma ................................................................................... C. curcumae
Conidia subhyaline to pale olivaceous, 1.5–3.5 µm wide; on Alpinia ........................................................... p. alpiniicola
Cercospora species on Zingiberaceae
Cercospora alpiniae-katsumadae S.Q. Chen &
P.K. Chi, J. S. China Agric. Univ. 11: 58 (1990); as
“alpini-katsumadae”.
(similar to Fig. 1)
Literature: Chi (1994: 157), Crous & Braun (2003: 52), Guo et
al. (2005: 308–309).
Illustrations: Chi (1994: 24, ig. 11), Guo et al. (2005: 309,
ig. 231).
Description: Leaf spots amphigenous, elliptical to irregular,
greyish white with brown border. Caespituli epiphyllous.
Mycelium internal. Stromata globose, 20–65 µm diam,
olivaceous-brown to brown. Conidiophores in fascicles, 5–25,
arising from stromata, erect, straight, cylindrical, to somewhat
curved, 0–3 times geniculate above, unbranched, 100–265 ×
3.5–5 µm, 3–13-septate, olivaceous-brown; conidiogenous
cells integrated, terminal and intercalary, conidiogenous
372
loci conspicuous, thickened and darkened. Conidia solitary,
acicular, 165–335 × 2.5–3.5 µm, 10–20-septate, hyaline, thinwalled, smooth, apex pointed, base truncate or subtruncate,
hila thickened and darkened.
Holotype: China: Guangdong: Yunan, on Alpinia hainanensis
[katsumadae] (Hb. S. China Agric. Univ., Guangzhou).
Host range and distribution: On Alpinia (hainanensis,
oxyphylla), Zingiberaceae, Asia (China).
Notes: This species has strictly acicular conidia and belongs
to the C. apii s. lat. complex.
Cercospora alpinigena To-anun, Meeboon & Hidayat,
sp. nov.
MycoBank MB809027
(Fig. 175)
Literature: To-anun et al. (2011: 82), as C. alpiniicola.
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
and intercalary, conidiogenous loci conspicuous, thickened
and darkened, 1.3–2.8 µm diam. Conidia solitary, obclavatesubcylindrical to subacicular, about 40–160 × 3.5–6.5 µm,
4–11-septate, hyaline, thin-walled, smooth, apex subacute or
subobtuse, base short obconically truncate, 2–2.5 µm wide,
hila thickened and darkened.
Holotype: thailand: Chiang Mai Province: Sanpatong
District, Mae Wang Sub-district, Tambol Mae Win, Bahn
Mae Sapok, Mae Sapok Royal Project, on Alpinia purpurata,
Zingiberaceae, 8 Feb. 2008, J. Meeboon & I. Hidayat (BBH
23684).
Host range and distribution: Only known from the type
collection.
Notes: The collection on Alpinia purpurata was previously
assigned to Cercospora alpiniicola, but differs in having
colourless, much broader, obclavate-cylindrical to subacicular
conidia and warrants to be considered a species of its own.
Cercospora alpiniicola S.Q. Chen & P.K. Chi, J. S.
China Agric. Univ. 11: 57 (1990); as “alpinicola”.
(Fig. 176)
Literature: Chi (1994: 33), Crous & Braun (2003: 53), Guo et
al. (2005: 308).
Illustrations: Chi (1994: 24, ig. 11), Guo et al. (2005: 308,
ig. 230).
Illustration: To-anun et al. (2011: 83, ig. 66), as C. alpiniicola.
Description: Leaf spots amphigenous, elliptical, pale
brown with narrow brown margin. Caespituli amphigenous.
Stromata 23–67 µm diam, olivaceous-brown. Conidiophores
fasciculate, 10–55, arising from stromata, erect,
subcylindrical, straight to geniculate, unbranched or rarely
branched, 40–65 × 3–3.6 µm, 2–5-septate, olivaceous;
conidiogenous cells integrated, terminal or intercalary,
with conspicuous conidiogenous loci, thickened and
darkened. Conidia solitary, obclavate, 40–80 × 1.5–3.5 µm,
3–11-septate, subhyaline to pale olivaceous, thin-walled,
smooth, apex obtuse, base short obconically truncate, hila
thickened and darkened.
Diagnosis: Morphologically resembling Cercospora alpiniicola
but conidia obclavate-cylindrical to subacicular, hyaline and
much broader, 3.5–6.5 µm,
Holotype: China: Guangdong: Gaoyao, on Alpinia oxyphylla,
Zingiberaceae, Apr. 1987, S. Q. Chen 103 (Hb., S. China
Agric. Univ., Guangzhou).
Description: Leaf spots amphigenous, subcircular to angular,
2–14 mm diam, vein-limited, conluent, brown to greyish
brown or centre becoming white with dark margin sometimes
forming distinct discolorations around the margin. Caespituli
amphigenous, mainly epiphyllous. Mycelium internal.
Stromata small to well-developed, about 10–40 µm, brown
to dark brown, composed of globose or subglobose cells.
Conidiophores in loose to dense fascicles, 3–8, arising
from stromata, erect, straight to decumbent, subcylindrical
to distinctly geniculate, unbranched, 48.5–100 × 4.5–6.5
µm, 1–3-septate, brown below, paler towards the tip, thinwalled, smooth; conidiogenous cells integrated, terminal
Host range and distribution: Only known from the type
collection.
Fig. 175. Cercospora alpinigena (BBH 23684). A. Conidiophore
fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.
VOLUME 5 · NO. 2
Notes: Type material could not be examined, but on account
of the conspicuous conidiogenous loci and obclavate, pale
olivaceous conidia, this species might rather belong to
Passalora. To-anun et al. (2011: 82) recorded this species
from Thailand on Alpinia purpurata, which differs, however,
in having acicular to obclavate-cylindrical, hyaline, broader
conidia and represents a distinct species of its own (see C.
alpinigena).
373
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Braun et al.
Fig. 177. Cercospora curcumae (based on Govindu & Thirumalachar
1956: plate IX, ig. 17). A. Conidiophore fascicle. B. Conidiophore tip.
C. Conidia. Bar = 10 µm.
Fig. 176. Cercospora alpiniicola (based on Chi 1994: 24, ig. 11). A.
Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
Cercospora curcumae Govindu & Thirum., Sydowia
10: 275 (1956).
(Fig. 177)
Literature: Vasudeva (1963: 94), Crous & Braun (2003: 148).
from stromata, erect, straight to usually distinctly geniculatesinuous, unbranched or rarely 1–2 times branched, about 15–70
× 3–5.5 µm, 1–6-septate, olivaceous-brown, paler towards the
apex, ultimate tips subhyaline; conidiogenous cells integrated,
terminal, with conspicuous conidiogenous loci, thickened and
darkened. Conidia solitary, obclavate-cylindrical, straight to
slightly curved, about 30–70 × 3–4 µm, 1–10-septate, hyaline,
thin-walled, smooth, apex pointed, base short obconically
truncate, hila thickened and darkened.
Illustration: Govindu & Thirumalachar (1956: plate IX, ig. 17).
Description: Leaf spots amphigenous, irregular, 3–5 mm diam,
dark brown to almost sooty black. Caespituli amphigenous.
Mycelium internal. Stromata globose, compact, 20–50 µm diam,
dark brown. Conidiophores loosely to densely fasciculate, arising
374
Holotype: India: Karnataka: Bangalore, Hebbal, on Curcuma
longa, 12 Apr. 1954, H. C. Govindu (probably not preserved).
Host range and distribution: On Curcuma (aromatica, longa),
Zingiberaceae, Asia (India, Myanmar, Nepal).
IMA FUNGUS
Cercosporoid fungi 2
Illustration: Chiddarwar (1960: 117, igs 4–6), Pavgi &
Upadhyay (1968: plate X, igs 4–6), Srivastava et al. (1994:
106, ig. 2).
ART I CLE
Literature: Crous & Braun (2003: 148, 218), Kamal (2010: 37,
51), Mall et al. (2013).
Description: Leaf spots amphigenous, numerous, circular
to angular-irregular, small, 2–5 mm diam, scattered,
occasionally conluent, brown. Caespituli amphigenous.
Mycelium internal. Stromata substomatal, about 15–60 µm
diam, brown to dark brown. Conidiophores in loose fascicles,
mostly 3–10, rarely solitary, arising from stromata, emerging
through stomata, erect, straight to curved, unbranched,
not to distinctly geniculate, about 40–130 × 3–6 µm,
2–6-septate, olivaceous-brown to brown, wall somewhat
thickened, smooth; conidiogenous cells integrated, terminal
and intercalary, sympodial, conidiogenous loci conspicuous,
about 2–2.5 µm diam, thickened and darkened. Conidia
solitary, acicular, shorter ones sometimes subcylindrical,
straight to curved, 50–190 × 2–4 µm, 3–19-septate, hyaline,
thin-walled, smooth, apex pointed, base truncate, about 1.5–
2.5 µm wide, hila thickened and darkened.
Holotype: India: Maharashtra: Mahabaleshwar, on Curcuma
caulina [Hitchenia caulina], Zingiberaceae, 19 Jan. 1955, P.
P. Chiddarwar (K(M) IMI 83175).
Host range and distribution: On Curcuma (angustifolia,
caulina, longa [domestica]), Zingiberaceae, Asia (India,
Madhya Pradesh, Uttar Pradesh; Nepal).
Notes: In the original paper, it was mentioned that portions of
the type of Cercospora cucurmae-longae were deposited at
HCIO and IMI, but it was not possible to locate any material
under this name at IMI (now at K). According to the original
description and illustration, this species belongs undoubtedly
to the Cercospora apii s. lat. complex. Kamal (2010)
examined type material of C. curcumina and stated that this
species is conspeciic with C. curcumae-longae. On the other
hand, the latter species is morphologically indistinguishable
from C. hitcheniae described from India on Curcuma caulina
[Hitchenia caulina], which is also a member of the C. apii s.
lat. complex.
Fig. 178. Cercospora hitcheniae (K(M) IMI 83175). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
Cercospora hitcheniae Chidd., Indian Phytopathol.
12: 113 “1959”(1960).
(Fig. 178)
Synonyms: Cercospora cucurmae-longae Pavgi & R.
Upadhyay, Sydowia 21: 102 “1967” (1968) [holotype:
India: Uttar Pradesh: Varanasi, on Curcuma longa, 3 Jan.
1964, R. Upadhyay (MSP, no. 341)].
Cercospora curcumina R.K. Srivast., N. Srivast. & A.K.
Srivast., Proc. Natl. Acad. Sci. India, Sect. B, Biol. Sci.
64: 107 (1994) [holotype: India: Madhya Pradesh: Sagar,
on Curcuma angustifolia, Aug. 1989, R. K. Srivastava 66
(GPU 1341); isotype: HCIO 30881].
VOLUME 5 · NO. 2
Cercospora stahlianthi Z.D. Jiang & P.K. Chi, in Chi,
Fungal Diseases of Medical Plants in Guangdong
Province: 162 (1994).
(Similar to Fig. 1)
Literature: Crous & Braun (2003: 386), Phensintham et al.
(2013b: 74), Guo et al. (2005: 309).
Illustrations: Chi (1994: 162, ig. 167), Guo et al. (2005: 310,
ig. 232), Phengsintham et al. (2013b: 75, igs 48–49).
Description: Leaf spots amphigenous, circular to irregular,
1–5 mm diam, pale to dark brown with yellowish to medium
brown margin. Caespituli amphigenous, punctiform, dark
brown, scattered. Mycelium internal; hyphae branched,
375
ART I CLE
Braun et al.
septate, with constrictions at the septa, pigmented, 2–6 µm
wide. Stromata substomatal, 10–70 µm diam, brown, cells
subglobose to angular-irregular in outline, 5–17 µm diam.
Conidiophores in small to moderately large fascicles, 2–27,
loose to dense, arising from stromata, though stomata,
unbranched, subcylindrical to somewhat geniculatesinuous, 20–260 × 4–6 µm, rarely longer, to 400 µm, 1- to
pluriseptate, brown, wall thin to somewhat thickened, smooth;
conidiogenous cells integrated, terminal and intercalary, 15–
65 µm long, conidiogenous loci thickened and darkened, 2–4
µm diam. Conidia solitary, acicular to somewhat obclavatesubcylindrical, 30–240 × 2–4 µm, 2–23-septate, colourless,
thin-walled, apex pointed, base truncate to obconically
truncate, 1.5–3 µm wide, thickened and darkened.
Holotype: China: Guangdong: Gaoming, on Stahlianthus
involucrata, Sep. 1986, Z. D. Jiang, no. 303 (SCHM [Hb.
South China Agriculture University, Guangzhou]).
Host range and distribution: On Stahlianthus (involucrata,
thorelii), Zingiberaceae, Asia (China, Laos).
Cercospora zingibericola A.K. Kar & M. Mandal,
Trans. Brit. Mycol. Soc. 53: 359 (1969).
(Fig. 179)
Literature: Crous & Braun (2003: 434), Kamal (2010: 99).
Illustration: Kar & Mandal (1969: 359, ig. 21).
Description: Leaf spots amphigenous, oblong between
veins to irregular, 0.5–3 mm diam, scattered to conluent,
greyish, margin indeinite. Caespituli amphigenous, mainly
hypophyllous, dark, punctiform. Mycelium internal. Stromata
small to moderately developed, 15–50 µm diam, subglobose
to irregular, pale to dark blackish brown. Conidiophores
loosely fasciculate, 2–16(–26), arising from stromata, erect,
straight, subcylindrical to geniculate-sinuous, unbranched
or rarely branched, 15–90 × 3.5–5.5 µm, 0–16-septate,
subhyaline to pale brown, paler towards the tip, smooth;
conidiogenous cells integrated, terminal, occasionally
intercalary or conidiophores reduced to conidiogenous cells,
about 10–30 µm long, conidiogenous loci conspicuous, 1–5
per cell, about 1.5–2.5 µm diam, thickened and darkened.
Conidia solitary, acicular, shorter ones sometimes obclavatecylindrical, 20–220 × 2–3.5 µm, 2–20-septate, hyaline, thinwalled, smooth, apex pointed to subobtuse, base truncate to
obconically truncate in short conidia, 1.5–2.5 µm wide, hila
thickened and darkened.
Holotype: India: West Bengal: Murshidabad, Khargram, on
Zingiber oficinale, 1 Feb. 1967, A. K. Kar & M. Mandal (K(M)
IMI 135186). Isotype: BPI 442646.
Host range and distribution: On Zingiber (oficinale, Zingiber
sp.), Zingiberaceae, Africa (Mauritius), Asia (Borneo, China
[Hong Kong], India, Myanmar).
Note: A true Cercospora s. str. belonging to the C. apii s. lat.
complex.
376
Fig. 179. Cercospora zingibericola (K(M) IMI 135186). A.
Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
Cercospora zingiberis Togashi & Katsuki, Bot. Mag.
Tokyo 65: 25 (1952); as “zingiberi”.
(Fig. 180)
Literature: Chupp (1954: 608), Katsuki (1965: 69), Crous &
Braun (2003: 434), Kamal (2010: 99).
Description: Leaf spots oblong, linear, between veins, 2–20
× 0.5–2 mm, occasionally conluent, pale brown to medium
brown, later greyish brown, often vein-limited, margin indistinct.
Caespituli mainly hypophyllous, punctiform, scattered, dark
IMA FUNGUS
Cercosporoid fungi 2
Holotype: Japan: Fukuoka Pref.: Takawa, Soeda, on Zingiber
mioga, 13 Sep. 1949, S. Katsuki (TNS-F-243831). Isotype:
CUP 40848.
ART I CLE
× 3.5–5 µm, 3–7-septate, subhyaline, thin-walled, smooth,
apex subacute or subobtuse, base short obconically truncate,
1.5–2 µm wide, hila thickened and darkened.
Host range and distribution: On Elettaria cardamomum,
Zingiber (mioga, oficinale), Zingiberaceae, Asia (India,
Karnataka; Japan), North America (USA, Florida).
Notes: The description of C. zingiberis in Nair (2011: 215)
with shorter conidiophores, 17.5–56 × 3.5 µm, and much
longer and narrowly linear conidia, 37–195 × 1.75–2.5 µm, is
undoubtedly based on a misidentiication and seems rather
to refer to C. zingibericola.
Passalora
Doubtful, excluded and insuficiently known
species
Passalora curcumae Purkay. & Mallik, Nova Hedwigia,
Beih. 63: 132 (1979).
Literature: Crous & Braun (2003: 450).
Illustration: Purkayastha & Mallik (1979: 131, Fig. 4).
Fig. 180. Cercospora zingiberis (CUP 40848). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
brown to blackish. Mycelium internal. Stromata lacking or
only formed as substomatal aggregations of swollen hyphal
cells, to 25 × 10 µm, dark brown. Conidiophores in divergent
to moderately dense fascicles, 4–15, effuse, erect, straight,
cylindrical, unbranched, barely geniculate-sinuous, 40–100 ×
3–8 µm, 3–6-septate, pale to medium brown or olivaceousbrown, tapered and paler towards the tip, wall thin, to about
0.8 µm, smooth; conidiogenous cells integrated, terminal, 10–
30 µm long, conidiogenous loci conspicuous, thickened and
darkened, 1–5, often aggregated at the tip, not or somewhat
protuberant, 1.5–2.5 µm diam. Conidia solitary, acicular to
obclavate-cylindrical, straight to somewhat curved, 25–90
VOLUME 5 · NO. 2
Description: Leaf spots amphigenous, circular to oval,
uniformly scattered, 3–10 mm diam, greyish white
with dark brown border, somewhat raised. Caespituli
hypophyllous. Mycelium internal. Stromata lacking or almost
so. Conidiophores in small fascicles, emerging through
stomata, erect, straight, subcylindrical, unbranched, about
110–240 × 4–9 µm, 2–6-septate, brown, swollen at the very
base; conidiogenous cells integrated, terminal, percurrent,
conidiogenous loci conspicuous, thickened. Conidia solitary,
obclavate-obovoid, 30–45 × 5–8 µm, (0–)1(–2)-septate,
olivaceous-brown, thin-walled, smooth, apex attenuated,
base obconically truncate, hila thickened.
Holotype: India: West Bengal: Baruipur, 24 Parganas,
on Curcuma reclinata, Zingiberaceae, 15 Dec. 1975 (not
preserved).
Host range and distribution: Only known from the type
collection.
Notes: Identity and status of this species are unclear. Type
material was said to be deposited at IMI (IMI 199984), but
could not be located. In the accession book of IMI, number
199984 is listed as discarded. It is rather doubtful that P.
curcumae represents a true Passalora.
377
Braun et al.
ART I CLE
Pseudocercospora
Key to Pseudocercospora species on Zingiberaceae
1
Conidiogenous cells rhachis-like, straight, subcylindrical, neither geniculate nor sinuous,
but with numerous distinct conical denticles, dactylaria-like; on Alpinia oficinarum
.................................................................. p. alpiniicola, see Doubtful, excluded and insuficiently known species]
Conidiogenous cells non-denticulate, at most subdenticulate, but then conidiogenous cells
geniculate-sinuous ................................................................................................................................................... 2
2 (1)
Stromata very large, 40–120 µm diam; conidiophores numerous, dense, in sporodochial conidiomata,
very short, 8–25 µm, 0–1-septate; conidia narrowly subacicular-iliform or long and
narrowly fusiform, 40–190 × 1.5–3 µm, subhyaline to pale yellowish; on Zingiber ................................ p. zingiberis
Stromata much smaller, 10–65 µm diam; conidiophores much longer, 10–150 µm
(if uniformly short, conidia obclavate and colourless), often with several septa;
conidia obclavate-cylindrical, pale olivaceous to olivaceous-brown ........................................................................ 3
3 (2)
Conidiophores very short, 3–10 µm; conidia obclavate, colourless; on Alpinia hainanensis
....................................................................................................................................... p. alpiniae-katsumadaicola
Conidiophores much longer, 10–150 µm; conidia pigmented ......................................................................................... 4
4 (3)
Mycelium in vivo internal and external; supericial hyphae with solitary conidiophores developed;
on Hedychium spp. .................................................................................................................................. p. hedychii
Mycelium in vivo internal; supericial hyphae and solitary conidiophores not developed;
on other hosts .......................................................................................................................................................... 5
5 (4)
Conidiophores long, 65–145 µm, fertile upper part distinctly and densely raduliform and annellate;
on Alpinia .................................................................................................................................................. p. alpiniae
Fertile part of conidiophores neither raduliform nor annellate, proliferation only sympodial;
on other hosts .......................................................................................................................................................... 6
6 (5)
Conidiophores very long, 30–150 µm, pluriseptate; on Etlingera .................................................................. p. nicolaiae
Conidiophores shorter, 10–75 µm, 0–6-septate; on other hosts ..................................................................................... 7
7 (6)
Conidiophores 10–75 × 3–5.5 µm; conidia 25–95 × 2.5–4.5 µm, 2–9-septate; on Amomum ............................ p. amomi
Conidiophores 12–26 × 2–3.2 µm; conidia 40–80 × 2–2.5 µm; on Curcuma ........................................... p. curcumicola
Pseudocercospora species on Zingiberaceae
pseudocercospora alpiniae S.Q. Chen & P.K. Chi, J.
S. China Agric. Univ. 11: 47 (1990).
(Fig. 181)
378
40–75 × 3–3.5 µm, 5–16-septate, pale olivaceous, thin-walled,
smooth, apex obtuse, base short obconically truncate, about 1
µm wide, hila unthickened, not darkened.
Literature: Chi (1994: 48), Guo et al. (1998: 402).
Holotype: China: Guangdong: Yunan, on Alpinia oficinarum,
Zingiberaceae, Oct. 1986, S. Q. Chen 105 (Hb. S. China
Agric. Univ., Guangzhou).
Illustrations: Chi (1994: 159, ig. 164), Guo et al. (1998: 403,
ig. 339).
Host range and distribution: Only known from the type
collection.
Description: Leaf spots amphigenous, irregular, greyish white
with brown border. Caespituli amphigenous. Mycelium internal.
Stromata small, subglobose, about 15–35 µm diam, olivaceousbrown. Conidiophores fasciculate, 5–15, divergent to dense,
arising from stromata, erect, straight, cylindrical, lexuous,
unbranched, fertile apex densely raduliform or annellate,
about 65–145 × 4.5–5 µm, 3–16-septate, olivaceous or pale
olivaceous, thin-walled, smooth; conidiogeous cells integrated,
terminal, narrowed towards the tip, sympodial and percurrent,
conidiogenous loci about 1 µm wide, unthickened, not darkened.
Conidia solitary, obclavate, straight to somewhat curved, about
pseudocercospora alpiniae-katsumadaicola (S.Q.
Chen & P.K. Chi) U. Braun & Crous, Mycosphaerella
and Anam.: 53 (2003).
(Fig. 182)
Basionym: Cercospora alpiniae-katsumadaicola S.Q. Chen &
P.K. Chi, J. S. China Agric. Univ. 11: 58 (1990); as “alpinikatsumadaicola”.
Synonym: Pseudocercospora alpiniae-katsumadaicola (S.Q.
Chen & P.K. Chi) P.K. Chi, Fungal Diseases of Cultivated
Medical Plants in Guangdong Province: 159 (1994),
comb. inval. (ICN, Art. 41.5).
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 182. Pseudocercospora alpiniae-katsumadaicola (based on Chi
1994: 159, ig. 164). A. Conidiophore fascicle. B. Conidia. Bar = 10
µm.
Fig. 181. Pseudocercospora alpiniae (based on Chi 1994: 48, ig.
29). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar
= 10 µm.
Literature: Guo et al. (1998: 404).
Illustrations: Chi (1994: 159, ig. 164), Guo et al. (1998: 404,
ig. 341).
Description: Leaf spots amphigenous, circular to elliptical,
grey-brown with yellowish to brown border. Caespituli
epiphyllous. Mycelium internal. Stromata small, globose,
20–50 µm diam, olivaceous. Conidiophores in small to
large fascicles, 5–50, arising from stromata, erect, straight,
subcylindrical to somewhat conical, 0–1-geniculate,
unbranched, short, 3–10 × 2.7–3.3 µm, 0–1-septate, pale
olivaceous or olivaceous, thin-walled, smooth; conidiophores
usually reduced to conidiogenous cells, conidiogenous loci
VOLUME 5 · NO. 2
indistinct, about 1 µm diam, unthickened, not darkened.
Conidia solitary, narrowly obclavate, 33–77 × 2.7–3.3 µm,
4–10-septate, hyaline, thin-walled, smooth, apex obtuse,
base short obconically truncate, about 1 µm wide, hila
unthickened, not darkened.
Holotype: China: Guangdong: Yunan, on Alpinia hainanensis
[katsumadae], Zingiberaceae, Oct. 1987, S. Q. Chen & P. K.
Chi, no. 117 (Hb. S. China Agric. Univ., Guangzhou).
Host range and distribution: Only known from the type
collection.
pseudocercospora amomi (A.K. Kar & M. Mandal)
Deighton, Trans. Brit. Mycol. Soc. 88: 388 (1987).
(Fig. 183)
Basionym: Cercospora amomi A.K. Kar & M. Mandal, Trans.
Brit. Mycol. Soc. 53: 358 (1969).
Literature: Guo et al. (1998: 405), Crous & Braun (2003: 55),
Kamal (2010: 148).
379
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Braun et al.
Fig. 184. Pseudocercospora curcumicola (TNS-F-60890). A.
Conidiophore fascicle. B. Conidia. Bar = 10 µm. U. Braun del.
Fig. 183. Pseudocercospora amomi (K(M) IMI 135184). A.
Conidiophore fascicles. B. Conidia. Bar = 10 µm.
Illustrations: Kar & Mandal (1969: 358, ig. 20), Guo et al.
(1998: 404, ig. 342).
Description: Leaf spots amphigenous, scattered to conluent,
circular to irregular, at irst smaller, 1–10 mm diam, later
expanding, covering large leaf segments, reddish brown to
greyish, margin indeinite or with darker border, purplish to
deep reddish brown. Caespituli amphigenous, punctiform,
dark, scattered. Mycelium internal. Stromata small to welldeveloped, substomatal, 10–65 µm diam, subglobose, oblong
to irregular, pale to dark olivaceous-brown. Conidiophores
in small to moderately large fascicles, mostly about 2–16,
arising from stromata, through stomata, erect, straight,
curved to somewhat geniculate-sinuous, unbranched, 10–75
× 3–5.5 µm, 0–6-septate, pale to dark olivaceous-brown,
thin-walled, smooth; conidiogenous cells integrated, terminal
or conidiophores reduced to conidiogenous cells, 10–25
380
µm long, conidiogenous loci inconspicuous, unthickened,
not darkened. Conidia solitary, obclavate to obclavatecylindrical, straight to somewhat curved, 25–95 × 2.5–4.5
µm, 2–9-septate, subhyaline to pale olivaceous-brown, thinwalled, smooth, apex pointed to subobtuse, base rounded
to short obconically truncate, about 1.5–2 µm wide, hila
unthickened, not darkened.
Holotype: India: West Bengal: Darjeeling, Tung, 1768 m
alt., on Amomum dealbatum, 12 May 1967, A. K. Kar & M.
Mandal (K(M) IMI 135184).
Host range and distribution: On Amomum (compactum,
dealbatum, villosum), Elettaria cardamomum [Amomum
cardamomum], Zingiberaceae, Asia (China, India, Indonesia).
pseudocercospora curcumicola A.Yoshida, I. Araki
& C. Nakash., sp. nov.
MycoBank MB809028
(Fig. 184)
Synonym: Pseudocercospora curcumae A.Yoshida, I. Araki
& C. Nakash., Fungal Flora of Diseases on Medical Plants in
Japan: 81 [Master thesis, Mie University, Tsu] (2008), nom.
inval. (ICN Art. 30.8).
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Leaf spots amphigenous, indistinct or distinct, scattered,
angular to irregular, small, 1–5 mm diam, later conluent, pale
brown to brown. Caespituli amphigenous. Stromata small to
well-developed, subglobular, substomatal to intraepidermal,
olivaceous-brown to brown, 10–40 µm diam. Conidiophores
in small to moderately large, loosely to densely fasciculate,
arising from stromata, emerging through stomata or
erumpent, erect, straight to slightly curved, geniculate,
subcylindrical, unbranched, pale olivaceous-brown, 12–26
× 2–3.2 µm, 1–4-septate, smooth; conidiogenous cells
integrated, terminal or intercalary, proliferating sympodially or
percurrently, with inconspicuous conidiogenous loci, 2–2.5 µm
diam, unthickened, not darkened. Conidia solitary, cylindrical
to obclavate, acicular, straight to curved or sinuous, 40–80 ×
2–2.5 µm, 1–6-septate, pale to pale olivaceous-brown, apex
obtuse, base truncate, 2–2.5 µm wide, hila unthickened, not
darkened.
Holotype: Japan: Wakayama Pref.: Hidakagawa, on Curcuma
longa, Zingiberaceae, 30 Oct. 2007, C. Nakashima & I. Araki
(MUMH 10859). Isotype: TNS-F-60890. Ex-holotype culture:
MUCC 733.
Host range and distribution: Only known from the type
collection.
Notes: Pseudocercospora curcumicola is distinguishable
from other Pseudocercospora species on hosts of the
Zingiberaceae by its moderately developed stromata,
geniculate, densely fasciculate conidiophores, and pale
conidia with truncate bases.
pseudocercospora hedychii (Boedijn) U. Braun,
Nova Hedwigia 73: 424 (2001).
(Fig. 185)
Basionym: Cercospora hedychii Boedijn, Nova Hedwigia 3:
432 (1961).
Literature: Thaung (1984), Crous & Braun (2003: 213),
Soares & Barreto (2008).
Illustrations: Boedijn (1961: plate 111, ig. 18), Braun (2001a:
426, ig. 7), Soares & Barreto (2008: 89, ig. 4).
Description: Leaf spots amphigenous, often marginal and
terminal, size and shape variable, oblong-elliptical, irregular,
often conluent, straw-coloured, pale greyish brown, margin
indeinite or narrow and brown. Caespituli amphigenous,
punctiform to subeffuse, dark. Mycelium internal and
external; supericial hyphae almost lacking or sparingly
developed, emerging through stomata, unbranched or
sparingly branched, septate, 1–4 µm wide, subhyaline to
pale olivaceous, thin-walled, smooth. Stromata substomatal
to intraepidermal, 10–60 µm diam, medium to dark
olivaceous-brown. Conidiophores in small to moderately
large fascicles, arising from stromata, through stomata or
erumpent or solitary, arising from supericial hyphae, lateral
or occasionally terminal, erect, straight, subcylindricalconical to geniculate-sinuous, unbranched, 5–60 × 2–4 µm,
0–3-septate, subhyaline to pale olivaceous or brownish, thin-
VOLUME 5 · NO. 2
Fig. 185. Pseudocercospora hedychi (L 53858). A. Conidiophore
fascicles. B. Supericial hypha with solitary conidiophores. C.
Conidiophores. d. Conidia. Bar = 10 µm.
walled, smooth; conidiogenous cells integrated, terminal or
conidiophores reduced to conidiogenous cells, 10–35 µm
long, conidiogenous loci inconspicuous, unthickened, not
darkened. Conidia solitary, narrowly obclavate-cylindrical,
subcylindrical, subacicular, 15–80 × 2–4 µm, 1–10-septate,
subhyaline to pale olivaceous or olivaceous-brown, thinwalled, smooth, apex subobtuse or subacute, base truncate
to obconically truncate, 1.5–2.5(–3.5) µm wide, hila
unthickened, not darkened.
Holotype: Indonesia: Java: Bogor, botanical garden, on
Hedychium coccineum, 28 June 1950, K. B. Boedijn (L
53858). Isotype: K(M) IMI 91602.
Host range and distribution: On Hedychium (coccineum,
coronarium, spicatum), Zingiberaceae, Asia (China [Hong
Kong], Indonesia, Myanmar), South America (Brazil).
pseudocercospora nicolaiae (Boedijn) U. Braun,
Nova Hedwigia 73: 427 (2001).
(Fig. 186)
Basionym: Cercospora nicolaiae Boedijn, Nova Hedwigia 3:
432 (1961).
Literature: Crous & Braun (2003: 291).
Illustrations: Boedijn (1961: plate 111, ig. 19), Braun (2001a:
426, ig. 11).
381
Braun et al.
ART I CLE
terminal, 10–40 µm long, conidiogenous loci inconspicuous,
unthickened, not darkened. Conidia solitary, obclavatecylindrical, 30–85 × 2.5–5 µm, 1–7-septate, pale olivaceous
to olivaceous-brown, thin-walled, smooth, apex obtuse to
subacute, base truncate to obconically truncate, 2–3 µm
wide, hila unthickened, not darkened.
Holotype: Indonesia: Java: Bogor, botanical garden, on
Etlingera sp. [Nicolaia sp.], Zingiberaceae, Feb. 1949, K. B.
Boedijn (L 53870). Isotype: K(M) IMI 91603.
Host range and distribution: Only known from the type
collection.
pseudocercospora zingiberis (Rathaia) Deighton,
Mycol. Pap. 151: 2 (1983).
(Fig. 187)
Basionym: Cercoseptoria zingiberis Rathaia, Mycologia 73:
774 (1981).
Illustration: Rathaia (1981: 775–777, igs 1–9).
Description: Leaf spots amphigenous, at irst oblong,
yellowish brown, parallel to the main veins, later oblongfusiform, 4–9 × 0.5–1.2 cm, centre becoming white with dark
brown margin and yellowish halo. Caespituli amphigenous,
punctiform, dark. Mycelium internal; hyphae dark brown, wall
thickened. Stromata well-developed, immersed, originating
from hyphae in the veins, brush-like, 40–120 µm diam, dark
brown. Conidiophores numerous in dense sporodochial
fascicles, arising from stromata, erumpent, subcylindrical or
somewhat wider towards the apex, straight to slightly curved
or somewhat geniculate-sinuous, unbranched, 8–25 × 2–3.5
µm, 0–1-septate, 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
and not darkened. Conidia solitary, subacicular-iliform,
long and narrow, fusoid, i.e. attenuated towards base and
apex, 40–190 × 1.5–3 µm, 6–24-septate, subhyaline to pale
yellowish, thin-walled, smooth, apex pointed, base short
to long obconically truncate, about 1–1.5 µm wide, hila
unthickened and not darkened.
Fig. 186. Pseudocercospora nicolaiae (L 53870). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
Description: Leaf spots lacking, sometimes with diffuse
ochraceous
discolorations.
Caespituli
amphigenous,
punctiform, often spreading between veins, dark brown.
Mycelium internal. Stromata substomatal to immersed,
variable, small to well-developed, 10–60 µm diam,
subglobose to oblong, medium to dark brown. Conidiophores
in small to very large fascicles, loose to dense, sometimes
subsynnematous, arising from stromata, through stomata
or erumpent, erect, straight, subcylindrical to somewhat
geniculate-sinuous, unbranched, 30–150 × 2.5–5 µm,
pluriseptate throughout, wall slightly thickened, pale to
medium brown, smooth; conidiogenous cells integrated,
382
Holotype: India: Assam: Halong, on Zingiber oficinale,
Zingiberaceae, 21 Jul. 1978, Y. Rathaia (K(M) IMI231501).
Host range and distribution: Only known from the type
collection.
Doubtful, excluded and insuficiently known
species
Pseudocercospora alpiniicola S.Q. Chen & P.K. Chi, J.
S. China Agric. Univ. 11: 48 (1990); as “alpinicola”.
Literature: Chi (1994: 49), Guo et al. (1998: 402).
Illustrations: Chi (1994: 49, ig. 30), Guo et al. (1998: 403,
ig. 340).
IMA FUNGUS
Cercosporoid fungi 2
ART I CLE
Fig. 188. Zasmidium alpinum (S-F23069). A. Supericial hypha.
B. Conidiophore fascicle. C. Conidiophore. d. Conidiophore tip. e.
Conidia. Bar = 10 µm.
µm, 5–19-septate, pale olivaceous, thin-walled, smooth,
apex obtuse to pointed, base short obconically truncate.
Holotype: China: Guangdong: Yunan, on Alpinia oficinarum,
Zingiberaceae, Oct. 1987, S. Q. Chen 106 (Hb. S. China
Agric. Univ., Guangzhou).
Fig. 187. Pseudocercospora zingiberis (K(M) IMI 231501). A.
Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
Description: Leaf spots amphigenous, irregular, greyish
white with brown margin. Caespituli amphigenous.
Mycelium immersed and external, with some supericial
hyphae, branched, septate, 1.7–3.3 µm wide, hyaline to
pale olivaceous. Stromata subglobose, 17–60 µm diam,
olivaceous-brown. Conidiophores loosely fasciculate, 7–18,
arising from stromata, or solitary, arising from supericial
hyphae, erect to decumbent, straight, cylindrical, nongeniculate, unbranched or rarely branched, 33–133 × 4 µm,
1–7-septate, pale olivaceous to olivaceous, thin-walled,
smooth; conidiogenous cells integrated, terminal, rhachislike, straight, cylindrical, neither geniculate nor sinuous,
denticulate, with numerous minute conical denticles. Conidia
solitary, obclavate, straight to slightly curved, 50–83 × 3.3
VOLUME 5 · NO. 2
Host range and distribution: Only known from the type
collection.
Notes: Due to the straight rhachis with numerous conical
denticles, this species is undoubtedly misplaced in
Pseudocercospora and rather belongs to the Dactylaria
complex, but a inal conclusion and reassessment have to be
based on an examination of type material.
Zasmidium
A single species.
Zasmidium alpiniae (Syd. & P. Syd.) U. Braun,
Schlechtendalia 20: 100 (2010).
(Fig. 188)
Basionym: Cercospora alpiniae Syd. & P. Syd., Ann. Mycol.
12: 202 (1914).
383
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Braun et al.
Synonyms: Helminthosporium alpiniae (Syd. & P. Syd.)
Chupp, Monograph of Cercospora: 607 (1954).
Stenella alpiniae (Syd. & P. Syd.) U. Braun, Fungal Diversity
8: 68 (2001).
Literature: Saccardo (1931: 894), Chupp (1954: 607), Katsuki
(1965: 69).
Illustration: Braun (2001b: 67, ig. 22).
Description: Leaf spots lacking or only with diffuse yellowish
discolorations on the upper leaf surface. Caespituli
hypophyllous, effuse, dark olivaceous to almost sooty.
Mycelium internal and external; supericial hyphae emerging
through stomata, sparingly branched, 1–3 µm wide, septate,
pale to medium olivaceous, thin-walled, verruculose.
Stromata lacking. Conidiophores in small, loose fascicles,
emerging through stomata (solitary conidiophores arising from
supericial hyphae lacking), erect to decumbent, lexuous,
strongly geniculate-sinuous, unbranched, very long, 50–300
× (3–)4–7(–8) µm, pluriseptate throughout, pale to medium
dark brown, wall thin to somewhat thickened; conidiogenous
cells integrated, terminal or intercalary, 10–50 µm long,
conidiogenous loci thickened and darkened, 1.5–2.5 µm diam.
Conidia solitary, rarely in short chains, obclavate-cylindrical,
20–80 × 4–8 µm, rarely longer, (1–)2–7(–9)-septate, pale
to medium dark olivaceous-brown, wall slightly thickened,
verruculose, apex obtuse, base obconically truncate, about 2
µm wide, hila somewhat thickened and darkened.
[Holotype: philippines: Prov. Laguna: near Los Baños,
Mt Maquiling, on leaves of Alpinia sp., 18 Dec. 1913, C. F.
Baker, no. 2221 (not preserved)]. Neotype (designated
here, MycoBank MBT178173): philippines: Prov. Laguna:
near Los Baños, Mt Maquiling, on leaves of Alpinia sp., Dec.
1914, C. F. Baker, no. 213 (S-F23069). Isoneotypes: BPI
432462, 432463, CUP-F-0213 and PC.
Host range and distribution: On Alpinia (galanga, japonica,
Alpinia sp.), Zingiberaceae, Asia (Japan, Malaysia [Sarawak],
Philippines).
Notes: The designated neotype is topotype material collected
in Dec. 1914.
ACKNowledgeMeNtS
We are very grateful to the directors and curators of B, BPI, CUP, DAOM,
FH, HBG, HMAS, ILL, ILLS, IUM, K, L, LE, LEP, LPS, MA, NIAES, NTU,
NY, NYS, PAD, PC, PDD, RREM, S, TFM, TNS and WSP for loaning
type material and other collections in their keeping during the course of
monographic studies of cercosporoids fungi on monocots.
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IMA FUNGUS
Uwe Braun