IMA FuNgus · 6(1): 25–97 (2015)
doi:10.5598/imafungus.2015.06.01.03
Uwe Braun1, Pedro W. Crous2,3, and Chiharu Nakashima4
ART I CLE
cercosporoid fungi (Mycosphaerellaceae) 3. species on monocots (Poaceae,
true grasses)
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
CBS-KNAW, Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands
3
Department of Microbiology and Plant Pathology, Forestry and Agricultural Biotechnology Institute, University of Pretoria, Pretoria 0002, South
Africa
4
Graduate School of Bioresources, Mie University, 1577 Kurima-machiya, Tsu, Mie 514-8507, Japan
Abstract: The third part of a series of monographic treatments of cercosporoid fungi (formerly Cercospora
s. lat., Mycosphaerellaceae, Ascomycota) continues with a treatment of taxa on monocots (Liliopsida;
Equisetopsida, Magnoliidae, Lilianae), covering asexual and holomorph species with mycosphaerellalike sexual morphs on true grasses (Poaceae), which were excluded from the second part. The species
concerned are keyed out, alphabetically listed, described, illustrated and supplemented by references
to previously published descriptions, illustrations, and exsiccatae. A key to the recognised genera and
a discussion of taxonomically relevant characters was published in the irst part of this series. Several
species are lecto- or neotypiied. The following taxonomic novelties are introduced: Cercospora barretoana
comb. nov., C. cymbopogonicola nom. nov., Cladosporium elymi comb. nov., Passalora agrostidicola sp.
nov., P. brachyelytri comb. nov., and P. dichanthii-annulati comb. nov.
Key words:
Ascomycota
Cercospora s. lat.
Liliopsida
Gramineae
hyphomycetes
Article info: Submitted: 4 January 2014; Accepted: 17 March 2015; Published: 9 April 2015.
INtroductIoN
The taxonomy of cercosporoid fungi (Cercospora s. lat.,
sensu Chupp 1954), especially the circumscription of
genera in this complex, has long been problematic. It has
been subjected to many changes in recent decades, due
to subjective assessments of morphology. The emergence
of molecular phylogenetic methods and their application to
the taxonomy of cercospora-like fungi has led to a better
understanding of the importance of morphological and
biological traits as well as more stable generic concepts
(Crous et al. 2013a, Groenewald et al. 2013). Cercosporoid
fungi are worldwide in distribution and embrace a wide range
of asexual morphs, asexual holomorphs and species with
mycosphaerella-like sexual morphs (Mycosphaerella s. str.
is now a heterotypic synonym of Ramularia, see Braun et
al. 2013), which are mostly leaf-spotting plant pathogens.
Numerous species cause serious diseases on cultivated
plants (crops, ornamental plants, forest trees) in agriculture
and forestry. The only comprehensive treatment of this fungal
complex (Chupp 1954) has become obsolete. Braun et al.
(2013) initiated a project to produce a modern monograph
of Cercospora and allied genera (Mycosphaerellaceae),
through a series of monographic papers rather than by a
comprehensive treatment in a single opus. A irst contribution
reviewed aspects of the taxonomy of cercospora-like fungi
(history, taxonomic value of traits, circumscriptions of genera,
key to genera) and dealt with cercosporoids on fungi, ferns
and gymnosperms (Braun et al. 2013). The second part of
this series encompassed a treatment of species occurring on
monocot families (Braun et al. 2014), but excluding taxa on
true grasses (Poaceae). The species of cercosporoid fungi
on Poaceae are treated in the present contribution, which
follows the principles outlined in previous parts of this series.
MAterIAl ANd Methods
The present work is a compilation based on papers and
our unpublished data, as well as global literature. Details
on methods are given in the papers cited. As far as new
examinations are concerned, fungal structures have been
examined with 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
© 2015 International Mycological Association
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VOLUME 6 · NO. 1
25
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Braun et al.
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).
tAxoNoMIc treAtMeNt
cercosporoid species on Poaceae (Gramineae, true grasses)
Cercospora
Key to Cercospora species on Poaceae
1
Conidia at least partly in chains ...................................................................................................................................... 2
Conidia consistently solitary ............................................................................................................................................ 7
2 (1)
Conidia solitary, rarely forming short chains, acicular to obclavate-cylindrical,
(15–)25–120(–320) × (1.5–)2.5–5.5 µm; on Sorghum spp. .............................................. c. sorghi var. ciccaronei
Conidia frequently in chains, not acicular; on other hosts ............................................................................................... 3
3 (2)
Conidiophores very long and rather broad, 100–275 × 4–6 µm, to 20-septate;
conidia very broad, 3–7.5 µm wide, but hila narrower, 1–1.5 µm;
on Eremochloa bimaculata, Australia .............................................................................................. c. eremochloae
Conidiophores much shorter, less than 100 µm, narrower and only 0–7-septate;
conidia narrower, 1.5–5 µm wide; on other hosts .................................................................................................... 4
4 (3)
Stromata lacking or almost so; conidiophores rather long, 25–215 × 3–5 µm, 0–7-septate;
conidia 2–4.5 µm wide, hila 1.5–2.5 µm wide; on Rottboellia cochinchinensis, Africa ........................ c. rottboelliae
Stromata developed and/or conidiophores much shorter, to 100 µm, and/or conidia narrower,
1.5–3 µm, and hila only 1–2 µm wide ...................................................................................................................... 5
5 (4)
Conidia (2.5–)3–5.5(–6) µm wide; on various hosts of the Panicoideae .................................................... c. barretoana
Conidia narrower, 1.5–3 µm wide (on hosts of Panicoideae) or 1.5–4 µm wide
(on Agrostis and Sphenopholis, Pooideae, Aveneae, in North America) ................................................................. 6
6 (5)
On numerous hosts of genera belonging to Panicoideae, almost worldwide ........................................ c. fusimaculans
On Agrostis and Sphenopholis spp. (Pooideae, Aveneae), North America ................................................. c. agrostidis
7 (1)
Conidial shape variable, narrowly subcylindrical, subacicular to somewhat cylindrical-obclavate,
fusiform, very narrow, 30–100 × 1–2 µm, hila narrow, 0.5–1.5 µm; loci of conidiogenous cells minute,
1–1.5 µm; on Digitaria spp., Africa, Oceania, South America ................................................................ c. digitariae
Conidia and hila broader; loci mostly larger than 1.5 µm diam; on other hosts .............................................................. 8
8 (7)
Conidiophores very short, 5–20 × 2–4 µm, aseptate; conidia narrowly iliform-acicular,
30–90 × 1.5–2.5 µm; conidiogenous loci and hila minute, 1–1.5 µm diam;
on Dactyloctenium ................................................................................................................................ c. tessellata
Conidiophores much longer and/or septate or conidiogenous loci broader;
conidia consistently acicular or acicular to obclavate-cylindrical or
uniformly obclavate-cylindrical or cylindrical, hila mostly broader; on other hosts ................................................... 9
9 (8)
Conidia obclavate, cylindrical, base obconically truncate, at most some conidia subacicular,
but acicular conidia with truncate base lacking ...................................................................................................... 10
Conidia consistently acicular (C. apii complex) or at least partly acicular, i.e. ranging from
acicular to obclavate-cylindrical (often only younger and/or smaller conidia obclavate-cylindrical,
base of the conidia truncate to obconically truncate) ............................................................................................. 28
10 (9) Conidia relatively broad, 4–9 µm wide, average ≥ 5 µm ................................................................................................ 11
Conidia narrower, (1.5–)2–5(–6) µm wide, average < 5 µm ......................................................................................... 14
11 (10) Conidiogenous loci minute, 0.5–1.5 µm diam; conidia (10–)15–65(–85) × 3–6.5 µm,
hila 1–2 µm wide; on Oryza (Ehrhartoideae, Oryzeae) .......................................................................... c. janseana
26
IMA FUNGUS
cercosporoid fungi 3
12 (11) On Saccharum spp. ....................................................................................................................................... c. longipes
On Zea mays ................................................................................................................................................................. 13
ART I CLE
Conidiogenous loci broader, more than 1.5 µm; conidia longer, at least partly more than 100 µm,
4–9 µm wide, hila larger, 2–3 µm diam; on hosts belonging to Panicoideae, Andropogoneae
(Saccharum, Zea) ................................................................................................................................................... 12
13 (12) Conidiophores 40–180 µm long; conidia broadly obclavate-cylindrical, 30–100 × 4–9 µm;
cultures not slow-growing, forming a red pigment (cercosporin) ..................................................... c. zeae-maydis
Conidiophores to about 100 µm in length; conidia broadly fusiform; cultures slow-growing,
without formation of red pigments (cercosporin) [morphologically barely distinguished
from C. zeae-maydis, but genetically clearly differentiated] .......................................................................... c. zeina
14 (10) Conidia long, to 150 µm, average > 50 µm (on Eragrostis, Ischaemum, Miscanthus, Paspalum,
Cenchrus (including Pennisetum), Setaria) ........................................................................................................... 15
Conidia relatively short, 15–65 µm long, average < 50 µm (on other hosts) ................................................................ 20
15 (14) Stromata small, little developed; conidiophores in relatively small fasciles, 2–15;
on Echinochloa, Ischaemum, Paspalum, [Cenchrus, including Pennisetum] and Setaria (Panicoideae)
................................................................................................................................................................................. 16
Stromata well-developed, large, to 80 µm diam; conidiophores numerous, to 30 per fascicle;
on Eragrostis or Miscanthus .................................................................................................................................. 19
16 (15) Conidiophore 0–1-septate; on Ischaemum australe or Cenchrus spicatus [Pennisetum glaucum] .............................. 17
Conidiophores 0–4-septate; on Echinochloa or Setaria ................................................................................................ 18
17 (16) Conidia obclavate-cylindrical, but long conidia may be almost acicular;
on Cenchrus spicatus [Pennisetum glaucum], India ............................................................................. c. typhoides
Conidia consistently obclavate; on Ischaemum australe, Australia ............................................................... c. ischaemi
18 (16) Conidiophores short, 8–45 µm; on Paspalum and Setaria ............................................................................. c. setariae
Conidiophores longer, 10–80 µm; on Echinochloa ................................................................................ c. echinochloae
19 (15) Conidiophores long, (15–)40–120 µm, 1–7-septate; on Miscanthus, Taiwan .............................................. c. miscanthi
Conidiophores shorter, 15–30(–50) µm, 0–1(–2)-septate; on Eragrostis brownei, New Zealand ............ c. eragrostidis
20 (14) Conidiogenous loci minute, 0.5–1.5 µm diam; conidia 3–6.5 µm wide, hila 1–2 µm wide; on Oryza ........... c. janseana
Conidiogenous loci larger, 1.5–2 µm, and/or conidia narrower, 2–5 µm ....................................................................... 21
21 (20) Conidiophores 0–1-septate; on Cymbopogon or Erharta .............................................................................................. 22
Conidiophores pluriseptate, at least partly with two or more septa ............................................................................... 23
22 (21) Conidiophores 28.5–70 µm long, 1-septate; conidia 1–10-septate; on Cymbopogon, India ......... c. cymbopogonicola
Conidiophores shorter, 15–30(–40) µm, 0(–1)-septate; on Ehrharta, New Zealand ................................ c. microlaenae
23 (21) Stromata well-developed, large, 20–300 µm diam; conidiophores long, 30–160 µm;
on Chusquea (Bambusoideae, Bambuseae) ...................................................................................... c. chusqueae
Stromata lacking or small, 10–30 µm diam; conidiophores shorter, to 80 µm;
on hosts of other subfamilies ................................................................................................................................. 24
24 (23) Conidiophores 10–60 µm long, 0–1-septate; on Bothryochloa saccharoides ...................................... c. bothryochloae
Conidiophores with to 4 septa; on other hosts .............................................................................................................. 25
25 (24) Conidiophores rather robust, broad, 10–80 × 3–8 µm, mostly pale, subhyaline,
yellowish or pale to medium olivaceous-brown; on Echinochloa .................................................... c. echinochloae
Conidiophores narrower, 2–5 µm wide, pigmentation usually darker; on other hosts ................................................... 26
26 (25) Stromata lacking or almost so; on Zizania (Ehrhartoideae, Oryzeae) ............................................................. c. zizaniae
Stromata to 30 µm diam; on Cynodon or Sporobolus (Chloridoideae) ......................................................................... 27
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Braun et al.
27 (26) On Cynodon (Chloridoideae, Cynodonteae), India .................................................................................... c. cynodontis
On Sporobolus (Chloridoideae, Eragrostideae), North America ....................................................................... c. seriata
28 (9) Conidia acicular to obclavate-cylindrical, base truncate to obconically truncate ........................................................... 29
Conidia consistently acicular, base truncate ................................................................................................................. 35
29 (28) Conidiophores relatively short, 10–67 µm, 0–1(–2)-septate; conidia mostly obclavate-cylindrical;
on Bromus spp. or Cenchrus spicatus [Pennisetum glaucum] .............................................................................. 30
Conidiophores pluriseptate (about 3–20 or even more); acicular conidia abundant ..................................................... 31
30 (29) Conidiophores cylindrical or only somewhat geniculate; conidia 2.5–5 µm wide;
on Cenchrus spicatus [Pennisetum glaucum], India ............................................................................. c. typhoides
Conidiophores distinctly, often strongly geniculate; conidia narrower, 1–3.5 µm wide;
on Bromus, North America ............................................................................................................... cercospora sp.
31 (29) Stromata well-developed, to 50 µm diam; conidiophores rather long, 25–300 µm,
conidiogenous loci minute, 1–1.5(–2) µm diam; conidia to 260 µm long; on Eleusine, Asia ................ c. eleusines
Stromata lacking or very small, to 20 µm diam; and/or conidiogenous loci 1.5–2 µm diam or
even larger; on other hosts .................................................................................................................................... 32
32 (31) Stromata 10–50 µm diam; conidiophores (10–)20–150(–220) × (2.5–)3–6.5(–7) µm;
conidiogenous loci and conidial hila 1.5–3 µm diam; on Sorghum .............................. c. sorghi and c. sorghicola
Stromata lacking or very small, to 20 µm diam; conidiogenous loci and hila 1.5–2 µm diam or,
if larger, conidiophores very long, to 800 µm ......................................................................................................... 33
33 (32) Conidiophores very long, 20–800 µm; conidia 30–300 µm long; conidiogenous loci (1.5–)2–3 µm;
on Festuca, North America ..................................................................................................................... c. festucae
Conidiophores much shorter, to 160 µm; conidiogenous loci 1.5–2 µm diam; on other hosts ...................................... 34
34 (33) Conidia 20–80 µm long, hyaline, subhyaline to very pale olivaceous;
on Bouteloua and Chondrosus spp., North America ........................................................................... c. boutelouae
Conidia longer, 40–235 µm long, consistently hyaline; on Rottboellia spp., Asia, South America ....... c. rottboelliigena
35 (28) Conidiophores consistently short, 6–28 µm long, pale olivaceous-brown; conidia narrow,
30–130 × 2–3 µm; on Oplismenus ........................................................................................................ c. oplismeni
Conidiophores much longer, 20–240 µm long, and darker, and/or conidia wider, 2–5 µm wide;
on other hosts ........................................................................................................................................................ 36
36 (35) Conidiophores short, 15–50 µm long; on Coix or Secale .............................................................................................. 37
Conidiophores much longer, > 50 µm long ................................................................................................................... 38
37 (36) Leaf spots circular to elliptical, 1–5 mm diam; on Coix, India ............................................................................. c. coicis
Leaf spots oblong, forming narrow lines, 0.5 mm wide or larger lesions to 35 × 3 mm; on Secale .................. c. secalis
38 (36) Stromata lacking or very small; conidiophores long, to 250 µm long;
conidiogenous loci relatively large, 2–4 µm; on Bromus and Cenchrus (including Pennisetum) ........................... 39
Stromata larger, 10–50 µm diam, and/or conidiophores much shorter, about 20–150 µm long;
and/or conidiogenous loci smaller, 2–2.5 µm diam; and/or conidia shorter, (15–)20–50(–100) µm ...................... 40
39 (38) Conidia to 240 µm long; on Cenchrus (including Pennisetum) spp. ............................................................. c. penniseti
Conidia shorter, to about 120 µm long; on Bromus inermis .................................................................... cercospora sp.
40 (38) Conidia relatively short, (15–)20–50(–100) µm long; on Secale ....................................................................... c. secalis
Conidia longer, 15–155 µm long; on Aristida or Arthraxon spp. .................................................................................... 41
41 (40) Leaf spots oval to oblong, 0.5–2 mm in length; on Aristida spp., North America ........................................... c. aristidae
Leaf spots circular to angular, 2–5 mm diam; on Arthraxon spp., India .................................................... c. arthraxonis
28
IMA FUNGUS
cercosporoid fungi 3
Tabular key to Cercospora species on Poaceae
On Agrostis ............................................................................................................................................................. c. agrostidis
ART I CLE
The species are listed in form of a tabular key based on host genera in alphabetical order.
On Aristida ................................................................................................................................................................ c. aristidae
On Arthraxon ........................................................................................................................................................ c. arthraxonis
On Avena .................................................................................................................................................................... ?c. secalis
On Beckeropsis ................................................................................................................................................ c. fusimaculans
On Bothriochloa .............................................................................................................................................. c. bothryochloae
On Bouteloua
1
Conidia solitary ............................................................................................................................................. c. bouteloae
Conidia catenate .................................................................................................................................. ?c. fusimaculans
On Brachiaria .................................................................................................................................................... c. fusimaculans
On Bromus
1
Conidia acicular, 3–4 µm wide; conidiophores 30–200 µm long, not or only slightly geniculate,
conidiogenous loci and hila 2–3.5 µm diam .......................................................................................... c. apii s. lat.
Conidia narrowly obclavate to acicular, shorter conidia sometimes fusiform, 1–3.5 µm wide;
conidiophores shorter, 10–40 µm, distinctly, often even strongly geniculate, conidiogenous loci
and hila 1–2 µm diam ...................................................................................................................... Cercospora sp.
On Cenchrus (including Pennisetum)
1
Conidia catenate .................................................................................................................................... c. fusimaculans
Conidia solitary ................................................................................................................................................................ 2
2 (1)
Conidiophores 50–250 µm long, pluriseptate; conidia acicular ..................................................................... c. penniseti
Conidiophores 17–67 µm long, 0–1-septate; conidia obclavate-cylindrical ................................................. c. typhoides
On Chasmopodium ........................................................................................................................................... c. fusimaculans
On Chondrosus ....................................................................................................................................................... c. bouteloae
On Chusquea ........................................................................................................................................................ c. chusqueae
On Coix ........................................................................................................................................................................... c. coicis
On Cymbopogon ....................................................................................................................................... c. cymbopogonicola
On Cynodon .......................................................................................................................................................... c. cynodontis
On Dactyloctenium .................................................................................................................................................................... c. tessellata
On Digitaria
1
Conidia solitary .............................................................................................................................................. c. digitariae
Conidia catenate .................................................................................................................................... c. fusimaculans
On Echinochloa
1
Conidia formed singly ............................................................................................................................. c. echinochloae
Conidia catenate ............................................................................................................................................................. 2
2 (1)
Conidia 1.5–3 µm wide .......................................................................................................................... c. fusimaculans
Conidia wider, (2.5–)3–5.5(–6) µm wide .................................................................................................... c. barretoana
VOLUME 6 · NO. 1
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Braun et al.
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On Ehrharta ......................................................................................................................................................... c. microlaenae
On Eleusine
1
Conidia catenate .................................................................................................................................. ?c. fusimaculans
Conidia solitary ................................................................................................................................................................ 2
2 (1)
Conidiophores 25–300 µm long, pluriseptate, conidiogenous loci 1–1.5(–2) µm diam;
conidia acicular to obclavate-cylindrical, 50–260 × 3–4 µm .................................................................. c. eleusines
Conidiophores very short, 5–20 × 2–4 µm, 0(–1)-septate; conidia narrowly iliform-acicular,
30–90 × 1.5–2.5 µm .............................................................................................................................. c. tessellata
On Entolasia ...................................................................................................................................................... c. fusimaculans
On Eragrostis ...................................................................................................................................................... c. eragrostidis
On Eremochloa .................................................................................................................................................. c. eremochloae
On Festuca ................................................................................................................................................................ c. festucae
On Hordeum ............................................................................................................................................................... ?c. secalis
On Hymenachne ................................................................................................................................................... c. barretoana
On Ischaemum ............................................................................................................................................................................. c. ischaemi
On Miscanthus ........................................................................................................................................................ c. miscanthi
On Muhlenbergia ............................................... c. muhlenbergiae (see Doubtful, excluded and insuficiently known species)
On Oplismenus
1
Conidia catenate .................................................................................................................................... c. fusimaculans
Conidia solitary ............................................................................................................................................. c. oplismeni
On Oryza .................................................................................................................................................................. c. janseana
On Panicum
1
Conidia 1.5–3 µm wide .......................................................................................................................... c. fusimaculans
Conidia wider, (2.5–)3–5.5(–6) µm wide .................................................................................................... c. barretoana
On Paspalidium ................................................................................................................................................. c. fusimaculans
On Paspalum .............................................................................................................................................................. c. setariae
On Pennisetum see Cenchrus
On Rottboellia
1
Conidia formed singly, acicular, 40–235 µm long, 3- to pluriseptate .................................................... c. rottboelliigena
Conidia catenate ............................................................................................................................................................. 2
2 (1)
Conidiophores to 215 µm long; conidia 15–55 × 2–4.5 µm, 1–7-septate ................................................. c. rottboelliae
Conidiophores much shorter, < 100 µm long; conidia 1.5–3 µm wide ................................................. ?c. fusimaculans
On Saccharum .......................................................................................................................................................... c. longipes
On Secale ..................................................................................................................................................................... c. secalis
On Setaria
1
Conidia catenate .................................................................................................................................... c. fusimaculans
Conidia solitary ................................................................................................................................................ c. setariae
30
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cercosporoid fungi 3
2 (1)
Conidia consistently solitary ............................................................................. c. sorghi var. sorghi and c. sorghicola
Conidia mostly solitary, but occasionally in short chains .......................................................... c. sorghi var. ciccaronei
ART I CLE
On Sorghum
1
Conidia frequently catenate, 1.5–3 µm wide .......................................................................................... c. fusimaculans
Conidia solitary or only rarely catenate, broader, usually 2.5–5.5 µm wide .................................................................... 2
On Sphenopholis .................................................................................................................................................... c. agrostidis
On Sporobolus ............................................................................................................................................................. c. seriata
On Stenotaphrum ............................................................................................................................................. c. fusimaculans
On Triticum ................................................................................................................................................................. ?c. secalis
On Urochloa ...................................................................................................................................................... c. fusimaculans
On Zea
1
Conidia catenate .................................................................................................................................... c. fusimaculans
Conidia solitary ................................................................................................................................................................ 2
2 (1)
Conidia acicular, narrow, < 5 µm wide .................................................................... c. apii s. lat. (C. sorghi var. maydis)
Conidia broadly obclavate-cylindrical to fusiform, 4–9 µm wide ..................................................................................... 3
3 (2)
Conidiophores 40–180 µm long; conidia broadly obclavate-cylindrical, 30–100 × 4–9 µm;
cultures not slow-growing, forming a red pigment (cercosporin) ..................................................... c. zeae-maydis
Conidiophores to about 100 µm in length; conidia broadly fusiform; cultures slow-growing,
without formation of red pigment (cercosporin) [morphologically barely distinguished
from C. zeae-maydis, but genetically clearly differentiated] .......................................................................... c. zeina
On Zizania ................................................................................................................................................................... c. zizaniae
List of Cercospora species on Poaceae
cercospora agrostidis G.F. Atk., J. Elisha Mitchell
Sci. Soc. 8: 44 (1892).
(Fig. 1)
Literature: Saccardo (1892: 656), Vassiljevsky & Karakulin
(1937: 271), Chupp (1954: 243), Braun & Mel’nik (1997: 61).
Illustration: Braun & Mel’nik (1997: ig. 84).
Description: Leaf spots elliptical, about 3–5 mm long and
1–3.5 mm wide, centre ochraceous to pale brown, with a
narrow to broad, purplish violet to dull reddish brown margin
or halo. Caespituli amphigenous, punctiform, dark brown to
blackish, scattered. Mycelium internal. Stromata substomatal,
almost lacking or relatively small, 10–30 µm diam, brown,
composed of swollen hyphal cells, 2–6 µm diam, circular to
slightly irregular-angular in outline. Conidiophores in small to
moderately large fascicles, divergent to dense, arising from
stromata, through stomata, erect, straight, subcylindrical or
somewhat narrowed towards the apex to slightly geniculatesinuous, unbranched, 20–70(–125) × 3–5 µm, continuous to
pluriseptate, olivaceous, yellowish to medium brown, often
paler towards the tip, thin-walled, smooth; conidiogenous cells
integrated, terminal or conidiophores occasionally reduced
to conidiogenous cells, 10–35 µm long, conidiogenous loci
VOLUME 6 · NO. 1
conspicuous, 1–2 µm diam, slightly thickened and darkened.
Conidia solitary or catenate, in simple chains, narrowly
cylindrical-obclavate, fusiform, straight to curved, (10–)20–
60 × 1.5–4 µm, (1–)2–5(–7)-septate, colourless, thin-walled,
smooth, apex subobtuse to subacute in solitary conidia,
conically truncate in catenate ones, base short to long
obconically truncate, 1–2 µm wide, hila somewhat thickened
and darkened.
Lectotype (designated here, MycoBank, MBT200446):
usA: Alabama: Lee County, Auburn, on Agrostis sp., 23 Jul.
1891, Newman & B. M. Duggar 2036 (CUP-A 2036#1(AL)).
Isolectotypes: CUP-A-2036#2(AL), CUP 40788.
Host range and distribution: Agrostis (gigantea, perennans,
scabra, Agrostis sp.), Sphenopholis obtusata, Poaceae
(Pooideae, Aveneae), ?Caucasus (Azerbaijan, Georgia),
North America (USA, Alabama, Idaho, North Dakota,
Oklahoma).
Notes: Due to the colourless conidia, we prefer to maintain C.
agrostidis as a species of Cercospora s. str. since results of
molecular sequence analyses have shown that species with
thickened, darkened conidiogenous loci and conidial hila
combined with colourless conidia, irrespective of whether
they are formed singly or in chains, belong to Cercospora
s. str. (Braun et al. 2013). Based on morphological similarity
31
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cercospora apii Fresen. s. lat. (sensu Crous & Braun
2003: 35).
(A) On Bromus inermis.
Notes: Cercospora collections on Bromus inermis and B.
marginatus were previously referred to as “C. festucae”
which is incorrect. The genera Bromus (Bromeae) and
Festuca (Poeae) are not closely allied and belong to distantly
related tribes of the Pooideae (Bouchenak-Khelladi et al.
2008). Material on Bromus marginatus was not available, but
two North Americal samples on B. inermis were examined.
One of them was a typical collection morphologically
assignable to Cercospora apii s. lat. (usA, Texas, College
Station, on Bromus inermis, Oct. 1949, M. D. Whitehead,
BPI 436347): Conidiophores in small to moderately large
fascicles, divergent to moderately dense, arising from small
stromatic hyphal aggregations, erect, straight, subcylindrical
or only slightly geniculate, unbranched, 30–200 × 3–6 µm,
pluriseptate, pale to medium brown throughout or tips paler,
occasionally subhyaline, thin-walled, smooth; conidiogenous
cells, integrated, terminal and occasionally intercalary, 10–40
µm long, with a single or only few conidiogenous loci, thickened
and darkened, 2–3.5 µm diam; conidia solitary, acicular, 40–
120 × 3–4 µm, 3–10-septate, hyaline, thin-walled, smooth,
apex subacute, base truncate, 2–3 µm wide, thickened and
darkened. The second sample is morphologically distinct
(see Cercospora sp.).
(B) On Zea mays.
(Fig. 2)
Synonym: Cercospora sorghi var. maydis Ellis & Everh.,
J. Mycol. 3: 15 (1887) [lectotype (designated here,
MycoBank, MBT200447): usA: Louisiana: Rapides
Parish, on Zea mays, 23 Jul. 1886, A. B. Langlois 613
(BPI 441565); isolectotypes: BPI 441551, NY 838620].
Cercospora sorghi f. maydis (Ellis & Everh.) Sacc., Syll.
Fung. 10: 656 (1892).
Fig. 1. Cercospora agrostidis (CUP-A 2036#1[AL], lectotype). A.
Conidiophore fascicle. B. Conidiophores. c. Conidia. Bar = 10 µm.
and re-examinations of type collections, Braun & Mel’nik
(1997) reduced Cercospora agrostidis to synonymy with
Passalora fusimaculans. The latter species is almost
circumglobal and undoubtedly conined to hosts of
Poaceae subfam. Panicoideae. Cercospora agrostidis is
conined to North America on a few species of Agrostis and
Sphenopholis obtusata, two genera belonging in subfamily
Pooideae (Aveneae). The two species are morphologically
barely distinguishable, although C. agrostidis tends to have
somewhat longer and wider conidia, but they are probably
biologically distinct, occurring on unrelated grasses, and not
conspeciic. We prefer to follow Chupp (1954) and maintain
two separate species. Cultures and results of molecular
sequence analyses are necessary for a inal taxonomic
conclusion. The identity of collections from Azerbaijan and
Georgia (Caucasus) are unconirmed (Braun & Mel’nik
1997).
32
Notes: Chupp (1954) mentioned that Cercospora sorghi var.
maydis from Zea mays is not able to infect Sorghum spp. and
possibly represents a separate species. Results of molecular
sequence analyses showed that sequences of Cercospora
sorghi var. maydis from Africa and North America cluster with
C. apii and C. beticola, respectively (Goodwin et al. 2001,
Crous et al. 2006), i.e. C. apii s. lat. can be transmitted to
maize. North American sequences clustered with C. apii s.
str., i.e. C. sorghi var. maydis must be considered a synonym
of the latter species, and a sequence from Africa with C.
beticola, both belonging to C. apii s. lat. The differentiation
between the two species is only possible on the basis of
multilocal sequence analyses (Groenewald et al. 2006,
2010). Type material of C. sorghi var. maydis has been reexamined and is characterised as follows: Often on faded
or necrotic leaves; caespituli scattered, punctiform, dark
brown to blackish; mycelium internal; stromata lacking or
small, brown, mostly substomatal; conidiophores in small
fascicles, divergent, emerging through stomata, erect,
straight, subcylindrical, not or only slightly geniculate near
the apex, unbranched, 20–180 × 4–6 µm, 2- to pluriseptate,
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Fig. 2. Cercospora apii [C. sorghi var. maydis] (BPI 441565,
lectotype). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.
Fig. 3. Cercospora aristidae (CUP 39097, holotype). A. Conidiophore
fascicle. B. Conidiophores. c. Conidia. Bar = 10 µm.
pale to medium brown throughout or tips somewhat paler,
thin-walled, smooth; conidiogenous cells integrated, terminal,
with a single or only few conidiogenous loci, 3–4 µm diam,
thickened and darkened; conidia solitary, acicular, 40–120
× 2–4.5 µm, pluriseptate, hyaline, thin-walled, smooth, apex
subacute, base truncate, 2–3 µm wide, hila thickened and
darkened.
Literature: Braun et al. (2002: 118), Crous & Braun (2003:
65).
cercospora aristidae Chupp,
Cercospora: 243 (1954).
(Fig. 3)
VOLUME 6 · NO. 1
Monograph
of
Illustration: Chupp (1954: 243, ig. 115).
Description: Leaf spots oval to oblong, 0.5–2 mm in length,
olivaceous to brown with reddish brown margin. Caespituli
amphigenous, but usually hypophyllous, punctiform, dark
brown. Mycelium internal. Stromata substomatal, 10–50
µm diam, subglobose, brown to dark brown. Conidiophores
in divergent fascicles, 5–20, arising from stromata, through
33
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Braun et al.
stomata, erect, straight to curved, subcylindrical to lexuous
or geniculate-sinuous in the fertile portion, unbranched, 20–
125 × 4–6 µm, pluriseptate, uniformly medium to medium
dark brown, thin-walled, smooth; conidiogenous loci 2–2.5
µm diam. Conidia solitary, acicular or subacicular, straight to
curved, 40–120 × 2.5–4 µm, pluriseptate, hyaline, thin-walled,
smooth, attenuated towards a pointed tip, base truncate or
only slightly attenuated at the base, about 2 µm wide, hila
somewhat thickened and darkened.
Holotype: usA: Alabama: Uniontown, on Aristida sp.,
Poaceae (Aristidoideae), 4 Sep. 1894, B. M. Duggar (CUP
39097).
Host range and distribution: Only known from the type
collection.
Note: This species belongs to the Cercospora apii complex.
cercospora arthraxonis M.S. Patil & Sawant, Indian
Phytopathol. 44: 15 (1991).
(Fig. 4)
Literature: Crous & Braun (2003: 67), Kamal (2010: 18).
Illustration: Patil & Sawant (1991: 16, igs 1–2).
Description: Leaf spots circular to angular, 2–5 mm diam.
Caespituli amphigenous. Mycelium internal. Stromata
substomatal, globose, pseudoparenchymatous, 30–45 µm
diam, brown. Conidiophores fasciculate, divergent, arising
from stromata, through stomata, erect, straight to curved,
geniculate in the upper fertile portion, unbranched, 37–155 ×
4.5–6 µm, narrowed and paler towards the tip, septate, brown,
thin-walled, smooth; conidiogenous cells integrated, terminal,
conidiogenous loci conspicuous, thickened and darkened.
Conidia solitary, acicular, straight to curved, 15–155 × 3 µm,
4–15-septate, hyaline, thin-walled, smooth, apex subobtuse
to pointed, base truncate, hila thickened and darkened.
Holotype: India: Maharashtra: Kolhapur, Amba, on Arthraxon
hispidus, 15 Oct. 1985, M. S. Patil (HCIO 39889).
Host range and distribution: On Arthraxon hispidus, Poaceae
(Panicoideae, Andropogoneae), Asia (India, Maharashtra).
Notes: This is undoubtedly a species of the Cercospora apii
s. lat. complex. The conidia were described as “cylindric,
attenuated towards the apex”, but the illustration clearly
shows acicular ones.
cercospora barretoana (U. Braun & Crous) U. Braun
& Crous, comb. nov.
MycoBank MB811240
(Fig. 5)
Basionym: Passalora fusimaculans var. barretoana U. Braun
& Crous, in Crous & Braun, Mycosphaerella and Anam.:
453 (2003).
Synonyms: Cladosporium piricularioides Dearn. & House,
Circ. New York State Mus. 24: 57 (1940), nom. inval.
34
Fig. 4. Cercospora arthraxonis (based on Patil & Sawant 1991: 16,
igs 1–2). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.
(Art. 39.1) [syntypes: usA: New York: Essex County,
Newcomb, on Panicum boreale, 17 Aug. 1924, H. D.
House (NYS 2365, DAOM 5741, NY 945783)].
Passalora barretoana (U. Braun & Crous) D.J. Soares, U.
Braun & R.W. Barreto, in Soares & Barreto, Australas. Pl.
Pathol. 35: 348 (2006).
Literature: Schubert (2005b: 220), Schubert & Braun (2005:
104), Bensch et al. (2012: 326), Phengsintham et al. (2013:
100).
Illustrations: Soares & Barreto (2006: 348, ig.
Phengsintham et al. (2013: 100, ig. 46, 101, ig. 47).
1),
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or olivaceous-brown, yellowish brown, sometimes paler
towards the apex, wall thin to somewhat thickened, 0.5–0.8
µm, smooth or almost so; conidiogenous cells integrated,
terminal or intercalary, occasionally conidiophores reduced
to conidiogenous cells, 10–65 µm long, conidiogenous loci
conspicuous, at irst terminal, later lateral, on shoulders formed
by sympodial proliferation, 2–10 per cell, 1.5–2(–3) µm diam,
apex truncate to slightly convex, somewhat thickened and
darkened. Conidia solitary or in unbranched chains, fusiform,
ellipsoid, subcylindrical to obclavate, 9–70 × (2.5–)3–5.5(–6)
µm, 0–4(–6)-septate, not constricted at the septa, hyaline or
subhyaline, wall thin or only slightly thickened, 0.3–0.5 µm,
smooth, apex rounded, attenuated to truncate, base truncate,
1–2 µm diam, hila somewhat thickened and darkened.
Holotype: Brazil: Rio de Janeiro, Comendaolos Venaricis
Reservoir, on Echinochloa polystachya, 13 Sep. 1989, R. W.
Barreto (K(M) IMI 345389).
Host range and distribution: On Echinochloa (esculenta,
polystachya),
Hymenachne
amplexicaulis,
Panicum
(boreale, Panicum sp.), Poaceae (Panicoideae, Paniceae),
Asia (Thailand), North America (USA, New York, Wisconsin),
South America (Brazil).
Notes: Due to the wider conidia, this species was originally
described as a variety of Passalora fusimaculans. Based
on several additional collections and clear morphological
differences, it was later raised to species rank (Soares &
Barreto 2006). Species with catenate, colourless conidia
and thickened, darkened conidiogenous loci and conidial
hila belong to Cercospora s. str. (see discussion under C.
fusimaculans).
cercospora bothriochloae U. Braun & Crous,
Mycotaxon 92: 396 (2005)
(Fig. 6)
Illustration: Braun & Crous (2005: 397, ig. 1).
Fig. 5. Cercospora barretoana (K(M) 345389, holotype). A.
Conidiophore fascicles. B. Conidiophore tip, c. Conidia. Bar = 10
µm.
Description: Leaf spots scattered, small, narrow, oval, fusoid
to oblong, 0.5–3 mm long and to 1 mm wide, or 3–8 mm diam,
yellowish brown, straw-coloured, reddish brown to dark brown,
often surrounded by a pale yellowish brown to olivacousbrown halo. Caespituli hypophyllous, scattered, but not effuse,
loose, pale brown. Mycelium internal, subcuticular; hyphae
2–4 µm wide. Stromata 10–45 µm diam, brown, composed
of swollen hyphal cells, circular to angular in outline, 4–10
µm diam, pale yellowish brown, brownish, amber-coloured,
smooth, walls slightly thickened. Conidiophores loosely
fasciculate, 3–19, arising from stromata, erumpent, erect,
straight to somewhat lexuous, unbranched, subcylindricaliliform to somewhat geniculate-sinuous, 20–180 × 3–6(–9)
µm, 0–6-septate, not constricted at the septa, pale brown
VOLUME 6 · NO. 1
Description: Leaf spots amphigenous, oblong, striate,
usually conined by veins, to 20 mm long and 1–2 mm
wide, brown, margin indeinite or occasionally with narrow
purplish violet border. Caespituli amphigenous, punctiform,
brown. Mycelium internal. Stromata substomatal, 10–30
µm diam, brown, composed of swollen hyphal cells, to 9
µm diam. Conidiophores in small to moderately large
fascicles, loose to moderately dense, arising from stromata,
emerging through stomata, erect, straight, subcylindrical to
slightly geniculate-sinuous, unbranched, 10–60 × 3–6 µm,
occasionally swollen at the base, to 10 µm, 0–1-septate,
subhyaline to pale olivaceous-brown, thin-walled, smooth;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, 10–40 µm long,
conidiogenous loci conspicuous, thickened and darkened,
1.5–2.5(–3) µm diam, thickened and darkened. Conidia
solitary, obclavate-subcylindrical, subfusiform, 15–45 × 3–4
µm, (0–)1–3(–4)-septate, colourless, thin-walled, smooth,
apex obtuse, base obconically truncate, 1.5–2 µm wide,
hilum somewhat thickened and darkened.
35
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Braun et al.
Fig. 6. Cercospora bothryochloae (NY 936943, holotype). A.
Conidiophore fascicle. B. Conidiophores. c. Conidia. Bar = 10 µm.
Holotype: usA: Kansas: Meade County, near State Lake, on
leaves of Bothriochloa saccharoides, Poaceae (Panicoideae,
Andropogoneae), 18 Jun. 1957, C. T. Rogerson R3803 (NY
936943).
Host range and distribution: Only known from the type
collection.
cercospora boutelouae Chupp & H.C. Greene,
Farlowia 1: 579 (1944).
(Fig. 7)
Literature: Chupp (1954: 244), Braun et al. (2002: 123),
Crous & Braun (2003: 86).
36
Fig. 7. Cercospora boutelouae (CUP 39229, lectotype). A.
Conidiophore fascicle. B. Conidiophore. c. Conidia. Bar = 10 µm.
Description: Leaf spots amphigenous, narrowly elliptical
to oblong, 0.5–4 mm in length, pale brown to blackish,
margin indeinite or with yellowish halo. Caespituli mainly
hypophyllous, in lines, dark. Mycelium internal. Stromata
lacking or small, composed of a few swollen hyphal cells,
brown. Conidiophores fasciculate, 2–14, divergent, arising
from internal hyphae or stromatic hyphal aggregations,
erect, straight to curved, subcylindrical, unbranched,
geniculate in the fertile apical portion, 20–100 × 3–6 µm,
sparingly septate, pale to medium brown, somewhat paler
towards the tip, thin-walled, smooth; conidiogenous cells
integrated, terminal, 10–30 µm long, conidiogenous loci
thickened and darkened, 1.5–2 µm diam. Conidia solitary,
acicular or subacicular to obclavate, straight to curved,
occasionally sigmoid, 20–80 × 3–5 µm, 2–8-septate, hyaline
IMA FUNGUS
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to subhyaline (pale olivaceous), thin-walled, smooth,
apex obtuse to subacute, base truncate or short to long
obconically truncate, about 2–2.5 µm wide, hila thickened
and darkened.
Lectotype (designated here, MycoBank, MBT200448):
usA: Wisconsin: Dane County, Madison, on Bouteloua
curtipendula [racemosa], 26 Jul. 1943, H. C. Greene (CUP
39229). Isolectotypes: BPI 433803, WIS.
Host range and distribution: On Bouteloua curtipendula,
Chondrosus (gracilis [Bouteloua gracilis], hirsutus [Bouteloua
hirsuta]), Poaceae (Chloridoideae, Cynodonteae), North
America (USA, Colorado, Illinois, Iowa, Oklahoma, South
Dakota, Virginia, Wisconsin).
Note: A true Cercospora s. str. distinct from C. apii s.lat. by
having obclavate conidia with obconically truncate base.
cercospora chusqueae Chupp,
Cercospora: 245 (1954).
(Fig. 8)
Monograph
of
Literature: Chupp (1954: 245), Crous & Braun (2003: 123).
Description: Leaf spots amphigenous, formed as small
specks, 0.5–1 mm diam, later forming long, narrow
streaks, 2–30 × 0.5–1 mm, medium to dark brown, margin
indeinite, but with yellow halo, surrounding tissue often
becoming necrotic, inally large leaf segments or entire
leaves discoloured, straw yellow. Caespituli hypophyllous,
punctiform to almost oblong pustulate, dark brown to
blackish. Mycelium internal. Stromata variable, almost
lacking to well-developed, large, oblong, 20–300 µm long,
dark brown to blackish. Conidiophores in moderately
large to large fascicles, arising from stromata, divergent
to dense, erect, straight to curved, cylindrical-iliform or
slightly to distinctly geniculate-sinuous, unbranched, 30–
160 × 3–5.5 µm, pluriseptate throughout, pale to medium
dark brown or olivaceous-brown, paler towards the tip,
wall thin to somewhat thickened, smooth; conidiogenous
cells integrated, terminal and intercalary, 10–40 µm long,
with a single or mostly several distinct conidiogenous loci,
1.5–2 µm diam. Conidia solitary, subcylindrical, obclavatecylindrical, fusiform, straight to curved, rarely sinuous, 20–
50 × 3–5 µm, 1–4(–5)-septate, hyaline, thin-walled, smooth,
apex obtuse to subacute, base obconically truncate, 1.5–2
µm wide, hila somewhat thickened and darkened.
Holotype: colombia: Lenquazaque, Cundinamarca, on
Chusquea sp., Poaceae (Bambusoideae, Bambuseae), 1
Dec. 1940, A. Franko (CUP 39412).
Host range and distribution: Only known from the type
collection.
Note: A true Cercospora s. str. distinct from C. apii s. lat.
by its small, cylindrical, obclavate-cylindrical to fusiform,
1–4(–5)-septate conidia.
VOLUME 6 · NO. 1
Fig. 8. Cercospora chusqueae (CUP 39412, holotype). A.
Conidiophore fascicle. B. Conidiophore. c. Conidia. Bar = 10 µm.
cercospora coicis N.D. Sharma & Mishra, J. Indian
Bot. Soc. 56: 131 (1977); as “coixii”.
(Fig. 9)
Synonyms: Cercospora coicis M.S. Patil & Sawant, Indian
Phytopathol. 44: 17 (1991), nom. illeg. (Art. 39.1)
[holotype: India: Maharashtra: Kolhapur, on Coix lacrymajobi, 22 Nov. 1985, M. S. Patil (HCIO 39892)].
Cercospora coicicola Kamal, Cercosporoid Fungi of India: 35
(2010), nom. nov.
Literature: Crous & Braun (2003: 131), Kamal (2010: 35–36).
Illustrations: Sharma & Mishra (1977: 132, igs 1–2), Patil &
Sawant (1991: 16, igs 3–4).
37
Braun et al.
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smooth, apex pointed or subobtuse, base truncate or only
slightly attenuated at the base, hila thickened and darkened.
Holotype: India: Madhya Pradesh: Jabalpur, Adhartal, on
Coix lacryma-jobi, Sep. 1975, N. D. Sharma (Herbarium,
Dept. of Mycol. & Pl. Pathol., J.N. Agric. Univ., Jabalpur,
India, No. 22).
Host range and distribution: On Coix lacryma-jobi
(Panicoideae, Andropogoneae), Poaceae, Asia (India,
Madhya Pradesh, Maharashtra).
Notes: Type material was not examined, but according to the
original description and illustration (Crous & Braun 2003),
this species may be a true Cercospora s. str. close to C. apii
s. lat. Kamal (2010) examined type material and conirmed
its position as a Cercospora species. He emphasized that
the homonymous C. coicis M.S. Patil & Sawant was distinct
from C. apii s. lat. by its smaller, broader, few-celled, almost
cylindrical conidia and introduced the new name C. coicicola
for this taxon. The latter species was probably based on
immature material with relatively short conidia. Young,
short, more or less cylindrical conidia were also described
and illustrated for C. coicis. The two species are probably
conspeciic and Cercospora on Coix lacryma-jobi is treated
as a single species.
cercospora cymbopogonicola U. Braun, nom. nov.
MycoBank MB811241
(Fig. 10)
Basionym: Cercospora sorghi var. cymbopogonis Govindu &
Thirum., Sydowia 8: 227 (1954), non C. cymbopogonis
J.M. Yen, 1977.
Literature: Crous & Braun (2003: 382), Kamal (2010: 88).
Illustration: Govindu & Thirumalachar (1954: plate 8, igs
33–34).
Fig. 9. Cercospora coicis (based on Sharma & Mishra 1977: 132, ig
1–2). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.
Description: Leaf spots amphigenous, circular to elliptical,
1–5 mm diam, with greyish brown to grey centre surrounded
by a darker margin, brownish to red. Caespituli amphigenous.
Mycelium internal. Stromata small, a few aggregated swollen
cells to prominent, substomatal, 15–35 µm diam, brown.
Conidiophores in divergent to sometimes dense fascicles,
2–10, arising from stromata, through stomata, erect,
subcylindrical to geniculate, unbranched, 15–45 × 3–5 µm,
0–3-septate, brown or olivaceous-brown, thin-walled, smooth;
conidiogenous cells intergrated, terminal, occasionally
intercalary or reduced to conidiogenous cells, conidiogenous
loci thickened and darkened. Conidia solitary, acicular,
shorter ones cylindrical or almost so, straight to somewhat
curved, 15–125 × 3–5 µm, 3–13-septate, hyaline, thin-walled,
38
Description: Leaf spots linear to irregular, conluent to
form long stripes often extending over the whole leaf
surface, medium brown or olivaceous. Caespituli mainly
epiphyllous. Mycelium internal. Stromata composed of a
few swollen hyphal cells, brown. Conidiophores fasciculate,
straight, subcylindrical to geniculate-sinuous, unbranched,
28.5–70 × 2.5–4 µm, 1-septate, olivaceous to deep brown;
conidiogenous cells integrated, terminal, conidiogenous
loci thickened and darkened. Conidia solitary, obclavatecylindrical, 20–50 × 2–3.5 µm, 1–10-septate, hyaline, thinwalled, smooth, apex pointed, base obconically truncate, hila
thickened and darkened.
Holotype: India: Karnataka: Bangalore, Hebbal, on
Cymbopogon caesius, Poaceae (Panicoideae, Andropogoneae), 10 Feb. 1953, H. C. Govindu (probably not
preserved).
Host range and distribution: Only known from the type
collection.
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Fig. 11. Cercospora cynodontis (based on Pavgi & Singh 1971: 118,
igs 8–10). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.
Illustration: Pavgi & Singh (1971: 118, igs 8–10).
Fig. 10. Cercospora cymbopogonicola (based on Govindu &
Thirumalachar 1954: plate 8, igs 33–34). A. Conidiophore fascicle.
B. Conidia. Bar = 10 µm.
Notes: Type material of Cercospora sorghi var. cymbopogonis
could not be traced and is probably not preserved, but
based on the original description this taxon is distinct from
Cercospora sorghi by having much shorter and narrower
conidia and warrants consideration as a distinct species. The
relation to collections of “C. sorghi” on other Cymbopogon
spp. from other parts of the world (see “host range” under C.
sorghi) is unclear.
cercospora cynodontis Pavgi & R.A.
Mycopathol. Mycol. Appl. 43: 120 (1971).
(Fig. 11)
Description: Leaf spots amphigenous, also on the leaf
sheath, oblong-linear, scattered, 0.5–2 × 0.2–0.3 mm, brown,
centre later grey to greyish white. Caespituli amphigenous.
Mycelium internal. Stromata small, poorly developed, brown.
Conidiophores fasciculate, 2–5, loose to moderately dense,
erect, straight to somewhat geniculate-sinuous, unbranched,
20–55 × 3–5 µm, 0–2-septate, olivaceous-brown, paler
towards the tip, thin-walled, smooth; conidiogenous
cells integrated, terminal or conidiophores reduced to
conidiogenous cells, conidiogenous loci probably somewhat
thickened and darkened. Conidia solitary, obclavatecylindrical to cylindrical, 17.5–65(–105) × 2.5–4 µm,
0–5-septate, subhyaline, thin-walled, smooth, apex obtuse,
base obconically truncate, hila probably somewhat thickened
and darkened.
Singh,
Types: India: Uttar Pradesh: Varanasi, on Cynodon dactylon,
10 Oct. 1965, R. A. Singh, MSP No. 344 (K(M) IMI 129666
and HCIO – syntypes).
Literature: Crous & Braun (2003: 150), Kamal (2010: 38).
VOLUME 6 · NO. 1
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Braun et al.
Host range and distribution: On Cynodon dactylon, Poaceae
(Chloridoideae, Cynodonteae), Asia (India, Uttar Pradesh;
Malaysia).
Notes: Type material of this species held at IMI was examined
and found to be in poor condition and without conidia. Only
a few conidiophores, as described in the original description
and illustration were found. Detailed examinations of the
conidiogenous loci were impossible. Syntype material
deposited at HCIO was not available. This species requires
leptotypiication. The colourless conidia of this species
indicate it belongs to Cercospora s. str.
cercospora digitariae J. Kranz, Sydowia 19: 74
“1965” (1966).
(Fig. 12)
Literature: Crous & Braun (2003: 161), Kamal (2010: 41).
Illustration: Kranz (1966: 75, ig. 2).
Description: Leaf spots elliptical to rounded, 3–4 mm diam,
brown, at irst indistinct, inally often conluent. Caespituli
amphigenous, punctiform, dark brown. Mycelium internal.
Stromata substomatal, 10–30 µm diam, almost colourless
to brown. Conidiophores in well-developed, large fascicles,
10–30, arising from stromata, through stomata, erect,
straight, subcylindrical-conical to somewhat geniculatesinuous, straight to curved, unbranched, 5–45 × 2–3 µm,
usually aseptate, occasionally with a single indistinct septum,
subhyaline to light brown, thin-walled, smooth; conidiophores
usually reduced to conidiogenous cells, conidiogenous loci
conspicuous, thickened and darkened, 1–1.5 µm diam.
Conidia solitary, narrowly subcylindrical, subacicular to
slightly cylindrical-obclavate or fusiform, straight to curved,
occasionally sigmoid, 30–100 × 1–2 µm, 3–8(–11)-septate,
hyaline, thin-walled, smooth, apex acute to subobtuse, base
short obconically truncate, 0.5–1.5 µm wide, hila slightly
thickened and darkened.
Holotype: guinea: Kindia, on Digitaria longilora, Jul. 1962, J.
Kranz (K(M) IMI 95634).
Host range and distribution: On Digitaria (abyssinica [mutica,
scalarum], exilis, insularis, longilora, stricta, Digitaria sp.),
Poaceae (Panicoideae, Paniceae), Africa (Guinea, Kenya,
Nigeria, Uganda), Asia (India, Chandigarh), Oceania (New
Caledonia), South America (Venezuela).
Note: A true Cercospora s. str. distinct from C. apii s. lat.
cercospora echinochloae Davis, Trans. Wisconsin
Acad. Sci. 18: 106 (1915).
(Fig. 13)
Literature: Chupp (1954: 245), Pavgi & Singh (1971: 119),
Crous & Braun (2003: 169), Kamal (2003: 42), Braun &
Urtiaga (2013: 592).
Exsiccatae: Davis, Fungi Wiscon. Exs. 14.
40
Fig. 12. Cercospora digitariae (K(M), IMI 95634, holotype). A.
Conidiophore fascicle. B. Conidiophore tip. c. Conidia. Bar = 10 µm.
Description: Leaf spots amphigenous, oblong, between
veins, 3–19 × 0.3–0.5 mm, reddish brown, later with pale
centre. Caespituli amphigenous, mainly hypophyllous,
punctiform, delicate, dark brown, often in lines. Mycelium
internal. Stromata lacking or small, substomatal, brown.
Conidiophores in small, loose to moderately dense
fascicles, 2–15, arising from stromata, emerging through
stomata, erect, subcylindrical to moderately geniculatesinuous, unbranched or occasionally with a short lateral
branchlet, 10–80 × 3–7(–8) µm, 0–4-septate, yellowish,
subhyaline, uniformly pale to medium olivaceous-brown,
thin-walled, smooth; conidiogenous cells integrated, terminal,
occasionally intercalary, occasionally conidiophores reduced
to conidiogenous cells, 10–30 µm long, sympodial, with a
single or several conidiogenous loci, conspicuous, somewhat
IMA FUNGUS
cercosporoid fungi 3
Host range and distribution: On Echinochloa (colona, crusgalli, polystachya, Echinochloa sp.), Poaceaea (Panicoideae,
Paniceae), Asia (India, Uttar Pradesh; Papua New Guinea),
Central and South America (Panama, Venezuela), North
America (USA, Iowa, Kansas, North Dakota, Wisconsin),
Oceania (Fiji, New Caledonia), West Indies (Cuba).
ART I CLE
galli, 9 Aug. 1913, J. J. Davis (BPI 436024). Isolectotypes:
CUP 39736, WIS and Davis, Fungi Wiscon. Exs. 14, e.g. BPI
436026, 868177 and CUP.
Notes: A true Cercospora s. str. distinct from C. apii s. lat.
by having consistently obclavate-cylindrical conidia. Records
on Cyperus rotundus are doubtful. Chupp (1954) mentioned
that he examined a collection on Echinochloa crus-galli from
North Dakota, which morphologically agreed with C. sorghi,
while other material from Venezuela deviated by longer and
narrower conidia.
cercospora eleusines Munjal, Lall & Chona, Indian
Phytopathol. 14: 181 (1961); as “eleusinis”.
(Fig. 14)
Literature: Crous & Braun (2003: 173), Kamal (2010: 43).
Illustration: Munjal et al. (1961: 183, ig. 3).
Description: Leaf spots oblong, 2–8 × 0.5–2 mm, also at
sheaths, centre greyish white, margin olivaceous-brown
to dark brown. Caespituli hypophyllous. Mycelium internal.
Stromata subglobose, small or to 50 µm diam, brown.
Conidiophores in fascicles, few to numerous, divergent
to dense, arising from stromata, erect, straight to curved,
subcylindrical to geniculate-sinuous or subnodulose,
unbranched, 25–300 × 4–5 µm, pluriseptate, brown, paler
towards the tip, thin-walled, smooth; conidiogenous cells
integrated, terminal, occasionally intercalary, 10–40 µm
long, conidiogenous loci thickened and darkened, 1–1.5(–2)
µm diam. Conidia solitary, acicular to narrowly obclavatecylindrical, 50–260 × 3–4 µm, pluriseptate, hyaline, apex
subacute to subobtuse, base truncate to obconically truncate,
1–2 µm wide, hila thickened and darkened.
Holotype: India: Uttarakhand: Nainital, Kathgodam, Eleusine
coracana, 23 Oct. 1959, J. N. Kapoor (HCIO 26848).
Fig. 13. Cercospora echinochloae (BPI 436024, lectotype). A.
Conidiophore fascicle. B. Conidiophores. c. Conidia. Bar = 10 µm.
Host range and distribution: On Eleusine (coracana, Eleusine
sp.), Poaceae (Chloridoideae, Eragrostideae), Asia (India,
Uttarakhand; Nepal, Thailand).
thickened and darkened, 1.5–2 µm diam. Conidia solitary,
obclavate-cylindrical, straight to somewhat curved, 20–65(–
95) × 2–5 µm, 0–7-septate, hyaline or subhyaline, with a
pale greenish tinge (i.e. wall colourless, but content of the
cells pale greenish), thin-walled, smooth, apex subacute or
subobtuse, base truncate 1.5–2 µm wide, hila thickened and
darkened.
Note: This is a true Cercospora s. str. distinct from C. apii s.
lat. by having smaller conidiogenous loci, 1–1.5(–2) µm wide,
and acicular to narrowly obclavate-cylindrical conidia.
Lectotype (designated here, MycoBank MBT200449): usA:
Wisconsin: Sauk County, Devil’s Lake, on Echinochloa crus-
VOLUME 6 · NO. 1
cercospora eragrostidis McKenzie & Latch,
New Zealand J. Agric. Res. 27: 113 (1984); as
“eragrostis”.
(Fig. 15)
Literature: Crous & Braun (2003: 175).
41
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Braun et al.
Fig. 15. Cercospora eragrostidis (PDD 43152, holotype). A.
Conidiophore fascicles. B. Conidia. Bar = 10 µm.
Fig. 14. Cercospora eleusines (based on Munjal et al. 1961: 183,
ig. 3). A. Conidiophore fascicles. B. Conidiophore tip. c. Conidia.
Bar = 10 µm.
Illustration: McKenzie & Latch (1984: 114, ig. 1A).
Description: Leaf spots amphigenous, elliptical-linear, centre
pale, margin pale brown. Caespituli hypophyllous. Mycelium
internal; hyphae branched, septate, pale olivaceous, 2–3 µm
wide. Stromata substomatal, 30–55 × 25–30 µm, medium
brown. Conidiophores in well-developed fascicles, to 30,
divergent to dense, erect, straight to lexuous, subcylindricalconical to somewhat geniculate in the fertile portion,
unbranched, 15–30(–50) × 3.5–5.5 µm, 0–1(–2)-septate,
olivaceous-brown, paler towards the tip, thin-walled, smooth;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, polyblastic, sympodial,
42
conidiogenous loci thickened and darkened, slightly
prominent, 1.25–2 µm diam. Conidia solitary, obclavatecylindrical, straight to usually somewhat curved, 30–120
× (2–)3–3.5(–4) µm, (0–)3–6(–8)-septate, hyaline, pink in
mass, thin-walled, smooth, apex rounded to subacute, base
short obconically truncate, 1.25–2 µm wide, hila somewhat
thickened and darkened.
Holotype: New Zealand: Auckland, Waitakere Range,
on Eragrostis brownei, 22 Apr. 1975, J. M. Dingley (PDD
43152).
Host range and distribution: On Eragrostis brownei, Poaceae
(Chloridoideae, Eragrostideae), New Zealand.
Note: A true Cercospora s. str. distinct from C. apii s. lat.
IMA FUNGUS
cercosporoid fungi 3
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cylindrical, ellipsoid, obovoid, obclavate, fusiform, straight or
almost so, 10–35 × 3–7.5 µm, (0–)1–4(–6) septate, hyaline
or subhyaline, smooth, apex rounded, base short obconically
truncate, 1–1.5 µm wide, hila somewhat thickened and
darkened-refractive.
Holotype: Australia: Queensland: Mareeba, on Eremochloa
bimaculata, 30 Apr. 1987, J. L. Alcorn (BRIP 15782). Isotype:
K(M) IMI 321201.
Host range and distribution: On Eremochloa bimaculata,
Poaceae
(Panicoideae,
Andropogoneae),
Australia
(Queensland).
Notes: Due to at least partly catenate conidia and short,
broad conidia with few septa, this species is passalora-like
and resembles former Phaeoramularia species. Its position in
Cercospora s. str. (Groenewald et al. 2013) was determined
by means of molecular sequence analyses (ITS and LSU),
which provides additional proof that passalora-like species
with colourless conidia belong to Cercospora, even in rare
cases when conidia are formed in chains.
cercospora festucae Hardison, Mycologia 37: 492
(1945).
(Fig. 17)
Literature: Chupp (1954: 246), Crous & Braun (2003: 183).
Fig. 16. Cercospora eremochloae (K(M) IMI 321201, isotype). A.
Conidiophores. B. Conidiophore tips. c. Conidia. Bar = 10 µm.
cercospora eremochloae R.G. Shivas & A.J. Young,
Persoonia 26: 111 (2011).
(Fig. 16)
Illustration: Shivas & Young, in Crous et al. (2011b: 110, plate,
without number).
Description: Leaf spots amphigenous, narrowly elliptical,
often elongated, to 7 cm long, 0.5–1.5 mm wide, smaller
leaf spots vein-limited, centre orange to pale brown with
darker reddish to purplish brown diffuse margin. Caespituli
hypophyllous, punctiform or inconspicuous, dark brown.
Mycelium internal. Stromata reddish brown, immersed,
erumpent, usually substomatal, to 40 µm diam. Conidiophores
in small, loose fascicles, 2–10, arising from stromata, usually
through stomata, erect, geniculate-sinuous, unbranched to
branched, 100–275 × 4–6 µm, somewhat attenuated towards
the apex, pluriseptate (to 20), reddish brown, paler towards
the tip, wall thin, smooth; conidiogenous cells integrated,
terminal, sympodial, geniculate, mono- to polyblastic,
conidiogenous loci thickened and darkened, 1–1.5 µm wide.
Conidia solitary or in short branched or unbranched chains,
VOLUME 6 · NO. 1
Description: Leaf spots oval to oblong, 0.5–4 mm in length,
centre grey, margin purplish. Caespituli amphigenous, but
mainly hypophyllous. Mycelium internal. Stromata lacking or
only formed as small aggregations of a few swollen hyphal
cells, brown. Conidiophores in small to moderately large,
loose to dense fascicles, mostly 3–8, arising from internal
hyphae or stromatic hyphal aggregations, erect, straight,
subcylindrical to somewhat geniculate-sinuous, unbranched,
20–800 × 3–5 µm, with few to numerous septa, pale to medium
olivaceous-brown, paler towards the tip; conidiogenous
cells integrated, terminal, conidiophores rarely reduced to
conidiogenous cells, about 10–30 µm long, conidiogenous
loci thickened and darkened, 2–3 µm diam. Conidia solitary,
acicular, shorter conidia subacicular, fusoid-obclavate,
straight to curved, occasionally somewhat sigmoid, 30–300
× 2–5 µm, 3- to pluriseptate, hyaline, apex pointed, base
truncate, subtruncate to short obconically truncate, (1.5–)2–3
µm wide, hila thickened and darkened.
Holotype: usA: Kentucky: Fayette County, Lexington, on
Festuca arundinacea, 23 Aug. 1943, J. R. Hardison (CUP
39807). Topotype (from July 1944): BPI 436348.
Host range and distribution: On Festuca arundinacea [elatior],
Poaceae (Pooideae, Bromeae, Poeae), North America (USA,
Georgia, Kentucky, Mississippi, Oklahoma, Oregon, Texas).
Notes: This is a true Cercospora s. str. close to C. apii s. lat.
The lengths of the conidiophores and conidia are variable,
ranging from uniformly short to long conidiophores to 800
µm, and conidia to 300 µm. Short conidia may be narrowly
43
Braun et al.
ART I CLE
is also incorrect and can currently only be referred to as C. apii
s. lat. Katsuki (1966) provided a description of this material:
stromata none; conidiophores solitary or 2–3 stalks, medium
olivaceous-brown near the base, paler and sometimes
narrower toward the tip, almost straight, not branched, tips
rounded, sparingly septate, 47–90 × 2–3um; conidia acicular,
curved or undulate, indistinctly 2–7 septate, base truncate,
tip acute, hyaline, 21–54 × 2–3 um. A part of the material
concerned had been sent to C. Chupp who considered it as
a collection belonging to C. apii s. str. A sample maintained
in Japan was recently re-examined by C. Nakashima, and
found to be devoid of any conidiophores and conidia.
cercospora fusimaculans G.F. Atk., J. Elisha Mitchell
Sci. Soc. 8: 50 (1892).
(Fig. 18)
Synonyms: Cercospora panici Davis, Trans. Wisconsin Acad.
Sci. 19: 714 (1919) [holotype: usA: Wisconsin: Shiocton,
on Panicum latifolium, 15 Aug. 1917, J. J. Davis (WIS)].
Cercosporina panici (Davis) Sacc., Syll. Fung. 25: 904 (1931).
Cercospora panici-miliacei Sawada, Rep. Gov. Agric. Res.
Inst. Formosa 51: 131 (1931) [syntype: taiwan: Taichung,
on Panicum miliaceum, 2 Aug. 1928, K. Sawada; 3 Aug.
1928, K. Sawada (TNS-F-220504)].
Phaeoramularia fusimaculans (G.F. Atk.) X.J. Liu & Y.L. Guo,
Acta Phytopathol. Sin. 12: 9 (1982).
Passalora fusimaculans (G.F. Atk.) U. Braun & Crous,
in Crous & Braun, Mycosphaerella and Anam.: 192
(2003).
Literature: Saccardo (1892: 655; 1931: 904), Vassiljevsky &
Karakulin (1937: 271), Chupp (1954: 246), Vasudeva (1963:
112), Katsuki (1965: 33), Ellis (1976: 260), McKenzie & Latch
(1984: 115), Hsieh & Goh (1990: 141–143), Crous & Braun
(1996: 272), Braun & Melnik (1997: 61), Guo et al. (2003:
144–146), Braun & Crous (2005: 410), Kamal (2010: 120–
121).
Illustrations: Chupp (1954: 243, ig. 116), Vasudeva (1963:
112, ig. 72), McKenzie & Latch (1984: 114, ig. 1D), Ellis
(1976: 260, ig. 197A), Hsieh & Goh (1990: 141, ig. 109, 144,
ig. 110, as Phaeoramularia sp.), Guo et al. (2003: 145, ig.
91).
Fig. 17. Cercospora festucae (BPI 436348, topotype). A.
Conidiophore fascicle. B. Conidiophore tips. c. Conidia. Bar = 10
µm.
obclavate-cylindrical, with truncate to obconically truncate
bases. Collections on Bromus inermis and B. marginatus
were previously referred to as C. festucae, which is incorrect.
The genera Bromus (Bromeae) and Festuca (Poeae) are
not closely allied. The two tribes belong to the Pooideae but
they are only distantly related (Bouchenak-Khelladi et al.
2008). Material on Bromus marginatus was not available,
but two collections on B. inermis from Texas were reexamined and found to belong to two different species, each
morphologically distinct from C. festucae (see Cercospora apii
[A] and Cercospora sp.). A record of C. festucae on Bromus
marginatus from Japan (Katsuki 1966, Crous & Braun 2003)
44
Description: Leaf spots amphigenous, oval, elliptical, fusiform
to oblong or irregular, 0.5–4 × 0.5–2 mm, when oblong or
conluent to 10 mm in length, centre brownish to dingy
grey, margin reddish to dark brown, sometimes entire spots
uniformly brown. Caespituli amphigenous, often epiphyllous,
delicate to distinctly punctiform, scattered, dark. Mycelium
internal; hyphae branched, septate, 1.5–5 µm wide,
subhyaline to pale olivaceous. Stromata lacking or almost so
to developed, but not very large, immersed to substomatal,
10–35 µm diam, brown. Conidiophores in divergent to
occasionally dense fascicles, 2–30, arising from internal
hyphae or stromata, through stomata or erumpent, erect,
straight, subcylindrical or somewhat attenuated towards
the tip to moderately geniculate-sinuous, unbranched,
10–70(–100) × 2.5–4(–5) µm, 0–3(–4)-septate, subhyaline,
pale olivaceous-brown to medium brown, pigmentation
IMA FUNGUS
cercosporoid fungi 3
Isolectotypes: CUP-A-2945#2(AL), CUP-A-2945#3(AL).
uniform throughout or paler towards the tip, thin-walled,
smooth; conidiophores reduced to conidiogenous cells or
conidiogenous cells integrated, terminal, 10–30 µm long,
with conspicuous conidiogenous loci, 1–1.5(–2) µm diam,
thickened and darkened. Conidia solitary or catenate, in
simple chains, narrowly obclavate-cylindrical, subacicular,
(10–)20–100 × 1.5–3 µm, 1–7-septate, colourless, thinwalled, smooth, apex subacute to subtruncate or conically
truncate in catenate conidia, base subtruncate to short
obconically truncate, 1–1.5 µm wide, hila slightly thickened
and darkened.
Lectotype (designated here, MycoBank, MBT200450): usA:
Alabama: Lee County, Auburn, on Panicum dichotomum,
15 Aug. 1891, B. M. Duggar 2054 (CUP-A-2054#1(AL)).
VOLUME 6 · NO. 1
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Fig. 18. Cercospora fusimaculans (CUP-A-205#1(AL), lectotype). A.
Conidiophore fascicles. B. Conidophore tips. c. Conidia. Bar = 10
µm.
Host range and distribution: On Brachiaria (brizantha,
decumbens, dictyoneura, eminii, fasciculata, humidicola,
jubata, reptans, ruziziensis, serrata, subquadripara
[miliformia]), Beckeropsis sp., Cenchrus (hordeoides
[Pennisetum hordeoides], pedicellatus [P. pedicellatum],
polystachion [P. polystachyon], purpureus [P. purpureum],
spicatus [P. glaucum]), Chasmopodium (caudatum,
Chasmopodium sp.), Digitaria (cognata [Leptoloma
cognatum], insularis, ischaemum, abyssinica [scalarum]),
Echinochloa (colona, crus-galli), Entolasia marginata,
Ichnanthus sp., Oplismenus undulatifolius, Panicum
(acuminatum [implicatum, paciicum], antidotale, boscii,
clandestinum, dichotomilorum, dichotomum, laetum,
latifolium, laxilorum [xalepense], leibergii, maximum,
mertensii,
miliaceum,
oligosanthes
[scribnerianum],
perlongum,
plicatum
[praecocius],
portoricense
[columbianum], virgatum, wilcoxianum, Panicum spp.),
Paspalidium geminatum, ?Rottboellia cochinchinensis
[exaltata], Setaria (barbata, homonyma [aequalis, lancea],
plicata, pumila [pallidifusca]), Sorghum (bicolor, halepense),
Stenotaphrum (pallens, secundatum), Urochloa panicoides
[Panicum javanicum], Zea mays, Poaceae (Panicoideae)
[unresolved records on Bouchloe dactyloides [Bouteloa
dactyloides] and Eleusine coracana, Chloroideae], Africa
(Botswana, Ethiopia, Ghana, Guinea, Ivory Coast, Kenya,
Malawi, Nigeria, Rwanda, Sierra Leone, South Africa,
Sudan, Tanzania, Togo, Uganda, Zambia, Zimbabwe), Asia
(Brunei, China, India, Japan, Korea, Malaysia, Papua New
Guinea, Philippines, Taiwan, Thailand), Australia, Caucasus
(Azerbaijan, Georgia), Central and South America (Bolivia,
Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French
Guiana, Guatemala, Guyana, Honduras, Nicaragua, Panama,
Peru, Venezuela), Europe (France, Russia), New Zealand,
North America (Mexico; USA, Alabama, Florida, Iowa, Idaho,
Illinois, Kansas, North Carolina, North Dakota, Oklahoma,
Oregon, Texas, Virginia, West Virginia, Wisconsin), Oceania
(Fiji, Guam, New Caledonia, Palau, Samoa, Solomon
Islands, Tonga, Vanuatu), and West Indies (Cuba, Dominican
Republ., French Antilles, Guadeloupe, Jamaica, Martinique,
Puerto Rico, Trinidad and Tobago, Virgin Islands).
Notes: Since the conidia of C. fusimaculans are colourless,
we prefer to maintain this taxon as a species of Cercospora
s. str. as results of molecular sequence analyses have shown
that species with thickened, darkened conidiogenous loci and
conidial hila combined with colourless conidia, irrespective
of whether they are formed singly or in chains, belong to
Cercospora s. str. (Braun et al. 2013). Cercospora agrostidis
on Agrostis and Sphenopholis spp. (Poaceae, Pooideae,
Aveneae) in North America, previously reduced to synonymy
with C. fusimaculans (Braun & Mel’nik 1997, Crous & Braun
2003), is now at least tentatively maintained as a separate
species (see comments under C. agrostidis). Hsieh & Goh
(1990) described conidia to 200 µm long, but conidia longer
than 100 µm have not been found in our examinations.
Rottboellia cochinchinensis [R. exaltata] has been recorded
as a host of C. fusimaculans. These records are doubtful and
probably represent C. rottboelliae.
45
Braun et al.
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scarce aerial mycelium, pale mouse grey, zonate, faintly
rosy-vinaceous towards the margin, reverse fuscous-black;
on malt extract agar 3 cm diam, lat, radially wrinkled, margin
entire, grey-olivaceous, reverse black.
Holotype: Australia: Northern Territory: Victoria River Downs,
S 15º 36’ 05”, E 131º 12’ 49”, on leaves of Ischaemum
australe, 20 Apr. 2012, R.G. Shivas (BRIP 56010, including
ex-type strain).
Host range and distribution: On Ischaemum australe,
Poaceae (Panicoideae, Andropogoneae), Australia (Northern
Territory, Western Australia).
Notes: Cercospora ischaemi is known from the type collection
and a second sample from Western Australia (BRIP 51367).
A specimen (BRIP 4473) on Ischaemum australe from
Queensland had comparable symptoms to C. ischaemi, but
according to Shivas et al. (2014) morphological examination
indicated the fungus differed and warranted further study.
Cercospora ischaemi, together with C. eremochloae, are
recently described species from native Australian tropical
grasses (Crous et al. 2011). Cercospora coniogrammes
(JX143583) on Coniogramme (Pteridaceae) from Australia
had the highest genetical identity (96%, 496/518 identical
base pairs) to C. ischaemi in a BLAST search of the ITS
region of rDNA.
Fig. 19. Cercospora ischaemi (based on Shivas et al. 2014: 4, ig. 2).
A. Conidiophore fascicles. B. Conidia. Bar = 10 µm.
cercospora ischaemi R.G. Shivas, Marney &
McTaggart, Fungal Biol. (2014), http://dx.doi.
org/10.1016/j.funbio.2014.09.004
(Fig. 19)
Illustration: Shivas et al. (2014: 4, ig. 2)
Description: Leaf spots amphigenous, linear to narrowly
ellipsoidal, bordered by parallel leaf veins, to 1 cm long and
1 mm wide, dark reddish brown to dark brown, with a narrow
yellowish diffuse halo to 0.5 mm wide, scattered, of similar
appearance on upper and lower leaf surfaces. Caespituli
epiphyllous, inconspicuous. Mycelium internal. Stromata
absent. Conidiophores erumpent through the cuticle, in
loose fascicles of 2–5, erect, subcylindrical, sometimes
geniculate, 10–40 × 3–5 μm, subhyaline to pale brown, thinwalled, smooth; conidiogenous cells terminal, subcylindrical,
sympodial, hyaline, smooth, polyblastic; conidiogenous loci
conspicuous, lat, circular, thickened and darkened, 1.5–
2.0 μm wide. Conidia solitary, obclavate, with a narrowly
obconically truncate base and then attenuated towards the
apex, 60–120 × 4–5 μm, 1–3-septate, hyaline, smooth, hila
thickened and darkened-refractive, 1.5–2.0 μm wide.
In vitro: (in the dark, 23°C, after 4 wk): Colonies on potatodextrose agar 5 cm diam, lat with scarce aerial mycelium,
pale mouse grey with irregular pale and darker patches,
margin irregularly crenate; reverse fuscous-black and paler
towards the margin. On oatmeal agar 2.5 cm diam, lat with
46
cercospora janseana (Racib.) O. Constant., Cryptog.
Mycol. 3: 63 (1982).
(Fig. 20)
Basionym: Napicladium janseanum Racib., Parasitische
Algen und Pilze Javas 2: 41 (1900).
Synonyms: Passalora janseana (Racib.) U. Braun,
Schlechtendalia 5: 39 (2000).
Cercospora oryzae Miyake, Bot Mag. Tokyo 23 (267): 139
(1909) [holotype: Japan: Ehime: Agricultural Experiment
Station, on Oryza sativa, Sep. 1907, I. Miyake (not traced,
probably not preserved)].
Sphaerulina oryzina Hara, Diseases of the rice plant (Japan):
144 (1918) [holotype: Japan: Gifu: Kawaue, on Oryza
sativa, 25 Oct. 1917 (not traced, probably not preserved)].
Cercospora oryzae var. ruipogonis R.A. Singh & Pavgi,
Sydowia 21: 176 “1967” (1968) [syntypes: India: Uttar
Pradesh: Varanasi, on Oryza ruipogon, 1 Nov. 1964, R.
A. Singh (HCIO, MSP no. 348].
Literature: Saccardo (1913: 1431), Vassiljevsky & Karakulin
(1937: 272), Chupp (1954: 249), Vaduseva (1963: 156),
Mulder & Holliday (1974c), Ellis (1976: 262), Sivanesan
(1984: 271), Hsieh & Goh (1990: 135), Teng (1996: 546),
Crous & Braun (1996: 280, 299), Braun & Sivapalan (1999:
5), Crous & Braun (2003: 231), Guo et al. (2003: 91–92),
Kamal (2010: 70, 124).
Illustrations: Mulder & Holliday (1974c, ig., unnumbered),
Ellis (1976: 261, ig. 198B), Constantinescu (1982: 64, ig. 1),
Sivanesan (1984: 271, ig. 152), Hsieh & Goh (1990: 137, ig.
104), Guo et al. (2003: 92, ig. 58).
IMA FUNGUS
cercosporoid fungi 3
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darkened, 0.5–1.5 µm diam. Conidia solitary, cylindrical to
obclavate, straight to curved, (10–)15–65(–85) × 3–6.5 µm,
(1–)3–5(–12)-septate, hyaline or subhyaline, thin-walled,
smooth, apex obtuse to subacute, base obconically truncate,
1–2 µm wide, hilum somewhat thickened and darkened.
Sexual morph: Ascomata scattered to gregarious,
immersed, 60–100 µm diam. Asci cylindrical-clavate, 50–60
× 10–13 µm, 8-spored. Ascospores fusoid, 20–33 × 4–5 µm,
3-septate, colourless.
Lectotype (designated by Constantinescu 1982): Indonesia:
Java: Bogor, on Oryza sativa, 1900, M. Raciborski (KRA).
Isolectotypes: BUCM 59761, ZT.
Host range and distribution: On Oryza (barthii, latifolia,
sativa, ruipogon), Poaceae (Ehrhartoideae, Oryzeae),
widely distributed, Africa (Angola, Chad, Congo, Gabon,
Gambia, Ghana, Kenya, Madagascar, Malawi, Mozambique,
Niger, Nigeria, Somalia, South Africa, Sudan, Tanzania, Togo,
Zambia, Zimbabwe), Asia (Afghanistan, Bangladesh, Brunei,
Cambodia, China, India, Indonesia, Japan, Korea, Laos,
Malaysia, Myanmar, Nepal, Pakistan, Papua New Guinea,
Philippines, Sri Lanka, Syria, Taiwan, Thailand, Vietnam),
Australia, Central and South America (Argentina, Bolivia,
Brazil, Colombia, Costa Rica, El Salvador, Guatemala,
Guyana, Honduras, Nicaragua, Panama, Suriname,
Venezuela), North America (Mexico; USA, Alabama,
Arkansas, Florida, Louisiana, South Carolina, Texas),
Oceania (Fiji, Solomon Islands), and West Indies (Cuba,
Dominican Republ., Haiti, Puerto Rico, Trinidad and Tobago,
Virgin Islands).
Fig. 20. Cercospora janseana (P [PS 60-114]). A. Conidiophore
fascicles. B. Conidiophores. c. Conidia. Bar = 10 µm.
Description: Leaf spots amphigenous, linear, oval or elliptical,
2–15 × 0.5–3 mm, pale to dark brown, paler towards the
periphery or centre paler, margin indeinite or darker.
Caespituli amphigenous, mainly hypophyllous, between
veins. Mycelium internal. Stromata lacking or small, about
10–20 µm diam, substomatal, brown. Conidiophores solitary
or in small, loose fascicles, about 2–15, arising from internal
hyphae or stromata, through stomata, erect, straight to
curved, subcylindrical to geniculate-sinuous, sometimes
strongly curved to sinuous, unbranched, length variable,
occasionally uniformly short, 10–140(–160) × 3–6(–7)
µm, continuous to pluriseptate (0–12), pale to medium
brown, rarely darker, sometimes paler towards the tip, thinwalled, smooth; conidiogenous cells integrated, terminal,
occasionally reduced to conidiogenous cells, about 10–40
µm long, conidiogenous loci somewhat thickened and
VOLUME 6 · NO. 1
Notes: Reports of this species on Leptochloa mucronata
[iliformis]
(Chloridoideae,
Eragrostideae),
Cenchrus
purpureus [Pennisetum purpureum], Coix lacryma-jobi,
Imperata cylindrica, Panicum maximum and P. repens
(Panicoideae) are not conspeciic with Cercospora janseana
and probably belong to other Cercospora species with
obclavate-cylindrical conidia. Type material of Sphaerulina
oryzina could not be traced, but several other collections are
deposited at NIAES.
cercospora longipes E.J. Butler, Mem. Dept. Agric.
India, bot. ser. 1: 41 (1906).
(Fig. 21)
Literature: Chupp (1954: 248), Ellis (1976: 261), Yen & Sun
(1978: 394), Deighton (1979: 22), Hsieh & Goh (1990: 134),
Crous & Braun (2003: 255), Guo et al. (2005: 121–122),
Kamal (2010: 60).
Illustrations: Ellis (1976: 261, ig. 198A), Yen & Sun (1978:
395, ig. 1D–E), Hsieh & Goh (1990: 135, ig. 102), Guo et al.
(2005: 122, ig. 83).
Description: Leaf spots amphigenous, oval to linear, at irst
narrowly oval and reddish, later elongated, with brown centre
and yellowish halo, about 1–8 × 0.5–2 mm, sometimes
conluent, forming larger reddish brown blotches, to 14 mm
in length. Caespituli amphigenous, but mainly hypophyllous.
47
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Braun et al.
Fig. 22. Cercospora microlaenae (PDD 43153, holotype). A.
Conidiophore fascicles. B. Conidia. Bar = 10 µm.
MBT200451): India: Bihar: Pusa, on Saccharum oficinarum,
11 Dec. 1922, M. Taslim (BPI 437895).
Fig. 21. Cercospora longipes (BPI 437895, neotype). A. Conidiophore
fascicle. B. Conidiophore tips. c. Conidia. Bar = 10 µm.
Mycelium internal. Stromata lacking or small aggregations of
swollen hyphal cells, 10–25 µm diam, brown. Conidiophores
in small to moderately large fascicles, to 18, usually divergent,
arising from stromata, erect, straight, subcylindrical,
geniculate, simple or rarely branched , 30–280 × 3–7 µm,
pluriseptate, pale to medium dark brown throughout or
paler towards the tip, wall thin to slightly thickened, smooth;
conidiogenous cells integrated, terminal and intercalary, 10–
50 µm long, with a single to mostly several conidiogenous
loci, thickened and darkened, 1–2.5 µm diam. Conidia
solitary, obclavate, straight to curved, 30–120 × 3–6(–7) µm,
3–10-septate, hyaline, thin-walled, smooth, apex subacute,
base obconically truncate, 1.5–2 µm wide, hila thickened and
darkened.
Host range and distribution: On Saccharum (oficinarum,
spontaneum), Poaceae (Panicoideae, Andropogoneae), widely
distributed, Africa (Ethiopia, Kenya, Madagascar, Malawi,
Mauritius, Somalia, South Africa, Tanzania, Uganda, Zambia,
Zimbabwe), Asia (Afghanistan, Bangladesh, India, Indonesia,
Myanmar, Nepal, Papua New Guinea, Philippines, Sri Lanka,
Taiwan, Thailand), Central and South America (Argentina,
Brazil, Colombia, Costa Rica, Nicaragua, Panama), North
America (Mexico; USA, Alabama, Florida, Louisiana), Oceania
(Hawaii, Solomon Islands), and West Indies (Cuba, Dominican
Republ., Jamaica, Puerto Rico, Virgin Islands).
Notes: Chupp (1954) reduced Cercospora longipes to
synonymy with C. koepkei, but according to Deighton (1979:
22), C. koepkei is not the same as C. longipes. The latter
species is a true Cercospora s. str. distinct from C. apii s. lat.
cercospora microlaenae McKenzie & Latch, New
Zealand J. Agric. Res. 27: 115 (1984).
(Fig. 22)
Literature: Crous & Braun (2003: 274).
[Holotype: India: on Saccharum oficinarum, E. J. Butler
(not preserved)]. Neotype (designated here, MycoBank,
48
Illustration: McKenzie & Latch (1984: 114, ig. 1B).
IMA FUNGUS
cercosporoid fungi 3
ART I CLE
Description: Leaf spots amphigenous, sometimes also on the
leaf sheath, linear, forming black streaks, often vein-limited,
becoming chlorotic around the spots, chlorosis may spread
to cover the whole width of leaves, often dying from the tip
downwards. Caespituli amphigenous. Mycelium internal;
hyphae mainly composed of swollen cells, 2–5 mm wide,
hyaline to pale brown, smooth. Stromata substomatal, 15–
60 × 15–50 µm, pale to dark brown. Conidiophores in welldeveloped fascicles, to 30, divergent, arising from stromata,
through stomata, erect, straight to lexuous, subcylindrical,
fertile portion geniculate-sinuous, unbranched, 15–30(–40)
× (3–)3.5–4 µm, 0(–1)-septate, pale olivaceous-brown, paler
towards the tip, thin-walled, smooth; conidiophores usually
reduced to conidiogenous cells, conidiogenous loci thickened
and darkened, slightly prominent, 1.25–1.75 µm diam. Conidia
solitary, obclavate-cylindrical, straight to curved, (20–)30–
50(–70) × 2–2.5(–3) µm, (1–)4–6(–8)-septate, colourless,
thin-walled, smooth, apex rounded, base short obconically
truncate, 1–1.5 µm wide, hila thickened and darkened.
Holotype: New Zealand: Auckland, Mt. Albert, on Ehrharta
stipoides [Microlaena stipoides], 1 Dec. 1982, E. H. C.
McKenzie (PDD 43153).
Host range and distribution: On Ehrharta stipoides, Poaceae
(Ehrhartoideae, Ehrharteae), New Zealand.
Note: A true Cercospora distinct from the C. apii s. lat. complex
by having obclavate-cylindrical conidia with obconically
truncate base.
cercospora miscanthi Goh & W.H. Hsieh, Trans.
Mycol. Soc. Republ. China 2: 127 (1987).
(Fig. 23)
Synonym: Cercospora miscanthi Sawada, Rep. Gov. Agric.
Res. Inst. Taiwan 87: 83 (1944), nom. inval. (Art. 39.1)
[syntypes: taiwan: Taichung, on Miscanthus loridulus,
8 Oct. 1910, K. Sawada (NTU-PPE, hb. Sawada;
TNS-F-220475].
Literature: Chupp (1954: 249), Hsieh & Goh (1990: 134),
Crous & Braun (2003: 277), Guo et al. (2005: 123–124).
Fig. 23. Cercospora miscanthi (NTU-PPE, holotype). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
Illustrations: Hsieh & Goh (1990: 136, ig. 103), Guo et al.
(2005: 123, ig. 84).
Description: Leaf spots amphigenous, elliptical, 3–20
mm diam, grey with purplish brown border. Caespituli
amphigenous. Mycelium internal. Stromata lacking to welldeveloped, to 80 µm diam, dark brown. Conidiophores in
divergent fascicles, (0–)2–20, arising from internal hyphae
or stromata, erect, subcylindrical, unbranched, geniculate,
(15–)40–120 × 4–6 µm, 1–7-septate, brown, paler towards
the tip, thin-walled, smooth; conidiogenous cells integrated,
terminal and intercalary, with conspicuous conidiogenous
loci, thickened and darkened. Conidia solitary, obclavate,
straight to mostly curved at the apex, 40–100 × 3–4.5(–5)
µm, 3–7-septate, hyaline, thin-walled, smooth, apex acute,
base obconically truncate.
Holotype: taiwan: Taichung, on Miscanthus loridulus, 8
Oct. 1910, K. Sawada (NTU-PPE [hb. Sawada]). Isotype:
TNS-F-220475.
Host range and distribution: On Miscanthus loridulus
[japonicus], Poaceae (Panicoideae, Andropogoneae), Asia
(Taiwan).
cercospora oplismeni Lall, H.S. Gill & Munjal, Indian
Phytopathol. 14: 117 (1962).
(Fig. 24)
Literature: Crous & Braun (2003: 299), Kamal (2010: 70).
Illustration: Lall et al. (1962: 119, ig. 3).
VOLUME 6 · NO. 1
49
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Braun et al.
Fig. 24. Cercospora oplismeni (based on Lall et al. 1962: 119, ig. 3).
A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.
Description: Leaf spots linear, 1–5 mm long, sometimes
conluent, tan. Caespituli amphigenous. Mycelium internal.
Stromata subglobose, dark brown, to about 45 µm diam.
Conidiophores in small to larger, loose to dense fascicles,
somewhat geniculate, irregular in width, unbranched,
short, 6–28 × 3–6 µm, septate, pale olivaceous-brown;
conidiogenous loci small, thickened and darkened. Conidia
solitary, acicular, straight to curved, 30–130 × 2–3 µm,
pluriseptate, hyaline, apex pointed, base truncate, hila
thickened and darkened.
Holotype: India: Himachal Pradesh: Shimla (Simal), on
Oplismenus sp., 7 May 1960, G. Lall (HCIO 27097).
Host range and distribution: On Oplismenus sp., Poaceae
(Panicoideae, Paniceae), Asia (India, Himachal Pradesh).
Notes: A true Cercospora distinct from common C. apii s.
lat. by uniformly short conidiophores and narrowly acicular
conidia. Type material was not available for re-examination.
50
Fig. 25. penniseti (CUP 40492, holotype). A. Conidiophore fascicle.
B. Conidiophores. c. Conidia. Bar = 10 µm.
cercospora penniseti Chupp,
Cercospora: 250 (1954).
(Fig. 25)
Monograph
of
Literature: Crous & Braun (2003: 313), Kamal (2010: 72).
Illustration: Chupp (1954: 250, ig. 117).
Description: Leaf spots at irst small, more or less elliptical,
medium to dark brown, later larger and sometimes conluent,
inally often large leaf segments or almost entire leaves
discoloured. Caespituli mainly hypophyllous, ine, dark brown.
Mycelium internal. Stromata lacking or small, subglobose,
dark brown to blackish brown. Conidiophores in small to
moderately large fascicles, 2–20, arising from internal
hyphae or stromata, through stomata or erumpent, erect,
IMA FUNGUS
cercosporoid fungi 3
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straight, subcylindrical, usually not or only once geniculate,
unbranched, 50–250 × 4–6 µm, pluriseptate, distance
between septa 10–25 µm, uniformly medium brown to dark
olivaceous-brown, wall thin to slightly thickened, smooth;
conidiogenous cells integrated, terminal, 10–30 µm long,
conidiogenous loci thickened and darkened, with a single
terminal locus or two, rarely several loci, 2.5–4 µm diam.
Conidia solitary, acicular, straight to curved, 30–240 × 2–5
µm, pluriseptate, hyaline, thin-walled, smooth, apex acute or
subacute, base truncate, 2–4 µm wide, hila thickened and
darkened.
Holotype: usA: Georgia: Tift County, Tifton, on Cenchrus
spicatus [Pennisetum glaucum], 19 Aug. 1943, C. L. Lefebvre
(CUP 40492). Isotype: K(M) IMI 103707 (slide). Topotypes:
BPI 439320, CUP 40493.
Host range and distribution: On Cenchrus (distachyus
[Pennisetum distachyum], spicatus [Pennisetum glaucum,
typhoides, Setaria glauca], orientalis [Pennisetum orientale],
purpureus [Pennisetum purpureum]), Poaceae (Panicoideae,
Paniceae), Africa (Malawi), Asia (India, Tamil Nadu, Uttar
Pradesh; Japan), Central America (Costa Rica), and North
America (USA, Georgia).
Notes: This species is part of the morphological Cercospora
apii s. lat. compex. Material on Cenchrus distachyus from
Costa Rica refers to an unpublished collection deposited as
BPI 439319. Cercospora typhoides, described from India on
Pennisetum glaucum, differs in having shorter, 0–1-septate
conidia and mainly obclavate-cylindrical conidia. The identity
of Indian records of C. penniseti is unclear. It is possible that
all of them belong to C. typhoides. Indian material was not
available for examination. The record of C. penniseti from
Japan on Pennisetum glaucum (Katsuki 1966) is uncertain
as material could not be traced. Katsuki’s (1966) description
agrees with this species, except for much larger stromata,
48–60 × 30–48 µm.
cercospora rottboelliae J. Kranz, Sydowia 19: 80
“1965” (1966).
(Fig. 26)
Synonyms: Passalora rottboelliae (J. Kranz) U. Braun &
Crous, in Crous & Braun, Mycosphaerella and Anam.:
358 (2003).
Cercospora rottboelliae J.M. Yen & Gilles, in Yen, Cah. Maboké
9: 114 “1971” (1973), nom. illeg. (Art. 53.1) [holotype:
gabon: Libreville, on Rottboellia cochinchinensis, 2 May
1971, G. Gilles 120 (PC); isotype: K(M) IMI 183413].
Cercospora rottboelliicola J.M. Yen, Bull. Soc. Mycol. France
91: 103 (1975), as nom. nov. for C. rottboelliae J.M. Yen
& Gilles 1973, non Kranz 1966.
Illustrations: Kranz (1966: 81, ig. 7), Yen (1973: 113, ig. 6).
Description: Leaf spots amphigenous, elliptical-fusiform to
irregular, 1–6 mm diam, at irst pale brown, later darker brown,
centre inally grey to greyish white, margin indeinite. Caespituli
amphigenous, but more abundant on the lower surface, indistinct
or ine, dark to blackish brown. Mycelium internal. Stromata
VOLUME 6 · NO. 1
Fig. 26. Cercospora rottboeliae (K(M) IMI 102274, holotype). A.
Conidiophore fascicle. B. Conidiophore. c. Conidiophore tips. d.
Conidia. Bar = 10 µm.
lacking or almost so, sometimes with small substomatal
aggregations of a few swollen hyphal cells, brown. Conidiophores
in loose fascicles, 2–8, arising from internal hyphae or swollen
hyphal cells, through stomata, erect to decumbent, straight
to distinctly geniculate-sinuous, unbranched, 25–215 × 3–5
µm, 0–7-septate, pale to medium brown, sometimes paler
towards the tip, thin-walled, smooth; conidiogenous cells
integrated, terminal, occasionally intercalary, about 10–30 µm
long, sometimes distinctly subdenticulate, conidiogenous loci
thickened and darkened, 2–2.5 µm diam. Conidia catenate, in
simple chains, cylindrical, subcylindrical to obclavate-cylindrical,
straight to somewhat curved, 15–55 × 2–4.5 µm, 1–7-septate,
hyaline, thin-walled, smooth, ends obtuse, rounded, truncate
to short obconically truncate, 1.5–2.5 µm wide, hila somewhat
thickened and darkened.
51
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Braun et al.
Holotype: guinea: Kindia, on Rottboellia cochinchinensis,
Aug. 1963, J. Kranz (K(M) IMI 102274).
Host range and distribution: On Rottboellia cochinchinensis
[exaltata], Poaceae (Panicoideae, Andropogoneae), Africa
(Gabon, Guinea, Ivory Coast).
Notes: The conidiogenous loci are thickened and darkened,
and, due to the catenate conidia, this species was
considered phaeoramularia-like and assigned to Passalora
s. lat. (Crous & Braun 2003). Results of molecular sequence
analyses has shown that passalora-like species with
colourless conidia belong to Cercospora (Groenewald et al.
2013), which also applies to species with catenate conidia
as recently demonstated for C. eremochloae (see Shivas
& Young, in Crous et al. 2011). Cercospora rottboelliae is
at present better maintained as species of Cercospora,
although sequence data is not yet available for this fungus.
Cercospora rottboelliae is morphologically close to C.
fusimaculans, which has also been recorded on Rottboellia
spp., but differs in having longer conidiophores and shorter,
broad conidia.
Yen (1975) introduced the new name Cercospora
rottboelliicola as replacement for C. rottboelliae J.M. Yen &
Gilles, non Kranz, and cited material from the Ivory Coast.
This material was preserved and has been re-examined
(Ivory Coast, Abidjan, on Rottboellia cochinchinensis, 16
Feb. 1974, G. Gilles 101, PC).
cercospora rottboelliigena Y.L. Guo & Y. Jiang,
Mycosystema 19: 447 (2000); as “rottboelligina”.
(Fig. 27)
Fig. 27. Cercospora rottboeliicola (HMAS 78800, holotype). A.
Conidiophore fascicles. B. Conidiophore tips. c. Conidia. Bar = 10
µm.
Literature: Guo & Jiang (2000: 447), Crous & Braun (2003:
358), Guo et al. (2005: 124), Braun & Urtiaga (2013: 593).
Illustrations: Guo & Jiang (2000b: 447, ig. 2), Guo et al.
(2005: 125, ig. 85).
Description: Leaf spots amphigenous, circular to oblong or
irregularly shaped, 2–10 × 1–4 mm or conluent and larger,
centre brown, on the upper side with darker brown border,
paler below. Caespituli amphigenous, ine, brown. Mycelium
internal. Stromata lacking or only formed as small stromatic
aggregations of a few swollen hyphal cell, 10–20 µm diam,
substomatal to intraepidermal, brown, cells to 6 µm diam.
Conidiophores in small to moderately large fascicles, 2–15,
arising from internal hyphae or stromatic hyphal aggregations,
through stomata or erumpent, erect, straight, subcylindrical
to strongly geniculate-sinuous, unbranched, (10–)25–160 ×
3.5–6.5 µm, (0–)1–5-septate, medium brown, paler towards
the tip, wall thin to slightly thickened, smooth; conidiogenous
cells integrated, terminal and intercalary, 10–50 µm long,
conidiogenous loci solitary to several, conspicuous, thickened
and darkened, 1.5–2 µm diam. Conidia solitary, acicular or
almost so to narrowly obclavate-subcylindrical, straight to
somewhat curved, 40–235 × 2.5–4.5 µm, 3- to pluriseptate,
hyaline, thin-walled, smooth, apex obtuse to subacute, base
subtruncate to obconically truncate, 1.5–2 µm wide, hila
thickened and darkened.
52
Holotype: china: Guangxi Province: Ningming, on Rottboellia
cochinchinensis, Oct. 1958, Z. C. Liang 1056 (HMAS 78800).
Host range and distribution: On Rottboellia cochinchinensis
[exaltata], Poaceae (Panicoideae, Andropogoneae), Asia
(China, Guangxi), South America (Venezuela).
Note: A true Cercospora s. str. distinct from C. apii s. lat.
by having acicular to obclavate-cylindrical conidia with
obconically truncate, narrower bases.
cercospora secalis Chupp, Monograph of Cercospora: 252 (1954).
(Fig. 28)
Literature: Chupp (1954: 252), Crous & Braun (2003: 371).
Description: Leaf spots oblong, narrow lines, 0.5 mm
wide or larger lesions to 35 × 3 mm, pale to dark brown,
sometimes with yellowish halo. Caespituli amphigenous,
punctiform, blackish, scattered, mostly arranged in lines.
Mycelium internal. Stromata small to medium in size,
mainly substomatal, 10–50 µm diam, sometimes oblong,
to 80 µm, brown, composed of swollen hyphal cells,
circular to somewhat angular-irregular in outline, 2–7 µm
IMA FUNGUS
cercosporoid fungi 3
ART I CLE
Fig. 29. Cercospora seriata (CUP-A 2009#1, lectotype). A.
Conidiophore fascicle. B. Conidiophores. c. Conidia. Bar = 10 µm.
Holotype: usA: Virginia: Fredericksburg Sherwood Forest
Farm, on Secale cereale, 9 Jun. 1947, C. L. Lefebvre & A. G.
Johnson (CUP 41183). Isotype: BPI 441070.
Host range and distribution: On Secale cereale, Poaceae
(Pooideae, Triticeae), North America (USA, Illinois, Virginia).
Fig. 28. Cercospora secalis (CUP 41181, holotype). A. Conidiophore
fascicle. B. Conidiophore. c. Conidia. Bar = 10 µm.
diam, brown, wall somewhat thickened. Conidiophores
in small to moderately large fascicles, 2–25, divergent to
moderately dense, arising from stromata, through stomata,
divergent, erect, straight, subcylindrical to somewhat
geniculate, unbranched, (15–)20–70(–100) × 3–5 µm,
sparingly septate, uniformly pale to medium brown, thinwalled, smooth; conidiogenous cells integrated, terminal
or conidiophores recuded to conidiogenous cells, 10–40
µm long, conidiogenous loci conspicuous, thickened and
darkened, (1.5–)2–3 µm wide. Conidia solitary, acicular,
shorter conidia sometimes subcylindrical, straight to
somewhat curved, 20–105 × 3–4.5 µm, pluriseptate,
hyaline, thin-walled, smooth, apex subobtuse, base truncate
to somewhat obconically truncate, (1.5–)2–2.5(–3) µm wide,
hila thickened and darkened.
VOLUME 6 · NO. 1
Notes: This species belongs to the Cercospora apii s. lat.
complex (Crous & Braun 2001: 330). The identity of records
from Malawi and Papua New Guinea on Triticum spp., and
reports on Avena sativa, Hordeum vulgare, and Triticum spp.
from North America (USA, Illinois) (Crous & Braun 2003) are
doubtful.
cercospora seriata G.F. Atk., J. Elisha Mitchell Sci.
Soc. 8: 59 (1892).
(Fig. 29)
Literature: Saccardo (1892: 657), Chupp (1954: 252), Crous
& Braun (2003: 374).
Leaf spots amphigenous, irregularly oblong, 1–3 × 0.5–1 mm,
yellowish brown to dingy grey, margin usually darker, brown,
sometimes with yellowish halo, inally sometimes entire
leaves turning brown, necrotic. Caespituli amphigenous,
punctiform, in lines, dark brown to blackish. Mycelium
internal. Stromata small, 10–30 µm diam, substomatal to
immersed, brown. Conidiophores in small to moderately large
fascicles, loose to moderately dense, arising from stromata,
through stomata or erumpent, erect, straight, subcylindrical
or attenuated towards the tip to moderately geniculate-
53
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Braun et al.
sinuous, unbranched or rarely once branched, 10–50 × 2–5
µm, 0–3-septate, pale to medium brown throughout or mostly
paler towards the tip, occasionally subhyaline at the very tip,
thin-walled, smooth; conidiogenous cells integrated, terminal
or conidiophores reduced to conidiogenous cells, 10–30 µm
long, conidiogenous loci thickened and darkened, 1.5–2 µm
diam. Conidia obclavate-cylindrical, straight to somewhat
curved, 20–65(–70) × 2–4 µm, 2–6-septate, hyaline, thinwalled, smooth, apex obtuse, base short obconically truncate,
1.5–2.5 µm wide, hila thickened and darkened.
Lectotype (designated here, MycoBank, MBT200452):
usA: Alabama: Auburn, on Sporobolus asper, 7 Aug. 1891,
Duggar & Newman 2009 (CUP-A 2009#1). Former syntype:
CUP-A 2009#2 (from 24 Jul. 1891). Topotypes: CUP 41196,
OSC 9905.
Host range and distribution: On Sporobolus (clandestinus
[compositus var. clandestinus], compositus [asper],
cryptandrus), Poaceae (Chloridoideae, Eragrostideae), North
America (USA, Alabama, Oklahoma, Wisconsin).
Note: A true Cercospora s. str. distinct from C. apii s. lat. by
having obclavate-cylindrical conidia.
cercospora setariae G.F. Atk., J. Elisha Mitchell Sci.
Soc. 8: 50 (1892).
(Fig. 30)
Synonyms: Cercosporina setariae (G.F. Atk.) Hori, J. Pl. Prot.
(Tokyo) 4: 1 (1917).
Cercospora setariicola Tehon & E.Y. Daniels, Mycologia 19:
128 (1927) [holotype: usA: Illinois: McDonough County,
Macomb, on Paspalum glauca, 16 Aug. 1924, P. A. Young
11542 (ILLS 11542); isotypes: CUP 41211, NY 945705;
paratype: ILLS 7905].
Cercospora paspali W.W. Ray, Mycologia 36: 173 (1944)
[holotype: usA: Oklaoma: Perkins, on Paspalum
stramineum, 26 Aug. 1942 (CUP 33134); isotypes: BPI
439257, CUP 40469, MICH 15347, NY 937110].
Literature: Saccardo (1892: 655; 1972: 1384), Vassiljevsky
& Karakulin (1937: 271–272), Chupp (1954: 253), Katsuki
(1965: 34), McKenzie & Latch (1984: 115), Hsieh & Goh
(1990: 136–137), Crous & Braun (2003: 375–376), Guo et al.
(2005: 125–127), Kamal (2010: 85).
Illustrations: Chupp (1954: 250, ig. 118), McKenzie & Latch
(1984: 114, ig. 1C), Hsieh & Goh (1990: 138, ig. 105), Guo
et al. (2005: 126, ig. 86).
Description: Leaf spots amphigenous, oval to elliptical, 1–12
× 0.5–2 mm, dark reddish brown or brown, later with grey
centre, often conluent. Caespituli amphigenous, but mainly
hypophyllous. Mycelium internal. Stromata small, substomatal,
brown. Conidiophores in divergent fascicles, 2–15, arising
from stromata, through stomata, erect, straight, subcylindrical
to somewhat geniculate-sinuous, unbranched, 8–45 × 2–5 µm,
rarely longer, continuous to septate, yellowish olivaceous to
medium olivaceous-brown, often paler towards the tip, thinwalled, smooth; conidiogenous cells integrated, terminal or
54
Fig. 30. Cercospora setariae (CUP-A 2120, lectotype). A.
Conidiophore fascicle. B. Conidiophores. c. Conidia. Bar = 10 µm.
conidiophores reduced to conidiogenous cells, about 5–25 µm
long, conidiogenous loci conspicuous, somewhat thickened and
darkened, 1–2 µm diam. Conidia solitary, narrowly obclavatecylindrical, longer ones sometimes almost subacicular, straight
to somewhat curved or slightly sigmoid, (20–)30–100(–150) ×
(1.5–)2–5(–6) µm, 1–13-septate, hyaline, thin-walled, smooth,
apex subacute or subobtuse, base subtruncate to usually
short to long obconically truncate, 1–2 µm wide, somewhat
thickened and darkened.
Lectotype (designated here, MycoBank, MBT200453): usA:
Alabama: Auburn, on Paspalum glaucum [Setaria glauca], 17
Sep. 1891, B. M. Duggar (CUP-A 2120). Isolectotype: CUP
41208.
IMA FUNGUS
cercosporoid fungi 3
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Host range and distribution: On Paspalum (conjugatum,
dilatatum,
glaucum
[lutescens,
Setaria
glauca],
scrobiculatum, stramineum), Setaria (italica, palmifolia,
parvilora [geniculata], poiretiana, pumila, sphacelata, viridis),
Poaceae (Panicoideae, Paniceae), Africa (Guinea, Mauritius,
Uganda), Asia (China, India, Japan, Korea, Russia, Taiwan),
Caucasus (Georgia), Central and South America (Argentina,
Brazil, Guatemala, Panama), Europe (Poland, Romania,
Russia, Ukraine), New Zealand, North America (USA,
Alabama, Florida, Iowa, Illinois, Kansas, Kentucky, Maryland,
Michigan, Minnesota, New Hampshire, North Dakota,
New York, Oklahoma, Pennsylvania, Texas, Virginia, West
Virginia, Wisconsin).
Notes: A true Cercospora s. str. distinct from C. apii s. lat. in
having obclavate-cylindrical conidia. Records of C. setariae
on Sporobolus cryptandrus from North America refer to C.
seriata.
cercospora sorghi Ellis & Everh., J. Mycol. 3: 15
(1887).
(Fig. 31)
var. sorghi
Synonym: ? Cercospora andropogonis Sawada, nom. inval.
(Art. 38.1), according to Sawada (1959: 226) and Hsieh
& Goh (1990: 137), see notes under Passalora fujikuroi.
Literature: Saccardo (1892: 656), Vassiljevsky & Karakulin
(1937: 270), Chupp (1954: 253), Sun (1955: 138), Vasudeva
(1963: 187), Katsuki (1965: 35), Mulder & Holliday (1974b),
Ellis (1976: 260), Hsieh & Goh (1990: 137), Crous & Braun
(1996: 308; 2003: 382), Okori et al. (2004), Guo et al. (2005:
127), Kamal (2010: 88), CMI Distribution Map No. 338.
Illustrations: Chupp (1954: 250, ig. 119), Sun (1955: 138, ig.
1), Mulder & Holliday (1974b: ig., unnumbered), Ellis (1976:
260, ig. 197B), Guo et al. (2005: 128, ig. 87).
Description: Leaf spots amphigenous, mostly oblong, 2–16
× 0.5–5 mm, at irst mostly dark purple to reddish, later with
tan to brown centre, occasionally deinite leaf spots lacking.
Caespituli amphigenous. Mycelium internal. Stromata absent
or almost so to well-developed, 10–50 µm diam, subglobose,
brown to dark brown. Conidiophores fasciculate, 3–20,
arising from internal hyphae or stromata, erect, straight,
subcylindrical to geniculate-sinuous in the upper half, width
sometimes somewhat irregular, unbranched, (10–)20–150(–
220) × (2.5–)3–6.5(–7) µm, pluriseptate, medium dark brown
or olivaceous-brown, somewhat paler towards the tip, thinwalled, smooth; conidiogenous cells integrated, terminal,
10–65 µm long, conidiogenous loci conspicuous, thickened
and darkened, 1.5–3 µm diam. Conidia solitary, acicular to
obclavate or obclavate-cylindrical, straight to somewhat
curved, (15–)25–120(–320) × (1.5–)2.5–5.5(–7) µm,
1–20-septate, hyaline, thin-walled, smooth, apex subacute,
base truncate to obconically truncate, 1.5–3 µm wide, hila
thickened and darkened.
Lectotype (designated here, MycoBank, MBT200454):
usA: Louisiana: Rapides Parish [Plaquemides County], on
VOLUME 6 · NO. 1
Fig. 31. Cercospora sorghi (NY 838621, lectotype). A. Conidiophore
fascicle. B. Conidiophore. d. Conidia. Bar = 10 µm.
Sorghum halepense, 1 Aug. 1886, A. B. Langlois 543 (NY
838621). Isolectotype: BPI 441532. Topotypes: NY 838618,
838619.
Host range and distribution: On Sorghum (×almum,
arundinaceum [verticillilorum], bicolor [dochna, roxburghii,
vulgare], drummondii [×sudanense], halepense, propinquum),
Poaceae (Panicoideae, Andropogoneae), widely distributed
in the tropics and subtropics, Africa (Benin, Burkina Faso,
Burundi, Cameroon, Central African Republ., Chad, Congo,
Ethiopia, Gabon, Gambia, Ghana, Ivory Coast, Kenya, Malawi,
Mauritius, Niger, Nigeria, Rwanda, Senegal, Sierra Leone,
Somalia, South Africa, Sudan, Tanzania, Togo, Uganda,
Zambia, Zimbabwe), Asia (Bangladesh, Bhutan, Brunei,
55
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Braun et al.
Cambodia, China, India, Indonesia, Japan, Korea, Laos,
Malaysia, Myanmar, Nepal, Pakistan, Papua New Guinea,
Philippines, Taiwan, Thailand, Yemen), Australia, Caucasus
(Armenia, Azerbaijan, Georgia), Central and South America
(Argentina, Brazil, Colombia, El Salvador, Guatemala, Guyana,
Honduras, Panama, Peru, Suriname, Venezuela), Europe
(Italy, Russia), North America (Mexico; USA, Alabama, Florida,
Georgia, North Dakota, Nebraska, Iowa, Indiana, Kansas,
Louisiana, Missouri, Mississippi, Nebraska, North Carolina,
Oklahoma, South Carolina, South Dakota, Tennessee, Texas,
Virginia, West Virginia), Oceania (American Samoa, Cook
Islands, Fiji, New Caledonia, Niue, Samoa, Solomon Islands,
Tonga, Vanuatu), and West Indies (Cuba, Dominican Republ.,
Jamaica, Puerto Rico, Trinidad and Tobago, Virgin Islands).
Notes: A true Cercospora s. str. distinct from C. apii s. lat. in
the at least partly obclavate or obclavate-cylindrical conidia.
Records of collections on other hosts than Sorghum spp.
[Bothriochloa pertusa [Amphilophis pertusa, Andropogon
pertusus], Cymbopogon (nardus [afronardus, validus], caesius
[excavatus], citratus, schoenanthus), Echinochloa (crusgalli, hispidula, pyramidalis), Holcus lanatus, Hyparrhenia
rufa, Panicum miliaceum, Cenchrus (spicatus [Pennisetum
glaucum, typhoides], purpureus [P. purpureum]), Setaria
geniculata, Sporobolus sp., Thelepogon sp., Zea mays (see
Crous & Braun 2003)] are doubtful and unproven and seem to
belong to other species. Records on Echinochloa spp. belong
undoubtedly to C. echinochloae, collections on Cenchrus
(including Pennisetum) spp. to C. penniseti and those on
Setaria geniculata and Sporobolus sp. to C. setariae. Records
on Zea mays refer to Cercospora sorghi var. maydis, which is
not conspeciic with C. sorghi (see C. apii). Available molecular
sequence analyses support C. sorghi as a species of its own
(Goodwin et al. 2001, Crous et al. 2006). Okori et al. (2004)
examined populations of C. sorghi in Africa by molecular
methods and showed that collections from wild and cultivated
Sorghum spp. are indistinguishable based on AFLP and ITS
data. They postulated that Cercospora on wild Sorghum spp.
might be sources of inoculum to cultivated species. Cercospora
sorghicola is a cryptic species described from Iran, which is
morphologically barely distinguishable from C. sorghi, but
genetically distinct (discussion see C. sorghicola).
Ellis & Everhart (1887) introduced Cercospora sorghi
var. maydis. Chupp (1954) emphasized that this Cercospora
from Zea mays does not infect Sorghum spp. and possibly
represents a separate species, which is supported by results
of molecular sequence analyses (Goodwin et al. 2001,
Crous et al. 2006). Sequences of Cercospora sorghi var.
maydis from Africa and North America cluster with C. apii
and C. beticola (Crous et al. 2006), i.e. C. apii s. lat. can be
transmitted to maize.
var. ciccaronei (N. Pons) U. Braun, Schlechtendalia
5: 48 (2000).
Basionym: Phaeoramularia ciccaronei N. Pons, Fitopatol.
Venez. 6: 2 (1993).
Literature: Crous & Braun (2003: 382).
Description: Conidia solitary and in short chains, otherwise
agreeing with var. sorghi.
Holotype: Venezuela: Borburata, Edo. Carabobo, on
Sorghum arundinaceum, 18 Feb. 1992, C. Rincones (VIA
5696). Isotype: K(M) IMI 364371.
Host range and distribution: On Sorghum arundinaceum
[verticillilorum], Poaceae (Panicoideae, Andropogoneae),
South America (Venezuela).
cercospora sorghicola M. Bakhshi, Babai-ahari,
Crous & U. Braun, Persoonia 34: 81 (2015).
(see Fig. 31)
Illustration: Bakhshi et al. (2015: 82, ig. 9).
Description: Leaf spots amphigenous, at irst forming dark
purple spots that gradually enlarge into linear-oblong lesions
with dark purple centre and dark red-purple border, 5–35 mm
long. Caespituli amphigenous, brown. Mycelium internal.
Stromata well-developed, substomatal or intraepidermal, to
50 µm diam, brown. Conidiophores in loose to dense fascicles,
5–40, arising from stromata, through stomata or erumpent,
erect, straight, subcylindrical to lexuous-geniculate as
result of sympodial proliferation, almost uniform in width,
unbranched, (45–)70–80(–100) × 4–5.5 µm, 1–8-septate,
pale brown to brown, paler towards the tip, thin-walled,
smooth; conidiogenous cells integrated, terminal, 20–40 µm
long, with a single to several conidiogenous loci, thickened
and darkened, terminal and lateral, protuberant, 2–4 µm diam.
Conidia solitary, acicular, obclavate to obclavate-cylindrical,
straight to curved, (20–)80–100(–150) × 3–4(–5) µm, (3–)8–
13(–17) µm, hyaline, thin-walled, smooth, apex subacute to
subobtuse, base truncate to obconically truncate, 1.5–2.5 µm
wide, hila thickened and darkened.
Holotype: Iran: Guilan Province: Kiashahr, on Sorghum
halapense, Aug. 2012, M. Bahkshi (IRAN 16457 F). Ex-type
culture: CCTU 1173 = CBS 136448.
Host range and distribution: On Sorghum halapense,
Poaceae, Asia (Iran).
Notes:
C.
sorghicola
is
morphologically
almost
indistinguishable from C. sorghi, yet genetically distinct,
at least from a North American C. sorghi ITS sequence
considered to be correctly identiied. Cercospora sorghi as
currently circumscribed is widespread in cultivated sorghum.
The actual distribution of C. sorghicola is unknown, since
molecular sequence data from different parts of the world are
lacking, although a wider distribution in Asia is likely.
cercospora sp.
(Fig. 32)
Material examined: usA: Texas: College Station, on Bromus
inermis, Poaceae (Pooideae, Bromeae), 1953, M. Whitehead
(BPI 436346).
Illustration: Pons (1993: 5, ig. 2).
56
IMA FUNGUS
cercosporoid fungi 3
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strongly so, unbranched, 10–40 × 3–6 µm, 0–1(–2)-septate,
pale to medium olivaceous, brownish to yellowish brown,
darker in mass, conidiogenous cells integrated, terminal or
conidiophores reduced to conidiogenous cells, usually with
several conidiogenous loci, often aggregated near the apex,
1–2 µm diam, thickened and darkened; conidia solitary,
narrowly obclavate to acicular, shorter conidia sometimes
fusiform, 20–100 × 1–3.5 µm, shorter conidia usually
1–4-septate, longer ones pluriseptate, hyaline, thin-walled,
smooth, apex acute to subobtuse, base truncate to short
obconically truncate, 1–2 µm wide, thickened and darkened.
This fungus is well characterized, but as it is currently only
known from a single collection we refrain from introducing a
formal description of this fungus as new species here.
cercospora tessellata G.F. Atk., J. Elisha Mitchell Sci.
Soc. 8: 59 (1892).
(Fig. 33)
Literature: Saccardo (1892: 656), Vassiljevsky & Karakulin
(1937: 273), Chupp (1954: 255), Crous & Braun (2003: 400).
Fig. 32. Cercospora sp. on Bromus (BPI 436346). A. Conidiophore
fascicles. B. Conidiophores. c. Conidia. Bar = 10 µm.
Notes: Cercospora collections on Bromus were previously
assigned to C. festucae, which is incorrect. A sample of “C.
festucae” on Bromus inermis from Texas was examined and
found to be distinct from the latter species as well as from
another collection on this host morphologically assigned
to C. apii s. lat. The irst collection differed from C. apii
and C. festucae in having shorter, strongly geniculate
conidiophores and narrower conidia, mostly obclavate with
obconically truncate base, 1–2.5 µm wide: Lesions variable,
ranging from small brown spots to often oblong to large
discoloured patches, brownish to dingy grey, inally large
leaf segments or almost entire leaves discoloured, necrotic;
caespituli amphigenous, punctiform, scattered, dark brown;
mycelium internal; stromata small or oblong, to 40 × 10 µm,
substomatal, brown; conidiophores in small to moderately
large fascicles, loose to dense, arising from stromata, through
stomata, erect, straight to usually distinctly geniculate, often
VOLUME 6 · NO. 1
Description: Leaf spots oblong, 3–5 × 0.5–1 mm, dark
brown to blackish, sometimes with bluish tinge. Caespituli
hypophyllous, pustulate, arrangement linear. Mycelium
internal. Stromata well-developed, substomatal, 20–50
µm diam, dark olivaceous-brown, composed of swollen
hyphal cells, circular to slightly angular-irregular in outline.
Conidiophores in small to moderately large fascicles, mostly
dense, arising from stromata, through stomata, erect, straight,
subcylindrical to conical, non-geniculate, unbranched,
short, 5–20 × 2–4 µm, aseptate, rarely with a single basal
septum, pale to dark olivaceous-brown, thin-walled, smooth;
conidiophores usually reduced to conidiogenous cells, with
1–2(–3) conspicuous conidiogenous loci, 1–1.5 µm diam,
thickened and darkened. Conidia solitary, iliform-acicular,
straight to curved, 30–90 × 1.5–2.5 µm, indistinctly 3- to
pluriseptate, hyaline, thin-walled, smooth, apex subacute,
base truncate to somewhat obconically truncate, 1–1.5 µm
wide, hila slightly thickened and darkened.
Lectotype (designated here, MycoBank MBT200455): usA:
Alabama: Auburn, on Dactyloctenium aegyptium, 6 Nov.
1891, G. F. Atkinson (CUP-A 2306). Isolectotype: CUP 41393.
Host range and distribution: On Dactyloctenium aegyptium
[Eleusine aegyptia], Eleusine (coracana, jaegeri), Poaceae
(Chloridoideae, Eragrostideae), Africa (Ethiopia, Kenya,
Nigeria), North America (USA, Alabama).
Note: A true species of Cercospora s. str. distinct from C. apii
s. lat. by having short conidiophores and narrow conidia.
cercospora typhoides O.P. Sharma & A.C. Jain,
J.N.K.V.V. Res. J. 1: 83 (1967).
Literature: Crous & Braun (2003: 414), Kamal (2010: 96).
Description: Leaf spots amphigenous, at irst visible as
minute dark brown spots, later circular or elliptical, 1.5–4.5 ×
57
Braun et al.
ART I CLE
Holotype: India: Madhya Pradesh: Gwalior, on Cenchrus
spicatus [Pennisetum glaucum, P. typhoides], Poaceae
(Panicoideae, Paniceae), 10 Oct. 1961, O. P. Sharma (Hb.
of Plant Pathology Department, Jawaharlal Nehru Krishni
Vishwa Vidyalaya, Jabalpur, India).
Host range and distribution: Only known from the type
collection.
Notes: According to the original description, Cercospora
typhoides differs from C. penniseti in having shorter,
0–1-septate conidiophores and usually cylindrical-obclavate
conidia. Type material was not available and it is unclear if
this material is preserved at all. Due to its acicular conidia,
Cercospora penniseti is part of the C. apii complex. This
species was also recorded from India, but material was not
available for examination. It remains unclear if these Indian
records indeed belong to C. penniseti or C. typhoides. The
citation of HCIO 26616 as type material (Crous & Braun 2003,
Kamal 2010) is doubtful and unveriied, but that collection is
not the holotype.
cercospora zeae-maydis Tehon & E.Y. Daniels,
Mycologia 17: 248 (1925).
(Fig. 34)
Literature: Chupp (1954: 256), Vassiljevsky & Karakulin
(1937: 270), Saccardo (1972: 1388), Crous & Braun (2003:
433), Kamal (2010: 99), Crous et al. (2006: 194), Groenewald
et al. (2013: 166).
Illustration: Crous et al. (2006: 194, ig. 4).
Fig. 33. Cercospora tesselata (CUP-A 2306, lectotype). A.
Conidiophore fascicles. B. Conidiophores. c. Conidia. Bar = 10 µm.
0.5–2 mm, longer axis parallel to veins, centre grey, margin
purplish to brown. Caespituli amphigenous, punctiform, in
parallel rows. Mycelium internal. Stromata lacking or small,
substomatal, to 35 µm diam, dark brown. Conidiophores in
fascicles, arising from internal hyphae or stromata, through
stomata, erect, straight to somewhat geniculate, unbranched,
17–67 × 3–5 µm, 0–1-septate, uniformly brown, tips conically
truncate; conidiogenous cells integrated, conidiogenous
loci thickened and darkened. Conidia solitary, cylindricalobclavate, narrowly obclavate, longer conidia almost
acicular, straight to slightly curved, 25.6–140 × 2.5–5 µm,
0–14-septate, hyaline, thin-walled, smooth, hila thickened
and darkened.
58
Description: Leaf spots oblong, forming linear lesions
parallel to the midrib, sometimes irregularly shaped, size
variable, brownish to greyish, often with a brown marginal
line or narrow border. Caespituli amphigenous, mostly
hypophyllous, punctiform to subeffuse, brown. Mycelium
internal. Stromata lacking or small, with a few substomatal
swollen hyphal cells, brown. Conidiophores in small
to moderately large fascicles, 3–14, divergent, arising
from internal hyphae or stromatic hyphal aggregations,
emerging through stomata, erect, straight, subcylindrical,
occasionally subclavate to lexuous, distinctly geniculatesinuous, unbranched, 40–180 × 4–8 µm, (0–)1–8-septate,
uniformly pale olivaceous to medium brown, thin-walled,
smooth; conidiogenous cells integrated, terminal,
occasionally intercalary, 10–40 µm long, often subclavate
when terminal, conidiogenous loci conspicuous, thickened
and darkened, 2–3 µm wide. Conidia solitary, broadly
obclavate-subcylindrical, 30–100 × 4–9 µm, 1–10-septate,
hyaline, thin-walled, smooth, apex obtuse, base obconically
truncate, 2–3 µm wide, hila thickened and darkened.
In vitro: Colonies on PDA reaching 15–25 mm diam
after 3 wk, forming ample spermatia. Colonies on MEA
erumpent, with sparse aerial mycelium, margin smooth, but
irregular, surface olivaceous-grey with irregular patches of
white to smoke-grey, reverse iron-grey, colonies fertile.
On OA colonies spreading with moderate aerial mycelium,
margin smooth but irregular, surface red with patches
IMA FUNGUS
cercosporoid fungi 3
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Fig. 34. Cercospora zeae-maydis (BPI 442569, isotype). A.
Conidiophore fascicle. B. Conidiophores. c. Conidia. Bar = 10 µm.
of white and pale olivaceous-grey, fertile. Formation of
cercosporin (red pigment) observed (Goodwin et al. 2001,
Crous et al. 2006).
Holotype: usA: Illinois: Alexander County, McClure, on Zea
mays, 29 Aug. 1924, P. A. Young (ILLS 4276). Isotype: BPI
442569. Epitype (designated by Crous et al. 2006): usA:
Wisconsin: Janesville, on Zea mays, 2002, B. Fleener (CBS
H-17774). Ex-epitype culture: CBS 117757.
Host range and distribution: On Zea mays, Poaceae
(Panicoideae, Andropogoneae), Africa (Cameroon, Congo,
Ethiopia, Kenya, Malawi, Mozambique, Nigeria, South
Africa, Swaziland, Tanzania, Uganda, Zambia, Zimbabwe),
Asia (India, China), Caucasus (Azerbaijan, Georgia),
Central and South America (Brazil, Colombia, Costa Rica,
Ecuador, Guatemala, Panama, Peru, Venezuela), West
Indies (Trinidad and Tobago), and North America (Canada,
Mexico; USA, Alabama, Colorado, Delaware, Iowa, Illinois,
Kansas, Kentucky, Maryland, Minnesota, North Carolina,
Ohio, Pennsylvania, South Dakota, Tennessee, Virginia,
Wisconsin, West Virginia).
Note: A true Cercospora s. str. distinct from C. apii s. lat.
by its broadly obclavate-cylindrical conidia, and molecularly
established as separate species (Goodwin et al. 2001, Crous
et al. 2006, Groenewald et al. 2013).
VOLUME 6 · NO. 1
Fig. 35. Cercospora zeina (CBS H-17775, holotype). A. Conidiophore
fascicle. B. Conidiophore tips. c. Conidia. Bar = 10 µm.
cercospora zeina Crous & U. Braun, Stud. Mycol. 55:
194 (2006).
(Fig. 35)
Literature: Liu & Xu (2013), Berger et al. (2014).
Illustration: Crous et al. (2006: 195, ig. 5).
Description: Leaf spots amphigenous, vein-limited, length
variable, 5–40 mm, width 2–3 mm, later conluent, pale grey
to pale brown, margin indistinct, chlorotic in younger leaf
spots. Caespituli amphigenous, punctiform to subeffuse, on
leaves grey to brown. Mycelium internal; hyphae branched,
septate, 3–4 µm wide, pale brown, thin-walled, smooth.
Stromata lacking or only formed as small substomatal
aggregations of swollen hyphal cells, to 30 µm diam, brown.
Conidiophores in small to moderately large fascicles, loose
59
ART I CLE
Braun et al.
to dense, arsing from internal hyphae or stromata, emerging
through stomata, erect, straight, subcylindrical to lexuous,
distinctly geniculate-sinuous, unbranched or occasionally
branched above, 40–100 × 5–7 µm, 1–5-septate, uniformly
pale olivaceous to medium brown, thin-walled, smooth;
conidiogenous cells integrated, terminal, 40–60 × 5–6 µm,
with several conspicuous conidiogenous loci, thickened
and darkened-refractive, 2–3 µm diam. Conidia solitary,
broadly fusiform, (40–)60–75(–100) × (6–)7–8(–9) µm,
(1–)3–5(–10)-septate, hyaline, thin-walled, apex subobtuse,
base subtruncate to obconically truncate, 2–3 µm wide, hila
thickened and darkened-refractive.
In vitro: Colonies on PDA reaching 10–15 mm after 3
wk, forming spermogonia. On MEA erumpent, with sparse
aerial mycelium, margin smooth, but irregular, surface
olivaceous-grey with irregular patches of white or iron-grey,
reverse iron-grey, colonies fertile. On OA colonies spreading
with moderate whitish aerial mycelium, margin smooth but
irregular, olivaceous-grey, fertile.
Holotype: south Africa: KwaZulu-Natal: Pietermaritzburg,
on Zea mays, 2005, P. Caldwell (CBS H-17775). Ex-type
culture: CBS 118820.
Host range and distribution: On Zea mays, Poaceae
(Panicoideae, Andropogoneae), Africa (Kenya, Rwanda,
South Africa, Uganda, Zambia, Zimbabwe), Asia (China),
North America (USA, North Carolina, New York, Ohio,
Virginia).
Notes: Besides obvious genetic differences, this species
differs from Cercospora zeae-maydis in having shorter
conidiophores, to about 100 µm in length, broadly fusiform
conidia, and slow-growing cultures without formation of red
pigments (cercosporin), but the differentiation between the
two species just based on morphology is dificult. Cercospora
zeae-maydis is the most common species on maize in
North America, although C. zeina also occurs in the USA.
Unambiguous identiication requires molecular sequence
analyses.
cercospora zizaniae Thirum. & Govindu, Sydowia 7:
49 (1953).
(Fig. 36)
Literature: Vasudeva (1963: 217), Crous & Braun (2003:
434), Kamal (2010: 100).
Illustration: Thirumalachar & Govindu (1953: pl. II, igs 9–10).
Description: Leaf spots linear, short to very long, to about
1 mm wide, to irregular, yellowish to brown, later conluent,
forming long stripes or almost entire leaves becoming
necrotic, margin infeinite. Caespituli mainly epiphyllous,
inely punctiform to effuse, dark brown to blackish. Mycelium
internal. Stromata lacking or only with aggregations of a
few swollen hyphal cells, brown. Conidiophores in small
fascicles, divergent, arising from internal hyphae or stromatic
hyphal aggregations, erect, divergent, straight, geniculate,
unbranched, 20–180 × (2–)3–5(–6) µm, 1–10-septate, light
60
Fig. 36. Cercospora zizaniae (BPI 442670, lectotype). A.
Conidiophore fascicle. B. Conidiophore tips. c. Conidia. Bar = 10
µm.
brown, paler towards the apex, tips sometimes subhyaline,
thin-walled, smooth; conidiogenous cells integrated,
terminal and intercalary, 10–50 µm long, conidiogenous loci
conspicuous, thickened and darkened, 1.5–2.5 µm diam.
Conidia solitary, obclavate-cylindrical to subacicular, straight
to curved, (25–)30–60 × 3–4 µm, 1–6-septate, hyaline, thinwalled, smooth, apex subacute or subobtuse, base truncate,
subtruncate to mostly obconically truncate, 1.5–2 µm wide,
hila thickened and darkened.
Lectotype (designated here, MycoBank MBT200456): India:
Bihar: Patna, on Zizania aquatica, Poaceae (Ehrhartoideae,
Oryzeae), 4 Oct. 1952, M. J. Thirumalachar (BPI 442670).
Isolectotypes: CUP 40775, HCIO, K(M) IMI 55519.
Host range and distribution: Only known from the type
collection.
IMA FUNGUS
cercosporoid fungi 3
Doubtful,
species
excluded
and
insuficiently
ART I CLE
Note: A true Cercospora s. str. distinct from C. apii s. lat. in
the relatively short conidia at least partly with an obconically
truncate base.
known
Cercospora acerosa Dickhoff & Arendsen-Hein, Arch.
Java Suikerindustr. 1901: 1009 (1901).
Literature: Saccardo (1906: 611), Chupp (1954: 243), Crous
& Braun (2003: 41–42).
Host range and distribution: On Saccharum (oficinarum,
Saccharum spp.), Poaceae, Africa (Madagascar), Asia
(Indonesia, Philippines).
Note: Not a Cercospora ide Chupp (1954). The status of this
species is unclear. The original description is meagre and
type material is not preserved.
Cercospora bromi R. Sprague, Mycologia 29: 204
(1937).
Synonyms: Ramulispora bromi (R. Sprague) R. Sprague,
Diseases of cereals and grasses in North America: 418
(1950).
Ansatospora bromi (R. Sprague) R. Sprague, Mycologia 38:
61 (1946).
Centrospora
bromi
(R.
Sprague) A.G.
Newhall,
Phytopathology 36: 895 (1946).
Centrospora bromi (R. Sprague) Vienn.-Bourg., Rev. Mycol.,
n.s. 10: 130 “1945” (1946), nom. illeg. (Art. 52.1).
Literature: Chupp (1954: 244), Braun (1995: 201), Braun &
Crous (2003: 89).
Illustration: Braun (1995: 204, Fig. 197).
Holotype: USA: Oregon: Wasco Co., near Tumwater, on
Bromus rigidus, 13 Mar. 1935, R. Sprague (OSC 10405).
Isotypes: BPI 433860, CUP 39251, NY 936944).
Host range and distribution: On Bromus (rigidus, secalinus,
vulgaris, Bromus sp.), Poaceae, Asia (Russia, Asian part),
North America (USA, Illinois, Oregon, Washington).
Cercospora eleusines Henn., in herb.
Material examined: Japan: Kōchi (Tosa): Inomachi, on
Eleusine indica, Poaceae, Yoshinaga (B).
Note: An unidentiied helminthosporioid fungus with broad,
distoseptate conidia possibly identical with Drechslera
nodulosa (Berk. & M.A. Curtis ex Sacc.) Subram. & B.L. Jain.
Cercospora elymi Rostr., Bot. Tidsskr. 22: 276 (1899).
(Fig. 37)
Synonym: Cladosporium elymi (Rostr.) U. Braun, comb.
nov.
MycoBank MB811242
VOLUME 6 · NO. 1
Fig. 37. Cladosporium elymi (C-F-92454, holotype). A. Conidiophore
fascicle. B. Conidiophores. c. Conidia. Bar = 10 µm.
Basionym: Cercospora elymi Rostr., Bot. Tidsskr. 22: 276
(1899).
Literature: Saccardo (1902: 1074), Lindau (1910: 87),
Vassiljevsky & Karakulin (1937: 273), Chupp (1954: 246),
Crous & Braun (2003: 173).
Description: Leaf spots oblong, formed as narrow streaks
between veins, 1–5 × 0.5–1 mm, sometimes conluent,
pale to medium dark brown, margin indeinite. Caespituli
amphigenous, mostly hypophyllous, scattered, punctiform,
dark brown to blackish. Mycelium internal. Stromata globular
to irregular, small aggregations of a few swollen hyphal cells
to large stromata, 10–80 µm diam, substomatal to immersed,
dark brown, composed of swollen hyphal cells, rounded to
somewhat angular-irregular in outline, 2–7 µm diam, walls
somewhat thickened. Conidiophores in small to mostly large,
usually dense fascicles, arising from stromata, erect, straight,
subcylindrical-conical to moderately geniculate in the upper
half, unbranched, 10–30 × 2–5 µm, 0–1(–2)-septate, pale
olivaceous to olivaceous-brown, slightly paler towards the
tip, dark brown in mass, thin-walled, smooth; conidiogenous
cells integrated, terminal or conidiophores reduced to
conidiogenous cells, conidiogenous loci coronate, darkenedrefractive, 1–2 µm diam. Conidia catenate, usually in simple
chains, broadly ellipsoid-ovoid, obovoid, occasionally
limoniform, straight, 5–15 × 3–7 µm, 0–1-septate, pale
olivaceous to olivaceous-brown, thin-walled, smooth or faintly
and irregularly verruculose, apex of primary conidia obtuse,
61
ART I CLE
Braun et al.
broadly rounded or conically truncate in catenate conidia,
base subtruncate to short obconically truncate, 1–2 µm wide,
hila cononate, somewhat darkened-refractive.
Holotype: denmark: Tisvilde, on Leymus arenarius, Poaceae
(Pooideae, Triticeae), 29 Jun. 1898, E. Rostrup (C-F-92454).
Host range and distribution: Only known from the type
collection.
Notes: The original description is confused and based on
heterogeneous elements. The described conidiophores
pertain to a Cladosporium. Abundant pigmented conidia with
hila agreeing with the coronate (cladosporioid) scar type of
the conidiogenous cells are present in the type material.
The described hyaline conidia, which Rostrup interpreted
as conidia of Cercospora elymi, were also found in the
recently re-examined type collection, but they do not belong
to the described conidiophores. The conidiogenesis of these
colourless, mostly 20–30 × 3–4 µm and 3-septate, fusarioid
conidia, is unclear, but they were deinitely not formed by the
described conidiophores. The hila are rounded to truncate,
unthickened and not darkened-refractive. The application of
the name C. elymi is herewith conined to the Cladosporium
element in the type material. This species is biotrophic.
Based on the recently published monograph of Cladosporium
(Bensch et al. 2012), all biotrophic and saprobic species of
this genus have been compared with C. elymi and were
morphologically distinguishable.
Cercospora muhlenbergiae G.F. Atk., Cornell Univ. Sci.
Bull. 3: 46 (1897).
(Fig. 38)
Fig. 38. Cercospora muhlenbergiae (CUP-A 33, lectotype). A.
Conidiophore fascicle. B. Conidiophore tip. c. Conidia. Bar = 10 µm.
Literature: Saccardo (1899: 1106), Chupp (1954: 249), Crous
& Braun (2003: 285).
Description: Leaf spots on leaves and the leaf sheath, forming
small straw-coloured to brown discolorations, later conluent
and extending, inally sometimes entire leaves discoloured,
necrotic. Caespituli amphigenous, punctiform, dark brown.
Mycelium internal. Stromata usually well-developed, 20–60
µm diam, brown, composed of swollen hyphal cells, rounded
to somewhat angular-iregular in outline. Conidiophores
in small to very large, loose to mostly dense fascicles,
occasionally almost coremioid, arising from stromata, erect,
straight, subcylindrical, geniculate-sinuous in the upper half
or sometimes throughout, unbranched, 40–200 × 3–6 µm,
pluriseptate, pale to medium olivaceous-brown throughout
or somewhat paler towards the tip, thin-walled, smooth;
conidiogenous cells integrated, terminal, 10–60 µm long,
subdenticulate to denticulate, denticles subcylindrical to
conical, 1.5–2 µm diam, wall of the conidiogenous loci
neither thickened nor darkened or somewhat refractive, in
front view visible as minute circle (only rim distinct, content
not darkened). Conidia solitary, fusiform to short obclavate,
20–35 × 3–5.5 µm, 1–3-septate, hyaline or subhyaline,
thin-walled, smooth, apex obtuse, base short obconically
truncate, 1.5–2 µm wide, hila unthickened or refractive to
slightly darkened-refractive.
62
Lectotype (designated here, MycoBank, MBT200457): usA:
Alabama: Lee County, Auburn, on Muhlenbergia diffusa,
3 Oct. 1891, G. F. Atkinson (CUP-A 33). Isolectotype: CUP
40345.
Host range and distribution: On Muhlenbergia (glomerata,
mexicana [foliosa], racemosa, schreberi [diffusa], sylvatica,
tenuifolia, Muhlenbergia sp.), Poaceae, North America (USA,
Alabama, Iowa, Kansas, North Dakota, Nebraska, New York,
Wisconsin).
Notes: Chupp (1954) excluded this species from Cercospora
since the conidia are often 1-septate. Crous & Braun (2003)
called the afinity of this species to cercosporoid genera into
question and, due to the denticle-like conidiogenous loci,
supposed a possible relation to the Dactylaria complex.
The phylogenetic afinity of this species is unknown.The
general habit is not dactylarioid but cercosporoid. The
small, 0–3-septate conidia arising from subdenticulate
to denticle-like conidiogenous loci are similar to those of
Denticularia spp. (Deighton 1972, Ellis 1976, Braun et al.
2013). Cercospora muhlenbergii differs in having hyaline
conidia formed singly (cf. pigmented catenate conidia
in Denticularia). Denticle-like conidiogenous loci are
IMA FUNGUS
cercosporoid fungi 3
Cercospora poae Baudyš & Picb., Prace Morav. Přír.
Společn. 1: 304 (1924).
Literature: Chupp (1954: 251), Vassiljevsky & Karakulin
(1937: 272), Saccardo (1972: 1381), Crous & Braun (2003:
329).
Description: Leaf spots amphigenous, often covering the
entire leaf furface, reddish. Caespituli mainly hypophyllous,
brown, minute, numerous. Conidiophores in fascicles, arising
from stromata, erect, curved, geniculate, wider towards the
apex, 66–89 × 6–7 µm, septate, grey-brown, paler towards
the tip. Conidia usually fusiform, about 20–40 × 4–6 µm
[according to Chupp’s (1954) estimation], 1–3-septate,
greenish to pale brownish, base broad, obconically truncate,
apex attenuated to rounded.
Holotype: czech republic: near Prague, Strasnice, on Poa
palustris [fertilis], Poaceae, 13 May 1911, Ed. Baudys.
Host range and distribution: Only known from the type
collection.
Notes: Not a Cercospora ide Chupp (1954). Type material
of this species was not available for re-examination. Based
on the original description, C. poae is probably a synonym
of Passalora graminis, which is known on Poa spp. as hosts,
but the identity has not yet been proven on the basis of type
material.
Cercospora sacchari Breda de Haan, Meded. Proefstat.
Suikerried. W. Java, Kagok-tegal 3: 15 (1892).
Synonyms: Helminthosporium sacchari E.J. Butler, Mem.
Dept. Agric. India, Bot. Ser., 6: 207 (1913).
Bipolaris sacchari (E.J. Butler) Shoemaker, Canad. J. Bot.
37: 884 (1959).
Bipolaris sacchari (Breda de Haan) Subram., Hyphomycetes
– An Account of Indian Species except Cercosporae: 769
(1971).
Literature: Saccardo (1895: 629), Chupp (1954: 251), Crous
& Braun (2003: 361).
Type: Indonesia: Jawa Barat (West-Java), on Saccharum
oficinarum (probably not preserved).
VOLUME 6 · NO. 1
Host range and distribution: On Brachiaria fasciculata
[Panicum
fasciculatum],
Cenchrus
(clandestinus
[Pennisetum clandestinum], purpureus [P. purpureum]),
Cymbopogon citratus, Cynodon dactylon, Digitaria insularis,
Echinochloa colona, Leptocoryphium lanatum, Panicum
maximum, Saccharum oficinarum, Zea mays, Poaceae,
Africa (Malawi, Mauritius, Nigeria, Sierra Leone, Senegal,
South Africa, Uganda), Asia (India, Indonesia, Malaysia,
Papua New Guinea, Philippines, Sri Lanka, Taiwan),
Australia, Central and South America (Brazil, Guatemala,
Venezuela), North America (Mexico; USA, Alabama,
Florida, Georgia, Gulf states, Louisiana), Oceania (Fiji,
Hawaii, Solomon Islands), and West Indies (Barbados,
Cuba, Dominican Republ., Haiti, Jamaica, Puerto Rico,
Virgin Islands).
ART I CLE
common in Pseudocercospora species, whereas colourless
conidia as well as amero- to phragmosporous conidia are
uncommon. The conidiogenous loci are unthickened, but
at least partly darkened by being refractive or even slightly
darkened-refractive, i.e. somewhat intermediate between
Passalora and Pseudocercospora. The structure of the
conidiogeneous loci and colourless conidia do not indicate
Passalora as currently circumscribed. Conidiogenous
loci and conidia are also reminiscent of species of
Distocercospora, although the conidia are euseptate. The
generic afinity of C. muhlenbergii is unclear, and possibly
a new genus is required pending molecular sequence
analyses.
Cercospora seminalis Ellis & Everh., J. Mycol. 4: 4
(1888).
Synonyms: Sporidesmium seminalis (Ellis & Everh.) U.
Braun, Cryptog. Mycol. 20: 175 (1999).
Porocercospora seminalis (Ellis & Everh.) Amaradasa,
Amundsen, Madrid & Crous, Mycologia 106: 81 (2014).
Literature: Saccardo (1892: 656), Chupp (1954: 252).
Illustrations: Braun (1999: 172, ig. 20), Amaradasa et al.
(2014: 83, ig. 2).
Lectotype (designated here, MycoBank MBT200458): usA:
Kansas: Manhattan, on seeds of Buchloë dactyloides, Jul.
1887, W. T. Swingle (BPI 441093B). Isolectotype: CUP
41186. Topotypes (from 8 Jun. 1888): BPI 441088, MICH
15365. Epitype (designated by Amaradasa et al. 2014): usA:
Nebraska: Mead, on seeds of Buchloë dactyloides, 5 Sep.
2011, B. S. Amaradasa (CBS H-21149). Ex-epitype culture:
CBS 134906.
Host range and distribution: On Buchloë dactyloides, Cynodon
dactylon, Poaceae, Central America (Guatemala), and North
America (USA, Colorado, Iowa, Kansas, North Dakota,
Nebraska, Oklahoma, South Dakota, Texas, Wisconsin).
Note: Type material was indicated as deposited at NY, but
was not found there, but two duplicates (isotypes) were
located which are used for lectotypiication here.
Cercospora striiformis G. Winter, Hedwigia 25: 103
(1886); as “striaeformis”.
Literature: Saccardo (1886: 383), Vassiljevsky & Karakulin
(1937: 270), Chupp (1954: 254), Crous & Braun (2003: 390).
Description (based on Winter 1886 and Chupp 1954): Leaf
spots linear, brown. Caespituli hypophyllous, punctiform.
Mycelium internal. Conidiophores in compact fascicles,
unbranched, slightly torulose, 120 × 4.5 µm, pluriseptate,
pale to medium brown. Conidia solitary, obclavate-cylindrical,
straight to curved, 45 × 1.5 µm, 3–5-septate, hyaline, base
subtruncate, apex obtuse.
63
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Braun et al.
Holotype: são tomé e Príncipe: on an unknown member of
Poaceae (not preserved).
Host range and distribution: Only known from the type
collection.
Notes: Application of the name Cercospora striiformis remains
uncertain as the identity of the type host of this species is
not known. The record of this species from Georgia on
Elymus elongatiformis [Elymus repens subsp. elongatiformis,
Elytrigia repens subsp. elongatiformis] is doubtful (Braun &
Mel’nik 1997). Type material could not be located. According
to Chupp (1954), this species is morphologically close to
Cercospora agrostidis, which also has short and narrow
conidia with few septa. Without type material the identity of
C. striiformis remains unclear.
Cercospora subulata (R. Sprague) R. Sprague,
Mycologia 29: 202 (1937).
Basionym: Cercosporella subulata R. Sprague, Mycologia
29: 202 (1937).
Synonym: Spermospora subulata (R. Sprague) R. Sprague,
Mycologia 40: 308 (1948).
Literature: Chupp (1954: 254), Braun (1995: 241), Crous &
Braun (2003: 393).
Illustration: Braun (1995: 242, ig. 224).
Holotype: usA: Oregon: Main Divide Trail, Ochoco National
Forest, on Melica subulata, 21 Aug. 1916, Ingram 606 (OSC
10.669). Isotype: NY 1042813). Topotype material: BPI
420956 (from 21 May 1916).
Host range and distribution: On Arrhenatherum elatius,
Bromus (inermis, marginatus, vulgaris), Calamagrostis
(rubescens, canadensis [scribneri]), Danthonia compressa,
Deschampsia cespitosa, Festuca rubra, Melica (bulbosa,
smithii, spectabilis, subulata), Oryza sativa, Poa nemoralis
[Agrostis alba], Puccinellia hauptiana, Sphenopholis obtusata
[Festuca obtusata], Trisetum spicatum, Poaceae, Caucasus
(Armenia), Europe (UK, Scotland), North America (Canada;
USA, Idaho, Montana, Oregon, Virginia, Washington,
Wyoming), and West Indies (Dominican Republic).
Note: The name Cercospora subulata is valid (alternative
names published before 1 Jan. 1953 are not invalid; Art.
36.2). This is most probably a North American species as
records on Festica rubra from Armenia (Simonyan 1981,
botanical garden) and Scotland (Foister 1961) as well as
Oryza sativa from the Dominican Republic (Ciferri 1961) are
unconirmed.
Passalora
Key to Passalora species on Poaceae
64
1
Supericial hyphae with solitary conidiophores in vivo developed (mycovellosiella-like) ................................................ 2
Supericial hyphae with solitary conidiophores in vivo lacking ........................................................................................ 7
2 (1)
Stromata present, 15–35 µm diam; conidiophores short, 9–37 µm; conidia long and narrow,
30–118 × 2.5–4 µm, hyaline; on Arthraxon hispidus ........................................................................... P. arthraxonis
Stromata absent; and/or conidiophores much longer and conidia much shorter and wider,
at least faintly pigmented ......................................................................................................................................... 3
3 (2)
Conidiophores in vivo consistently solitary, arising from supericial hyphae;
conidia subhyaline and narrow, 10–30 × 1–3 µm, 0–1-septate;
on Sorghum bicolor .................................................................................................................................. P. fujikuroi
Conidiophores in vivo fasciculate and solitary, arising from supericial hyphae,
and/or conidia wider, at least slightly pigmented ...................................................................................................... 4
4 (3)
Conidiophores 20–200 µm long; on Miscanthus, Saccharum and Sorghum spp. .......................................................... 5
Conidiophores shorter, 10–75 µm long; on Hystrix and Imperata spp. ........................................................................... 6
5 (4)
Stromata well-developed, large, 10–75 µm diam; conidiophores strictly solitary,
arising from supericial hyphae; conidiogenous loci small, 1–1.5 µm diam;
on Saccharum spp. ................................................................................................................................... P. vaginae
Stromata lacking or small, 10–20 µm diam; conidiophores in small fascicles and solitary;
conidiogenous loci larger, 1.5–2 µm diam; on Miscanthus, Saccharum and Sorghum spp.
....................................................................................................................................................................P. koepkei
6 (4)
Conidiophores consistently solitary, arsing from supericial hyphae, 6–8 µm wide;
conidia 4–6 µm wide, hyaline; on Imperata cylindrica .......................................................................... P. imperatae
Conidiophores solitary as well as in small, loose fascicles, 3–6 µm wide;
conidia 3–5 µm wide, subhyaline to pale olivaceous; on Hystrix patula ................................................. P. asprellae
IMA FUNGUS
cercosporoid fungi 3
Conidia in chains ............................................................................................................................................................. 8
Conidia solitary .............................................................................................................................................................. 14
8 (7)
Stromata large, 30–350 µm diam; conidiophores short, 5–15 × 3–6 µm, aseptate;
conidia 15–60 × 4–6.5 µm, 1–4-septate; on Paspalum clavuliferum .................................................. P. paspalicola
Stromata smaller, 10–60 µm diam; and/or conidiophores much longer and septate ...................................................... 9
9 (8)
Conidia short and broad, 11–23 × 5–11 µm, aseptate .................................................................................................. 10
Conidia longer and/or narrower, septate, at least 1-septate .......................................................................................... 11
ART I CLE
7 (1)
10 (9) Conidiogenous loci 1–2 µm diam; on Phragmites australis ........................................................................ P. maculicola
Conidiogenous loci larger, (1–)1.5–2.5(–3) µm diam; on Phalaris arundinacea .......................................... P. phalaridis
11 (9) Stromata large, 30–100 µm diam; conidiophores long, 80–160 µm; conidia 3.5–7 µm wide;
on Cenchrus bambusiformis [Pennisetum bambusiforme] ......................................................... P. tungurahuensis
Stromata smaller, to 50 µm diam; conidiophores shorter, 10–90 µm; conidia narrower,
1.5–5 µm wide ....................................................................................................................................................... 12
12 (11) Conidia narrow, 18–38 × 1.5–2 µm; on Agrostis sp. ............................................................................... P. agrostidicola
Conidia much wider, 2–5 µm wide ................................................................................................................................. 13
13 (12) Conidiophores in small fascicles of to 10, 15–45 µm long; conidia often in branched chains;
on Dichanthium annulatum .................................................................................................... P. dichanthii-annulati
Conidiophores in larger fascicles of 13–25, 30–90 µm long; conidia mostly in unbranched chains;
on Digitaria diagonalis ............................................................................................................................ P. digitariae
14 (7) Stromata well-developed, large, 100–500 µm diam; conidiophores long, 40–150 µm; conidia narrow,
20–65 × 1.5–4 µm, mostly 1-septate; on Chloris and Eustachys spp. ................................................. P. caespitosa
Stromata lacking or smaller, 10–130 µm diam and/or conidia much broader, 3–8 µm; on other hosts ........................ 15
15 (14) Stromata large, 50–150 µm diam; conidiophores 50–300 µm long; conidia 20–55 × 5–8 µm,
(1–)3(–4)-septate; on Arundinaria spp. .................................................................................................. P. compacta
Stromata lacking or smaller, usually < 100 µm; conidiophores mostly < 100 µm in length;
conidia 0–2-septate or broader, (5–)6–12(–14) µm ............................................................................................... 16
16 (15) Conidia 3–6 µm wide .................................................................................................................................................... 17
Conidia broader, 5–14 µm wide .................................................................................................................................... 18
17 (16) Stromata absent; conidiophores solitary or in loose groups; conidia subhyaline;
on Brachyelytrum erectum ................................................................................................................. P. brachyelytri
Stromata present, small, 10–25 µm diam; conidiophores distinctly fasciculate;
conidia subhyaline to olivaceous-brown; on Leersia oryzoides ...................................................... P. ramularioides
18 (16) Stromata variable in shape and size, 20–130 µm diam; conidia (15–)20–50(–60) × (5–)6–12(–14) µm,
smooth or almost so; on a wide range of grasses .................................................................................. P. graminis
Stromata applanate to oblong, to 60 × 20 µm; conidia 12–38 × 5–10.5 µm, verruculose;
on Milium effusum ........................................................................................................................................... P. milii
Tabular key to Passalora species on Poaceae
The species are listed in form of a tabular key based on host genera in alphabetical order.
ˣAgrohordeum .......................................................................................................................................................... P. graminis
Agropyron ................................................................................................................................................................. P. graminis
ˣAgrositanion ............................................................................................................................................................ P. graminis
VOLUME 6 · NO. 1
65
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Braun et al.
Agrostis
1
Conidia formed singly, 15–60 × 5–14 µm, (0–)1(–3)-septate ......................................................................... P. graminis
Conidia solitary and in chains, 18–38 × 1.5–2 µm, 3–4-septate ............................................................. P. agrostidicola
Alopecurus ................................................................................................................................................................ P. graminis
Ammophila ................................................................................................................................................................ P. graminis
Anthoxanthum .......................................................................................................................................................... P. graminis
Arctagrostis .............................................................................................................................................................. P. graminis
Arrhenatherum .......................................................................................................................................................... P. graminis
Arthraxon .............................................................................................................................................................. P. arthraxonis
Arundinaria
1
Conidia (0–)1(–3)-septate .............................................................................................................................. P. graminis
Conidia (1–)3(–4)-septate ............................................................................................................................. P. compacta
Avena ........................................................................................................................................................................ P. graminis
Beckmannia .............................................................................................................................................................. P. graminis
Brachyelytrum ..................................................................................................................................................... P. brachyelytri
Bromus ...................................................................................................................................................................... P. graminis
Calamagrostis ........................................................................................................................................................... P. graminis
Cenchrus (incl. Pennisetum)
1
Conidia solitary and in chains, 15–60 × 3.5–7 µm, (0–)1–4(–5)-septate ........................................... P. tungurahuensis
Conidia solitary, 15–60 × 5–14 µm, (0–)1(–3)-septate ................................................................................... P. graminis
Chloris .................................................................................................................................................................... P. caespitosa
Cinna ........................................................................................................................................................................ P. graminis
Cynodon .................................................................................................................................................................... P. graminis
Cynosurus ................................................................................................................................................................. P. graminis
Dactylis ...................................................................................................................................................................... P. graminis
Danthonia .................................................................................................................................................................. P. graminis
Deschampsia ............................................................................................................................................................ P. graminis
Dichanthium ............................................................................................................................................. P. dichanthii-annulati
Digitaria
1
Conidia formed singly, 15–60 × 5–14 µm, (0–)1(–3)-septate ......................................................................... P. graminis
Conidia solitary and in chains, 20–55 × 2.5–3 µm, 1–5-septate ................................................................... P. digitariae
ˣElyhordeum ............................................................................................................................................................. P. graminis
Elymus ....................................................................................................................................................................... P. graminis
ˣElysitanion ............................................................................................................................................................... P. graminis
66
IMA FUNGUS
cercosporoid fungi 3
Eragrostis .................................................................................................................................................................. P. graminis
Festuca ...................................................................................................................................................................... P. graminis
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Eustachys ............................................................................................................................................................... P. caespitosa
Glyceria ..................................................................................................................................................................... P. graminis
Helictotrichon ........................................................................................................................................................... P. graminis
Hierochloe ................................................................................................................................................................. P. graminis
Homalotrichon .......................................................................................................................................................... P. graminis
Hordeum .................................................................................................................................................................... P. graminis
Hystrix
1
Mycelium in vivo internal; stromata developed; conidia 15–60 × 5–14 µm, (0–)1(–3)-septate ...................... P. graminis
Mycelium in vivo internal and external; stromata lacking or almost so; conidia 20–75 × 3–5 µm,
2–6-septate ............................................................................................................................................. P. asprellae
Imperata ................................................................................................................................................................... P. imperatae
Koeleria ..................................................................................................................................................................... P. graminis
Leersia ....................................................................................................................................................................... P. graminis
Leucopoa ................................................................................................................................................................... P. graminis
Leymus ...................................................................................................................................................................... P. graminis
Lolium ........................................................................................................................................................................ P. graminis
Leersia ............................................................................................................................................................... P. ramularioides
Melica ........................................................................................................................................................................ P. graminis
Milium
1
Conidia verruculose ............................................................................................................................................... P. milii
Conidia smooth or almost so .......................................................................................................................... P. graminis
Miscanthus
1
Mycelium in vivo internal and external; solitary conidiophores arising from supericial hyphae;
conidia 20–65 × 4–6.5 µm ........................................................................................................................ P. koepkei
Mycelium in vivo internal; solitary conidiophores lacking; conidia 15–60 × 5–14 µm, (0–)1(–3)-septate ....... P. graminis
Muhlenbergia ............................................................................................................................................................ P. graminis
Nassella ..................................................................................................................................................................... P. graminis
Oryzopsis .................................................................................................................................................................. P. graminis
Panicum .......................................................................................................................................................................................... P. graminis
Pennisetum, see Cenchrus
Phalaris
1
Conidia solitary, 15–60 × 5–14 µm, (0–)1(–3)-septate ................................................................................... P. graminis
Conidia catenate, 12–23 × 5–10.5 µm, aseptate ......................................................................................... P. phalaridis
VOLUME 6 · NO. 1
67
Braun et al.
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Phleum ....................................................................................................................................................................... P. graminis
Phragmites
1
Conidia solitary, 15–60 × 5–14 µm, (0–)1(–3)-septate ................................................................................... P. graminis
Conidia catenate, 11–23 × 5–11 µm, aseptate ............................................................................................ P. maculicola
Poa ............................................................................................................................................................................. P. graminis
Pseudosclerochloa ................................................................................................................................................... P. graminis
Puccinellia ..................................................................................................................................................................................... P. graminis
Paspalum ............................................................................................................................................................... P. paspalicola
Saccharum
1
Stromata well-developed, large, 10–75 µm diam; conidiophores strictly solitary,
arising from supericial hyphae; conidiogenous loci small, 1–1.5 µm diam .............................................. P. vaginae
Stromata absent or small, 10–20 µm diam; conidiophores in small fascicles and solitary;
conidiogenous loci larger, 1.5–2 µm diam ................................................................................................ P. koepkei
Secale ........................................................................................................................................................................ P. graminis
Sorghum
1
Conidiophores 15–50 × 3–5 µm; conidia 10–30 µm long, 1–3 µm wide at the ends,
0–1-septate, subhyaline ............................................................................................................................ P. koepkei
Conidiophores longer, 30–200 × 4–7 µm; conidia 20–65 × 4–6.5 µm, 1–7-septate,
almost colourless to olivaceous-brown .................................................................................................... P. fujikuroi
Spartina ..................................................................................................................................................................... P. graminis
Stenotaphrum ........................................................................................................................................................... P. graminis
Stipa ........................................................................................................................................................................... P. graminis
Syagrus ..................................................................................................................................................................... P. graminis
Torreyochloa ............................................................................................................................................................. P. graminis
Trisetum .................................................................................................................................................................... P. graminis
Triticum ..................................................................................................................................................................... P. graminis
List of Passalora species on Poaceae
Passalora agrostidicola Phengs. & U. Braun, sp.
nov.
MycoBank MB811239
(Fig. 39)
Literature: Phengsintham et al. (2013: 103), as Passalora
fusimaculans.
Illustration: Phengsintham et al. (2013: 103, igs 50–51), as
Passalora fusimaculans.
Diagnosis: Distinguished from all species of Passalora on
grasses by the much narrower conidia (1.5–2 µm wide),
except for P. fujikuroi, which is mycovellosielloid, i.e. in vivo
with solitary conidiophores arising from supericial hyphae.
68
Description: Leaf spots circular to irregular, 1–5 mm diam,
reddish to medium brown in the centre, margin brown to dark
brown. Caespituli amphigenous, scattered. Mycelium internal,
inconspicuous. Stromata well-developed, substomatal,
subglobose, 20–50 µm diam, brown to dark brown, composed
of swollen hyphal cells, oval, ellipsoid to angular in outline,
wall 0.3–0.5 µm wide. Conidiophores fasciculate, 6–24, arising
from substomatal stromata, emerging through stomata, or
arising from almost supericial stromatic hyphal aggregations,
erect, straight to curved, unbranched, subcylindrical or slightly
narrowed towards the tip, 10–52 × 3–5 µm, 0–1-septate, pale to
moderately olivaceous-brown, wall 0.3–0.5 µm wide, smooth;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, 10–30 µm long, 3–5 µm wide
below and 1.5–2 µm above, conidiogenous loci conspicuous,
thickened and darkened, 1.5–2 µm diam. Conidia solitary or
catenate, in simple chains, cylindrical or subcylindrical, straight
IMA FUNGUS
cercosporoid fungi 3
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Fig. 39. Passalora agrostidicola (MFLU10-0317, holotype). A.
Conidiophore fascicles. B. Conidia. Bar = 10 µm.
to somewhat curved, 18–38 × 1.5–2 µm, 3–4-septate, slightly
constricted at the septa, pale olivaceous, wall thin (0.2–0.3
µm), smooth to inely verruculose, both ends subtruncate
when catenate, apex bluntly rounded in solitary and primary
conidia, apical hila 0.5–0.8 µm wide, basal hila 0.5–1.5 µm
wide, somewhat thickened and darkened.
Holotype: thailand: Chiang Rai Province: Doi Tung National
Park, on living leaves of Agrostis sp., Poaceae (Pooideae,
Aveneae), 18 Aug. 2009, P. Phengsintham (MFLU10-0317).
Host range and distribution: Only known from the type
collection.
Notes: Phengsintham et al. (2013) identiied the type
collection of this species as Passalora fusimaculans, including
Cercospora agrostidis as a synonym, which is now treated
as separate species. Passalora agrostidicola is distinguished
from C. agrostidis by its circular to irregular leaf spots,
larger stromata, 0–1-septate conidiophores, and shorter
and narrower, olivaceous, partly verruculose, cylindrical to
subcylindrical conidia. The collection on Agrostis from Thailand
is not conspeciic with C. agrostidis (as well as C. fusimaculans
s. lat.). Based on its pigmented conidia, the fungus from
Thailand is better placed in Passalora. Cercospora agrostidis
and C. fusimaculans are now treated as species of Cercospora
s. str. due to their colourless conidia.
VOLUME 6 · NO. 1
Fig. 40. Passalora arthraxonis (HMAS 51952, holotype). A.
Conidiophore fascicle. B. Solitary conidiophores arising from
supericial hyphae. c. Conidia. Bar = 10 µm.
Passalora arthraxonis (Y.L. Guo) U. Braun & Crous,
in Crous & Braun, Mycosphaerella and Anam.: 438
(2003)
(Fig. 40)
Basionym: Mycovellosiella arthraxonis Y.L. Guo, Mycosystema 21: 497 (2002).
Literature: Guo et al. (2003: 30–31).
Illustrations: Guo & Xu (2002: 498. ig. 1), Guo et al. (2003:
31, ig. 16).
Description: Leaf spots amphigenous, subcircular to elliptical,
0.5–1.5 mm diam, often conluent, margin indeinite, at irst
olivaceous-brown, later brown to dark brown, with yellowish
69
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Braun et al.
brown halo on the upper surface, brown to greyish brown
below. Caespituli amphigenous. Mycelium internal and
external; superical hyphae emerging through stomata,
sometimes forming loose ropes, often climbing leaf hairs,
branched, septate, 1.5–3.2 µm wide, subhyaline, thin-walled,
smooth. Stromata lacking or substomatal, subglobose, 15–
35 µm diam, pale olivaceous-brown. Conidiophores loosely
fasciculate, 5–10, arising from stromata, through stomata
or solitary, arising from supericial hyphae, lateral, straight
to curved, subcylindrical to attenuated towards the tip,
unbranched, 1–3 times geniculate, about 9–37 × 3–5.5 µm,
0–2-septate, pale olivaceous to olivaceous or brown at the
base, thin-walled, smooth; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells,
conidiogenous loci conspicuous, thickened and darkened,
1.5–2.5 µm diam. Conidia solitary, narrowly obclavate to
acicular, straight to curved, 30–118 × 2.5–4 µm, 3–12-septate,
hyaline, thin-walled, smooth, apex acute to obtuse, base
obconically truncate, hila thickened and darkened.
Holotype: china: Zhejiang Province: Hangzhou, on Arthraxon
hispidus, Poaceae (Panicoideae, Andropogoneae), 23 Sep.
1961, Q. M. Ma & X. J. Liu 408 (HMAS 51952).
Host range and distribution: Only known from the type
collection.
Notes: The generic afinity of this species is not clear. The
general habit of P. arthraxonis is mycovellosielloid, but the
conidia are formed singly, cercospora-like and colourless. It
may belongs to Cercospora s. str., but this question remains
unresolved.
Passalora asprellae (U. Braun) U. Braun & Crous,
in Crous & Braun, Mycosphaerella and Anam.: 69
(2003).
(Fig. 41)
Basionym: Mycovellosiella asprellae U. Braun, Sydowia 48:
206 (1996).
Synonym: Cercospora asprellae Ellis & Galloway, in herb.
Literature: Chupp (1954: 256).
Illustration: Braun (1996: 207, ig. 2).
Description: Leaf spots amphigenous, narrow, oblong, 1–2
× 0.5 mm, later conluent, forming narrow streaks, to about
10 mm long, dark, blackish, margin indeinite. Caespituli
amphigenous, mostly hypophyllous, dense, velvety, dull
greyish brown. Mycelium internal and external; supericial
hyphae branched, septate, 2–6 µm wide, subhyaline
to olivaceous-brown, thin-walled, smooth, often dense,
intertwined or forming ropes. Stromata lacking or only with
small, brown hyphal aggregations. Conidiophores in small,
loose fascicles, arising from stromatic hyphal aggregations
or solitary, arising from supericial hyphae, erect, straight,
subcylindrical to geniculate-sinuous, unbranched, 10–70 ×
3–6 µm, aseptate or only sparingly septate, pale olivaceous
to olivaceous-brown, thin-walled, smooth; conidiogenous
cells integrated, terminal or conidiophores reduced to
70
Fig. 41. Passalora asprellae (NY 985492, holotype). A. Supericial
hyphae. B. Solitary conidiophores arising from supericial hyphae. c.
Conidiophore fascicle. d. Conidiophores. e. Conidia. Bar = 10 µm.
conidiogenous cells, 10–30 µm long, conidiogenous loci
conspicuous, 1–2 µm diam, thickened and darkened. Conidia
solitary, fusiform, obclavate, 20–75 × 3–5 µm, 2–6-septate,
rarely somewhat constricted at the septa, subhyaline to
pale olivaceous, thin-walled, smooth, apex pointed, base
obconically truncate, 1.5–2 µm wide, hila thickened and
darkened.
Holotype: usA: Oregon: on Hystrix patula, 10 Sep. 1889, M.
B. Waite 199 (NY 985492).
IMA FUNGUS
cercosporoid fungi 3
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Host range and distribution: On Hystrix patula [Asprella
hystrix, Elymus hystrix], Poaceae (Pooideae, Triticeae),
North America (USA, Illinois, Oregon).
Passalora brachyelytri (H.C. Greene) U. Braun &
Bensch, comb. nov.
MycoBank MB811243
(Fig. 42)
Basionym: Cladosporium brachyelytri H.C. Greene, Trans.
Wisconsin Acad. Sci. 53: 214 (1964).
Literature: Bensch et al. (2012: 302).
Illustrations: Schubert (2005a: 64, ig. 5), Bensch et al. (2012:
302, ig. 358).
Description: Leaf spots distinct, numerous, scattered, narrow,
oblong to oblong-elliptical, 1–3(–7) mm long and to 1 mm
wide, reddish brown, surrounded by a paler reddish brown
halo, often along leaf veins, occasionally conluent. Caespituli
hypophyllous, scattered, loosely villose, pale, whitish to pale
brown. Mycelium internal. Stromata lacking. Conidiophores
solitary or in small, loose groups, arising from internal hyphae,
emerging through stomata, erect, straight to somewhat
lexuous, geniculate-sinuous, mostly unbranched, rarely
apically branched, 48–130 × 3.5–5.5 µm, slightly attenuated
towards the apex, septate, not or somewhat constricted at
the septa, pale brown or olivaceous-brown, wall somewhat
thickened, smooth; conidiogenous cells integrated, terminal
and intercalary, 12–35 µm long, proliferation sympodial,
geniculate, with a single to several conspicuous conidiogenous
loci, protuberant, subdenticulate, planate, 1.5–2 µm diam,
somewhat thickened and darkened-refractive. Conidia in
unbranched chains, straight, subcylindrical, subfusiform,
rarely somewhat obclavate, (6–)14–27 × 3–5(–6) µm,
1-septate, septum more or less median, not or only slightly
constricted at the septum, subhyaline, thin-walled, smooth,
apex rounded, slightly pointed or mostly truncate, base
truncate, 1–2 µm wide, with protuberant hilum, thickened and
darkened-refractive.
Lectotype (designated in Bensch et al. 2012: 302): usA:
Wisconsin: Sawyer Co., Flambeau State Forest near Oxbow,
on Brachyelytrum erectum, 22 Jul. 1964, H.C. Greene (WIS).
Isolectotype: BPI 426166.
Fig. 42. Passalora brachyelytri (WIS, holotype). A. Leaf spots. B.
Conidiophore fascicles. c. Conidiophores. d. Conidia. Bar = 10 µm.
U. Braun del.
of Cercospora s. str. The generic afinity of Cladosporium
brachyelytri is not clear, but due to the consistently
didymosporous, almost colourless conidia, we prefer to place
this species in Passalora s. lat. pending molecular studies.
Passalora caespitosa (Ellis & Everh.) U. Braun,
Cryptog. Mycol. 20: 165 (1999).
(Fig. 43)
Basionym: Cercospora caespitosa Ellis & Everh., Proc. Acad.
Sci. Philadelphia I, 43: 88 (1891).
Host range and distribution: On Brachyelytrum erectum,
Poaceae (Pooideae, Brachyelytreae), North America (USA,
Wisconsin).
Literature: Chupp (1954: 244), Crous & Braun (2003: 92).
Notes: This species is readily distinguishable from Cercospora
fusimaculans (syn. Passalora fusimaculans) by its much
broader conidia. Schubert (2005a) reduced Cladosporium
brachyelytri to synonymy with Passalora fusimaculans
var. barretoana (now Cercospora barretoana).Cercospora
barretoana, conined to hosts in the Panicoideae, differs from
P. brachyelytri (on Brachyelytrum, Pooideae, Brachyelytreae)
in having 0–4(–7)-septate, much longer conidia, 9–70 µm.
Based on colourless, cercosporoid conidia, Passalora
barretoana and P. fusimaculans are now treated as species
Exsiccatae: Ellis & Everh., North Amer. Fungi 2590, 3192.
VOLUME 6 · NO. 1
Illustration: Braun (1999: 163, ig. 10).
Description: Leaf spots lacking. Caespituli amphigenous,
mostly hypophyllous, punctiform to pustulate, blackish.
Mycelium internal. Stromata immersed to somewhat
erumpent, large, about 100–500 µm diam, dark brown to
blackish, composed of swollen hyphal cells, 2–8 µm diam,
pale to medium dark brown. Conidiophores numerous, in
dense fascicles, arising from stromata, erumpent, forming
sporodochial conidiomata, straight, subcylindrical to slightly
71
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Lectotype (designated here, MycoBank, MBT200459): usA:
Mississippi: Ocean Springs, on Eustachys petraea, 30 Aug.
1881, S. M. Tracy 1215 (BPI 433902). Isolectotype: MICH
15265. Former syntypes: Ocean Springs, on Eustachys
petraea, 16 Sep. 1889, S. M. Tracy (CUP 39255) and Ellis
& Everh., North Amer. Fungi 3190 (e.g. BPI 433904, CUP,
OSC 53156).
Host range and distribution: On Chloris gayana, Eustachys
(neglecta, petraea [Chloris petraea, swartziana]), Poaceae
(Chloridoideae), North America (USA, Florida, Michigan,
Mississippi, Wisconsin).
Notes: The “lectotypiication” cited in Braun (1999) is incorrect
since the proposed material (Ellis & Everh., North Amer.
Fungi 3192) was collected in 1893, i.e. after the publication
of this species name.
Passalora compacta (Berk. & M.A. Curtis) U. Braun
& Crous, in Crous & Braun, Mycosphaerella and
Anam.: 133 (2003).
(Fig. 44)
Basionym: Cladosporium compactum Berk. & M.A. Curtis,
Grevillea 3: 106 (1875).
Synonyms: Cercosporidium compactum (Berk. & M.A. Curtis)
Deighton, Mycol. Pap. 112: 59 (1967).
Cercospora scolecotrichoides G.F. Atk., Cornell Univ. Bull.
3: 46 (1897) [lectotype (designated here, MycoBank,
MBT200461): usA: Alabama: Lee Co., Auburn, on
Arundinaria gigantea subsp. tecta, 28 Oct. 1891, B. M.
Duggan 2293 (CUP-A-2293); isolectotypes: CUP 41180,
K(M) IMI 95405].
Literature: Saccardo (1886: 364; 1899: 1106), Lindau (1907:
833), Chupp (1954: 251), Deighton (1967: 59), Crous &
Braun (2003: 133), Schubert (2005b: 202), Bensch et al.
(2013: 307).
Illustrations: Deighton (1967: 59, ig. 30, 61, ig. 31).
Fig. 43. Passalora caespitosa (BPI 433902, lectotype). A.
Conidiophore fascicle. B. Conidiophore tips. c. Conidia. Bar = 10
µm.
lexuous-sinuous, barely to slightly geniculate, unbranched,
40–150 × 3–6 µm, width uniform, 0–2-septate, pale
olivaceous to medium brown, medium dark brown in mass,
tips often paler, wall thin to slightly thickened, smooth;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, 20–120 µm long,
conidiogenous loci conspicuous, thickened and darkened,
1.5–2 µm diam, not or barely protuberant. Conidia solitary,
cylindrical, cylindrical-obclavate, fusiform, 20–65 × 1.5–4
µm, (0–)1(–3)-septate, subhyaline to pale yellowish brown,
thin-walled, smooth, apex obtuse, base short obconically
truncate, 1–2 µm wide, hilum barely thickened, somewhat
darkened.
72
Description: Leaf spots lacking or almost so. Caespituli
hypophyllous, more or less evenly scattered, punctiform,
dark. Mycelium internal. Stromata well-developed, oblong,
50–150 µm, dark brown. Conidiophores in large, compact
fascicles (to 100 or even more), looser when young, very
dense and sometimes even subcoremioid when older, arising
from stromata, erumpent, rupturing the cuticle, erect, straight
to somewhat curved-sinuous, mostly geniculate-sinuous
above, often strongly so, unbranched or rarely branched,
50–300 × 3–6.5 µm, loosely pluriseptate, pale, olivaceous,
thin-walled, smooth; conidiogenous cells integrated, terminal
and intercalary, 10–45 µm long, geniculate, conidiogenous
loci conspicuous, thickened and darkened, prominent, 1.5–
2.5 µm diam. Conidia solitary, broadly ellipsoid-fusiform
to obclavate, straight to slightly curved, 20–55 × 5–8 µm,
(1–)3(–4)-septate, pale olivaceous, thin-walled, smooth,
apex usually attenuated towards an obtuse or subobtuse
tip, occasionally broadly rounded, base short obconically
truncate, about 2 µm wide, hila somewhat thickened and
darkened.
IMA FUNGUS
cercosporoid fungi 3
ART I CLE
Fig. 44. Passalora compacta (K(M) IMI 193890, lectotype). A.
Conidiophore fascicle. B. Conidiophore tips. c. Conidia. Bar = 10
µm.
Lectotype (designated here, MycoBank, MBT200460):
usA: “N. America, no. 3767”, on Arundinaria gigantea subsp.
tecta, ex herb. Broome (K(M) 193890). Isolectotypes: K(M)
193891, K(M) IMI 69771, STR (ex herb. Curtis).
Host range and distribution: On Arundinaria (gigantea subsp.
tecta [tecta], Arundinaria sp.), Poaceae (Bambusoideae,
Bambuseae), North America (USA, Alabama, Florida).
Passalora dichanthii-annulati (Chaudhary,
Singh & P.N. Singh) U. Braun, comb. nov.
MycoBank MB811244
(Fig. 45)
S.K.
Basionym: Phaeoramularia dichanthii-annulati Chaudhary,
S.K. Singh & P.N. Singh, Indian Phytopathol. 55: 469
(2002); as “dicanthii-annulatae”.
Illustration: Chaudhary et al. (2002: 469, ig. 1).
Description: Leaf spots amphigenous, small, dark
brown. Caespituli hypophyllous, effuse, brown. Mycelium
internal.
Stromata
well-developed,
subepidermal,
pseudoparenchymatous, about 20–25 µm diam, pale
olivaceous. Conidiophores in fascicles, to 10, arising from
VOLUME 6 · NO. 1
Fig. 45. Passalora dichanthii-annulati (based on Chaudhary et al.
2002: 469, ig. 1). A. Conidiophore fascicle. B. Conidiophore tip. c.
Conidia. Bar = 10 µm.
stromata, erect, straight to lexuous, geniculate, unbranched,
15–45 × 3–4 µm, 1–5-septate, light olivaceous, thinwalled, smooth; conidiogenous cells integrated, terminal,
conidiogenous loci conspicuous, thickened and darkened.
Conidia solitary to catenate, in simple or often branched
chains, cylindrical or subcylindrical, straight to curved, 13–68
× 2–5 µm, 2–7-septate, light olivaceous, thin-walled, smooth,
apex subacute to obtuse, base obconically truncate, hila
thickened and darkened.
Holotype: India: Uttar Pradesh: Gorakhpur, on Dichanthium
annulatum, Poaceae (Panicoideae, Andropogoneae), Nov.
1994, R. K. Chaudhary (HCIO 42571).
Host range and distribution: Only known from the type
collection.
73
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Notes: This species is morphologically close to Passalora
barretoana, but differs in having more cylindrical conidia often
formed in branched chains. The genus Dichanthium belongs
in the Andropogoneae. All known hosts of P. barretoana are
members of the Paniceae.
Passalora digitariae (Crous & B. Sutton) Crous & U.
Braun, Mycosphaerella and Anam.: 451 (2003).
(Fig. 46)
Basionym: Phaeoramularia digitariae Crous & B. Sutton, S.
Afr. J. Bot. 63: 282 (1997).
Illustration: Crous & Sutton (1997: 282, ig. 4).
Description: Leaf spots amphigenous, narrowly elliptical, 3–8 mm
long and 0.5–2 mm wide, light brown. Caespituli amphigenous,
scattered, distinct, punctiform, dark brown. Mycelium internal;
hyphae branched, septate, 2–4 µm wide, hyaline to olivaceous,
thin-walled, smooth. Stromata well-developed, substomatal,
20–40 µm diam, dark brown. Conidiophores in fascicles, 13–
25, arising from stromata, through stomata, erect, divergent,
straight, subcylindrical to slightly sinuous, unbranched, 30–90
× 3–5 µm (combined length of stromata and conidiophores
40–110 µm), 1–5-septate, medium brown, paler towards the tip,
thin-walled, smooth; conidiogenous cells integrated, terminal,
15–35 µm long, conidiogenous loci conspicuous, thickened and
darkened, about 1.5–2 µm diam. Conidia solitary to catenate,
in simple chains, subcylindrical-obclavate, 20–55 × 2.5–4 µm,
1–5-septate, pale olivaceous, thin-walled, smooth, apex obtuse
to subtruncate in catenate conidia, base short obconically
truncate, about 1.5–2 µm wide, hila somewhat thickened and
darkened.
Holotype: south Africa: KwaZulu-Natal: Pietermaritzburg,
Nottingham Rd., on Digitaria diagonalis, Poaceae
(Panicoideae, Paniceae), Mar. 1939, A. P. D. McClean
(PREM 33113).
Host range and distribution: Only known from the type
collection.
Passalora fujikuroi (N. Pons) U. Braun & Crous,
Mycosphaerella and Anam.: 190 (2003).
(Fig. 47)
Basionym: Mycovellosiella fujikuroi N. Pons, Ernstia 6: 42
(1996).
Synonym: Cercospora andropogonis Sawada, Special
Publ. Coll. Agric. Natl. Taiwan Univ. 8: 226 (1959), nom.
inval. (Art. 38.1) [authentic material: taiwan: Taipei, on
Sorghum bicolor [vulgare], 6 Nov. 1909, K. Fujiroko (NTUPPE, hb. Sawada; TNS-F-218232; BPI 432655)].
Illustration: Pons (1996: 46, ig. 1).
Description: Lesions not distinct. Caespituli epiphyllous.
Mycelium internal and external; supericial hyphae emerging
through stomata, branched, 1.5–3 µm wide, subhyaline,
thin-walled, smooth. Stromata lacking or small, substomatal,
textura angularis, pigmented. Conidiophores solitary,
arising from supericial hyphae, lateral, or arising from small
74
Fig. 46. Passalora digitariae (PREM 33113, holotype). A.
Conidiophore fascicle. B. Conidia. Bar = 10 µm.
stromata in small, loose fascicles, through stomata, erect,
straight or lexuous, subcylindrical, unbranched, 15–50 × 3–5
µm, aseptate or 1–2-septate, pale to dark brown, thin-walled,
smooth; conidiogenous cells integrated, terminal, intercalary
or conidiophores reduced to conidiogenous cells, proliferation
sympodial, 7–30 µm long, 3–5 µm wide below and 2.5–4 µm
wide above; conidiogenous loci conspicuous, unthickened to
somewhat thickened. Conidia solitary, subcylindrical to short
obclavate, straight to curved, 10–45 µm long, base and apex
1–3 µm wide, 0–4-septate, subhyaline to pale olivaceous,
thin-walled, smooth, apex obtuse, base obconically truncate,
basal hilum thickened and darkened.
Holotype: taiwan: Taipei, on Sorghum bicolor, 6 Nov. 1909, Y.
Fujikuro (NTU-PPE, hb. Sawada). Isotype: TNS-F-218232).
Host range and distribution: On Sorghum bicolor [vulgare],
Poaceae (Panicoideae, Andropogoneae), Asia (Taiwan).
IMA FUNGUS
cercosporoid fungi 3
ART I CLE
Fig. 47. Passalora fujikuroi (NTU-PPE, holotype). A. Solitary
conidiophores. B. Conidia. Bar = 10 µm.
Fig. 48. Passalora graminis (HAL, lectotype). A. Conidiophore
fascicles. B. Conidiophores. c. Conidia. Bar = 10 µm.
Notes: The combination of morphological characters in this
species is unusual. Its generic afinity is unclear, as its conidia
are subhyaline or pale and thus cercospora-like. On the other
hand, solitary conidiophores in vivo arising from supericial
hyphae are not typical of Cercospora s. str. Presently this
species is maintained in Passalora s. lat. Based on results
of the examination of isotype material, the assignment of
this species to the latter genus was conirmed. Sawada
(1959) and Goh & Hsieh (1990) cited C. andropogonis as
a synonym of Cercospora sorghi, a true Cercospora s. str.,
which is in conlict with the examinations of Pons (1996) and
our own observations. It is possible that authentic material
of C. andropogonis (nom. nud.) originally encompassed
conidiophores and conidia of two cercosporoid fungi,
although this is not proven.
VOLUME 6 · NO. 1
Passalora graminis (Fuckel) Höhn., Zentralbl.
Bakteriol. Parasitenk., Abt. 2, 60: 6 (1923).
(Fig. 48)
Basionym: Scolicotrichum graminis Fuckel, Hedwigia 2(15):
134 (1863).
Synonyms: Passalora hordei G.H. Otth, Mitth. Naturf. Ges.
Bern 1868: 66 (1868) [holotype: switzerland: Bern, on
Hordeum distichum, G. H. Otth (ZT)].
Passalora punctiformis G.H. Otth, Mitth. Naturf. Ges. Bern
1868: 67 (1868) [syntypes: switzerland: Bern, on
Dactylis glomerata and Melica unilora, G. H. Otth (?ZT,
host as “Arrhenaterum elatius”).
Cladosporium sphaeroideum Cooke, Grevillea 8(46): 60
(1879) [holotype: New Zealand: Counterbury Alps, on
75
ART I CLE
Braun et al.
Poa foliosa, ex herb. M. C. Cooke 398 (K(M) 121569)].
Cercospora graminicola Tracy & Earle, Bull. Torrey Bot.
Club 22: 179 (1895) [lectotype (designated here,
MycoBank, MBT200462): usA: Mississippi: Starkville, on
Phleum pratense, 4 Nov. 1894, F. S. Earle (BPI 436794);
isolectotypes: BPI 436793, CUP 39906, NY 937032,
937033].
Scolicotrichum compressum Allesch., in Syd., Mycoth. March.
4388 (1895) and Hedwigia 35: (34) (1896) [lectotype
(designated here, MycoBank, MBT200463): germany:
Berlin, Lichterfelde, on Poa compressa, Sep. 1895, P.
Sydow (BPI 425109); isolectotypes: Syd., Mycoth. March.
4388, e.g. B, PAD].
Scolicotrichum graminis var. nanum Sacc., Ann. Mycol. 3:
515 (1905) [holotype: Italy: Belluno, Agordo, on Dactylis
glomerata, D. Saccardo (not preserved in PAD)].
Scolicotrichum graminis var. brachypodum Speg., An. Mus.
Nac. Buenos Aires, ser. 3, 13: 436 (1911) [syntypes:
Argentina: on Hordeum jubatum, Bromus unioloides,
etc., not speciied].
Cercospora graminis (Fuckel) Horsfall, Mem. Cornell Univ.
Agric. Exp. Sta. 130: 100 (1930).
Heterosporium secalis Dippen., South African J. Sci. 28:
286 (1931) [holotype: south Africa: Cape Province:
Stellenbosch, on Secale cereale, 25 Sep. 1929, B. J.
Dippenar 31 (PREM 46907)].
Cercosporidium graminis (Fuckel) Deighton, Mycol. Pap.
112: 62 (1967).
Passalora compressa (Allesch.) Petr., Reliquiae Petrakianae
1: 50 (No. 192) (1977), comb. inval. (Art. 41.5).
Passalora graminis (Fuckel) Poonam Srivast., J. Living World
1: 116 (1994), comb. inval. et illegit. (Art. 41.5).
Literature: Saccardo (1886: 348; 1895: 617; 1897: 682; 1911:
774; 1913: 1374), Vassiljevsky & Karakulin (1937: 212, 272),
Sprague (1950: 424–429), Chupp (1954: 247), Deighton
(1967: 62), Ellis (1971: 281), Crous & Braun (1996: 273),
David (1997: 121), Crous & Braun (2003: 203), Guo et al.
(2003: 90–91).
Illustrations: Vassiljevsky & Karakulin (1937: 212, ig. 19),
Ellis (1971: 280, ig. 192 B), Guo et al. (2003: 91, ig. 57).
Exsiccatae: Barthol., Fungi Columb. 2685. Clements &
Clements, Crypt. Format. Colorad. 505. W.B. Cooke,
Mycobiota N. Amer. 445b. Ellis & Everh., Fungi Columb.
991, 1980, 2169, 2170. Ellis & Everh., N. Amer. Fungi 1988,
2600. Erikss., Fungi Paras. Scand. Exs. 186B. Fuckel,
Fungi Rhen Exs. 130. Kabát & Bubák, Fungi Imperf. Exs.
94, 442. Kellerm., Ohio Fungi 97. Krieger, Fungi Saxon.
Exs. 939, 1938–1940. Kunze, Fungi Sel. Exs. 395. Petr.,
Fl. Bohem. Morav. Exs. Pilze 498. Petr., Mycoth. Gen. 300.
Poelt & Scheuer, Reliqu. Petrak. 1594. Rabenh., Fung.
Eur. Exs. 4200. Roum., Fungi Sel. Gall. Exs. 740. D. Sacc.,
Mycoth. Ital. 1739. Săvul., Herb. Mycol. Rom. 1346. Siem.,
Fungi Bialowiez. Exs. 199. Solh., Mycol. Saximont. Exs.
200, 500, 1197, 1199. Syd., Mycoth. Germ. 288, 848. Syd.,
Mycoth. March. 4888. Triebel, Microf. Exs. 169. Verstergr.,
Micromyc. Rar. Sel. Praec. Scand. 149. Winter, Fungi
Helvet. 95. Zahlbruckner, Krypt. Exs. 1191.
76
Description: Leaf spots lacking or indeinite, i.e. colonies on
necrotic or faded leaves, or with lesions of variable shape
and size, mostly oblong, at irst 2–12 × 1–4 mm, later
forming long necrotic streaks, inally large leaf segments
or almost entire leaves necrotic, yellowish, ochraceous,
straw-coloured, pale brown to dark brown, greyish brown,
dingy grey, sometimes with diffuse yellowish halo. Caespituli
usually hypophyllous, scattered, punctiform, dark brown to
blackish. Mycelium internal. Stromata variable in shape and
size, but usually well-developed, substomatal to immersed,
20–130 µm diam, brown, cells circular to somewhat angularirregular in outline, 3–12(–15) µm diam. Conidiophores in
small to very large fascicles, arising from stromata, through
stomata or erumpent, divergent to dense, erect, straight,
cylindrical, subcylindrical to somewhat curved, sinuous,
slightly geniculate, unbranched, apex usually obtuse,
sometimes hooked, 20–105 × 3–8 µm, 0–4-septate, individual
conidiophores pale to medium brown, medium to dark brown
in mass, thin-walled, smooth to somewhat rough-walled;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, 15–60 µm long, with a
single to several conspicuous conidiogenous cells, circular in
outline, 2.5–4 µm diam, somewhat thickened and darkened,
usually barely protuberant. Conidia formed singly, ellipsoidovoid, obovoid, short obclavate, (15–)20–50(–60) × (5–)6–
12(–14) µm, 0–3-septate, mostly 1-septate, occasionally
slightly constricted at the septa, subhyaline to pale brownish,
thin-walled, smooth to somewhat rough-walled, apex obtuse,
often broadly rounded, base rounded to short obconically
truncate, sometimes somewhat peg-like, 3–5 µm wide, hila
somewhat thickened and darkened.
Lectotype (designated here, MycoBank, MBT200464):
germany: Rheinland-Pfalz: Mt. Rabenkopf, on grass leaves
(exact identity unclear), Fuckel [Fungi Rhen. Exs. 130] (HAL).
Isolectotypes: Fuckel, Fungi Rhen. Exs. 130 (e.g. FH, G).
Host range and distribution: On ×Agrohordeum macounii,
Agropyron cristatum, ×Agrositanion sp., Agrostis (capillaris,
castellana, exarata, gigantea, hallii, oregonensis, rossiae,
scabra,
stolonifera
[palustris],
tenuis),
Alopecurus
(aequalis, alpinus, carolinianus, geniculatus, pratensis,
textilis [tilisiensis]), Ammophila (arenaria, Ammophila sp.),
Anthoxanthum odoratum, Arctagrostis (latifolia, Arctagrostis
sp.), Arrhenatherum (album [erianthum], elatius subsp. elatius,
elatius subsp. bulbosum [Avena bulbosa]), ?Arundinaria
(gigantea subsp. gigantea, gigantea subsp. tecta [tecta],
Arundinaria sp.), Avena sativa, Beckmannia (eruciformis,
syzigachne, Beckmannia sp.), Bromus (anomalus, carinatus,
catharticus [willdenowii], ciliatus, frondosus, hordeaceus
[mollis], inermis, japonicus, kalmii [purgans], laevipes,
marginalis [breviaristatus], orcuttianus, paciicus, secalinus,
sitchensis, vulgaris), Calamagrostis (canadensis, inexpansa),
Cenchrus (purpureus [Pennisetum purpureum], Cenchrus
sp.), Cinna (arundinacea, latifolia, Cinna sp.), Cynodon
(dactylon, Cynodon sp.), Cynosurus (cristatus, echinatus),
Dactylis (glomerata subsp. glomerata, glomerata subsp.
hispanica [hispanica]), Danthonia (californica, intermedia,
Danthonia sp.), Deschampsia (atropurpurea, cespitosa,
danthonioides, elongata, Deschampsia sp.), Digitaria
IMA FUNGUS
cercosporoid fungi 3
Notes: Passalora graminis is widespread on a wide range of
grasses. Deighton (1967) considered P. graminis an aggregate
species possibly composed of several taxa. Collections on
various hosts, however, are morphologically uniform. The
biology of this species is not well known. It often occurs on
VOLUME 6 · NO. 1
necrotic or faded leaves and is sometimes considered a
parasite of weakened grasses. Inoculation experiments and
results of molecular sequence analyses are necessary to
elucidate the biology and taxonomy of this species. In the
interim, we prefer to maintain Passalora graminis in its current
wide sense (s. lat.). Records of P. graminis on Arundinaria spp.
are doubtful and possibly refer to P. compactum. Collections of
P. graminis on Milium effusum may belong to P. milii.
Data on asexual/sexual morph connections of
Scolicotrichum graminis are confusing and unsubstantiated.
Scolicotrichum graminis is often listed as the asexual morph and
synonym of Mycosphaerella recutita (Fr.) Johanson 1884 (see
Index Fungorum database), which goes back to Fuckel (1870:
107) who cited this species as a “conidial form” of Sphaerella
recutita (Fr.) Rabenh. (syn. Sphaeria recutita Fr. 1823), which
was conirmed by Cooke (1871: 921). Arx (1949: 67) provided
a comprehensive description of M. recutita. According to
Eriksson (1992), the type material of Sphaeria recutita does not
contain a Mycosphaerella, i.e. the common application of this
name for a Mycosphaerella on grasses must be considered
erroneous. The oldest valid name for M. recutita sensu Arx is
not Mycosphaerella wichuriana (J. Schröt.) Johanson 1884,
as suggested by Eriksson (1992), but Sphaerella disseminata
De Not. & Carestia 1871 (i.e. Mycosphaerella disseminata
(De Not. & Carestia) Tomilin 1967), which was re-combined
as Davidiella disseminata (De Not. & Carestia) Aptroot 2006
(Aptroot 2006: 80). Davidiella is a synonym of Cladosporium,
i.e. the asexual morphs are entirely cladosporioid with catenate
conidia and a speciic coronate type of conidiogenous loci and
conidial hila. A genetic connection between Passalora graminis
and Davidiella disseminata (M. recutita sensu Arx) is doubtful.
Detailed examinations of the life-cycle of P. graminis, cultures
and molecular examinations are necessary to elucidate the
true biology and life cycle of this species.
The fungal herbarium of G.H. Otth was originally
deposited at BERN. Some years ago, the fungus
collections of BERN were tranferred to ZT. Type material of
Passalora hordei is now preserved in ZT, but syntypes of P.
punctiformis, described from Dactylis glomerata and Melica
unilora, have not been traced. There is a single specimen
on Arrhenatherum elatius (Bern, G.H. Otth) deposited at ZT
under the name P. punctiformis. It is unclear if this collection
represents type material of this species. The reference to
“Dactylis and Melica” in the protologue indicates uncertainty
in the identiication of the host species.
ART I CLE
(eriantha [smutsii], sanguinalis, Digitaria sp.), ×Elyhordeum
stebbinsianum [Elymus aristatus], Elymus (albicans
[Agropyron grifithii], angulatus [antarcticus], canadensis
[robustus], caninus, enysii, elymoides [Sitanion hystrix],
glaucus, ×hansenii [Sitanion ×hansenii], hispidus [Agropyron
trichophorum], lanceolatus [Agropyron dasystachyum,
A. riparium], macrourus [Agropyron sericeum], mollis,
multisetum [Sitanion jubatum], repens, ×saxicola, sibiricus,
smithii, spicatus, trachycaulus [Agropyron latiglume,
richardsonii, subsecundum], vancouverensis, virginicus),
×Elysitanion sp., Eragrostis (secundilora, Eragrostis sp.),
Festuca (arundinacea [elatior], kingii, nigrescens, rubra,
subulata, varia), Glyceria (borealis, canadensis, elata,
luitans, grandis, leptostachya, maxima, notata, remota,
septentrionalis, Glyceria sp.), Helictotrichon canescens
[Trisetum canescens], Hierochloe (redolens, Hierochloe sp.),
Homalotrichon pubescens [Avenula pubescens], Hordeum
(brachyantherum, bulbosum [nodosum], jubatum, pusillum,
vulgare), Hystrix (patula, Hystrix sp.), Koeleria (micranthera
[cristata], Koeleria sp.), Leersia (oryzoides, Leersia sp.),
Leucopoa sp., Leymus (condensatus [Elymus condensatus],
mollis [Elymus mollis], triticoides [Elymus triticoides]),
Lolium (multilorum, perenne, Lolium sp.), Melica (bulbosa,
geyeri, smithii, spectabilis, subulata, Melica sp.), ?Milium
(effusum, Milium sp.), Miscanthus (sinensis, Miscanthus
sp.), Muhlenbergia (iliformis, mexicana, racemosa,
sylvatica, Muhlenbergia sp.), Nassella viridula [Stipa
viridula], Oryzopsis hymenoides, Oryzopsis sp.), Panicum
(antidotale, Panicum sp.), Phalaris arundinacea, Phleum
(alpinum, pratense), Phragmites (australis, sp.), Poa (alpinia,
annua, arida, chaixii, compressa, cusickii [epilis], foliosa,
interior, juncifolia [ampla, nevadensis], longiligula, nemoralis,
nervosa, palustris, pratensis, remota [Glyceria remota],
secunda, stenantha, trivialis), Pseudosclerochloa rupestris
[Poa canbyi, gracillima, sandbergii], Puccinellia (distans,
Puccinellia sp.), Secale (cereale, montana), Spartina gracilis,
Stenotaphrum sp., Stipa (comata, coronata, lemmonii,
lettermannii, nelsonii [williamsii], occidentalis [californica,
columbiana, elmeri], pinetorum, speciosa, thurberiana, Stipa
sp.), Torreyochloa paucilora [Glyceria paucilora], Trisetum
(lavescens, montanum, spicatum), Triticum (aestivum,
Triticum sp.), Zea mays, Poaceae, widespread, Africa (Canary
Islands, South Africa), Asia (China, Iran, Japan, Taiwan,
Thailand), Australia, Caucasus (Armenia), Europe (Belgium,
Finland, France, Germany, Italy, Poland, Russia, UK,
Ukraine), New Zealand, North America (Canada, Manitoba,
Ontario; USA, Alaska, Arizona, California, Colorado, Idaho,
Illinois, Kansas, Kentucky, Main, Maryland, Massachusetts,
Michigan, Minnesota, Mississippi, Montana, Nebraska, New
Hampshire, New Mexico, New York, Nevada, North Dakota,
Ohio, Oregon, South Carolina, South Dakota, Texas, Utah,
Vermont, Virginia, Washington, Wisconsin, Wyoming), South
America (Argentina, Chile, Colombia), and West Indies
(Cuba, Puerto Rico, Virgin Islands).
Passalora imperatae (Syd. & P. Syd.) U. Braun &
Crous, in Crous & Braun, Mycosphaerella and
Anam.: 225 (2003).
(Fig. 49)
Basionym: Cercosporina imperatae Syd. & P. Syd., Ann.
Mycol. 14: 372 (1916).
Synonyms: Cercospora imperatae (Syd. & P. Syd.)
Vassiljevsky, in Vassiljevsky & Karakulin, Fungi imperfecti
parasitici. 1. Hyphomycetes: 270 (1937).
Cercospora imperatae (Syd. & P. Syd.) Sawada, Taiwan
Agric. Rev. 38: 697 (1942).
Mycovellosiella imperatae (Syd. & P. Syd.) Goh & W.H.
Hsieh, in Hsieh & Goh, Cercospora and similar fungi from
Taiwan: 139 (1990).
77
Braun et al.
ART I CLE
obclavate-cylindrical, straight or almost so, 30–65 × 4–6 µm,
1–7-septate, colourless, thin-walled, smooth, apex obtuse,
base rounded to obconically truncate, about 2 µm wide, hila
thickened and darkened.
Holotype: Philippines: Los Baños, on Imperata cylindrica, 17
Sep. 1913, M. B. Raimundo 1717 (S-F20471). Isotype: CUP
40054.
Host range and distribution: On Imperata cylindrica
[arundinacea], Poaceae (Panicoideae, Andropogoneae),
Asia (China, India, Philippines, Taiwan).
Notes: The generic afinity of this species is unclear. The
colourless conidia support placement in Cercospora s. str.,
but the mycovellosiella-like habit with solitary conidiophores
arising from supericial hyphae are in conlict. We prefer to
retain this species in Passalora.
Passalora koepkei (W. Krüger) U. Braun & Crous, in
Crous & Braun, Mycosphaerella and Anam.: 238
(2003).
(Fig. 50)
Fig. 49. Passalora imperatae (S-F20471). A. Solitary conidiophores.
B. Conidia. Bar = 10 µm.
Literature: Chupp (1954: 247), Vasudeva (1963: 123), Hsieh
& Goh (1990: 139), Crous & Braun (2003: 225), Guo et al.
(2003: 31–32), Kamal (2010: 124).
Illustrations: Hsieh & Goh (1990: 139, ig. 106), Guo et al.
(2003: 32, ig. 17).
Description: Leaf spots amphigenous, subcircular to
elliptical, 2–10 mm diam, often conluent, yellowish to
pale brown, margin indeinite. Caespituli amphigenous,
mainly hypophyllous, unevenly scattered, diffuse, dark.
Mycelium internal and external; supericial hyphae sparingly
branched, septate, about 1–3.5 µm wide, pigmented, pale,
thin-walled, smooth. Stromata absent. Conidiophores
solitary, arising from supericial hyphae, lateral, occasionally
arising from hyphal aggregations, in loose groups, to 4,
subfasciculate, erect, straight, subcylindrical to somewhat
curved or geniculate-sinuous, unbranched, 20–75 × 6–8 µm,
0–3-septate, pale to medium brown, paler towards the tip,
thin-walled, smooth; conidiogenous cells integrated, terminal
or conidiophores reduced to conidiogenous cells, about
10–35 µm long, conidiogenous loci conspicuous, thickened
and darkened, 1–2 µm diam. Conidia solitary, cylindrical to
78
Basionym: Cercospora koepkei W. Krüger, Ber. Versuchsstat.
Zuckerrohr W.-Java, Kagok-Tegal 1: 115 (1890).
Synonyms: Mycovellosiella koepkei (W. Krüger) Deighton,
Mycol. Pap. 144: 20 (1979).
Pseudocercospora miscanthi Katsuki, J. Jap. Bot. 31:
372 (1956) [lectotype (designated here, MycoBank,
MBT200465): Japan: Pref. Kagoshima: Mikata-mura,
Amami Island, on Miscanthus sinensis, 6 Oct. 1954, S.
Katsuki (CUP 41022). Isolectotype: K(M) IMI 68966)].
Cercospora koepkei var. sorghi K. Goto, K. Hirano & Fukatsu,
Ann. Phytopathol. Soc. Japan 27: 52 (1962) [syntypes:
Japan: Chiba, Sep. 1940, on Sorghum bicolor [Holcus
sorghum var. japonicus]; Saitama, 1955, on S. bicolor
(not traced)].
Literature: Saccardo (1892: 656), Matsumoto & Yamamoto
(1934: 591–594), Chupp (1954: 248), Sun (1955: 163),
Vasudeva (1963: 656), Katsuki (1965: 34–35, 76), Mulder
& Holliday (1974a), Ellis (1976: 262), Deighton (1979: 20),
Hsieh & Goh (1990: 140), Crous & Braun (1996: 280; 2003:
238), Guo et al. (2003: 33–34).
Illustrations: Krüger (1890: plates VI, VIII B). Matsumoto
& Yamamoto (1934: 592, ig. 3), Sun (1955: 164, ig. 21),
Mulder & Holliday (1974a, ig., unnumbered). Ellis (1976:
263, ig. 199B), Hsieh & Goh (1990: 140, ig. 107), Guo et al.
(2003: 33, ig. 18).
Description: Leaf spots irregularly shaped to almost elliptical,
size variable, at irst small, later oblong, to 5 mm in length,
at irst yellowish, later reddish to purplish brown, leaves
with heavy infections inally often becoming straw-coloured,
necrotic. Caespituli amphigenous, mainly hypophyllous, forming
mould-like colonies or covers. Mycelium internal and external;
supericial hyphae emerging through stomata, sparingly
branched, 2–3 µm wide, pale olivaceous, thin-walled, smooth.
Stromata almost lacking, only with a few swollen hyphal cells or
IMA FUNGUS
cercosporoid fungi 3
ART I CLE
Kenya, Mauritius, Reunion, Sierra Leone, Somalia, South
Africa, Tanzania, Zimbabwe, Uganda), Asia (Brunei,
Cambodia, China, India, Indonesia, Japan, Malaysia,
Myanmar, Nepal, Pakistan, Papua New Guinea, Philippines,
Sabah, Sri Lanka, Taiwan, Thailand), Australia, Central and
South America (Brazil, Colombia, Costa Rica, El Salvador,
Guatemala, Guyana, Honduras, Panama, Suriname,
Venezuela), North America (Mexico; USA, Alabama,
Florida, Louisiana), Oceania (American Samoa, Fiji, French
Polynesia, Guam, Hawaii, Micronesia, New Caledonia,
Palau, Samoa, Solomon Islands, Tonga, Vanuatu), and West
Indies (Cuba, Dominican Republic, Puerto Rico, Trinidad and
Tobago, Virgin Islands).
Notes: The status of Cercospora koepkei var. sorghi is not
clear. Re-examination of the type material and additional
collections are necessary, but type material was not traced.
This taxon is tentatively treated as a synonym of P. koepkei.
A neotypiication of P. koepkei is not proposed since suitable
material from Java has not yet been found.
Passalora maculicola (Ellis & Kellerm.) U. Braun,
Schlechtendalia 5: 39 (2000).
(Fig. 51)
Basionym: Scolicotrichum maculicola Ellis & Kellerm., J.
Mycol. 3: 103 (1887); as “maculicolum”.
Synonym: Fusicladium maculicola (Ellis & Kellerm.) Ondřej,
Česká Mykol. 25: 337 (1971).
Literature: Saccardo (1892: 601), Schubert et al. (2003: 117),
Crous & Braun (2003: 458–459).
Fig. 50. Passalora koepkei (based on Hsieh & Goh 1990: 140, ig.
107). A. Conidiophore fascicle. B. Solitary conidiophores arising from
a supericial hypha. c. Conidiophore tip. d. Conidia. Bar = 10 µm.
small, 10–20 µm diam, brown. Conidiophores in small, mostly
loose fascicles, 2–15, arising from internal hyphae or stromata,
through stomata, and solitary, arising from supericial hyphae,
lateral, rarely terminal, erect, straight, subcylindrical to distinctly
geniculate-sinuous, unbranched, 30–200 × 4–7 µm, aseptate
to pluriseptate, pale to medium olivaceous-brown, thin-walled,
smooth; conidiogenous cells integrated, terminal or intercalary,
occasionally conidiophores reduced to conidiogenous cells,
about 10–40 µm long, with a single to several conspicuous
conidiogenous loci, thickened and darkened, 1.5–2 µm diam.
Conidia solitary, fusiform-obclavate, mostly straight, occasionally
somewhat curved, 20–65 × 4–6.5 µm, 1–7-septate, often
3-septate, without constrictions, colourless or almost so to pale
olivaceous, thin-walled, smooth, apex obtuse to subacute, base
short obconically truncate, 1.5–2.5 µm wide, hila thickened and
darkened.
Syntypes: Indonesia: Java: on Saccharum spp., W. Krüger
(probably not preserved).
Host range and distribution: On Miscanthus (loridulus
[japonicus], sinensis), Saccharum (oficinarum, robustum
[edule],
spontaneum),
Sorghum
bicolor,
Poaceae
(Panicoideae, Andropogoneae), Africa (Gabon, Ghana,
VOLUME 6 · NO. 1
Illustrations: Ondřej (1971: 238, igs 1–2), Braun (2000: 37,
ig. 6).
Exsiccatae: Ellis & Everh., N. Amer. Fungi 1989, 2789.
Kellerman & Swingle, Kansas Fungi 20. Rabenh., Fungi Eur.
Exs. 3800. Roum., Fungi Sel. Gall. Exs. 5580.
Description: Leaf spots amphigenous, oblong, fusiform,
ellipsoid, 3–15 × 1–2(–3) mm, yellowish, ochraceous,
dingy brownish, margin narrow, dull medium to dark brown.
Caespituli amphigenous, mainly hypophyllous, punctiform,
mostly dense, dark brown to blackish. Mycelium internal.
Stromata small to well-developed, substomatal, globose to
oblong, 10–60 µm diam, brown, composed of swollen hyphal
cells, subcircular to somewhat irregular in outline, about 3–7
µm diam. Conidiophores in small to fairly large, loose to dense
fascicles, arising from stromata, through stomata, erect,
straight and subcylindrical to usually moderately to strongly
geniculate-sinuous, unbranched or only rarely branched,
(20–)30–80(–100) × (3–)4–7(–8) µm, continuous to septate,
mostly sparingly septate, pale to medium brown or olivaceousbrown throughout or paler towards the tip, wall thin to slightly
thickened, smooth, occasionally somewhat verruculose;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, 10–50 µm long, with a single
to several conspicuous conidiogenous loci, non-protuberant,
truncate, slightly thickened and somewhat darkened, 1–2
µm diam, non-coronate (i.e., not cladosporioid). Conidia
79
Braun et al.
ART I CLE
USA, Iowa, Kansas, North Dakota, Nebraska, Oklahoma,
Oregon, South Dakota, Washington, Wisconsin).
Notes: North American records of this species on Arundo
donax are unproven and unclear. The generic afinity of P.
maculicola is intricate and can only be veriied with certainty
by using cultures and results from molecular sequence
analyses. The general habit of this species is cladosporioid
and reminiscent of heterosporium-like Cladosporium species
(Bensch et al. 2012), but the conidiogenous loci and conidial
hila are not coronate (not cladosporioid) but truncate and
somewhat thickened and darkende, i.e. cercospora-like. Due
to verruculose conidia, P. maculicola is also comparable with
Asperisporium, which is possibly a synonym of Passalora s.
lat. (Braun et al. 2013). The conidia in Asperisporium spp.
are usually formed singly. At present this species is best
maintained in Passalora. Deightoniella roumeguerei (Cavara)
Constant. 1983 (syn. Scolicotrichum roumeguerei Cavara
1890, as “roumegueri”) is another species on Phragmites,
which is distinct and not congeneric.
Passalora milii (Syd.) G.A. de Vries, Contrib.
Knowledge of the Genus Cladosporium Link ex
Fries: 94 (1952).
(Fig. 52)
Basionym: Cladosporium milii Syd., Ann. Mycol. 12: 538
(1914).
Literature: Saccardo (1931: 792), Crous & Braun (2003: 460),
Schubert (2005b: 213), Bensch et al. (2013: 321).
Exsiccatae: Syd., Mycoth. Germ. 1295, 1296.
Fig. 51. Passalora maculicola (NY 830653, lectotype). A.
Conidiophore fascicles. B. Conidiophores. c. Conidia. Bar = 10 µm.
solitary or in short, simple or rarely branched chains, broadly
ellipsoid-ovoid, obovoid, subfusiform, rarely subcylindrical
or subglobose, 11–23 × 5–11 µm, aseptate, subhyaline,
pale yellowish green to olivaceous, verruculose, apex
broadly rounded in solitary conidia, or attenuated-truncate in
catenate ones, bases rounded, subtruncate, truncate or short
obconically truncate, 1–2.5 µm wide, hila slightly thickened
and darkened.
Lectotype (designated by Braun 2000): usA: Kansas:
Manhattan, on Phragmites australis, 1 Jun. 1887, W. A.
Kellerman 934 (NY 830653). Isolectotypes: BPI 425144, NY
266874, 266875, 830656–830660.
Host range and distribution: On Phragmites australis,
Poaceae (Arundinoideae), North America (Canada, Ontario;
80
Description: Leaf spots on living and faded leaves, indistinct
or linear, about 1 mm wide, length variable, pale, margin
indeinite. Caespituli hypophyllous, scattered, punctiform,
dark brown to blackish. Mycelium internal. Stromata
substomatal to intraepidermal, applanate to oblong, to 60 ×
20 µm, brown, composed of swollen hyphal cells, subcircular
to irregular in outline, 3–8 µm diam. Conidiophores in
moderately large to very large fascicles, moderately dense,
arising from stromata, through stomata or erumpent, erect,
straight, subcylindrical to somewhat curved-sinuous,
usually not geniculate, unbranched, 40–75 × 5–8 µm,
aseptate or 1–2-septate, septa more or less near the base,
brownish, thin-walled, smooth; conidiophores reduced to
conidiogenous cells or integrated, terminal, to 60 µm long,
tips often curved, with a single or several conspicuous
conidiogenous loci near the tip, circular in outline, 2–3 µm
diam, slightly thickened and darkened. Conidia solitary,
ellipsoid-ovoid, broadly obovoid or short cylindrical, 12–38
× 5–10.5 µm, 0–2-septate, not constricted, subhyaline, pale
olivaceous, olivaceous to pale greyish brown, thin-walled,
verruculose, apex broadly rounded, base subtruncate
to short obconically truncate, 2.5–3 µm wide, hila slightly
thickened and darkened.
Lectotype (designated here, MycoBank, MBT200466):
France: Lorraine: Forbach (“Wald am Öttinger Tälchen”), on
Milium effusum, 22 Jun. 1913, A. Ludwig (HBG). Isolectotypes:
IMA FUNGUS
cercosporoid fungi 3
ART I CLE
Fig. 53. Passalora paspalicola (W-1978-07621, lectotype). A.
Conidiophore fascicle. B. Conidiophores. c. Conidia. Bar = 10 µm.
Passalora paspalicola (Petr. & Cif.) U. Braun, Fungal
Diversity 8: 56 (2001).
(Fig. 53)
Basionym: Cercospora paspalicola Petr. & Cif., Ann. Mycol.
30: 226 (1932).
Literature: Chupp (1954: 250), Crous & Braun (2003: 308).
Fig. 52. Passalora milii (HBG, lectotype). A. Conidiophore fascicle.
B. Conidiophores. c. Conidia. Bar = 10 µm.
Illustration: Braun (2001: 53, ig. 12).
Exsiccatae: Cif., Mycol. Doming. Exs. 331.
BPI 427262, F 1093516. Former syntypes: Syd., Mycoth.
Germ. 1295, e.g. CUP, M, MICH 15428; Syd., Mycoth. Germ.
1296, e.g. BPI 427263, CUP, M, MICH 15429).
Host range and distribution: On Milium effusum, Poaceae
(Pooideae, Aveneae), Europe (France, Germany, Latvia,
Russia).
Notes: Several collections from Germany (B700006628,
700006630–70006636, 70006482) and two samples from
Latvia (B700006627, 700006629) have been examined.
Based on the verruculose conidia, this species is reminiscent
of Asperisporium.
VOLUME 6 · NO. 1
Description: Leaf spots indistinct, later irregular, brown
discolorations, usually 1–3 mm diam. Caespituli
amphigenous, punctiform, subcircular to oblong in outline,
blackish. Mycelium internal. Stromata immersed, large,
30–350 µm diam, dark brown. Conidiophores numerous,
densely fasciculate, forming well-developed sporodochial
conidiomata, conidiophores little differentiated, reduced
to conidiogenous cells, only developed as somewhat
elongated peripherical cells of the stromata, subcylindricalconical, 5–15 × 3–6 µm (sometimes with persistent conidia
resembling longer conidiophores), aseptate, brownish, thinwalled, smooth, conidiogenous loci conspicuous, slightly
thickened and darkened, 1–1.5 µm diam. Conidia solitary
81
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Braun et al.
Fig. 54. Passalora phalaridis (NY 1042610, holotype). A. Conidiophore fascicles. B. Conidia. Bar = 10 µm.
to catenate, occasionally in branched chains, cylindrical,
obclavate-subcylindrical, ellipsoid-ovoid, 15–60 × 4–6.5
µm, 1–4-septate, subhyaline to pale olivaceous, thin-walled,
smooth, apex obtuse, base rounded to obconically truncate,
1–1.5 µm wide, hila slightly thickened and darkened.
Lectotype (designated by Braun 2001): dominical republic:
Prov. Azua: Valle de San Juan, San Juan de la Maguana,
on living leaves of Paspalum clavuliferum, Poaceae
(Panicoideae, Paniceae), 22 Aug. 1929, E. L. Ekman [Cif.,
Mycol. Doming. Exs. 331] (W-1978-07621). Isolectotypes:
BPI 845246, CUP (Ciferri, M.Dom. 0331), ILL 33101, MICH
15348. Topotypes: CUP 40470 (Aug. 1926), NY 937111,
937112 (1 Aug. 1929).
Host range and distribution: Only known from the type
collections.
Passalora phalaridis K. Schub. & U. Braun, Nova
Hedwigia 84: 197 (2007).
(Fig. 54)
Illustration: Schubert & Braun (2007: 198, ig. 5).
82
Description: Leaf spots amphigenous, elliptical to oblong,
to 10 mm long and 1–2 mm wide, scattered, but often
aggregated, occasionally conluent, pale brown, surrounded
by a narrow brown margin, surrounding leaf tissue often
discoloured, brownish to somewhat reddish brown, often
somewhat paler below. Caespituli usually hypophyllous,
occasionally epiphyllous, scattered to effuse, often in lines,
at irst covered by the white detached cuticle, later erumpent,
loose to dense, caespitose, pale brown, velvety. Mycelium
internal, subcuticular to intraepidermal; hyphae sparingly
branched, 3–5 µm wide, septate, pale yellowish brown,
smooth, wall only slightly thickened. Stromata substomatal,
dense, several layers deep, composed of swollen hyphal
cells, subglobose to angular, pale yellowish brown, smooth,
walls slightly thickened. Conidiophores loosely to densely
fasciculate, arising from stromata, emerging through stomata,
erect, straight to somewhat lexuous, cylindrical, sometimes
geniculate towards the apex, unbranched, 35–80 × 4–5.5(–6)
µm, (0–)1–4-septate, subhyaline to pale brown, smooth, wall
thin or almost so; conidiogenous cells integrated, terminal,
rarely intercalary, cylindrical, 20–50 µm long, sympodial,
with a single or several conspicuous conidiogenous loci,
somewhat crowded near the apex, protuberant, truncate, (1–
)1.5–2.5(–3) µm diam, thickened and somewhat darkenedIMA FUNGUS
cercosporoid fungi 3
ART I CLE
refractive. Conidia catenate, in unbranched chains, broadly
ellipsoid-fusiform, 12–23 × 5–10.5 µm, aseptate, almost
hyaline to pale olivaceous, smooth or almost so, walls
unthickened, attenuated towards the apex and base, hila
protuberant, truncate, 1–2.5 µm diam., somewhat thickened
and darkened-refractive.
Holotype: usA: New York: Genese Co., Bergen Swamp,
on Phalaris arundinacea, Poaceae (Pooideae, Aveneae),
19 Jul. 1946, W. C. Muenscher and C. T. Rogerson (NY
1042610), originally deposited as Cladosporium velutinum
Ellis & Tracy.
Host range and distribution: Only known from the type
collection.
Note: Resembling Cercospora barretoana, but conidia much
shorter, broader, aseptate and at least slightly pigmented.
Passalora ramularioides (Sacc. & Fautrey) U. Braun,
Schlechtendalia 5: 40 (2000).
(Fig. 55)
Basionym: Scolicotrichum ramularioides Sacc. & Fautrey,
Bull. Soc. Mycol. France 16: 24 (1900).
Literature: Saccardo (1902: 1057), Vassiljevsky & Karakulin
(1937: 213), Crous & Braun (2003: 467), Braun & Crous
(2005: 413).
Illustration: Braun (2000: 41, ig. 9).
Description: Leaf spots amphigenous, fusiform, elliptical,
1–5 × 0.5–1.5 mm, centre pale, yellowish to ochraceous,
later greyish white, margin narrow, dark. Caespituli
amphigenous, inely punctiform, effuse to dense, dark
brown. Mycelium internal. Stromata almost absent or small,
10–25 µm diam, brown, substomatal to intraepidermal,
cells globose to somewhat angular-irregular in outline,
2–8 µm diam, walls somewhat thickened. Conidiophores
solitary or in small fascicles, loose to dense, arising from
stromata, through stomata or erumpent, erect, straight,
subcylindrical to geniculate-sinuous, unbranched, 25–100
× 3–7 µm, septate, pale to medium brown throughout or
apex somewhat paler, walls somewhat thickened, smooth;
conidiogenous cells integrated, terminal, 20–50 µm long,
conidiogenous loci somewhat thickened and darkened,
1.5–2 µm diam. Conidia solitary, rarely in short chains,
subcylindrical, subfusiform, ellipsoid-ovoid, 15–30 × 3–6
µm, (0–)1-septate, subhyaline to pale yellowish, ochraceous
or olivaceous-brown, thin-walled, smooth, apex obtuse,
rounded, base short obconically truncate, 1–2 µm wide, hila
slighty thickened and darkened.
Holotype: France: Côte-d’Or, on Leersia oryzoides, F.
Fautrey 23 (PAD).
Host range and distribution: On Leersia (oryzoides, Leersia
sp.), Poaceae (Ehrhartoideae, Ehrharteae), Europe (France,
Ukraine), North America (USA, Iowa).
Fig. 55. Passalora ramularioides (PAD, holotype). A. Conidiophore
fascicle. B. Conidiophores. c. Conidia. Bar = 10 µm.
Passalora tungurahuensis (Petr.) U. Braun & Crous,
in Crous & Braun, Mycosphaerella and Anam.: 413
(2003).
(Fig. 56)
Basionym: Cercospora tungurahuensis Petr., Sydowia 4: 574
(1950).
Literature: Chupp (1954: 255).
VOLUME 6 · NO. 1
83
Braun et al.
ART I CLE
short obconically truncate, 1.5–2 µm wide, hila somewhat
thickened and darkened.
Lectotype (designated here, MycoBank, MBT200467):
ecuador: Tungurahua: Baños, Hacienda San Antonio, on
Cenchrus bambusiformis [Pennisetum bambusiforme], 6
Dec. 1937, H. Sydow [hb. Petrak 32194] (W-1974-0003437).
Isolectotype: B 700016007.
Host range and distribution: On Cenchrus bambusiformis
[Pennisetum bambusiforme], Poaceae (Panicoideae,
Paniceae), South America (Brazil, Ecuador).
Passalora vaginae (W. Krüger) U. Braun & Crous, in
Crous & Braun, Mycosphaerella and Anam.: 417
(2003).
(Fig. 57)
Basionym: Cercospora vaginae W. Krüger, Meded. Proefstn.
Suikerriet W. Java, Kagok-Tegal 3: 29 (1896).
Synonym: Mycovellosiella vaginae (W. Krüger) Deighton,
Mycol. Pap. 144: 26 (1979).
Literature: Saccardo (1899: 1106), Chupp (1954: 256), Sun
(1955: 168), Vasudeva (1963: 208), Abbott (1964: 49–50),
Katsuki (1965: 34), Kirk (1973), Ellis (1976: 262), Sivanesan
& Waller (1986: 49–50), Hsieh & Goh (1990: 141), Guo et al.
(2003: 34–35).
Illustrations: Sun (1955: 169, ig. 25), Ellis (1976: 263, ig.
199B), Hsieh & Goh (1990: 142, ig. 108), Guo et al. (2003:
35, ig. 19).
Fig. 56. Passalora tungurahuensis (W-1974-0003437, lectotype). A.
Conidiophore fascicle. B. Conidiophore tips. c. Conidia. Bar = 10
µm.
Description: Leaf spots amphigenous, large, large leaf
segments or later almost entire leaves discoloured, greyish
brown, margin indeinite or with a diffuse yellowish halo.
Caespituli epiphyllous, punctiform, scattered, dark brown to
blackish. Mycelium internal. Stromata immersed, large, 30–
100 µm diam, dark brown, composed of swollen hyphal cells,
circular to somewhat angular-irregular in outline, about 3–6
µm diam. Conidiophores in large fascicles, loose to mostly
dense, arising from stromata, erumpent, erect, straight,
subcylindrical to somewhat geniculate-sinuous, unbranched,
80–160 × 4–7.5 µm, pluriseptate throughout, pale to medium
dark brown, tips paler, wall somewhat thickened, smooth;
conidiogenous cells integrated, terminal, occasionally
intercalary, 10–40 µm long, with a single to several
conspicuous conidiogenous loci, somewhat thickened and
darkened, 1.5–2 µm diam. Conidia solitary or in short chains,
ellipsoid-ovoid, subcylindrical, almost obclavate, fusiform,
15–60 × 3.5–7 µm, (0–)1–4(–5)-septate, hyaline, subhyaline
to pale greenish olivaceous, thin-walled, smooth, apex
obtuse to short conically truncate in catenate conidia, base
84
Description: Spots mainly on sheaths, sometimes also
formed as leaf spots, at irst small, subcircular to elliptical,
red, margin conspicuous, spots later conluent or increasing,
to about 15 mm diam, on leaves dark reddish above, indistinct
below. Caespituli amphigenous, effuse, dark greyish brown,
velvety, mostly in the centre of the lesion. Mycelium internal
and external; supericial hyphae sparingly branched, septate,
pale, thin-walled, smooth. Stromata sometimes developed,
substomatal, 10–75 µm diam, dark brown, but without
conidiophore fascicles. Conidiophores solitary, arising
from supericial hyphae, lateral, at the top of mother cells,
occasionally terminal, i.e. at the end of procumbent hyphae,
erect to ascending, straight to curved, subcylindrical, conical
to geniculate-sinuous, simple or sometimes branched,
occasionally entangled, 20–200 × 3–5 µm, 1–5-septate, pale
olivaceous-brown to darker brown, paler towards the tip, thinwalled, smooth; conidiogenous cells integrated, terminal,
with conspicuous conidiogenous loci, about 1–1.5 µm diam.
Conidia solitary, cylindrical or obclavate-cylindrical, straight
to somewhat curved, 15–55 × 3–6.5 µm, 0–5-septate,
occasionally slightly constricted at the septa, hyaline to
olivaceous, thin-walled, smooth, apex obtuse, base short
obconically truncate, 1–2 µm wide, somewhat thickened and
darkened.
Holotype: Indonesia: Java: on Saccharum oficinarum
(details not recorded, probably not preserved).
IMA FUNGUS
cercosporoid fungi 3
excluded
and
insuficiently
known
Mycovellosiella oryzae (Deighton & D. Shaw) Deighton,
Mycol. Pap. 144: 25 (1979).
Basionym: Ramularia oryzae Deighton & D. Shaw, Trans.
Brit. Mycol. Soc. 43: 516 (1960).
ART I CLE
Doubtful,
species
Literature: Braun (1998: 201), Crous & Braun (2003: 485).
Illustrations: Braun (1998: 203, ig. 470).
Holotype: New guinea: Madang, on Oryza sativa, Poaceae,
15 Mar. 1958, D. Shaw (K(M) IMI) 73536).
Mycovellosiella paspali Deighton, Mycol. Pap. 144: 24
(1979).
Synonym: Ramularia paspali (Deighton) U. Braun, Nova
Hedwigia 50: 513 (1990).
Literature: Braun (1998: 201), Crous & Braun (2003: 485).
Illustrations: Deighton (1979: 24, ig. 12), Braun (1998: 203,
ig. 471).
Holotype: trinidad: Botany Island, I.C.T.A., on Paspalum
sp., Poaceae, 8 Apr. 1960, C. L. A. Leakey (K(M) IMI)
86339a).
Mycovellosiella sacchari Sarbajna, see Pseudocercospora sacchari.
Fig. 57. Passalora vaginae (based on Hsieh & Goh 1990: 142, ig.
108). A. Solitary conidiophores arising from supericial hyphae.
Conidiophore fascicle. B. Conidiophore. c. Conidia. Bars = 10 µm.
Mycovellosiella taiwanensis (T. Matsumoto & W.
Yamam.) X.J. Liu & Y.L. Guo, see Pseudocercospora
taiwanansis.
Passalora aterrima Bres., Ann. Mycol. 18: 57 (1920).
Host range and distribution: On Saccharum (oficinarum,
spontaneum), Poaceae (Panicoideae, Anthropogoneae),
Africa (Ghana, Madagascar, Malawi, Mauritius, Mozambique,
Senegal, Sierra Leone, South Africa, Togo, Zimbabwe), Asia
(Afghanistan, China, India, Indonesia, Japan, Malaysia,
Pilippines, Taiwan, Thailand, Vietnam), Central and South
America (Brazil, Costa Rica, El Salvador, Guatemana,
Guyana, Honduras, Panama, Peru, Venezuela), North
America (Mexico; USA, Florida, Georgia, Louisiana, Texas),
Oceania (Hawaii), and West Indies (Barbados, Cuba,
Dominican Republ., Haiti, Jamaica, Puerto Rico, Trinidad and
Tobago, Virgin Island).
Notes: Chupp (1954) referred to “Ber. Vers. Stat. Zuckerr.
West Java 1: 64 (1890)” as the place and date of publication
of the basionym. Type material of this species is probably
not preserved, but a neotypiication is postponed since
appropriate material from Java has not yet been found.
VOLUME 6 · NO. 1
Literature: Saccardo (1931: 799), Crous & Braun (2003: 477),
Guo et al. (2003: 89–90).
Illustration: Guo et al. (2003: 89, ig. 57).
Description: Colonies densely gregarious, velutinous,
blackish. Hyphae 5–6 µm wide. Conidiophores solitary, erect,
straight, cylindrical-iliform, 380–500 × 5–6 µm, septate,
pigmented, apex obtuse. Conidia solitary, ellipsoid, 14–19 ×
8–10 µm, 1-septate, brown, apex broadly rounded [according
to Guo et al. (2003), conidiophores 140–700 × 4–5 µm, and
conidia 15–18 × 8–10 µm].
Holotype: Brazil: Rio Grande do Sul: São Leopoldo, on
hymenium (rarely stalk) of Thelephora sp., on bamboo,
Poaceae, Rick (not traced).
Notes: This species, known from the type collection in
Brazil and on Bambusa sp. in China (Guo et al. 2003), is
undoubtedly not congeneric with Passalora in the current
sense, but its generic afinity is unclear.
85
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Braun et al.
Passalora bambusicola (Sawada) Poonam Srivast., J.
Living World 1: 113 (1994), nom. inval. (ICN, Art.
39.1).
Type: Not clearly indicated, there are cultures under IMI
173304 and in the “Indian Type Culture Collection” and a
dried culture in “Herbarium, New Delhi University, Dept. of
Botany”.
Basionym: Cercosporidium bambusicola Sawada, Taiwan
Agric. Res. Inst. Rep. 87: 77 (1944), nom. inval. (Art.
39.1); as “bambusicolum”.
Synonym: Pseudospiropes bambusicola Goh & W.H.
Hsieh, in Hsieh & Goh, Cercospora and similar fungi from
Taiwan: 147 (1990).
Notes: This species was isolated from green leaves of
Triticum. The material in IMI (now K) has been examined and
the taxonomic afinity of P. graminicola was unclear, but this
species is not cercosporoid.
Holotype: taiwan: Taipei, on Bambusa sp., Poaceae, 6 Mar.
1913, Y. Fujikuro (NTU-PPE, herb. Sawada).
Phaeoramularia kellermaniana Marasas & I.H. Bredell,
Bothalia 11: 217 (1974).
Passalora eragrostidis Viégas, Bragantia 6: 386 (1946).
Literature: Crous & Braun (2003: 452).
Illustration: Viégas (1946: plate 26).
Description: Leaf spots hypophyllous, oblong, 5–10 mm, 1–2
mm wide, between veins, brown. Colonies effuse. Mycelium
internal and external; supericial hyphae septate, hyaline.
Conidiophores solitary, arising from supericial hyphae, erect,
straight to curved-sinuous, unbranched, 50–180 × 4–4.5 µm,
pluriseptate, brown below, subhyaline above; conidiogenous
cells integrated, terminal, barely geniculate, but with obtuse,
truncate denticles. Conidia solitary, ellipsoid, 10–16 × 6–7
µm, 1-septate, at irst hyaline, later brown, thin-walled,
asperulate, ends more or less rounded.
Holotype: Brazil: Matto Grosso: Jupiá, Rio Paraná, on
Eragrostis ciliaris, Poaceae, 20 Apr. 1943, R. O. Botero (not
traced).
Notes: Type material of this species has not been examined.
The afinity of P. eragrostidis is unclear, but it does not belong
to the complex of cercosporoid fungi. Viégas (1946) described
asperulate conidia and illustrated supericial mycelium, so
that this species could also be a member of Cladosporium
(subgen. Heterosporium).
Phaeoramularia graminicola Mukerji & Khanna, in
Mukerji et al., Bibl. Mycol. 91: 291 (1983).
Synonyms: Cladosporium gossypii Jacz., Holopkovoe Delo
1929 (5–6): 567 (1929), non Alternaria gossypii (Jacz.) Y.
Nisik., K. Kimura & Miyaw., 1940.
Cladosporium malorum Rühle, Phytopathology 21: 1146
(1931).
Cladosporium porophorum Matsush., Icones Microfungorum
a Matsushima lectorum: 36 (1975).
Cladophialophora kellermaniana (Marasas & I.H. Bredell) U.
Braun & Feiler, Microbiol. Res. 150: 83 (1995).
Pseudocladosporium kellermanianum (Marasas & I.H.
Bredell) U. Braun, A monograph of Cercosporella,
Ramularia and allied genera 2: 393 (1998).
Alternaria malorum (Rühle) U. Braun, Crous & Dugan,
Mycol. Progr. 2: 5 (2003).
Chalastospora gossypii (Jacz.) U. Braun & Crous, Persoonia
22: 144 (2009).
Holotype: south Africa: Cape Prov.: Kopgat, Calvinia,
isolated from wheat straw, Triticum aestivum, Poaceae, Feb.
1972, W. F. O. Marasas OP-76 (PREM 44703). Isotype: K(M)
IMI 165252; ex-type culture: CBS 266.75.
Notes: This is a saprobic species which was placed in the
genus Chalastospora E.G. Simmons, Pleosporales (Crous
et al. 2009). In a new phylogenetic/taxonomic concept of
Alternaria s. lat., recently introduced by Woudenberg et
al. (2013), Chalastospora was reduced to synonymy with
Alternaria and treated as section of this genus. Based on this
concept, Alternaria malorum is the nomenclaturally correct
name of this species. Cladosporium gossypii is the oldest
valid name for this species, but the epithet “gossypii” is preoccupied in Alternaria so cannot be taken up.
Literature: Crous & Braun (2003: 481).
Pseudocercospora
Key to Pseudocercospora species on Poaceae
86
1
Mycelium in vivo internal; stromata large, to 230 µm diam; conidia (23–)30–38.5(–42.5) × 5.8–7.7 µm,
4–7-septate; on Bambusa tulda ............................................................................................................ P. bambusae
Mycelium in vivo internal and external, supericial, with solitary conidiophores arising from
supericial hyphae; stromata not developed; on other hosts .................................................................................... 2
2 (1)
Leaf spots absent; conidia cylindrical-iliform, 20–90 × 2–3 µm, pale olivaceous-brown;
on Saccharum .................................................................................................................................. P. whalianensis
Leaf spots present, distinct; conidia at least partly obclavate or acicular,
and/or hyaline or subhyaline .................................................................................................................................... 3
IMA FUNGUS
cercosporoid fungi 3
Conidia pigmented, at least pale olivaceous-brown; on Cymbopogon or Saccharum .................................................... 4
Conidia hyaline or subhyaline; on Saccharum ................................................................................................................ 5
4 (3)
Conidiophores narrow, 4–65 × 1–3.5 µm; conidia narrowly obclavate-cylindrical, 15–80 × 1–3.5 µm;
on Saccharum .......................................................................................................................................... P. sacchari
Conidiophores somewhat wider, 16–55 × 3–4 µm; conidia obclavate-cylindrical, 25–95 × 3–4 µm;
on Cymbopogon ............................................................................................................................ P. cymbopogonis
5 (3)
Older leaf spots subcircular to irregular, 0.5–12 mm wide, reddish to dark purple,
most distinct on the upper leaf surface (referred to as “Purple spot of sugarcane”);
conidiophores short, 12–35 µm long, 1–3-septate ........................................................................ P. rubropurpurea
Leaf spots different, characteristic purple spots not formed; conidiophores longer, 10–156 µm long,
0–7-septate .............................................................................................................................................................. 6
6 (5)
Lesions developed as characteristic black stripes; conidia long, obclavate-iliform,
14–212 × 2–4.5 µm, 1–22-septate ................................................................................................... P. atroiliformis
Lesions different, black stripes not formed, leaf spots either developed as ring spots
or as elliptical to oblong patches, brown, reddish or straw-coloured ....................................................................... 7
7 (6)
Lesions developed as characteristic ring spots (circular to irregular, 4–21 × 1.5–7.5 µm, or conluent,
to 35 mm diam, brown, often with a reddish brown to dark reddish brown narrow border,
inally with a grey centre surrounded by a purplish brown margin);
conidia 36–127 × 2–3.5 µm .............................................................................................................. P. saccharicola
Lesions narrowly elliptical to oblong, 2–10 × 1–1.5 mm; conidia longer, 20–275 × 2–4 µm ..................... P. taiwanensis
ART I CLE
3 (2)
Tabular key to Passalora species on Poaceae
Bambusa ................................................................................................................................................................. P. bambusae
Cymbopogon .................................................................................................................................................. P. cymbopogonis
Saccharum
1
Leaf spots lacking; conidia cylindrical-iliform, 20–90 × 2–3 µm, pale olivaceous-brown ....................... P. whalianensis
Leaf spots developed, distinct; conidia at least partly obclavate or acicular,
and/or hyaline or subhyaline .................................................................................................................................... 2
2 (1)
Conidia pigmented, pale olivaceous-brown .................................................................................................... P. sacchari
Conidia hyaline or subhyaline ......................................................................................................................................... 3
3 (2)
Older leaf spots subcircular to irregular, 0.5–12 mm wide, reddish to dark purple,
most distinct on the upper leaf surface (referred to as “Purple spot of sugarcane”);
conidiophores short, 12–35 µm, 1–3-septate ............................................................................... P. rubropurpurea
Leaf spots different, characteristic purple spots not formed; conidiophores longer,
10–156 µm, 0–7-septate .......................................................................................................................................... 4
4 (3)
Lesions developed as characteristic black stripes; conidia long, obclavate-iliform,
14–212 × 2–4.5 µm, 1–22-septate ................................................................................................... P. atroiliformis
Lesions different, black stripes not formed, leaf spots either developed as ring spots
or as elliptical to oblong patches, brown, reddish or straw-coloured ....................................................................... 5
5 (4)
Lesions developed as characteristic ring spots (circular to irregular, 4–21 × 1.5–7.5 µm, or conluent,
to 35 mm diam, brown, often with a reddish brown to dark reddish brown narrow border,
inally with a grey centre surrounded by a purplish brown margin);
conidia 36–127 × 2–3.5 µm .............................................................................................................. P. saccharicola
Lesions narrowly elliptical to oblong, 2–10 × 1–1.5 mm; conidia longer, 20–275 × 2–4 µm ..................... P. taiwanensis
VOLUME 6 · NO. 1
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Illustration: Yen et al. (1953: 4, ig. 1), Sun (1955: 162, ig.21),
Guo et al. (1998: 383, ig. 315).
Description: Leaf spots amphigenous, at irst yellow, ovoid
to rounded, later developing into narrow dark brown to black
streaks, 5–36 mm long and 0.5–1.2 mm wide, between
veins (referred to as “Black stripe of sugarcane”). Caespituli
amphigenous, mostly hypophyllous. Mycelium internal
and external, supericial. Stromata lacking. Conidiophores
solitary, arising from supericial hyphae, lateral, at the top
of mother cells, erect, subcylindrical, narrowed towards the
tip, geniculate-tortuous, unbranched, 20–80 × 3–4.5 µm,
2–6(–9)-septate, dark olivaceous-brown, paler towards the
tip, thin-walled, smooth; conidiogenous cells integrated,
terminal, conidiogenous loci inconspicuous, unthickened,
not darkened. Conidia solitary, narrowly obclavate-iliform,
slighty to strongly curved, 14–212 × 2–4.5 µm, 1–22-septate,
hyaline or subhyaline, hila neither thickened nor darkened.
Syntypes: taiwan: Taichung, on Saccharum oficinarum, 8
Dec. 1952, W. Y. Yen; Puli, on S. oficinarum, 19 Jan. 1953,
W. Y. Yen; Hualian, on S. oficinarum, 19 Jan. 1953, W. Y.
Yen (not traced).
Host range and distribution: On Saccharum oficinarum,
Poaceae (Panicoideae, Andropogoneae), Asia (China,
Taiwan).
Pseudocercospora bambusae Saika & A.K. Sarbhoy,
Indian Phytopathol. 38: 432 “1985” (1986).
(Fig. 59)
Illustration: Saika & Sarbhoy (1986: 433, ig. 1).
Fig. 58. Pseudocercospora atroiliformis (based on Sun 1955: 162,
ig. 21). A. Solitary conidiophores arising from supericial hyphae. B.
Conidia. Bar = 10 µm.
List of Pseudocercospora species on Poaceae
Pseudocercospora atroiliformis (W.Y. Yen, T.C. Lo
& C.C. Chi) J.M. Yen, Bull. Trimestriel Soc. Mycol.
France 97: 152 (1981).
(Fig. 58)
Basionym: Cercospora atroiliformis W.Y. Yen, T.C. Lo & C.C.
Chi, J. Sugarcane Res. Taiwan 7: 15 (1953).
Literature: Sun (1955: 162), Abbott (1964: 21–23), Sivanesan
& Waller (1986: 51), Hsieh & Goh (1990: 144), Guo et al.
(1998: 383), Crous & Braun (2003: 71).
88
Description: Caespituli hypophyllous, at irst scattered,
punctiform, later effuse, velvety, forming circular, subcircular to
elliptical colonies, brown to dark brown. Mycelium immersed;
hyphae branched, septate, about 2.5–4 µm wide, subhyaline to
pale brown. Stromata epidermal to subepidermal, subcircular
to lattended, to 230 µm diam, pigmented. Conidiophores
numerous, arising from stromatic hyphal aggregations, erect,
divergent, almost straight below, lexuous to geniculate
above, unbranched, about 87.5–180 µm long, 3–4.7 µm wide
below and 4.7–6.3 µm wide above at the subclavate apex,
4–7-septate, pale to medium brown, thin-walled, smooth;
conidiogenous cells integrated, terminal, conidiogenous loci
inconspicuous. Conidia solitary, broadly obclavate-fusiform,
straight to slightly curved, about (23–)30–38.5(–42.5) × 5.8–
7.7 µm, 4–7-septate, pale to medium brown, thin-walled,
smooth, apex obtuse, base short obconically truncate, about
1.5–2.5 µm wide, hila unthickened, not darkened.
Holotype: India: Assam: on Bambusa tulda, Poaceae
(Bambusoideae), 3 May 1978, U. N. Saikia (HCIO 32704).
Host range and distribution: Only known from the type
collection.
Notes: Pseudocercosporella bambusae Deighton 1973 is a
different cercosporoid fungus on Bambusa spp., distinguished
IMA FUNGUS
cercosporoid fungi 3
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Fig. 59. Pseudocercospora bambusae (based on Saika & Sarbhoy
1986: 433, ig. 1). A. Conidiophore fascicle. B. Conidiophore tip. c.
Conidia. Bar = 10 µm.
in the short, colourless conidiophores and colourless conidia.
Hsieh & Goh (1990: 147) re-examined type material of
Cercosporella dendrocalami Sawada 1944 (nom. inval., Art.
39.1) and reduced it to synonymy with Deighton’s species.
Pseudocercospora cymbopogonis (J.M. Yen) J.M.
Yen, Bull. Trimestriel Soc. Mycol. France 94: 386
“1978” (1979).
(Fig. 60)
Basionym: Cercospora cymbopogonis J.M. Yen, Bull.
Trimestriel Soc. Mycol. France 93: 148 (1977).
VOLUME 6 · NO. 1
Fig. 60. Pseudocercospora cymbopogonis (based on Yen 1977: 149,
ig. 2). A. Supericial hyphae emerging through a stoma. B. Solitary
conidiophores arising from supericial hyphae. c. Conidiophore. d.
Conidia. Bar = 10 µm.
Literature: Hsieh & Goh (1990: 145), Guo et al. (1998: 384),
Crous & Braun (2003: 150).
Illustrations: Yen (1977: 149, ig. 2), Guo et al. (1998: 384,
ig. 316).
Description: Leaf spots scattered or conluent, brown, margin
indeinite. Caespituli amphigenous or only hypophyllous.
Mycelium internal and external; supericial hyhae emerging
through stomata, branched, septate, pale olivaceous-brown,
2–3 µm wide. Stromata lacking. Conidiophores solitary,
arising from supericlal hyphae, lateral, at the top of mother
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cells, erect, straight to tortuous-geniculate, unbranched,
16–55 × 3–4 µm, 1–6-septate, olivaceous-brown, apex
rounded to truncate, thin-walled, smooth; conidiogenous
cells integrated, terminal, conidiogenous loci inconspicuous.
Conidia solitary, obclavate-cylindrical, somewhat curvedsinuous, 25–95 × 3–4 µm, 4–11-septate, pale olivaceousbrown, thin-walled, smooth, apex rounded, base slightly short
obconically truncate, hila unthickened, not darkened.
Holotype: taiwan: Taichung, garden, on Cymbopogon sp.,
Poaceae (Panicoideae, Andropogoneae), 29 Oct. 1971, J. M.
Yen 71258 (not traced).
Host range and distribution: Only known from the type
collection.
Pseudocercospora rubropurpurea (S.H. Sun) J.M.
Yen, Bull. Trimestriel Soc. Mycol. France 97: 154
(1981).
(Fig. 61)
Basionym: Cercospora rubropurpurea S.H. Sun, J. Agric.
Forest. (Taichung) 4: 182 (1955).
Literature: Sivanesan & Waller (1986: 40–42), Hsieh & Goh
(1990: 145), Guo et al. (1998: 385), Crous & Braun (2003:
360).
Illustration: Sun (1955: 165, ig. 22), Guo et al. (1998: 385,
ig. 317).
Description: Leaf spots at irst only visible as indistinct
yellowish stripes, later subcircular to irregular, 0.5–12 mm
wide, reddish to dark purple, most distinct on the upper leaf
surface (referred to as “Purple spot of sugarcane”). Caespituli
amphigenous, mainly epiphyllous. Mycelium internal and
external, supericial. Stromata lacking. Conidiophores
solitary, arising from supericial hyphae, lateral, at the top
of mother cells, erect, straight to slightly curved, geniculate,
unbranched, 12–35 × 3.5–4 µm, 1–3-septate, olivaceousbrown, thin-walled, smooth; conidiogenous cells integrated,
terminal, conidiogenous loci inconspicuous. Conidia solitary,
obclavate, cylindrical, acicular, straight to slightly curved,
45–108 × 3–3.5 µm, 3–7(–9)-septate, hyaline, thin-walled,
smooth, apex subacute or subobtuse, base subtruncate to
long obconically truncate, hila unthickened, not darkened.
Holoype: taiwan: Taichung, on Saccharum oficinarum, 9
Aug. 1955, S. H. Sun (not traced).
Host range and distribution: On Saccharum oficinarum,
Poaceae (Panicoideae, Andropogoneae), Asia (China,
Taiwan).
Pseudocercospora sacchari K. Bhalla & A.K.
Sarbhoy, Indian Phytopathol. 53: 265 (2000), nom.
nov. (as “(Sarbajna) K. Bhalla & A.K. Sarbhoy,
comb. nov.”).
(Fig. 62)
Basionym: Mycovellosiella sacchari Sarbajna, J. Mycopathol.
Res. 28: 162 (1990), nom. inval. (Art. 37.5).
90
Fig. 61. Pseudocercospora rubropurpurea (based on Sun 1955: 165,
ig. 22). A. Solitary conidiophores arising from supericial hyphae. B.
Conidia. Bar = 10 µm.
Synonym: Pseudocercospora sacchari U. Braun & Crous, in
Crous & Braun, Mycosphaerella and Anam.: 488 (2003),
nom. illegit. (Art. 52.1)
Literature: Braun & Crous (2007: 66), Kamal (2010: 216).
Illustration: Sarbajna (1990: 161, ig. 1), Bhalla & Sarbhoy
(2000: 264, ig. 4).
IMA FUNGUS
cercosporoid fungi 3
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ropes. Stromata lacking or almost so. Conidiophores solitary,
arising from supericial hyphae, lateral, at the top of mother
cells, erect, straight, subcylindrical to strongly curved,
sinuous, geniculate, unbranched or branched, 4–65 × 1–3.5
µm, 0–6-septate, pale olivaceous-brown, thin-walled, smooth;
conidiogenous cells integrated, terminal or conidiophores
sometimes reduced to conidiogenous cells, about 10–25 µm
long, conidiogenous loci inconspicuous or visible by being
denticle-like, but always unthickened and not darkened.
Conidia solitary, obclavate-cylindrical (often with short lateral
branchlets, sometimes giving rise to secondary conidia),
15–80 × 1–3.5 µm, 2–13-septate, often slightly constricted
at septa, pale olivaceous-brown, thin-walled, smooth, apex
obtuse or subobtuse, base short obconically truncate, about
1.5–2 µm wide, hila unthickened, not darkened.
Holotype: India: West Bengal: Baduria, on Saccharum
oficinarum, 18 Oct. 1986, K. K. Sarbajna (K(M) IMI 311125).
Host range and distribution: On Saccharum oficinarum,
Poaceae (Panicoideae, Andropogoneae), India (West
Bengal).
Pseudocercospora saccharicola (S.H. Sun) J.M.
Yen, Bull. Trimestriel Soc. Mycol. France 97: 154
(1981).
(Fig. 63)
Basionym: Cercospora saccharicola S.H. Sun, J. Agric.
Forest. Taiwan 4: 183 (1955).
Literature: Sutton & Waller (1986: 51), Hsieh & Goh (1990:
146), Guo et al. (1998: 395), Crous & Braun (2003: 362).
Illustrations: Yen et al. (1953: 5, ig. 2), Sun (1955: 167, ig.
23), Guo et al. (1998: 386, ig. 318).
Fig. 62. Pseudocercospora sacchari (K(M) IM)I 311125, holotype).
A. Supericial hyphae emerging through a stoma. B. Conidiophores
arising from supericial hyphae. c. Conidiophores. d. Conidia. Bar
= 10 µm.
Description: Leaf spots amphigenous, circular, elliptical to eyeshaped, with greyish centre, surrounded by a reddish brown
margin, 3–11 mm diam, sometimes conluent. Caespituli
amphigenous, mainly hypophyllous, deep olivaceous,
somewhat loccose. Mycelium internal and external; supericial
hyphae emerging through stomata, branched, septate, pale
brownish, 1–2.5 µm wide, sometimes intertwined, forming
VOLUME 6 · NO. 1
Description: Leaf spots at irst small, circular to irregular, dark
green to yellowish, later larger and subcircular to irregular,
4–21 × 1.5–7.5 mm or conluent and larger, to 35 mm diam,
brown, often with a reddish brown to dark reddish brown narrow
border, inally with a grey centre surrounded by a purplish
brown margin (referred to as “Ring spot of sugarcane”).
Caespituli hypophyllous. Mycelium internal and external.
Stromata lacking. Conidiophores solitary, arising from
supericial hyphae, lateral, at the top of mother cells, erect,
distinctly geniculate, simple, rarely branched, 34–126 × 3–3.5
µm, 2–4-septate, olivaceous-brown, thin-walled, smooth;
conidiogenous cells integrated, terminal, conidiogenous loci
inconspicuous. Conidia solitary, obclavate, acicular, straight
to slightly curved, 36–127 × 2–3.5 µm, 3–9-septate, hyaline,
thin-walled, smooth, apex subacute, base subtruncate, hila
neither thickened nor darkened.
Syntypes: taiwan: Taichung, on Saccharum oficinarum, 10
Aug. 1955, S. H. Sun; Taichung, Nantu, Puli, Wufeng, on S.
oficinarum, 1935, W. Y. Yen (not traced).
Host range and distribution: On Saccharum oficinarum,
Poaceae (Panicoideae, Andropogoneae), Asia (China,
Taiwan).
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Fig. 64. Pseudocercospora taiwanensis (CUP 41374, lectotype).
A. Supericial hyphae. B. Solitary conidiophores arising from a
supericial hypha. c. Conidia. Bar = 10 µm.
Illustrations: Matsumoto & Yamamoto (1934: 585, ig. 1 and
pl. 3 [upper left and right]), Sun (1955: 168, ig. 24), Sivanesan
(1976: ig., unnumbered).
Fig. 63. Pseudocercospora saccharicola (based on Sun 1955: 167,
ig. 23). A. Solitary conidiophores arising from supericial hyphae. B.
Conidia. Bar = 10 µm.
Pseudocercospora taiwanensis (T. Matsumoto &
W. Yamam.) J.M. Yen, Bull. Trimestriel Soc. Mycol.
France 97: 154 (1981).
(Fig. 64)
Basionym: Cercospora taiwanensis T. Matsumoto & W.
Yamam., J. Soc. Trop. Agric. Taiwan 6: 590 (1934).
Synonym: Mycovellosiella taiwanensis (T. Matsumoto & W.
Yamam.) X.J. Liu & Y.L. Guo, Mycosystema 1: 262 (1988).
Literature: Chupp (1954: 255), Sun (1955: 167), Abbott
(1964: 33–36), Katsuki (1965: 74), Sivanesan (1976; 1984:
480), Sivanesan & Waller (1986: 51–52), Hsieh & Goh (1990:
146), Crous & Braun (2003: 396).
92
Description: Leaf spots elliptical to oblong, 2–10 × 1–1.5
mm, yellowish to reddish brown, centre later straw-coloured.
Caespituli amphigenous. Mycelium internal and external,
supericial; hyphae branched, septate, 1–3.5 µm wide,
subhyaline to pale olivaceous or brownish, thin-walled,
smooth. Stromata lacking. Conidiophores solitary, arising
from supericial hyphae, lateral, at the top of mother cells,
erect, subcylindrical to geniculate, somewhat attenuated
towards the tip, unbranched, 5–100(–155) × 2.5–4.5 µm,
0–7-septate, subhyaline, pale olivaceous, yellowish brown to
olivaceous brown, paler towards the tip, thin-walled, smooth;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, conidiogenous loci
inconspicuous, neither thickened nor darkened. Conidia
solitary, narrowly obclavate to acicular, straight, curved
to somewhat sinuous, 20–275 × 2–4 µm, indistinctly
1–15-septate, hyaline or subhyaline, thin-walled, smooth,
apex subacute, base subtruncate, hila neither thickened nor
darkened.
IMA FUNGUS
cercosporoid fungi 3
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Lectotype (designated here, MycoBank, MBT200468):
taiwan: Hualien Kang (Karenkô), on Saccharum oficinarum,
13 May 1934, W. Yamamoto (CUP 41374). Isolectotypes: BPI
441849, 441850.
Host range and distribution: On Saccharum oficinarum
Poaceae (Panicoideae, Andropogoneae), Asia (China,
Japan, Taiwan).
Notes: Leptosphaeria taiwanensis W.Y. Yen & C.C. Chi (Yen
& Chi 1952) was originally proposed as sexual morph of
Cercospora taiwanensis, but Hsieh (1979) linked this species
to Stagonospora taiwanensis W.H. Hsieh and Phoma sp.
Eriksson & Hawksworth (2003) introduced the combination
Saccharicola taiwanensis (W.Y. Yen & C.C. Chi) Erikss. &
D. Hawksw. 2003 and emphasized further examination was
required to establish the asexual/sexual morph connexion
of S. saccharicola. Matsumoto & Yamamoto (1934) cited
“in foliis Sacchari oficinarum, Karenkô, Taiwan (Formosa)”
(without date and collector) as type material. They mentioned
that the fungus was brought to their attention in spring 1933
by I. Okamoto, but the material concerned was sterile. They
obtained a second sample in 1934 and recollected this fungus
in that year themtheves, i.e. all original samples collected at
Karenkô, including duplicates preserved at BPI and CUP, are
syntypes.
Pseudocercospora whalianensis (J.M. Yen &
S.K. Sun) U. Braun & Crous, in Crous & Braun,
Mycosphaerella and Anam.: 430 (2003).
(Fig. 65)
Basionym: Cercospora whalianensis J.M. Yen & S.K. Sun,
Mycotaxon 7: 394 (1978).
Synonym: Cercoseptoria whalianensis (J.M. Yen & S.K. Sun)
J.M. Yen, Bull. Trimestriel Soc. Mycol. France 97: 93
(1981).
Literature: Hsieh & Goh (1990: 133).
Illustration: Yen & Sun (1978: 395, ig. 1 A–C).
Description: Leaf spots lacking. Caespituli amphigenous.
Mycelium internal and external; supericial hyphae branched,
septate, 2–2.5 µm wide, pale olivaceous-brown, thin-walled,
smooth. Stromata lacking. Conidiophores solitary, arising
from supericial hyphae, lateral, at the top of mother cells,
erect, straight to curved, subcylindrical or once geniculate,
10–45 × 3–3.5 µm, 1–4-septate, pale olivaceous-brown,
thin-walled, smooth; conidiogenous cells integrated,
terminal, conidiogenous loci inconspicuous, unthickened,
not darkened. Conidia solitary, cylindrical-iliform, straight
to somewhat curved, 20–90 × 2–3 µm, 1–11-septate, pale
olivaceous-brown, thin-walled, smooth, apex rounded, base
subtruncate, hila unthickened, not darkened.
Holotype: taiwan: Whalian, Kungfu, on Saccharum
oficinarum, 12 Aug. 1977, S.K. Sun 114 (not traced).
Host range and distribution: On Saccharum oficinarum,
Poaceae (Panicoideae, Andropogoneae), Asia (Taiwan).
VOLUME 6 · NO. 1
Fig. 65. Pseudocercospora whalianensis (based on Yen & Sun 1978:
395, ig. 1 A–C). A. Solitary conidiophores arising from supericial
hyphae. B. Conidia. Bar = 10 µm.
Zasmidium
A single species.
Zasmidium dichanthii (S.A. Khan & Kamal) U. Braun
& Crous, Schlechtendalia 20: 100 (2010).
(Fig. 66)
Basionym Cercospora dichanthii S.A. Khan & Kamal,
Mycopathol. Mycol. Appl. 39: 200 (1969); as “dicanthii”.
Synonym: Stenella dichanthii (S.A. Khan & Kamal) U. Braun
& Crous, in Crous & Braun, Mycosphaerella and Anam.:
159 (2003).
Illustration: Khan & Kamal (1969: 198, ig. 2).
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Host range and distribution: Only known from the type
collection.
Notes: This species is a typical stenella-like fungus with
verruculose supericial hyphae and solitary conidiophores.
The solitary conidia are also verruculose. The conidigenous
loci are often inconspicuous, 1–1.5 µm wide, unthickened
and slightly darkened-refractive.
AcKNowledgeMeNts
We are very grateful to the directors and curators of B, BPI, C, CBS,
CUP, DAOM, HBG, K, NIAES, NTU, NY, NYS, PAD, PDD, PREM,
S, TNS, W and WIS for loaning type material and other collections
in their keeping during the course of our monographic studies
of cercosporoid fungi on true grasses. This study was partially
supported by JSPS KAKENHI Grant #24780149 & 24380079 and
IFO (Osaka) to C. Nakashima.
reFereNces
Fig. 66. Zasmidium dichanthi (K(M) IMI 104699, holotype). A.
A. Solitary conidiophores arising from supericial hyphae. B.
Conidiophore tip. c. Conidia. Bar = 10 µm.
Description: Leaf spots variable, often oblong and marginal,
dark reddish brown. Caespituli amphigenous, mainly
hypophyllous. Mycelium internal and external; supericial
hyphae sparingly branched, septate, 1–3 µm wide,
subhyaline or pale, thin-walled, verruculose. Stromata lacking
or almost so. Conidiophores solitary, arising from supericial
hyphae, lateral, erect, straight, subcylindrical to geniculatesinuous, unbranched, 20–60(–75) × 2–4 µm, 0–4-septate,
pale olivaceous to olivaceous-brown, thin-walled, smooth;
conidiogenous cells integrated, terminal or conidiophores
reduced to conidiogenous cells, about 10–25 µm long,
conidiogenous loci inconspicuous to somewhat conspicuous
by being slightly darkened-refractive, barely thickened, 1–1.5
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