New Species and New Records of Cercosporoids Hyphomycetes from Cuba and Venezuela (Part 2).

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Mycosphere Doi 10.5943/mycosphere/4/2/3
176
New species and new records of cercosporoid
hyphomycetes from Cuba and Venezuela (Part
2)
Braun U1* and Urtiaga R2
1Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik und Botanischer Garten, Herbarium, Neuwerk 21,
06099 Halle (Saale), Germany
2Apartado 546, Barquisimeto, Lara, Venezuela.
Braun U, Urtiaga R 2013 – New species and new records of cercosporoid hyphomycetes from Cuba
and Venezuela (Part 2). Mycosphere 4(2), 174–214, Doi 10.5943/mycosphere/4/2/3
Examination of specimens of cercosporoid leaf-spotting hyphomycetes made between 1966 and
1970 in Cuba and Venezuela, now housed at K (previously deposited at IMI as “Cercospora sp.”),
have been continued. Additionally examined Venezuelan collections, made between 2006 and 2010,
are now deposited at HAL. Several species are new to Cuba and Venezuela, some new host plants
are included, and the following new species and a new variety are introduced: Cercospororella
ambrosiae-artemisiifoliae, Passalora crotonis-gossypiifolii, P. solaniphila, P. stigmaphyllicola,
Pseudocercospora calycophylli, P. coremioides, P. lonchocarpicola, P. lonchocarpigena, P.
paulliniae, P. picramniae, P. psidii var. varians, P. solanacea, P. teramnicola, P. trichiliae-hirtae,
P. zuelaniae, Pseudocercosporella leonotidis, Zasmidium cubense, Z. genipae-americanae. The
new name Pseudocercospora toonae-ciliatae and the new combination Zasmidium hyptiantherae
are proposed.
Key words – Ascomycota – Cercospora – Cercosporella – Mycosphaerellaceae – Passalora –
Pseudocercospora – South America – West Indies – Zasmidium
Article Information
Received 6 November 2012
Accepted 8 February 2013
Published online 16 March 2013
*Corresponding author: U. Braun – e-mail – uwe.braun@botanik.uni-halle.de
Introduction
Cercosporoid fungi are anamorphic
ascomycetes [Ascomycota, Pezizomycotina,
Dothideomycetidae, Capnodiales, Mycosphae-
rellaceae (Schoch et al. 2006)] and represent
one of the largest and most diverse groups of
hyphomycetes, causing a wide range of
diseases of wild as well as numerous cultivated
plants. The second author of the this paper has
collected cercosporoid anamorphs in Cuba and
Venezuela since about 1966. Early collections
were deposited at IMI as Cercospora sp.
(recently completely transferred to K). These
specimens have recently been sent on loan to
the first author to be determined and for further
treatment. Venezuelan collections made bet-
ween about 1990 and 2012 (most of them since
2006) have been directly sent to the first author
and are now deposited at HAL. First results of
examinations of the samples concerned have
already been published (Braun & Urtiaga 2008,
Braun et al. 2010). Braun & Urtiaga (2012)
published results of examinations of further
collections from Cuba and Venezuela, which
are continued in the present paper. These
results also supplement first contributions to

Mycosphere Doi 10.5943/mycosphere/4/2/3
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the knowledge of cercosporoid fungi of Cuba
(Arnold 1986, Castañeda & Braun 1989, Braun
& Castañeda 1991, Vilaró et al. 2006) and
Venezuela (Chupp 1934, Pons 1984, 1988,
1993, 2004, 2007, Urtiaga 1986, García et al.
1996, Itturiaga & Minter 2006). Older data are
summarized in Crous & Braun (2003).
Methods
Sporulating structures were mounted in
distilled water without any staining, and
examined using oil immersion (bright field and
phase contrast), with standard light microscopy
(Olympus BX 50, Hamburg, Germany). Thirty
measurements ( 1000 magnification) of
conidia and other structures were made, with
the extremes given in parentheses. All
drawings have been prepared by the first
author.
Results and discussion
New records of cercosporoid hyphomy-
cetes from Cuba and Venezuela and descrip-
tions of new species and new varieties are
listed in alphabetical order by genus and
species. Discussion and comments are added to
each taxon.
Cercospora achyranthina Thirum. & Chupp
Material examined – VENEZUELA,
Lara, Barquisimeto, on leaves of Achyranthes
aspera var. indica L. [ A. indica (L.) Mill.]
(Amaranthaceae), Feb. 2008, R. Urtiaga 108
(HAL 2529 F).
Notes – New to Venezuela. This species
belongs to the C. apii s. lat. complex (Crous &
Braun 2003).
Cercospora apii Fresen. s. lat. (C. apii complex,
sensu Crous & Braun 2003)
Material examined – CUBA, Bayamo, on
leaves of Aloysia virgata (Ruiz & Pav.) Pers.
(Verbenaceae), 14 Sep. 1967, R. Urtiaga 918
(IMI 129468 = K(M) 176126); l.c., on leaves
of Atkinsia cubensis (Britton & P. Wilson) R.A.
Howard (Malvaceae), 13 Nov. 1966, R. Urtiaga
1018 (IMI 130165 = K(M) 176145); l.c., on
leaves of Eryngium foetidum L. (Apiaceae), 9
Dec. 1965, R. Urtiaga 5 (IMI 116898 = K(M)
176129); l.c., 21 Oct. 1966, R. Urtiaga (IMI
123279 = K(M) 176130); l.c., 9 Dec. 1966, R.
Urtiaga (IMI 124025 = K(M) 176131); l.c., 5
June 1967, R. Urtiaga A-604 (IMI 128002 =
K(M) 176128); l.c., 5 June 1967, R. Urtiaga B-
605 (IMI 128003 = K(M) 176127); l.c., on
leaves of Kallstroemia maxima (L.) Hook. &
Arn. (Zygophyllaceae), 4 Nov. 1966, R. Urtiaga
(IMI 123393 = K(M) 176120); l.c., on leaves
of Polypodium punctatum Thunb. (Polypo-
diaceae), 2 Jan. 1967, R. Urtiaga 1 (IMI
124326 = K(M) 176141); l.c., on dead stems of
Sesbania emerus (Aubl.) Urb. (Fabaceae), 6
Mar. 1967, R. Urtiaga (IMI 126239 = K(M)
176149); Bayamo-Ote, on leaves of Zantho-
xylum martinicense (Lam.) DC. (Rutaceae), 27
Apr. 1967, R. Urtiaga (IMI 127343 = K(M)
176136). VENEZUELA, Lara, Rio Claro, La
Cuchilla, on leaves of Scoparia sp. (Planta-
ginaceae), June 2010, R. Urtiaga 390 (HAL
2524 F).
Notes – All collections from Cuba have
been recorded by Urtiaga (1986) as Cercospora
sp. Aloysia virgata is listed as host of C. apii s.
lat. in Crous & Braun (2003). The Cuban
material is characterized by fasciclate coni-
diophores, up to 250 µm long, and hyaline
acicular conidia. Atkinsia cubensis, an endemic
Cuban plant, is cited as host of C. apii s. lat. by
Crous & Braun (2003). This collection is
characterized as follows: leaf spots brown,
greyish brown to dingy grey, shape and size
variable, up to 50 mm diam., margin indefinite;
caespituli hypophyllous, punctiform, dark
brown to blackish; conidiophores fasciculate,
divergent, 80–250  3–7 µm, pluriseptate,
cylindrical to slightly geniculate towards the
apex, medium brown throughout or usually
paler towards the tip, thin-walled, smooth;
conidiogenous cells integrated, terminal or
intercalary, 15–30 µm long, conidiogenous loci
thickened and darkened, 2.5–4 µm diam.;
conidia solitary, acicular, up to 150  3–5 µm,
hyaline, thin-walled, smooth, pluriseptate,
distance between septa usually 4–12 µm, base
truncate, hila thickened and darkened, 3–5 µm
wide. Kallstroemia maxima was cited as host
of Cercospora sp. in Arnold (1986). In this
collection the fasciculate brown conidiophores
are 30–140  3–6 µm and produce acicular
colourless conidia, 70–150  3–4.5 µm.
Polypodium punctatum and Zanthoxylum mar-
tinicense have not yet been listed as hosts of
cercosporoid fungi (Crous & Braun 2003). The
collection on Zanthoxylum martinicense is

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characterized as follows: leaf spots amphi-
genous, subcircular to angular-irregular, 1–8
mm diam., at first yellowish to ochraceous or
brownish, later greyish white, with narrow dark
margin; stromata 10–50 µm diam.; conidio-
phores fasciculate, 20–120  3–6 µm, usually
distinctly geniculate, olivaceous-brown or paler
towards the tip; conidiogenous loci 1.5–2.5 µm
diam.; conidia acicular, up to about 100 µm
long and 2–3 µm wide. Pavgi & Singh (1970)
described Cercospora oxyphylli Pavgi & U.P.
Singh from India on Zanthoxylum oxyphyllum
Edgew. The taxonomic affinity of this species
is unclear (type material not available), but due
to its conidia described as pigmented, and the
original illustration, this species seems rather to
belong to Pseudocercospora. The collection of
C. apii s. lat. on Scoparia does not belong to
the Indian C. scopariae Thirum. & Lacy, which
is characterized by having very short conidio-
phores, 6–15 µm, and narrowly obclavate
conidia (Chupp 1954, Vasudeva 1963). The
conidiophores in the material from Venezuela
are up to 120 µm long and the conidia are
acicular.
Cercospora bidentis Tharp
Material examined – VENEZUELA, La-
ra, Rio Claro, La Cuchilla, on leaves of Bidens
squarrosa Kunth (Asteraceae), Apr. 2009, R.
Urtiaga 200 (HAL 2555 F).
Notes – Known from Venezuela on
Bidens pilosa L. (Urtiaga 1986, Crous & Braun
2003, Iturriaga & Minter 2006), but on a new
host species.
Cercospora brachiata Ellis & Everh.
Material examined – VENEZUELA, La-
ra, Barquisimeto, Parque Macuto, on leaves of
Amaranthus viridis L. (Amaranthaceae), Oct.
2009, R. Urtiaga 294 (HAL 2535 F).
Notes – This is the second collection of
this species on A. viridis from Venezuela
(Braun & Urtiaga 2012). C. brachiata is known
from Venezuela on Amaranthus spinosus L.
and A. tricolor L. (= A. dubius Mart., nom.
inval.).
Cercospora colei Boedijn
Material examined – CUBA, Bayamo, on
leaves of Plectranthus scutellarioides (L.) R.
Br. [= Coleus blumei Benth.] (Lamiaceae), 6
Mar. 1967, R. Urtiaga (IMI 126249 = K(M)
176155).
Notes – Cercospora colei was listed from
Cuba in Crous & Braun (2003). This species is
a true Cercospora s. str. distinct from the C.
apii complex and well characterized by its
small stromata, 10–50 µm diam., short, fasci-
culate conidiophores, 10–60  2–7 µm, and
relatively short conidia, narrowly obclavate,
subcylindrical to short acicular, 25–75  2.5–5
µm. C. colei was described from Indonesia on
Coleus atropurpureus Benth. and C. hybridus
Voss (Boedijn 1961), but both names are now
considered to be synonyms of Plectranthus
scutellarioides. Braun (2001) re-examined type
material of this species and published an
illustration (Braun 2001: 422, Fig. 4). C.
coleicola Chupp & A.S. Mull. (Chupp 1954),
described from Brazil on Coleus sp. (type
material examined: CUP-MG 1109) is quite
distinct from C. colei and well characterized by
its lesions formed as dark to black spots on
stems, large pustulate stromata, 30–80 µm
diam. and acicular conidia. The identity of the
type host, Coleus sp., and its relation to
Plectranthus scutellarioides are unclear. C.
coleana J.M. Yen & Lim (Yen & Lim 1980) on
Coleus sp. in Singapore is also characterized by
the formation of acicular conidia, but stromata
are lacking or rudimentary and the coni-
diophores are formed in small fascicles. The
latter species is a typical member of the C. apii
complex. Two examined collections on Plec-
tranthus scutellarioides (as Coleus blumei)
coincide with C. coleana (Solomon Islands, 14
May 1978, E.H.C. McKenzie, GZU; Vanuatu,
Santo, 4 May 1983, E.H.C. McKenzie, PDD
44145). A second collection from Cuba on P.
scutellarioides (Bayamo, 6 Mar. 1967, R.
Urtiaga, IMI 126250 = K(M) 176154) belongs
possibly to C. coleana or it is a mixed infection
of the latter species and C. colei.
Key to Cercospora spp. on former Coleus spp.
(now Plectranthus):
1. Stromata medium-sized, 10–50 µm diam.;
conidiophores relatively short, 10–60  2–7
µm; conidia narrowly obclavate-subcylin-
drical, occasionally some conidia short
acicular, 25–75  2.5–5 µm ............ C. colei
1* Stromata either lacking or rudimentary or
very large, 30–80 µm diam.; conidiophores

Mycosphere Doi 10.5943/mycosphere/4/2/3
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much longer, up to about 250 µm; conidia
acicular ..................................................... 2
2. Lesions on stems, dark to blackish; stromata
very large, 30–80 µm diam. .... C. coleicola
2* Lesions formed as small, at first brown,
later greyish white leaf spots; stromata
lacking, rudimentary or small, 10–25 µm
diam. .......................................... C. coleana
Cercospora conyzoides Thirum. & Govindu
Fig. 1
Material examined – VENEZUELA, La-
ra, Sanare, Sabana Redonda Arriba, on leaves
of Ageratum conyzoides L. (Asteraceae), Sep.
2010, R. Urtiaga 391 (HAL 2528 F).
Fig. 1 – Cercospora conyzoides. Based on
HAL 2528 F. a Conidiophore fascicle. b
Conidiophore tips. c Conidia. – Bar = 10 µm.
Notes – This species, hitherto only
known from the original description based on
Indian material (Thirumalachar & Govindu
1957), is new to Venezuela. Ageratum cony-
zoides is native in tropical America and an
invasive weed in different parts of the world,
including Southeast Asia. The following
description is based on the material from
Venezuela:
Leaf spots at first rather indistinct,
greyish green to olivaceous discolorations, later
forming distinct leaf spots, subcircular to
angular-irregular, yellowish brown to brown,
1–10 mm diam., margin indefinite. Caespituli
amphigenous, not very conspicuous. Mycelium
internal; stromata lacking, only with a few
swollen hyphal cells, substomatal, subcircular
in outline, brown, 2–6 µm diam. Conidiophores
solitary or in small, divergent fascicles, 2–5,
erect, straight to usually distinctly pluri-
geniculate-sinuous, often with constrictions
and swellings, simple or occasionally branched,
30–110  3–6 µm, pluriseptate, cells 5–15 µm
long, pale to medium olivaceous-brown or
brown, paler towards the tip (pale olivaceous or
subhyaline), thin-walled (up to 0.8 µm),
smooth; conidiogenous cells integrated, ter-
minal to intercalary, 10–30 µm long, with a
single to several conidiogenous loci per cell,
small, 0.8–1 µm diam., slightly thickened and
darkened, less conspicuous than in most other
Cercospora spp. Conidia solitary, narrowly
obclavate, shorter conidia subcylindrical-ellip-
soid to fusiform, long conidia almost acicular,
(10–)15–80  (1.5–)2–3(–3.5) µm, (0–)1–7-
septate, hyaline, thin-walled, smooth, apex
obtuse to subacute, base obconically truncate,
hila about 1 µm wide, slightly thickened and
darkened.
Cercospora ageraticola Goh & W.H.
Hsieh, described on Ageratum houstonianum
Mill. from Taiwan, is very similar, but the
conidiophores are much longer, up to 200 µm,
and less geniculate, the conidiogenous loci are
broader and the conidia are wider, 3–4.5 µm
(Hsieh & Goh 1990). C. agerati-conyzoidis
Bagyan., Jagad. & U. Braun (Bagyanarayana et
al. 1991) on A. conyzoides in India belongs to
the C. apii s. lat. complex (Crous & Braun
2003). The conidia are acicular with truncate
base and the conidiophores are rather long, up
to 250 µm. The following key comprises

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Cercospora spp. on Ageratum:
1. Conidiophores up to 110 µm long, usually
distinctly and often strongly geniciculate-
sinuous; conidiogenous loci not very con-
spicuous, small, about 0.8–1 µm diam.;
conidia narrow, (1.5–)2–3(–3.5) µm; on
Ageratum conyzoides ............. C. conyzoides
1* Conidiophores longer, up to 250 µm, not or
less geniculate-sinuous; conidiogenous loci
broader, > 1 µm; conidia wider, 3–4.5 µm . 2
2. Conidia acicular, base truncate; on Ageratum
conyzoides .................. C. agerati-conyzoidis
2* Conidia acicular to obclavate, base truncate
to obconically truncate; on Ageratum hous-
tonianum ................................ C. ageraticola
Cercospora erythrinicola Tharp
Material examined – VENEZUELA, La-
ra, Parque Macuto, on leaves of Erythrina
berteroana Urb. (Fabaceae), Mar. 2008, R.
Urtiaga 110 (HAL 2534 F).
Notes – New to Venezuela. Urtiaga (1986)
recorded Cercospora sp. on this host from
Venezuela.
Cercospora geraisensis Chupp
Material examined – Cuba, Bayamo, on
leaves of Terminalia catappa L. (Combret-
aceae), 23 Dec. 1966, R. Urtiaga (IMI 124076
= K(M) 176159).
Notes – Crous & Braun (2003) referred C.
geraisensis to the C. apii complex and recorded
this species from Cuba. Arnold (1986) listed T.
catappae from Cuba as host of Cercospora sp.
Cercospora guatemalensis A.S. Mull. &
Chupp
Material examined – CUBA, Bayamo, on
leaves of Ocimum sanctum L. (Lamiaceae), 5
June 1967, R. Urtiaga E-608 (IMI 128006a =
K(M) 176153).
Notes – C. guatemalensis is known from
Cuba (Crous & Braun 2003). This species
belongs to the morphological C. apii s. lat.
complex.
Cercospora hyalospora A.S. Mull. & Chupp
(nom. inval.)
(= C. apii s. lat.)
Material examined – VENEZUELA, La-
ra, Sanare, Sabana Redonda Arriba, on leaves
of Sida urens L. (Malvaceae), June 2010, R.
Urtiaga 392 (HAL 2527 F).
Notes – This species has been recorded
from Venezuela by Dennis (1970) on Sida sp. It
differs from C. sidicola in forming distinct leaf
spots.
Cercospora mikaniicola F. Stevens
Material examined – VENEZUELA, La-
ra, Rio Claro, La Cuchilla, on leaves of
Mikania cordifolia (L. f.) Willd. (Asteraceae),
Apr. 2009, R. Urtiaga 201 (HAL 2551 F); Lara,
Sanare, Sabana Redonda Arriba, on M. cordi-
folia, Apr. 2009, R. Urtiaga 259 (HAL 2562 F).
Notes – New to Venezuela (Crous &
Braun 2003).
Cercospora mucunicola Gonz. Frag & Cif.
(= C. apii s. lat.)
Material examined – VENEZUELA, La-
ra, Rio Claro, La Cuchilla, on leaves of
Dalechampia scandens L. (Euphorbiaceae),
Apr. 2009, R. Urtiaga 246 (HAL 2551 F).
Notes – see Pseudocercospora euphor-
biacearum.
Cercospora ricinella Sacc. & Berl.
Material examined – VENEZUELA, La-
ra, Sanare, Rio Claro, La Cuchilla, on leaves of
Ricinus communis L. [seedlings and adult
plants] (Euphorbiaceae), Apr. 2008, R. Urtiaga
211 & 248 (HAL 2542 F, 2543 F).
Notes – Known from Venezuela (Urtiaga
1986, Crous & Braun 2003, Iturriaga & Minter
2006).
Cercospora sp. (1) Fig. 2
Material examined – VENEZUELA, La-
ra, Villanueva, on leaves of Vismia sp. (Hyperi-
caceae), Nov. 2008, R. Urtiaga 150 (HAL 2515
F).
Notes – Cercospora vismiae Syd. and C.
vismiicola Chupp have been reallocated to the
genus Pseudocercospora. The present collec-
tion belongs to a true Cercospora (s. str.), but
not to C. apii s. lat. It is characterized as
follows: Leaf spots amphigenous, subcircular
to somewhat angular-irregular or oblong, 2–25
mm diam., light brown, ochraceous, later
almost greyish white, surrounded by a narrow
somewhat darker marginal line and a diffuse
darker halo; caespituli hypophyllous, not very
conspicuous; mycelium internal; stromata

Mycosphere Doi 10.5943/mycosphere/4/2/3
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lacking or only small, substomatal to intra-
epidermal, 10–25 µm diam., brown; coni-
diophores solitary or in small, divergent
fascicles (2–8), arising from internal hyphae or
stromata, through stomata or erumpent, erect,
straight, subcylindrical to usually strongly
geniculate-sinuous, unbranched, 20–250  3–7
µm, pluriseptate, thin-walled, pale to medium
brown throughout or paler towards the tip,
smooth; conidiogenous cells integrated, ter-
minal and intercalary, 10–25 µm long, with a
single to usually several, sometimes numerous
conidiogenous loci, thickened and darkened,
1.5–2.5 µm diam.; conidia solitary, narrowly
obclavate, acicular, 20–80  1.5–3 µm, pluri-
septate, hyaline, thin-walled, smooth, apex
subacute, base truncate to obconically truncate,
hila (1–)1.5–2 µm wide, thickened and
darkened. The material is not sufficient for a
final description. Furthermore, Cercospora
species with acicular conidia should only be
described as new species when the identity as
separate species has been proven by cultures
and molecular sequence analyses (Groenewald
et al. 2012). The taxonomy of this complex is
intricate and many plurivorous species with
insufficiently known host range are involved.
Cercospora sp. (2)
Material examined – VENEZUELA, La-
ra, Sanare, Sabana Redonda Arriba, on leaves
of Conyza canadensis (L.) Cronquist (Aster-
aceae), June 2009, R. Urtiaga 261 (HAL 2563
F).
Notes – This collection is characterized
as follows: Conidiophores in small to mode-
rately large fascicles, distinctly geniculate, 50–
160  3–7 µm, pluriseptate, medium brown;
conidiogenous cells terminal and intercalary,
conidiogenous loci 2.5–4 µm diam.; conidia
acicular, 40–180  2.5–5 µm, relatively densely
pluriseptate, hyaline, apex subacute to mostly
obtuse, base truncate, hila 2–4 µm wide.
Cercospora nilghirensis Govindu & Thirum. on
Conyza spp. in India is similar, but has densely
aggregated conidiophores and the conidia are at
least partly obclavate with obconically truncate
base (Vasudeva 1963). The North American
Cercospora erigeronicola U. Braun & Roger-
son (Braun & Rogerson 1993) on Erigeron
divergens Torr. & A. Gray is a quite distinct
species with cylindrical conidia. Furthermore,
C. bidentis Tharp has been recorded on Conyza
sp. Identity and relation between Cercospora
on Bidens and Conyza are, however, unclear.
Cultures and molecular sequence data are
necessary to elucidate the taxonomy of the taxa
concerned.
Cercospora sidicola Ellis & Everh.
Material examined – VENEZUELA, La-
ra, Sanare, Sabana Redonda Arriba, on Sida
acuta Burm. f. (Malvaceae), June 2010, R.
Urtiaga 385 (HAL 2526 F).
Notes – New to Venezuela (Iturriaga &
Minter 2006, Crous & Braun 2003).
Fig. 2 – Cercospora sp. on Vismia sp. Based on
HAL 2515 F. a Conidiophore fascicle. b
Conidiophore tips. c Conidia. – Bar = 10 µm.
Cercospora solanicola G.F. Atk.
Material examined – CUBA, Rio Cauto,
on leaves of Solanum nigrum L. (Solanaceae),
25 Aug. 1967, R. Urtiaga 788 (IMI 129025 =
K(M) 176132).
Notes – Colonies in this collection are
confined to stems and can morphologically be
assigned to C. solanicola (Chupp 1954), a

Mycosphere Doi 10.5943/mycosphere/4/2/3
182
species that belongs to the C. apii compex
(Crous & Braun 2003).
Cercospora talini Syd. & P. Syd.
Material examined – VENEZUELA, La-
ra, Barquisimeto, on leaves of Talinum pani-
culatum (Jacq.) Gaertn. [= T. patens (L.) Juss.]
(Portulacaceae), Apr. 2009, R. Urtiaga 250
(HAL 2539 F).
Notes – This species, based on type
material from Argentina, was first recorded
from Venezuela in Chupp (1954). In the
present collection, this fungus is possibly a
secondary invader, associated with Clado-
sporium and additional fungi.
Fig. 3 – Cercosporella ambrosiae-artemi-
siifoliae. Based on type material. a Conidio-
phore fascicles. b Solitary conidiophores
arising from superficial hyphae. c Conidio-
phores. d Conidia. – Bar = 10 µm.
Cercospora volkameriae Speg.
Material examined – CUBA, Bayamo, on
leaves of Clerodendrum speciosissimum C.
Morren (Lamiaceae), 21 Jan. 1967, R. Urtiaga
3 (IMI 124815 = K(M) 176125).
Notes – C. volkameriae is a species of the
C. apii s. lat. complex (Crous & Braun 2003),
which is known from Cuba on Clerodendrum
splendens G. Don (Arnold 1986).
Cercosporella ambrosiae-artemisiifoliae U.
Braun & Urtiaga, sp. nov. Fig. 3
MycoBank, MB 801952.
Etymology – epithet derived from the
host species, Ambrosia artemisiifolia.
Differt ab omnibus speciebus Cerco-
sporellae ad hospites Asteracearum conidio-
phoris fasciculatis et etiam solitariis, ex hyphis
superficialibus oriundis, subhyalinis, pallide
olivaceis, pallide flavo-brunneis vel aureo-
brunneis. Praeterea Passalorae ambrosiae
superficiale similis, sed stromatibus minoribus,
ad 30 µm diam., conidiophoris fasciculatis et
etiam solitariis, cicatricibus conidialibus con-
vexis, non fuscatis et conidiis angustioribus, 3–
6 µm latis.
Leaf spots amphigenous, shape and size
variable, subcircular to irregular, 2–10 mm
diam. or expanded and larger, yellowish to
ochraceous or greyish brown to medium
brown, margin indefinite. Caespituli mainly
hypophyllous, not very conspicuous, puncti-
form to subeffuse, greyish to brownish. Myce-
lium internal and external; superficial hyphae
emerging through stomata, sparingly branched,
1–4 µm wide, septate, thin-walled, smooth,
subhyaline; stromata lacking or small, sub-
stomatal, 10–30 µm diam., brownish. Conidio-
phores in small to moderately large fascicles,
loose to moderately dense, arising from in-
ternal hyphae or stromata, emerging through
stomata, or solitary, arising from superficial
hyphae, lateral, erect, straight, subcylindrical to
moderately geniculate-sinuous, simple or
occasionally branched, 20–90 × 3–7 µm, 0–5-
septate, subhyaline to pale olivaceous, yellow-
ish to golden brown below and paler towards
the tip or faintly pigmented throughout, darker
in mass, thin-walled, smooth; conidiogenous
cells integrated, terminal or conidiophores
occasionally reduced to conidiogenous cells,
15–30 µm long, proliferation sympodial, rarely

Mycosphere Doi 10.5943/mycosphere/4/2/3
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percurrent, conidiogenous loci (scars) con-
spicuous, 1.5–2 µm diam., somewhat bulging
(convex), slightly thickened and refractive, but
not darkened. Conidia formed singly, ob-
clavate-cylindrical, 20–80 × 3–6 µm, 1–8-
septate, subhyaline or with a very pale
yellowish, greenish or olivaceous tinge, thin-
walled, smooth, apex obtuse, base obconically
truncate to rounded, hila 1.5–2 µm wide,
slightly thickened and somewhat refractive.
Material examined – CUBA, Bayamo, on
leaves of Ambrosia artemisiifolia L. (Aster-
aceae), 6 June 1966, R. Urtiaga (IMI 119623 =
K(M) 176119, holotype).
Notes – Cercospora ambrosiae Chupp
was described from Colombia on Ambrosia
peruviana Willd. (Chupp 1954). Crous &
Braun (2001) re-examined type material and
reallocated this species to Passalora. This
species is known on A. peruviana from
Colombia, Dominican Republic, Puerto Rico
and Venezuela (Chupp 1954, Crous & Braun
2001, 2003). Records of Passalora ambrosiae
(Chupp) Crous & U. Braun on Ambrosia
artemisiifolia from Cuba (Crous & Braun
2003) are based on misidentifications and refer
to a different species, described above as
Cercosporella ambrosiae-artemisiifoliae. P.
ambrosiae is only superficially similar to the
latter species and easily distinguishable by its
larger stromata, up to 60 µm diam., lacking
superficial hyphae, consistently fasciculate
conidiophores (solitary conidiophores lacking),
thickened and darkened conidiogenous loci and
much broader conidia (6–10 µm). The fungus
on Ambrosia artemisiifolia from Cuba repre-
sents a new undescribed species, but its generic
affinity is difficult and must be discussed in
detail. At first glance, it seems that this species
might be ascribable to Passalora based on
pigmented conidiophores, conspicuous coni-
diogenous loci and obclavate-cylindrical coni-
dia. However, the structure of the coni-
diogenous loci does not agree with Passalora
scars, which are planate, thickened and
darkened throughout. The scars of the fungus
on Ambrosia artemisiifolia are bulging, re-
fractive, but not darkened, i.e. the wall of the
loci is not darker than the surrounding wall of
the conidiogenous cell, and rather coincide
with conidiogenous loci of the genus Cerco-
sporella. Pigmented and solitary conidio-
phores arising from superficial hyphae are
unusual in Cercosporella, but known in
several, mainly tropical-subtropical species,
e.g. C. crataevae (Berk. & Broome) Petch, C.
hypoestis Hansf., C. polysciatis (Henn.) Hansf.,
C. rosea G. Winter, C. pseudoidium Speg., C.
pyri (Farl.) Karak. and C. pyrina Ellis & Everh.
(Braun 1995b). Monophyly of Cercosporella in
its current circumscription is still unclear. Only
a few cultures of Cercosporella spp. are
available, and comprehensive phylogenetic
examinations of Cercosporella, including pig-
mented taxa and species with superficial
mycelium, have not yet been done. Thus, we
follow the current taxonomic concept of the
latter genus as outlined in Braun (1995b).
Numerous Cercosporella species on hosts
belonging to the Asteraceae are known,
including C. cana (Sacc.) Sacc. [= C.
virgaureae (Thüm.) Allesch.], the type species,
but all of them differ from C. ambrosiae-
artemisiifoliae in having colourless conidio-
phores, consistently formed in fascicles, i.e.
superficial hyphae with solitary conidiophores
are not developed (Braun 1995b).
Cercosporella virgaureae (Thüm.) Allesch.
Material examined – VENEZUELA, La-
ra, Rio Claro, La Cuchilla, on leaves of Conyza
canadensis (L.) Cronquist (Asteraceae), Apr.
2009, R. Urtiaga 228 (HAL 2553 F); Lara,
Sanare, Sabana Redonda Arriba, on C.
canadensis, June 2009, R. Urtiaga 260 (HAL
2564 F).
Notes – This species is new to Vene-
zuela (Braun 1995b, Iturriaga & Minter 2006).
Passalora crotonis-gossypiifolii U. Braun &
Urtiaga, sp. nov. Fig. 4
MycoBank, MB 801953.
Etymology – epithet derived from the
host species, Croton gossypiifolius.
Passalorae rubidae similis, sed conidio-
phoris semper non fasciculatis, hilis 0.8–1.5
µm diam. et conidiis brevioribus et angusti-
oribus, (8–)12–40(–50)  (2.5–)3–5(–5.5) µm,
0–3-septatis.
Leaf spots lacking, on the lower leaf
surface only visible as dingy greyish brown to
sooty patches caused by fungal colonies in the
tomentum, subcircular to irregular, 2–8 mm
diam. or confluent and larger. Mycelium

Mycosphere Doi 10.5943/mycosphere/4/2/3
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internal and external; superficial hyphae
emerging through stomata, climbing leaf hairs,
branched, septate, 1–5 µm wide, subhyaline to
medium olivaceous-brown, smooth or almost
so. Stromata lacking. Conidiophores solitary,
arising from superficial hyphae, lateral or
terminal, sometimes loosely aggregated, but
true fascicles not formed, erect to decumbent,
simple or branched, straight, subcylindrical to
strongly geniculate-sinuous, 5–100  3–6 µm,
aseptate to pluriseptate, pale to medium
olivaceous-brown or brown, thin-walled,
smooth or almost so; conidiogenous cells
integrated, terminal and intercalary, 8–25 µm
long, with a single to mostly several con-
spicuous conidiogenous loci, slightly thickened
and somewhat darkened, 0.8–1.5 µm diam.
Conidia catenate, often in branched chains,
mostly cylindrical or subcylindrical, short
conidia sometimes ellipsoid-ovoid, longer
conidia occasionally almost obclavate, (8–)12–
40(–50)  (2.5–)3–5(–5.5) µm, 0–3-septate,
pale olivaceous to olivaceous-brown, thin-
walled, smooth or almost so, ends rounded to
short obconically truncate, hila 0.8–1.5 µm
wide, slightly thickened and darkened.
Material examined – VENEZUELA, La-
ra, Villanueva, on leaves of Croton gossy-
piifolius Vahl (Euphorbiaceae), Nov. 2008, R.
Urtiaga 146 (HAL 2522 F, holotype).
Notes – Numerous Passalora species
have been described on hosts of the genus
Croton. The new species on C. gossypiifolius
from Venezuela is characterized by the
formation of superficial hyphae with solitary
conidiophores, i.e. it pertains to a group of
Passalora species previously assigned to the
genus Mycovellosiella, which is now con-
sidered a synonym of Passalora (Crous &
Braun 2003). Three Mycovellosiella-like
Passalora species are known on Croton spp.,
viz. Passalora crotoniphila (Speg.) Crous, P.
manaosensis (Henn.) U. Braun & Crous and P.
rubida Crous, Alfenas & R.W. Barreto, but all
of them have much longer, pluriseptate conidia
and except for the latter species also well-
developed stromata and fasciculate conidio-
phores (Chupp 1954, Crous et al. 1999, 2000,
Crous & Braun 2003). Superficial mycelium
and solitary conidiophores are lacking in all
other Passalora spp. on Croton, i.e. they are
characterized by having conidiophores only
formed in fascicles. The conidia in Passalora
crotonifolia (Cook) Crous, U. Braun & Alfenas
are formed singly (Chupp 1954, Crous et al.
1999), whereas the conidia in all other species
of this group are formed in chains, i.e. they are
Phaeoramularia-like: Passalora crotonis (Ellis
& Everh.) Crous & U. Braun, P. crotonis-
oligandri (J.M. Yen & Gilles) Crous, U. Braun
& Alfenas and P. maritima (Tracy & Earle)
Crous & U. Braun (Chupp 1954, Yen 1971,
Crous et al. 1999, Crous & Braun 2003). The
species concerned are keyed out as follows
1. Superficial hyphae with solitary conidio-
phores developed ........................................ 2
1* Superficial hyphae with solitary conidio-
phores lacking (conidiophores only formed in
fascicles) ..................................................... 5
2. Conidia (8–)12–40(–50)  (2.5–)3–5(–5.5)
µm, 0–3-septate; on Croton gossypiifolius,
Venezuela ............... P. crotonis-gossypiifolii
2* Conidia much longer, 20–170 µm, pluri-
septate, (0–)1–13 ........................................ 3
3. Stromata and conidiophore fascicles lacking;
conidia very long, 25–170 µm, up to 13-
septate; on Croton floribundus and C.
peruvianus, Brazil, Peru ................ P. rubida
3* Stromata and fasciculate conidiophores
developed; conidia shorter, up to about 100
µm, with up to 9 septa ................................ 4
4. Stromata very large, 30–150 µm diam.;
conidiophores 25–200 µm long; conidia 3–8
µm wide; on Croton sp., Brazil, Venezuela
.............................................. P. manaosensis
4* Stromata smaller, up to about 40 µm diam.;
conidiophores only 10–30 µm long; conidia
narrower, (2.5–)3–4(–5) µm; on Croton
gossypiifolius, C. glandulosus, Croton sp.,
Brazil, Paraguay, USA, Venezuela
............................................... P. crotoniphila
5. Conidia formed singly; on Croton glandu-
losus, USA ............................ P. crotonifolia
5* Conidia in chains ...................................... 6
6. Caespituli epiphyllous; conidiophores short,
15–45 µm; conidia hyaline and small, 14–22
 2–3 µm; on Croton oligandrum, Gabon
..................................... P. crotonis-oligandri
6* Caespituli amphigenous, often hypo-
phyllous; conidiophores much longer, up to
130 µm; conidia much larger, up to 120  7

Mycosphere Doi 10.5943/mycosphere/4/2/3
185
µm; on other species .................................. 7
7. Conidiophores loosely fasciculate; conidia
smooth, 2–8-septate; on Croton capitatus, C.
lobatus, C. texensis, Croton sp., Cuba,
Ghana, Sudan, Trinidad and Tobago, USA,
Venezuela ................................... P. crotonis
7* Conidiophores in dense, often coremioid
fascicles; conidia finely verruculose, 1–5-
septate; on Croton glandulosus, C. mariti-
mus, C. punctatus and Croton sp., Dominican
Republic, USA, Venezuela ....... P. maritima
Fig. 4 – Passalora crotonis-gossypiifolii.
Based on type material. a Hyphae. b Solitary
conidiophores arising from superficial hyphae.
c Conidia. – Bar = 10 µm.
Passalora henningsii (Allesch.) R.F. Casta-
ñeda & U. Braun
 Cercospora henningsii Allesch.
Material examined – VENEZUELA, La-
ra, Villanueva, on leaves of Manihot esculenta
Crantz (Euphorbiaceae), Nov. 2008, R. Urtiaga
140 (HAL 2520 F).
Notes – This species is listed from
Venezuela in Crous & Braun (2003), but not
included in Iturriaga & Minter (2006).
Passalora lemnischea (Cif.) U. Braun & Crous
 Cercospora lemnischea Cif.
= Cercospora mikaniae Ellis & Everh.,
non Passalora mikaniae (F. Stevens) U. Braun
& F.O. Freire.
 Passalora mikaniigena U. Braun &
Crous, nom. superfl.
Material examined – VENEZUELA, La-
ra, Sanare, Sabana Redonda Arriba, on leaves
of Mikania cordifolia (L. f.) Willd. (Astera-
ceae), June 2009, R. Urtiaga 264 (HAL 2562
F).
Notes – New to Venezuela and new host
species (Crous & Braun 2003, Iturriaga &
Minter 2006).
Passalora solaniphila U. Braun & Urtiaga, sp.
nov. Fig. 5
MycoBank, MB 801954.
Etymology – epithet derived from the
host genus Solanum and -philus (-loving,
Greek).
Passalorae aratai paulum similis, sed
conidiis angustioribus, (2.5–)3–5 µm, et hilis
quoque angustioribus, 1–2 µm.
Leaf spots amphigenous, subcircular to
angular-irregular, 1–10 mm diam. or confluent
and larger, on the upper leaf side conspicuous,
at first greenish grey, olivaceous, later yel-
lowish, ochraceous to light brown, margin
indefinite or narrow and somewhat darker,
hypophyllous spots less conspicuous or almost
lacking, yellowish, later olivaceous, brown to
greyish white by abundant fructification (fas-
cicles of conidiophores and conidia). Caespituli
hypophyllous, punctiform to dense, olivaceous,
brown to greyish white by abundant conidial
formation. Mycelium internal. Stromata lack-
ing to well-developed, 10–70 µm diam., sub-
stomatal or occasionally intraepidermal, pale to
medium brown or olivaceous-brown, cells 2–5

Mycosphere Doi 10.5943/mycosphere/4/2/3
186
µm diam. Conidiophores in small to usually
large fascicles, divergent to mostly dense,
arising from internal hyphae or stromata,
through stomata or occasionally erumpent,
erect, straight, subcylindrical, subclavate or
somewhat narrowed towards the tip, not to
moderately geniculate-sinuous, unbranched,
10–50  (2.5–)3–5(–6) µm, 0–2(–3)-septate,
subhyaline, pale olivaceous to olivaceous-
brown, medium olivaceous-brown in mass,
thin-walled, smooth; conidiogenous cells inte-
grated, terminal or conidiophores reduced to
conidiogenous cells, 5–30 µm long, with a
single to mostly several conspicuous conidio-
genous loci, 1–2 µm diam., slightly thickened
and somewhat darkened. Conidia solitary,
cylindrical or obclavate-cylindrical, (10–)15–
60  (2.5–)3–5 µm, (0–)1–4(–5)-septate, sub-
hyaline to pale olivaceous, thin-walled, smooth
to faintly rough-walled, apex obtuse, base
rounded to short obconically truncate, hila 1–2
µm broad, slightly thickened and somewhat
darkened.
Fig. 5 – Passalora solaniphila. Based on type
material. a Conidiophore fascicle. b Conidio-
phores. c Conidia. – Bar = 10 µm.
Material examined – VENEZUELA, La-
ra, Barquisimeto, on leaves of Solanum nigrum
L. (Solanaceae), Mar. 2008, R. Urtiaga 112
(HAL 2532 F, holotype).
Notes – Due to conspicuous conidio-
genous loci and pigmented cylindrical to
obclavate-cylindrical conidia, this fungus on
Solanum nigrum from Venezuela belongs in
the genus Passalora. It is morphologically
superficially similar to Cercospora solanacea
Sacc. & Berl., but the latter species belongs in
Pseudocercospora (see under P. solanacea in
this paper). Two other comparable Passalora
species with fasciculate conidiophores and
conidia formed singly are known on Solanum
spp., but the species concerned have quite
distinct conidia which are above all much
broader [Passalora aratai (Speg.) U. Braun, R.
Delhey & M. Kiehr – conidia 6–14 µm wide,
hila 1.5–3 µm diam. (Chupp 1954, Braun et al.
2001); P. solani (Seaver) U. Braun – conidia
12–32  6–12 mm, 0–1-septate (Braun 1992)].
P. bruchiana (Speg.) U. Braun & Crous is
Phaeoramularia-like, i.e. the conidia are
formed in chains (Chupp 1954, Crous & Braun
2003). There are numerous additional species
of Passalora on Solanum spp., but all of them
are quite distinct from P. solaniphila in
forming superficial mycelium with solitary
conidiophores, i.e. they are Mycovellosiella-
like: Passalora brachycarpa (Syd.) U. Braun &
Crous, P. concors (Casp.) U. Braun & Crous,
P. dulcamarae (Peck) U. Braun & Crous, P.
incarnata (Deighton) U. Braun & Crous, P.
nattrassii (Deighton) U. Braun & Crous, P.
paradoxa (Munt.-Cvetk.) U. Braun & Crous, P.
solanacearum (K. Bhalla, S.K. Singh & A.K.
Srivast.) U. Braun & Crous, P. solani-torvi
(Gonz. Frag. & Cif.) U. Braun & Crous, P.
tarrii (Deighton) U. Braun & Crous (Chupp
1954, Deighton 1974, Crous & Braun 2003).
Passalora stigmaphyllicola U. Braun & Urti-
aga, sp. nov. Fig. 6
MycoBank, MB 801956.
Etymology – epithet derived from the
host genus, Stigmaphyllon.
Passalorae peixotoae paulum similis, sed
maculis foliorum et stromatibus cum conidio-
phoris fasciculatis formantibus et conidiis 20–
55  3–5.5 µm, 1–5-septatis.

Mycosphere Doi 10.5943/mycosphere/4/2/3
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Fig. 6 – Passalora stigmaphyllicola. Based on
type material. a Conidiophore fascicles. b
Conidiophores arising from superficial hyphae.
c Conidiophores. d Conidia. – Bar = 10 µm.
Leaf spots amphigenous, subcircular to
usually angular-irregular, 2–8 mm diam. or
occasionally confluent and larger, pale to dark
brown, later greyish brown to dingy grey,
margin indefinite or narrow and darker, usually
with a narrow to moderately broad diffuse
purplish halo. Caespituli hypophyllous, incon-
spicuous. Mycelium internal and external,
superficial, emerging through stomata; hyphae
sparingly branched, 1–2.5 µm wide, subhyaline
to pale olivaceous, septate, smooth thin-walled.
Stromata lacking or small to moderately large,
10–50 µm, substomatal to intraepidermal,
circular to somewhat angular-irregular in out-
line, brown, cells 2–8 µm diam. Conidiophores
in small to moderately large fascicles, loose to
moderately dense, arising from internal hyphae
or stromata, emerging through stomata or
erumpent, or solitary, arising from superficial
hyphae, lateral, erect, straight, subcylindrical-
conical to slightly geniculate, unbranched or
branched, 5–60 × 2–6 µm, 0–3-septate, pale
olivaceous to olivaceous-brown, thin-walled,
smooth; conidiogenous cells integrated, termi-
nal or conidiophores occasionally reduced to
conidiogenous cells, 5–25 µm long, conidio-
genous loci conspicuous, 0.8–2 µm diam.,
somewhat thickened and darkened. Conidia
solitary to catenate, in simple or occasionally
branched chains, cylindrical, ellipsoid-fusi-
form, short obclavate, 15–25 × 2–3 µm, 1–3-
septate, subhyaline to very pale olivaceous,
thin-walled, smooth, apex obtuse, subacute to
subtruncate in catenate conidia, base short ob-
conically truncate, hila 0.8–1.2 µm diam.,
slightly thickened and darkened.
Material examined – CUBA, Bayamo, on
leaves of Stigmaphyllon sagraeanum A. Juss.
(Malpighiaceae), 26 Mar. 1966, R. Urtiaga
(IMI 118051 = K(M) 176146, holotype); l.c.,
20 Oct. 1966, R. Urtiaga (IMI 122811 = K(M)
176147, paratype).
Notes – Passalora stigmaphyllicola is a
Mycovellosiella-like species, with solitary
conidiophores arising from superficial myce-
lium. P. stigmaphylli (R.E.D. Baker & W.T.
Dale) U. Braun & Crous on Stigmaphyllon
species in Cuba and Trinidad is quite distinct
from P. stigmaphyllicola by its very large
stromata, fasciculate conidiophores, up to 100
µm long (superficial hyphae and solitary
conidiophores lacking) and conidia formed
singly, 30–75  3.5–6 µm, 1–5-septate (Chupp
1954, Crous & Braun 2003). Several other
Passalora species on hosts of other genera of
the Malpighiaceae have been described. P.
peixotoae (Chupp & Viégas) U. Braun & Crous
on Peixotoa reticulata Griseb. (= G. macro-
phylla Griseb.) in Brazil is the only comparable
species. However, leaf spots, stromata and
fasciculate conidiophores are lacking in the
latter species and the 1–5-septate conidia are
20–55  3–5.5 µm (Chupp 1954). Superficial
hyphae with solitary conidiophores are lacking
in all other species. In Passalora bunchosiae
U. Braun & Crous ( Cercospora bunchosiae
Chupp & A.S. Mull., nom. inval.) on
Bunchosia glandulifera in Venezuela, P.
cornifoliae (Chupp) U. Braun & Crous on
Bunchosia nitida (Jacq.) A. Rich. (= B.
cornifolia Kunth) in Colombia, P. kreiseliana
U. Braun & Crous on Malpighia glabra L. in
Jamaica and P. malpighiae-glabrae U. Braun
& Crous on M. glabra in Florida, USA, the
conidia are formed singly (Chupp 1954, Braun

Mycosphere Doi 10.5943/mycosphere/4/2/3
188
et al. 2002, Crous & Braun 2003). P.
malphigiae (U. Braun & Mouch.) U. Braun &
Crous on Malpighia glabra (= M. punicifolia
L.) in French Polynesia is characterized by
catenate conidia which are shorter, 6–18  1.5–
3 µm, hyaline or subhyaline and verruculose
(Braun et al. 1999).
Fig. 7 – Pseudocercospora calycophylli. Based
on type material. a Superficial hyphae. b
Solitary conidiophores arising from superficial
hyphae. c – Conidiophore fascicles. e Conidio-
phores. e Conidia. – Bar = 10 µm.
Pseudocercospora borreriae (Ellis & Everh.)
Deighton
 Cercospora borreriae Ellis & Everh.
Material examined – VENEZUELA, La-
ra, Rio Claro, La Cuchilla, on leaves of Sper-
macoce sp. (Borreria sp.) (Rubiaceae), Apr.
2009 R. Urtiaga 226 (HAL 2538 F).
Notes – This species was not listed from
Venezuela in Crous & Braun (2003), but
recorded from this country by Dennis (1970)
on Spermacoce latifolia Aubl. The present
collection is characterized by having small to
very large, loose to dense fasciles of long
conidiophores, up to 200 µm and cylindrical to
obclavate-cylindrical conidia, 30–100  3.5–
5(–5.5) µm, 3–10-septate.
Pseudocercospora calycophylli U. Braun &
Urtiaga, sp. nov. Fig. 7
MycoBank, MB 801957.
Etymology – epithet derived from the
host genus, Calycophyllum.
Pseudocercosporae cinchonicolae similis,
sed hyphis superficialibus cum conidiophoris
solitariis formantibus et hilis conidiorum
angustioribus, 1–2 µm diam.
Leaf spots amphigenous, subcircular to
usually angular-irregular, 2–12 mm diam. or
confluent and larger, medium to dark brown on
the upper leaf side, later brown below, margin
indefinite or surrounded by somewhat darker
veins. Caespitili amphigenous, finely puncti-
form on the upper side, less conspicuous
below, scattered, dark brown to blackish.
Mycelium internal and external, superficial
hyphae only on the lower leaf surface,
emerging through stomata, sparingly branched,
1.5–3 µm wide, septate, subhyaline to pale
olivaceous, thin-walled, smooth; stromata
lacking to well-developed, above all on the
upper leaf side, immersed or substomatal, 10–
40 µm diam., medium to dark brown,
composed of swollen hyphal cells, 2–6 µm
diam. Conidiophores in small to moderately
large fascicles, loose to dense, arising from
internal hyphae or usually from stromata,
erumpent or emerging through stomata, occa-
sionally with some solitary conidiophores
arising from superficial hyphae, lateral, erect,
straight, subcylindrical to distinctly geniculate-
sinuous, mostly unbranched, only occasionally
irregularly branched, 5–60 × 2–5 µm, 0–3-

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septate, sometimes constricted at the septa, pale
olivaceous to medium olivaceous-brown, thin-
walled, smooth to faintly rough-walled; coni-
diophores reduced to conidiogenous cells or
conidiogenous cells integrated, terminal, 5–30
µm long, proliferation sympodial, occasionally
percurrent, conidiogenous loci inconspicuous
or subdenticulate, but wall of the loci always
unthickened and not darkened. Conidia formed
singly, narrowly obclavate to subcylindrical,
(10–)20–95(–120) × 2.5–4 µm, (0–)1–8(–10)-
septate, subhyaline to pale olivaceous, thin-
walled, smooth to faintly rough-walled, apex
obtuse to subacute, base usually short ob-
conically truncate, occasionally truncate, 1–2
µm wide, hila unthickened, not darkened.
Material examined – CUBA, Bayamo, on
leaves of Calycophyllum candidissimum (Vahl)
DC. (Rubiaceae, Cinchonoideae, Calyco-
phylleae), 12 Apr. 1967, R. Urtiaga (IMI
126874a = K(M) 176139, holotype); CUBA,
without locality, on C. candidissimum, 10 Jan.
1972, L.H. Isla 8 (IMI 163712 = K(M)
176140).
Notes – P. calycophylli is the first species
of Pseudocercospora on a host of the genus
Calycophyllum. Comparable species on allied
genera of the Calycophylleae (Rubiaceae, Cin-
chonoideae) are unknown. Several Pseudo-
cercospora species have been described on
more distantly related hosts belonging to other
tribes of the Cinchonoideae, including the
morphological similar species P. cinchonicola
(Boedijn) U. Braun on Cinchona sp. in
Indonesia (Braun 2001), which differs from P.
calycophylli in having consistently fasciculate
conidiophores (superficial hyphae with solitary
conidiophores lacking) and conidia with wider
hila, (1.5–)2–2.5(–3) µm, and P. hymenodictyi
(Petr.) Y.L. Guo & X.J. Liu on Hymenodictyon
orixense (Roxb.) Mabb. (= H. excelsum
(Roxb.) DC.) in Asia, characterized by its
much wider conidia, 30–70 × 4–6.5 µm (Braun
1995a, Guo & Hsieh 1995). Other species are
quite distinct [P. cinchonae (Ellis & Everh.) U.
Braun & Crous on Cinchona spp. in Africa and
North America, superficial hyphae lacking,
conidiophores very short, conidia narrowly
obclavate-cylindrical to linear, 25–80 × 2–3
µm (Chupp 1954, Crous & Braun 2003); P.
mussaendae Katsuki on Mussaenda parviflora
Miq. in Japan, stromata lacking, conidiophores
arising from decumbent threads, conidia 6–7
µm wide (Katsuki 1956, 1965); P. philip-
pinensis (Tak. Kobay. & E.D. Guzman) U.
Braun & Crous on Mussaenda philippica A.
Rich., Philippines, superficial hyphae lacking,
conidia wider, 4.5–5.5 µm (Kobayashi &
Guzman 1988, Crous & Braun 2003)].
Pseudocercospora catappae (Henn.) X.J. Liu
& Y.L. Guo
 Cercospora catappae Henn.
Material examined – VENEZUELA, La-
ra, Barquisimeto, zoological garden, on leaves
of Terminalia catappa L. (Combretaceae), Jan.
2010, R. Urtiaga 314 (HAL 2558 F).
Notes – New to Venezuela (not listed in
Urtiaga 1986, Crous & Braun 2003 and
Iturriaga & Minter 2006).
Pseudocercospora cordiae-alliodorae U.
Braun & Urtiaga
Material examined – VENEZUELA, La-
ra, Barquisimeto, zoological garden, on leaves
of Cordia toqueve Aubl. (Boraginaceae), Apr.
2008, R. Urtiaga 126 (HAL 2513 F); l.c., on
leaves of Cordia alliodora (Ruiz & Pav.)
Oken, Mar. 2008, R. Urtiaga 118 (HAL 2523
F) and Apr. 2008, R. Urtiaga 127 (HAL 2514
F).
Notes – This species was described by
Braun & Urtiaga (2012) based on material on
Cordia alliodora collected in the zoological
garden of Barquisimeto, Venezuela. The
specimen on Cordia toqueve, collected in the
zoological garden of Barquisimeto as well, is
sparingly developed, but some superficial
hyphae with solitary conidiophores, found in
this material, and small conidia, 15–25  2.5–3
µm, 1–3(–4)-septate, agree well with type
material of P. cordiae-alliodorae. Cordia
toqueve is a new host for this species (Braun &
Urtiaga 2012).
Pseudocercospora cordiana U. Braun &
Urtiaga
Material examined – VENEZUELA, La-
ra, Parque Macuto, on leaves of Cordia alba
(Jacq.) Roem. & Schult. [= C. dentata Poir.]
(Boraginaceae), Mar. 2008, R. Urtiaga 114
(HAL 2533 F).

Mycosphere Doi 10.5943/mycosphere/4/2/3
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Notes – This species, recently described
from Cuba (Braun & Urtiaga 2012), is new to
Venezuela.
Pseudocercospora coremioides U. Braun &
Urtiaga, sp. nov. Fig. 8
MycoBank, MB 801958.
Etymology – epithet referring to the
coremium-like fascicles of conidiophores.
Pseudocercosporae richarsoniicolae valde
similis, sed stromatibus nullis vel minoribus,
10–40 µm diam., fasciculis procerioribus et
conidiophoris solitariis ex hyphis superficia-
libus oriundis bene evolutis.
Leaf spots amphigenous, about 5–10 mm
diam., subcircular to somewhat irregular,
yellowish, ochraceous to brown, margin
indefinite. Caespituli hypophyllous, conspi-
cuous, scattered, dark olivaceous-brown to
almost blackish, visible as minute brush-like
tufts or coremioid aggregations of conidio-
phores when viewed with a stereomicroscope.
Mycelium internal and external; superficial
hyphae emerging through stomata, sparingly
branched, 1.5–5 µm wide, septate, subhyaline
to pale olivaceous-brown, thin-walled, smooth
or almost so; stromata lacking to moderately
large, substomatal, 10–40 µm diam., medium
to dark brown. Conidiophores in small to large
fascicles, loose to very dense, coremium-like,
densely appressed almost through or only in
the lower half and splaying out in the upper
half, sometimes solitary, arising from
superficial hyphae, erect, straight, sub-
cylindrical, filiform, barely geniculate-sinuous
or only slightly to moderately so near the apex,
40–300  2.5–6 µm, pluriseptate, individual
threads pale to medium olivaceous or
olivaceous-brown, much darker in mass, thin-
walled, smooth to faintly rough; conidiogenous
cells integrated, terminal, occasionally inter-
calary, 10–30 µm long, conidiogenous loci
inconspicuous, unthickened and not darkened,
occasionally subdenticulate. Conidia solitary,
obclavate-cylindrical, 25–100  4–6.5 µm, 3–
12-septate, subhyaline to pale olivaceous or
olivaceous-brown, thin-walled, smooth or
almost so, apex obtuse, base obconically
truncate, (1.5–)2–2.5(–3) µm wide, hila neither
thickened nor darkened.
Material examined – VENEZUELA, La-
ra, Villanueva, on leaves of Diodia sp.
(Rubiaceae, Rubioideae, Spermacoceae), Nov.
2008, R. Urtiaga 141 (HAL 2517 F, holotype).
Notes – The identification of the host
plant caused some difficulties. In any case, it
belongs to a genus and species of tribe
Spermacoceae as currently circumscribed
(Groeninckx et al. 2009). Richardia (incl.
Richardsonia) can be ruled out as the calyx of
the flowers is 4-lobed. The host looks like a
species of Diodia or Spermacoce, but since the
capsules seem to be indehiscent, as far as
discernable, it is rather a species of Diodia.
Several morphologically similar Pseudo-
cercospora spp. are known on related hosts of
the Spermacoceae. P. richardsoniicola Crous
& A.P.S. Câmara [ Cercospora richardsoniae
Henn., non Pseudocercospora richardsoniae
Crous & A.P.S. Câmara (as “(Ellis & Everh.)
Crous & A.P.S. Câmara)”] on Richardia spp.
in Brazil is morphologically very close, but the
stromata are very large, up to 100 µm,
pustulate, the conidiophore fascicles are also
very large, up to 100 µm wide, and solitary
conidiophores arising from superficial hyphae
are not formed (Chupp 1954, Crous & Câmara
1998). P. borreriae (Ellis & Everh.) Deighton,
widespread on numerous host species of
Mitracarpus and Spermacoce, is also
comparable, but superficial mycelium with
solitary conidiophores and coremium-like
fascicles are lacking in this species and the
conidia are much narrower, 2–4.5 µm (Chupp
1954, Vasudeva 1963). Several collections of
P. borreriae on Mitracarpus sp. and
Spermacoce spp. from Brazil and Cuba, now
deposited at HAL, have been examined. P.
hedyotis (S. Singh) B. Sutton on Hedyotis spp.
in India and Nepal is the third similar species,
which is also distinguished by lacking solitary
conidiophores and verruculose conidiophores
and conidia (Singh 1980, Sutton 1994).
Pseudocercospora costi (F. Stevens) U. Braun
& Crous
 Cercospora costi F. Stevens.
Material examined – VENEZUELA,
Miranda, Guatope Pk., on leaves of Costus sp.
(Costaceae), Feb. 1971, R. Urtiaga 1407 (IMI
156498); Lara, Villanueva, on leaves of Costus
sp., Nov. 2008, R. Urtiaga 135 (HAL 2518 F).
Notes – Listed from Venezuela in
Crous & Braun (2003) and recorded by Braun

Mycosphere Doi 10.5943/mycosphere/4/2/3
191
& Urtiaga (2012), but lacking in Iturriaga &
Minter (2006). The collection from 2008 is
very rich (superficial hyphae with solitary
conidioiphores developed, conidiophores 5–30
 2–5 µm, conidia 25–110  2–4 µm). Super-
ficial hyphae with solitary conidiophores have
also been observed in type material of this
species that has been re-examined (on Costus
sp., Panama, Gatun, 24 Aug. 1923, F.L.
Stevens 1343, ILL 15148). P. costina (Syd. &
P. Syd.) Deighton differs in having very broad,
strongly curved conidia (Deighton 1976).
Fig. 8 – Pseudocercospora coremioides. Based
on type material. a Superficial hyphae with
solitary conidiophores. b Conidiophore
fascicles. c Conidiophore tips. d Conidia. – Bar
= 10 µm.
Pseudocercospora eupatorii-formosani U.
Braun & Bagyan.
 Cercospora eupatorii-formosani
Sawada, nom. inval.
Material examined – VENEZUELA, La-
ra, Barquisimeto, zoological garden, on leaves
of Chromolaena odorata (L.) R.M. King & H.
Rob. (Asteraceae), Dec. 2009, R. Urtiaga 320
(HAL 2561 F).
Notes – New to Venezuela (Crous &
Braun 2003, Iturriaga & Minter 2006). Super-
ficial hyphae with solitary conidiophores may
be present or lacking in this species (Bagy-
anarayana & Braun 1999). In the new
collection from Venezuela, superficial myce-
lium has not been observed. The North
American Pseudocercospora eupatorii (Peck)
U. Braun & R.F. Castañeda is very similar and
confusable, but differs in having much broader,
robust conidiophores, 5–30  3–8 µm, and
longer conidia, up to 190 µm, with up to 14
septa (detailed descriptions, illustrations and
discussion in Bagyanarayana & Braun 1999).
Pseudocercospora euphorbiacearum U. Braun
= Cercospora mucunicola sensu Chupp
(1954: 226–227).
= Pseudocercospora mucunicola (Gonz.
Frag. & Cif.) Deighton, sensu Deighton (1976:
148).
Material examined – VENEZUELA, La-
ra, Rio Claro, La Cuchilla, on leaves of Dale-
champia scandens L. (Euphorbiaceae), Apr.
2009, R. Urtiaga 246 (HAL 2551 F), mixed
infection with Cercospora mucunicola.
Notes – The type host of Cercospora
mucunicola was original determined as
“Mucuna pruriens”, but later corrected to
Dalechampia scandens (Chupp 1954). Braun
(2003) re-examined type material deposited at
MA and found a mixed infection of a true
Cercospora s. str. and a Pseudocercospora,
confined the name C. mucunicola to the
Cercospora element by lectotypification and
introduced the new species Pseudocercospora
euphorbiacearum for the Pseudocercospora
involved. C. mucunicola is known from
Venezuela (Dennis 1970, Urtiaga 1986,
Iturriaga & Minter 2006). However, it is
unclear if these records refer to the Cercospora
or Pseudocercospora on Dalechampia or both
species. The present record is, in any case, the
first unequivocal record of P. euphorbiacearum
from Venezuela.
Pseudocercospora genipicola U. Braun &
Freire
Material examined – CUBA, Bayamo,
on leaves of Genipa americana L. (Rubiaceae),

Mycosphere Doi 10.5943/mycosphere/4/2/3
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18 Mar. 1968, R. Urtiaga 1215 (IMI 132561 =
K(M) 176138). VENEZUELA, Lara, Barqui-
simeto, zoological garden, on leaves of G.
americana, Jan. 2008, R. Urtiaga 103 (HAL
2530 F).
Notes – Pseudocercospora genipicola
was described from Brazil on Genipa ameri-
cana (Braun & Freire 2002). Caespituli in the
type material are mainly epiphyllous. Hypo-
phyllous colonies, which are often deviating
from epiphyllous ones in Pseudocercospora
species, are almost lacking in the type
collection from Brazil. The sample from Cuba
is characterized by its well-developed fungal
colonies on both sides of host leaves. Epi-
phyllous caespituli agree well with type
material, but hypophyllous colonies are dis-
tinct. The material from Venezuela agrees well
with type material from Brazil. An emended
description of P. genipicola based on type
material as well as sample from Cuba and
Venezuela is necessary:
Leaf spots amphigenous, subcircular to
angular-irregular, 2–10 mm diam., yellowish,
ochraceous, brownish, later whitish, above all
on the upper side of leaves, margin indefinite
or narrow and darker. Caespituli amphigenous,
distinctly punctiform on the upper leaf surface,
dark brown to blackish, more delicate and less
conspicuous below. Mycelium internal, occasi-
onally with a few superficial hyphae on the
lower leaf surface, sparingly branched, 1–3 µm
wide, septate, thin-walled, smooth, pale oli-
vaceous. Stromata well-developed, 10–100 µm
diam., immersed and larger on the upper side,
substomatal to intraepidermal and smaller
below, olivaceous-brown, composed of swollen
hyphal cells, 2–6 µm diam. Conidiophores in
small to very large, sporodochial fascicles,
loose to usually dense, arising from stromata,
erumpent or (on the lower side) emerging
through stomata, occasionally solitary, arising
from superficial hyphae, erect, straight, sub-
cylindrical to somewhat geniculate-sinuous,
unbranched, 5–40  2–5 µm, 0–2-septate, pale
olivaceous to olivaceous-brown, thin-walled,
smooth; conidiogenous cells integrated, either
terminal or conidiophores reduced to conidio-
genous cells, 5–25 µm long, conidiogenous loci
inconspicuous, neither thickened nor darkened.
Conidia solitary, obclavate-cylindrical, fusi-
form, small conidia sometimes ellipsoid-ovoid,
10–65  2.5–4.5 µm, (0–)1–6-septate, sub-
hyaline to pale olivaceous-brown, thin-walled,
smooth, apex obtuse to subacute, base truncate
to usually short obconically truncate, 1–2 µm
wide, hila unthickened, not darkened.
Pseudocercospora jatropharum (Speg.) U.
Braun
 Cercospora jatropharum Speg.
Material examined – CUBA, Bayamo, on
leaves of Jatropha sp. [as “integrifolia”]
(Euphorbiaceae), 1 Dec. 1966, R. Urtiaga (IMI
139309 = K(M) 176157); l.c., 23 Jul. 1966, R.
Urtiaga (IMI 120950).
Notes – Braun (2000) examined the poor,
almost exhausted type material of this species
(on Jatropha macrocarpa Griseb., LPS 943),
confirmed that it belongs to Pseudocercospora,
but nothing could be added to Chupp’s (1954)
description of this species. The present re-
description is based on K(M) 176157: Leaf
spots amphigenous, subcircular to angular-
irregular, 1–5 mm diam., brown or with dingy
grey center and brown border, somtimes vein-
limed. Caespituli amphigenous, mainly hypo-
phyllous, punctiform, scattered to dense, dark
brown. Mycelium internal; stromata 10–60 µm
diam., substomatal to erumpent, brown, cells
3–8 µm diam., thick-walled Conidiophores in
small to moderately large fascicles, loose to
usually dense, arising from stromata, emerging
through stomata, erect, straight, subcylindrical
to only slightly geniculate-sinuous, usually not
branched, 10–150 × 4–10 µm, continuous to
pluriseptate, olivaceous, olivaceous-brown or
pale brown, wall thin to slightly thickened,
smooth; conidiogenous cells integrated, usually
terminal, 10–30 µm long, conidiogenous loci
(scars) inconspicuous. Conidia solitary, ob-
clavate-subcylindrical, 30–100 × 4.5–9 µm, 1–
6-septate, usually pale olivaceous or brownish,
thin-walled, smooth, apex obtuse, base ob-
conically truncate, hila 1.5–2.5 µm wide, un-
thickened, not darkened.
Pseudocercospora jussiaeae (G.F. Atk.)
Deighton
Material examined – VENEZUELA, La-
ra, Rio Claro, La Cuchilla, on leaves of Lud-
wigia erecta (L.) H. Hara [ Jussiaea erecta
L.] (Onagraceae), Apr. 2009, R. Urtiaga 210
(HAL 2552 F).

Mycosphere Doi 10.5943/mycosphere/4/2/3
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Notes – This species is known from
Venezuela (Chupp 1954, Crous & Braun 2003,
Iturriaga & Minter 2006).
Pseudocercospora lippiae-albae U. Braun &
R.F. Castañeda
Material examined – CUBA, without
locality, on leaves of Lippia alba (Mill.) N.E.
Br. ex Britton & P. Wilson (Lamiaceae), 1966,
R. Urtiaga H 288/66 (IMI 120604 = K(M)
176124); Bayamo (Plant Pathology Labora-
tory), 15 Dec. 1965, R. Urtiaga (IMI 116895 =
K(M) 176123); Bayamo, 8 May 1967, R.
Urtiaga (IMI 127443 = K(M) 176122). VENE-
ZUELA, Lara, Barquisimeto, on leaves of
Lippia nodiflora L. ( Phyla nodiflora (L.)
Greene), Nov. 2006, R. Urtiaga (HAL 2170 F).
Notes – Pseudocercospora lippiae-
albae was described by Castañeda & Braun
(1989) from Cuba on Lippia alba. These
collections are additional records of this
species from Cuba and a first from Venezuela
on a new host species. P. lippiae-albae is also
known on Lippia alba (incl. L. geminata
Kunth) from Brazil (Braun & Freire 2002) and
Uruguay (material examined – Monte Video,
Atahualpa, as L. geminata, Sep. 1930, Herter,
Pl. Uruguayensis Exs. 1483, HBG, NY). The
conidia in the present Cuban collection are
somewhat longer than in all other specimens
that have been examined (up to 110 µm long,
with up to 11 septa).
Pseudocercospora lonchocarpicola U. Braun
& Urtiaga, sp. nov. Fig. 9
MycoBank, MB 801959.
P. lonchocarpi similis, sed conidiophoris
longioribus et latioribus, ad 200 × 10 µm, cel-
lulis conidiogenis sympodialiter proliferan-
tibus et conidiis brevioribus et latioribus, ad 80
× 10 µm.
Leaf spots lacking or only with diffuse
yellowish to brownish discolorations. Caes-
pituli hypophyllous, formed on diffuse dis-
colorations of the leaves, punctiform to effuse,
loose to dense, brown. Mycelium internal and
external, superficial; hyphae emerging through
stomata, sparingly branched, usually straight,
2–5 µm wide, septate, thin-walled, smooth,
subhyaline to olivaceous-brown. Stromata
lacking or only with aggregations of a few
swollen hyphal cells, small, substomatal to
intraepidermal, brown, about 10–20 µm diam.,
cells 4–10 µm diam., brown.
Fig. 9 – Pseudocercospora lonchocarpicola.
Based on type material. a Superficial hypha. b
Conidiophores arising from superficial hyphae
and swollen hyphal cells. c Conidiophore tips.
d Conidia. – Bar = 10 µm.
Conidiophores solitary, arising from superficial
hyphae, or in usually small and divergent,
occasionally large and denser fascicles, arising
from internal hyphae or aggregations of
swollen hyphal cells, emerging through sto-
mata or erumpent, erect, usually rather straight
to slightly geniculate-sinuous, subcylindrical to
subclavate, i.e. somewhat increasing in width
towards the apex, unbranched or branched,
above all near the apex, 30–200  5–10 µm,
pluriseptate, pale to medium dark brown or
paler towards the tip, wall smooth, up to 1.5
µm wide, above all below; conidiogenous cells
integrated, terminal, occasionally pleuro-
genous, 10–55 µm long, conidiogenous loci
neither thickened nor darkened. Conidia
solitary, obclavate-cylindrical, subclavate to
broadly fusoid, straight to distinctly curved,
(30–)40–70(–80)  (5–)6–9(–10) µm, 1–5-,
mostly 3-septate, subhyaline, pale olivaceous
to brownish, thin-walled, smooth or almost so,

Mycosphere Doi 10.5943/mycosphere/4/2/3
194
apex broadly rounded, base short obconically
truncate, 2–3 µm wide, hila unthickened, not
darkened.
Material examined – CUBA, Bayamo, on
leaves of Lonchocarpus domingensis (Turpin
ex Pers.) DC. (Fabaceae, Millettieae), 8 May
1967, R. Urtiaga 17 (IMI 127478 = K(M)
176151, holotype).
Notes – The genus Lonchocarpus
belongs to the Fabaceae tribe Millettieae (Silva
et al. 2012). Pseudocercospora lonchocarpi
(J.A. Stev.) Crous & M.P.S. Câmara, known
from Brazil, Guyana and Peru on various
Lonchocarpus spp. (Crous & Braun 2003), is
easily distinguishable from P. lonchocarpicola
by its much shorter and narrower conidio-
phores, percurrently proliferating conidio-
genous cells and much longer and narrower
finely verruculose conidia, (30–)50–100(–120)
 3–3.5(–4.5) µm with (1–)3–7(–13) septa
(Chupp 1954, Crous & Câmara 1998). Passa-
lora amazonica U. Braun (Braun 2003) is
another cercosporoid hyphomycete described
from Brazil on Lonchocarpus sp. The conidio-
genous loci of this species are, however,
thickened and darkened and the conidia are
formed in chains. A few additional species of
Pseudocercospora have been described on
hosts of allied genera of the Millettieae, but all
of them are morphologically quite distinct. P.
hardwarensis (Naras.) U. Braun & Crous on
Tephrosia purpurea (L.) Pers. in India and
Myanmar (Crous & Braun 2003) is well
characterized by having internal mycelium,
consistently fasciculate conidiophores and
narrower conidia, 20–80  3–6 µm (Vasudeva
1963; several Indian collections examined, IMI
155984, 180785, 264177 and 91412), P.
ichthyomethiae (Dearn. & Barthol.) U. Braun
& Crous on Piscidia piscipula (L.) Sarg. in
Cuba and Florida (Crous & Braun 2003) has
uniformly fasciculate, often verruculose
conidiophores with subdenticulate conidio-
genous loci and much smaller conidia, 20–55 
2–3.5 µm (Chupp 1954; type material, DAOM,
Dearness 5042, examined). P. millettiae Goh &
W.H. Hsieh (Hsieh & Goh 1990), only known
from Taiwan on Callerya reticulata (Benth.)
Schot ( Millettia reticulata Benth), possesses
consistently fasciculate conidiophores, per-
currently proliferating conidiogenous cells and
much narrower, cylindrical conidia, 28–90  3–
4.5 µm. P. vataireae (Henn.) U. Braun & F.O.
Freire (Braun & Freire 2002) on Derris spp. in
Brazil is characterized by having large stro-
mata, 10–150 µm diam., conidiophores with
monoblastic, determinate to mostly per-
currently, rarely sympodially proliferating co-
nidiogenous cells and narrower conidia, 30–90
 3–6 µm.
Fig. 10 – Pseudocercospora lonchocarpigena.
Based on type material. a Conidiophore
fascicles. b Conidiophores. c Conidia. – Bar =
10 µm.
Pseudocercospora lonchocarpigena U. Braun
& Urtiaga, sp. nov. Fig. 10
MycoBank, MB 801960.

Mycosphere Doi 10.5943/mycosphere/4/2/3
195
P. ichthyomethiae similis, sed stroma-
tibus formantibus, 20–80 µm diam., conidio-
phoris laevibus et cellulis conidiogenis non
denticulatis.
Leaf spots amphigenous, subcircular to
irregular, 3–25 mm diam., often at tips and
marginal, pale to medium brown, greyish
brown to dingy grey, margin indefinite or with
a very narrow darker marginal line. Mycelium
internal. Stromata well-developed, 20–80 µm
diam., olivaceous-brown, composed of swollen
hyphal cells, 1.5–5 µm diam., immersed or
substomatal on the lower leaf side. Caespituli
amphigenous, dark brown to blackish, punc-
tiform and conspicuous on the upper leaf
surface, less conspicuous and finer below.
Conidiophores numerous, in moderately large
to large, almost sporodochial fascicles, loose to
mostly dense, arising from stromata, erumpent
or emerging through stomata, erect, sub-
cylindrical or narrower towards the tip, straight
to somewhat geniculate-sinuous, unbranched,
5–25  1–3 µm, 0–1-septate, subhyaline to pale
olivaceous, thin-walled, smooth; conidiophores
usually reduced to conidiogenous cells or
occasionally integrated, terminal, 5–20 µm
long, conidiogenous loci inconspicuous, oc-
casionally visible as truncate tip or lateral
shoulder, but always unthickened and not
darkened. Conidia solitary, subcylindrical,
cylindrical-obclavate, 15–60  (1–)1.5–2.5(–3)
µm, 1–5-septate, subhyaline to very pale
olivaceous, tips obtuse to subacute, base
truncate to short obconically truncate, 1–2 µm
wide, hila neither thickened nor darkened.
Material examined – CUBA, Bayamo, on
leaves of Lonchocarpus longipes Urb. &
Ekman (Fabaceae, Millettieae), 29 May 1967,
R. Urtiaga M-566 (IMI 127924 = K(M)
176152, holotype).
Notes – Pseudocercospora lonchocarpi
differs from P. lonchocarpigena in having
percurrently proliferating conidiogenous cell
and finely verruculose, much longer and wider
conidia, (30–)50–100(–120)  3–3.5(–4.5) µm
(Crous & Câmara 1998). P. lonchocarpicola,
described above, is quite distinct by superficial
hyphae with solitary conidiophores and much
wider conidia. P. ichthyomethiae is the only
morphologically comparable species of Pseu-
docercospora on hosts of other genera
belonging to the Millettieae. However, large
stromata are lacking in the latter species, the
conidiophores are often verruculose and the
conidiogenous loci are subdenticulate (Chupp
1954, Crous & Braun 2003). All other species
– P. hardwarensis, P. millettiae and P.
vataireae – are quite distinct by their much
wider conidia (Braun & Freire 2002, Hsieh &
Goh 1990, Vasudeva 1963).
Pseudocercospora lythracearum (Heald &
F.A. Wolf) X.J. Liu & Y.L. Guo
 Cercospora lythracearum Heald &
F.A. Wolf.
Material examined – VENEZUELA, La-
ra, Barquisimeto, on leaves of Lagerstroemia
speciosa (L.) Pers. (Lythraceae), Dec.. 2009,
R. Urtiaga 323 (HAL 2570 F).
Notes – New to Venezuela (not listed in
Urtiaga 1986, Crous & Braun 2003 and
Iturriaga & Minter 2006).
Pseudocercospora mirandensis (Chupp) R.F.
Castañeda & U. Braun
Material examined – VENEZUELA, La-
ra, Dto. Moran, Santa Marta, on leaves of
Clidemia sp. (Melastomataceae), Apr. 2009, R.
Urtiaga 302 (HAL 2536 F).
Notes – First records of this species on
Clidemia hirta (L.) D. Don were published in
Braun & Urtiaga (2008).
Pseudocercospora mombin (Petr. & Cif.)
Deighton
 Cercospora mombin Petr. & Cif.
Material examined – VENEZUELA, La-
ra, Macuto, Bosque, on leaves of Spondias
mombin L. (Anacardiaceae), Dec. 2009, R.
Urtiaga 313 (HAL 2571 F).
Notes – This is an additional collection of
this species, which is known from Venezuela
(Crous & Braun 2003, Iturriaga & Minter
2006, Braun & Urtiaga 2008), from a new
region in this country.
Pseudocercospora musae (Zimm.) Deighton
 Cercospora musae Zimm.
Material examined – VENEZUELA, La-
ra, Villanueva, on leaves of Musa acuminata
Colla (Musaceae), Nov. 2008, R. Urtiaga 148
(HAL 2519 F).
Notes – This species is known from
Venezuela (Crous & Braun 2003), but M.

Mycosphere Doi 10.5943/mycosphere/4/2/3
196
acuminata is probably a new host for this
country. P. musae is not listed in Iturriaga &
Minter (2006).
The identification of Pseudocercospora
species on banana is not possible solely based
on symptoms, and a differentiation using
microscopical characters is also not easy.
Braun et al. (1999) discussed Pseudocer-
cospora on banana and provided a key to the
species. Further detailed discussions on banana
leaf spot diseases caused by Pseudocer-
cospora have been published by Crous &
Mourichon (2002) and Arzanlou et al. (2008)
in connection with molecular studies on this
group of species and in the latter case with the
introduction of several morphologically very
similar, but genetically clearly distinct new
species. However, a supplemented key to the
species concerned has not yet been published.
As such a key may be useful for microscopical
routine examinations and identification pur-
poses, the following supplemented key, based
on the key of Braun et al. (1999), has been
prepared:
Key to Pseudocercospora spp. on Musa spp.
1. Conidia broadly obclavate-cylindrical (-fusi-
form), (20–)40–75  (4–)5–8 µm
.................................... P. musicola U. Braun
1* Conidia narrower, 2–5(–6) µm ................ 2
2. Superficial hyphae with solitary conidio-
phores present; fasciculate conidiophores
very long, 30–120 µm and pluriseptate .........
P. musae-sapienti (A.K. Kar & M. Mandal)
U. Braun & Mouch.
2* Superficial hyphae and solitary conidio-
phores lacking ............................................ 3
3. Caespituli predominantly epiphyllous, sporo-
dochial, up to 100 µm wide, composed of
large stromata, up to 70 µm diam., and
numerous densely arranged short conidio-
phores, 10–25 µm long, sympodially or up to
4 times percurrently proliferating; conidia
subcylindrical, mostly 30–50  2.5–3 µm
........................... P. eumusae Crous & Mour.
3* Conidiophores fasciculate, large sporodo-
chia not formed, stromata smaller, percurrent
proliferations lacking, conidia longer if
cylindrical ................................................... 4
4. Conidia obclavate-cylindrical, (2–)2.5–5(–6)
µm wide ..................................................... 5
4* Conidia consistently cylindrical or
subcylindrical, narrow, 2–3 µm ................. 7
5. Conidiogenous loci visible as minute circles,
rim very slightly thickened and darkened, hila
with similar structure; conidia subhyaline ....
P. fijiensis (M. Morelet) Deighton (
Paracercospora fijiensis (M. Morelet)
Deighton)
5* Conidiogenous loci and hila neither
thickened nor darkened; conidia pale oliva-
ceous to brownish ....................................... 6
6. Conidiophores 0–1-septate, barely geniculate
.......................... P. musae (Zimm.) Deighton
6* Conidiophores 1–5-septate, usually pluri-
septate throughout and geniculate-sinuous ...
P. fengshanensis (T.Y. Lin & J.M. Yen) J.M.
Yen
7. Conidia very long, 80–120  2.5–4 µm
................... P. longispora Arzanlou & Crous
7* Conidia shorter, usually  100 µm, and
narrower, 2–3 µm ....................................... 8
8. Conidia (30–)60–70(–83) µm long
.................. P. assamensis Arzanlou & Crous
8* Conidia longer, (40–)78–95(–120) µm
................. P. indonesiana Arzanlou & Crous
Pseudocercospora nigricans (Cooke) Deigh-
ton
 Cercospora nigricans Cooke.
Material examined – VENEZUELA, La-
ra, Barquisimeto, on leaves of Senna tora (L.)
Roxb. (Fabaceae), Jan. 2009, R. Urtiaga 268
(HAL 2565 F).
Notes – Listed from Venezuela in Crous
& Braun (2003), but lacking in Iturriaga &
Minter (2006).
Pseudocercospora panacis (Thirum. & Chupp)
Y.L. Guo & X.J. Liu
 Cercospora panacis Thirum. & Chupp.
Material examined – VENEZUELA, La-
ra, Barquisimeto, on leaves of Polyscias
guilfoylei (W. Ball) L.H. Bailey (Araliaceae),
June 2008, R. Urtiaga 130 (HAL 2511 F).
Notes – New to Venezuela and new host
species (Crous & Braun 2003). P. panacis is
widespread on various hosts of the genus

Mycosphere Doi 10.5943/mycosphere/4/2/3
197
Polyscias in Africa, Asia, Australia and
Oceania (New Caledonia), but this species is
new on P. guilfoylei and new for South
America (Crous & Braun 2003). P. guilfoylei is
a common host of Pseudocercospora poly-
sciatis (S.H. Sun) J.M. Yen, known from
Africa [Mauritius], Asia, Oceania and West
Indies [Cuba] (Crous & Braun 2003). The latter
species is distinguished from P. panacis by
longer, frequently branched, pluriseptate coni-
diophores and the formation of superficial
hyphae with solitary conidiophores. Braun
(2003) discussed, described and illustrated
Pseudocercospora spp. on Polyscias spp. in
detail.
Pseudocercospora paulliniae U. Braun &
Urtiaga, sp. nov. Fig. 11
MycoBank, MB 801961.
Etymology – epithet derived from the
host genus, Paullinia.
Fig. 11 – Pseudocercospora pauliniae. Based
on type material. a Conidiophore fascicles. b
Conidiophores. c Conidia. – Bar = 10 µm.
Pseudocercosporae pometiae valde simi-
lis, sed conidiis angustioribus, 2–4 µm.
Leaf spots amphigenous, angular to
irregular, partly vein-limited, 2–10 mm diam.,
medium brown, margin indefinite, occasionally
with a diffuse yellowish halo. Caespituli
amphigenous, punctiform, dark brown to
blackish. Mycelium internal. Stromata on the
upper leaf side immersed, on the lower side
immersed to substomatal, occasionally some-
what erumpent, small to moderately large, 10–
60 µm diam. on the upper side, almost lacking
or smaller below, olivaceous-brown, cells 2–7
µm diam. Conidiophores in small, loose to
moderately large and dense fascicles, arising
from stromata, through stomata or erumpent,
erect, straight, subcylindrical-conical to some-
what geniculate-sinuous, unbranched, 5–50 ×
2.5–5 µm, often uniformly short (5–25 µm), 0–
3-septate, pale to medium olivaceous-brown,
thin-walled, smooth; conidiogenous cells inte-
grated, terminal or conidiophores reduced to
conidiogenous cells, 10–20 µm long, conidio-
genous loci inconspicuous. Conidia formed
singly, narrowly obclavate, shorter conidia
sometimes subcylindrical or cylindrical, 20–80
× 2–4 µm, (1–)3–7(–8)-septate, hyaline, sub-
hyaline, pale olivaceous to olivaceous-brown,
thin-walled, smooth, apex ± pointed in ob-
clavate conidia, subobtuse in cylindrical ones,
base unthickened, not darkened.
Material examined – CUBA, Bayamo, on
leaves of Paullinia fuscescens Kunth (Sapin-
daceae), 19 Apr. 1967, R. Urtiaga 22 (IMI
127210 = K(M) 176135, holotype).
Notes – The genus Paullinia belongs to
the family Sapindaceae subfam. Sapindoideae.
Some Pseudocercospora spp. on host of this
family are morphologically rather similar. P.
pometiae U. Braun & R.G. Shivas on Pometia
pinnata J.R. Forst. & G. Forst. in Vanuatu is
very similar, but has somewhat wider conidia,
3–5 µm (McTaggart et al. 2008). P. cupaniae
(Syd.) U. Braun & F.O. Freire on Cupania
guatemalensis (Turcz.) Radlk. in Costa Rica
(stromata larger, up to 100 µm, conidia only
15–55 µm long) and P. talisiae U. Braun &
F.O. Freire on Talisia esculenta (A.St.-Hil.)
Radlk. in Brazil (lesions quite distinct, conidia
partly in chains, shorter and wider, 15–55  3–
5 µm) are two additional species with obvious
morphological similarity (Chupp 1954, Braun

Mycosphere Doi 10.5943/mycosphere/4/2/3
198
& Freire 2002). P. mitteriana Goh & W.H.
Hsieh (= Cercospora mitteriana Syd.) on
Dodonaea viscosa Jacq. (subfam. Dodo-
naeoideae) in Asia is also similar, but the
longer conidia are distinctly linear and possess
obtuse tips (Chupp 1954, Hsieh & Goh 1990,
Crous & Braun 1996). P. dodonaeae Boesew.
on the same host in New Zealand has quite
distinct lesions and pale brown conidia up to
150 µm long (Crous & Braun 1996). All other
Pseudocercospora spp. on hosts of the
Sapindaceae subgen. Sapindoideae are mor-
phologically easily distinguishable: P. allophyli
(Hansf.) Deighton on Allophylus spp. [with
superficial mycelium and solitary conidio-
phores, fascicles lacking], P. allophylina Goh
& W.H. Hsieh on Allophylus timorensis (DC.)
Blume, Taiwan [conidiophores up to 500 µm
long], P. allophylorum (A.K. Kar & M.
Mandal) Bagyan., U. Braun & Jagad. (= P.
allophylicola Deighton) on Allophylus cobbe
(L.) Raeusch. and A. africanus P. Beauv. in
Asia and Africa [conidiophores very long, up
to 150 µm], P. deinbolliae Deighton on
Deinbollia spp. in Africa [lesions distinct,
conidia ± cylindrical, up to 130 µm long], P.
nephelii B. Sutton & Peregrine on Nephelium
lappaceum L. in Brunei [conidia ± cylindrical,
verruculose, 60–85 µm long], P. sapindi-
emarginati (T.S. Ramakr. & K. Ramakr.) U.
Braun, Bagyan. & Jagad. on Sapindus spp. and
Koelreuteria paniculata Laxm. [conidia 3–7
µm wide], P. thouiniae (F. Stevens) U. Braun
& Crous on Thouinia stricta Radlk., Puerto
Rico and Virgin Islands [conidiophores very
long, up to 500 µm, conidia 5–8 µm wide], and
P. zambalesica (Petr.) U. Braun on Allophylus
macrostachys Radlk. [with superficial hyphae
and solitary conidiophores] (Chupp 1954,
Deighton 1987, Hsieh & Goh 1990, Guo &
Hsieh 1995, Braun 1999, Crous & Braun 1996,
Peregrine et al. 2000, Braun & Freire 2002,
McTaggart et al. 2008). Cercospora sapindi
F.L. Tai is a genuine species of Cercospora s.
str. (Guo et al. 2005). The generic affinity of
Cercospora sapindi Obreg.-Bot. on Sapindus
saponaria L., known from Colomia and
Panama, is unclear (type material not
available), but according to olivaceous conidia
this species seems to belong in Pseudo-
cercospora. The later species differs from P.
paulliniae in having quite distinct lesions,
conidiophores with hyaline tips and conidia
with obtuse apex (Chupp 1954, Crous & Braun
2003). Furthermore, there are two comparable
species on maple (Sapindaceae, subfam. Hip-
pocastanoideae): P. acericola (Woron.) Y.L.
Guo & X.J. Liu on Acer spp. in Asia and
Europe [stromata absent, external hyphae with
solitary conidiophores developed] and P.
sphendamnophila Kirschner on Acer oblongum
Wall. ex DC. (incl. A. albo-purpurascens
Hayata) in Taiwan [conidia ± cylindrical, only
2 µm wide] (Chupp 1954, Guo & Hsieh 1995,
Kirschner et al. 2009).
Fig. 12 – Pseudocercospora picramniae. Based
on type material. a Superficial hyphae. b
Solitary conidiophores arising from a super-
ficial hypha. c Conidiophore fascicles. d
Conidiophore. e Conidia. – Bar = 10 µm.
Pseudocercospora picramniae U. Braun &
Urtiaga, sp. nov. Fig. 12
MycoBank, MB 801963.
Etymology – epithet derived from the
host genus, Picramnia.

Mycosphere Doi 10.5943/mycosphere/4/2/3
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Differt a paene omnibus speciebus Pseu-
docercosporae ad Simaroubaceas hyphis super-
ficialibus cum conidiophoris solitariis evolutis
et a P. bruceae (cum hyphis superficialis)
stromatibus evolutis, conidiophoris fasciculatis
et conidiis angustioribus, 2–4.5 µm.
Leaf spots amphigenous, subcircular to
angular-irregular, 1–20 mm diam., ochraceous,
pale to medium dark brown, later greyish
brown to dingy greyish white, margin in-
definite or with a narrow darker border,
occasionally somewhat raised, sometimes with
a narrow, dark publish to violet halo. Myce-
lium internal and external; superficial hyphae
emerging through stomata, branched, septate,
1–3.5 µm wide, thin-walled, smooth or almost
so, subhyaline to olivaceous-brown. Stromata
lacking to well-developed, immersed to sub-
stomatal, 10–60 µm diam., medium brown or
olivaceous-brown, cells 2–7 µm diam. Coni-
diophores solitary, arising from superficial
hyphae, lateral, or in small, loose to large and
dense fascicles, arising from internal hyphae or
stromata, through stomata or erumpent, erect,
straight, subcylindrical-conical to distinctly
geniculate-sinuous, simple or often irregularly
branched, 5–60  2–5 µm, 0–4-septate, pale to
medium olivaceous-brown or brown, thin-
walled, smooth; conidiogenous cells integrated,
terminal, 5–25 µm long; conidiogenous loci
inconspicuous to subdenticulate, but always
unthickened and not darkened. Conidia soli-
tary, narrowly obclavate to obclavate-cylin-
drical, 18–85  2–4.5 µm, 1–8-septate, sub-
hyaline to pale olivaceous-brown, smooth or
almost so, apex obtuse, occasionally subacute,
base short obconically truncate, 1–1.5(–2) µm
wide, hila neither thickened nor darkened.
Material examined – CUBA, Bayamo, on
leaves of Picramnia pentandra Sw. (Picram-
niaceae), 12 Apr. 1967, R. Urtiaga (IMI
126873 = K(M) 176133, holotype).
Notes – This is the first species of
Pseudocercospora on a host of the family
Picramniaceae. Previously Picramnia was
assigned to the family Simaroubaceae. P.
bruceae (Petch) Y.L. Guo & X.J. Liu on
Brucea spp. in Asia is the only species of
Pseudocercospora on a host belonging to the
latter family with formation of superficial
mycelium with solitary conidiophores, but it
differs from P. picramniae in having broader
conidia, 3–5.5 µm. Furthermore, stromata and
fasciculate conidiophores are lacking in this
species (Chupp 1954, Guo & Hsieh 1995). All
other Pseudocercospora spp. on hosts of the
Simaroubaceae are distinguished from P.
picramniae by lacking superficial mycelium
and consistently fasciculate conidiophores [P.
ailanthicola (Patwardhan) Deighton, P.
ailanthigena H.S. Rao, Archana Singh &
Kamal, P. quinlingensis Y.L. Guo, P. sima-
roubae U. Braun & F.O. Freire (Chupp 1954,
Guo & Hsieh 1995, Rao et al. 1996, Liu et al.
1998, Braun & Freire 2002)].
Fig. 13 – Pseudocercospora psidii var. varians.
Based on type material. a Hypha. b Solitary
conidiophores arising from superficial hyphae.
c Conidiophore and hyphae emerging through
stomata. d Conidiophore fascicle. e Conidia. –
Bar = 10 µm.

Mycosphere Doi 10.5943/mycosphere/4/2/3
200
Pseudocercospora piperis (Pat.) Deighton
 Cercospora piperis Pat.
Material examined – VENEZUELA, La-
ra, Villanueva, on leaves of Piper sp.
(Piperaceae), Nov. 2008, R. Urtiaga 134 (HAL
2516 F).
Notes – This species is known from
Venezuela (Chupp 1954, Crous & Braun
2003).
Pseudocercospora psidii (Rangel) R.F.
Castañeda & U. Braun var. varians U. Braun &
Urtiaga var. nov. Fig. 13
MycoBank, MB 801967.
Differt a var. psidii hyphis superficialibus
cum conidiophoris solitariis evolutis, hypo-
phyllis.
Distinct from var. psidii by forming
superficial hyphae with solitary conidiophores
on the lower leaf surface.
Etymology – referring to the variable
fructification ranging from solitary conidio-
phores arising from superficial hyphae to
fascicles of conidiophores.
Material examined – VENEZUELA, La-
ra, Barquisimeto, market, on leaves of Psidium
guajava L. (Myrtaceae), Mar. 2012, R. Urtiaga
460 (HAL 2510 F, holotype); Lara, Barqui-
simeto, zoological garden, Jan. 2010, R.
Urtiaga 315 (HAL 2556 F). INDIA, Andhra
Pradesh, Yellore, Mahabubnagar, Jan. 1992,
Bagyanarayana & Jagadeeswar (HAL, s.n.).
Notes – Pseudocercospora psidii s. lat. is
redescribed as follows: Leaf spots amphi-
genous, subcircular, angular-iregular to some-
what diffuse, 1–7 mm diam or confluent and
larger, brown, greyish brown, reddish brown to
dingy grey or greyish white, margin indefinite
or narrow, dark brown, blackish or purple,
occasionally vein-limited. Caespituli amphi-
genous, punctiform, dark brown to blackish,
distinct or not very conspicuous. Mycelium
internal, external hyphae lacking (var. psidii) or
internal and external, with well-developed
superficial hyphae on the lower leaf surface
(var. varians), branched, 1.5–4 µm wide,
subhyaline to pale olivaceous or olivaceous-
brown, thin-walled, smooth; stromata lacking
to well-developed, 10–50 µm diam., sub-
globose to somewhat irregular, immersed or
substomatal, medium to dark brown, cells 2–5
µm diam. Conidiophores strictly in small, loose
to moderately large and dense fascicles, arising
from internal hyphae or stromata, erumpent or
through stomata (var. psidii), or also with
solitary conidiophores, arising from superficial
hyphae, lateral, rarely terminal (var. varians),
erect, straight, subcylindrical-conical to geni-
culate-sinuous, mostly not branched, 5–45 
(2–)2.5–5(–6) µm, 0–3-septate, pale olivaceous
to medium olivaceous-brown, thin-walled,
smooth; conidiogenous cells integrated, termi-
nal or conidiophores reduced to conidiogenous
cells, 5–20 µm long, conidiogenous loci
inconspicuous or occasionally subdenticulate.
Conidia solitary narrowly obclavate-cylin-
drical, (15–)20–70(–80)  (1.5–)2–3.5(–4) µm,
(1–)3–7(–9)-septate, subhyaline to pale oliva-
ceous, thin-walled, smooth, apex obtuse to
subacute, base obconically truncate, 1–1.5(–2)
µm wide, hila neither thickened nor darkened.
Pseudocercospora psidii is not listed
from Venezuela in Crous & Braun (2003).
Abundant superficial hyphae with solitary
conidiophores are formed in the present
collection of P. psidii from Venezuela. In the
type material of this species from Brazil, in a
collection on common guava from Cuba
(Castañeda & U. Braun 1989), in Chinese
samples described by Guo & Hsieh (1995), and
in additional samples examined by Crous
(1999), superficial mycelium is absent and
conidiophores are only formed in fascicles
arising from stromata. Sarbajna & Chatto-
padhyay (1991) recorded an Indian collection
on P. guajava and described and illustrated
superficial hyphae with solitary conidiophores.
Due to the formation of superficial hyphae and
solitary conidiophores, Bagyanarayana et al
(1995) identified an Indian collection on
common guava (Andhra Pradesh, Yellore,
Mahabubnagar, Jan. 1992, Bagyanarayana &
Jagadeeswar, HAL, s.n.) as Pseudocercospora
sawadae (W. Yamam.) Goh & W.H. Hsieh
(record also cited in Kamal 2010), which is,
however, a misidentification. This collection
and probably most or even all Indian col-
lections referred to as P. sawadae seem to
belong to P. psidii and represent sampes with
superficial mycelium. Such deviating collec-
tions with hypophyllous fructification, super-
ficial mycelium and solitary conidiophores are
described as a new variety of P. psidii.
Absence or presence of superficial mycelium

Mycosphere Doi 10.5943/mycosphere/4/2/3
201
and solitary conidiophores render the differen-
tiation between P. psidii and P. sawadae
difficult and cause confusion. The true P. sa-
wadae is distingished from P. psidii by the
following characters: distinct leaf spots,
stromata and well-developed fascicles of
conidiophores lacking, conidia wider, 3–5 µm,
and hila wider, 1.5–2 µm [conidia (1.5–)2–
3.5(–4) µm and hila 1–1.5 µm wide in P.
psidii]. The present redescription of P. psidii,
which is based on collections from Cuba, India
and Venezuela, shall help to avoid further
confusion.
Pseudocercospora rhinocarpi U. Braun &
Crous
 Cercospora rhinocarpi Chupp & A.S.
Mull., nom. inval.
Material examined – VENEZUELA, La-
ra, Macuto, Bosque, on leaves of Anacardium
excelsum (Bertero ex Kunth) Skeels [ A.
rhinocarpus DC., nom. illeg.] (Anacar-
diaceae), Dec. 2009, R. Urtiaga 310 (HAL
2557 F).
Notes – The original description of this
species, published by Braun and Crous in
Crous & Braun (2003), was based on type
material from Venezuela (CUP-VZ 2047). This
is the second collection of this species.
Pseudocercospora riachuelii (Speg.) Deighton
var. riachuelii
Material examined – VENEZUELA, La-
ra, Rio Claro, Cuchilla, on leaves of Cissus
verticillata (L.) Nicolson & C.E. Jarvis [= C.
sicyoides L.] (Vitaceae), Apr. 2009, R. Urtiaga
203 (HAL 2550); l.c., June 2010, R. Urtiaga
379 (HAL 2525 F).
Notes – Known from Venezuela (Urtiaga
1986, Crous & Braun 2003). These collections
belong to the typical variety characterized by
lacking superficial mycelium.
Pseudocercospora solanacea U. Braun, sp.
nov.
MycoBank, MB 493087.
 Pseudocercosporella solanacea B.K.
Gupta & Kamal, Indian Phytopathol. 42: 391,
“1989” 1990, nom. inval. (Art. 37.6).
 Pseudocercospora solanacea (B.K.
Gupta & Kamal) U. Braun, A monograph of
Cercosporella, Ramularia and allied genera
(phytopathogenic hyphomycetes), Vol. 1: 198,
1995 [as P. solanacea B.K. Gupta & Kamal ex
U. Braun, in MycoBank].
Latin description – Gupta & Kamal,
Indian Phytopathol. 42: 391, “1989” 1990.
Material examined – INDIA, U.P.,
Gorakhpur, on Solanum nigrum L. (Solana-
ceae), Kamal, KS 150 (IMI 238212, holotype,
now K[M]).
Fig. 14 – Pseudocercospora trichophila var.
punctata (= Cercospora solanacea). Based on
CUP-VZ 2927. a Hyphae. b Solitary coni-
diophores arising from superficial hyphae. c
Conidiophore fascicles. d Conidiophores. e
Conidia. – Bar = 10 µm.

Mycosphere Doi 10.5943/mycosphere/4/2/3
202
Notes – Gupta & Kamal (1990) described
Pseudocercosporella solanacea, cited a single
collection (IMI 237162), but failed to indicate
“holotype” (or “type”), which is necessary
since 1990 for genera and taxa below this rank
in order to be validly published [in the original
publication of Gupta & Kamal, “1989” was
given as year, but it was effectively issued in
1990]. Braun (1995b) examined the latter
material, introduced the combination Pseudo-
cercospora solanacea (B.K. Gupta & Kamal)
U. Braun and cited IMI 237162 as holotype.
With regard to the validity of the latter name,
there is a discrepancy between Index Fun-
gorum, in which this combination is classified
to be invalid, and MycoBank, in which this
name is corrected to “Pseudocercospora sola-
nacea” cited as B.K. Gupta & Kamal ex U.
Braun” and classified as “orthographic variant”
and valid name. A formal validation of this
name was not performed by Braun (1995b)
since it was unknown at that time that this
species had not been published in 1989. In
order to clarify the confused nomenclatural
situation, the latter name is formally validated
once more. Pseudocercosporella leonotidis
B.K. Gupta & Kamal is another invalid name
published by Gupta & Kamal (1990). Braun
(1995b) re-examined potential type material of
this species and published a full description
and new illustration, but failed to validate this
name formally:
Pseudocercosporella leonotidis B.K. Gupta &
Kamal ex U. Braun, sp. nov.
MycoBank, MB 125145.
 Pseudocercosporella leonotidis B.K.
Gupta & Kamal, Indian Phytopathol. 42: 389,
“1989” 1990, nom. inval. (Art. 37.6).
Latin and English description: Gupta &
Kamal (1990: 389).
Material examined – INDIA, U.P.,
Gorakhpur, on Leonotis nepetifolia R. Br.
(Solanaceae), Kamal, KS 84 (IMI 237162,
holotype, now K[M]).
The name Pseudocercospora solanacea
influences the nomenclature of Cercospora
solanacea Sacc. & Berl. This name has been
misinterpreted for a long time due to Chupp’s
(1954) insufficient description and interpre-
tation. Unfortunately type material of C. sola-
nacea, deposited at PAD, is currently not
available for a re-examination. However, sev-
eral collections determined by Chupp as C.
solanacea, deposited at CUP, have been exa-
mined. Chupp (1954) cited type material of this
species. It is not quite clear but possible that he
had examined and compared this collection.
Chupp’s specimens, including a sample from
Venezuela, have been examined. C. solanacea
proved to be a species of the genus Pseudo-
cercospora (conidiogenous loci and hila of the
conidia neither thickened nor darkened), con-
specific with P. trichophila (F. Stevens)
Deighton. Braun & Urtiaga (2012) introduced
P. trichophila var. punctata U. Braun &
Urtiaga for collections with epiphyllous puncti-
form caespitili and well-developed stromata.
This variety is identical with C. solanacea, i.e.
the latter name must be reduced to synonymy
with P. trichophila var. punctata. C. solanacea
is older than C. trichophila, but its epithet is
blocked under Pseudocercospora:
Pseudocercospora trichophila var. punctata
U. Braun & Urtiaga
= Cercospora solanacea Sacc. & Berl.,
Atti Reale Ist. Veneto Sci. Lett. Arti VI, 3: 721,
1885, syn. nov.
Material examined – VENEZUELA, road La
Guira–Caracas, on leaves of Solanum
erianthum var. adulterinum (Ham. ex G. Don)
R.E.D. Baker & N.W. Simmonds [= S. hazenii
Britton] (Solanaceae), 3 Mar. 1939, A.S.
Muller & H.H. Whetzel (CUP-VZ 2927).
PUERTO RICO, Rio Pietras, on S. erianthum
D. Don (= S. verbascifolium L.), 7 Jan. 1917,
J.A. Stevenson (CUP 41246). TRINIDAD,
North Coast Road, on S. erianthum, 15 Nov.
1947, R.E.D. Baker (CUP 37506). USA,
Florida, Miami, on S. erianthum, 4 Feb. 1921,
J.A. Stevenson (CUP 41247).
The examined collections of P. tricho-
phila var. punctata (= C. solanacea) from CUP
are characterized as follows: Leaf spots
amphigenous, conspicuous on the upper leaf
side, less conspicuous below, angular-irregular,
1–6 mm diam. or confluent and larger, pale to
medium dark brown, reddish brown to almost
black, often vein-limited. Caespituli amphi-
genous, conspicuously punctiform on the upper
leaf surface, dark bown to blackish, incon-
spicuous below. Mycelium internal and ex-
ternal; hyphae subhyaline to pale olivaceous,
1.5–5 µm wide. Stromata epiphyllous, 20–60

Mycosphere Doi 10.5943/mycosphere/4/2/3
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µm diam., subcircular to somewhat irregular in
outline, brown. Conidiophores on the upper
side in small to moderately large fascicles,
loose to dense, arising from stromata, erum-
pent, erect, or solitary, arising from superficial
hyphae, straight, subcylindrical or somewhat
attenuated towards the tip to moderately
geniculate-sinuous, usually unbranched, 10–60
× 3–7 µm, 0–3-septate, pale to medium oli-
vaceous or olivaceous-brown, darker in mass,
thin-walled, smooth; conidiogenous cells inte-
grated, terminal, 10–25 µm long, conidio-
genous loci inconspicuous, occasionally sub-
denticulate. Conidia cylindrical or obclavate-
cylindrical, 20–60 × 4–6 µm, 1–6-sptate,
subhyaline to pale olivaceous, thin-walled,
smooth, apex obtuse to subacute, base short
obconically truncate, hila 1–2 µm wide, neither
thickened nor darkened.
Records of C. solanacea from India on
Solanum melongena L. (material examined:
BPI 441388 and CUP s.n., Herb. Crypt. Ind.
Orient. Exs., III. Indian Cercosporae, Fasc. 1,
No. 41) and S. nigrum L. (material examined:
BPI 457102 and CUP s.n., Herb. Crypt. Ind.
Orient. Exs., III. Indian Cercosporae, Fasc. 1,
No. 42) are based on misidentifications and
must be corrected to Paracercospora egenula
(Syd.) Deighton ( Pseudocercospora egenula
(Syd.) U. Braun & Crous) and Pseudocerco-
spora atromarginalis (G.F. Atk.) Deighton,
respectively. A collection on Solanum eri-
anthum (= S. verbascifolium) from Taiwan
(material examined: Taipeh, 26 Oct. 1933, W.
Yamamoto ex herb. Univ. Imp. Taihokuensis,
BPI 441389), determined as C. solanacea,
proved to be Pseudocercospora trichophila
var. trichophila as already stated by Hsieh &
Goh (1990), based on a duplicate of this
specimen deposited at NTU-PPE.
An additional specimen recently col-
lected in Venezuela [Rio Claro, Cuchilla, on
leaves of Solanum erianthum, Apr. 2009, R.
Urtiaga 197 (HAL 2541 F)] proved to be P.
trichophila var. trichophila showing that both
varieties may occur on the same host species in
the same region.
Pseudocercospora struthanthi U. Braun, F.O.
Freire & N. Pons
 Cercospora struthanthi Chupp & A.S.
Mull., nom. inval.
Material examined – VENEZUELA, La-
ra, Las Brujitas, on leaves of Phthirusa stelis
(L.) Kuijt [incl. P. anastyla Rizz.] (Loran-
thaceae), Apr. 2010, R. Urtiaga 124 (HAL
2512 F).
Notes – Phthirusa stelis is a new host
species. The description of Pseudocercospora
struthanthi was based on type material of the
invalid Cercospora struthanthi (on Struth-
anthus sp. from Venezuela) and a collection
from Brazil, also on Struthanthus sp. (Braun &
Freire 2002). Braun & Freire (2004) added new
Brazilian specimens on Phoradendron sp.
(Viscaceae) and Struthanthus sp. (distributed as
“U. Braun, Fungi sel. exs. 27”), and Braun et
al. (2010) published a record of this species
from Brazil on Tripodanthus sp. (also Loran-
thaceae). Phthirusa stelis has numerous
synonyms in Loranthus as well as Struth-
anthus. It is possible that the type material
(Struthanthus sp.) represents a species of
Phthirusa. The present material agrees very
well with type material and the original
description (conidia 15–65  2.5–4.5 µm),
except for a few superficial hyphae formed on
the leaf surface and many germinated conidia,
partly with microcyclic conidiogenesis.
Pseudocercospora taichungensis Goh & W.H.
Hsieh Fig. 15
Leaf spots amphigenous, subcircular to
somewhat angular-irregular, about 5–20 mm
diam., often zonate, pale to medium brown or
greyish brown, with a narrow dark border or
marginal line, dark brown to blackish.
Caespituli amphigenous, mainly hypophyllous,
distinctly punctiform, dark brown to blackish.
Mycelium internal; hyphae branched, 1.5–2.5
µm wide, septate, subhyaline or pale, thin-
walled, smooth. Stromata well-developed, sub-
stomatal to intraepidermal, erumpent, sub-
circular to somewhat irregular in outline, 20–
80 µm diam., olivaceous-brown to brown, cells
1.5–6 µm diam., outline circular to angular.
Conidiophores very numerous, in dense, almost
sporodochial fascicles, arising from stromata,
erect, straight, subcylindrical-conical to slighty
geniculate, unbranched, 5–25  1.5–3.5 µm,
0(–1)-septate, subhyaline to very pale oliva-
ceous, medium olivaceous in mass, thin-
walled, smooth; conidiophores usually reduced
to conidiogenous cells, conidiogenous loci

Mycosphere Doi 10.5943/mycosphere/4/2/3
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inconspicuous. Conidia obclavate-cylindrical,
15–45  2–3 µm, 0–4-septate, subhyaline to
very pale olivaceous, thin-walled, smooth, apex
acute to subobtuse, base truncate to usually
short obconically truncate, 1–1.5(–2) µm wide,
hila neither thickened nor darkened.
Material examined – VENEZUELA, La-
ra; Barquisimeto, zoological garden, on leaves
of Senna atomaria (L.) H.S. Irwin & Barneby
(Fabaceae), Jan. 2010, R. Urtiaga 312 (HAL
2559 F).
Fig. 15 – Pseudocercospora taichungensis.
Based on HAL 2559 F. a Conidiophore
fascicles. b Conidiophores. c Conidia. – Bar =
10 µm.
Notes – Several species of Pseudo-
cercospora have been described on hosts
belonging to Cassia and Senna, and have been
discussed, illustrated and keyed out in Braun
(1989). P. cassiigena (J.M. Yen & Lim) Yen
on Senna alata (L.) Roxb. in Singapore (Yen &
Lim 1980) is the only comparable, morpho-
logically similar species treated in Braun
(1989), but differs from the present collection
from Venezuela in having quite distinct leaf
spots (angular-irregular, vein-limited, brown,
not zonate, only 1–4 mm diam), indistinct
caepituli, smaller stromata (25–40 µm diam.),
pale brown conidiophores, only 6–11 µm long,
and pale olivaceous-brown conidia. Hsieh &
Goh (1990) described Pseudocercospora tai-
chungensis on Cassia fistula from Taiwan.
Caespituli (conidiomata), conidiophores and
conidia of the collections on Senna atomaria
from Venezuela agree well with P. taichun-
gensis, except for much larger leaf spots. In
spite of distinct lesions, different host species
and localities in Asia and South America,
respectively, this collection is tentatively
assigned to the latter species.
Pseudocercospora teramnicola U. Braun &
Urtiaga, sp. nov. Fig. 16
MycoBank, MB 801968.
Etymology: epithet derived from the host
genus, Teramnus.
Pseudocercosporae calopogonii et P. pue-
rariicolae similis, sed lesionibus valde distinctis
et hyphis superficialibus cum conidiophoris
solitariis evolutis.
Lesions amphigenous, formed as small
angular-irregular darker discolorations mainly
caused by the fructification. Colonies amphi-
genous, punctiform to subeffuse, dark brown to
blackish. Mycelium internal and external;
superficial hyphae branched, septate, 2–5 µm
wide, subhyaline to medium brown, thin-
walled, smooth; stromata lacking to well-
developed, immersed to substomatal, 10–60
µm diam. or sometimes even larger, brown,
cells 2–6 µm wide. Conidiophores in small to
large, loose to dense fascicles, arising from
internal hyphal or stromata, or associated with
immature ascomata (initials), sometimes
arising from them, or conidiophores solitary,
arising from superficial hyphae, lateral, erect to
decumbent, shape and size variable, straight,

Mycosphere Doi 10.5943/mycosphere/4/2/3
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cylindrical-conical to strongly geniculate-
sinuous, usually unbranched, 5–110  3–5 µm,
continuous to pluriseptate, pale to medium
olivaceous-brown or brown throughout or paler
towards the tip, thin-walled, smooth; conidio-
genous cells intergrated, terminal or inter-
calary, 5–30 µm long, conidiogenous loci
inconspicuous to subdenticulate, but wall of the
loci always unthickened and not darkened.
Conidia solitary, obclavate(-cylindrical), short
conidia sometimes cylindrical-ellipsoid or
almost obovoid, 20–80  2.5–5 µm, 1–7-
septate, subhyaline to pale olivaceous, apex
obtuse to subacute, base short obconically
truncate, 1–2 µm wide, hila neither thickened
nor darkened.
Material examined – CUBA, Bayamo, on
leaves of Teramnus labialis (L. f.) Spreng.
(Fabaceae), Dec. 1967, R. Urtiaga 1066 (IMI
131169 = K(M) 176148, holotype).
Notes – The genus Teramnus belongs to
the phaseoloid legumes (tribe Phaseoleae), and
is, according to molecular phylogenetic studies
(Lee & Hymowitz 2001, Stefanović et al.
2009), part of subtribe Glycininae. Cercospora
teramnicola Ragun. & K. Ramakr. (Ragu-
nathan & Ramakrishnan 1965) is insufficiently
known, and type material of this species could
not be traced. However, based on the original
description and illustration, the latter species is
in any case not identical with the new
Pseudocercospora on this host from Cuba
(stromata lacking or almost so, superficial
mycelium not developed, conidiophores very
long, about 70–150 µm, often branched,
conidia 12–45  3–6 µm, subhyaline). The
conidiogenous loci were described to be
prominent, which was used in this paper in
various other cases as circumscription for
thickened and darkened scars of true
Cercospora species, whereas the loci in former
Cercospora spp. which now belong to
Pseudocercospora were described to be
indistinct or not described at all. Hence, C.
teramnicola is undoubtedly a true Cercospora
or maybe a Passalora due to small obclavate
conidia. Most Pseudocercospora species on
hosts of allied genera are easily distinguishable
from P. teramnicola by lacking superficial
hyphae and some additional differences: P.
boringuensis (E. Young) Deighton on Calo-
pogonium spp. (conidiophores up to 200 µm
long, conidia 30–55  4–7 µm), P. calopogonii
(F. Stevens & Solh.) Deighton on Calo-
pogonium spp. (conidiophores and conidia
rather similar, lesions quite distinct), P. cruenta
(Sacc.) Deighton (also recorded on Calo-
pogonium, Glycine and Pueraria; conidio-
phores shorter, lesions quite distinct), P.
glycines (Cooke) Deighton on Glycine spp.
(numerous, short, densely arranged conidio-
phores arising from large stromata in
sporodochial conidiomata, conidia only 1.5–3
µm wide), P. monoica (Ellis & Everh.)
Deighton on Amphicarpaea spp. (also with
sporodochial conidiomata), P. pachyrrhizi
Sawada & Katsuku on Pachyrhizus erosus (L.)
Urb. (similar to P. glycines, but conidia only
1.5–2.5 µm wide), P. puerariae (Syd. & P.
Syd.) Deighton (conidiophores up to 350 µm
long, conidia 4–9 µm wide), P. puerariicola
(W. Yamam.) Deighton on Pueraria spp.
(lesions quite distinct) [Chupp 1954, Deighton
1976, Yen & Lim 1980, Hsieh & Goh 1990,
Guo & Hsieh 1995].
Fig. 16 – Pseudocercospora teramnicola.
Based on type material. a Hypha. b Solitary
conidiophore arising from superficial hyphae. c
Conidiophore fascicles. d Conidiophore
fascicles. e Conidia. – Bar = 10 µm.

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P. puerariina (J.M. Yen) Y.M. Yen on
Pueraria phaseoloides (Roxb.) Benth. in
Singapore is characterized by having well-
developed superficial mycelium with solitary
conidiophores, but stromata are lacking and the
acicular-obclavate conidia are 90–200 µm long.
Among more distantly related Pseudocerco-
spora species on hosts of Phaseolus, Vigna and
allied genera, there are a few with superficial
hyphae and solitary conidiophores, but all of
them are clealy distinct from P. teramnicola.
Stromata and fasciculate conidiophores are
absent in P. phaseolicola Goh & W.H. Hsieh,
fascicles of conidiophores are formed in P.
shihmensis (J.M. Yen) J.M. Yen but stromata
are lacking, P. vignicola (J.-M. Yen, A.K. Kar
& B.K. Das) U. Braun has short conidiophores
and narrowly cylindrical-filiform conidia, and
P. vignigena J.M. Yen, A.K. Kar & B.K. Das is
characterized by its cylindrical conidia, 4–6 µm
wide (Yen & Lim 1980, Yen et al. 1982a,
Hsieh & Goh 1990, Braun et al 1999). The
latter authors provided a key to Pseudo-
cercospora spp. on Phaseolus and allied
genera.
Pseudocercospora trichiliae-hirtae U. Braun
& Urtiaga, sp. nov. Fig. 17
MycoBank, MB 801969.
Etymology: name derived from the host
species, Trichilia hirta.
Pseudocercosporae swieteniae valde si-
milis, sed maculis foliorum indistinctis, coni-
diophoris saepe ramosis, conidiis latioribus,
(2.5–)3–5(–5.5) µm, hilis latioribus, 1.5–2 µm.
Differt a P. amooriae et P. meliicola maculis
foliorum indistinctis, conidiophoris saepe ra-
mosis et conidiis brevioribus, (10–)20–70 µm,
1–7-septatis.
Leaf spots lacking or almost so, or only
forming diffuse, irregular discolorations. Colo-
nies amphigenous, mainly hypophyllous, thin,
subcircular to irregular in shape or effuse, dark
greyish brown to sooty. Mycelium internal and
external; superficial hyphae emerging through
stomata, sparingly developed to abundant,
sparingly branched, 1.5–3 µm wide, septate,
hyaline, subhyaline to pale olivaceous thin-
walled, smooth; stromata lacking or small,
substomatal, 10–25 µm diam., olivaceous-
brown, or somewhat erumpent aggregations of
swollen hyphal cells, up to 40 µm diam.
Conidiophores in small to moderately large
fascicles, mostly loose, arising from internal
hyphae or stromata, through stomata, or
solitary, arising from superficial hyphae, erect,
straight, subcylindrical to usually slightly to
strongly geniculate-sinuous, often branched, 5–
70 × 2.5–6 µm, pale olivaceous to medium
olivaceous-brown, 0–4-septate, often con-
stricted at the septa, thin-walled (wall up to
0.75 µm wide), smooth; conidiogenous cells
integrated, terminal or conidiophores reduced
to conidiogenous cells, 5–25 µm long, conidio-
genous loci inconspicuous or subdenticulate,
but always unthickened and not darkened.
Conidia solitary, oblavate-cylindrical, (10–)20–
70 × (2.5–)3–5(–5.5) µm, 1–7-septate, distance
between septa about 8–15 µm, subhyaline to
pale olivaceous, smooth or almost so, apex
obtuse to subacute, base short obconically
truncate, 1.5–2 µm wide, hila neither thickened
nor darkened.
Material examined – CUBA, Bayamo, on
leaves of Trichilia hirta L. (Meliaceae), 28 Jan.
1967, R. Urtiaga 9 (IMI 125049 = K(M)
176142, holotype); l.c., on T. hirta, without
date, R. Urtiaga 1161 (IMI 131876 = K(M)
176143, paratype).
Notes – Pseudocercospora trichilae (U.
Braun) U. Braun ( Cercostigmina trichiliae U.
Braun) on Heynea trijuga Roxb. (= Trichilia
connaroides (Wight & Arn.) Bentv.) is a quite
distinct species with sporodochial conidiomata,
percurrently proliferating conidiogenous cells
and pale conidiophores and conidia (Mehrotra
& Verma 1994, Braun 1995a). Several other
Pseudocercospora spp. have been described on
hosts of various genera belonging to the
Meliaceae. The South African P. ekebergiae
(Crous & B. Sutton) U. Braun & Crous is also
a former Cercostigmina with percurrently
proliferating conidiogenous cells (Braun & Hill
2002). Pseudocercospora aphanamixidis M.K.
Khan, M.S. Khan & Kamal on Aphanamixis sp.
in India (Khan et al. 1994), P. cedrelae (S.
Chowdhury) T. De on Toona ciliata M. Roem.
( Cedrella toona Roxb. ex Rottler & Willd.)
in India (Chowdhury 1961, Braun et al. 2003),
P. cedrelae-mexicanae U. Braun & Crous on
Cedrela odorata L. (= C. mexicana M. Roem.)
in Venezuela (Braun et al. 2003), P.
didymochetonis (Wakef.) Y.L. Guo on
Dysoxylum richii C. DC., Fiji (Chupp 1954,

Mycosphere Doi 10.5943/mycosphere/4/2/3
207
Guo 1999), P. subsessilis (Syd. & P. Syd.)
Deighton (incl. Pseudocercospora meliae A.N.
Rai & Kamal, ?P. meliacearum C.D. Sharma et
al., P. sagarensis S.A. Firdousi et al., Pseudo-
cercosporella indica A.N. Rai, B. Rai &
Kamal, P. meliae S.R. Chowdhury & Chandal),
widespread on Azadirachta spp. (Chupp 1954,
Hsieh & Goh 1990, Guo & Hsieh 1995, Kamal
2010), and P. toonae M.D. Mehrotra & R.K.
Verma (= P. toonae Y.L. Guo & X.J. Liu) on
Toona sinensis (A. Juss.) M. Roem. in China
and India (Guo & Hsieh 1995) are
distinguished from P. trichiliae-hirtae by
consistently fasciculate conidiophores, i.e.
superficial hyphae with solitary conidiophores
are lacking.
Four comparable Pseudocercospora spp.
on Meliaceous hosts characterized by the
formation of superficial hyphae with solitary
conidiophores are known, but all of them form
distinct leaf spots. P. swieteniae U. Braun &
Urtiaga on Swietenia macrophylla King in
Venezuela (Braun & Urtiaga 2008) is
morphologically very close to P. trichiliae-
hirtae but differs in having unbranched coni-
diophores and narrower conidia, 2.5–4 µm
wide, with narrower hila, 1–1.5 µm. P.
amoorae A.K. Das on Aphanamixis poly-
stachya (Wall.) R. Parker (= Amoora rohituca
(Roxb.) Wight & Arn.) in India (Das 1991) and
P. meliicola J.M. Yen, A.K. Kar & B.K. Das
on Azadirachta indica A. Juss. (= Melia
azadirachta L.) in India (Yen et al. 1982c) are
characterized by usually unbranched coni-
diophores and longer, pluriseptate conidia, up
to 120 µm. Pseudocercospora toonae-ciliatae
U. Braun, nom. nov., MycoBank, MB 801970
(Bas.: Cercoseptoria cedrelae J.M. Yen, A.K.
Kar & B.K. Das, Mycotaxon 16: 82, 1982, non
Pseudocercospora cedrelae (S. Chowdhury)
T.K. De, 1992) on Toona ciliata is easily
distinguishable from P. trichiliae-hirtae by its
intraepidermal stromata and cylindrical-
filiform conidia with truncate base that are
longer and narrower, 50–80  2–2.5 µm (Yen
et al. 1982c). Cercoseptoria is an established
synonym of Pseudocercospora, but Cerco-
septoria cedrelae has not yet been reallocated
to the latter genus, although morphologically
quite distinct from all other Pseudocercospora
species on Toona and other Meliaceous hosts.
Fig. 17 – Pseudocercospora trichiliae-hirtae.
Based on type material. a Hypha. b Solitary
conidiophore arising from superficial hyphae. c
Conidiophore fascicle. d Conidiophores. e
Conidia. – Bar = 10 µm.

Mycosphere Doi 10.5943/mycosphere/4/2/3
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Pseudocercospora trichostigmatis (F. Stevens)
Deighton
 Cercospora trichostigmatis F. Stevens.
Material examined – VENEZUELA, La-
ra, Barquisimeto, zoological garden, on leaves
of Trichostigma octandrum (L.) H. Walter
(Phytolaccaceae), Jan. 2008, R. Urtiaga 102
(HAL 2531 F); l.c., Nov. 2009, R. Urtiaga 297
(HAL 2537 F).
Notes – New to Venezuela (Crous &
Braun 2003, Iturriaga & Minter 2006).
Pseudocercospora zuelaniae U. Braun &
Urtiaga, sp. nov. Fig. 18
MycoBank, MB 801971.
Etymology – epithet derived from the
host genus, Zuelania.
Differt ab omnibus speciebus Pseudo-
cercosporae ad Samydaceas (P. caseariae, P.
caseariigena, P. cylindrosporioides, P. dovya-
lidis, P. samydacearum) hyphis superficialibus
evolutis.
Leaf spots amphigenous, subcircular to
angular-irregular, 2–20 mm diam., pale to dark
brown, later greyish brown to dingy greyish
white, margin indefinite or limited by darker
veins or a narrow dark border, dark brown to
blackish, in addition often with a diffuse
reddish to purple-violet halo. Caespituli epi-
phyllous, punctiform, dark brown to blackish,
scattered. Mycelium internal and external;
superficial hyphae amphigenous, emerging
through stomata, straight to sinuous, branched,
1–3.5 µm wide, septate, subhyaline to pale
olivaceous-brown, smooth, thin-walled; stro-
mata on the upper side, intraepidermal, brown,
10–40 µm diam., immersed to slightly
erumpent, subcircular to somewhat irregular in
outline, cells 2–5 µm diam. Conidiophores in
small to moderately large, usually loose
fascicles, arising from internal hyphae or
stromata on the upper leaf side, erumpent
(solitary conidiophores arising from superficial
hyphae not observed), subcylindrical or
somewhat attenuated towards the tip, straight
to strongly geniculate-sinuous, unbranched or
occasionally irregularly branched, 10–70 ×
2.5–5 µm, 0–6-septate, thin-walled, smooth,
pale to medium olivaceous or olivaceous-
brown; conidiogenous cells integrated, ter-
minal, occasionally conidiophores reduced to
conidiogenous cells, 5–25 µm long, coni-
diogenous loci inconspicuous to subden-
ticulate, but always unthickened and not
darkened. Conidia formed singly, obclavate-
cylindrical, short conidia sometimes ellipsoid-
subcylindrical or fusiform, (10–)20–65 × 2–4.5
µm, (0–)1–5(–6)-septate, subhyaline to very
pale olivaceous or olivaceous-brown, smooth,
thin-walled, apex obtuse to subacute, base short
obconically truncate, 1–2 µm wide, hila neither
thickened nor darkened.
Material examined – CUBA, Alegria del
Pio, Niquero, on leaves of Zuelania guidonia
(Sw.) Britton & Millsp. (Samydaceae [=
Salicaceae tribe Samydeae]), 18 Mar. 1968, R.
Urtiaga 1167 (IMI 132514 = K(M) 176156,
holotype).
Fig. 18 – Pseudocercospora zuelaniae. Based
on type material. a Conidiophore fascicles. b
Conidiophores. c Conidia. – Bar = 10 µm.

Mycosphere Doi 10.5943/mycosphere/4/2/3
209
Notes – The genus Zuelania was
previously assigned to the family Flacourti-
aceae, which is now considered a synonym of
the Salicaceae. Within the latter family it
belongs in the tribe Samydeae. In a recently
published phylogenetic examination, William
(2012) emphasized to recognize this group as a
separate family – Samydaceae. Pseudocerco-
spora zuelaniae is characterized by the
formation of epiphyllous as well as hypo-
phyllous superficial mycelium, but without any
solitary conidiophores arising from such
external hyphae. On the lower side, additional
distinctly verruculose, Zasmidium-like hyphae
have been observed. Superficial mycelium is
lacking in all other Pseudocercospora spp. on
hosts belonging to this tribe or family.
Furthermore, P. caseariae (F. Stevens) U.
Braun & Sivapalan differs in having much
shorter conidiophores (5–20 µm) and wider
conidia (4–6.5 µm), P. caseariigena H.S.G.
Rao, S. Chandra & Kamal is well characterized
by its very long (up to 126.5 µm), pluriseptate,
often branched conidiophores and very long
conidia (up to 140 µm), P. cylindrosporioides
(Solheim & Chupp) Y.L. Guo & X.J. Liu has
densely arranged, short conidiophores arising
from well-developed stromata and cylindrical
to almost acicular conidia, P. dovyalidis
(Chupp & Doidge) Deighton, recorded on
Guidonia tomentosa (Roxb.) Kurz [ Casearia
tomentosa Roxb.] (Crous & Braun 2003),
possesses narrowly linear-cylindrical to
subacicular conidia, and P samydacearum A.K.
Singh, Kamal & S.K. Singh (incl. Cerco-
septoria caseariae Abbasi, R. Kumar &
Kamal) is easily distinguishable by its large
stromata, up to 80 µm diam., shorter coni-
diophores (5–30 µm) and longer pluriseptate
conidia (Chupp 1954, Singh et al. 1985, Rao et
al. 1995, Braun & Sivapalan 1999). P.
flacourtiicola U. Braun & Kamal on Flacourtia
indica (Burm. f.) Merrill in Nepal is a rather
similar species, but superficial mycelium is not
formed (Braun et al. 2003). Other Pseudo-
cercospora species on former flacourtiaceous
hosts are also distinct from P. zuelaniae by
lacking superfical hyphae and some additional
differences: P. kiggelariae (Syd.) Crous & U.
Braun on Kiggelaria africana L. in South
Africa forms large stromata with numerous
densely fasciclate conidiophores and narrowly
cylindrical-filiform conidia and P. pangiicola
U. Braun & Sivapalan from Brunei on
Pangium edule Reinw. has much broader hila,
2–3.5 µm (Chupp 1954, Crous & Braun 1994,
Braun & Sivapalan 1999).
Ramularia occidentalis Ellis & Kellerm.
= Ramularia pratensis var. angustiformis
U. Braun.
Material examined – VENEZUELA, Rio
Claro, La Cunchilla, on leaves of Rumex
patienta L. (Polygonaceae), Apr. 2009, R.
Urtiaga 239 (HAL 2540 F).
Notes – New host and new to Venezuela
(Braun 1998). The conidiophores in this
collection on Rumex patientia are only 1.5–3
µm wide, and the conidia are 5–30  (1–)1.5–
2(–2.5) µm, which agrees well with Ramularia
occidentalis. However, in typical collections of
the latter species, leaf spots are lacking or
diffuse, expanded, yellowish, ochraceous to
brownish, finally grey. The specimen on
Rumex patientia has distinct leaf spots, rather
agreeing with lesions formed by Ramularia
pratensis Sacc. Based on distinct, Ramularia
pratensis-like leaf spots and narrow conidia, a
North American collection on Rumex
acetosella was described as R. pratensis var.
angustiformis (Braun 1998). This variety
should be reduced to synonymy with
Ramularia occidentalis. Typical collections of
the latter species without distinct leaf spots are
also known from North America on Rumex
acetosella (Braun 1998). The taxonomic value
of differences in the lesions is unclear and may
be influenced by reactions of particular hosts.
Ramularia indica K.L. Kothari, M.K. Bhatn. &
N.S. Bhatt (= R. pakistanica S.A. Khan & M.
Kamal), known from India and Pakistan on
Rumex dentatus L., has narrow conidiophores
and conidia, agreeing with those of Ramularia
occidentalis, but due to distinct leaf spots
Braun (1998) treated this fungus as variety of
the later species. The relation between
Ramularia indica and R. occidentalis is not
quite clear, but the Asian species should be
maintained as a separate species, at least
tentatively.
Stenella araguata Syd.
Material examined – VENEZUELA, La-

Mycosphere Doi 10.5943/mycosphere/4/2/3
210
ra, Rio Claro, La Cuchilla, on leaves of Pithe-
cellobium dulce (Roxb.) Benth. (Fabaceae),
Apr. 2009, R. Urtiaga 235 (HAL 2554 F).
Notes – This is a new collection of this
species, which was described on type material
from Venezuela.
Zasmidium cubense U. Braun & Urtiaga, sp.
nov. Fig. 19
MycoBank, MB 801972.
Etymology – epithet referring to Cuba.
Zasmidii oliganthis valde similis sed
maculis foliorum formantibus et conidiophoris
aequabiliter latis vel apicem versus attenuatis.
Leaf spots amphigenous, subcircular to
angular-irregular, 1–5 mm diam. or confluent
and larger, darker brown on the upper leaf
surface, paler brown below, margin indefinite,
sometimes vein-limited. Colonies hypho-
phyllous, inconspicuous. Mycelium internal
and external; hyphae emerging through sto-
mata, superficial, sometimes climbing leaf
hairs, branched, 1.5–4 µm wide, septate,
subhyaline to pale olivaceous, thin-walled,
verruculose. Stromata lacking. Conidiophores
solitary, arising from superficial hyphae,
lateral, rarely terminal, erect, straight, conical,
subcylindrical and straight to distinctly geni-
culate-sinuous, unbranched, width uniform
throughout or attenuated towards the tip, 5–45
 2–5 µm, 0–3-septate, pale olivaceous to
olivaceous-brown, thin-walled, smooth to
faintly rough-walled; conidiophores reduced to
conidiogenous cells or integrated, terminal to
intercalary, 5–30 µm long, conidiogenous loci
conspicuous, usually several per cell, slightly
thickened and darkened, in front view visible
as minute dark circle, 1–1.5 µm diam. Conidia
solitary, cylindrical, subacicular to almost
obclavate, 15–60  2.5–4 µm, (0–)1–6-septate,
subhyaline to pale olivaceous or brownish,
thin-walled, verruculose, apex obtuse to sub-
acute, base truncate to short obconically
truncate, 1.5–3 µm wide, hilum very slightly
thickened and somewhat darkened-refractive.
Material examined – CUBA, Bayamo, on
leaves of Koanophyllon villosum (Sw.) R.M.
King & H. Rob. [ Eupatorium villosum Sw.]
(Asteraceae), 18 May 1967, R. Urtiaga C-503
(IMI 127702 = K(M) 176158, holotype).
Fig. 19 – Zasmidium cubense. Based on type
material. a Hyphae. b Solitary conidiophores
arising from superficial hyphae. c
Conidiophores. d Conidia. – Bar = 10 µm.
Notes – Several Zasmidium species
have been described on host plants belonging
to the Asteraceae. Z. oliganthis (R.E.D. Baker
& W.T. Dale) U. Braun & Crous, described on
Oliganthes condensata (Less.) Sch. Bip. from
Trinidad, is morphologically close to Z.

Mycosphere Doi 10.5943/mycosphere/4/2/3
211
cubense but leaf spots are lacking in the latter
species and the conidiophores are wider near
the apex (Chupp 1954, Braun et al. 2010). The
other species are quite distinct from Z. cubense
and easily distinguishable by obvious dif-
ferences in the characters of conidiophores and
conidia. Z. haematitica U. Braun & Crous on
Solidago flexicaulis L. [= S. latifolia L.] in
North America is characterized by its very
long, pluriseptate conidia, 50–260  2–4.5 µm
(Braun et al. 2003); Z. kansense (Syd. & P.
Syd.) U. Braun on Cirsium spp. in North
America is distinguished by its very long
conidiophores, 200–500 µm, and broad
conidia, 5–7 µm (Chupp 1954, Braun et al.
2010); Z. oligoneuri (H.C. Greene) U. Braun &
Crous on Solidago riddellii Frank in the USA
has long conidiophores, up to 115 µm,
emerging through stomata, and long, uniformly
obclavate conidia, 40–70  2.5–4 µm (Crous &
Braun 2003); and in Z. solidaginis (Chupp &
H.C. Greene) U. Braun on several Solidago
spp. in Asia (India and Japan) and North
America, leaf spots are lacking and stromata as
well as fasciculate conidiophores are formed
(Chupp 1954, Crous & Braun 2001, Kamal
2010).
Zasmidium genipae-americanae U. Braun &
Urtiaga, sp. nov. Fig. 20
MycoBank, MB 801973.
Etymology – epithet derived from the
host species, Genipa americana.
Zasmidii coffeae, Z. naucleae, Z. plec-
troniae, Z. xeromphigeni et Z. hyptiantherae
similis, sed conidiis valde longioribus, ad 220
µm, et ad 14-septatis.
On living leaves, lesions amphigenous,
subcircular to mostly angular-irregular, some-
times confluent, 1–5(–10) mm diam., ochra-
ceous, pale to medium brown, later becoming
pale, margin indefinite or limited by somewhat
darker veins. Colonies amphigenous, puncti-
form, scattered, delicate, dark brown to
blackish. Mycelium internal and external;
superficial hyphae emerging through stomata
or arising from stromata, richly developed on
the lower leaf surface, less abundant above,
sparingly branched, straight and rigid to
somewhat sinuous, 1–3 µm wide, septate, thin-
walled, subhyaline to pale olivaceous-brown,
verruculose. Stromata well-developed, im-
mersed to substomatal, 10–60 µm diam.,
brown, composed of swollen hyphal cells, 2–8
µm diam., wall somewhat thickened. Conidio-
phores solitary, arising from superficial
hyphae, lateral, rarely terminal, with 1–2
conidiophores per cell, as well as aggregated in
small to moderately large fascicles, loose,
arising from stromata, through stomata or
erumpent, erect, straight, subcylindrical to
moderately geniculate-sinuous, unbranched, 5–
35 × 2–4 µm, 0–1-septate, pale to medium
olivaceous or olivaceous-brown throughout or
with paler tips, thin-walled, almost smooth to
somewhat rough-walled; conidiophores mostly
reduced to conidiogenous cells, or conidio-
genous cells occasionally integrated, terminal,
5–25 µm long, with a single to usually several
conspicuous conidiogenous loci, circular in
outline, slightly thickened and darkened, 1–1.5
µm diam. Conidia formed singly, narrowly
obclavate, cylindrical to acicular, 15–80 × 2–4
µm, (0–)1–7-septate, subhyaline to olivaceous-
brown, thin-walled, almost smooth to verru-
culose, apex subacute to obtuse, base truncate
to short obconically truncate, rarely long ob-
conically truncate, 1–2 µm wide, hila slightly
thickened and darkened.
Material examined – VENEZUELA, La-
ra, on leaves of Genipa americana L. (Rubia-
ceae), 22 Jan. 1970, R. Urtiaga 928 (IMI
146238 = K(M) 176137, holotype).
Notes – Several species of Zasmidium
have been described on hosts belonging to the
Rubiaceae, but all of them occur on hosts of
other genera, are only known from Asia (India
and Nepal) and morphologically quite distinct.
Stromata and fasciculate conidiophores are
lacking in the Indian species Zasmidium canthii
(J.M. Yen, A.K. Kar & B.K. Das) Kamal on
Canthium didymum C.F. Gaert. (Yen et al.
1982a, Kamal 2010: 238) and Z. wend-
landiicola (U. Braun & Crous) Kamal & U.
Braun on Wendlandia exserta (Roxb.) DC.
(Crous & Braun 2003, Kamal 2010: 238). Z.
rubiacearum (S. Chaudhary, N. Sharma &
Kamal) Kamal (Chaudhary et al. 2002, Kamal
2010: 247) on Wendlandia tinctoria (Roxb.)
DC. has very small stromata composed of a
few swollen hyphal cells, much longer
conidiophores, up to 113.5 µm, and longer
conidia, up to 100.5 µm. Z. meynae-laxiflorae
(K. Srivast., A.K. Srivast. & Kamal) P.

Mycosphere Doi 10.5943/mycosphere/4/2/3
212
Fig. 20 – Zasmidium genipae-americanae.
Based on type material. a Hyphae. b Solitary
conidiophores arising from superficial hyphae.
c Conidiophore fascicle. d Conidia. – Bar = 10
µm.
Phengsintham, K.D. Hyde & U. Braun on
Meyna laxiflora Robyns in Asia (Srivastava et
al. 1995, Phengsintham et al. 2009) is distin-
guished from the new species by its much
longer conidiophores, up to about 100 µm and
conidia partly formed in chains. Z. vangueriae
(M.J. Thirum. & Mishra) Kamal (Kamal 2010:
252) on Meyna spinosa Roxb. ex Link is quite
distinct by having catenate cladosporioid coni-
dia. All other species of Zasmidium on rubia-
ceous hosts form well-developed stromata, but
differ from Z. genipae-americanae in having
much longer, pluriseptate conidia, up to about
220 µm and about 14 septa [viz. the India
species Z. coffeae (J.M. Yen, A.K. Kar & B.K.
Das) Kamal on Coffea bengalensis B. Heyne
ex Schult. (Yen et al. 1982b, Kamal 2010), Z
naucleae (A.K. Das) Kamal on Nauclea
excelsa Blume (Das 1990, Kamal 2010: 238),
Z. plectroniae (K.M. Ponnappa) Kamal on
Canthium coromandelicum (Burm. f.) Alston
(= Plectronia parviflora (Lam.) Bedd.)
(Ponappa 1968, Kamal 2010: 246) and Z.
xeromphigenum (Y.M. Yen, A.K. Kar & B.K.
Das) Kamal on Tamilnadia uliginosa (Retz.)
Tirveng. & Sastre (Yen et al. 1982b, Kamal
2010: 252)]. Stenella hyptiantherae, described
from Nepal on Hyptianthera stricta (Roxb. ex
Schult.) Wight & Arn., belongs to this group of
species with very long, pluriseptate conidia as
well. Based on cercosporoid conidia and
planate conidiogenous loci and hila, this
species undoubtedly belongs to Zasmidium, but
has not yet been reallocated:
Zasmidium hyptiantherae (S.K. Singh, Arch.
Singh & Kamal) U. Braun, comb. nov.
Mycobank MB 801974.
Bas.: Stenella hyptiantherae S.K. Singh,
Arch. Singh & Kamal, Mycol. Res. 101(5):
533, 1997.
Acknowledgements
We are much obliged to the mycology
curator of the Kew Herbarium (K) for the
possibility to examine collections previously
deposited by R. Urtiaga at IMI.
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