SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
1
A monograph of Fusicladium s.lat. (Hyphomycetes)
Konstanze SCHUBERT, Anja RITSCHEL & Uwe BRAUN
Abstract: S CHUBERT , K., R ITSCHEL, A. & B RAUN , U.: A monograph of Fusicladium s.lat.
(Hyphomycetes). Schlechtendalia 9: 1–132.
The genus Fusicladium s.lat. is monographed. Pollaccia and Spilocaea are reduced to synonymy
with Fusicladium. The latter genus has been proposed to be conserved. The history, phylogeny,
taxonomy, circumscription and delimitation of this genus are discussed in detail, a key to Fusicladium
and morphologically similar genera and a key-like list of Fusicladium species by host genera are
included. Individual species are then described, illustrated and discussed. Doubtful, ill-defined and
excluded taxa are listed and discussed at the end of the paper. The new species Fusicladium asperatum,
F. caulicola, F. junci and F. nashicola are described and the new combinations F. ahmadii, F.
byrsonimatis, F. catenosporum, F. elegans, F. mandshuricum, F. nebulosum, F. oleagineum, F.
phillyreae, F. radiosum var. lethiferum, F. radiosum var. populi-albae and Pseudocladosporium
caruanianum are introduced.
Zusammenfassung: SCHUBERT, K., RITSCHEL , A. & BRAUN , U.: Monographie der Gattung
Fusicladium s.lat. (Hyphomyceten). Schlechtendalia 9: 1–132.
Die Gattung Fusicladium s.lat. wird monographisch bearbeitet. Pollaccia und Spilocaea werden als
Synonyme von Fusicladium betrachtet. Fusicladium wurde zur Konservierung vorgeschlagen. Geschichte, Phylogenie, Taxonomie, Umschreibung und Abgrenzung der Gattung werden im Detail
diskutiert. Ein Schlüssel zu Fusicladium und ähnlichen Gattungen und eine schlüsselartige Liste zu
den Arten, auf Grundlage der Wirtsgattungen, werden geboten. Die einzelnen Arten werden ausführlich beschrieben, abgebildet und diskutiert. Zweifelhafte, unklare und ausgeschlossene Taxa werden
am Ende dieser Arbeit aufgelistet und diskutiert. Die neuen Arten Fusicladium asperatum, F. caulicola,
F. junci und F. nashicola werden beschrieben und die neuen Kombinationen F. ahmadii, F.
byrsonimatis, F. catenosporum, F. elegans, F. mandshuricum, F. nebulosum, F. oleagineum, F.
phillyreae, F. radiosum var. lethiferum, F. radiosum var. populi-albae und Pseudocladosporium
caruanianum werden eingeführt.
Contents
1.
Introduction ................................................................................................... 2
2.
Materials and methods .................................................................................. 2
3.
History of Fusicladium, Pollaccia and Spilocaea ......................................... 3
3.1. Spilocaea ....................................................................................................... 3
3.2. Fusicladium .................................................................................................. 3
3.3. Pollaccia ....................................................................................................... 4
4.
Molecular examinations and phylogeny ....................................................... 5
5.
The taxonomic value of morphological features ........................................... 6
6.
Taxonomy based on morphology and molecular data ................................... 8
7.
The species concept ...................................................................................... 9
8.
Key to Fusicladium emend. and similar genera .......................................... 10
9.
Index/key to Fusicladium species by host genera ....................................... 11
10.
The species of Fusicladium emend. ............................................................. 15
10.1. Abbreviations ................................................................................................ 15
2
10.2.
11.
12.
13.
14.
15.
15.1.
15.2.
1.
Schlechtendalia 9 (2003)
Descriptions of species ................................................................................. 17
Doubtful and unclear species of Fusicladium s.lat. .................................... 109
Excluded species ........................................................................................... 112
References ..................................................................................................... 118
Acknowledgements ....................................................................................... 125
Index ............................................................................................................. 126
Index of fungal names .................................................................................. 126
Index of host genera ..................................................................................... 131
Introduction
The genera Fusicladium Bonord., Pollaccia E. Bald. & Cif. and Spilocaea Fr.
(Hyphomycetes) are, as far as known, anamorphs of Venturia Sacc. (Ascomycota,
Venturiaceae E. Müll. & Arx ex M.E. Barr). These fungi are plant pathogens causing
characteristic leaf spots, necroses and scab diseases as well as leaf and fruit deformations. They overwinter as mycelia in fallen fruits, leaves and twigs, where they form
pseudothecia with asci and ascospores in the following spring. Mature two-celled
ascospores are forcibly ejected after dehiscence of the apical ascus wall and are dispersed by the wind. They germinate on suitable host organs, form germ tubes which
enter the host through stomata or directly through the cuticle. Intramatrical mycelia
then develop and new infections become established. First symptoms are often visible
as small leaf discolorations or spots, which are later covered by dark punctiform to
effuse fungal colonies, composed of conidiophores, conidiomata and conidia. Conidia
are spread throughout the whole growing season by rain and wind, and are transferred
to healthy fruit, leaves and twigs, where they cause new infections.
The anamorphs are the phytopathologically relevant and diagnostically important phases
in the life cycles of these fungi. Since monographs of these fungi do not exist, comprehensive examinations of Fusicladium, Pollaccia and Spilocaea species by means of molecular, morphological and scanning electron microscopic methods have recently been carried
out, with the aim to elucidate the phylogenetic and taxonomic significance of these
anamorphs for Venturia. All taxa assigned to these genera have been considered and reassessed, i.e., either excluded from or recognised in Fusicladium s.lat. The latter species are
keyed out, redescribed, illustrated and supplemented with host range and distribution
data, so that this work can be considered a monograph of this fungal genus.
2.
Materials and methods
Collections from numerous herbaria (BRA, DAVFP, G, HAL, HBG, IMI, JE, K, LE,
M, NTU-PPE (TAI), NY, PAD, PC, PDD, TLF, WIS, VPRI) and some fresh specimens were examined by standard light microscopy (oil immersion). Drawings and
photomicrographs were also made where necessary from prepared material, including type collections. Electronmicrographs were prepared at the “Interdisziplinäres
Wissenschaftliches Zentrum für Materialwissenschaften” at the Martin-Luther-University, Halle, using an Environmental Scanning Electron Microscope (ESEM). Molecular investigations were carried out at the “Botanische Staatssammlung” (Munich).
Detailed results of these studies will be published separately.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
3.
History of Fusicladium, Pollaccia and Spilocaea
3.1.
Spilocaea
3
The name Spilocaea was introduced by FRIES (1819) in connection with Spilocaea
pomi, the type species of this genus, but the first detailed description dates back to
FRIES (1825). LINK (1825) added Spilocaea scirpi, described from stems of Scirpus
sp. CORDA (1829) redescribed the latter species, but its generic affinity and the
status of Spilocaea was not discussed. In 1832, FRIES described Spilocaea epiphylla
from leaves of Malus and Pyrus species and mentioned S. scirpi in a foot-note. S.
epiphylla and S. scirpi have not been recorded or otherwise considered since.
SACCARDO (1886) assigned Spilocaea (with S. concentria Schwein., S. epiphylla
Fr., S. pomi Fr., S. opuntiae Rabenh. and S. scirpi Link) to a list of doubtful and
excluded genera (“Genera dubia vel excludenda“). SACCARDO (1897), ADERHOLD
(1896, 1897), LINDAU (1907) and FERRARIS (1912) treated Spilocaea pomi as a
synonym of Fusicladium dendriticum (Wallr.) Fuckel. LIND (1913) proposed the
combination Fusicladium pomi (Fr.) Lind, although S. pomi was the type species of
the older genus Spilocaea. For a long time, Spilocaea had been a forgotten name.
Most species morphologically belonging to the Spilocaea type were treated under
Fusicladium or other names like Cladosporium Link and Cycloconium Castagne
(VASSILJEVSKY & KARAKULIN 1937). In 1953, HUGHES re-introduced Spilocaea
for Venturia anamorphs with annellate conidiogenous cells, reduced Cycloconium
Castagne (CASTAGNE 1845) and Basiascum Cavara (CAVARA 1888) to synonymy
with this genus, and confined Fusicladium to taxa with sympodially proliferating
conidiogenous cells. Most subsequent authors followed his new taxonomic concept
(e.g., BARR 1968; ELLIS 1976; SIVANESAN 1977, 1984a).
3.2.
Fusicladium
Fusicladium was introduced by BONORDEN (1851) as a monotypic hyphomycetous
genus with F. virescens Bonord. on apple leaves as type species. Based on Malus as
type host, HÖHNEL (1923) considered F. virescens to be identical with F. dendriticum
(= Spilocaea pomi), although BONORDEN (1851) described and depicted denticulate (sympodial) conidiogenous cells. SACCARDO (1897) and LINDAU (1907) reduced F. virescens to a synonym of F. pyrorum (Lib.) Fuckel. HUGHES (1953) later
suggested that the type host of the former species could have been misidentified.
Type material of Bonorden´s species is not preserved, so it is not possible now to
check the identification of the host. Since Fusicladium pyrorum and Spilocaea pomi
are pathogens of Malus spp. as well as Pyrus spp., it is irrelevant if F. virescens had
been described from apple or pear leaves. The identity of this species can only be
proven on the basis of the morphological features of the conidiophores given in the
original description. SACCARDO (1897), LINDAU (1907), FERRARIS (1912) and other
previous authors used Fusicladium s.lat. for taxa with sympodial (denticulate) as
well as percurrent (annellate) conidiogenous cells, including Pollaccia-like species, e.g., F. radiosum (Lib.) Lind, and Spilocaea-like taxa, e.g., F. dendriticum.
BALDACCI & CIFERRI (1937) excluded Fusicladium radiosum and placed it in the
new genus Pollaccia. V IENNOT -B OURGIN (1949) introduced the new genus
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Schlechtendalia 9 (2003)
Megacladosporium for Fusicladium-like species with denticulate (sympodial)
conidiogenous cells and confined Fusicladium s.str. to taxa with annellate
conidiogenous cells, being undoubtedly influenced by HÖHNEL´s (1923) treatment of
F. virescens, the type species of Fusicladium, as a synonym of F. dendriticum.
Megacladosporium was introduced without indicating a type species and has to be
considered a superfluous name since the type species of Fusicladium was included in
the protologue of its original description. Based on BONORDEN´s (1851) original description and illustration, HUGHES (1953) confined Fusicladium s.str. to species with
sympodially proliferating conidiogenous cells and more or less denticle-like
conidiogenous loci, and placed all taxa with distinctly annellate conidiogenous cells
in Spilocaea. Several authors assigned Fusicladium species with catenate conidia to
Cladosporium, e.g., F. carpophilum (Thüm.) Oudem., F. cerasi (Rabenh.) Erikss. and
F. effusum G. Winter [as Cladosporium caryigenum (Ellis & Langl.) Gottwald]
(BENSANDE & KEITT 1928; ELLIS 1976; GOTTWALD 1982). Previous authors, e.g.,
SACCARDO (1897) and LINDAU (1907), treated these species under Fusicladium s.lat.
HÖHNEL (1923) and VASSILJEVSKY & KARAKULIN (1937) excluded some species
with catenate conidia from Fusicladium as well as Cladosporium and introduced the
new genera Hormocladium and Fusicladiopsis, respectively. The latter name, which
is a younger homomyn of Fusicladiopsis R. Maire, 1906, was replaced by Karakulinia
nom. nov. (GOLOVINA 1964). ONDÌEJ (1971) retained species with catenate conidia
in Fusicladium, but placed these taxa in the new subgenus Pseudofusicladium. BATISTA
(1957) described the new genus Ramalia with R. veronicae Bat. as type species.
SUTTON & PASCOE (1988) re-examined type material of this species and reduced
Ramalia to synonymy with Fusicladium. Ramalia veronicae is also characterised by
having catenate conidia. PARTRIDGE & MORGAN-JONES (2003) described the new
genus Fusicladosporium [type species: F. carpophilum (Thüm.) Partridge & MorganJones, ≡ Cladosporium carpophilum Thüm., Fusicladium carpophilum] for Venturia
anamorphs with catenate conidia. They discussed Karakulinia N.P. Golovina and
Megacladosporium Vienn.-Bourg., but failed to take into consideration that the older
genera Hormocladium and Ramalia had also been introduced for Fusicladium-like
anamorphs with catenate conidia.
3.3.
Pollaccia
BALDACCI & CIFERRI (1937) described the monotypic genus Pollaccia for P. radiosa
(Lib.) E. Bald. & Cif. (≡ Oidium radiosum Lib., Fusicladium radiosum) distinguished
from Fusicladium species by having short, monoblastic, determinate to percurrent,
annellate conidiogenous cells. SERVAZZI (1939) distinguished two Pollaccia species
on Populus leaves, viz., P. radiosa, which he considered the anamorph of Venturia
tremulae Aderh., and P. elegans Servazzi, the anamorph of V. populina (Vuill.) Fabric. HUGHES (1953) recognised Pollaccia as a separate genus well-distinguished from
Fusicladium and Spilocaea, and most subsequent authors followed his taxonomy (e.g.,
BARR 1968; ELLIS 1971, 1976; SIVANESAN 1977, 1984a; BRANDENBURGER 1985).
MORELET (1972, 1985) and MORELET & SIGAUD (1996) published important contributions to biology and taxonomy of Pollaccia species.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
4.
5
Molecular examinations and phylogeny
SCHNABEL et al. (1999) were the first to publish molecular analyses (rDNA, ITS) of
Venturia species. Their analyses, which were restricted to species that occur on rosaceous hosts, were later supplemented by KASANEN et al. (2001) who examined Venturia
ditricha (Fr.) P. Karst., V. populina and V. tremulae. Sequence data deposited at the
NCBI (National Center for Biotechnology Information) have been used for a new,
more comprehensive molecular analysis of Venturia species and their anamorphs together with additional sequences of Fusicladium, Pollaccia and Spilocaea species
obtained during the course of molecular examinations carried out in Munich. The
research group there used Cladosporium [C. cladosporioides (Fresen.) G.A. de Vries],
often confused with Fusicladium, as the outgroup, and included data of “human pathogenic Cladosporium species“ (= Cladophialophora Borelli) and Botryosphaeria
dothidea (Moug.) Ces. & de Not. The data coming out of these studies are still limited, so only preliminary conclusions are possible. The detailed results of these examinations will be published in a separate paper. A cladogram based on all the data
mentioned above provides the first comprehensive molecular results for Venturia and
its anamorphs and a first insight into phylogenetic connections. In any case, Venturia
species and its anamorphs (Venturiaceae) have been proven to form a monophyletic
clade. This was confirmed by BRAUN et al. (2003) who put rDNA ITS data of
cladosporioid Venturia anamorphs in a more comprehensive context of Cladosporium-like fungi, i.e., dematiaceous hyphomycetes with amero- to phragmosporous
conidia formed in acropetal chains.
The cladogram published by SCHNABEL et al. (1999) is less useful for phylogenetic
analyses and taxonomic interpretations since “Cladosporium caryigenum“ was taken
to serve as the outgroup. The latter species is, however, a true anamorph of the
Venturiaceae and this has been confirmed by molecular analysis and a study of the
morphology of the fungus.
The Venturia clade is composed of several small subclades, which, at least partly, seem
to indicate tendencies of co-evolutions between hosts and Venturia species. V. asperata
Samuels & Sivan., V. carpophila E.E. Fisher and V. cerasi Aderh. form a subclade of
closely allied species on various hosts of the Rosaceae that are characterised by having
anamorphs with catenate conidia. The Rosaceae also appear to have been colonised by
Venturia on several different occasions. Venturia inaequalis (Cooke) G. Winter s.lat. as
well as Venturia pyrina Aderh. and V. nashicola S. Tanaka & S. Yamam. form separate
subclades. Another group is formed by species with Pollaccia anamorphs on hosts of
the Salicaceae. Various morphological features of Venturia anamorphs, e.g., conidia
formed singly or in chains and proliferation of conidiogenous cells percurrently or
sympodially, have probably been acquired several times in the past in separate lineages
in co-evolution with several groups of hosts. However, a clear separation of the Venturia
clade into uniform subclades based on morphological features of the anamorphs is not
evident. Species with sympodial conidiogenous cells occur in several subclades, and
taxa with percurrent proliferations form two different groups separated by a subcluster
with Fusicladium anamorphs (sympodial conidiogenous cells). Species with catenate
conidia are also not confined to a single subcluster.
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Schlechtendalia 9 (2003)
These results also show that Fusicladium convolvularum OndÍej and F. effusum, two
species with unknown teleomorphs, are true members of the Venturiaceae. The treatment of Venturia pyrina and V. nashicola as two distinct species is also supported by
the present molecular data.
5.
The taxonomic value of morphological features
Fusicladium, Pollaccia and Spilocaea are characterised by having immersed, subcuticular to intraepidermal mycelia, often forming radiating strands or loose to dense
stromatic aggregations, which give rise to conidiogenous cells or conidiophores penetrating the cuticle. A superficial (secondary) mycelium is rarely formed, but is present,
for example, in Fusicladium veronicae (Bat.) B. Sutton & Pascoe, F. scillae (Deighton)
U. Braun & K. Schub. and Spilocaea oleaginea (Castagne) S. Hughes. In Spilocaea
pomi, a superficial mycelium is usually absent, but in a single collection some external hyphae have been observed. The conidiophores are usually erumpent through the
cuticle, but in Fusicladium levieri Magnus, F. junci Sawada ex K. Schub. & U. Braun
and F. convolvularum they are, at least partly, fasciculate and emerge through stomata. In Pollaccia species, superficial hyphae are unknown.
The arrangement of the conidiophores within the three anamorphic genera of the
Venturiaceae is fairly variable, ranging from conidiophores formed singly or in loose
to dense fascicles, to distinct sporodochia. In Spilocaea nebulosa (Ellis & Everh.) S.
Hughes & Piroz., all conidiophores are formed singly and are erumpent through the
cuticle, in S. oleaginea they are solitary to fasciculate, and S. pomi is characterised by
having loosely to densely fasciculate conidiophores. In Fusicladium species, the
conidiophores are usually formed singly (e.g., F. caricinum Bres., F. veronicae) or in
fascicles (e.g., F. pyrorum), but some taxa with sporodochial conidiomata are also
known (e.g., F. fraxini Aderh., F. romellianum OndÍej). In F. martianoffianum (Thüm.)
K. Schub. & U. Braun, all kinds of conidiophore arrangements, ranging from solitary
conidiophores to sporodochia, can be observed. Pollaccia species are usually characterised by having conidiophores in fascicles or sporodochia. Hence, the arrangement
of conidiophores and the location of the mycelium are not suitable for the differentiation of Fusicladium, Pollaccia and Spilocaea. These genera have been traditionally
distinguished by the mode of proliferation of the conidiogenous cells, viz., Fusicladium
with sympodial proliferation, Pollaccia with monoblastic, determinate to percurrent
conidiogenous cells (with few rather inconspicuous annellations) and Spilocaea with
percurrent proliferation and numerous conspicuous annellations. However, there are
numerous transitional cases, e.g., in Fusicladium obducens Pat. and F. veronicae, two
species with sympodial proliferations which are occasionally mixed with some
annellations caused by percurrent proliferations. The conidiogenous cells of
Fusicladium caulicola U. Braun & K. Schub. and F. romellianum are usually
monoblastic, determinate. In F. fraxini, the conidiogenous cells range from being
unilocal, percurrent to multilocal, sympodial and sometimes even mixed in the same
collection. Pollaccia and Spilocaea species possess unilocal, determinate to percurrent
(annellate) conidiogenous cells, except for Spilocaea oleaginea and S. nebulosa in
which conidiogenous cells with several loci had been observed (HUGHES 1953). The
differences between Pollaccia and Spilocaea are weak and only gradual, so that a
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
7
separation of these genera cannot be maintained. Both genera have monoblastic (unilocal)
conidiogenous cells, frequently determinate or only with few annellations in Pollaccia
and usually with numerous annellations in Spilocaea. The discrimination of Fusicladium
on the one hand and Pollaccia/Spilocaea on the other hand, based on the presence of
percurrent and sympodial conidiogenous cells, is also not tenable. This can be demonstrated by the existence of numerous taxa with mixed types of conidiogenous cells.
This phenomenon has also been observed in various other hyphomycetous genera, e.g.,
in cercosporoid Mycosphaerella anamorphs. CROUS et al. (2000, 2001) showed that
combinations of sympodially and percurrently proliferating conidiogenous cells are not
uncommon in Pseudocercospora, and that the separation of the latter genus and
Cercostigmina U. Braun is not tenable. This view is also supported by data from molecular studies of these genera. Percurrent and sympodial conidiogenous cells are also
known to occur together in Septoria Sacc. species (VERKELEY 1997).
Different structures of the conidiogenous loci, reflecting differences in the mode of
conidiogenesis, proved to be meaningful for genera of cercosporoid anamorphs (CROUS
et al. 2000, 2001; CROUS & BRAUN 2003). The conidiogenous loci and conidial hila
of Fusicladium, Pollaccia and Spilocaea species have been examined by means of
light and scanning electron microscopy (RITSCHEL 2001; SCHUBERT 2001). The basic structure of the loci and hila is, however, rather uniform by being more or less
truncate, unthickened or almost so and non-pigmented to slightly darkened–refractive. These scars agree well with those of Pseudocercospora species. There are not
any fundamental differences in the structures of the conidiogenous loci and conidial
hila of Fusicladium, Pollaccia and Spilocaea species. The conidiogenous loci of
Fusicladium species are often more or less denticle-like, and there are gradual differences in the width of loci and hila, which are mostly somewhat narrower in Fusicladium
and wider in Pollaccia and Spilocaea, although there are numerous exceptions and
transitions, e.g., in Fusicladium caricinum with relatively broad loci.
The conidiogenesis in all Venturia anamorphs is consistently holoblastic, and the
structures of the conidiogenous loci and conidial hila are uniform, and, consequently,
these features cannot be used to separate these genera.
The conidia of Fusicladium, Pollaccia and Spilocaea species are also fairly uniform
by being amero-, didymo- to phragmosporous and pigmented. They are usually ellipsoid–ovoid to fusiform and more or less smooth, although there are some taxa with
verruculose conidia [e.g., Fusicladium psoraleae (Ellis & Barthol.) S. Hughes & Piroz.
and F. pisicola Linford].
It has been suggested that the formation of the conidia in chains or singly is a feature
that could be used to separate Fusicladium into smaller units (H ÖHNEL 1923,
GOLOVINA 1964, ONDÌEJ 1971, see chapter 3.2.). Taxa with catenate conidia occur in
Fusicladium as well as Pollaccia, viz., P. catenospora Butin, and there are even some
species in which the conidia are formed singly as well as in chains, e.g., F. cerasi.
Conidial formed in chains or singly is a good, useful feature for differentiating between species, but is is not one that can be used at the generic level. This is also true
with other groups of hyphomycetes, e.g., Ramularia Unger (BRAUN 1998) and
cercosporoid genera (CROUS et al. 2000, 2001). A separation of species with catenate
conidia from Fusicladium is not tenable and this is also supported by molecular data.
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6.
Taxonomy based on morphology and molecular data
The results of the examinations recently carried out by RITSCHEL (2001) and SCHUBERT
(2001) have shown that, within the Venturia anamorphs (Fusicladium, Pollaccia and
Spilocaea), features such as the type and growth of the mycelium, arrangement of the
conidiophores (solitary, fasciculate and sporodochial), proliferation of conidiogenous
cells (percurrent and sympodial), structure of the conidiogenous loci and shape and
size and formation of conidia (solitary, catenate) cannot be used to define the various
genera. These features are only useful for distinguishing between species. RITSCHEL
(2001) and SCHUBERT (2001) using molecular analyses has also demonstrated that
Venturia species form a monophyletic clade. These two sets of results mean that the
separation of Venturia anamorphs into several genera cannot be maintained and that
Fusicladium, Pollaccia and Spilocaea must be merged into a single anamorph genus.
The conidiogenesis and structure of the conidiogenous loci and conidial hila are uniform in these genera and resemble those found in Pseudocercospora and allied genera (Mycosphaerella anamorphs). The situation in the Venturiaceae reminds one of
the Erysiphales (powdery mildew fungi), which is characterised by the anamorph
genus Oidium Link. There is, however, a nomenclatural problem since the less known,
smaller genus Spilocaea (five recognised species) is older than the much larger genus
Fusicladium with 40 recognised species. Fusicladium is well-known to mycologists
and phytopathologists and readily associated with Venturia. Merging the three
anamorph genera of Venturia under the oldest name Spilocaea would result in more
than 40 new combinations. Therefore, it has been proposed to maintain the wellknown genus Fusicladium (BRAUN et al. 2002). In doing so, the old wide concept of
Fusicladium, including all kinds of Venturia anamorphs (see chapter 2.2.), is reintroduced. The new circumscription of Fusicladium can be given as follows:
Fusicladium Bonord., Handb. Mykol.: 80 (1851), emend.
(nom. cons. prop., anamorphs of Acantharia Theiss. & Syd., Apiosporina Höhn.,
Protoventuria Berl. & Sacc., Venturia Sacc., Venturiaceae).
Type species: F. virescens Bonord. (= F. pyrorum).
= Spilocaea Fr., Novit. fl. svec. 5: 79 (1819), nom. rej. prop., type species: S. pomi Fr.
= Cycloconium Castagne, Cat. pl. Marseille: 220 (1845), nom. rej. prop., type species: C. oleagineum Castagne.
= Napicladium Thüm., Hedwigia 14: 3 (1875), type species: N. soraueri Thüm.
= Basiascum Cavara, Atti Ist. Bot. Univ. Pavia, Ser. 2, 1: 433 (1888), type species:
B. eriobotryae Cavara.
= Hormocladium Höhn., Ber. Deutsch. Bot. Ges. 37: 156 (1919), type species:
Fusicladium kaki Hori & Yoshino.
= Fusicladiopsis Karak. & Vassiljevsky, in Vassiljevsky & Karakulin, Parazitnye
nesovershennye griby, Ch. 1. Gifomitsety: 209 (1937), nom. illeg. (homonym),
type species: not indicated.
= Pollaccia E. Bald. & Cif., Atti Ist. Bot. “Giovanni Briosi“ 10: 71 (1937), type
species: P. radiosa (Lib.) E. Bald. & Cif.
= Megacladosporium Vienn.-Bourg., Les champignons parasites des plantes cultivées
1: 489 (1949), nom. superfl.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
9
= Ramalia Bat., Revista Biol. (Lisbon) 1: 111 (1957), type species: R. veronicae Bat.
= Karakulinia N.P. Golovina, Novosti Sist. Nizsh. Rast. 1: 213 (1964), type species:
K. cerasi (Rabenh.) N.P. Golovina.
= Fusicladosporium Partridge & Morgan-Jones, Mycotaxon 85: 360 (2003), type
species: F. carpophilum (Thüm.) Partridge & Morgan-Jones.
On leaves, fruits and twigs, causing leaf spots, scab diseases, necroses and deformations. Colonies punctiform, scattered, caespitose or dendritic, olivaceous, olivaceousbrown, dingy grey to blackish. Mycelium internal, subcuticular, intraepidermal to intercellular, sometimes substomatal, often forming expanded radiating hyphal or stromatic
strands or plates, rarely external, superficial. Stromata lacking to well-developed,
pseudostromatic, composed of rounded to isodiametric swollen hyphal cells, pigmented,
wall often somewhat thickened. Conidiophores solitary, arising from internal or rarely
external hyphae or stromatic hyphal aggregations, or fasciculate, arising from internal
hyphae or stromata, erumpent, occasionally emerging through stomata, sometimes forming well-developed sporodochial conidiomata, conidiophores often reduced to
conidiogenous cells or composed of several cells, erect, cylindrical, pyriform, subclavate,
narrowly obclavate, slightly to distinctly geniculate–sinuous, unbranched or occasionally branched, continuous to pluriseptate throughout, pigmented, pale olivaceous to
dark brown, tips often paler, smooth to verruculose, wall thin to somewhat thickened.
Conidiogenous cells integrated, terminal or intercalary or conidiophores reduced to
single conidiogenous cells, unilocal (monoblastic), determinate to multilocal (polyblastic), proliferation percurrent or sympodial, occasionally occurring together;
conidiogenous loci terminal or lateral, often denticle-like, somewhat protuberant, apex
truncate to slightly convex, wall unthickened or almost so, non-pigmented or slightly
darkened–refractive. Conidia solitary or catenate, in simple or branched acropetal chains,
amero-, didymo- to phragmosporous, ellipsoid–ovoid, obovoid, fusiform, obclavate–
subcylindrical, straight to curved, 0–3(–4)-euseptate, subhyaline to medium brown, but
mostly olivaceous, usually non-constricted at the septa, smooth to verruculose, ends
pointed or rounded to truncate, wall thin to somewhat thickened, hila unthickened or
almost so, occasionally somewhat darkened–refractive.
7.
The species concept
Venturia species and their anamorphs are, as far as known, host specific, mostly confined to a single host genus or at least allied host genera in a single host family.
Plurivorous taxa are unknown and this has been confirmed by preliminary molecular
examinations. Hence, the morphological differentiation between and keys to the species concerned can be based on host plant families. The following features are useful
and applicable at species rank:
- Symptoms, lesions.
- Mycelium (internal, external).
- Arrangement of conidiophores (solitary, fasciculate, in sporodochia).
- Shape, length and septation of conidiophores.
- Conidiogenous cells (terminal, intercalary, conidiophores reduced to conidiogenous
cells; determinate or proliferation percurrent, sympodial; number of loci, width).
10
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-
Conidia (formation, solitary or catenate; shape, size, septation, wall smooth
or verruculose).
The following characteristics are either more or less uniform within Venturia anamorphs
or very variable and thus less appropriate for the discrimination of species:
- Structure of the mycelium.
- Width of the conidiophores.
- Structure of the conidiogenous loci (uniform).
- Degree of pigmentation of conidiophores and conidia.
8.
Key to Fusicladium emend. and similar genera
1 Conidiophores and conidia colourless; conidiogenous loci conspicuous, somewhat thickened and darkened; conidia solitary or catenate, hila also slightly thickened and darkened ………………………………………………...…. Ramularia
1* Conidiophores and conidia pigmented ………………………..................…….. 2
2 Conidiogenous loci inconspicuous or denticle-like, but wall of the loci always
unthickened ………………….……………………………………….………….. 3
2* Conidiogenous loci conspicuous, thickened and darkened ……...…..…………. 6
3 Mycelium and stromata usually subcuticular to intraepidermal, hyphae often radiating, forming hyphal or stromatic strands or plates, membranous (Fusicladiumlike growth), rarely substomatal; conidiophores solitary, fasciculate or in
sporodochial conidiomata, usually erumpent through the cuticle, rarely emerging
through stomata or arising from superficial hyphae; conidia amero-, didymo- to
phragmosporous …………..............................................................…………... 4
3* Mycelium not Fusicladium-like, immersed or superficial, neither radiating nor
forming hyphal plates; stromata usually substomatal; conidiophores usually emerging through stomata ………....………....................………………...………….. 5
4 Mycelium and stromata membranous; older conidiophores usually curved by having unequally thickened walls; conidiogenous cells unilocal (monoblastic), usually determinate, rarely percurrent, conidiogenous loci distinctly thickened and
darkened; conidia solitary, 1-septate, broad (6–14 µm) ….…..…..… Fusicladiella
4* Mycelium and stromata usually radiating; conidiogenous cells uni- to multilocal
(mono- to polyblastic), but wall of the conidiophores equally thickened, not distinctly curved, percurrent or sympodial; loci always unthickened, at most somewhat darkened–refractive; conidia solitary or catenate, amero- to phragmosporous,
0–2(–4)-septate, < 10 µm …………………………………… Fusicladium emend.
5 Conidia scolecosporous, usually pluriseptate ………….……... Pseudocercospora
5* Conidia amero- to phragmosporous, 0–1(–3)-septate ……………….. Denticularia
6 Conidiogenous loci more or less protuberant, coronate (with a central convex
‘dome’ surrounded by a raised rim, i.e., Cladosporium type) ……. Cladosporium
6* Conidiogenous loci uniformly thickened and darkened, more or less truncate
(planate), without raised rim ………..…………………………………………. 7
7 Conidia formed singly, coarsely verrucose–echinulate …………… Asperisporium
7* Conidia formed singly or in chains, smooth to faintly rough-walled …… Passalora
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
11
9.
Index/key to Fusicladium species by host genera
This list contains all species of Fusicladium emend., including Pollaccia and Spilocaea
species. The host genera are alphabetically arranged under the names of their host families.
Aceraceae:
On Acer, Fusicladium humile
Apiaceae:
On Angelica, Fusicladium peucedani
On Cicuta, Fusicladium peucedani
On Glehnia, Fusicladium peucedani
On Lomatium, Fusicladium peucedani
On Peucedanum, Fusicladium peucedani
On Sphaenosciadium, Fusicladium peucedani
Asteraceae:
On Solidago, Fusicladium virgaureae
Betulaceae:
On Betula
1 Conidiophores yellowish brown to dark brown; loci unthickened, not darkened; conidia consistently solitary; on different species of Betula ..… Fusicladium betulae
1* Conidiophores olivaceous to pale brown; loci unthickened, but darkened; conidia
solitary or sometimes in short chains; on B. populifolia .... Fusicladium scribnerianum
Convolvulaceae:
On Calystegia, Fusicladium convolvularum
On Convolvulus, Fusicladium convolvularum
Corylaceae:
On Carpinus, Fusicladium carpineum
Crassulaceae:
On Sedum, Fusicladium caulicola
Cyperaceae:
On Carex, Fusicladium caricinum
Dipsacaceae:
On Succisa, Fusicladium consors
Ebenaceae:
On Diospyros, Fusicladium levieri
Euphorbiaceae:
On Euphorbia
1 Conidiophores in small, loose fascicles, arising from hyphae, 37–175 × 4–5 µm;
conidia 8.5–16 × (3–)4–6.5 µm, mostly aseptate, rarely with a single septum
……………………………..…….......…………...…... Fusicladium fasciculatum
1* Conidiophores in dense fascicles, arising from stromata, shorter, 10–90 × 4–6
µm; conidia longer, 10–22(–40) × 3–6 µm, 0–3 septa ......................................... 2
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Schlechtendalia 9 (2003)
2 Conidia solitary or in short, unbranched chains, rarely in branched chains, 0–2(–
3)-septate; conidiophores thin-walled; on different Euphorbia-species, not on
Euphorbia brittingeri .....................................………….. Fusicladium euphorbiae
2* Conidia solitary, 0–1-septate, lower cell mostly larger than the upper cell, rarely
with 3 septa; stromatic cells and conidiophores darker (olivaceous-brown) and
wall thicker; on Euphorbia brittingeri ......…………….…..... Fusicladium fautreyi
On Hevea, Fusicladium heveae
Fabaceae:
On Astragalus, Fusicladium brevipes
On Lathyrus, Fusicladium lathyrinum
On Pisum, Fusicladium pisicola
On Psoralea, Fusicladium psoraleae
Fagaceae:
On Quercus, Fusicladium-state of Acantharia echinata
Juglandaceae:
On Carya, Fusicladium effusum
Juncaceae:
On Juncus, Fusicladium junci
Liliaceae:
On Scilla, Fusicladium scillae
Malpighiaceae:
On Byrsonima, Fusicladium byrsonimatis
Oleaceae:
On Fraxinus
1 Conidiophores solitary, effuse, subglobose to ampulliform, campanulate, 5–12 ×
5–7 µm, yellowish brown; proliferation percurrent; conidia verruculose or minutely
echinulate ………………………………………………. Fusicladium nebulosum
1* Conidiophores in loose to dense fascicles, sometimes sporodochial, subcylindrical
to conical, 12–35 × 3–5 µm, pale olivaceous; proliferation sympodial and
percurrent; conidia smooth …………....................................... Fusicladium fraxini
On Olea, Fusicladium oleagineum
On Phillyrea, Fusicladium phillyreae
On Syringa, Fusicladium diedickeanum
Rosaceae:
On Amelanchier
1 Conidiophores 30–225 × 4 µm; proliferation sympodial; anamorph immediately formed
on the teleomorph; North America ............. Fusicladium-state of Apiosporina collinsii
1* Conidiophores 10–23 × 5–6 µm; proliferation percurrent; anamorph separately
formed from the teleomorph; Europe, New Zealand ..........….... Fusicladium pomi
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
13
On Aronia,
1 Proliferation sympodial; loci 1–3 µm wide ......................... Fusicladium pyrorum
1* Proliferation percurrent; loci 4–5 µm wide ............................... Fusicladium pomi
On Chaenomeles, Fusicladium pyrorum
On Cotoneaster, Fusicladium pomi
On Crataegus, Fusicladium crataegi
On Docynia, Fusicladium pomi
On Eriobotrya,
1 Proliferation sympodial; loci 1–3 µm wide .......…................ Fusicladium pyrorum
1* Proliferation percurrent; loci 4–5 µm wide ........................……. Fusicladium pomi
On Heteromeles, Fusicladium pomi
On Kageneckia, Fusicladium pomi
On Malus,
1 Conidiogenous cells sympodial .......................................................................... 2
1* Conidiogenous cells percurrent; conidia solitary, not catenate, 12–30 × 6–10 µm;
cosmopolitan ………...........................................….................... Fusicladium pomi
2 Conidia solitary, 10–34 × 5–11 µm; cosmopolitan ................ Fusicladium pyrorum
2* Conidia solitary or in short, sometimes branched chains, 9.5–17 × 3–5 µm; New
Zealand, North America....................………………......... Fusicladium asperatum
On Photinia, Fusicladium pomi
On Prunus,
1 Conidiogenous cells percurrent ..................……....................... Fusicladium pomi
1* Conidiogenous cells sympodial .......................................................................... 2
2 On twigs of the host plant, anamorph formed on the teleomorph; conidia ovoid,
obovoid, ellipsoid or irregular, 4–19 × (2–)3–6 µm, often laterally fused in pairs;
North America .........………………... Fusicladium-state of Apiosporina morbosa
2* On leaves and fruits, anamorph formed separately from the teleomorph; conidia
mostly fusiform, cylindrical or obclavate, larger, 9–30 × 4–7(–10) µm, not fused
in pairs ..............................................................................................………...... 3
3 Colonies consistently formed on leaves, dendritic; conidiophores 40–120 × 6–8
µm; conidia solitary, hila 2–4.5 µm wide; South America; on Prunus capollin and
P. serotina .......................................................................…. Fusicladium obducens
3* Colonies formed on fruits and leaves, velvety, caespitose; conidiophores narrower,
3–6(–7) µm wide; conidia solitary or in chains, hila 1–2(–2.5) µm wide …........ 4
4 Conidia solitary, rarely in chains; conidiophores (10–)20–40(–60) × (3–)4–6(–7)
µm, on a few hosts only, mostly on Prunus cerasus ………..... Fusicladium cerasi
4* Conidia consistently catenate; conidiophores 25–100 × 3–6 µm; with a wide host
range, but very rarely on Prunus cerasus ...................…. Fusicladium carpophilum
On Pyracantha, Fusicladium pomi
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SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
15
On Pyrus,
1 Conidiogenous cells percurrent ..................…………........................................ 2
1* Conidiogenous cells sympodial ..........................……....................................... 3
7* Conidia often curved; occurring on various species of Populus, incl. P. alba, P.
grandidentata and P. tremuloides; North America
.................................................................... Fusicladium radiosum var. lethiferum
2 Conidia obclavate, 25–45 × 6–8 µm, 1(–2)-septate; on P. pashia; Pakistan
................................................................................................ Fusicladium ahmadii
2* Conidia obpyriform to clavate, shorter and wider, 12–30 × 6–10 µm, 0–1-septate;
cosmopolitan ………......……………………………...…......... Fusicladium pomi
On Salix,
1 Conidiogenous cells polyblastic, sympodial ...…….............…………............... 2
1* Conidiogenous cells monoblastic or percurrent ................………...............…... 3
3 Conidia 9–20(–28) × 5.5–10 µm; on Chinese and Japanese species of Pyrus
........................................………......................................... Fusicladium nashicola
3* Conidia significantly longer, 10–34 × 5–11 µm; only on European species of Pyrus
……………………...….………………............................... Fusicladium pyrorum
On Rubus, Fusicladium grayianum
On Sorbus, Fusicladium pomi
On Spiraea, Fusicladium spiraeae
Salicaceae:
On Populus,
1 Conidiogenous cells polyblastic, with a single to several conidiogenous loci,
subdenticulate, 5–35 × 3–10 µm ......…................... Fusicladium martianoffianum
1* Conidiogenous cells mostly monoblastic (unilocal), not or only slightly denticulate, shorter and narrower, 5–25 × 2–5 µm .....…………..................................... 2
2 Conidiophores monoblastic, rarely with two or several conidiogenous loci, loci
1–3 µm wide; conidia solitary or in unbranched or branched chains ................... 3
2* Conidiophores always monoblastic, loci ca. 5 µm wide; conidia always solitary
............................................................................................................................. 4
3 Conidiomata mostly sporodochial; conidiogenous cells sympodial, not percurrent,
loci 1–2 µm wide; conidia aseptate, very rarely with a single septum
........................................................................................ Fusicladium romellianum
3* Conidiophores in small to large fascicles, rarely sporodochial; conidiogenous cells
sympodial as well as often percurrent with a single to several annellations, loci
1.5–3 µm wide; conidia (0–)1(–3)-septate …....................... Fusicladium subsessile
4 Conidia relatively large, 23–39 µm long ............................................................ 5
4* Conidia smaller, up to 28 µm long (Fusicladium radiosum s.lat.) …..............…. 6
5 Conidia often curved, fusiform to obclavate, 7–10 µm wide, (0–)2–3(–4)-septate;
on P. simonii × P. nigra; China ……..………………. Fusicladium mandshuricum
5* Conidia straight, rarely curved, ellipsoid to broadly fusiform, 9–14 µm, (0–)2(–3)septate; on P. balsamifera and P. nigra; Europe, North America ... Fusicladium elegans
6 Conidia (0–)1(–2)-septate, 8–10 µm wide; on P. alba; Europe, North Africa
...........……….........................………... Fusicladium radiosum var. populi-albae
6* Conidia mostly 1–2-septate, narrower, 5–8 µm wide ..……...........…................. 7
7 Conidia straight, rarely curved, on various species of Populus, but not on P.
grandidentata and P. tremuloides ..............… Fusicladium radiosum var. radiosum
2 Conidiophores 40–95(–130) × 3–5 µm, arising from stromata; conidia (10–)12–
20 × 3–4(–5) µm .………..…...........…………..........………. Fusicladium sp. (1)
2* Conidiophores arising as short lateral branchlets from hyphae; conidia 20–25 ×
5–8 µm ..............................………...….........……..........…… Fusicladium sp. (2)
3 Conidia in unbranched, rarely branched chains, mostly 0(–2)-septate
...................................................................................... Fusicladium catenosporum
3* Conidia solitary, 1–2-septate ...........….....…………….. Fusicladium saliciperdum
Scrophulariaceae:
On Parahebe, Fusicladium veronicae
Verbenaceae:
On Vitex, Fusicladium viticis
10.
The species of Fusicladium emend.
The species of Fusicladium emend. (incl. Pollaccia and Spilocaea) are alphabetically arranged. References to the original descriptions, type material, synonyms,
teleomorphs (as far as known), references to important descriptions and illustrations
in literature, exsiccatae, comprehensive descriptions, host range and distribution,
material examined and notes are given for each species. The drawings, mostly based
on type or other original material, have been prepared at a ration of 1 : 100 (bar = 10
µm). The exsiccatae cited have generally been examined. If only few collections
have been examined (up to five), all of them are listed, whereas numerous specimens
(more than five) are not cited in order to save space. In the latter case, the acronyms
of the herbaria, in which the numerous collections examined are housed, are mentioned. Herbarium acronyms are based on HOLMGREN et al. (1990), abbreviations of
author names follow BRUMMITT & POWELL (1992), and those of journals agree with
the system introduced by LAWRENCE et al. (1968), supplemented by BRIDSON & SMITH
(1991). The names of European countries are symbolised by the international standard abbreviations for the particular countries that are used for vehicles. Abbreviations
of the particular states of the USA follow FARR et al. (1989), and those of Canadian
provinces and territories are based on GINNS (1986). Names of all other countries are
given in full.
10.1.
Abbreviations
General: auct. = auctorum, ca. = circa, comb. nov. = new combination, comb. superfl. = superfluous
combination, fig. = figure, herb. = herbarium, incl. = inclusive, l.c. = locus citatus, nom. cons. =
nomen conservandum, nom. illeg. = nomen illegitimum, nom. inval. = nomen invalidum, nom. nov.
16
Schlechtendalia 9 (2003)
= nomen novum, nom. nud. = nomen nudum, p.p. = pro parte, s.lat. = sensu lato, s.str. = sensu stricto,
spp. = species. CMI Descr. = CMI Descriptions of Pathogenic Fungi and Bacteria, IMI Descr. = IMI
Descriptions of Fungi and Bacteria.
European countries: A = Austria, AL = Albania, B = Belgium, BG = Bulgaria, CH = Switzerland,
CS = former Czechoslovakia (incl. Czech Republic and Slovakia), CZ = Czech Republic, D = Germany, DK = Denmark (incl. Bornholm), E = Spain, EW = Estonia, F = France (incl. Corsica and, on
geographical grounds, the Channel Islands and Monaco,), FR = Faeroe Islands, GB = United Kingdom (with all islands, but excluding Northern Ireland and the Channel Islands), GR = Greece, H =
Hungary, HR = Croatia, I = Italy (with Sardinia and Sicily), IRL = Ireland (the whole island, north
and south), LT = Lithuania, LV = Latvia, N = Norway, NL = the Netherlands, P = Portugal (incl.
Azores), PL = Poland, RO = Romania, RUS = Russia, S = Sweden, SF = Finland, SK = Slovakia,
SLO = Slovenia, TR = Turkey (European part), Ukr. = Ukraine, YU = former Yugoslavia (incl.
Slovenia, Croatia, Serbia, etc.). Byelorussia and Moldavia are not abbreviated. Older records have
often been published from Yugoslavia and Czechoslovakia. Sometimes it is difficult or even impossible to refer such records to the particular succeeding countries, so the old names have to be maintained. They are to be understood as summarising abbreviations.
USA: AL = Alabama, AK = Alaska, CA = California, CO = Colorado, CT = Connecticut, DE =
Delaware, FL = Florida, GA = Georgia, IA = Iowa, ID = Idaho, IL = Illinois, KS = Kansas, KY =
Kentucky, LA = Louisiana, ME = Maine, MD = Maryland, MA = Massachusetts, MI = Michigan,
MN = Minnesota, MS = Mississippi, MO = Missouri, MT = Montana, NE = Nebraska, NH = New
Hampshire, NJ = New Jersey, NM = New Mexico, NY = New York, NC = North Carolina, ND =
North Dakota, NV = Nevada, OH = Ohio, OK = Oklahoma, OR = Oregon, PA = Pennsylvania, SC =
South Carolina, SD = South Dakota, TN = Tennessee, TX = Texas, UT = Utah, VA = Virginia, VT =
Vermont, WA = Washington, WI = Wisconsin, WY = Wyoming.
Canada: NWT = Northwest Territories, BC. = British Columbia, Alta. = Alberta, Sask. = Saskatchewan, Man. = Manitoba, Ont. = Ontario, Que. = Quebec, NB. = New Brunswick, NS. = Nova Scotia,
PEI = Prince Edward Island, Nfld. = Newfoundland, Yukon = Yukon Territory. (Labr. = Labrador).
Exsiccata:
Allesch. & Schn., F. bavar. = Allescher & Schnabl, Fungi bavarici.
Barthol., F. Columb. = Bartholomew, Fungi Columbiani.
Brenckle, F. Dakot. = Brenkle, Fungi Dakotenses.
Briosi & Cav., F. paras. = Briosi & Cavara, I funghi parassiti delle piante coltivate od utili, essicati,
delineati e descritti.
Calif. F. = California Fungi (distributed by the herbarium of the University of California).
Cooke, F. brit. exs. = Cooke, Fungi britannici exsiccati.
Crypt. exs. = Cryptogamae exsiccatae editae a Museo Hist. Nat. Vindobonensi.
Ellis or Ellis & Everh., N. Am. F. = Ellis or Ellis & Everh., North American Fungi, Ser. I or Ser. II.
Ellis & Everh., F. Columb. = Ellis & Everh., Fungi Columbiani.
Erb. Critt. Ital. = Erbario Crittogamico Italiano.
Erikss., F. paras. scand. = Eriksson, Fungi parasitici scandinavici exsiccati.
Fl. bav. = Starcs, Flora bavarica.
Fl. Suec. = Flora Suecica.
Fuckel, F. rhen. = Fuckel, Fungi rhenani.
F. est. = Fungi estonici.
F. latv. exs. = Smarods, Fungi latvici exsiccati.
Fl. Gall. Germ. exs. = Flora Gallici Germaniae exsiccatae.
Fl. Hung. exs. = Flora Hungarica exsiccata.
Fl. Olten. exs. = Flora Oltenia exsiccata.
Griffiths, West Am. F. = Griffiths, West American Fungi.
Herb. Mycol. Rom. = Herbarium Mycologicum Romanicum.
Jaap, F. sel. exs. = Jaap, Fungi selecti exsiccati.
Kab. & Bub., F. imp. exs. = Kabát & Bubák, Fungi imperfecti exsiccati.
Kellerm., Ohio Fungi = Kellerman, Ohio Fungi.
Kellerm. & Sw., Kans. F. = Kellerman & Swingle, Kansas Fungi.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
17
Krieger, F. sax. = Krieger, Fungi saxonici.
Krypt. exs. = Kryptogamae exsiccatae.
Lib., Pl. crypt. ard. = Libert, Plantae cryptogamicae guas in Arduenna collegit.
Lin., F. hung. = Linhart, Fungi hungarici.
Migula, Crypt. Germ. Austr. Helv. exs. = Migula, Cryptogamae Germaniae, Austriae et Helveticae exsiccatae.
Mycoth. Fenn. = Liro, Mycotheca Fennica.
Mycoth. Ross. = TranÓel’ et al., Mycotheca Rossica.
Neger, Forstschäd. P. = Neger, Forstschädliche Pilze.
Oudem., F. neerl. exs. = Oudemans, Fungi neerlandici exsiccati.
Petr., F. alban. bosn. exs. = Petrak, Fungi albanici et bosniae exsiccati.
Petr., F. polon. = Petrak, Fungi polonici.
Petr., Fl. Bohem. Morav. exs. = Petrak, Flora Bohemica et Moravica exsiccata.
Petr., Mycoth. gen. = Petrak, Mycotheca generalis.
Pilzfl. Sib. = Pilzflora Sibiriens.
Rabenh., F. eur. = Rabenhorst, Fungi europaei exsiccati (also Rabenhorst & Winter, F. eur. or
Rabenhorst, Winter & Pazschke, Fungi eur. et extraeur.).
Rabenh., Herb. mycol. = Rabenhorst, Herbarium mycologicum.
Rehm, Ascomyc. = Rehm, Ascomyceten.
Reliqu. Petrak. = Reliquiae Petrakianae.
Romell, F. exs. = Romell, Fungi exsiccati.
Roum., F. gall. exs. = Roumeguère, Fungi gallici exsiccati.
Roum., F. sel. exs. = Roumeguère, Fungi selecti gallici exsiccati.
Sacc., Mycoth. ital. = Saccardo, Mycotheca italica.
Sacc., Mycoth. Ven. = Saccardo, Mycotheca Veneta.
Schmarotzerp. Ruhrg. = Schmarotzerpilze des Ruhrgebiets.
Seym. & Earle, Econ. F. = Seymour & Earle, Economic Fungi.
Siem., F. bialow. exs. = Siemaszko, Fungi bialowiezenses exsiccati.
Solh., Mycofl. Sax. exs. = Solheim, Mycoflora Saximontanensis exsiccatae.
Syd., F. exot. exs. = H. Sydow, Fungi exotici exsiccati.
Syd., Mycoth. germ. = H. & P. Sydow, Mycotheca germanica.
Syd., Mycoth. march. = P. Sydow, Mycotheca marchica.
Thüm., F. austr. = de Thümen, Fungi austriaci.
Thüm., Herb. myc. oec. = de Thümen, Herbarium mycologicum oeconomicum.
Thüm., Mycoth. univ. = de Thümen, Mycotheca universalis.
Vestergr., Micromyc. rar. sel. exs. = Vestergren, Micromycetes rariores selecti exsiccati.
Westend. & Wall., Herb. crypt. belg. = Westendorp & Wallays, Herbier cryptogamique belge.
10.2.
Descriptions of species
10.2.1.
Fusicladium ahmadii (M.B. Ellis) Ritschel & U. Braun comb. nov.
Fig. 1
≡ Spilocaea ahmadii M.B. Ellis, More Dematiaceous Hyphomycetes: 114 (1976).
Holotype: on Pyrus pashia, Pakistan, Swat, Mingora, 10 Aug. 1952, S. Ahmad (IMI 81091).
Teleomorph: Unknown.
Ill.: ELLIS (1976: 113, Fig. 81).
On living leaves, leaf spots amphigenous, subcircular to irregular, 5–10 mm diam.,
whitish, surrounded by a brown, shinning margin. Colonies amphigenous, punctiform, aggregated in the centre, olivaceous to dark brown. Mycelium immersed. Stromata subcuticular, composed of subcircular to slightly angular, pale to medium
olivaceous-brown, somewhat thick-walled cells, 5–8 µm diam. Conidiophores loosely
to densely fasciculate, arising from the upper cells of the stromata, erumpent through
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Fig. 1: Fusicladium ahmadii. A – conidia, B – loose fascicle of conidiogenous cells with one to
several conspicuous annellations, scale = 10 µm, A. Ritschel del.
the cuticle, erect, straight to slightly flexuous, cylindrical to ampulliform, unbranched,
15–60 × 5–6 µm, 0–1-septate, pale to medium olivaceous-brown, smooth, thick-walled.
Conidiogenous cells integrated, terminal, with a single conidiogenous locus, proliferation percurrent, with up to eight conspicuous annellations at the distal end, loci 4–
5 µm wide, unthickened, not darkened. Conidia solitary, obclavate, straight to slightly
curved, (23–)25–38 × 6–8 µm, 1(–2)-septate, septum median or somewhat in the
lower half, more or less constricted at the septum, pale to medium olivaceous-brown,
smooth, walls of the lower cells somewhat thicker than those of the upper one, apex
narrowly pointed, base truncate, hila 4–5 µm wide, unthickened, not darkened.
Hosts and Distribution: on Pyrus spp. (Rosaceae), Asia – Pyrus pashia (Pakistan).
Notes: This species seems to be closely allied to F. pomi, but differs in having significantly longer
and narrower conidia. Conidiogenous cells illustrated by ELLIS (1976) seem to show up to two loci,
which could not be confirmed by a re-examination of type material.
Fig. 2
Fig. 2: Fusicladium asperatum. A – conidia, B – conidiogenous cells. C – small fascicle of
conidiophores, scale = 10 µm, K. Schubert del.
Holotype: on Malus sylvestris ‘Dougherty’, New Zealand, Distr. Auckland Prov., Waitemata Co.,
Oratia, Experimental Station, Sept. 1973, P.J. Brook (PDD 31846), anamorph only.
Teleomorph: Venturia asperata Samuels & Sivan., New Zealand J. Bot. 13: 646 (1975).
Lit.: Fungi Canadenses (No. 291), SIVANESAN (1977: 38–39; 1984a: 608).
Ill.: Fungi Canadenses (No. 291, Fig. 5), SAMUELS & SIVANESAN (1975: Figs 11, 12; Pl. 8, E),
SIVANESAN (1977: 39, Fig. 12 C, D; 1984a: 608, Fig. 365).
Maculae saepe epiphyllae, griseo-brunneae, margine irregulari. Coloniae caespitosae, atro-brunneae
vel nigrae. Hyphae subcuticulares. Stromata 35–60 µm diam., ex cellulis olivaceis vel brunneis,
crassitunicatis, pseudoparenchymaticis composita. Conidiophora solitaria vel subfasciculata, pauca,
10.2.2.
Fusicladium asperatum K. Schub. & U. Braun sp. nov.
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erecta vel flexuosa, non-ramosa, 30–150 × 4–5 µm, pluriseptata, olivacea vel brunnea, apicem versus pallidiora, levia, leniter crassitunicata, basi leniter inflata. Cellulae conidiogenae integratae, terminales, sympodiales. Cicatrices conidiales terminales, aggregatae, denticulatae, 1–2 µm latae, nonincrassatae, non-fuscatae–refractivae. Conidia solitaria vel catenata, fusiformes vel cylindrica, recta,
9.5–17 × 3–5 µm, 0(–1)-septata, flavissima vel pallide olivacea, levia vel verruculosa, leniter
crassitunicata, apice rotundato vel truncato, basi truncata, 1–2 µm lata, non-incrassata, non- vel
leviter fuscata–refractiva.
On living leaves causing distinct leaf spots, sometimes almost symptomless, spots
epiphyllous, greyish brown, margin irregular. Colonies caespitose, dark brown to
black. Hyphae subcuticular. Stromata 35–60 µm diam., composed of olivaceous to
brown, thick-walled, pseudoparenchymatous cells. Conidiophores solitary or in small
groups, straight, erect to flexuous, unbranched, 30–150 × 4–5 µm, pluriseptate,
olivaceous to brown, paler towards the apex, smooth, with somewhat thickened walls,
sometimes base somewhat swollen. Conidiogenous cells integrated, terminal, with
several denticle-like conidiogenous loci, mostly crowded at the apex, proliferation
sympodial, loci 1–2 µm wide, unthickened, slightly darkened–refractive. Conidia solitary or catenate, in short, simple or sometimes branched chains, fusiform to cylindrical, straight, 9.5–17 × 3–5 µm, 0(–1)-septate, yellowish to pale olivaceous, sometimes subhyaline, smooth to verruculose, walls somewhat thickened, apex rounded or
truncate, base truncate, hila 1–2 µm wide, unthickened, not or very slightly darkened–refractive.
Hosts and Distribution: on Malus spp. (Rosaceae), New Zealand, North America –
Malus pumila (New Zealand), M. sylvestris (New Zealand; North America, Canada,
Ont.).
Notes: In culture this fungus grows Cladosporium-like. The conidiophores are often determinate
and form conidia in short, branched chains (Fungi Canadenses No. 291). The type material of F.
asperatum is also paratype material of Venturia asperata. Since the anamorph may occur independently of the teleomorph, we prefer to propose a separate name for this stage.
10.2.3.
Fusicladium betulae Aderh., Centralbl. Bakteriol., 2. Abth., 2: 57 (1896)
and Hedwigia 36: 80 (1897)
Fig. 3
Neotype: on Betula pendula (= Betula alba), Germany, Berlin, Grunewaldmoor, 13 Jul. 1923, Laubert
(B), selected here.
Teleomorph: Venturia ditricha (Fr.) P. Karst., Mycol. fenn. 2: 188 (1873).
Lit.: LINDAU (1907: 778), VASSILJEVSKY & KARAKULIN (1937: 13), BARR (1968: 813–814),
SIVANESAN (1977: 61–63; 1984a: 613–614), SAGDULLAEVA (1990: 54).
Ill.: KARSTEN (1873: 188, Figs 29, 30; Pl. 1, A), ADERHOLD (1897: Tab. 4, Fig. 1), SIVANESAN
(1977: 62, Fig. 30; 1984a: 614, Fig. 370 B).
Exs.: Krieger, F. sax. 232 b; Rehm, Ascomyc. 597; Siem., F. bialow. exs. 197; Syd., Mycoth. march.
982; Thüm., Mycoth. univ. 350.
On living leaves, occasionally also on petioles, spots amphigenous, subcircular to
somewhat irregular, punctiform, 1–4 mm wide, scattered, brown, later darker to black,
sometimes with a paler margin or a paler centre. Colonies amphigenous, punctiform,
scattered, black. Mycelium subcuticular, radiating, hyphae 3–4 µm wide, septate,
hyaline to pale yellowish, stroma composed of thick-walled, brown cells, forming up
to three layers. Conidiophores solitary or in loose fascicles, arising from stromata,
Fig. 3: Fusicladium betulae. A – conidia, B – conidiophore germinating with hyphae, C – fasciculate
conidiophores, scale = 10 µm, K. Schubert del.
erumpent through the cuticle, erect, straight to flexuous, unbranched, 25–100 × (3–)
4–6 µm, 0(–1)-septate, yellowish brown to dark brown, paler towards the apex, apex
sometimes subhyaline, smooth, later occasionally rough-walled, wall somewhat thickened, base mostly swollen, up to 10 µm. Conidiogenous cells integrated, terminal,
with a single or several conidiogenous loci, subdenticulate, proliferation sympodial,
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loci unthickened, not darkened–refractive, 2–3 µm wide. Conidia solitary, fusiform
to obclavate or clavate, straight to slightly curved, 12–30(–34) × 5–9 µm, yellowish to
pale olivaceous, (0–)1(–2)-septate, often constricted at the septa, septum mostly somewhat in the upper half, smooth to verruculose, apex obtuse, rounded to pointed, base often
elongated, hila truncate or slightly convex, unthickened, not darkened, 2–3 µm wide.
Hosts and Distribution: on Betula spp. (Betulaceae), Asia, Caucasus, Europe, North
America – Betula glandulosa (North America, Canada, Que.), B. litwinowii (Caucasus, Georgia), B. nana (Europe, DK, N, RO), B. papyrifera (North America, Canada,
Ont.), B. pendula (Asia, Kazakhstan, Uzbekistan; Europe, A, D, DK, GB, PL, RO,
RUS, Ukr.), B. populifolia (North America, USA, ME, NY), B. pubescens (Europe,
D, DK, S), Betula spp. (Asia, Kirghizia, Uzbekistan; Caucasus, Armenia; Europe, D,
EW, GB, H, RUS, Byelorussia; North America, Canada, Ont., Que.).
Material examined: collections from B, LE.
Notes: Parts of Aderhold’s herbarium, previously housed at the “Biologische Bundesanstalt Berlin”
(BBA), were transferred to the herbarium of the “Botanical Garden Berlin-Dahlem” (B), but the type
material of Fusicladium betulae unfortunately was not preserved. Consequently, it is necessary to
propose a neotype for this species.
SIVANESAN (1977) reduced Asteroma betulae Roberge ex Desm., Ann. Sci. Nat. Bot. 19: 349 (1843),
to synonymy with Fusicladium betulae. The examination of lectotype material (Desm., Pl. Crypt. N.
France, 1346; PC) showed that only immature ascomata of the teleomorph, Venturia ditricha, with
setae, stromata and some remnant of conidiophores were present, but conidia could not be traced.
Hence, Asteroma betulae should rather be considered a synonym of Venturia ditricha.
In one collection, conidiophores germinating with hyphae have been seen.
LIND (1937) described Venturia ditricha from Alnus incana and cited STRASSER (1907: 314), who
also recorded it from Berberis vulgaris. These records are very doubtful.
Fig. 4: Fusicladium brevipes. A – conidia, B – conidiogenous cells, scale = 10 µm, K. Schubert del.
Hosts and Distribution: on Astragalus spp. (Fabaceae), North America, in the western parts of the USA – Astragalus bisulcatus (USA, MT), A. canadensis (USA, WA),
A. hypoglottis (USA, CO).
10.2.5.
Fusicladium byrsonimatis (U. Braun & Mouch.) U. Braun comb. nov.
Fig. 5
Holotype: on Astragalus hypoglottis, USA, Colorado, Musie Pass, Sangre de Christo Range, 16 Jul.
1888, C.H. Demetrio (NY).
Teleomorph: Unknown.
Lit.: SACCARDO (1892: 598).
Ramalia byrsonimatis U. Braun & Mouch., Mycol. Res. 104(8): 1010 (2000); holotype: on
Byrsonima sp., Brazil, Minas Gerais, Belo Horizonte, 8 Sept. 1913, A. Maublanc, Fungi
Brasilienses 180 p.p. (PC, ex herb. ‘Station Centrale de Pathologie Végétale, Versailles’, mixed
infection with Pseudocercospora byrsonimatis).
Teleomorph: Unknown.
Ill.: BRAUN & MOUCHACCA (2000: 1010, Figs 4–5).
On living leaves, lesions inconspicuous or causing diffuse discolorations. Colonies
hypophyllous, variable in shape and size, effuse, dense, sometimes confluent, dark greyish brown to blackish. Mycelium internal, forming well-developed, loose to dense stromatic hyphal aggregations in thin layers, subcuticular to intraepidermal, cells oblong to
subglobose, 2–15 µm diam., at first subhyaline, pale yellowish-greenish, later brown.
Conidiophores arising from stroma cells, solitary, in loose to dense fascicles or in effuse
layers, erumpent, erect, straight, subcylindrical, conical to flexuous, somewhat geniculate–sinuous, unbranched, 5–25(–30) × 3–9 µm, 0–1-septate, pale to medium brown,
smooth, thin-walled, conidiophores usually reduced to conidiogenous cells, unilocal, determinate, occasionally with 2–3 loci, proliferation sympodial, loci subtruncate to convex, 2–4 µm wide, unthickened, not darkened. Conidia solitary, ellipsoid to ovoid, oblong, 15–35 × 7–13 µm, aseptate (according to the original description later with 1–2
septa), pale olivaceous to olivaceous-brown, almost smooth to finely asperulate, apex
broadly rounded, base subtruncate to rounded, hila unthickened, not darkened.
Leaf spots amphigenous, subcircular to slightly angular–irregular, small, 0.5–2 mm
wide, grey, greyish brown, margin narrow, blackish brown, somewhat raised. Colonies mainly hypophyllous, punctiform, dense, medium to dark brown. Mycelium internal, subcuticular to intraepidermal. Stromata well-developed, composed of swollen hyphal cells, 2–8 µm wide, yellowish to medium brown. Conidiophores solitary
or in small loose groups, arising from stromata, erect to decumbent, straight,
subcylindrical to somewhat geniculate–sinuous, subnodulose, apically subdenticulate
to conspicuously denticulate, unbranched, 20–100 × 3–7 µm, pluriseptate, pale to
medium brown, smooth, wall somewhat thickened, paler and thin-walled towards the
apex. Conidiogenous cells integrated, terminal, proliferation sympodial, 10–30 µm
long, subdenticulate to denticulate, denticles minute, conically truncate, wall of the
loci neither thickened nor darkened. Conidia solitary or catenate, occasionally in
branched chains, solitary conidia obovoid, catenate conidia ellipsoid–fusiform, straight,
8–18 × 4–7 µm, 0–1-septate, light brown, smooth, wall thin or somewhat thickened,
10.2.4.
Fusicladium brevipes Ellis & Everh., J. Mycol. 5: 69 (1889)
Fig. 4
≡
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25
Fig. 5: Fusicladium byrsonimatis.
A – conidia, B – conidiophores, scale
= 20 µm, U. Braun del.
apex broadly rounded or
obconically truncate, base
obconically truncate, 1–2.5 µm
wide, hila unthickened, not
darkened.
Hosts and Distribution: only
known from the type.
10.2.6.
Fusicladium caricinum Bres., in P. Syd., Mycoth. march., Cent. XLI, No.
4065 (1894)
Fig. 6
Lectotype: on Carex acutiformis, Germany, Berlin, near Zehlendorf, Sept. 1893, P. Sydow, Mycoth.
march. 4065 (HBG), selected here; isolectotypes: on “Carex ampullacea“, Germany, Berlin, between Zehlendorf and Klein-Machnow, 1893, P. Sydow (B) and Syd., Mycoth. march. 4065.
Teleomorph: Unknown.
Lit.: SACCARDO (1895: 618), LINDAU (1907: 775), IMI Descr. (No. 1511).
Ill.: IMI Descr. (No. 1511, Figs A–B).
Exs.: Fl. bav. 2831; Rabenh., F. eur. 4294; Syd., Mycoth. march. 4065.
Leaf spots amphigenous, elliptical–oblong to irregular, 2–5 × 1–5 mm, pale yellowish-olivaceous, brownish, surrounded by a narrow, medium to dark brown or blackish margin. Colonies amphigenous, effuse, punctiform, dark, fructification mostly
sparse. Conidiophores solitary or up to three in small fascicles, erumpent, erect, straight
to slightly curved, subglobose, broadly ampulliform to short cylindrical, unbranched,
8–23 × (6–)10–19 µm, aseptate, medium to dark brown, smooth, walls thin to somewhat thickened, broadly truncate at the apex, conidiophores reduced to conidiogenous
cells, unilocal, determinate, conidiogenous loci truncate to slightly convex, occasionally rounded, 3–5 µm wide, unthickened, not or only very slightly darkened–
refractive. Conidia solitary, fusiform to broadly obclavate, straight to flexuous, (20–)
30–50(–57) × 10–15(–17) µm, (0–)1(–2)-septate, sometimes slightly constricted at
the septa, septa more or less median or often in the lower half, pale olivaceous to pale
brown, smooth, wall thin to somewhat thickened, pointed at the apex, base mostly
slightly oblique, hila 3–6(–7) µm wide, unthickened, not darkened.
Fig. 6: Fusicladium caricinum. A – conidia, B – conidiogenous cells, scale = 10 µm, K. Schubert del.
Hosts and Distribution: on Carex spp. (Cyperaceae), Europe – Carex acutiformis
(D, LV), C. riparia (D), C. vesicaria (LV), Carex spp. (H).
Material examined: collections from B, HBG, JE, LE, M.
Notes: In the original diagnosis, the conidiophores were described as hyaline but the examination of
type material showed that they are medium to dark brown.
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10.2.7.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
27
Fusicladium carpineum (Ellis & Everh.) U. Braun & K. Schub., IMI
Descriptions of Fungi and Bacteria 152, No. 1512 (2002)
Fig. 7
Fusicladium effusum [G. Winter] var. carpineum Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 1891: 91 (1891); lectotype: on Carpinus americana, Canada, London, Oct. 1889, J. Dearness (NY), selected here; isolectotypes: DAOM, M.
≡ Cladosporium caryigenum [(Ellis & Langl.) Gottwald] var. carpineum (Ellis & Everh.) Gottwald,
Mycologia 74(3): 389 (1982), comb. inval.
Teleomorph: Unknown.
Lit.: SACCARDO (1892: 598).
Ill.: IMI Descr. (No. 1512, Figs A–B).
Exs.: Ellis & Everh., N. Am. F. 2793.
≡
Leaf spots mainly epiphyllous, subcircular to angular–irregular, 1–5 mm wide, dingy
yellowish to greyish brown, scattered. Colonies punctiform to subeffuse, dark brown.
Mycelium internal, mainly subcuticular, hyphae about 2 µm wide, septate, hyaline to
pale yellowish. Stromata variable, small to large, up to 150 µm diam., composed of
swollen hyphal cells, 5–12 µm diam., brown, thick-walled. Conidiophores in loose
fascicles, arising from stromata, erect, straight to curved or somewhat geniculate–
sinuous, unbranched, 90–300(–340) × 5–7(–8) µm, pluriseptate, brown, paler towards
the apex, smooth, thick-walled, occasionally with a percurrent proliferation which is
not connected with conidiogenesis. Conidiogenous cells integrated, terminal, 10–40
µm long, with a single or several inconspicuous to subdenticulate conidiogenous loci,
1.5–3 µm wide, wall of the loci unthickened, not darkened. Conidia catenate, in simple
or branched chains, pyriform, ellipsoid, fusiform, straight to slightly curved, 10–21×
(5–)6–8(–10) µm, aseptate, pale olivaceous to pale brown, smooth, thin-walled, attenuated towards apex and base, apex rounded, pointed or truncate, base truncate,
hila 1.5–3 µm wide, unthickened, not darkened.
Hosts and Distribution: on Carpinus spp. (Corylaceae), North America – Carpinus
americana (Canada), C. caroliniana (USA, GA, WI).
Material examined: collections from B, M, NY.
Notes: This fungus is morphologically closely allied to Fusicladium effusum G. Winter on Carya
species (Juglandaceae), but differs in its occurrence on unrelated hosts (on Carpinus species,
Corylaceae), distinct lesions, much longer and wider conidiophores with pale tips and less conspicuous conidiogenous loci. Based on these differences, and since Venturia species with phytopathogenic anamorphs are generally confined to related hosts of a single host plant family, F. effusum var.
carpineum is considered a separate species.
Fusicladium carpini Osipyan (OSIPYAN 1971), described from Armenia on Carpinus caucasica, is a
quite distinct species with very short conidiophores (10–17 × 3.5–7.5 µm) and narrower, 0–1-septate
conidia (18.1–26.4 × 4.9–6 µm).
10.2.8.
≡
≡
≡
Fusicladium carpophilum (Thüm.) Oudem., Verh. Kon. Ned. Akad.
Wetensch., Afd. Natuurk. 1900: 388 (1900)
Fig. 8
Cladosporium carpophilum Thüm., Oesterr. Bot. Z. 27: 12 (1877); syntype: on Prunus persica
(= Persica vulgaris), Austria, Wien, 1877, Thüm., Herb. myc. oec. 599 (LE).
Megacladosporium carpophilum (Thüm.) Vienn.-Bourg., Les champignons parasites des plantes
cultivées 1: 489 (1949).
Fusicladosporium carpophilum (Thüm.) Partridge & Morgan-Jones, Mycotaxon 85: 362 (2003).
Fig. 7: Fusicladium carpineum. A – conidia, B – small fascicle of conidiophores, scale = 10 µm, K.
Schubert del.
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=
Fusicladium pruni Ducomet, Thèse Fac. Sci. Paris: 137 (1907); type: on fruits of Prunus domestica
var. “Ente“, France, Villeneuve (not seen).
= Fusicladium amygdali Ducomet, Ann. École Natl. Agric. Rennes 4: 11 (1911); type: on Prunus
dulcis (= Amygdalus communis), France, prope Ecole Nat. Agric. Rennes (not seen).
Teleomorph: Venturia carpophila E.E. Fisher, Trans. Brit. Mycol. Soc. 44: 339 (1961).
Lit.: BENSANDE & KEITT (1928: 313–329), HUGHES (1953: 568–569), ELLIS (1971: 315–317),
ONDÌEJ (1971: 167–168), CMI Descr. (No. 402), SUBRAMANIAN (1971: 235), SIVANESAN (1977:
45–47; 1984a: 609), PARTRIDGE & MORGAN-JONES (2003: 362).
Ill.: HUGHES (1953: 569, Fig. 9), SCHWEIZER (1958: 80, Fig. 20), ELLIS (1971: 316, Fig. 218),
ONDÌEJ (1971: 167, Fig. 1), CMI Descr. (No. 402, Figs D, E), SIVANESAN (1977: 46, Fig. 18; 1984a:
609, Fig. 366), PARTRIDGE & MORGAN-JONES (2003: 361, Fig. 1).
Exs.: Barthol., F. Columb. 2009; Ellis & Everh., F. Columb. 1164; Ellis & Everh., N. Am. F. 3588;
Roum., F. gall. exs. 3991; Thüm., Herb. myc. oec. 559.
On leaves, fruits and twigs, patches on the fruits superficial, on the exposed surface, circular to oval, small, brown, often confluent, forming large, brown areas,
leaf spots hypophyllous, brown. Colonies effuse or punctiform, dark olivaceous,
velvety. Mycelium subcuticular or subepidermal. Hyphae branched, 3–6 µm wide,
septate, olivaceous, forming a pseudoparenchymatous layer below the cuticle, stromatic cells thick-walled. Conidiophores solitary or in loose fascicles, erumpent,
erect, straight or somewhat flexuous, unbranched or occasionally branched, 25–
100 × 3–6 µm, septate, slightly constricted at the septa, olivaceous or medium to
dark brown, paler towards the apex, smooth, sometimes swollen at the base, walls
somewhat thickened. Conidiogenous cells integrated, terminal or intercalary, 10–
25 µm long, with one to several denticulate loci, (1–)1.5–2(–2.5) µm wide,
unthickened, not darkened. Conidia mostly in simple or branched chains, rarely
solitary, cylindrical to fusiform, sometimes short obclavate, straight, (9–)12–20(–
30) × (3–)4–5(–6) µm, 0(–1)-septate, then slightly constricted at the septum, pale
olivaceous, smooth to verruculose, hila truncate, (1–)1.5–2(–2.5) µm wide,
unthickened, not darkened.
Fig. 8: Fusicladium carpophilum. A – conidia, B – conidiophores, scale = 10 µm, K. Schubert del.
Hosts and Distribution: on Prunus s.lat. (Rosaceae), Asia, Caucasus, Europe, Africa, North America, South America, Australia, New Zealand (cosmopolitan) – P.
americana (North America, USA, FL), P. angustifolia (USA, FL), P. armeniaca
[Asia, China, Kirghizia; Caucasus, Armenia, Azerbaijan, Georgia; Europe, GB, RUS;
Africa, Morocco, Rhodesia (Zimbabwe), South Africa; North America, USA, CA,
FL, OK, TX; South America, Chile; Australia, Queensland, Tasmania, Victoria;
New Zealand], P. capollin (South America, Columbia), P. caroliniana (North
America, USA, FL), P. cerasifera (New Zealand), P. cerasus (Asia, Uzbekistan;
North America, USA, IA, NJ, NY ), P. domestica (Asia, Afghanistan; Caucasus,
Armenia; Europe, F, GB, RO; Africa, Kenya, Morocco; North America, USA, CA,
FL, IA; South America; Australia, New South Wales), P. dulcis (Asia, Lebanon,
Kazakhstan, Kirghizia, Uzbekistan; Caucasus, Armenia, Azerbaijan; Europe, F, GB,
Moldavia, RO; Africa, Libya, Morocco; North America, USA, CA, CT, OR, WA;
South America, Chile; New Zealand), P. laurocerasus (North America, USA, FL),
P. mahaleb (Asia, Lebanon), P. mandshurica (Asia, China), P. mume (Asia, China,
Guangdong Prov., Taiwan, Japan; New Zealand), P. munsoniana (North America,
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USA, FL, IA), P. padus (North America, USA, AK), P. persica [Asia, Afghanistan,
Jordan, Lebanon, Uzbekistan, China, India, Japan, Korea, Taiwan; Caucasus, Georgia; Europe, A, F, GB; Africa, Ethiopia, Kenya, Morocco, Mozambique, Zambia,
Rhodesia (Zimbabwe), South Africa; North America, Canada, Ont., USA, AL, CA,
CT, DE, IL, NC, NY, PA, TX; Central America, Guatemala; South America, Chile,
Columbia, Uruguay; Australia, Queensland, Tasmania, Victoria; New Zealand], P.
salicina (Africa, South Africa; South Australia; New Zealand), P. serotina (North
America, USA, FL), P. serrulata (North America, USA, MS), P. spinosa (Europe,
GB; South America), Prunus spp. (Asia, Japan; Europe, H; North America, Canada,
Que., USA, OK; South America).
Material examined: collections from JE, HBG, LE.
Notes: Previous authors, e.g., ELLIS (1976) and SIVANESAN (1977), often referred Fusicladium
carpophilum to Cladosporium. However, this species is an anamorph of Venturia with fusicladioid
conidiogenous loci and conidial hila, which has also been confirmed by molecular data. Records of
F. carpophilum from various other hosts are very doubtful, e.g., on species of Potentilla, Filipendula
and Rosa from Kazakhstan (SHVARTSMAN et al. 1975), Salix sp. from Romania (BONTEA 1985),
Bromus inermis and Hordeum vulgare from Estonia (JÄRVA et al. 1998) and species of Acacia,
Cheirodendron and Metrosideros from the USA (FARR et al. 1989). Records from South America are
also uncertain because Fusicladium carpophilum and Coryneum carpophilum (Clasterosporium
carpophilum) have often been confused in this area (VIÉGAS 1961).
10.2.9.
Fusicladium catenosporum (Butin) Ritschel & U. Braun comb. nov.
Fig. 9
Pollaccia catenospora Butin, Mycol. Res. 96(8): 658 (1992); holotype: on Salix triandra, Germany, Berlin, Eberswalde-Finow, 8 Jul. 1990, Butin (IMI 349857).
= Fusicladium salicis Moesz & Smarods, in herb.
Teleomorph: Unknown.
Ill.: BUTIN (1992: 659, Figs 1–6).
Exs.: Krieger, F. sax. 2090.
≡
Leaf spots amphigenous, scattered, at first subcircular, later irregular, 2–10 mm wide,
reddish brown, margin dark brown to blackish, often causing distortions at the leaf
margin, incurved or entire leaves bent, on young twigs forming punctiform, pale brown
swellings. Colonies amphigenous, dense, oblong or circular, medium olivaceousbrown, sometimes confluent. Mycelium immersed, subcuticular, forming colourless,
circular hyphal plates. Stroma 100–300 µm diam., composed of pale brown, thickwalled cells, 4–8 µm diam., 15–25 µm deep. Conidiophores in dense fascicles, arising from the upper cells of the stromata, forming sporodochial conidiomata, erumpent
through the cuticle, erect, ovoid to doliiform, unbranched, 10–16 × 6–8 µm, aseptate,
medium olivaceous-brown, smooth, walls somewhat thickened, conidiophores reduced
to conidiogenous cells, unilocal, determinate or rarely percurrent, with up to two
inconspicuous annellations, loci truncate or slightly convex, (2–)3–4(–5) wide, not
to slightly thickened, not darkened. Conidia catenate, in unbranched or rarely branched
chains, ellipsoid, limoniform or fusiform, straight to sometimes slightly curved, 10–
21(–27) × 5–9 µm, mostly aseptate, rarely 1–2-septate, pale olivaceous-brown, smooth,
walls not or only slightly thickened, truncate at the apex and base, hila (2–)3–4(–5)
µm wide, unthickened to occasionally very slightly thickened, not darkened.
Fig. 9: Fusicladium catenosporum. A – conidia, B – conidiogenous cells with catenate conidia
arranged in a sporodochium, scale = 10 µm, A. Ritschel del.
Hosts and Distribution: on Salix spp. (Salicaceae), Europe – Salix amygdalina (D),
S. purpurea (LV), S. triandra (D).
Material examined: on Salix purpurea, Latvia, Kandara, May 1936, Smarods (M), as Fusicladium
salicis Moesz & Smarods.
Notes: BUTIN (1992) described conidiogenous cells with only a few, inconspicuous annellations. In
Fig. 1 (drawing) and Fig. 3 (micrograph), conidiogenous cells possibly with two conidiogenous loci
are shown, indicating an affinity to species of Fusicladium s.str. with catenate conidia. During the
course of monographic studies, carried out by RITSCHEL (2001), conidiogenous cells with up to two
annellations were found, but two loci were not observed.
ONDÌEJ (1973) described Fusicladium sp. from Salix purpurea, characterised by having catenate
conidia, but this fungus was distinguished by having longer, pluriseptate conidia and longer, narrower conidia formed in unbranched chains.
10.2.10.
Fusicladium caulicola U. Braun & K. Schub. sp. nov.
Fig. 10
Holotype: on dry stems of Sedum maximum, Germany, Bavaria, Gerolzhofen, May 1906, herb. P.
Magnus (HBG).
Teleomorph: Unknown.
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SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
33
Ad caules siccos, sine lesionibus. Coloniae punctiformes, effusae, dispersae vel confluentes. Stromata 30–120 µm diam. vel confluentes, modice vel atro-brunnea, ex cellulis subglobosis vel angularis–
irregularis, 3–15 µm latis, crassitunicatis composita. Conidiophora laxe vel dense fasciculata, pauca
vel numerosa, conidiomata sporodochiales vel conidiophora dispersa, caespitosa, recta, cylindrica
vel flexuosa, geniculata–sinuosa, apicem versus saepe attenulata, non-ramosa, 10–50 × 4–8 µm,
septata, pallide vel modice brunnea, apicem versus saepe pallidiora, apice interdum subhyalino,
levia, leniter crassitunicata; cellulae conidiogenae integratae, terminales, saepe monoblasticae,
determinatae vel polyblasticae, sympodiales; loci truncati vel leniter convexi, 2–3 µm lati, nonincrassati, non- vel interdum lenissime fuscati–refractivi. Conidia solitaria, ellipsoidea, ovoidea,
obovoidea, recta, 14–22 × (3–)4–6 µm, 0–1-septata, ad septa non- vel leniter constricta, apice obtuso
vel subacuto, basi truncata vel subconvexa, 2–3 µm lata, hila non-incrassata, non-fuscata.
On dry stems without conspicuous lesions or discolorations. Colonies punctiform,
effuse, scattered to confluent, black. Stromata 30–120 µm diam. or confluent and
larger, sometimes forming expanded layers, medium to dark brown, cells subcircular
to angular–irregular in outline, 3–15 µm diam., thick-walled. Conidiophores in small
to large, loose to dense fascicles, sporodochial or spread, caespitose, forming layers;
conidiophores straight, subcylindrical to flexuous, geniculate–sinuous, usually attenuated, unbranched, 10–50 × 4–8 µm, septate, septa mostly in the lower half, pale
to medium brown throughout or usually paler towards the apex, tips sometimes very
pale, subhyaline, smooth, walls somewhat thickened. Conidiogenous cells integrated,
terminal, mostly unilocal (monoblastic), determinate, sometimes with two or three
loci (polyblastic, sympodial), loci flat, truncate to slightly convex, 2–3 µm wide,
unthickened or almost so, not darkened, occasionally somewhat refractive or slightly
darkened. Conidia solitary, ellipsoid, ovoid, obovoid, straight, 14–22 × (3–)4–6 µm,
0–1-septate, not or only slightly constricted at the septum, pale olivaceous or brownish, verruculose, thin-walled to slightly thickened, apex blunt to somewhat pointed,
base truncate to slightly convex, hila 2–3 µm wide, unthickened, not darkened.
Hosts and Distribution: only known from the type collection.
10.2.11.
Acrosporium cerasi Rabenh., in A. Braun, Verh. Vereins Beförd. Gartenbaues Königl. Preuss.
Staaten 1: 176 (1853); type: on fruits of Prunus cerasus, Borussia [Iconotype: Braun (l.c.: Pl. 1,
B, 1–2)].
≡ Fusicladium cerasi (Rabenh.) Sacc., Syll. Fung. 4: 346 (1886), comb. superfl.
≡ Cladosporium cerasi (Rabenh.) Aderh., Centralbl. Bakteriol., 2. Abth., 7: 656 (1901).
≡ Fusicladiopsis cerasi (Rabenh.) Karak. & Vassiljevsky, in Vassiljevsky & Karakulin, Parazitnye
nesovershennye griby, Ch. I. Gifomitsety: 210 (1937).
≡ Megacladosporium cerasi (Rabenh.) Vienn.-Bourg., Les champignons parasites des plantes
cultivées 1: 537 (1949).
≡ Karakulinia cerasi (Rabenh.) N.P. Golovina, Novosti Sist. Nizsh. Rast. 1: 213 (1964).
Teleomorph: Venturia cerasi Aderh., Landw. Jahrb. 29: 541 (1900).
Lit.: LINDAU (1907: 783–784), ONDÌEJ (1971: 168–169), CMI Descr. (No. 706), SIVANESAN (1977:
50–53; 1984a: 610–611).
Ill.: ADERHOLD (1900: Pl. 9, Fig. 22), VASSILJEVSKY & KARAKULIN (1937: 210, Fig. 18), HUGHES
(1953: 568, Fig. 7), SCHWEIZER (1958: Figs 5–13, 16, 20), ONDÌEJ (1971: 167, Fig. 2), CMI Descr.
(No. 706, Figs), SIVANESAN (1977: 51, Fig. 22; 1984a: 610, Fig. 367).
Exs.: Petr., Fl. Bohem. Morav. exs. 578; Vestergr., Micromyc. rar. sel. exs. 138.
≡
Fig. 10: Fusicladium caulicola. A – conidia, B – conidiogenous cells with a single or two loci, C –
dense fascicles of conidiophores, scale = 10 µm, K. Schubert del.
Fusicladium cerasi (Rabenh.) Erikss., Meddeland. Kongl. Lantbruksakad.
Exp.-fält 1: 73 (1885)
Figs 11, 12
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SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
35
Fig. 11: Fusicladium cerasi. Iconotype (from BRAUN 1853). 1 – conidiophores with conidia, 2 –
conidia.
Colonies on fruits as greyish-brownish or black tufts, caespitose, sooty, on leaves
mostly epiphyllous, causing small, greyish black spots. Mycelium intra- or intercellular, hyphae 2–3 µm wide, hyaline or yellowish. Stromata subcuticular, composed of
unthickened or only slightly thickened polygonal cells, 3–6 µm diam. Conidiophores
solitary or in loose fascicles, arising from stromata, erect, straight or somewhat
flexuous, unbranched or rarely branched at the base, (10–)20–40(–60) × (3–)4–6(–7)
µm, 0–1(–2)-septate, septa in the lower half, pale to medium brown or olivaceous,
paler towards the apex, smooth, walls slightly thickened, mostly somewhat swollen
at the base. Conidiogenous cells integrated, terminal, with several conidiogenous loci,
crowded at the apex, proliferation sympodial, loci denticulate, 1–2(–2.5) µm wide,
unthickened, not or only slightly darkened–refractive. Conidia solitary, rarely in short,
unbranched or occasionally branched chains, fusiform, subcylindrical, ellipsoid or
obclavate, 11–25(–28) × 4–7 µm, 0–1(–3)-septate, constricted at the septa, yellowish
to medium brown, smooth or somewhat roughened to wrinkled, pointed or rounded
at the apex, truncate at the base, hilum truncate to slightly convex, 1–2(–2.5) µm
wide, unthickened, not or only slightly darkened–refractive.
Hosts and Distribution: on species of Prunus s.lat. (Rosaceae), Asia, Caucasus,
Europe, North America, South America, Australia, New Zealand – Prunus armeniaca
(Europe, D, RO), P. avium (Asia, Kirghizia, Kazakhstan, Uzbekistan; Caucasus,
Azerbaijan, Georgia; Europe A, CS, D, DK, I, LV, Moldavia, NL, RO, RUS, SLO,
Ukr.; America; Australia, Victoria; New Zealand), P. capollin (South America, Columbia), P. cerasifera (Europe, RO), P. cerasus (Asia, Iran, Kazakhstan, Kirghizia,
Uzbekistan; Caucasus, Armenia, Azerbaijan, Georgia; Europe, A, CZ, D, DK, EW,
GB, LV, Moldavia, NL, RO, RUS, S, Ukr.; North America, USA, NC, NE, WA; South
America, Brazil; Australia, Victoria; New Zealand), P. domestica (Asia, Iran), P. persica
(Europe, D, RO), Prunus spp. (Europe, BG, HR, NL, SK, Cyprus; Asia, Japan).
Fig. 12: Fusicladium cerasi. A – conidia, B – conidiophores, scale = 10 µm, K. Schubert del.
Material examined: collections from B, HBG, JE, LE.
Notes: MENON (1956), ONDÌEJ (1971) and SIVANESAN (1977) described very short conidiophores,
up to 20 µm in length. Fusicladium cerasi and F. carpophilum are two allied species on various hosts
of the genus Prunus s.lat., but they are morphologically (conidia usually formed singly in F. cerasi,
always catenate in F. carpophilum), physiologically (differences in the temperature tolerance), bio-
logically (BENSANDE & KEITT 1928, differences in the host range) and genetically (SCHUBERT 2001;
chapter 3.5) distinguished. Furthermore, they are connected with two different teleomorphs of the
genus Venturia. The two species have often been confused. FERRARIS (1912) and LIND (1913) even
reduced F. carpophilum to synonymy with F. cerasi.
36
10.2.12.
Schlechtendalia 9 (2003)
Fusicladium consors Sacc., Ann. Mycol. 4: 491 (1906)
Fig. 13
Holotype: on stems of Succisa pratensis (= Scabiosa succisa), France, Bais de Meudon, May 1900,
M. Ludwig, comm. P. Hariot (PC), [together with Didymosphaeria perexigua].
Teleomorph: Unknown.
Lit.: SACCARDO (1910: 732; 1913: 1375).
Lesions lacking. Mycelium internal. Hyphae sparingly branched, 2–4 µm wide, septate, pale or pigmented, smooth, often with swollen cells and constrictions at the septa.
Conidiophores solitary or in small, loose fascicles, arising from internal hyphae or swollen
hyphal cells, erumpent, erect, straight, subcylindrical, barely or only slightly geniculate–sinuous, 10–40 × 3–5 µm, 0(–1)-septate, brownish, wall thin to slightly thickened,
smooth or almost so; conidiophores usually reduced to conidiogenous cells,
conidiogenous loci subdenticulate, 1.5–2 µm wide, unthickened, not darkened. Conidia
formed singly or in short, simple chains, ellipsoid–obovoid, pyriform, 12.5–18 × 3.5–5
µm, (0–)1-septate, pale olivacous-brown, smooth or almost so, wall thin or slightly
thickened, apex obtuse, truncate, rarely subacute, base obconically truncate, 1.5–2 µm
wide, hila unthickened, but mostly somewhat darkened–refractive.
Hosts and Distribution: only known from the type collection.
Notes: This species is tentatively maintained in Fusicladium since it is morphologically indistinguishable from other species of this genus. The biology of F. consors is unclear. Lesions are not
formed, so that a saprobic habit may be supposed.
A
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
10.2.13.
37
Fusicladium convolvularum OndÍej, „eská Mykol. 25(3): 171 (1971) Fig. 14
Holotype: on Convolvulus arvensis, Czech Republic, Libina, okraj pole pod nadrazim (okr. Sumperk),
7 Sept. 1970, OndÍej (BRA).
Teleomorph: Unknown.
Lit.: IMI Descr. (No. 1513).
Ill.: ONDÌEJ (1971: 170, Fig. 5), IMI Descr. (No. 1513, Figs A–D).
Leaf spots amphigenous, 1–5 mm wide, subcircular, brown, later with greyish brown
to greyish white centre and brown margin. Mycelium usually subcuticular. Hyphae
brown, forming flat stromatic layers of angular–irregular, subhyaline to brown cells,
often radiating. Conidiophores solitary or in small loose groups, arising from stromatic cells or hyphae, mostly erumpent through the cuticle, rarely emerging through
stomata, erect, straight or curved at the apex, subcylindrical to geniculate–sinuous,
unbranched, 10–60 × 4–7 µm, 0–1(–2)-septate, pale to medium brown, smooth,
wall somewhat thickened, but often irregularly thickened, often with a single or
occasionally several percurrent proliferations which are not connected with
conidiogenesis. Conidiogenous cells integrated, terminal, 10–25 µm long, with a
single or several, often denticle-like conidiogenous loci, proliferation sympodial,
loci 1.5–3 µm wide, unthickened, not or often somewhat darkened–refractive. Conidia solitary or occasionally in unbranched or branched chains, ellipsoid–ovoid,
fusiform, subcylindrical, 10–27 × 3–6 µm, 0–3-septate, mostly constricted at the
septa, subhyaline to pale olivaceous, smooth to rough-walled, usually somewhat
attenuated towards apex and base, apex rounded, pointed or truncate, base truncate,
hila 1.5–3 µm wide, flat, unthickened or almost so, not or somewhat darkened–
refractive.
Hosts and Distribution: on Calystegia and Convolvulus spp. (Convolvulaceae),
Europe, New Zealand – Calystegia sepium (Europe, CZ), C. soldanella (Europe, GB),
Convolvulus arvensis (Europe, CZ; New Zealand).
Material examined: on Calystegia soldanella (= Convolvulus soldanella), England, Kent, Camber,
8 Oct. 1963, Sutton and Pirozynski (IMI 102675a); on Convolvulus arvensis, Czech Republ., Mlade
(Litorel), 5 Sept.1970, M. OndÍej (BRA); on Convolvulus arvensis, New Zealand, 7 Nov. 2000, C.F.
Hill, dried culture (ex herb. IMI); on Calystegia sepium, Czech Republ., Libina, 23 Sept. 1970, M.
OndÍej (BRA).
Notes: In the collections examined some conidium-like structures with a length of up to 60 µm have
been observed, but it is not quite clear if they belong to the present species. In molecular studies, it
has been demonstrated that F. convolvularum is a genuine member of Fusicladium, since this species
clustered near to F. effusum within a large monophyletic Venturia clade.
B
C
Fig. 13: Fusicladium consors. A – conidia, B – conidiophores, C – mycelium, scale = 10 µm, U.
Braun del.
10.2.14.
Fusicladium crataegi Aderh., Ber. Deutsch. Bot. Ges. 20: 200 (1902) Fig. 15
Lectotype: on Crataegus laevigata (= C. oxyacantha auct.), Germany, Erfurt, 15 Mar. 1902, Aderhold,
Syd., Mycoth. germ. 45 (HBG), selected here; isolectotypes: Syd., Mycoth. germ. 45 (e.g., JE, LE).
≡ Megacladosporium crataegi (Aderh.) Vienn.-Bourg., Les champignons parasites des plantes
cultivées 1: 539 (1949).
Teleomorph: Venturia crataegi Aderh., Ber. Deutsch. Bot. Ges. 20: 200 (1902).
Lit.: LINDAU (1907: 778–779), VASSILJEVSKY & KARAKULIN (1937: 199–200), HUGHES (1953:
567–568), SIVANESAN (1977: 59–60; 1984a: 612).
38
Schlechtendalia 9 (2003)
Fig. 14: Fusicladium convolvularum. A – conidia, B – conidiogenous cells, C – conidiophore arising from hyphae or swollen hyphal cells, D – conidiophore emerging through a stoma, scale = 10
µm, K. Schubert del.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
39
Fig. 15: Fusicladium crataegi. A – conidia, B – conidiophores solitary or in small groups, scale = 10
µm, K. Schubert del.
40
Schlechtendalia 9 (2003)
Ill.: ADERHOLD (1902: Tab. IX), LINDAU (1907: 779, Fig. 680), HUGHES (1953: 568, Fig. 8),
SIVANESAN (1977: 59, Fig. 28; 1984a: 613, Fig. 369).
Exs.: Syd., Mycoth. germ. 45.
On living leaves and fruits, dark, crustaceous, small, subcircular spots, 1–2 mm wide
or confluent and larger, sometimes covering the entire surface of the leaves or fruits.
Colonies brown to blackish. Mycelium immersed, subcuticular, hyphae branched,
2.5–3 µm wide, septate, pale olivaceous. Stromata pseudoparenchymatous, forming
few layers, composed of pale olivaceous to brown, thick-walled, polygonal cells, 5–
13 µm diam. Conidiophores solitary or in loose to dense fascicles, arising from stromata, erumpent through the cuticle, erect, straight, at the distal end mostly somewhat
curved, unbranched, 20–54(–80) × (3.5–)4–5(–6) µm, 0–2-septate, mainly in the lower
half, pale olivaceous to chestnut-brown, paler towards the apex, smooth, walls somewhat thickened. Conidiogenous cells integrated, terminal, proliferation sympodial,
with a single or several conidiogenous loci, mostly at the apex, denticulate, (1–)1.5–
2(–2.5) µm wide, unthickened, not darkened. Conidia solitary, fusiform, obclavate,
10–25 × 4–6(–8.5) µm, pale olivaceous, 0–1-septate, septum more or less median,
rarely 2-septate, more or less constricted at the septa, smooth to verruculose, apex
usually acute or rounded, truncate at the base, walls only slightly thickened, hila 1.5–
2 µm wide, unthickened, not darkened.
Hosts and Distribution: on Crataegus spp. (Rosaceae), Asia, Caucasus, Europe, North
America, Australia – Crataegus crenulata (≡ Pyracantha crenulata) (South Australia),
C. laevigata (= C. oxyacantha auct.) (Europe, D, DK, F, RO), C. monogyna (Europe,
RO), C. pentagyna (Asia, Iran), C. subvillosa (Europe, RO), Crataegus spp. (Caucasus,
Armenia, Georgia; Europe, H, LT, RO, RUS, SK; North America, USA, FL).
Material examined: collections from B, HBG, JE, LE, M.
Notes: VIENNOT-BOURGIN (1949) described conidia formed singly as well as in short chains, with
one or two septa.
10.2.15.
Fusicladium diedickeanum U. Braun, Nova Hedwigia 55(1–2): 211 (1992)
Fig. 16
Holotype: on Syringa vulgaris, Germany, Thuringia, Erfurt, Oct. 1897, Diedicke (JE).
Teleomorph: Venturia syringae (Syd.) M.E. Barr, Canad. J. Bot. 46: 815 (1968).
Lit.: SIVANESAN (1977: 106).
Ill.: SIVANESAN (1977: 107, Fig. 57; 1984a: 611, Fig. 368 B); BRAUN (1992: 212, Fig. 2).
Leaf spots amphigenous, subcircular, angular to irregular, often vein-limited,
brownish, margin indefinite or centre later often greyish white, with narrow, brown
margin, occasionally somewhat raised. Mycelium internal. Stromata almost absent or small, substomatal or intraepidermal composed of swollen hyphae, 3–8
µm diam., pale brown. Conidiophores solitary or in small groups, usually 2–8,
erumpent through stomata or the cuticle, erect, geniculate–sinuous, unbranched,
25–100 × 4–7 µm, septate, brown, paler towards the apex, smooth. Conidiogenous
cells integrated, terminal or intercalary, 10–30 µm long, proliferation sympodial,
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
41
Fig. 16: Fusicladium diedickeanum. A – conidia, B –
conidiophores, scale = 20 µm, U. Braun del.
with one to several conidiogenous loci,
subdenticulate, loci unthickened or almost so,
not or only slightly darkened–refractive. Conidia solitary, ellipsoid, ovoid, subcylindrical,
8–18 × 4–7 µm, 0–3-septate, olivaceous to
olivaceous-brown, verruculose, apex more or
less rounded, often somewhat attenuated towards the base, base more or less truncate, hila
unthickened or almost so, not or only very
slightly darkened–refractive.
Hosts and Distribution: on Syringa spp.
(Oleaceae), Europe, North America – Syringa
vulgaris (Europe, D; North America, USA,
MA).
10.2.16.
Fusicladium effusum G. Winter, J. Mycol. 1: 101–102 (1885)
Fig. 17
Holotype: on Carya tomentosa (= Carya alba), North America, USA, Illinois, Cobden Zels., 1 Oct.
1882, F.S. Earle (B).
≡ Cladosporium effusum (G. Winter) Demaree, J. Agric. Res. 37: 186 (1928), homonym, non Berk.
& M.A. Curtis, Grevillea 3(27): 106 (1875).
≡ Fusicladosporium effusum (G. Winter) Partrigde & Morgan-Jones, Mycotaxon 85: 364 (2003).
= Fusicladium caryigenum Ellis & Langl., J. Mycol. 4: 124 (1888); lectotype: on leaves of Carya
illinoensis, USA, Louisiana, St. Martin, 3 Sept. 1888, A.B. Langlois, Fl. Ludov. 1499 (NY),
selected here; isolectotypes: on leaves of Carya illinoensis (= Carya olivaeformis), USA, Louisiana, St. Martinsville, Sept. 1888, A.B. Langlois (BPI 426315, 426333; M).
≡ Cladosporium caryigenum (Ellis & Langl.) Gottwald, Mycologia 74(3): 388 (1982).
Teleomorph: Unknown.
Lit.: SACCARDO (1886: 346), GOTTWALD (1982), IMI Descr. (No. 1514), PARTRIDGE & MORGANJONES (2003: 364).
Ill.: GOTTWALD (1982: Figs 1–3), IMI Descr. (No. 1514, Figs A–D), PARTRIDGE & MORGAN-JONES
(2003: 363, Fig. 2).
Exs.: Ellis & Everh., N. Am. F. 545; Kellerm. & Sw., Kans. F. 39.
Leaf spots amphigenous, subcircular to angular, 1–3 mm wide, often confluent, diffuse, numerous, mostly spread along leaf veins, dark brown to black, with an irregular margin. Colonies amphigenous, caespitose, dark brown to blackish. Mycelium
mainly subcuticular. Stromata variable in size, composed of pale olivaceous to brown,
angular to rounded, thick-walled, pseudoparenchymatous cells, 4–8 µm diam., forming up to three layers. Conidiophores solitary or loosely fasciculate, arising from
stromata or from hyphae, erect, straight, sometimes flexuous at the apex, unbranched
or apically branched, 22–130(–170) × 4–6 µm, septate, pale to dark brown, smooth,
with somewhat thickened walls. Conidiogenous cells integrated, terminal or intercalary, or conidiophores reduced to conidiogenous cells, 10–40 µm long, with a single
42
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SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
43
or several denticle-like conidiogenous loci, proliferation sympodial, loci unthickened,
not or only somewhat darkened–refractive, 1.5–3 µm wide. Conidia in simple or
branched chains, pyriform, subcylindrical, ellipsoid, fusiform, (8.5–)10–24 × 5–10
µm, pale brown, 0(–1)-septate, smooth, attenuated towards apex and base, apex mostly
truncate, occasionally rounded or pointed, base truncate, hila unthickened, but often
somewhat darkened–refractive, 1.5–3 µm wide.
Hosts and Distribution: on Carya spp. and ?Juglans spp. (Juglandaceae), Africa,
North, Central and South America, New Zealand – Carya aquatica (North America,
USA, FL), C. cordiformis (= C. amara) (North America, USA, KS, WI), C. glabra
(North America, USA, FL), C. illinoensis (= C. pecan, = C. olivaeformis) (Africa,
South Africa; North America, USA, AL, LA, MO, TX; South America, Brazil; New
Zealand), C. ovata (North America, USA, WI), C. tomentosa (= C. alba) (North
America, USA, IL, LA, KS), Carya spp. (North America, USA, AL, FL, NC, OK;
Central America, Mexico; South America, Paraguay), ?Juglans regia (South America,
Brazil).
Material examined: collections from B, M, NY.
Notes: On account of the catenate conidia, GOTTWALD (1982), assigned Fusicladium effusum to
Cladosporium, but the conidiogenous loci of this species agree well with those of Fusicladium in
that they are denticle-like and have unthickened walls. The conidiogenous loci in species of Cladosporium, described and illustrated in detail by DAVID (1997), are quite distinct. Therefore, F. effusum
belongs in Fusicladium, as has recently been confirmed by molecular studies of rDNA ITS sequences
(SCHNABEL et al., 1999; SCHUBERT 2001) in which this species clustered close to various Venturia
species with Fusicladium and Pollaccia anamorphs within a monophyletic Venturia clade (see chapter 3) In addition to its occurrence on an unrelated host, Fusicladium effusum var. carpineum Ellis &
Everh. on Carpinus species (Corylaceae) in North America is distinguished from F. effusum on Carya
species (Juglandaceae) in causing distinct lesions and having much longer and wider conidiophores
with paler conidiogenous cells and less conspicuous conidiogenous loci. This variety is now considered to be a separate species of Fusicladium. Records on Juglans regia from Brazil (MENDEZ et al.
1998) are uncertain (no material seen).
10.2.17.
Pollaccia elegans Servazzi, Boll. Lab. Sperim. Osserv. Fitopatol. 15(3–4): 64 (1939); neotype:
on Populus nigra, Germany, Geesthacht, 10 Jul. 1904, Jaap (B), as Napicladium asteroma (Fuckel)
Allesch., selected here; isoneotype: H.
= Fusicladium radiosum [(Lib.) Lind] var. balsamiferae Davis, Trans. Wisconsin Acad. Sci. 20:
402 (1922); lectotype: on Populus balsamifera, USA, Wisconsin, Sturgeon Bay, 23 Jun. 1913,
J.J. Davis (WIS); topotype: on Populus balsamifera, USA, Wisconsin, Sturgeon Bay, 19 Jul.
1918, J.J. Davis (WIS).
≡ Pollaccia balsamiferae (Davis) M. Morelet, Bull. Soc. Sci. Nat. Archéol. Toulon Var 4: 3 (1972).
= Fusicladium tremulae A.B. Frank, Hedwigia 22: 127 (1883) p. p.
= Napicladium tremulae auct. p.p.
= Stigmina radiosa auct. p.p.
= Fusicladium radiosum auct. p.p.
Teleomorph: Venturia populina (Vuill.) Fabric., Jahresber. Neuerung Pflanzenkrankh. 5: 282 (1902).
Lit.: DANCE (1961: 875–890), ONDÌEJ (1972: 145), CMI Descr. (No. 483), ELLIS (1976: 110),
SIVANESAN (1977: 89–93; 1984a: 619–620), BRANDENBURGER (1985: 39), ELLIS & ELLIS (1997:
192), KHAN & MISRA (1989).
≡
Fig. 17: Fusicladium effusum. A – conidia, B – microcyclic conidiogenesis, C – conidiophore arising from a hypha, D – conidiophores in a loose fascicle, scale = 10 µm, K. Schubert del.
Fusicladium elegans (Servazzi) Ritschel & U. Braun comb. nov. Fig. 18
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45
loci 4–7 µm wide, not to very slightly thickened, somewhat refractive. Conidia solitary, ellipsoid to broadly fusiform, straight, rarely slightly curved, (21–)25–40(–45)
× 9–13(–16) µm, 1–2(–3)-septate, with a large central cell and two smaller cells at the
ends with somewhat thinner walls, more or less constricted at the septa, dark
olivaceous-brown, smooth, thick-walled, rounded at the apex, base truncate to slightly
convex, sometimes oblique, hila 4–7 µm wide, not to very slightly thickened, somewhat refractive.
Hosts and Distribution: on Populus spp. (Salicaceae), Asia, Europe, North America
– Populus balsamifera [Asia, China; Europe, GB; North America, Canada, Alta., BC.,
Man., NB., Nfld., NS., NWT (Mack.), Ont., PEI, Que., Sask., Yukon, USA, WI], P. ×
canadensis (Europe, I; North America, Canada, Ont.), P. ciliata (Asia, India), P.
deltoides (North America, Canada, Que.) P. nigra (incl. P. pyramidalis) (Asia, India;
Europe, CS, D, F, I, SLO; North America, Canada, NB., USA, northern central states,
north-eastern states, OR, WA), P. trichocarpa (North America, Canada, BC., USA,
AK, OR), Populus spp. (Europe, CH, GB; North America, Canada, Alta., BC., Man.,
Nfld., Ont., Sask., USA, northern central states, north-eastern states, WA).
Fig. 18: Fusicladium elegans. A – conidia, B – conidiogenous cells arranged in a sporodochium,
scale = 10 µm, A. Ritschel del.
Ill.: SERVAZZI (1939: Pl. III, Figs 10–15; Pl. IV, Figs 16–22; Pl. V, Figs 23–25; Pl. VI, Figs 26–31; Pl.
VII, Figs 32–36), DANCE (1961: Pl. I, Figs 1–9; Pl. II, 10–18, Figs 1–9), BARR (1968: 805, Fig. 20),
ONDÌEJ (1972: 145, Figs 6, 7), CMI Descr. (1976: No. 483, Fig. B), ELLIS (1976: 110, Fig. 77 B),
SIVANESAN (1977: 91, Fig. 49; 1984a: 620, Fig. 374 B), LIU, CHEN & SHAO (1981: Pl. II, Fig. 6; 23,
Figs 1–2), WU & SUTTON (1995: 985, Fig. 6), ELLIS & ELLIS (1997: Pl. 82, Fig. 850).
Leaf spots amphigenous, subcircular to irregular, 5–20 mm wide, pale brown, surrounded by a narrow, reddish brown, somewhat raised margin, limited by the leaf
margin or by veins, later confluent and larger, sometimes covering large leaf segments, occasionally causing distortions of the leaves. Colonies amphigenous, punctiform to confluent, fructification often spread along veins, on the upper leaf surface
dark brown to almost black, below pale brown to dark brown, also on twigs. Mycelium
immersed. Stroma intraepidermal to subcuticular, composed of subcircular to slightly
angular, thick-walled cells, 5–10 µm diam., forming up to three layers. Conidiophores
usually in dense fascicles, arising from the upper cells of stromata, forming
sporodochial conidiomata, sometimes solitary, erumpent through the cuticle,
subglobose to cylindrical, 10–16 × 4–10 µm, 0–2-septate, olivaceous-brown, smooth,
relatively thick-walled, conidiophores usually reduced to conidiogenous cells, unilocal,
determinate or occasionally percurrent, with a single or two inconspicuous annellations,
Notes: PRILLIEUX (1892) examined Fusicladium on Populus nigra from France, compared it with F.
tremulae (= Pollaccia radiosa), and found differences in the conidial shape and size but maintained
this fungus under the latter species (as Napicladium tremulae). LIND (1905) followed the treatment
of PRILLIEUX (1892), but called this species Fusicladium radiosum. SERVAZZI (1939) introduced
Pollaccia elegans and discussed the differences to P. radiosa in detail. Unfortunately, type material
of P. elegans could not be traced and is probably not preserved, so a neotype is proposed in this
paper. D ANCE (1961) showed that Venturia populina only attacked Populus species of sect.
Tacamahaca and sect. Aigeiros and this was confirmed during the course of the monographic examinations by RITSCHEL (2001). Records from Populus species of sect. Leuce (GINNS 1986, ONDÌEJ
1972) have probably been based on misidentifications of the fungi or hosts.
10.2.18.
Fusicladium euphorbiae Karak., Bolezni Rast.13: 132 (1924)
Fig. 19
Lectotype: on stems of Euphorbia virgata, Russia, prov. Leningrad, Ropsha, 5 Sept. 1924, B.N.
Karakulin (LE 40571), selected here; isolectotypes: HAL 1629, IMI 92541, LE 40957, LE 40958.
≡ Fusicladiopsis euphorbiae (Karak.) Karak., in Vassiljevsky & Karakulin, Parazitnye
nesovershennye griby, Ch. I. Gifomitsety: 209 (1937).
≡ Karakulinia euphorbiae (Karak.) N.P. Golovina, Novosti Sist. Nizsh. Rast. 1: 213 (1964).
Teleomorph: Unknown.
Lit.: DEIGHTON (1967: 23–25), ONDÌEJ (1971: 169–170).
Ill.: KARAKULIN (l.c.: Fig. 11), VASSILJEVSKY & KARAKULIN (1937: 209, Figs 16–17), DEIGHTON
(1967: 24, Fig. 11), ONDÌEJ (1971: 168, Fig. 3).
On living leaves and stems without conspicuous lesions. Colonies caespitose, varying in shape and size, dark brown to black, scattered. Mycelium both inter- and intracellular, subcuticular. Stromata oblong, 50–150 × 10–50 µm diam., composed of swollen, pale olivaceous to brown, somewhat thick-walled cells, 4–8 µm diam.,
pseudoparenchymatous, forming few layers. Conidiophores in small to usually large,
loose to dense fascicles, arising from stromata, erect, more or less straight, unbranched,
10–90 × (3–)4–6 µm, septate, olivaceous to pale brown, paler towards the apex, smooth,
walls relatively thin, occasionally swollen at the base. Conidiogenous cells integrated,
terminal or intercalary, with one to several conidiogenous loci, denticulate, prolifera-
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47
tion sympodial, loci flat, truncate or slightly convex, 1.5–3(–4) µm wide, not or only
very slightly thickened, not or somewhat darkened–refractive. Conidia solitary or
catenate, in short, simple or sometimes branched chains, fusiform, subcylindrical,
obclavate or clavate, straight, 10–22(–40) × 3–6 µm, 0–2(–3)-septate, often slightly
constricted at the septa, pale olivaceous, smooth to minutely verruculose, walls only
slightly thickened, apex more or less obtuse or truncate, sometimes slightly papillate,
base truncate, hila 1.5–3(–4) µm wide, unthickened or only very slightly thickened,
not or only slightly darkened–refractive.
Hosts and Distribution: on Euphorbia spp. (Euphorbiaceae), Asia, Caucasus,
Europe – Euphorbia amygdaloides (Europe, RO), E. cyparissias (Europe, RO),
E. esula (Europe, CZ), E. exigua (Europe, RO), E. lamprocarpa (Asia,
Kazakhstan; Europe, CZ, RUS), E. villosa (Europe, RO), E. virgata (Europe,
RUS), Euphorbia spp. (Asia, Central Asia, Turkmenistan; Caucasus, Armenia;
Europe, CZ, RUS).
Material examined: on Euphorbia sp., Czech Republic, Mlade… (Litorel), 5 Sept. 1970, M. OndÍej
(BRA); on E. sp., Czech Republic, H. Libina, near Òumperk, 18 Sept. 1970, M. OndÍej (BRA); on E.
villosa, RO, Distr. Suceava, Gr|niceÕti, 26 May 1972, A. Manulin (M).
10.2.19.1. Fusicladium fasciculatum Cooke & Ellis, Grevillea 6: 88 (1878) var.
fasciculatum
Fig. 20
Holotype: on stems of Euphorbia nutans, USA, New Jersey, Newfield, J.B. Ellis no. 2774 (90098 b)
(K); isotype: NY (mixed infection with Cladosporium chaetomium).
≡ Scolecotrichum fasciculatum (Cooke & Ellis) Shear, Bull. Torrey Bot. Club 29: 499 (1902).
≡ Passalora fasciculata (Cooke & Ellis) Earle, Torreya 2: 60 (1902).
≡ Cercosporidium fasciculatum (Cooke & Ellis) Höhn., Centralbl. Bakteriol., 2. Abth., 60: 4 (1923).
Teleomorph: Unknown.
Lit.: DEIGHTON (1967: 16–21).
Ill.: DEIGHTON (1967: 20, Fig. 9).
Exs.: Ellis, N. Am. F. 545.
Fig. 19: Fusicladium euphorbiae. A – conidia, B – dense fascicle of conidiophores (LE 40957,
type), C – dense fascicle of conidiophores (collection from M), scale = 10 µm, K. Schubert del.
On stems, without conspicuous lesions. Colonies caespitose, effuse, up to 2 cm
long, black, velvety, rarely on leaves, amphigenous. Mycelium internal, intercellular, subcuticular, hyphae branched, 1.5–4 µm wide, septate, almost colourless
to very pale olivaceous. Conidiophores in small, loose fascicles, up to 6, arising
from hyphae, consisting of two or three swollen cells, erumpent through the cuticle, erect or slightly divergent, more or less straight, usually somewhat flexuous,
unbranched, very rarely branched, 37–175 × (3–)4–5 µm, septate, septa very thin,
not always conspicuous, dark olivaceous, paler towards the apex, smooth, thickwalled, base often slightly swollen, sometimes with percurrent proliferations which
are not connected with conidiogenesis. Conidiogenous cells integrated, terminal
or intercalary, numerous and often crowded, proliferation sympodial, loci conspicuous and prominent, sometimes situated at the end of short lateral projections, denticulate, 1.5–2 µm wide, wall not or only very slightly thickened, somewhat darkened–refractive. Conidia solitary, fusiform, ellipsoid or subcylindrical,
straight, 8.5–16 × (3–)4–6.5 µm, 0(–1)-septate, pale olivaceous, smooth to
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verruculose, wall relatively thin-walled, apex rounded or pointed, attenuated at
the base, truncate, hila 1.5–2 µm wide, not or only very slightly thickened, but
somewhat darkened–refractive.
Hosts and Distribution: on Euphorbia spp. (Euphorbiaceae), North America, South
America – Euphorbia corollata (North America, USA, WI), E. glyptosperma (North
America, USA, WI), E. nutans (North America, USA, NJ), E. serpyllifolia (North
America, USA, WI), Euphoria spp. (North America, USA, KS, MD, NJ; South
America, Brazil).
Material examined: collections from B, M.
Notes: FARR et al. (1989) listed species of Ammophila and Alopecurus as hosts of this species, which
is very doubtful.
10.2.19.2. Fusicladium fasciculatum [Cooke & Ellis] var. didymum Deighton,
Mycol. Pap. 112: 23 (1967)
Fig. 21
Holotype: on Euphorbia corollata, USA, Iowa, Decorah, 5 Aug. 1884, E.W.D. Holway, as Fusicladium
fasciculatum (NY).
Teleomorph: Unknown.
Ill.: DEIGHTON (1967: 22, Fig. 10).
Leaf spots indefinite or sometimes small, subcircular, up to 3 mm wide, somewhat orange to yellowish brown, or sometimes consisting of dark punctate areas
of irregular shape, which may cover a large area of the leaf. Colonies amphigenous,
dark, velvety. Mycelium immersed, intercellular; hyphae sparingly branched, 1.5–
3 µm wide, septate, almost colourless. Conidiophores solitary or in small groups
of 2–3, arising from hyphae, erumpent through the cuticle, erect, more or less
flexuous or curved, unbranched or rarely branched near the apex, 65–260 × 2.5–
5 µm, septate, dark olivaceous, paler towards the apex, smooth, walls somewhat
thickened. Conidiogenous cells integrated, terminal or intercalary, proliferation
sympodial, loci numerous, prominent, often on short nodulose projections, denticulate, 1.5–2.5 µm wide, conspicuous, but not or only very slightly thickened,
somewhat darkened–refractive. Conidia solitary, broadly fusiform to short clavate, (9–)13–19 × (4.5–)5–7 µm, (0–)1(–2)-septate, usually slightly or distinctly
constricted at the septum, sometimes not constricted, pale olivaceous, verruculose,
apex rounded or shortly papillate, base attenuate, truncate, hila 1.5–2.5 µm wide,
unthickened or only very slightly thickened, not or only slightly darkened–refractive.
Hosts and Distribution: on Euphorbia spp. (Euphorbiaceae), North America –
Euphorbia corollata (USA, IA).
Notes: Var. didymum differs from var. fasciculatum in having usually 1-septate conidia (more
than 90 %).
Fig. 20: Fusicladium fasciculatum var. fasciculatum. A – conidia, B – microcyclic conidiogenesis,
C – unbranched and branched conidiophores, D – conidiophores in a small fascicle, scale = 10 µm,
K. Schubert del.
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10.2.20.
Fusicladium fautreyi Deighton, Mycol. Pap. 112: 25 (1967)
51
Fig. 22
Holotype: on Euphorbia brittingeri (= Euphorbia verrucosa), France, May 1895, F. Fautrey, Roum.,
F. sel. exs. 6829 (G).
Teleomorph: Unknown.
Ill.: DEIGHTON (1967: 26, Fig. 12).
Exs.: Roum., F. sel. exs. 6829.
On dry stems, numerous discrete, oblong, greyish brown dots, densely scattered over
the whole surface. Colonies forming small blackish tufts on the greyish brown spots.
Mycelium subcuticular. Stromata 60–110 µm diam., composed of dark olivaceousbrown, thick-walled cells. Conidiophores densely fasciculate, arising from stromata,
erumpent through the cuticle, erect, more or less straight, unbranched, 30–90 × 4–6
µm, septate, dark olivaceous-brown, somewhat paler towards the apex, smooth.
Conidiogenous cells integrated, terminal, with a single to several conidiogenous loci,
proliferation sympodial, loci conspicuous, slightly prominent, not or only very slightly
thickened, not darkened. Conidia solitary, obclavate to fusiform, straight, 11–20.5 ×
4–5.5 µm, 0–1-septate, lower cell mostly larger than the upper one, very rarely with
three septa, not or only very slightly constricted at the septa, pale olivaceous, slightly
verruculose, apex pointed, truncate at the base, hilum truncate to slightly convex,
1.5–2.5 µm wide, unthickened, not darkened.
Hosts and Distribution: only known from the type collection.
Notes: This species resembles Fusicladium euphorbiae, but differs in having more robust, thickwalled, darker conidiophores, thick-walled, dark brown stromatic cells and somewhat shorter and
wider conidia, consistently formed singly. Additional collections and molecular data are necessary
to prove the true status of F. fautreyi and its affinity to F. euphorbiae.
Fig. 21: Fusicladium fasciculatum var. didymum. A – conidia, B – conidiophores, C – mycelium and
base of conidiophores, scale = 10 µm, K. Schubert del.
Fig. 22: Fusicladium fautreyi. A – conidia, B – conidiophores, scale = 10 µm, K. Schubert del.
52
10.2.21.
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53
Fusicladium fraxini Aderh., Hedwigia 36: 74, 83* (1897) [*erroneously
as Fusicladium tremulae]
Fig. 23
Neotype: on leaves of Fraxinus ornus, Italy, prov. Verona, Tregnago, May 1913, C. Massalongo, Kab.
& Bub., F. imp. exs. 794 (B), selected here; isoneotypes: Kab. & Bub., F. imp. exs. 794 (e.g., BPI, W).
≡ Spilocaea fraxini (Aderh.) Sivan., Biblioth. Mycol. 59: 65 (1977).
= Scolecotrichum fraxini Pass., Erb. Critt. Ital., Ser. II. 1395 (1884); syntype: on leaves of Fraxinus
ornus, Italy, Parma, Vigheffio, Estate, G. Passerini, Erb. Critt. Ital., Ser II. 1395 (e.g., M).
= Actinonema fraxini Allesch., Bot. Centralbl. 2: 44 (1890); syntype: on leaves of Fraxinus excelsior,
Germany, Munich, Isardamm, Sept. 1897, Allescher (M).
= Fusicladium granulosum Pass., in herb. (B).
Teleomorph: Venturia fraxini Aderh., Hedwigia 36: 83 (1897).
Lit.: LINDAU (1907: 787, 798), FERRARIS (1912: 326), VASSILJEVSKY & KARAKULIN (1937: 193),
SIVANESAN (1984a: 614), SAGDULLAEVA (1990: 54).
Ill.: ADERHOLD (1897: Tab. 4, Fig. 6), SIVANESAN (1977: 67, Fig. 33; 1984a: 615, Fig. 371).
Exs.: Briosi & Cav., F. paras. 297; Erb. Critt. Ital. 1395; Herb. Mycol. Rom. 293; Kab. & Bub., F.
imp. exs. 93, 794; Petr., Fl. Bohem. Morav. exs. 782b; Rabenh., F. eur. 943; Syd., Mycoth. march.
2928; Vestergr., Micromyc. rar. sel. exs. 950.
Leaf spots amphigenous, circular, oval to angular–irregular, up to 10 mm wide,
ochraceous, yellowish to olivaceous-brown on the upper leaf surface, paler below, surrounded by a medium to dark brown, narrow margin, often marginal and
fragile. Colonies amphigenous, punctiform, mainly spread along leaf veins, dark
olivaceous to blackish. Stroma subcuticular to intraepidermal, 10–100 µm diam.,
composed of relatively large, olivaceous to medium brown swollen cells, 2–7 µm
diam., forming expanded layers. Conidiophores aggregated in loose to dense fascicles, sometimes sporodochial or solitary, arising from the stromata, mostly erect,
straight to geniculate, flexuous, subcylindrical to conical, unbranched, (5–)12–
35(–100) × 3–5 µm, 0(–1)-septate, pale olivaceous to medium brown, smooth,
often swollen at the base, up to 7 µm wide. Conidiogenous cells integrated, terminal or conidiophores usually reduced to conidiogenous cells, proliferation
sympodial with one to several loci, or percurrent with several distinct, transverse
annellations or both types of proliferation mixed, loci subdenticulate, 1–2 µm
wide, unthickened, not or at most slightly darkened–refractive. Conidia solitary,
fusiform to obclavate, straight to slightly curved, 12–28 × 4–6(–7) µm, (0–)1(–
3)-septate, not or only slightly constricted at the septa, septa more or less median
or somewhat in the lower half, subhyaline, pale olivaceous to olivaceous-brown,
walls somewhat thickened, attenuated towards apex and base, apex often oblong–
pointed, truncate at the base, hila 1–2 µm wide, unthickened, slightly darkened–
refractive.
Hosts and Distribution: on Fraxinus spp. (Oleaceae), Asia, Caucasus, Europe,
North America – Fraxinus angustifolia (Europe, RO), F. excelsior (Asia, Central
Asia, Uzbekistan; Caucasus, Armenia, Georgia; Europe, CZ, D, EW, GB, LV,
RO, RUS, Ukr.), F. ornus (Europe, I, RO), F. raibocarpa (Asia, Russia,
Tadzhikistan), F. sogdiana (Central Asia, Uzbekistan), Fraxinus spp. (Central Asia,
Uzbekistan; Caucasus, Armenia; Europe, EW, H, LV, RO, Ukr.; North America,
USA, FL).
Material examined: collections from B, HBG, IMI, LE, M.
Fig. 23: Fusicladium fraxini. A – conidia, B – sympodially proliferating conidiogenous cells, C –
conidiophores, arranged in a sporodochium, mostly with a single locus, D – percurrently proliferating conidiogenous cells, scale = 10 µm, K. Schubert del.
Notes: ADERHOLD (1897) described a new species of Fusicladium on Fraxinus and called it repeatedly (pp. 74–76) Fusicladium fraxini, but the original diagnosis was erroneously connected with the
name F. tremulae, which is a synonym of Pollaccia radiosa, a Venturia-anamorph occurring on
Populus spp. This is undoubtedly a correctable mistake. A neotype is proposed since Aderhold´s
type material is not preserved. Detailed studies on the structure of the conidiogenous cells and loci
showed that percurrent and sympodial proliferations can occur within a single collection, so that this
species has to be considered intermediate between Fusicladium s.str. and Spilocaea. Fusicladium
fraxini var. phillyreae Trotter, in Pamp. (Nuovo Giorn. Bot. Ital. 31: 233, 1924), described on Phillyrea
media from Libya, is probably not conspecific with Spilocaea phillyreae, but type material could not
be traced.
54
10.2.22.
Schlechtendalia 9 (2003)
Fusicladium grayianum (Ellis) Deighton & M.B. Ellis, in Deighton,
Mycol. Res. 94(8): 1097 (1990)
Fig. 24
Isariopsis grayiana Ellis, Bull. Torrey Bot. Club 9: 98 (1882); holotype: on dead stems of Rubus
villosus, USA, Pa., West Chester, Oct. 1881, J.B. Gray (NY); isotypes: Ell., N. Am. F. 818 (e.g.,
IMI 92619).
≡ Phaeoisariopsis grayiana (Ellis) Ferraris, Ann. Mycol. 7: 280 (1909), as ‘grayana’.
Teleomorph: Unknown.
Ill.: DEIGHTON (1990: 1097, Fig. 1).
Exs.: Ellis, N. Am. F. 818.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
10.2.23.
≡
Colonies on stems, effuse, tufted, dark blackish brown. Mycelium immersed, hyphae
branched, 1.5–3 µm wide, septate, pale brown, wall smooth. Stromata mostly immersed, pseudoparenchymatous, 16–60 µm wide, 20–40 µm high, brown or dark
brown. Conidiophores usually aggregated in dense fascicles, arising from the upper
cells of the stromata, erect, flexuous, unbranched, 80–280 × 2.5–4 µm, pluriseptate,
brown, dark near the base, paler towards the apex, mostly smooth, the upper part
occasionally wrinkled or verruculose. Conidiogenous cells integrated, terminal, with
a single or several scattered, subdenticulate conidiogenous loci, proliferation
sympodial, loci unthickened, not or only very slightly darkened. Conidia solitary,
cylindrical to short obclavate, straight or slightly curved, 6–14 × 2.5–4 µm, 0–1septate, pale olivaceous, smooth to verruculose, rounded at the apex, base 0.5–1 µm
wide, hila unthickened, not or only very slightly darkened–refractive.
Hosts and Distribution: on Rubus spp. (Rosaceae), North America – Rubus villosus
(USA, PA), Rubus spp. (USA, PA).
Material examined: on Rubus sp., USA, Pa., Catoctin, Jun. 1940, J.A. Stevenson (M).
55
Fusicladium heveae K. Schub. & U. Braun, in Crous & Braun, Mycosphaerella
and its anamorphs: 1. Names published in Cercospora and Passalora. CBS
Biodiversity Series 1: 481 (2003)
Fig. 25
Fusicladium macrosporum Kuyper, Recueil Trav. Bot. Néerl. 8: 374 (1911), non Fusicladium
macrosporium Bonord., 1864; neotype: on Hevea brasiliensis, British Guiana, Araka R., Issorora,
Feb. 1926, Alison (IMI 18583), selected here.
= ?Passalora heveae Massee (nom. nud.) sensu Stahel, Bull. Dept. Landb. Suriname 34: 34 (1917).
Teleomorph: Microcyclus ulei (Henn.) Arx, in Müller & Arx, Beitr. Kryptogamenfl. Schweiz 11:
373 (1962).
Lit.: CMI Descr. (No. 225), ELLIS (1976: 239), SIVANESAN (1984a: 180–182).
Ill.: KUYPER (1912, Fig. 93), STAHEL (1917: Pl. 12, 1–2, Pl. 14, 1–10, Pl. 18, 1, Pl. 25, 1–3), CMI
Descr. (No. 225, Fig. A), ELLIS (1976: 240, Fig. 180), SIVANESAN (1984a: 183, Fig. 93).
≡
On leaves, stems, petioles, inflorescences, flowers and young fruits, leaf spots amphigenous,
variable in shape and size, up to 15 mm wide, greyish brown, scattered on the leaf surface,
occasionally confluent, sometimes somewhat raised. Colonies effuse, greyish, powdery.
Mycelium immersed, subepidermal. Stromata composed of loosely to densely aggregated,
yellowish brown, thick-walled cells, 6–11 µm diam. Conidiophores loosely to densely
fasciculate, arising from stromata, erect, straight or somewhat flexuous, sometimes geniculate, unbranched, 15–140 µm long, but usually less than 50 µm, 4–7 µm wide, at first nonseptate with a subglobose base, later septate, pale olivaceous, mostly smooth, rarely somewhat rough-walled. Conidiogenous cells integrated, terminal, with 1–4 loci, proliferation
sympodial, loci only slightly denticulate, 2–4 µm wide, often somewhat convex, unthickened,
not darkened. Conidia solitary, obclavate, sometimes straight, usually curved to sigmoid,
15–65 × 6–11 µm, usually (0–)1-septate, somewhat constricted at the septa, hyaline to pale
olivaceous, smooth to minutely verruculose, walls thin-walled or only slightly thickened,
broadest part in the lower third, attenuated towards the apex, truncate at the base, hilum 2–
4 µm wide, unthickened, not or only very slightly darkened–refractive.
Hosts and Distribution: on Hevea spp. (Euphorbiaceae), South America – Hevea
benthamiana (Brazil), H. brasiliensis (Brazil, British Guiana, Columbia, Peru, Trinidad), H. guianensis (CMI Descr. 225: South America), H. spruceana (CMI Descr.
225: South America), Hevea spp. (South America, Trinidad).
Material examined: on Hevea sp., Trinidad, 1 Feb. 1980, Kheng Hoy Chee (IMI 247065); on Hevea
brasiliensis, Trinidad, 6 Feb. 1969, T. Mungal (IMI 137417); on Hevea brasiliensis, Trinidad, 22
Jun. 1962, P. Holliday (IMI 134624); on Hevea brasiliensis, Trinidad, Balandra, Mar. 1961, C.L.A.
Leakey (IMI 87945).
Fig. 24: Fusicladium grayianum. A – conidia, B – dense fascicle of conidiophores, scale = 10 µm,
K. Schubert del.
Notes: Type material of this species could not be traced and is probably not preserved, so that a
neotypification is proposed. The epitheta “macrosporum“ and “macrosporium“ are confusable and
must be treated as homonyms (ICBN, Art. 64.2). Therefore, a new name was introduced for
Fusicladium macrosporum Kuyper. STAHEL (1917) and MÜLLER & ARX (1962) listed Scolecotrichum
heveae Vincens (Cah. Pathol. Vég. Entomol. Agric. 2: 17, 1915) as synonym, but, according to the
original description, the latter name does not agree with Fusicladium heveae (conidiophores much
longer, 150–200 µm, and conidia formed in chains).
Fusicladium heveae is an unusual species, since its teleomorph, Microcyclus ulei, is placed into the
Mycosphaerellaceae (KIRK et al. 2001) and not into the Venturiaceae. Additional examinations, including molecular studies, are necessary to find the true affinity of this fungus. The anamorph of
Microcyclus ulei is tentatively maintained in Fusicladium since it is morphologically indistinguishable from other species of this genus.
56
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10.2.24.
57
Fusicladium humile (Davis) K. Schub. & U. Braun, IMI Descriptions of
Fungi and Bacteria 152, No. 1520 (2002)
Fig. 26
Cladosporium humile Davis, Trans. Wisconsin Acad. Sci. 19: 702 (1919); lectotype: on Acer rubrum,
USA, Wisconsin, Luck, 25 Aug. 1916, J.J. Davis (WIS), selected here; isolectotype: BPI 427214.
≡ Fusicladosporium humile (Davis) Partridge & Morgan-Jones, Mycotaxon 85: 366 (2003).
Teleomorph: Venturia acerina Plakidas ex M.E. Barr, Canad. J. Bot. 46: 814 (1968).
Lit.: PLAKIDAS (1942: 35), ELLIS (1976: 340), SIVANESAN (1977: 26–27), PARTRIDGE & MORGANJONES (2003: 366).
Ill.: PLAKIDAS (1942: 30, Fig. 2), ELLIS (1976: 340, Fig. 258), SIVANESAN (1977: 26, Fig. 2; 1984a:
607, Fig. 364), IMI Descr. (No. 1520, Figs A–C), PARTRIDGE & MORGAN-JONES (2003: 365, Fig. 3).
≡
Leaf spots (only associated with the anamorph) amphigenous, variable in shape and
size, subcircular to angular, 0.5–20 mm wide, dark reddish brown on upper leaf surface, greyish below, sometimes zonate, margin irregular, pale grey. Mycelium internal, hyphae subhyaline to pale brown, rough-walled, septate, forming mainly subcuticular net-like aggregations, effuse. Colonies amphigenous, punctiform to subeffuse,
dark. Stromata 35–80 µm diam., composed of thick-walled, brown cells, 4–10 µm
wide. Conidiophores solitary, arising from internal or external, superficial hyphae or
swollen hyphal cells, or formed in loose fascicles arising from stromata, erect to
flexuous, straight to geniculate, unbranched or occasionally branched, 15–70 × 3.5–
6 µm, septate, olivaceous to pale brown, smooth, wall somewhat thickened.
Conidiogenous cells integrated, terminal or intercalary, with a single or several denticle-like conidiogenous loci, proliferation sympodial, loci unthickened, not or only
slightly darkened–refractive. Conidia catenate, in simple or branched chains, cylindrical or fusiform, straight or slightly curved, 10–29 × 4–7 µm, 0–2(–3)-septate, not
or somewhat constricted at the septum, pale olivaceous to medium brown, smooth,
attenuated towards apex and base, apex pointed or truncate, base truncate, hila 1–3(–
3.5) µm wide, unthickened, not darkened.
Hosts and Distribution: on Acer spp. (Aceraceae), North America – Acer negundo
(USA), A. nigrum (USA), A. rubrum (Canada, NB., Nfld., Ont.; USA, AL, MI, NY,
WI), A. saccharinum (USA, NC, WI), A. saccharum (Canada, Ont.), A. spicatum
(BARR 1968: USA).
Material examined: collections from WIS.
Notes: The teleomorph was described and illustrated in detail by BARR (1968) and SIVANESAN (1977,
1984a). Due to the structure of the conidiogenous loci and conidial hila, Cladosporium humile has to
be excluded from Cladosporium and assigned to Fusicladium, which is consistent with its connection with a Venturia teleomorph. The report of Cladosporium humile from India (Kashmir) on Populus
(BEIG & KHAN 1999) probably refers to either Fusicladium martianoffianum or F. romellianum.
10.2.25.
Fig. 27
Fusicladium junci Sawada, Rep. Gov. Res. Inst. Formosa 86: 162 (1943), nom. inval. (without
Latin diagnosis).
Holotype: on Juncus prismatocarpus, Taiwan, K. Sawada (NTU-PPE) [= National Taiwan University, Dept. of Plant Pathology and Entomology].
Teleomorph: Unknown.
Ill.: SAWADA (l.c.: Figs 34–35).
≡
Fig. 25: Fusicladium heveae. A – conidia, B – conidiogenous cells, C – conidiophores in loose
fascicles, scale = 10 µm, K. Schubert del.
Fusicladium junci Sawada ex K. Schub. & U. Braun sp. nov.
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59
Fig. 27: Fusicladium junci. A – conidia, B – stromata substomatal arranged, C – dense fascicles of
conidiophores, scale = 10 µm, K. Schubert del.
Maculae indefinitae, diffusae, brunneae. Coloniae amphigenae, punctiformes, brunneae. Mycelium immersum.
Stromata nulla vel bene evoluta, substomatalia, oblonga, 20–40 × 10–20 µm, brunnea, ex cellulis, 3–6 µm
latis, brunneis, leniter crassitunicatis composita. Conidiophora fasciculata, pauca, per stoma emergentia,
cylindrica–conica, recta, erecta, non-ramosa, 5–15(–38) × 4–7(–9) µm, pallide brunnea, levia. Cellulae
conidiogenae monoblasticae, determinatae vel polyblasticae, sympodiales; loci truncati, 1.5–2.5 µm lati,
non-incrasati, non-fuscati, interdum refractivi. Conidia solitaria, obclavata vel cylindrica, recta, (26–)30–
50(–57) × 4–6 µm, 1-septata, leniter constricta, subhyalina vel pallide flavissima-viridula vel olivacea, levia,
apice obtuso, rotundato, basi obconice truncata, 1.5–2.5 µm lata, hila non-incrassata, non-fuscata.
On necrotic brown leaves, sheaths and stems, leaf spots indefinite, diffuse, brown. Colonies amphigenous, punctiform, brown. Mycelium internal. Stromata absent to welldeveloped, substomatal, oblong, 20–40 × 10–20 µm, brown, stromatic cells 3–6 µm
diam., brown, with slightly thickened walls. Conidiophores in small, more or less dense
fascicles, emerging through stomata, cylindrical–conic, straight, erect, unbranched, 5–
15(–38) × 4–7(–9) µm, aseptate, pale brown, smooth, conidiophores usually reduced to
conidiogenous cells. Conidiogenous cells unilocal (monoblastic), determinate or with
two conidiogenous loci, sympodial, loci flat, 1.5–2.5 µm wide, unthickened, not darkened, but sometimes somewhat refractive. Conidia solitary, obclavate to cylindrical,
straight, (26–)30–50(–57) × 4–6 µm, 1-septate, slightly constricted at the septa,
subhyaline to very pale yellowish-greenish or olivaceous, smooth, apex obtuse, rounded,
base obconically truncate, hila flat, 1.5–2.5 µm wide, unthickened, not darkened.
Fig. 26: Fusicladium humile. A – conidia, B – conidiophores arising from hyphae, C – loose fascicle
of conidiophores arising from stromata, scale = 10 µm, K. Schubert del.
Hosts and Distribution: on Juncus spp. (Juncaceae), Taiwan – Juncus prismatocarpus
(Taiwan).
60
10.2.26.
Schlechtendalia 9 (2003)
Fusicladium lathyrinum (Ellis & Galloway) S. Hughes & Piroz., Canad.
J. Bot. 50(12): 2528 (1972)
Fig. 28
Dicoccum lathyrinum Ellis & Galloway, J. Mycol. 5: 65 (1889), as ‘lathyrmum’; holotype: on
Lathyrus ochroleucus, USA, Montana, Highwood Mts., Highwood Canyon, 18 Jun. 1888, R.S.
Williams, Parasitic Fungi of Montana 301 (NY); isotype: DAOM 130903 (permanent slide).
Teleomorph: Unknown.
Ill.: HUGHES & PIROZYNSKI (1972: 2528, Fig. 3).
≡
Leaf spots amphigenous, on the upper leaf surface forming yellowish
discolorations, on the lower side almost white, sunken. Colonies hypophyllous,
dense, caespitose, dark yellowish brown, velvety. Mycelium immersed, subcuticular to intraepidermal, composed of irregular, colourless, thin-walled hyphae,
branched in the mesophyll, aggregated in the epidermis, and giving rise to widely
extended, compact layers of conidiogenous cells. Conidiophores reduced to
conidiogenous cells, erumpent through the outer wall of the epidermis and the
cuticle, cylindrical, narrowly clavate or ovoid, erect, unbranched, 4–18 × 4–7
µm, aseptate, subhyaline to very pale brown, smooth, thin-walled, except for the
apex, which has a thicker brown wall, mostly with two or three loci, proliferation
sympodial, loci denticulate, flat, about 3 µm wide, not or only slightly thickened
and darkened. Conidia solitary, ellipsoid, subcylindrical to obclavate, straight,
15–28 × 6–10 µm, 0–1-septate, constricted at the septa, septa thin, inconspicuous, at first hyaline, later pale olivaceous, coarsely verrucose, rounded at the
base, with a flat, broad, not or only slightly thickened, not or only very slightly
darkened hilum, 3 µm wide.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
10.2.27.
61
Fusicladium levieri Magnus, in Sommier & Lévier, Trudy Imp. S.Peterburgsk. Bot. Sada 16: 543 (1900)
Fig. 29
Holotype: on Diospyros lotus, Caucasus, Georgia, Batum, in Silvis litoris Euscini, 16 Jun. 1890
(HBG).
≡ Ragnhildiana levieri (Magnus) Vassiljevsky, in Vassiljevsky & Karakulin, Parazitnye
nesovershennye griby, Ch. I. Gifomitsety: 373 (1937).
≡ Cladosporium levieri (Magnus) Hara, Agric. & Hort. 12: 2706 (1937).
≡ Phaeoramularia levieri (Magnus) U. Braun, in Braun & Melnik, Trudy Bot. Inst. Komarova (St.
Petersburg) 20: 69 (1997).
= Fusicladium kaki Hori & Yoshino, Bot. Mag. (Tokyo) 19: 220 (1905).
= Fusicladium diospyri Chona, Munjal & J.N. Kapoor, Indian Phytopathol. 9: 129 (1956); type:
on Diospyros kaki, India, U. P., Saharanpur (HCIO?).
= Fusicladium diospyri Hori & Yoshino in herb. (B).
Teleomorph: Unknown.
Lit.: SACCARDO (1902: 1056), SUBRAMANIAN (1971: 235), SCHOLLER et al. (2003).
Ill.: VASSILJEVSKY & KARAKULIN (1937: 373, Fig. 32), BRAUN & MELNIK (1997: Fig. 38), SCHOLLER
et al. (2003: Figs 1–2).
Exs.: Herb. Mycol. Rom. 1195; Kab. & Bub., F. imp. exs. 845.
Leaf spots amphigenous, subcircular to angular–irregular, 1–5 mm wide, centre greenish brown or ochraceous to greyish white, surrounded by a small to fairly broad,
Hosts and Distribution: only known from the type collection.
Fig. 28: Fusicladium lathyrinum. A – conidia, B – conidiogenous cells, scale = 10 µm, K.
Schubert del.
Fig. 29: Fusicladium levieri. A – conidia, B – conidiophores emerging through a stoma, C –
conidiophores arising from stromatic hyphal aggregations, scale = 10 µm, K. Schubert del.
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63
dark, often almost blackish margin, sometimes with a wide, somewhat discoloured
halo, scattered on the leaf surface. Colonies amphigenous, mainly hypophyllous, inconspicuous to punctiform, dark, fructification mostly sparse. Mycelium internal,
immersed, hyphae branched, septate, pigmented. Stromata absent or as small, brown
hyphal aggregations, substomatal or intraepidermal. Conidiophores solitary or usually in small, loose to dense fascicles, arising from internal hyphae or stromatic hyphal
aggregations, erumpent through the cuticle or emerging through stomata on the lower
leaf surface, erect to flexuous, straight to geniculate–sinuous, unbranched or rarely
branched, 10–70 × 3–8 µm, 0–3-septate, pale olivaceous to pale brown, smooth, walls
somewhat thickened. Conidiogenous cells integrated, terminal, with a single to only
few conidiogenous loci, proliferation sympodial, loci 1.5–3 µm wide, inconspicuous,
unthickened or almost so, occasionally somewhat darkened. Conidia catenate, in simple, occasionally in branched chains, subcylindrical, ellipsoid to fusiform, straight,
13–40 × 3–7 µm, 0–2-septate, sometimes slightly constricted at the septa, pale
olivaceous, smooth, ends obtuse or obconically truncate, hila 1.5–3 µm wide,
unthickened or almost so, occasionally somewhat darkened.
Hosts and Distribution: on Diospyros spp. (Ebenaceae), Asia, Caucasus, Europe, North
America – Diospyros kaki (Asia, China, India, Japan; Caucasus, Georgia; Europe, RO),
D. lotus (Caucasus, Georgia), D. virginiana (North America, USA, CT, FL, IN, MS).
Material examined: on Diospyros kaki, Japan, Miyazaki, Houzyô-mati, 30 May 1935, M. Ebihara
(B); on Diospyros lotus, Anbrevsky, 2 Aug. 1915, V. Semaschk (LE 161233); on Diospyros virginiana,
USA, Indiana, Vigo County, Terre Haute, Persimmon Street, J. Lehman, 26 Jun. 2002 (PUR 1680).
Notes: HÖHNEL (1919: 156) introduced the new genus Hormocladium based on Fusicladium kaki.
Braun, in BRAUN & MELNIK (1997), transferred Fusicladium levieri to Phaeoramularia, but detailed examinations of additional collections clearly showed that the fungus from Diospyros spp.
must be maintained in Fusicladium. The conidiogenous loci and conidial hila are occasionally somewhat darkened–refractive, but consistently truncate and unthickened as in other species of Fusicladium.
10.2.28.
Fusicladium mandshuricum (M. Morelet) Ritschel & U. Braun comb.
nov.
Fig. 30
Pollaccia mandshurica M. Morelet, Ann. Soc. Sci. Nat. Archéol. Toulon Var 45(3): 218 (1993);
holotype: on leaves of Populus simonii × P. nigra, north-eastern China, Liaoning, 17 Jun. 1992,
M. Morelet [PC (PFN 1466)].
= Pollaccia sinensis W.P. Wu & B. Sutton, in herb. (IMI).
Teleomorph: Venturia mandshurica M. Morelet, Ann. Soc. Sci. Nat. Archéol. Toulon Var 45(3): 219
(1993).
Lit.: MORELET (1993: 218–219), WU & SUTTON (1995: 983–986), MORELET & SIGAUD (1996: 11–20).
Ill.: WU & SUTTON (1995: 984, Figs 1–4), MORELET & SIGAUD (1996: 16, Fig. 4; 17, Fig. 6).
≡
On living leaves, petioles and twigs, spots circular to irregular, 5–10 mm wide, at
first punctiform, later confluent and larger, often vein-limited, at first brown, later
silvery white to grey, margin conspicuous, yellowish brown, on the lower leaf surface 3–8 mm wide, grey, margin yellowish brown, infected necrotic tips of shoots
often curved, hook-like. Colonies amphigenous, punctiform, scattered to aggregated
in groups, dark olivaceous-green to blackish. Mycelium immersed, hyphae branched,
Fig. 30: Fusicladium mandshuricum. A – conidia. B – conidiophores with a somewhat irregular rim
at the conidiogenous tip, arranged in a sporodochium, scale = 10 µm, A. Ritschel del.
2–3.5 µm wide, septate, brownish. Stroma at first intraepidermal, later subcuticular,
40–80 µm diam., composed of subglobose to slightly angular, brown, relatively thickwalled cells, 4–8 µm diam., aggregated, forming up to three layers. Conidiophores
densely fasciculate, arising from the upper cells of the stromata, forming sporodochial
conidiomata, erumpent through the cuticle, straight, cylindrical to ampulliform,
unbranched, 5–7 × 6–7.5 µm, aseptate, pale to medium olivaceous-brown, smooth,
walls somewhat thickened, occasionally swollen at the base, up to 10 µm wide,
conidiophores reduced to conidiogenous cells, unilocal, determinate or percurrent,
with a single annellation (in culture up to three annellations), annellations with a
somewhat irregular, more or less dark rim, loci truncate to slightly convex, 4(–5) µm
wide, not to slightly thickened, not darkened. Conidia solitary, fusiform, more or less
curved, (27–)30–39(–42) × 6–9(–10) µm, (0–)2–3(–4)-septate, central cell often bulging, more or less constricted at the septa, pale to dark olivaceous-brown, smooth,
thick-walled, attenuated towards apex and base, apex pointed, base (hila) truncate to
slightly convex, 4(–5) µm wide, not to very slightly thickened, not darkened, occasionally with a small, lateral, subhyaline foot-like projection.
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65
Hosts and Distribution: on Populus spp. (Salicaceae), Asia – P. simonii × P. nigra
and Populus spp. (north-eastern China).
Material examined: on Populus sp., Asia, China, Waling Shan, 2 Aug. 1986, Wu (IMI 362777), as
Pollaccia sinensis.
Notes: This species, described as “grey spot disease“ of Populus spp. in northeast China, was erroneously considered in the older Chinese literature to be Coryneum populinum Bres. and the anamorph
of Mycosphaerella mandshurica Miura. However, MORELET (1993) and MORELET & SIGAUD (1996)
examined this fungus in vivo and in vitro and demonstrated that it was an undescribed anamorph of
a new species of Venturia. W U & SUTTON (1995) depicted the conidiogenous loci with irregular,
darkened rims. In the material examined during the course of the present monographic studies, the
rims of the loci were less conspicuous. It is to be supposed that this species is more widespread in
China and possibly also in the Far East of Russia.
10.2.29.
Fusicladium martianoffianum (Thüm.) K. Schub. & U. Braun, IMI
Descriptions of Fungi and Bacteria 152, No. 1515 (2002)
Fig. 31
Cladosporium martianoffianum Thüm., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 55(1):
74 (1880); lectotype: on Populus laurifolia, Russia, Siberia, Minussinsk, near river Jenissei,
Aug. 1879, N. Martianoff (M), selected here; isolectotype: on Populus laurifolia, Russia, Siberia, Minussinsk, Aug. 1879, N. Martianoff, Thüm., Mycoth. univ. 2067 (HAL).
= Fusicladium asiaticum OndÍej, „eská Mykol. 27(4): 237 (1973); holotype: on Populus sp.,
Turkmenistan, Tashkentskij U., 1914, Zaprometov (LE 161361).
Teleomorph: Unknown.
Ill.: ONDÌEJ (1973: 238, Fig. 6), IMI Descr. (No. 1515, Figs A–C).
Exs.: Pilzfl. Sib. 474, 653; Thüm., Mycoth. univ. 2067.
≡
Leaf spots amphigenous, subcircular to somewhat angular–irregular, 1–10 mm wide,
dirty greenish to dark brown by abundant fructification, margin indefinite. Colonies
epiphyllous, punctiform, scattered to dense, sometimes confluent, dark brown. Mycelium subcuticular to intraepidermal, forming swollen, yellowish-brownish, thickwalled cells, 3–10 µm diam., which form conidiophores. Conidiophores solitary or in
loose to dense sporodochial aggregations, often confluent, forming dense fascicles or
layers, cylindrical to conical or irregularly shaped, erect, straight, 5–35 × 3–10 µm,
0(–1)-septate, olivaceous or medium brown, smooth, wall thin to somewhat thickened, conidiophores mainly reduced to conidiogenous cells. Conidiogenous cells with
a single or several conidiogenous loci, subdenticulate, proliferation sympodial, loci
1.5–2.5(–4) µm wide, apex truncate to slightly convex, unthickened, not or only very
slightly darkened–refractive. Conidia solitary (primary conidia) or mostly in short,
sometimes branched chains, ellipsoid to ovoid, obovoid, subglobose to irregular, 10–
22 × (4–)5–9 µm, 0–1(–3)-septate, pale olivaceous to olivaceous, smooth, rarely somewhat rough-walled, thin-walled, attenuated towards apex and base, apex rounded,
truncate or pointed, base obconically truncate, with a single hilum, apex with one or
two hila, 1–3 µm wide, unthickened, not or only very slightly darkened–refractive.
Hosts and Distribution: on Populus spp. (Salicaceae), Asia – Populus afghanica
(Tadzhikistan), P. alba (Kazakhstan, India), P. ciliata (India), P. deltoides (Kazakhstan,
India), P. italica (Kazakhstan), P. laurifolia (Russia, Siberia), P. nigra (Kazakhstan),
P. pruinosa (Tadzhikistan), P. suaveolens (Russia, Siberia), P. tremula (Kazakhstan),
Populus spp. (Kazakhstan, Kirghizia, Uzbekistan, Turkmenistan).
Fig. 31: Fusicladium martianoffianum. A – conidia, B – conidiogenous cells, C – dense fascicles of
conidiophores, scale = 10 µm, K. Schubert del.
Material examined: on Populus deltoides, India, Kashmir, 14 Jul. 1979, A.M. Shah (IMI 240137);
on Populus alba, India, Chogul, 16 Jun. 1985, A.K. Kuul (IMI 304892).
Notes: The examination of type material of Cladosporium martianoffianum and Fusicladium asiaticum
showed that the two taxa are conspecific. The conidiogenous loci are quite distinct from those of
Cladosporium species, described and illustrated in detail by DAVID (1997), in that they are denticlelike with an unthickened, non-pigmented wall. S AGDULLAEVA et al. (1990) described ‘C.
martianoffianum’ with very long conidiophores, up to 110 µm, but the material on which this description was based is undoubted not the present species.
10.2.30.
Fusicladium nashicola K. Schub. & U. Braun sp. nov.
Fig. 32
Holotype: on leaves of Pyrus pyrifolia, Japan, Tsukuba, Ibaraki, Orchard of the National Institute of
Agro-Environmental Sciences, 18 Aug. 2000, H. Ishii (HAL 1749).
Teleomorph: Venturia nashicola S. Tanaka & S. Yamam., Ann. Phytopathol. Soc. Japan 29: 136
(1964).
Lit.: ISHII et al. (1992: 293–298), ISHII et al. (1997: 130–133), ISHII & YANASE (2000: 755–759).
Ill.: ISHII & YANASE (2000: 757, Figs 1–2).
Differt a F. pyrorum conidiis brevioribus.
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67
Leaf spots amphigenous, pale brown to dark brown, even blackish by abundant
fructification. Colonies mostly smaller than 1 mm, but confluent and finally sometimes covering the entire surface of the leaves, often occurring along leaf veins. Mycelium subcuticular. Stromata composed of brown, thick-walled cells, 4–8 µm diam.,
forming few layers, hyphae branched, 2–3 µm wide, septate, subhyaline to pale
olivaceous. Conidiophores loosely to densely fasciculate, arising from stromata, erect
to flexuous, geniculate–subnodulose, sinuous, unbranched or rarely branched, 20–70
× 4–6.5 µm, continuous or septate, brown, paler towards the apex, smooth, thickwalled. Conidiogenous cells integrated, terminal or intercalary, with numerous
conidiogenous loci, proliferation sympodial, loci denticulate, 1.5–3 µm wide, truncate to slightly convex, unthickened, somewhat darkened–refractive, sometimes with
a single percurrent proliferation which is not connected with conidiogenesis. Conidia
solitary, subcircular to fusiform, sometimes cylindrical, straight or slightly curved, 9
–20(–28) × 5.5–10 µm, aseptate, pale brown, smooth, apex rounded or obtuse to
pointed, base truncate or slightly convex, hilum 1.5–3 µm wide, unthickened, somewhat darkened–refractive.
Hosts and Distribution: on leaves of Pyrus spp. (Rosaceae), Asia – Pyrus betulaefolia
(Asia, China), P. bretschneideri (China), P. lindleyi (China, Taiwan), P. pyrifolia (China,
Japan, Korea, Taiwan), P. ussuriensis (China).
Material examined: on leaves of Pyrus lindleyi (= P. sinensis), Japan, Tokyo, Shirai, as F. dentriticum (B).
Notes: SIVANESAN (1977, 1984a) reduced Venturia nashicola to a synonym of V. pyrina, but ISHII et
al. (1992, 1997) and ISHII & YANASE (2000) found clear morphological, pathological and physiological differences between the fungi on Asian and European pears. Venturia nashicola differs from
V. pyrina in having significantly shorter conidia and shorter, narrower lower cells of the ascospores.
The two taxa are also biologically differentiated. V. nashicola is confined to Asian pears, and V.
pyrina occurs on European pears. Molecular data obtained by SCHNABEL et al. (1999) showed that
these two taxa are closely allied. However, the data available are not sufficient to determine whether
the fungi are two distinct species or two races of a single species. The morphological differences
between the two taxa on pears were confirmed during the course of the present monographic studies
and, therefore, we prefer to follow the taxonomy of ISHII & YANASE (2000) in keeping two separate
species. The anamorph of Venturia nashicola has not yet been denominated formally. Since it occurs
independently from the teleomorph, we prefer to introduce a separate name for this anamorph.
10.2.31.
Fusicladium nebulosum (Ellis & Everh.) Ritschel & U. Braun comb.
nov.
Fig. 33
Dicoccum nebulosum Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 1893: 463 (1893);
holotype: on Fraxinus americana, USA, Wisconsin, Kinosha, 10 Sept. 1893, Davis (NY); isotype:
BPI 423603.
≡ Spilocaea nebulosa (Ellis & Everh.) S. Hughes & Piroz., Canad. J. Bot. 50(12): 2530 (1972).
Teleomorph: Unknown.
Lit.: SACCARDO (1895: 616).
Ill.: HUGHES & PIROZYNSKI (1972: 2529, Figs 4, 8).
≡
Fig. 32: Fusicladium nashicola. A – conidia, B – conidiogenous cells, C – fasciculate conidiophores,
scale = 10 µm, K. Schubert del.
Leaf spots hypophyllous, formed as irregular discolorations. Colonies punctiform,
irregular, effuse, confluent, 1–5 mm wide, often formed along veins, at first pale
olivaceous-brown, later dark greyish brown to blackish. Mycelium immersed, subcuticular or intraepidermal, hyphae colourless. Conidiophores solitary, erumpent through
68
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SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
69
Fig. 33: Fusicladium nebulosum. A – conidia, B – sympodially proliferating conidiogenous cell
with two loci, C – conidiogenous cell with a single locus erumpent through the epidermis, D –
conidiogenous cells, scale = 10 µm, A. Ritschel del.
the cuticle, subglobose, broadly ampulliform to campanulate, 5–12 × 5–7 µm, 0–1septate, medium olivaceous-brown, smooth, somewhat thick-walled. Conidiophores
reduced to conidiogenous cells or conidiogenous cells integrated, terminal, with a
single, rarely with two conidiogenous loci, proliferation percurrent, producing a succession of up to 10 single conidia, but annellations not very conspicuous because the
conidia may arise and secede at about the same level or even at progressive lower
levels. Conidia solitary, fusiform to navicular, straight to somewhat curved, (10–)12–
15(–16) × (4–)5–6 µm, 0–1(–2)-septate, septum near the base, pale olivaceous-brown,
verruculose to minutely echinulate, pointed at the apex, truncate to somewhat convex
at the base, hila (3–)4 µm wide, unthickened, not or only slightly darkened.
Hosts and Distribution: on Fraxinus spp. (Oleaceae), North America – Fraxinus
americana (USA, WI).
Material examined: on Fraxinus americana, USA, Wisconsin, Domers, 10 Sept. 1893, J.J. Davis (NY).
Notes: HUGHES & PIROZYNSKI (1972) described conidiogenous cells with up to two conidiogenous
loci with percurrent proliferation and this was confirmed during the re-examination of the type material. In this respect, F. nebulosum is intermediate between Fusicladium s.str. (multilocal, sympodial)
and Spilocaea (unilocal, percurrent).
10.2.32.
Fusicladium obducens Pat., Bull. Soc. Mycol. France 9: 161 (1893)
Fig. 34
Type: on leaves of Prunus serotina (= P. salicifolia), South America, Ecuador, Cotocollao, Lagerheim
(not seen).
Teleomorph: Unknown.
Lit.: SACCARDO (1895: 618), IMI Descr. (No. 1516).
Ill.: IMI Descr. (No. 1516, Figs A–C).
Exs.: Syd., F. exot. exs. 1234.
Fig. 34: Fusicladium obducens. A – conidia, B – conidiogenous cells, C – conidiophores arising
from hyphae, D – branched conidiophore, scale = 10 µm, K. Schubert del.
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SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
71
Leaf spots lacking or almost so. Colonies epiphyllous, effuse, dendritic,
olivaceous-brown to blackish. Mycelium immersed, subcuticular, hyphae
branched, 2–3 µm wide, septate, hyaline. Stromata composed of relatively small
brown, thick-walled, rough-walled cells, 5–7 µm diam. Conidiophores in loose
fascicles arising from stromatic cells or from hyphae, erect, mostly somewhat
flexuous or geniculate, unbranched or branched, 40–120 × 6–8 µm, septate,
olivaceous to brown, paler towards the apex, smooth, wall somewhat thickened, often with percurrent proliferations which are not connected with
conidiogenesis. Conidiogenous cells integrated, terminal or intercalary, proliferation sympodial, with a single or several conidiogenous loci, subdenticulate,
2–4.5 µm wide, wall unthickened, slightly convex, non-pigmented, often with
percurrent proliferations which are not connected with conidiogenesis. Conidia
solitary, fusiform, obovoid, ellipsoid, straight, (9.5–)12–26 × 6–10 µm, pale to
medium brown, 0–1(–2)-septate, not or only slightly constricted at the septa,
smooth, apex obtuse, rounded or pointed, obconically truncate at the base, hila
2–4.5 µm wide, unthickened, non-pigmented.
Hosts and Distribution: on Prunus spp. (Rosaceae), South America – Prunus capollin
(Ecuador), P. serotina (South America).
Material examined: some duplicates of ‘Syd., F. exot. exs. 1234’ from herb. B, HBG, M.
Notes: Type material of this species could not be traced at either FH or PC. The present description and
illustration are based on duplicates of ‘Sydow, Fungi exotici exsiccati 1234’ from B, HBG and M.
10.2.33.
Fusicladium oleagineum (Castagne) Ritschel & U. Braun comb. nov.
Fig. 35
Cycloconium oleagineum Castagne, Cat. pl. Marseille: 220 (1845), as ‘oleaginum’; lectotype:
on leaves of Olea europaea, France, Marseille, Castagne (STR), selected here; isolectotypes: on
leaves of Olea europaea, France, Marseille, Castagne (M; IMI 69757, slide).
≡ Spilocaea oleaginea (Castagne) S. Hughes, Canad. J. Bot. 31: 564 (1953).
Teleomorph: Unknown.
Lit.: BRIOSI & CAVARA (F. paras. 223, 1893), SACCARDO (1886: 343; 1892: 596; 1895: 616), LINDAU
(1907: 769–772), GONZÁLES FRAGOSO (1927: 176–179), ELLIS (1971: 143), BRANDENBURGER (1985: 489).
Ill.: BRIOSI & CAVARA (F. paras. 223, Fig.), LINDAU (1907: 771, Figs 1–4), GONZÁLES FRAGOSO
(1927: 177, Fig. 36; 178, Fig. 37), HUGHES (1953: 564, Fig. 4), ELLIS (1971: 142, Fig. B).
Exs.: Briosi & Cav., F. paras. 223; Kab. & Bub., F. imp. exs. 144.
≡
On living leaves, fruit stalks and fruits, leaf spots amphigenous, subcircular, 5–10
mm wide, pale to greyish brown. Colonies mainly epiphyllous, punctiform or radiating, 3–10 mm wide, below punctiform, 0.1–0.5 mm wide, grey to blackish. Mycelium intraepidermal to mostly subcuticular or superficial, hyphae occasionally
branched, 2.5–8 µm wide, septate, hyaline to medium brown, thin-walled, sometimes
slightly thick-walled. Conidiophores solitary, arising from hyphal cells, erumpent
through the cuticle, subglobose, 8–10 µm diam, or ampulliform, 10–25(–30) × 5–7
µm, or up to 15 µm wide at the base, erect, straight, unbranched, mostly aseptate,
medium to dark brown, paler towards the apex, sometimes smooth, usually roughwalled, thick-walled, conidiophores reduced to conidiogenous cells, with a single or
Fig. 35: Fusicladium oleagineum. A – conidia. B – superficial septate hyphae, C – germinating
conidium, D – conidiogenous cells with several conspicuous annellations, E – percurrently proliferating conidiogenous cell with two loci, F – conidiogenous cell arising from a hypha, scale = 10 µm, A.
Ritschel del.
rarely with two or three conidiogenous loci, proliferation percurrent, with up to seven
conspicuous annellations, loci 5 µm wide, unthickened, not darkened. Conidia solitary, obclavate, straight or occasionally slightly curved, (15–)18–25(–28) × (9–)10–
11(–12) µm, 1-septate, septum median or somewhat in the lower half, sometimes 2septate, often slightly constricted at the septum, medium to dark olivaceous-brown,
verrucose, thick-walled, apex pointed to rounded, truncate at the base, hila 5 µm
wide, unthickened, not darkened.
Hosts and Distribution: on Olea spp. (Oleaceae), Asia, Europe (Mediterranean),
Africa, North America, South America, New Zealand – Olea europaea [Asia, India,
Iran, Israel (Palestine), Jordan, Lebanon, Turkey; Europe, E, F, I, Malta, Cyprus;
Africa, Egypt, Algeria, Libya, Morocco, Somalia, South Africa, Tunisia; North
America, USA, CA; South America, Chile; New Zealand].
Material examined: on Olea europaea, Italy, Siena (1889), Como (1892), Briosi & Cav., F. paras.
223 (HAL, M); Italy, Verona, Tregano, May 1903, Massalongo, Kab. & Bub., F. imp. exs. 144 (M);
USA, California, Berkeley, Feb. 1895, Berletti (M).
Notes: Fusicladium oleagineum differs from almost all other Spilocaea-like Fusicladium species in
having more than one conidiogenous locus. “Striate” conidia described by HUGHES (1953) have not
been observed in the collections examined. The occurrence of this fungus on fruits and stalks was
described by LINDAU (1907).
72
10.2.34.
Schlechtendalia 9 (2003)
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
73
Fusicladium peucedani Ellis & Holw., Bull. Lab. Nat. Hist. Iowa State
Univ. Ia, 3(3): 42 (1895)
Fig. 36
Holotype: on leaves of Peucedanum simplex (= Lomatium simplex), USA, California, Modoc Co.,
13 Jun. 1894, Nutting (NY).
≡ Asperisporium peucedani (Ellis & Holw.) Maubl., Lavoura 16: 207, 211 (1913) and Bull. Soc.
Mycol. France 29: 357 (1913).
≡ Pollaccia peucedani (Ellis & Holw.) Deighton, in Deighton & Piroz., Mycol. Pap. 101: 41 (1965).
Teleomorph: Unknown.
Lit.: SACCARDO (1895: 618; 1913: 1375), ELLIS (1976: 110).
Ill.: DEIGHTON & PIROZYNSKI (1965: 42, Figs 17 A–D), ELLIS (1976: 110, Fig. 77 A).
Exs.: Solh., Mycofl. Saximont. exs. 499.
Leaf spots amphigenous, more or less angular, 2–7 mm wide, grey to pale brownish, vein-limited. Colonies amphigenous, punctiform, dark olivaceous-brown, in
most collections (but not the type collection) associated with black, densely arranged pycnidia. Mycelium immersed, hyphae branched, septate, colourless, densely
intricated. Stromata intraepidermal to subcuticular, up to 100 µm diam., forming
two to four layers. Conidiophores in dense fascicles, arising from the upper cells of
the stromata, forming sporodochial conidiomata, at first subcuticular, later erumpent
through the cuticle, protruding, 5.5–6.5 µm wide, attenuated towards the apex, about
4 µm wide above, erect, straight to slightly flexuous, cylindrical, unbranched, 10–
30 × 5–8 µm, aseptate, at first more or less colourless, with a slightly greenish
tinge, and smooth, later pale to medium olivaceous-brown, loosely verrucose, walls
barely thickened, conidiophores reduced to conidiogenous cells, unilocal, determinate or proliferation percurrent, with up to six conspicuous annellations, loci slightly
convex, 3–5 µm wide, unthickened, not darkened. Conidia solitary, oblong–ellipsoid, straight, 20–29 × (7–)9–11 µm, 0–1-septate, with a median septum, usually
somewhat constricted in the middle, pale olivaceous, loosely verrucose, walls somewhat thickened, apex rounded, hilum slightly convex, 3–5 µm wide, unthickened,
not darkened.
Hosts and Distribution: on Angelica, Cicuta, Glehnia, Lomatium and
Sphaenosciadium spp. (Apiaceae), North America – Angelica arguta (Canada, Alta.,
BC.), A. breweri (USA, CA), Angelica spp. (USA, WA, WY), Cicuta douglasii (USA,
ID), C. occidentalis (USA, ID), Glehnia leiocarpa (USA, WA), G. litoralis (Canada,
BC., USA, CA), Lomatium brandegei (Canada, BC., USA, WA), L. macrocarpum
(Canada, Alta.), L. martindalei (Canada, BC.), L. nudicaule (USA, ID, OR, WA), L.
simplex (= Peucedanum simplex) (USA, CA), L. triternatum (USA, OR),
Sphaenosciadium capitellatum (USA, NV).
Notes: Based on percurrent conidiogenous cells, Deighton, in DEIGHTON & P IROZYNSKY (1965)
placed Fusicladium peucedani Ellis & Holw. in Pollaccia. He supposed that pycnidia, often
found in association with sporodochia of F. peucedani, may belong into the life cycle of this
species. Records on Peucedanum decursivum from Japan [e.g., SHIRAI & H ARA (1927)] have to
be excluded since they belong to Fusicladium peucedani Syd. [≡ Passalora depressa (Berk. &
Broome) Sacc.].
Fig. 36: Fusicladium peucedani. A – conidia, B – conidiogenous cells, one with several conspicuous
annellations, arranged in a sporodochium, C – solitary conidiophore, scale = 10 µm, A. Ritschel del.
10.2.35.
Fusicladium phillyreae (Nicolas & Aggéry) Ritschel & U. Braun comb.
nov.
Fig. 37
Cycloconium phillyreae Nicolas & Aggéry, Bull. Soc. Mycol. France 44: 303 (1928); type: on
Phillyrea angustifolia, France.
≡ Spilocaea phillyreae (Nicolas & Aggéry) M.B. Ellis, More Dematiaceous Hyphomycetes: 111
(1976).
Teleomorph: Unknown.
Ill.: ELLIS (1976: 111, Fig. 78).
≡
On living leaves and petioles, spots amphigenous, subcircular, 1–4 mm wide, on the
upper leaf surface medium brown and somewhat shinning, below pale brown. Colonies punctiform, diffuse, olivaceous to blackish brown. Mycelium subcuticular. Stromata composed of loosely aggregated, subcircular, pale to dark olivaceous-brown
cells, 6–9 µm diam. Conidiophores solitary or in small groups, arising from the upper
cells of the stromata, erumpent through the cuticle, ampulliform or doliiform, erect,
straight to slightly flexuous, unbranched, 10–30 × 5–7 µm, aseptate, medium to dark
olivaceous-brown, smooth, thick-walled, swollen at the base, up to 11 µm wide,
conidiophores usually reduced to conidiogenous cells, unilocal, proliferation
percurrent, with up to six conspicuous annellations, loci truncate, 5–6 µm wide,
unthickened, not darkened. Conidia solitary, navicular to broadly ovate or obclavate,
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SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
75
Fig. 37: Fusicladium phillyreae. A – conidia, B – aggregated conidiophores with several conspicuous annellations, scale = 10 µm, A. Ritschel del.
Fig. 38: Fusicladium pisicola. A – conidia, B – conidiophores, scale = 10 µm, K. Schubert del.
straight to slightly curved, 17–40 × 7–11 µm, (0–)1(–3)-septate, septum median or
somewhat in the lower half, often slightly constricted at the septa, medium to dark
olivaceous-brown, verruculose, thick-walled, apex pointed or somewhat rounded, truncate at the base, hilum 5–6 µm wide, not or only very slightly thickened and darkened.
Hosts and Distribution: on Phillyrea spp. (Oleaceae), Europe – Phillyrea angustifolia
(F), P. media var. ligustrifolia (CH).
Material examined: on Phillyrea media var. ligustrifolia, Europe, Switzerland, Brissago, Lago
Maggiore, Ticino, 15 May 1966, Deighton (IMI 119435).
Notes: Type material of this species could not be traced, but a collection from Switzerland has been
examined. According to the original diagnosis, Fusicladium fraxini var. phillyreae Trotter, described
from galls of Braueriella phillyreae on Phillyrea media, differs from F. phillyreae in having much
smaller conidia (11–12 × 4–5.5 µm) and seems to be quite distinct. However, type material of this
variety could not be traced and examined.
10.2.36.
Fusicladium pisicola Linford, Phytopathology 16(8): 549 (1926)
Fig. 38
Lectotype: on Pisum sativum, USA, Utah, Exp. Station Logan, 21 Aug. 1923, M.R. Linford (M), as
F. brevipes Ellis & Everh., selected here; isolectotypes: BPI 424334 A, B.
Teleomorph: Unknown.
Ill.: LINFORD (1926: Pl. 27, C).
On leaflets, stipules and tendrils, leaf spots hypophyllous, varying in shape and size,
irregular to oblong, up to 10 mm wide, pale to dark brown, dark grey, sometimes
sooty black, paler on the upper leaf surface, yellowish grey to pale brown, sometimes
confluent, vein-limited, without conspicuous margin. Colonies hypophyllous, rarely
amphigenous, dark, velvety. Mycelium intercellular, subcuticular to intraepidermal,
hyphae branched, 3–5 µm wide, septate, subhyaline to pale olivaceous. Stromata composed of slightly thick-walled, rough-walled cells, 5–8 µm diam., forming stromatic
layers. Conidiophores scattered to aggregated, but not fasciculate, erumpent through
the cuticle, very short, conical, erect, unbranched, 5–24 × (4–)6–8 µm, aseptate,
olivaceous to brown, smooth, walls somewhat thickened, slightly swollen at the base,
truncate at the apex, conidiophores usually reduced to conidiogenous cells, unilocal,
determinate, with a single, subdenticulate locus, flat to slightly convex, 2–4 µm wide,
unthickened or almost so, somewhat darkened–refractive. Conidia solitary, ellipsoid–
ovoid, obovoid to short cylindrical, 12–28(–32) × 6–10(–14) µm, 0–1-septate, more
or less constricted at the septum, septa more or less median or somewhat in the upper
half, asperulate, coarsely verrucose to echinulate, walls somewhat thickened, apex
broadly rounded, base rounded to truncate, hila 2–4 µm wide, unthickened or almost
so, somewhat darkened–refractive.
76
Schlechtendalia 9 (2003)
Hosts and Distribution: on Pisum spp. (Fabaceae), North America – Pisum sativum
(USA, ID, UT).
Notes: The lectotype is a part of the material cited in the original publication.
10.2.37.
≡
≡
=
≡
≡
≡
≡
=
=
=
≡
≡
=
=
=
=
=
≡
≡
≡
=
=
≡
≡
=
=
≡
=
=
=
=
=
=
=
Fusicladium pomi (Fr.) Lind, Dan. fung.: 521 (1913)
Fig. 39
Spilocaea pomi Fr., Novit. fl. svec. 5: 79 (1819); syntypes: on Malus sylvestris, Sweden, Fr.,
Scler. exs. 260 (e.g., B, UPS).
Spilocaea pomi Fr.: Fr., Syst. mycol. 3: 504 (1832).
Actinonema crataegi Pers., Mycol. eur. 1: 52 (1822); type: on Crataegus torminalis (= Sorbus
torminalis), Switzerland, Neuchâtel (L).
Capillaria crataegi (Pers.) Link, in Willd., Sp. pl. ed. 4, T. 6(1): 22 (1824).
Phlyctidium crataegi (Pers.) Wallr., Fl. crypt. Germ. 2: 418 (1833).
Asteroma crataegi (Pers.) Rabenh., Deutschl. Krypt.-Fl. 1: 139 (1844).
Spilocaea crataegi (Pers.) S. Hughes, Canad. J. Bot. 36: 807 (1958).
Fumago mali Pers., Mycol. eur. 1: 9 (1822).
Helminthosporium pyrorum Lib. (p.p. in Pyri mali), Lib., Pl. crypt. ard., Fasc. 2, 188 (1832).
Cladosporium dendriticum Wallr., Fl. crypt. Germ. 2: 169 (1833); syntypes: B, STR.
Fusicladium dendriticum (Wallr.) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 357 ‘1869’
(1870).
Passalora dendritica (Wallr.) Sacc., Mycoth. ven., Cent. XII, 1246, Padua 1876 [Michelia 1:
265 (1878)].
Asteroma mali Desm., Ann. Sci. Nat. Bot., Sér. 2, 15: 141 (1841).
Asteroma crataegi var. pomi Desm., Ann. Sci. Nat. Bot. Sér. 3, 8: 36 (1846).
Asteroma crataegi var. sorbi Desm., Ann. Sci. Nat. Bot. Sér. 3, 8: 35 (1846).
Actinonema pomi Lév., Ann. Sci. Nat. Bot. Sér. 3, 9: 260 (1847).
Cladosporium orbiculatum Desm., Ann. Sci. Nat. Bot., Sér. 3, 11: 275 (1849); lectotype: on
living leaves of Sorbus domestica, herb. Desmazières (PC).
Fusicladium orbiculatum (Desm.) Thüm., Fungi austr., Cent. VIII, 774, Teplitz 1873.
Passalora dendritica var. orbiculata (Desm.) Berk., in Sacc., Mycoth. ven., Cent. XII, 1246,
Padua 1876 [Michelia 1: 265 (1878)].
Fusicladium dendriticum var. orbiculatum (Desm.) Sacc., Syll. fung. 4: 345 (1886).
Scolecotrichum venosum Bonord., in Rabenh., Fungi eur., Cent. VI, 582, Dresden 1863; lectotype:
on leaves of Malus sp., Germany, Westphalia, Rabenh., F. eur. 582 (HAL, M).
Passalora pyracanthae G.H. Otth, Mitth. Naturf. Ges. Bern 1868: 66 (1868).
Fusicladium pyracanthae (G.H. Otth) Vienn.-Bourg., Rev. Mycol. (Paris) 6: 155 (1941).
Spilocaea pyracanthae (G.H. Otth) Arx, Tijdschr. Plantenziekten 63: 198 (1957).
Fusicladium dendriticum var. opuli Thüm., Fungi austr., Cent. XI, 1091, Bayreuth 1874, nom.
nud.
Napicladium soraueri Thüm., Mycoth. univ., Cent. I, 91, Bayreuth 1875.
Fusicladium dendriticum var. soraueri (Thüm.) Sacc., Syll. fung. 4: 346 (1886).
Actinonema crataegi f. sorbi-torminalis Thüm., Herb. myc. oec., Fasc. XI, 527, Klosterneuburg
1877, nom. nud.
Fusicladium pirinum var. pyracanthae Thüm., Mycoth. univ., Cent. IX, 874, Klosterneuburg
1877.
Actinonema crataegi var. arachnoideum f. sorbi-torminalis Thüm., Mycoth. univ., Cent. XV,
1487, Klosterneuburg 1879, nom. nud.
Actinonema crataegi f. sorbi ariae Thüm., Mycoth. univ., Cent. XIV, 1372, Klosterneuburg
1879, nom. nud.
Fusicladium dendriticum var. pyracanthae Thüm., Hedwigia 18: 155 (1879).
Cladosporium dendriticum var. heteromeles Harkn. (1881), in herb.
Fusicladium pirinum var. amelanchieris Sacc., Syll. fung. 4: 346 (1886).
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
=
77
Basiascum eriobotryae Cavara, Atti Ist. Bot. Univ. Pavia 2(1): 433 (1888); neotype: on leaves of
Eriobotrya japonica, Italy (Caserto) and Portugal (Lisbona), 1891, J. Verissimo d’ Almeida,
Briosi & Cav., F. paras. 186 (IMI 7582), selected here; isoneotypes: H, HAL, LE, M.
≡ Fusicladium eriobotryae (Cavara) Cavara, in Briosi & Cav., F. paras. 186 (1892).
≡ Fusicladium melanconioides Ferraris, Ann. Mycol. 7: 284 (1909) (nom. nov.).
≡ Spilocaea eriobotryae (Cavara) S. Hughes, Canad. J. Bot. 31: 563 (1953).
= Fusicladium dendriticum f. microsperma Roum., Fungi sel. exs., Cent. LXI, 5592, Toulouse
1891.
= Fusicladium dendriticum var. (ã) eriobotryae Scalia, Boll. Accad. Gioenia Sci. Nat. Catania
1901: 5 (1901).
= Fusicladium dendriticum var. sorbinum Sacc., Ann. Mycol. 3: 170 (1905); syntype: on leaves of
Sorbus aucuparia, Italy, Selva (Treviso), Aug. 1904, Sacc., Mycoth. ital. 1582 (B).
= Fusicladium dendriticum var. sorbinum f. fruticola Ferraris, Fl. Ital. Crypt., Pars I. Fungi, Fasc.
6: 879 (1910).
= Fusicladium photinicola McClain, Phytopathology 15: 181 (1925); syntypes: on Photinia (=
Heteromeles) arbutifolia, USA, California, Riverside, McClain (M, NY).
≡ Spilocaea photinicola (McClain) M.B. Ellis, More Dematiaceous Hyphomycetes: 112 (1976).
= Fusicladium lalandi É.J. Marchal & Verpl., Bull. Soc. Roy. Bot. Belgique 59: 24 (1926).
= Fusicladium dendriticum var. sorbi-torminalis S|vul. & Sandu, Hedwigia 73: 24 (1933).
= Coniosporum mali Dearn. & A.C. Foster, Canad. J. Res., Sect. C, Bot. Sci. 16: 274 (1938); type:
DAOM.
= Spilocaea amelanchieris I.C. Harv., in I.C. Harv. & Braithwaite, New Zealand J. Agric. Res. 25
(3): 441 (1982); holotype: on Amelanchier sp., New Zealand, Canterbury, 10 Nov. 1978, Harvey
(IMI 233721).
Teleomorph: Venturia inaequalis (Cooke) G. Winter, Hedwigia 36: 81 (1897).
Lit.: SACCARDO (1884: 206; 1886: 345, 346, 348; 1895: 617; 1906: 579; 1913: 1376), ADERHOLD
(1886: 876–913; 1897: 67–63), LINDAU (1907: 779–783), FERRARIS (1912: 315–317, 318–319),
GONZÁLES FRAGOSO (1927: 184–186), VASSILJEVSKY & KARAKULIN (1937: 194–195, 196–199),
BARR (1968: 808–809), ELLIS (1971: 143; 1976: 111–113), SUBRAMANIAN (1971: 361–363), RAABE
& G ARDENER (1972: 914–916), Fungi Canadenses (No. 35), OSIPYAN (1975: 433–439, 441),
SHVARTSMAN et al. (1975: 118–120, 140), CMI Descr. (No. 401), SIVANESAN (1977: 71–76; 1984a:
615–616), BRANDENBURGER (1985: 256, 258), MELNIK & POPUSCHOI (1992: 176), ELLIS & ELLIS
(1997: 164).
Ill.: ADERHOLD (1886: Pl. XXIX, Figs 1, 2, Pl. XXX, Figs 1, 2 a, b, c, 5, 6, 10, 11; 1897: Pl. IV, Fig.
4), LINDAU (1907: 780, Fig.), MCCLAIN (1925: 181, Fig. 1; 182, Fig. 2 A, B), VASSILJEVSKY &
KARAKULIN (1937: 194, Fig. 12), Arx (1952: 263, Fig. 2), BARR (1968: 805, Fig. 18), ELLIS (1971:
142, Fig. 95 A), SUBRAMANIAN (1971: 362, Fig. 260), Fungi Canadenses (No. 35, Figs 2, 4, 6, 7),
SHVARTSMAN et al. (1975: 119, Fig. 59), CMI Descr. (No. 401, Figs D, E), ELLIS (1976: 112, Figs
79, 80), SIVANESAN (1977: 74, Fig. 38 A, B; 1984a: 617, Fig. 372 B), BRANDENBURGER (1985:
1109, Fig. 287), MELNIK & POPUSCHOI (1992: 177, Fig. 130 a, b), ELLIS & ELLIS (1997: Pl. 85, Fig.
882).
Exs.: Allesch. & Schn., F. bavar. 594; Barthol., F. Columb. 3326; Briosi & Cav., F. paras. 140,
186; Calif. F. 636; Cooke, F. brit. exs. 645; Ellis, N. Am. F. 2792; Fuckel, F. rhen. 115, 456; F.
latv. exs. 600; Jaap, F. sel. exs. 513; Kab. & Bub., F. imp. exs. 45, 46; Krieger, F. sax. 198, 1748;
Krypt. exs. 1190, 1473; Lib., Pl. crypt. ard. 188; Migula, Crypt. Germ. Austr. Helv. exs. 63;
Mycoth. Fenn. 297; Mycoth. Ross. 100; Neger, Forstschäd. P. 163; Oudem., F. neerl. ex. 190,
198; Petr., F. polon. 49; Rabenh., F. eur. 582, 1168, 1764, 3997; Rabenh., Herb. mycol. Ed. II,
766; Reliqu. Petrak. 2566; Roum., F. gall. exs. 322, 323, 438; Roum., F. sel. exs. 5592; Sacc.,
Mycoth. ital. 1582; Sacc., Mycoth. Ven. 1067, 1246; Seym. & Earle, Econ. F. 38; Syd., Mycoth.
germ. 2249; Syd., Mycoth. march. 1494, 2790, 3793; Thüm., F. austr. 277, 774, 1091; Thüm.,
Herb. myc. oec. 42, 128, 178, 527, 722; Thüm., Mycoth. univ. 91, 261, 802, 874, 1174, 1372,
1487, 1891; Vestergr., Micromyc. rar. sel. exs. 1242; Westend. & Wall., Herb. crypt. belg. 695,
1090.
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SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
79
On living leaves, petioles, twigs and fruits, spots amphigenous, subcircular to irregular, 1–10 mm wide, at first pale olivaceous-brown, later greyish black, on leaves
sometimes with a brown, slightly raised margin, confluent, occasionally covering
large leaf segments, on fruits forming small, circular to larger and than irregular spots,
margin indefinite to whitish. Colonies amphigenous, in the center of the spots densely
caespitose, at the margin dendritic, radiating, olivaceous-brown to blackish, on young
twigs blister-like. Mycelium immersed, very rarely superficial, then conidiophores
solitary, arising from superficial hyphae. Stromata more or less well-developed, subcuticular, composed of subglobose cells, 5–10 µm diam., olivaceous-brown, relatively thick-walled. Conidiophores mostly in loose to dense fascicles, arising from
the upper cells of the stromata, erumpent through the cuticle, erect, straight to slightly
flexuous, cylindrical to ampulliform, unbranched, 10–50(–90) × 5–6 µm, variable in
length, depending on the age of the conidiophores, 0–1-septate, rarely pluriseptate,
pale to dark olivaceous-brown, paler towards the apex, smooth, walls thickened, often swollen at the base, up to 10 µm wide. Conidiogenous cells integrated, terminal,
with a single locus, proliferation percurrent, with a few to many conspicuous
annellations, loci truncate, 4–5(–6) µm wide, unthickened, not darkened. Conidia
solitary, shape variable, ovoid to obpyriform or obclavate, straight, (11–)14–23(–32)
× (4–)7–10(–13) µm, 0–1(–2)-septate, more or less constricted at the septa, pale to
medium olivaceous-brown, smooth, walls somewhat thickened, narrowly pointed or
broadly rounded at the apex, truncate at the base, hilum truncate, 4–5(–6) µm wide,
unthickened to occasionally very slightly thickened, not darkened.
Fig. 39: Fusicladium pomi. A – on Pyracantha sp., A1 – conidia, A2 – dense fascicle of conidiophores
with several annellations, B – on Malus sp., B1 – conidia, B2 – long septate conidiophore, B3 –
conidiophores, C – on Sorbus sp., C1 – conidia, C2 – conidiophores, C3 – conidiophore arising from
superficial hyphae, D – on Eriobotrya japonica, D1 – conidia, D2 – conidiophores, E – on Heteromeles
arbutifolia, E1 – conidia, E2 – conidiophores, F – on Amelanchier sp., F1 – conidia, C2 –
conidiophores, scale = 10 µm, A. Ritschel del.
Hosts and Distribution: on species of Amelanchier, Aronia, Cotoneaster, Docynia,
Eriobotrya, Heteromeles, Kageneckia, Malus, Prunus, Pyracantha, Pyrus, Sorbus
(Rosaceae), cosmopolitan – Amelanchier sp. (Europe, F; New Zealand), Aronia
prunifolia (= Pyrus floribunda) (Europe, D), Cotoneaster aitchisoni (Asia, India), C.
integerrimus (Europe, CH), C. suavis (Caucasus, Georgia), Docynia indica (Asia,
India), Eriobotrya japonica [Asia, China, Israel (Palestine), Iran, Japan, Jordan, Lebanon, Russia, Turkey, Uzbekistan; Caucasus, Armenia, Georgia; Europe, CH, D, E,
GB, GR, F, I, P, RUS, SLO, Cyprus; Africa, Libya, Morocco, South Africa; North
America, USA, CA, FL, OH, WA; South America, Chile; Australia, New South Wales,
Tasmania; New Zealand], Heteromeles arbutifolia (= Photinia arbutifolia) (North
America, USA, CA), Heteromeles sp. (= Photinia sp.) (Europe, GB; Australia, Tasmania), Kageneckia oblonga (North America, USA, CA), Malus domestica (incl. all
of the cultivars) (Asia, Afghanistan, China, India, Iran, Kazakhstan, Russia, Turkey,
Turkmenistan, Uzbekistan; Caucasus, Armenia; Europe, CS, D, I, E, EW, GB, H, RO,
RUS, S, SF, Cyprus; North Africa, Libya; North America, Canada, Alta., BC., Man.,
NB., Nfld., NS., Ont., PEI, Que., Sask., USA, AK, CA, GA, FL, IL, MA, WI, VA;
South America, Chile), M. sylvestris (= Pyrus malus) [Asia, China, India, Israel (Palestine), Jordan, Nepal, Pakistan, Russia, Turkey, Turkmenistan, Uzbekistan; Europe,
A, BG, CS, D, DK, E, EW, GB, GR, I, IRL, LV, RO, RUS, S, SF, SLO, Cyprus;
Africa, Ethiopia, Libya, Madagascar, Morocco, Mozambique, South Africa; North
America, USA, AK, AL, CA, CT, DE, FL, ID, KS, MI, MS, MT, NC, ND, NE, OK,
OR, SD, TN, WA, WI; South America, Argentina, Chile, Columbia, Peru; Australia,
80
Schlechtendalia 9 (2003)
New South Wales, Queensland, Tasmania, Victoria; New Zealand], Malus sp. (Asia,
Afghanistan, China, India, Iran, Iraq, Japan, Jordan, Kazakhstan, Korea, Pakistan,
Russia, Saudi-Arabia, Syria, Taiwan; Caucasus, Armenia, Azerbaijan, Georgia; Europe, B, BG, CH, D, DK, EW, F, FR, GB, H, I, LT, LV, RO, Ukr., YU; Africa, Egypt,
Ethiopia, Libya, Zimbabwe; North America, Canada, Alta., BC., NB., NS., Ont., Que.,
Sask., USA, CT, MA, NC, NH, OH, PA, RI, SD, WI; Central America, Panama;
South America; New Zealand), Prunus spp. (Europe, GB), Pyracantha angustifolia
(New Zealand), P. coccinea (incl. cv. lalandei) (Caucasus, Georgia; Europe, B, D,
DK, F, GB, NL, RO, SLO, TR, Ukr.; South Africa; North America, Canada, BC.,
USA, AL, CA, DE, FL, GA, NC, OK, OR, WA), P. crenato-serrata (North America,
USA, IL), Pyracantha spp. (Europe, F; North America, Canada, BC., USA, FL, MO,
NC; Australia, New South Wales), Pyrus communis (Europe, D, EW, GB, TR), P.
serotina (Asia, Korea), P. sinensis (Asia, Korea), Pyrus spp. (Asia, Pakistan; North
America, USA, WI), Sorbus americana (North America, USA, IL, MN, NY, WA), S.
aria (Asia, Russia, Turkmenistan; Europe, A, CH, D, GB, SLO), S. aucuparia (Asia,
Russia; Europe, CS, D, DK, EW, I, SLO; North America, USA, CT, IL), S. boissieri
(Caucasus, Armenia), S. chamaemespilus (Europe, CH), S. domestica (Europe, D, E,
F, I, LV, SLO, TR), S. intermedia (= Pyrus suecica, = S. scandica) (Europe, D, EW,
FR), S. lanata (Asia, India), S. persica (Asia, Kazakhstan, Uzbekistan), S. tianschanica
(Asia, Kazakhstan, Uzbekistan), S. torminalis (Asia, Russia; Caucasus, Armenia,
Georgia; Europe, A, CS, D, GB, RO), Sorbus spp. (Asia, Russia; Caucasus, Armenia,
Georgia; Europe, CS, GB, H, RO, RUS, TR; North America, USA, WA).
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
81
species seems to be the same as Fusicladium pomi. The name of the type host “Crataegus lalandi”
does not exist and seems to refer to Pyracantha coccinea “Lalandiae” (cv. lalandei).
Records of this species from Docynia indica are from BILGRAMI, JAMALUDDIN & RIZWI (1991).
RAABE & GARDENER (1972) recorded Kageneckia oblonga as host of this species. The record of
Arum korolkovi (Araceae) as host in SAGDULLAEVA et al. (1990) is undoubtedly wrong, and records
from Viburnum opulus (e.g., Fusicladium dendriticum var. opuli Thüm., F. austr. 1091) have been
checked and proved to be based on misidentifications of the hosts.
10.2.38.
Fusicladium psoraleae (Ellis & Barthol.) S. Hughes & Piroz., Canad. J.
Bot. 50(12): 2532 (1972)
Fig. 40
Dicoccum psoraleae Ellis & Barthol., in Ellis & Everh., Fungi Columb. 1820 (1903); lectotype:
on Psoralea tenuiflora, USA, Kansas, Stockton, 27 Jun. 1903, E. Bartholomew 3044, ex herb.
Ellis (NY), selected here; isolectotype: DAOM and F. Columb. 1820 (BPI 423604, NY).
Teleomorph: Unknown.
Ill.: HUGHES & PIROZYNSKI (1972: 2532, Fig. 6).
Exs.: Ellis & Everh., F. Columb. 1820.
≡
Colonies on discoloured areas of leaves, petioles and stems, dense, effuse, olivaceous,
velvety. Mycelium immersed, composed of branched, colourless, thin-walled hyphae.
Stromata subcuticular, sometimes intraepidermal, punctiform to effuse,
pseudoparenchymatous, composed of rounded to polygonal, pale brown, thin-walled
cells, 5 µm diam., few layers thick. Conidiophores solitary or in loose to dense groups,
Material examined: collections from herb. B, HAL, HBG, IMI, JE, LE, M, NY, PC.
Notes: Spilocaea pomi was originally used for the anamorph of Venturia inaequalis on Pyrus spp.
SIVANESAN (1977, 1984a) recorded the latter species from Cotoneaster integerrimus, Malus,
Pyracantha, Pyrus and Sorbus species. Morphologically very similar fungi occur on some other
hosts of the Rosaceae (Amelanchier, Heteromeles, Eriobotrya, Kageneckia). Some of them have
been considered to be distinct species. Detailed morphological examinations of Spilocaea pomi-like
fungi have been carried out by RITSCHEL (2001). Collections from Malus, Sorbus and Amelanchier
species are indistinguishable from each other. There are no differences in the conidial shape and size
(RITSCHEL 2001). Conidia from Eriobotrya and Pyracantha species are slightly shorter on an average, and collections from Heteromeles are somewhat larger in general, but these differences are not
significant. Therefore, Spilocaea amelanchieris, S. eriobotryae, S. photinicola and S. pyracanthae
are reduced to synonym with Fusicladium pomi. Spilocaea ahmadii on Pyrus pashia is the only
species in this complex which is morphologically clearly distinct from F. pomi by having much
longer and narrower conida. RAABE & GARDENER (1972) carried out inoculation experiments with
collections from Eriobotrya, Kageneckia, Heteromeles and Pyracantha, and proposed to refer all of
them to Spilocaea pyracanthae. Based on inoculation experiments, MENON (1956) proposed some
“formae speciales“ of Venturia inaequalis for races on Pyrus malus, Sorbus aucuparia, Cotoneaster
integerrimus and Crataegus oxyacantha (the latter one was misapplied and referred to Venturia
crataegi Aderh.). Molecular examinations carried out during the course of monographic studies
(RITSCHEL 2001) showed that V. inaequalis and Spilocaea pyracanthae are very closely allied or
even indistinguishable. Collections from Sorbus species are also indistinguishable from F. pomi. The
fungus on Eriobotrya was originally described as Basiascum eriobotryae Cavara. HUGHES (1953)
proposed the combination Spilocaea eriobotryae, and RAABE & GARDENER (1972) reduced it to
synonymy with Spilocaea pyracanthae.
HUGHES (1958) examined type material of Spilocaea pomi from UPS. Type material of Fusicladium
lalandi É.J. Marchal & Verpl. could not be traced but, according to the original description, this
Fig. 40: Fusicladium psoraleae. A – conidia. B – dense fascicles of conidiophores, scale = 10 µm,
K. Schubert del.
82
Schlechtendalia 9 (2003)
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
83
sometimes sporodochial, forming large expanded layers, arising from stromata,
cylindrical to narrowly ovoid or obpyriform, 6–40 × 3–5 µm, aseptate, thinwalled, pale brown, smooth to verruculose, mainly in the upper half of the
conidiophores, conidiophores usually reduced to conidiogenous cells.
Conidiogenous cells unilocal, determinate or with two or only few conidiogenous
loci, proliferation sympodial, loci flat, truncate, 2–3 µm wide, unthickened, not
darkened. Conidia solitary, broadly ellipsoid, ovoid to clavate, straight, sometimes slightly curved, 10–27(–34) × 6–10(–11) µm, 0–1(–3)-septate, not or only
slightly constricted at the septa, at first hyaline, becoming pale brown, smooth
to coarsely verrucose, rugose, rounded at the apex, hila flat, truncate, 2–3 µm
wide, unthickened, not darkened.
Hosts and Distribution: on Psoralea spp. (Fabaceae), North America – Psoralea
argophylla (Canada, Man.), P. tenuiflora (USA, KS).
Material examined: on Psoralea tenuiflora (= P. floribunda), USA, Kansas, Reley Co., 19 Jun.
1951, C.T. Rogerson (NY).
10.2.39.
Fusicladium pyrorum (Lib.) Fuckel, Jahrb. Nassauischen Vereins Naturk.
23–24: 357 ‘1869’ (1870), as ‘Fusicladium pyrinum’
Fig. 41
Helminthosporium pyrorum Lib. (p.p.), Pl. crypt. ard., Fasc. 2, 188 (1832); lectotype: on leaves
of Pyrus communis, Pl. crypt. ard. 188 (DAOM).
≡ Passalora ‘pyrina’ (Lib.) Sacc., Michelia 1: 537 (1879).
≡ Megacladosporium pyrorum (Lib.) Vienn.-Bourg., Les Champignons parasites des plantes
cultivées 1: 489 (1949), as ‘Megacladosporium pirinum’.
= Arthrinium pyrinum Wallr., Fl. crypt. Germ. 2: 163 (1833); holotype: herb. Wallroth (STR);
isotype: IMI 68300.
= Fusidium pyrinum Corda, Icon. fung. 1: 3 (1837); type: PR.
= Fusicladium virescens Bonord., Handb. Mykol.: 80 (1851); iconotype: Bonorden, l.c.: Fig. 94.
= Cladosporium polymorphum Peyl, Lotos 15: 18 (1865).
= Fusicladium fuscescens Rabenh., Bot. Zeitung (Berlin) 15: 430 (1857); syntypes: on leaves of
Malus domestica (= Pyrus malus), Germany, Dresden, autumno, L. Rabenhorst, Herb. mycol.
588 (HAL, HBG).
= Passalora pomi G.H. Otth, Mitth. Naturf. Ges. Bern 1868: 66 (1868); type: on leaves of Pyrus
coronarius, Swiss, Bern.
= Fusicladium pyrorum [(Lib.) Fuckel] var. cladophilum Ellis & Everh., North Am. Fungi 2791
(1892); syntypes: on twigs of Pyrus communis, USA, Wisconsin, Pewankee, May 1890, Geo P.
Pfeiffer, Ellis & Everh., N. Am. F. 2791 (M, NY).
= Cercospora porrigo Speg., Anales Mus. Nac. Buenos Aires. II. 3: 341 (1899); holotype: on
fruits of Malus domestica (= Pyrus malus), Argentina, prov. Buenos Aires, La Plata, Nov. 1894,
C. Spegazzini, no. 934 (LPS).
= Fusicladium pyrorum [(Lib.) Fuckel] f. carpophila Sacc., Mycoth. ital. 992 (1901); syntype: on
fruits of Pyrus communis, Italy, Selva, Treviso, Aug. 1901, D. Saccardo, Mycoth. ital. 992 (B).
= Acrotheca dearnessiana Sacc., Ann. Mycol. 10: 314 (1912); syntype: on Aronia melanocarpa
(= Pyrus melanocarpa), Canada, Ont., London, Aug. 1910, J. Dearness, Barthol., F. Columb.
5001 (IMI 7073).
≡ Fusicladium dearnessianum (Sacc.) M.B. Ellis, in herb.
Teleomorph: Venturia pyrina Aderh., Landw. Jahrb. 25: 875 (1896), as ‘pirina’.
Lit.: LINDAU (1907: 781–782), FERRARIS (1912: 313), VASSILJEVSKY & KARAKULIN (1937: 195–
196), HUGHES (1953: 566–567; 1958: 768), BARR (1968: 811–812), SUBRAMANIAN (1971: 234),
Fungi Canadenses (No. 36), CMI Descr. (No. 404), SIVANESAN (1977: 94–99; 1984a: 620–621).
≡
Fig. 41: Fusicladium pyrorum. A – conidia, B – microcyclic conidiogenesis, C – solitary or fasciculate
conidiophores, scale = 10 µm, K. Schubert del.
84
Schlechtendalia 9 (2003)
Ill.: BONORDEN (1851: 80, Fig. 94), ADERHOLD (1896: Figs 51–53, Pl. 1, E & F), BRIOSI & CAVARA
(F. paras. 43, Figs 1–3), FERRARIS (1910: 313, Fig. 99), VASSILJEVSKY & KARAKULIN (1937: 196,
Fig. 13), HUGHES (1953: 565, Fig. 5; 567, Fig. 6), BARR (1968: 805, Fig. 21), ELLIS (1971: 272, Fig.
186), SUBRAMANIAN (1971: 234, Fig. 203), Fungi Canadenses (No. 36), CMI Descr. (No. 404: 1,
Figs D, E), SIVANESAN (1977: 98, Fig. 53; 1984a: 621, Fig. 375), ARX (1987: 59, Fig. 28),
SAGDULLAEVA et al. (1990: 52, Fig. 6).
Exs.: Barthol., F. Columb. 4700, 5001; Briosi & Cav., F. paras. 43; Crypt. exs. 4104; Ellis, N. Am. F.
372; Ellis & Everh., N. Am. F. 2791; Erb. Critt. Ital. 696; Fl. Gall. Germ. exs. 597; Fl. Olten. exs.
568; Fuckel, F. rhen. 1517; F. est. 28825; Herb. Mycol. Rom. 1196, 1457; Jaap, F. sel. exs. 683;
Krieger, F. sax. 344, 2447; Krypt. exs. 1496, 4104; Lin., F. hung. 293; Migula, Crypt. Germ. Austr.
Helv. exs. 382; Petr., F. alban. bosn. exs. 13; Rabenh., F. eur. 1168; Rabenh., Herb. mycol. 588;
Reliqu. Petrak. 2159; Roum., F. gall. exs. 1868; Sacc., Mycoth. ital. 992; Sacc., Mycoth. Ven. 585;
Schmarotzerp. Ruhrg. 58, 114; Seym. & Earle, Econ. F. 39; Syd., Mycoth. germ. 1782; Syd., Mycoth.
march. 800; Thüm., F. austr. 276; Thüm., Herb. myc. oec. 224.
On leaves and fruits, rarely on young twigs and buds, leaf spots scab-like, amphigenous,
diffuse, subcircular, olivaceous to dark brown or almost black, surrounded by a paler
brown halo. Colonies amphigenous, caespitose, velvety, dark brown to olivaceousbrown. Mycelium subcuticular, hyphae branched, 3 µm wide, septate, pale brown.
Stroma almost lacking to well developed, stromatic cells 4–11 µm diam., dark brown,
thick-walled. Conidiophores solitary or in loose fascicles, arising from stromata, erect,
straight to flexuous, geniculate–subnodulose, unbranched, 11–70(–90) × 4–11 µm,
0–1-septate, olivaceous to dark brown, paler towards the apex, smooth, slightly roughwalled with age, thick-walled. Conidiogenous cells integrated, terminal or intercalary, with numerous conidiogenous loci, proliferation sympodial, loci denticulate,
short cylindrical, truncate or somewhat convex, 1–3 µm wide, walls unthickened or
almost so, slightly darkened–refractive. Conidia solitary, very rarely in unbranched
chains, fusiform to pyriform, ellipsoid to obovoid, straight to slightly curved, 10–34
× 5–11 µm, 0–1(–2)-septate, olivaceous to pale brown, smooth, later somewhat roughwalled, pointed at the apex, truncate at the base, hila 1–3(–4) µm wide, unthickened
or slightly thickened, somewhat darkened–refractive.
Hosts and Distribution: on Aronia, Chaenomeles, Eriobotrya, Malus and Pyrus
spp. (Rosaceae), cosmopolitan – Aronia melanocarpa (North America, Canada,
Ont.), Chaenomeles speciosa (North America, USA, OK), Eriobotrya japonica
(Europe, GB), Malus domestica (Europe, D; South America, Argentina), Pyrus
amygdaliformis (Europe, F, TR), P. bucharica (Central Asia, Tadzhikistan,
Uzbekistan), P. caucasica (Caucasus, Georgia), P. communis (= P. sativa) (Asia,
Israel, Lebanon, Iran, Iraq, Afghanistan, Russia, Kazakhstan, Kirghizia, Uzbekistan,
Turkmenistan, Tadzhikistan, China, India, Korea, Japan, Taiwan; Caucasus, Armenia, Azerbaijan, Georgia; Europe, A, AL, BG, CH, CZ, D, DK, EW, F, GB, H, I, LT,
LV, P, RO, RUS, SLO, TR, Ukr., YU, Byelorussia, Moldavia, Malta, Cyprus; Canary Islands; Africa, Libyan, Morocco, Mozambique, South Africa, Madagascar;
North America, Canada, BC., NB., NS., Ont., Que., PEI, USA, FL, MA, WI; South
America, Columbia, Argentina, Chile; Australia, New South Wales, Queensland,
Tasmania, Victoria; New Zealand;), P. coronarius (Europe, CH), P. korshinskyi
(Central Asia, Uzbekistan), P. mamorensis (Africa, Morocco), P. pyraster (Europe,
RO), Pyrus spp. (Asia, China; Europa, EW).
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
85
Material examined: on Eriobotrya japonica, Europe, Great Britain, Surrey, Wisley, 23 Dec. 1966,
A.V. Brook (IMI 123946), on Pyrus spp. collections from B, G, HBG, IMI, J, LE, M.
Notes: In a few cases, conidia forming secondary conidiophores and conidia by microcyclic
conidiogenesis have been found. This phenomenon usually occurs under high humidity. BONTEA
(1985) recorded Sorbus domestica as host of this species. Malus spp. are also known to be hosts of
Fusicladium pyrorum, e.g., in the types of F. fuscescens and Cercospora porrigo, but the correct
determinations of these hosts could not be proven.
10.2.40.1. Fusicladium radiosum (Lib.) Lind, Ann. Mycol. 3: 430 (1905) var.
radiosum
Fig. 42
Oidium radiosum Lib., Pl. crypt. ard., Fasc. 3, 285 (1834); lectotype: on Populus tremula, Belgium, Belgian Ardennes, 1834, Libert (BR), selected by MORELET 1985; isolectotypes: Lib., Pl.
crypt. ard., Fasc. 3, 285.
≡ Fusicladium radiosum (Lib.) Lindau, in Rabenh., Krypt.-Fl., ed. 2, 1(8): 777 (1907).
≡ Stigmina radiosa (Lib.) Goid., Ann. Bot. (Rome) 21: 11 (1936).
≡ Pollaccia radiosa (Lib.) E. Bald. & Cif., in E. Bald., Atti Ist. Bot. “Giovanni Briosi“ 10: 61 (1937).
≡ Venturia radiosa (Lib.) Ferd. & C.A. Jørg., Skovtraeernes Sygdomme 1: 125 (1938) (nom. anamorph.).
= Cladosporium ramulosum Roberge ex Desm., Ann. Sci. Nat. Bot. Sér. 2, 18: 361 (1852), non
Reisseck 1851; holotype: on Populus alba, France, Paris, Parc du Libisy, May 1851, Roberge
(PC 1518); isotype: herb. Desmazières 2135 (PC).
≡ Fusicladium ramulosum Rostr., Tidsskr. Skovbr. 6: 294 (1883), nom. nov., as ‘(Roberge, in Desm.)
Rostr.’.
≡ Pollaccia ramulosa (Rostr.) OndÍej, Eur. J. Forest Pathol. 2: 143 (1972), nom. nov., as ‘(Desm.) OndÍej’.
= Cladosporium asteroma Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 355 ‘1869’ (1870).
≡ Napicladium asteroma (Fuckel) Allesch., Ber. Bayer. Bot. Ges. 5: 25 (1897).
≡ Napicladium asteroma (Fuckel) Sacc., Malpighia 17: 421 (1902).
= ? Cladosporium asteroma [Fuckel] var. macrosporum Sacc., Michelia 2: 126 (1882).
= Fusicladium tremulae A.B. Frank, Hedwigia 22: 127 (1883); type: on Populus tremula, Germany, Berlin, Tegel, A.B. Frank (B).
≡ Napicladium tremulae (A.B. Frank) Sacc., Syll. fung. 4: 482 (1886).
= ? Cladosporium asteroma [Fuckel] var. microsporum Sacc., Syll. fung. 4: 357 (1886).
≡ Fusicladium radiosum [(Lib.) Lindau] var. microsporum (Sacc.) Lindau, in Rabenh. Krypt.-Fl.,
ed. 2, 1(8): 777 (1907).
= Fusariella populi Garb., Bull. Soc. Mycol. France 33: 89 (1917).
Teleomorph: Venturia tremulae Aderh., Hedwigia 36: 81 (1897) var. tremulae [MORELET (1985: 108)].
Lit.: ADERHOLD (1897: 81), SACCARDO (1886: 357; 1913: 1376), LINDAU (1907: 777, 778), GONZÁLES
FRAGOSO (1927: 188–190), SERVAZZI (1939: 49–153), VASSILJEVSKY & KARAKULIN (1937: 200),
ELLIS (1971: 142), ONDÌEJ (1972: 142), SIVANESAN (1977: 80), BRANDENBURGER (1985: 39), ELLIS
& ELLIS (1997: 192).
Ill.: ADERHOLD (1897: Pl. IV, Fig. 3), VASSILJEVSKY & KARAKULIN (1937: 201, Fig. 14), SERVAZZI
(1939: Pl. IX, Figs 44–47, Pl. X, Figs 48–53, Pl. XI, Figs 54–57), ELLIS (1971: 141, Fig. 94),
SUBRAMANIAN (1971: 86, Fig. 69), ONDÌEJ (1972: 142, Figs 1, 2), BRANDENBURGER (1985: 1101,
Fig. 273), SIVANESAN (1984a: 618, Fig. 373 B), MORELET (1985: 111, Fig. 5), WU & SUTTON (1995:
985, Fig. 5), ELLIS & ELLIS (1997: Pl. 82, Fig. 849).
Exs.: Allesch. & Schn., F. bavar. 597; Crypt. exs. 3625, 1499; Erikss., F. paras. scand. 298, 399;
Fuckel, F. rhen. 2208; Krieger, F. sax. 194; Lib., Pl. crypt. ard. 285; Mycoth. Ross. 247 a; Petr.,
Mycoth. gen. 1381; Romell, F. exs. 50; Syd., Mycoth. germ. 1783; Syd., Mycoth. march. 3095;
Thüm., F. austr. 536; Thüm., Herb. myc. oec. 340; Thüm., Mycoth. univ. 1170.
≡
On living leaves, petioles and twigs, spots amphigenous, circular to irregular, 5–20
mm wide, sometimes confluent, covering large parts of the leaf, at first olivaceous to
medium brown, later reddish brown with a black-brown, narrow to broad margin,
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Material examined: collections from herb. B, HAL, HBG, JE, M, PC.
Notes: Pollaccia species on poplars were often confused in the past, since it was unknown that
different species of Populus are inhabited by allied, but morphologically distinct species of this
genus. ADERHOLD (1897) found the teleomorph of Fusicladium tremulae and described it as Venturia
tremulae. LIND (1905) discussed the synonymy of this fungus, adopted the oldest valid name and
introduced the combination Fusicladium radiosum. BALDACCI & CIFERRI (1937) described the new
genus Pollaccia with P. radiosa as type species. SERVAZZI (1938) recognised differences between
Pollaccia collections on Populus tremula and P. nigra, and introduced the name Pollaccia elegans
for the latter fungus. MÜLLER & ARX (1950) considered P. radiosa the anamorph of Venturia macularis
(Fr.) E. Müll. & Arx, a species on Populus spp. of sect. Leuce. MORELET (1985) pointed out that
three Venturia species occur on Populus spp. of this section, and that F. radiosum has to be considered
the anamorph of Venturia tremulae.
Older records of Pollacia radiosa in the literature must be considered with caution. They often refer
to P. radiosa s.lat., making it impossible to assign them to any one of the varieties of F. radiosum.
Also, the identifications of the hosts are often uncertain and doubtful.
10.2.40.2. Fusicladium radiosum var. lethiferum (Peck) Ritschel & U. Braun comb.
nov.
Fig. 43
Cladosporium lethiferum Peck, Rep. (Annual) New York State Mus. Nat. Hist. 40: 64 (1887);
holotype: on Populus tremuloides, USA, New Hampshire, Keene, Jun. 1887, Peck (NYS); isotype:
BPI 427241.
≡ Pollaccia lethifera (Peck) M. Morelet, Bull. Soc. Sci. Nat. Archéol. Toulon Var 34 (219): 12
(1978).
≡ Pollaccia radiosa [(Lib.) E. Bald. & Cif.] var. lethifera (Peck) M. Morelet, Cryptog. Mycol. 6:
113 (1985).
= Clasterosporium populi Ellis & Everh., J. Mycol. 7: 134 (1892), non Saccardo 1886; types: on
Populus tremuloides, Canada, London, Dearness (DAOM, NY).
≡ Stigmina populi Pound & Clem., Bull. Geol. Nat. Hist. Surv. 9: 662 (1896), nom. nov., as ‘(Ellis
& Everh.) Pound & Clem.’.
≡ Stigmina populi Peck, Bull. New York State Mus. Nat. Hist. 157: 34 (1912), nom. nov., as ‘(Ellis
& Everh.) Peck’.
= Dicoccum populinum Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 1893: 462 (1894);
holotype: on Populus grandidentata, USA, Iowa, Iowa City, Jun. 1889 (NY).
= Fusicladium lageniforme Solheim & Hadfield, in Hadfield, Univ. Wyoming Publ. 1946: 18–19
(1946), nom. inval.
= Pollaccia americana OndÍej, Eur. J. Forest Pathol. 2: 144 (1972); holotype: on Populus
grandidentata, Canada, Kentville, 16 Jun. 1952, Creelman (DAOM), as Fusicladium radiosum
(Lib.) Lind.
Teleomorph: Venturia tremulae [Aderh.] var. grandidentatae M. Morelet, Cryptog. Mycol. 6: 113
(1985) [≡ Venturia moreletii Rulamort, Bull. Soc. Bot. Centre-Ouest, N.S., 17: 191 (1986)].
Lit.: SACCARDO (1892: 604; 1895: 621; 1899: 1083), ONDÌEJ (1972: 144), MORELET (1978: 12;
1985: 113–115).
Ill.: ONDÌEJ (1972: 143, Fig. 3; 144, Fig. 5), MORELET (1985: 114, Fig. 7).
≡
Fig. 42: Fusicladium radiosum var. radiosum. A – conidia, B – conidiogenous cells arranged in a
sporodochium, scale = 10 µm, A. Ritschel del.
sometimes leaf margin incurved, twigs often becoming necrotic, causing twig dieback. Colonies amphigenous, pale to dark olivaceous-brown, fructification at first formed in the center
of the leaf spots, dendritically expanding. Mycelium immersed, hyphae branched, septate,
smooth. Stromata intraepidermal to subcuticular, composed of rounded to somewhat angular, pale to medium olivaceous-brown, relatively thick-walled cells, 5–8 µm diam.
Conidiophores densely fasciculate, arising from the upper cells of the stromata, forming
sporodochial conidiomata, erumpent through the cuticle, erect, cylindrical to doliiform or
somewhat ampulliform, unbranched, short, 10–17 × 4–6(–8) µm, usually aseptate, medium
olivaceous-brown, smooth, walls somewhat thickened, conidiophores reduced to
conidiogenous cells, unilocal, determinate or occasionally percurrent, with up to two inconspicuous annellations, loci (2–)3–4(–5) µm wide, neither thickened nor darkened. Conidia
solitary, ellipsoid to obovoid, straight, sometimes slightly curved, (13–)18–26(–37) × 5–8
µm, mostly 1-septate, septum in the upper third, to 2-septate, second septum in the lower
third, rarely aseptate, more or less constricted at the septum, pale to dark olivaceous-brown,
smooth, wall somewhat thickened, rounded to pointed at the apex, obconically truncate at
the base, hilum truncate, (2–)3–4(–5) µm wide, unthickened, not darkened.
Hosts and Distribution: on Populus spp. (Salicaceae), Asia, Caucasus, Europe – Populus
alba (Europe, F), P. × canescens (Europe, F), P. tremula (Asia, Russia, Turkmenistan, Uzbekistan;
Caucasus, Armenia, Europe, BG, CS, D, DK, E, F, GB, I, LT, LV, N, PL, RO, RUS, S, SF, Ukr.,
YU), P. tremula × P. alba (Europe, F), P. tremula × P. tremuloides (Europe, F).
On living leaves, petioles and twigs, spots circular to irregular, 5–20 mm wide, yellowish brown, surrounded by a pale to later dark brown margin. Colonies amphigenous,
pale to dark olivaceous-brown, sometimes almost black, dendritically spreading.
Mycelium immersed. Stromata intraepidermal to subcuticular, composed of
subglobose, pale to dark olivaceous-brown, relatively thick-walled cells, 4–9 µm diam.
Conidiophores in dense fascicles, arising from the upper cells of stromata, forming
sporodochial conidiomata, erumpent through the cuticle, erect, short cylindrical to
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Material examined: on Populus grandidentata, Canada, Nova Scotia, New Glasgow, 26 Aug. 1908
(NYS), as Cladosporium lethiferum Peck; on Populus grandidentata, USA, Maryland, Prince
George’s, 26 Jun. 1951, Waterman (B), as Fusicladium tremulae A.B. Frank.
Notes: Based on the original diagnosis, SERVAZZI (1937) considered Cladosporium lethiferum to be
a species probably belonging in Pollaccia, but he did not introduce a formal combination. ONDÌEJ
(1972) described this fungus as Pollaccia americana, and MORELET (1978) published the combination P. lethifera. In 1985, MORELET found the teleomorph of this North American species which he
considered a variety of Venturia tremulae (var. grandidentatae), and he introduced the combination
Pollaccia radiosa var. lethifera for its anamorph. RULAMORT (1986) reintroduced Pollaccia lethifera
and proposed the new name Venturia moreletii for the teleomorph. However, this re-assessment was
not based on any new examinations or any other new data.
Morphological analyses of the whole Fusicladium radiosum complex (RITSCHEL 2001) showed that
the conidial sizes of var. radiosum and var. lethiferum are not significantly distinct. The conidia in
var. lethiferum are only distinguished from those of var. radiosum by being frequently curved. These
minor differences are not sufficient to treat this taxon as a separate species. We prefer to follow
MORELET (1978) and maintain its status as a variety. A molecular approach would be the only way to
re-assess the whole F. radiosum complex with all taxa included.
10.2.40.3. Fusicladium radiosum var. populi-albae (M. Morelet) Ritschel & U. Braun
comb. nov.
Fig. 44
Pollaccia radiosa [(Lib.) E. Bald. & Cif.] var. populi-albae M. Morelet, Cryptog. Mycol. 6: 112
(1985); holotype: on living leaves of Populus alba var. bolleana, Poland, Dabroszyn (the former
“Tamsel“, ca. 5 km north-east of Kostrzyn), 21 Sept. 1904, Sydow (PAD), as Cladosporium
asteroma [Fuckel] var. microsporum Sacc.
≡ Pollaccia populi-albae (M. Morelet) Rulamort, Bull. Soc. Bot. Centre-Ouest, N.S., 17: 191
(1986).
= Napicladium asteroma var. microsperma Sacc., in herb.
Teleomorph: Venturia tremulae [Aderh.] var. populi-albae M. Morelet, Cryptog. Mycol. 6: 112
(1985).
Lit.: SACCARDO (1886: 357), ONDÌEJ (1972: 143, 144), MORELET (1985: 112–113).
Ill.: ONDÌEJ (1972: 143, Figs 3, 4), MORELET (1985: 112, Fig. 6).
Exs.: Petr., Mycoth. gen. 1744; Petr., Fl. Bohem. Morav. exs. 1456; Syd., Mycoth. germ. 444.
≡
Fig. 43: Fusicladium radiosum var. lethiferum. A – conidia, B – conidiogenous cells arranged in a
sporodochium, scale = 10 µm, A. Ritschel del.
doliiform or ampulliform, unbranched, 8–12 × 4–6 µm, aseptate, pale to medium
olivaceous-brown, smooth, walls thickened, conidiophores reduced to conidiogenous cells,
unilocal, determinate or occasionally percurrently proliferating, with a single or two inconspicuous annellations, loci truncate, (2–)3–4(–5) µm wide, neither thickened, nor darkened. Conidia solitary, oblong–ellipsoid to fusiform, often slightly curved, sometimes
straight, 17–26(–33) × (5–)6–8(–10) µm, mostly 1-septate, septum in the upper third, to
2-septate, second septum in the lower third, rarely aseptate, often more or less constricted
at the septa, pale to medium olivaceous-brown, smooth, walls somewhat thickened, apex
pointed or rounded, truncate at the base, sometimes somewhat oblique, hila truncate to
slightly convex, (2–)3–4(–5) µm wide, not or only slightly thickened, not darkened.
Hosts and Distribution: on Populus spp. (Salicaceae), Europe, North America –
Populus alba (North America, Canada, Ont.), P. alba × P. grandidentata (North
America, Canada, Ont.), P. alba × P. jackii (North America, Canada, Que.), P.
grandidentata (North America, Canada, NB., NS., Ont., PEI, USA, MD, NY, WI), P.
tremuloides (Europe, BG, GB; North America, Canada, Alta., BC., Labr., Man., NB.,
Nfld., NS., Ont., PEI, Que., Sask., USA, AK, ID, ME, MT, NH, NY, PA, SD, WI).
On living leaves, petioles and twigs, leaf spots circular to irregular, 10–30 mm wide,
or confluent and larger, pale to reddish brown, with a dark brown, narrow to broad
margin, sometimes causing twig dieback. Colonies amphigenous, pale to later dark
olivaceous-brown, on the upper leaf surface often dendritic. Mycelium immersed.
Stromata intraepidermal to subcuticular, composed of subglobose, thick-walled cells,
5–9 µm diam. Conidiophores in loose to dense fascicles, arising from the upper cells
of stromata, forming sporodochial conidiomata, erumpent through the cuticle, erect,
straight to slightly flexuous, cylindrical to ampulliform or doliiform, unbranched,
short, 6–10 × 6–7(–7.5) µm, aseptate, medium olivaceous-brown, smooth, walls somewhat thickened, swollen at the base, up to 15 µm wide, conidiophores reduced to
conidiogenous cells, unilocal, determinate or sometimes percurrent, with up to two
inconspicuous annellations, loci truncate, 3–6 µm wide, unthickened, not darkened.
Conidia solitary, ellipsoid to pyriform, straight, 12–21(–25) × (6–)7–9(–12) µm,
(0–)1(–2)-septate, septum in the upper third, often constricted at the septa, pale to
medium olivaceous-brown, smooth, walls slightly thickened, apex rounded to pointed,
truncate at the base, hilum truncate, 3–6 µm wide, unthickened, not darkened.
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Fig. 45: Fusicladium romellianum. A – conidia, B – conidiogenous cells with few conidiogenous
loci, C – conidiophores arranged in a sporodochium, scale = 10 µm, A. Ritschel del.
Fig. 44: Fusicladium radiosum var. populi-albae. A – conidia, B – conidiogenous cells arranged in
a sporodochium, scale = 10 µm, A. Ritschel del.
Phaeoramularia maculicola (Romell & Sacc.) B. Sutton, Canad. J. Bot. 48: 471 (1970).
Cladosporium maculicola (Romell & Sacc.) M. Morelet, Bull. Soc. Sci. Nat. Archéol. Toulon
Var 201: 4 (1972).
= Pollaccia borealis A. Funk, Canad. J. Bot. 67: 776 (1989); holotype: on leaves of Populus
tremuloides, Canada, BC., Cassier, 13 Sept. 1987, A. Funk (DAVFP 23609).
Teleomorph: Venturia borealis A. Funk, Canad. J. Pl. Pathol. 11: 355 (1989).
Lit.: ELLIS (1976: 322).
Ill.: ONDÌEJ (1973: 238, Fig. 4), ELLIS (1976: 323, Fig. 243 A), FUNK (1989a: Figs 1–8).
Exs.: Fl. Suec. 17688; Mycoth. Ross. 247; Reliqu. Petrak. 328; Syd., Mycoth. germ. 2248.
≡
≡
Hosts and Distribution: on Populus spp. (Salicaceae), Europe, North America –
Populus alba (Europe, CH, CS, D, DK, H, F, I, LV, PL; North America, USA, NY).
Material examined: collections from herb. B, H, JE, M, PAD.
Notes: ONDÌEJ (1972) used the name Cladosporium ramulosum for this fungus from Populus alba, but
this name is a synonym of Pollaccia radiosa var. radiosa. MORELET (1985) introduced the name P.
radiosa var. populi-albae for this taxon, which was later raised to species rank by RULAMORT (1986).
Morphological and morphometric examinations, carried out by RITSCHEL (2001), confirmed the observations of MORELET (1985) that the conidial size in collections from Populus alba is only slightly different
from those of other varieties of P. radiosa. Therefore, we prefer to follow MORELET´s (1985) taxonomy,
in which this fungus was treated as variety of the latter species. Furthermore, it is noteworthy that Populus
alba is infected by all varieties of this species, i.e., a strict host separation is not evident.
10.2.41.
≡
Fusicladium romellianum ONDÌEJ, „eská Mykol. 27(4): 237 (1973) Fig. 45
Torula maculicola Romell & Sacc., in Sacc., Grevillea 21: 69 (1893), non Fusicladium maculicola
(Ellis & Kellerm.) OndÍej; lectotype: on Populus tremula, Sweden, Nacka Vikdalen, L. Romell,
24 Jun. 1890 (S), selected by SUTTON (1970); isolectotypes: WINF(M) 11082 (slide), IMI 17008
and ‘ad Holmiae Cap.’ 1890/1891, Romell (PAD).
Leaf spots more or less circular, 1.5–3 mm wide, brown, later with a whitish to pale
grey centre, often surrounded by a dark reddish brown, sometimes somewhat raised
margin. Colonies mainly hypophyllous, pale brownish, fructification spread over the
whitish centre. Mycelium immersed, hyphae branched, septate, brown. Stromata
intraepidermal, olivaceous to yellowish brown, 20–100 µm diam., composed of loosely
aggregated cells. Conidiophores in dense fascicles, arising from stromata, forming
sporodochial conidiomata, erumpent through the cuticle, erect, straight or flexuous,
cylindrical to conical, unbranched, 5–25 × 2–4 µm, aseptate, pale brown, olivaceous,
smooth, conidiophores usually reduced to conidiogenous cells, unilocal, determinate
or with two or only few conidiogenous loci, subdenticulate, unthickened, not or only
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very slightly darkened–refractive. Conidia solitary or catenate, in simple or branched
chains, oblong, cylindrical or ellipsoid, straight or slightly curved, (7–)11–21(–28) ×
3–7 µm, 0(–1)-septate, pale brown, sometimes subhyaline, smooth, walls thin to somewhat thickened, attenuated towards apex and base, apex pointed or truncate, base truncate, hila 1–2 µm wide, unthickened, not or only very slightly darkened–refractive.
Hosts and Distribution: on Populus spp. (Salicaceae), Asia, Europe, North America
– Populus alba (Asia, Kazakhstan), P. cataracti (Asia, Kazakhstan), P. deltoides (Asia,
India), P. grandidentata (North America, Canada, Ont., Que.), P. italica (Asia,
Kazakhstan), P. laurifolia (Asia, Kazakhstan), P. nigra (Asia, Kazakhstan), P. pilosa
(Asia, Kazakhstan), P. pruinosa (Asia, Kazakhstan), P. talassica (Asia, Kazakhstan),
P. tremula (Asia, Kazakhstan; Europe, D, LV, S, SF), P. tremuloides (North America,
Canada, Alta., BC., Man., Nfld., NWT, Ont., Que., Sask., USA, CO, UT, WI), P.
trichocarpa (North America, Canada, BC.), Populus spp. (Asia, Kazakhstan; North
America, Canada, Ont., USA, AK).
Material examined: collections from B, DAVFP, HBG, M.
Notes: Fusicladium romellianum has often been confused with Pollaccia radiosa, the anamorph of
Venturia tremulae. Both species can easily be distinguished by their symptoms and the morphology
of the conidia. Pollaccia radiosa causes irregular, large, blackish leaf spots on species of Populus.
The conidia are formed singly, they are larger than those of F. romellianum (15–42 × 6–11 µm) and
often septate (with 1–2 septa). Pollaccia borealis is morphologically indistinguishable from F.
romellianum. The teleomorph, Venturia borealis, has only been found and proven in connection with
Pollaccia borealis (FUNK 1989b).
10.2.42.
Fusicladium saliciperdum (Allesch. & Tubeuf) Tubeuf, Arbeiten Biol.
Reichsanst. Land-Forstw. 2: 568 (1902)
Fig. 46
Fig. 46: Fusicladium saliciperdum. A – conidia, B – conidiogenous cells arranged in a sporodochium,
scale = 10 µm, A. Ritschel del.
≡
Septogloeum saliciperdum Allesch. & Tubeuf, in Allesch. & Schn., Fungi bavar. 485 (1895);
lectotype: on Salix bicolor (= S. laurina), Germany, Bavaria, Tutzing, Jun. 1895, Tubeuf (M),
selected here; isolectotypes: Allesch. & Schn., F. bavar. 485.
≡ Fusicladium saliciperdum (Allesch. & Tubeuf) Lind, Ann. Mycol. 3: 430 (1905).
≡ Pollaccia saliciperda (Allesch. & Tubeuf) Arx, Tijdschr. Plantenziekten 63: 233 (1957).
Teleomorph: Venturia saliciperda J. Nüesch, Phytopathol. Z. 39: 349 (1960).
Lit.: ADERHOLD (1897: 82–83, as ‘Fusicladium ramulosum Rostr.’), SACCARDO (1899: 1031; 1913:
1376), LINDAU (1907: 776), VASSILJEVSKY & KARAKULIN (1937: 201–203), CMI Descr. (No. 482),
E LLIS (1976: 110), SIVANESAN (1977: 102–105; 1984a: 622), Fungi Canadenses (No. 247),
BRANDENBURGER (1985: 45), ELLIS & ELLIS (1997: 246).
Ill.: ADERHOLD (1897: Pl. IV, Fig. 4), VASSILJEVSKY & KARAKULIN (1937: 201, Fig. 14; 202, Fig.
15), ONDÌEJ (1973: 238, Fig. 8), CMI Descr. (No. 482, Fig.), ELLIS (1976: 110, Fig. 77 C), Fungi
Canadenses (No. 247, Figs 1–3), SIVANESAN (1984a: 618, Fig. 373 C).
Exs.: Allesch. & Schn., F. bavar. 485; Kab. & Bub., F. imp. exs. 642; Mycoth. Ross. 146; Petr., Fl.
Bohem. Morav. exs. 925; Petr., Mycoth. gen. 735; Syd., Mycoth. germ. 2796.
30 µm deep, composed of subcircular to slightly angular, somewhat thick-walled
cells, 5–10 µm diam. Conidiophores in dense fascicles, arising from the upper cells
of stromata, forming sporodochial conidiomata, erumpent through the cuticle, erect,
cylindrical to slightly ampulliform, unbranched, short, 8–15 × 5–8 µm, aseptate, pale
to medium olivaceous-brown, smooth, walls somewhat thickened, conidiophores usually reduced to conidiogenous cells, unilocal, determinate or occasionally percurrently
proliferating, with a single or two, more or less conspicuous annellations, loci truncate, 3–6 µm wide, unthickened. Conidia solitary, cylindrical to ellipsoid, often inflated in the middle, straight, sometimes slightly curved, (12–)16–23(–30) × (5–)7–
9(–11) µm, mostly 1-septate, septum in the upper half, occasionally aseptate or with
a second septum in the lower half, often slightly constricted at the septa, olivaceous
to yellowish brown, smooth, walls somewhat thickened, apex rounded, base (hilum)
truncate to slightly convex, 3–6 µm wide, unthickened, not darkened.
On living leaves, petioles and twigs, spots mostly hypophyllous, but also epiphyllous
along leaf veins, irregularly shaped, 1–3 mm wide, brown, surrounded by a dark brown,
somewhat raised, shining margin, often causing lateral distortions and necroses as
well as twig dieback. Colonies amphigenous, punctiform, scattered to dendritic, mainly
along leaf veins, dark brown to blackish. Mycelium immersed, pale to medium
olivaceous-brown. Stromata intraepidermal to subcuticular, 10–60 µm wide and 10–
Hosts and Distribution: on Salix spp. (Salicaceae), Asia, Caucasus, Europe, North
America – Salix alba (Europe, CZ, DK, GB, RUS, RO; north-eastern North America),
S. americana (Europe, D, PL), S. alba (incl. var. pendula) (North America, Canada,
NS., Que., USA, CT, MA, ME, NH, NY), S. alba × S. babylonica (Europe, D), S.
amygdalina (Europe, D, RUS), S. aurita (Europe, GB, DK), S. babylonica (Europe,
D, GB, RUS; North America, Canada, BC., NB., NS., Que., USA, CT, MA, NY, PA),
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S. bicolor (= S. laurina) (Europe, D), S. blanda (Europe, RUS; North America, Canada,
NS.), S. caprea (Europe, D, DK), S. cinerea (Asia, Kazakhstan; Europe, DK, RUS),
S. cordata (North America, Canada, Que.), S. cuspidata (Europe, DK), S. discolor
(North America, Canada, Que.), S. fragilis (Europe, D, DK, RO; North America,
Canada, NS., USA, MA, NY), S. fragilis × S. pentandra (Europe, DK), S. gracilis
(Europe, D), S. japonica (Europe, D), S. lucida (North America, USA, CT, ME), S.
mollissima (Europe, DK), S. nigra (North America, Canada, NS., USA, CT, MA,
NY), S. pentandra (Europe, D; North America, Canada, Que.), S. purpurea (incl. var.
nana) (Europe, RO, RUS; North America, Canada, Que.), S. sericea (North America,
USA, CT, NY), S. silesiaca (Europe, RO), S. × smithiana (North America, Canada,
BC.), S. viminalis (Europe, RUS), S. vitellina (Europe, D; North America, Canada
BC., NS.), Salix spp. (Asia, Japan, Russia, Uzbekistan; Caucasus, Armenia, Azerbaijan,
Georgia; Europe, CZ, D, DK, GB, LT, LV, NL, RUS; North America, Canada, BC.,
Man., NB., Nfld., NS., Ont., PEI, Que., USA, north-eastern states, NC, PA, WA).
Material examined: collections from B, HBG, JE, M.
Notes: ROSTRUP (1883) used the name Fusicladium ramulosum for collections on species of Salix as
well as Populus, suggesting that they were the same as Cladosporium ramulosum, described by
Desmazières from Populus alba. ADERHOLD (1897) disagreed, assigned the fungus from Populus
spp. to Fusicladium tremulae and confined the name Fusicladium ramulosum to the species on Salix,
which he erroneously considered to be the anamorph of Venturia chlorospora. The first valid description of the Fusicladium on Salix spp. dates back to Allescher & Tubeuf, in ALLESCHER &
SCHNABL, Fungi bavarici 485, 1895, who introduced Septogloeum saliciperdum, which was later
transferred to Fusicladium by TUBEUF (1902). NÜESCH (1960) examined this fungus in vitro, found
the teleomorph and described it as Venturia saliciperda.
10.2.43.
Fusicladium scillae (Deighton) U. Braun & K. Schub., IMI Descriptions
of Fungi and Bacteria 152, no. 1518 (2002)
Fig. 47
Cladosporium scillae Deighton, in Laundon, New Zealand J. Bot. 8: 55 (1970); holotype: on
Scilla peruviana, New Zealand, Levin, 21 Dec. 1965, G.F. Laundon (IMI 116997).
Teleomorph: Unknown.
Ill.: LAUNDON (1970: 57, Fig. 4), IMI Descr. (No. 1518, Figs A–D).
≡
Leaf spots amphigenous, elliptical, small or occasionally up to 10 × 5 mm, dark brown,
finally greyish brown, margin indefinite. Colonies epiphyllous, effuse, brown, loose
to dense. Primary mycelium internal, inter- and intracellular in the mesophyll and
epidermal cells, composed of pale olivaceous, septate, sparingly branched hyphae,
1.5–2 µm wide, and subcuticular swollen hyphal cells, 8–12 × 4–8 µm, aggregated,
forming expanded subcuticular radiating strands, stromatic hyphal aggregations variable, almost lacking to well-developed. Secondary mycelium superficial, almost lacking to well-developed, hyphae creeping, septate, sparingly branched, 1.5–2.5 µm wide,
pale olivaceous, smooth. Conidiophores in small to large fascicles (up to 100 or even
more), loose to moderately dense, often caespitose, arising from internal hyphae or
stromatic hyphal aggregations, erumpent through the cuticle, sometimes emerging
through stomata, occasionally solitary, arising from creeping superficial hyphae, erect,
straight, subcylindrical to flexuous, geniculate–sinuous, usually unbranched, rarely
branched, 19–70(–120) × (1.5–)2–4 µm, septate, pale olivaceous, smooth, wall thin
Fig. 47: Fusicladium scillae. A – conidia, B – solitary conidiophores arising from creeping hyphae, C – conidiophores and conidiogenous cells, D – fasciculate conidiophores, scale = 10 µm,
K. Schubert del.
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97
to slightly thickened. Conidiogenous cells integrated, terminal, 10–25 µm long, with
a single or several conidiogenous loci, subdenticulate, proliferation sympodial, loci
1–1.5 µm wide, wall unthickened, non-pigmented, at most somewhat refractive. Conidia catenate, in simple or branched chains, subcylindrical, ellipsoid–ovoid (–fusiform), 7–22 × 2.5–4 µm, 0–3-septate, often somewhat constricted at the septa, pale
olivaceous, occasionally subhyaline, smooth to faintly rough-walled, apex obtuse in
primary conidia, truncate in secondary conidia, base obconically truncate, hila 1–1.5
µm wide, unthickened, not darkened.
Hosts and Distribution: on Scilla spp. (Liliaceae), New Zealand – Scilla peruviana
(New Zealand).
Material examined: on Scilla peruviana, New Zealand, Levin, 27 May 1965, E.A. Way (IMI 115480).
Notes: The conidiogenous loci in this species are quite distinct from those of Cladosporium species,
described in detail by DAVID (1997), and agree with Fusicladium by being subdenticulate, unthickened
and not darkened. Based on the structure of the conidiogenous loci and the features of the conidia,
Cladosporium scillae was transferred to Fusicladium.
10.2.44.
Fusicladium scribnerianum (Cavara) M.B. Ellis, More Dematiaceous
Hyphomycetes: 238 (1976)
Fig. 48
Cladosporium scribnerianum Cavara, Hedwigia 31: 143 (1892) and in Briosi & Cavara, F. paras.
187 (1892); syntype: on Betula populifolia, Italy, Pavia, 1890, F.L. Scribner, Briosi & Cavara, F.
paras. 187 (HAL).
Teleomorph: Unknown.
Lit.: FERRARIS (1912: 340).
Ill.: BRIOSI & CAVARA (F. paras. 187, Figs 1–4), ELLIS (1976: 238, Fig. 178).
Exs.: Briosi & Cavara, F. paras. 187.
≡
Leaf spots amphigenous, numerous, subcircular, 3–8 mm wide, on the upper leaf
surface olivaceous-greyish to dark brown, pale greenish below, centre of the spots
darker as the surrounding halo, sometimes confluent, spreading over the leaf veins.
Colonies only epiphyllous, effuse, velvety, olivaceous. Mycelium subcuticular,
hyphae 2–3 µm wide, septate, pale olivaceous. Stromata composed of brown, thickwalled cells, 5–8 µm diam. Conidiophores solitary or loosely to densely fasciculate,
arising from stromata, erumpent through the cuticle, erect, straight to slightly
flexuous, unbranched, 18–90 × 4–5.5 µm, 0–1-septate, septa not very conspicuous,
olivaceous to pale brown, paler towards the apex, smooth, walls somewhat thickened. Conidiogenous cells integrated, terminal, with a single to several conidiogenous
loci, proliferation sympodial, loci 2–3 µm wide, unthickened, but somewhat darkened–refractive, sometimes with percurrent proliferations which are not connected
with conidiogenesis. Conidia solitary or sometimes catenate, in unbranched or
branched chains, fusiform to obclavate or cylindrical, straight, 11.5–29.5 × 4.5–7
µm, 0–3-septate, slightly constricted at the septum, pale olivaceous, smooth to
verruculose, truncate at the base, hila 2–3 µm wide, unthickened, but somewhat
darkened–refractive.
Hosts and Distribution: on Betula spp. (Betulaceae), Asia, Europe – Betula pendula
(Asia, Kazakhstan; Europe, D, I), B. populifolia (Europe, I).
Fig. 48: Fusicladium scribnerianum. A – conidia, B – catenate conidia, C – conidiophores, scale =
10 µm, K. Schubert del.
Notes: Branched conidiophores, as described by FERRARIS (1912), have not been observed.
Fusicladium scribnerianum is morphologically close to the anamorph of Venturia ditricha, but differs in having conidia formed singly and in chains as well as darkened–refractive conidiogenous loci
and hila. Additional collections from Betula spp. and molecular data would be helpful to find the true
taxonomic position and affinity of F. scribnerianum.
10.2.45.
Fusicladium spiraeae Karak., Mater. Mikol. Obsl. Rossii 2: 82 (1915)
Fig. 49
Lectotype: on leaves of Spiraea crenifolia, Russia, Bashkortostan, Sterlitamak, 23 Jul. 1913, Karakulin
(LE), selected here.
≡ Scolecotrichum spiraeae (Karak.) Karak., in Vassiljevsky & Karakulin, Parazitnye
nesovershennye griby. Ch. I. Gifomitsety 1: 214 (1937).
≡ Pollaccia spiraeae (Karak.) OndÍej, „eská Mykol. 38(1): 46 (1984).
Teleomorph: Unknown.
Lit.: SHVARTSMAN et al. (1975: 132–133), BRANDENBURGER (1985: 226).
Ill.: ONDÌEJ (1984a: 47, Fig.).
Leaf spots amphigenous, irregular, 3–5 mm diam., reddish brown, with a somewhat
darker, slightly raised, shining margin, sometimes confluent. Colonies epiphyllous,
punctiform, scattered, dark brown to blackish. Mycelium immersed. Stroma subcuticular, 50–80 µm diam., composed of subcircular, olivaceous-brown cells, 5–10 µm
diam. Conidiophores in dense fascicles, arising from the upper cells of stromata, form-
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Fig. 49: Fusicladium spiraeae.
A – conidia, B – dense fascicle
of conidiophores, scale = 10
µm, A. Ritschel del.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
10.2.46.
99
Fusicladium subsessile (Ellis & Barthol.) K. Schub. & U. Braun, IMI
Descriptions of Fungi and Bacteria 152, No. 1519 (2002)
Fig. 50
Cladosporium subsessile Ellis & Barthol., Erythea 4: 83 (1896); lectotype: on leaves of Populus
deltoides ssp. monilifera (= Populus monilifera), USA, Kansas, 18 Sept. 1894, Bartholomew
(NY), selected here; isolectotypes: on leaves of Populus deltoides ssp. monilifera, USA, Kansas, Rockport, Sept. 1894, E. Bartholomew, Ellis & Everh., N. Am. F. 3288 (M, NY).
≡ Cladosporium brevipes Ellis & Barthol., Erythea 4: 27 (1896), homonym, non Peck, 1887.
Teleomorph: Unknown.
Ill.: IMI Descr. (No. 1519, Figs A–C).
Exs.: Ellis & Everh., N. Am. F. 3288.
≡
Leaf spots amphigenous, subcircular to irregular, 2–10 mm wide, sometimes confluent, greyish brown, margin darker brown, narrow. Colonies amphigenous, punctiform, dark brown to blackish, scattered. Mycelium internal, hyphae sparingly branched,
1.5–2 µm wide, septate, olivaceous. Stromata variable in size and shape, composed
of swollen hyphal cells, 4–7 µm diam., pale olivaceous, thick-walled. Conidiophores
aggregated in loose to dense fascicles or caespitose, rarely forming sporodochial
conidiomata, arising from stromatic hyphal aggregations, erect, straight,
subcylindrical–conical, slightly geniculate–sinuous, unbranched, (3–)9–26 × 3.5–5
ing sporodochial conidiomata, erumpent through the cuticle, erect, straight to sometimes slightly flexuous, cylindrical, unbranched, 5–15(–30) × 3–4 µm, 0–1-septate,
subhyaline, pale yellowish to pale olivaceous-brown, smooth, walls slightly thickened, often swollen at the base, up to 7 µm wide. Conidiogenous cells integrated,
unilocal, determinate or occasionally percurrently proliferating, with a single, inconspicuous annellation, locus truncate, (2–)3–4 µm wide, unthickened, not darkened.
Conidia solitary, narrowly cylindrical to obclavate, (15–)18–29(–35) × (3.5–)5–
6(–7) µm, 0–1-septate, sometimes slightly constricted at the septum, subhyaline, pale
yellowish to pale olivaceous-brown, smooth, walls somewhat thickened, apex pointed
or rounded, base (hilum) truncate, (2–)3–4 µm wide, not or only slightly thickened,
not darkened.
Hosts and Distribution: on Spiraea spp. (Rosaceae), Europe – Spiraea crenifolia
(RUS), S. hypericifolia (RUS), Spiraea spp. (RUS).
Material examined: on leaves of Spiraea hypericifolia, Russia, Bashkortostan, Micher., 24 May
1916, Poreunij (LE); on leaves of Spiraea crenifolia, Russia, Bashkortostan, Saratova, 9 Aug. 1919,
Karakulin (LE).
Notes: Karakulin, in VASSILJEVSKY & KARAKULIN (1937) described amphigenous colonies with
fasciculate conidiophores emerging through stomata, which could not be confirmed during the course
of the present monographic studies. The conidiophores observed were epiphyllous, erumpent through
the cuticle, which was also confirmed by examinations of ONDÌEJ (1984).
Fig. 50: Fusicladium subsessile. A – conidia, B – solitary and fasciculate conidiophores and
sympodially proliferating conidiogenous cells, C – fasciculate conidiophores and percurrently proliferating conidiogenous cells, scale = 10 µm, K. Schubert del.
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SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
101
µm, 0(–1)-septate, pale olivaceous, smooth to faintly rough-walled, wall thin to slightly
thickened. Conidiogenous cells integrated, terminal or conidiophores usually reduced
to conidiogenous cells, with 1–3, rarely more conidiogenous loci, subdenticulate,
proliferation sympodial as well as percurrent with 1–3 fine annellations, conidiogenous
loci often denticle-like, 1.5–2.5 µm wide, but wall of the loci unthickened, not darkened or at most slightly darkened–refractive. Conidia solitary or catenate, in simple
or occasionally branched chains, fusiform–subcylindrical, straight to somewhat curved,
9–24(–36) × 3.5–5(–6) µm, (0–)1(–3)-septate, pale olivaceous, smooth or faintly
rough-walled, wall thin to slightly thickened, attenuated towards apex and base, apex
rounded or truncate, base truncate or somewhat convex, hila 1.5–3 µm diam.,
unthickened, not darkened.
Hosts and Distribution: on Populus spp. (Salicaceae), North America – Populus
deltoides ssp. monilifera (= Populus monilifera) (USA, KS).
Notes: This species was reduced to synonymy with Phaeoramularia maculicola (Romell. & Sacc.)
B. Sutton (≡ Fusicladium romellianum OndÍej) by SUTTON (1970) and ELLIS (1976). However, F.
romellianum differs from F. subsessile in having conidiophores, which are usually formed in
sporodochial conidiomata, smaller conidiogenous loci and conidial hila (1–2 µm wide) and conidia,
which are usually aseptate, rarely 1-septate. F. romellianum occurs on a wide range of Populus
species, whereas F. subsessile is, as far as known, confined to Populus deltoides var. monilifera. It
may be supposed that the latter species has often been confused with other species of Fusicladium
and Pollaccia on Populus species. It is probably much more common than indicated under ‘hosts
and distribution’.
10.2.47.
Fusicladium veronicae (Bat.) B. Sutton & Pascoe, Austral. Syst. Bot. 1:
81 (1988)
Fig. 51
Ramalia veronicae Bat., Revista Biol. (Lisbon) 1(2): 111 (1957); holotype: on Parahebe
derwentiana, Australia, Clyde Mtn., N.S.W., Jan. 1937, L. Fraser (DAR 3568).
Teleomorph: Protoventuria parahebicola Pascoe & B. Sutton, Austral. Syst. Bot. 3: 281 (1990).
Ill.: SUTTON & PASCOE (1988: 81–85, Figs 1–7).
≡
On leaves, petioles and stems, leaf spots amphigenous, mainly hypophyllous,
subcircular, variable in size, up to 5 mm wide, dark olivaceous-brown, margin darker,
scattered over the leaf surface, frequently confluent, sometimes covering the entire
surface. Primary mycelium immersed, epidermal and subcuticular, in the upper epidermal cells intracellular, hyphae branched, 1–3 µm wide, septate, hyaline, between
the cuticle and upper epidermal wall plates of fan-like, hyaline, septate hyphae, forming one cell-layer, composed of irregularly lobed cells, 2.5–6.5 µm wide. Secondary
(superficial) mycelium not extensively developed, when present composed of irregularly branched, 3–5 µm wide, pale to medium brown, thin- or thick-walled, smooth
hyphae. Stromata absent. Conidiophores solitary, arising from the subcuticular mycelium, erumpent through the cuticle, erect, straight or slightly curved, flexuous, cylindrical, unbranched, 25–60(–70) × 5–8 µm, 1–5-septate, pale to medium brown,
paler towards the apex, verruculose, thick-walled, slightly attenuated towards the
apex. Conidiogenous cells integrated, terminal, with 1–2, rarely more loci, proliferation sympodial, loci 2–4 µm wide, unthickened or almost so, not darkened, often with
1–3 percurrent proliferations which are not connected with conidiogenesis. Conidia
Fig. 51: Fusicladium veronicae. A – conidia, B – germinating conidia and secondary conidia (microcyclic
conidiogenesis), C – conidiophores arising from hyphae, scale = 10 µm, K. Schubert del.
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solitary or catenate, in unbranched, short chains, fusiform, straight, 11.5–20 × 4.5–8
µm, 0–3-septate, mostly with one septum, slightly constricted at the septum, pale
brown, verruculose, attenuated to a conical apex and to a truncate base, hila 2–4 µm
wide, unthickened or almost so, not darkened.
Hosts and Distribution: on Parahebe spp. (Scrophulariaceae), Australia – Parahebe
derwentiana, P. formosa, P. perfoliata.
Material examined: on Parahebe perfoliata, Australia, Victoria, Burnley, P. R. I. Gardens, 4 Jun.
1986, H.Y. Yip (VPRI 13987); on P. perfoliata, Australia, Victoria, Burnley, P. R. I., 5 Dec. 1985, S.
Isaacs (VPRI 13120, DAR 55930).
10.2.48.
Fusicladium virgaureae OndÍej, „eská Mykol. 25(3): 170 (1971)
Fig. 52
Holotype: on Solidago virgaurea, Moravia, Hrubý Jesenik, „ervenohorské, 29 Jul. 1969, M. OndÍej
(BRA).
Teleomorph: Unknown.
Ill.: ONDÌEJ (1971: 168, Fig. 4).
Leaf spots amphigenous, shape and size variable, irregular, varying from small spots,
2–3 mm wide, to large blotches, 10–30 mm wide or oblong, up to 50 mm, yellowish,
olivaceous to brown, sometimes purple-brown. Stromata absent, only with small subcuticular aggregations of swollen hyphae. Conidiophores solitary or in small, loose
fascicles, erect, straight to somewhat flexuous, subcylindrical to slightly sinuous,
unbranched or rarely branched, 20–100 × 3–6 µm, septate, yellowish or olivaceous-
Fig. 52: Fusicladium virgaureae. A – conidia, B – branched conidiophore, C – conidiophores, scale
= 10 µm, K. Schubert del.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
103
brown to medium brown, paler towards the apex, smooth, occasionally minutely
rough-walled, walls somewhat thickened. Conidiogenous cells integrated, terminal,
with a single or up to four loci, proliferation sympodial, loci truncate to slightly convex, 1–1.5 µm wide, neither thickened nor darkened, rarely with percurrent proliferations, which are not connected with conidiogenesis. Conidia in unbranched chains,
cylindrical to fusiform or obclavate, straight, 8–16 × 3–6 µm, 0–1-septate, not or
only slightly constricted at the septum, yellowish or olivaceous-brown, smooth, hila
truncate to slightly convex, 1–1.5 µm wide, neither thickened nor darkened.
Hosts and Distribution: on Solidago spp. (Asteraceae), Europe – Solidago gigantea
(A), S. virgaurea (CZ, SK).
Material examined: on Solidago virgaurea, Moravia, Hrubý Jesenik, 16 Aug. 1981, M. OndÍej
(BRA); on S. virgaurea, Slovakia, Slovensko Vysoke Tatry, 7 Sept. 1974, M. OndÍej (BRA); on
Solidago gigantea, Austria, Steiermark, Grazer Feld, S of Graz, Kaiserwald, near Wundschuh, 16
Aug. 1995, P. Zwetko (GZU), see SCHEUER (2003).
10.2.49.
Fusicladium viticis M.B. Ellis, More Dematiaceous Hyphomycetes: 238
(1976)
Fig. 53
Holotype: on leaves of Vitex cienkowskii, Uganda, Mulanga Grassland, Sept. 1919, R.A. Dummer
4269 (IMI 102079b).
Teleomorph: Unknown.
Ill.: ELLIS (1976: 237, Fig. 177).
Definite leaf spots lacking or only with irregular discolorations, dark brown to
greyish brown. Colonies hypophyllous, effuse, dark brown, often spread along leaf
veins. Mycelium internal, subcuticular, hyphae branched, 2–3 µm wide, septate,
Fig. 53: Fusicladium viticis. A – conidia, B – conidiophores, scale = 10 µm, K. Schubert del.
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pale olivaceous. Stroma with flat, only one-layered aggregations of swollen, pale brown,
thick-walled cells, 3–6 µm diam., forming layers or radially spreading. Conidiophores
solitary or fasciculate, arising from stromata, erect, straight or slightly flexuous,
unbranched, 40–100 × 3–5 µm, 0–1-septate, brown, paler towards the apex, smooth,
thick-walled. Conidiogenous cells integrated, terminal or intercalary, with numerous,
small conidiogenous loci, crowded at the apex, proliferation sympodial, loci denticulate, 1 µm wide, unthickened, somewhat darkened–refractive. Conidia solitary, fusiform or limoniform, 8–14 × 5–7 µm, aseptate, yellowish to pale olivaceous, smooth to
minutely verruculose, attenuated towards apex and base, apex pointed, base truncate,
hila 1 µm wide, unthickened, somewhat darkened–refractive.
Hosts and Distribution: on Vitex spp. (Verbenaceae), Africa – Vitex cienkowskii
(Uganda).
10.2.50.
Fusicladium-state of Acantharia echinata (Ellis & Everh.) Theiss. & Syd.
Fig. 54
Teleomorph: Acantharia echinata (Ellis & Everh.) Theiss. & Syd., Ann. Mycol. 16: 15 (1918).
Ill.: SIVANESAN (1984b: 508–510, Figs 1–3).
On leaves, mycelium superficial, setose, composed of olivaceous-brown, densely
branched, septate hyphae up to 8.5 µm thick, sometimes forming a multicellular compact mass of hyphopodium-like structures functioning as anchoring organs on the
leaf. Stromata subcuticular, composed of thick-walled, dark brown cells, eventually
bursting through the cuticle. Conidiophores and setae arise from the stromata as well
as from the base and outer wall of the ascostromata. Setae simple, 70–210 × 6.5–9
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
105
µm, septate, dark brown, smooth to rough-walled, thick-walled. Conidiophores simple, occasionally branched, 16–46 × 5.5–5.8 µm, septate, brown, paler towards the
apex, smooth to rough-walled. Conidiogenous cells integrated, terminal, holoblastic,
proliferation sympodial, cicatrised. Conidia solitary, dry, blastic, broadly fusiform to
broadly ellipsoid, 15–23 × 8–10 µm, 0–2-septate, commonly with one septum, brown,
smooth to rugulose, tapering towards the apex, truncate at the base.
Hosts and Distribution: on Quercus spp. (Fagaceae), North America – Quercus
chrysolepis (USA, CA), Q. vacciniifolia (USA, CA).
Notes: SIVANESAN (1984b) described the anamorph of Acantharia echinata as Fusicladium sp. Material of this fungus has not been examined, and the descriptions and illustration are based on
SIVANESAN (1984b). It is not quite clear if the position of this anamorph in Fusicladium is correct.
The setae formed on the host are very unusual in the latter genus. Additional material should be
examined, and molecular data would also help any re-assessment of this fungus.
10.2.51.
Fusicladium-state of Apiosporina collinsii (Schwein.) Höhn.
Fig. 55
Teleomorph: Apiosporina collinsii (Schwein.) Höhn., Sitzungsber. Kaiserl. Akad. Wiss., Math.Naturwiss. Cl., Abt. 1, 119: 439 (1910).
Lit.: BARR (1968: 855), SIVANESAN (1984a: 598).
Ill.: BARR (1968: 854, Fig. 94), SIVANESAN (1984a: 599, Fig. 360).
Exs.: Barthol., F. Columb. 2927, 2320, 5013; Brenckle, F. Dakot. 505; Ellis, N. Am. F. 488a (as
Sphaeria collinsii); Ellis & Everh., F. Columb. 1431; Griffiths, West Am. F. 177; Kellerman, Ohio
Fungi 182; Rabenh., F. eur. 3541; Seym. & Earle, Econ. F. 125a, 125b; Thüm., Mycoth. univ. 849 (as
Dimerosporium collinsii).
On leaves the teleomorph causes crustaceous layers, mainly on the lower surface,
anamorph only formed on the teleomorph, caespitose, brown. Conidiophores arising
from superficial hyphae of the subiculum, hyphae branched, 3–5 µm wide, pale
olivaceous. Conidiophores erect, straight to flexuous, cylindrical, unbranched or sometimes branched, 30–225 × 4 µm, septate, somewhat constricted at the septa, medium
to dark brown, verruculose, walls only slightly thickened. Conidiogenous cells integrated, terminal or intercalary, with several conidiogenous loci, proliferation
sympodial, loci denticulate, apex truncate, 1–2 µm wide, unthickened, slightly darkened, refractive. Conidia in short, unbranched or branched chains, ellipsoid, ovoid to
subcircular, straight, 8–18 × 5–9 µm, aseptate, olivaceous to pale brown, verruculose,
walls somewhat thickened, often with 2–3 denticles at the distal end, truncate at the
base, hila 1–2 µm wide, unthickened, slightly darkened–refractive.
Hosts and Distribution: on Amelanchier spp. (Rosaceae), North America –
Amelanchier alnifolia (Canada, Alta., BC., Man., NWT, Ont., Sask.; USA, ID, ND,
NV, WY), A. canadensis (Canada, NB., NS., Ont.; USA, CT, MA, WI), A. pallida
(USA, OR), A. pumila (USA, NM), A. utahensis (North America), Amelanchier spp.
(Canada, Alta., Man., NB., NS., Nfld., Ont., Sask.; USA, ID, MA, VA).
Material examined: collections from M.
Fig. 54: Fusicladium-state of Acantharia
echinata (from SIVANESAN 1984b).
Notes: This anamorph was usually referred to Cladosporium sp., but on account of the structure of
the conidiogenous loci and conidial hila and its connection to Apiosporina collinsii (Venturiaceae),
it has to be placed in Fusicladium. Since this anamorph is strictly confined to and connected with the
teleomorph, it is not necessary to propose a separate, formal name for it.
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Schlechtendalia 9 (2003)
Fig. 55: Fusicladium-state of Apiosporina collinsii. A – conidia, B – conidiophores, C – base of
conidiophore arising from superficial hyphae, scale = 10 µm, K. Schubert del.
10.2.52.
Fusicladium-state of Apiosporina morbosa (Schwein.) Arx
Fig. 56
Teleomorph: Apiosporina morbosa (Schwein.) Arx, Acta Bot. Neerl. 3: 86 (1954).
Lit.: BARR (1968: 855–856), ELLIS (1976: 238–239), Fungi Canadenses (No. 84), SIVANESAN (1984a:
599–600).
Ill.: ARX (1954: 86, Figs 17, 95), ELLIS (1976: 239, Fig. 179), Fungi Canadenses (No. 84, Fig. 4),
SIVANESAN (1984a: 600, Fig. 361).
Exs.: Barthol., F. Columb. 4336; Brenckle, F. Dakot. 97, 410; Syd., F. exot. exs. 515.
Stromata on twigs, erumpent, variable in shape and size, at first olivaceous-green,
later blackish and firm, consisting of fungal hyphae on the surface and a mixture of
hyphae and host cells inside, sometimes anamorph abundant, forming brown covers
on the blackish surface of the teleomorph, colonies effuse to caespitose, brown, stromatic cells 4–12 µm diam., pale to medium brown, thick-walled. Conidiophores arising from the upper cells of stroma, in loose to dense fascicles, erect, flexuous, geniculate
or apex curved, unbranched or branched at the base, 20–95 × 3–6(–7) µm, septate,
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
107
Fig. 56: Fusicladium-state of Apiosporina morbosa. A – conidia, B – conidiophores, scale = 10 µm,
K. Schubert del.
pale olivaceous to pale brown, paler towards the apex, smooth, thick-walled, somewhat swollen at the base. Conidiogenous cells integrated, terminal or intercalary,
with several conidiogenous loci, proliferation sympodial, loci denticulate, 1–1.5 µm
wide, unthickened, somewhat darkened–refractive. Conidia solitary or rarely in short
chains, often laterally fused in pairs, ovoid, obovoid, ellipsoid or irregular, 4–19 × 3–
6 µm, 0–1-septate, pale olivaceous, smooth, thick-walled, hilum 1–1.5 µm wide,
unthickened, slightly darkened–refractive.
Hosts and Distribution: on Prunus spp. (Rosaceae), North America – Prunus
americana (USA, AL, FL, MS, NC, ND, OK), P. angustifolia (USA, FL, GA), P.
armeniaca (Canada, BC.; USA, CO, FL, IA, NY), P. avium (Canada, BC., Nfld.;
USA, NC, NE, TX), P. besseyi (USA, ND), P. cerasus (Canada, BC., NB., PEI, Que.;
USA, GA, NC and the eastern states), P. domestica (Canada, BC., NB., NS., Nfld.,
Ont., PEI, Que.; USA, FL, KY, MI, MS, NC, OH), P. dulcis (Canada, BC.), P.
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Schlechtendalia 9 (2003)
emarginata (USA, ID, MT, OR, WA), P. maritima (USA, AL, MA, NY, OR), P.
munsoniana (USA, AL, FL, GA, MS), P. nigra (Canada, NS., Ont.), P. padus (Canada,
Alta., Man., NWT), P. pensylvanica (Canada, Alta., Man., NB., NS., Nfld., NWT,
Ont., PEI, Que., Sask.; USA, NC), P. persica (Canada, NS.), P. pumila (USA, MI,
ND, WI), P. serotina (Canada, NB., NS., Ont., PEI, Que.; USA, GA, MS, NC, VA,
WI), P. spinosa (Canada, BC.), P. subcordata (Canada, BC.; USA, OR), P. umbellata
(USA, GA), P. virginiana (Canada, Alta., BC., Man., NB., NS., Nfld., NWT, Ont.,
PEI, Que., Sask.; USA, CA, CO, ID, MS, MT, NC, SC, SD, WA, WI), Prunus spp.
(Canada, Alta., BC., Man., NB., NS., Nfld., NWT, Ont., PEI, Que., Sask.; USA, GA,
MA, MD, NC, ND, OK, SD, VT, WI).
Material examined: all collections from IMI.
Notes: This anamorph is always associated with the teleomorph and does not occur separately. Therefore, we decline to propose a formal, separate name for it.
10.2.53.
Fusicladium sp. (1)
Fig. 57
Teleomorph: Unknown.
Lit.: ONDÌEJ (1973: 239).
Ill.: ONDÌEJ (1973: 238, Fig. 7).
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
10.2.54.
109
Fusicladium sp. (2)
Teleomorph: Venturia chlorospora (Ces.) P. Karst., Mycol. fenn. 2: 189 (1873).
Lit.: SIVANESAN (1977: 55).
On leaves. Conidiophores arising as lateral branches of brown hyphae. Conidiogenous
cells terminal or intercalary, with numerous conidiogenous loci, proliferation
sympodial. Conidia in unbranched chains, ellipsoid to cylindrical, 20–25 × 5–8 µm,
mostly 1-septate, truncate at the base.
Hosts and Distribution: on Salix spp. (Salicaceae), Europe – Salix acutifolia (RUS,
SF), S. alba (D, I), S. caprea (D), S. cinerea (D), S. fragilis (PL), S. triandra (I), S.
viminalis (D), Salix spp. (D).
Notes: This anamorph, which has not yet been observed in nature, was found by NÜESCH (1960) in
cultures isolated from ascospores of Venturia. Based on conidia formed in chains, SIVANESAN (1977)
considered it to be a species of Cladosporium.
11.
Doubtful and unclear species of Fusicladium s.lat.
Type material of the following species could not be traced or was not available and other collections have
not yet been found, so that the generic affinity and taxonomic status of these taxa could not be proven.
Leaf spots on fading or mainly necrotic leaves, irregular, brown, often confluent.
Mycelium internal, subepidermal. Conidiophores solitary or in small groups of up to
6, arising from stromata, erumpent through the cuticle, straight, simple or rarely
branched, 40–95(–130) × 3–5 µm, septate, dark coloured. Conidiogenous cells integrated, terminal, with a single or up to three conidiogenous loci, proliferation
sympodial. Conidia catenate, in unbranched or branched chains, cylindrical or fusiform, (10–)12–20 × 3–4(–5) µm, 0–1(–2)-septate, dark, pigmented.
Hosts and Distribution: on Salix spp. (Salicaceae), North America – Salix purpurea
(USA, NY).
Notes: ONDÌEJ (1973) described this fungus based on material from North America (on Salix purpurea,
USA, NY, 22 Jun. 1949), which was originally identified as Fusicladium saliciperdum. Since this
material could not be traced, it was impossible to re-assess this fungus. The anamorph of Venturia
chlorospora [Fusicladium sp., NÜESCH (1960) and SIVANESAN (1977)] is probably allied, but distinguished by having broader conidia.
Fusicladium aconiti Bres., Ann. Mycol. 18: 58 (1920).
Type: on leaves of Aconitum clusii (Ranunculaceae), Hungary, Tatra, Greschik.
Original description (SACCARDO 1931: 802): Caespitulis dense gregariis, in macula fusca, epiphylla
nidulantibus, hyphis unicellularibus, cylindraceis, olivaceis, 45–50 × 7–9 µm; conidiis olivaceis, 1septatis, subclavatis, basi truncata, ad septa subconstrictis, 40–48 × 8–10 µm.
Fusicladium aplectri Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 1894: 378
(1894).
Type: on leaves of Aplectrum hyemale (Orchidaceae), USA, Delaware, Commons.
Original description (ELLIS & EVERHART l.c.): Spores irregular, whitish, with a shaded, purple border, 1cm diam. Hyphae fasciculate, olivaceous, simple, 2–3-septate, 65–75 × 5–6 µm, mostly twisted
or abruptly bent at the tip, olive-brown. Conidia terminal, elliptical, greenish, granular, continuous
at first, becoming 1-septate, 12–15 × 6–7 µm.
Fusicladium butyrospermi Griffon & Maubl., Bull. Soc. Mycol. France 29: 249
(1913).
Type: on leaves of Butyrospermum parkii (Sapotaceae), Africa, Koulikoro, Vuillet.
Original description (S ACCARDO 1931: 803): Maculis rotundatis vel angulosis, 2–3 mm diam.
amphigenis, superne brunneis inferne fulvis, margine atropurpureo cinctis; caespitulis obscure
brunneis, amphigenis, minutis, in centro macularum dense gregariis; hyphis fertilibus, caespitosis,
simplicibus, cylindraceis apice rotundatis vel paulum attenuatis, continuis vel uniseptatis, fuligineis,
15–20 × 4–5 µm; conidiis acrogenis, ovoideis, medio septatis et subinde constrictulis, fuligineis, 10
× 6 µm; mycelio in epidermide evoluto, cellulis globosis vel ellipticis formato.
Fusicladium carpini Osipyan, Mikol. Fitopatol. 5(1): 88 (1971).
Fig. 57: Fusicladium sp. (from
ONDÌEJ 1973).
Type: on leaves of Carpinus caucasica (Corylaceae), Caucasus, Armenia, Goris-Kafan, 30 Jul. 1954,
A. Grossh. (ERCB).
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Schlechtendalia 9 (2003)
Description (OSIPJAN 1971): Maculae amphigenae, irregulares, 2–6 mm in diam., initio flavae, deinde
fuscescentes, saepe centro aquilae, diffusae, interdum nervis, subtus haud raro nigrescentibus limitatae.
Pruina conidialis vix evoluta. Conidiophora in fasciculos compactos vix prominentes pulviniformes congesta,
levia, pallide olivacea, irregulariter cylindrica, vix clavata, erecta, eseptata, non ramosa, 10–17 × 3.5–7.2
µm. Conidia catenulata, pallide olivacea, fusiformia, vix obclavata, interdum unilateralia, medio vix angustata,
unicellularia vel uniseptata, ad septum subconstricta, extremitatibus late obtusatis, 18.1–26.4 × 4.9–6 µm.
Fusicladium cephalanthi Speg., Anales Mus. Nac. Buenos Aires 6: 339 (1899).
Type: on twigs, leaves and buds of Cephalantus sarandi, Uruguay, Montevideo; Argentina, pr. Quilmes
et La Plata.
Original description (SACCARDO 1902: 1056): Rami-foliicolum, late denseque effusum, olivaceum;
caespitulis superficialibus, densissime gregariis, ex hyphis paucis, 5–12 µm, efformatis; hyphis e
basi sclerotiacea parenchymatico-olivacea matrici innata orientibus, chlorinis, subteretibus, 40–50 ×
3–4 µm, apice vix attenuatis, 1–5-septatis, non constrictis; conidiis acrogenis, cylindraceis, non v.
vix clavulatis, magnitudine ludentibus, 10–30 × 3–5 µm, chlorinis, 1-septatis, non constrictis.
Fusicladium chanousii Ferraris, Malpighia 16: 474 (1902).
Type: on anthers of Gentiana lutea, Italy, Picollo S. Bernardo.
Original description (FERRARIS l.c.): Caespitulis, minutis, velutinis, effusis, olivaceis, hyphis fuscis,
erectis, simplicibus, non vel 1-septatis, apice subdenticulatis, 35–50 × 4.5–6 µm; conidiis olivaceis,
ellipticis, continuis, dein. 1-septatis non constrictis, 14–19 × 6–7 µm.
Fusicladium ephedrae Cruchet, Bull. Soc. Vaud. Sci. Nat. 55: 157 (1925).
Type: on Ephedra sp., Switzerland, Montagny provenant de Sion (Valais), D. Cruchet.
Teleomorph: Venturia ephedrae Cruchet, l.c.
Ill.: CRUCHET (1925: 158, Fig. 2/1).
Original description (CRUCHET l.c.): Spores fusiformes, peu arquées, arrondies atténuées deux extrémités,
unibiseptées, légèrement resserrées à la choison, très nombreuses, brunes, se détachant fascilement de
basides fasciculées, cylindriques, ténues, peu apparantes, presquet hyalines, entourant les périthèces.
Fusicladium fici Bacc., Ann. Bot. (Rome) 4: 277 (1906).
Type: on leaves of Ficus sp., Eritrea, Valle Catalaben, Mensa 1900, Pappi.
Original description (BACCARINI 1906): Maculis orbicularibus amphigenis, supra pallidis et rubro-cinctis,
subtus rufo-ferrugineis; caespitulis amphigenis punctiformibus; hyphis fertilibus simplicibus, continuis,
brevibus, fuscidulis, ad apicem pallidioribus; conidiis piriformibus, fuscidulis, septatis, 20 × 10 µm.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
111
4 µm, cellulae conidiogenae sympodiales, ad apicem conico–truncatae, pallido-brunneae, cicatricosa
conspicua; conidia singularia, raro catenulata, longiovoidea, pallido-brunnea, 1-septata, utrinque
attenuata, ad basin obconico–truncata, 7–8 × 3–4 µm.
Fusicladium hippophaës Vasyag. & Byzova, in Shvartsman et al., Flora Sporovych
Rastenij Kazachstana 8(2): 142 (1975).
Type: on leaves and twigs of Hippophaë rhamnoides, Russia, Kazakhstan, region Alma-Ata, in promontoriis
Ala-Tau Transiliensis, prope pagum Saty, 23 Jul. 1956, B.K. Kalymbetov (AA).
Ill.: SHVARTSMAN et al. (l.c., Fig. 63).
Original description (SHVARTSMAN et al. l.c.): Conidiophora 0–3-septata, interdum subconstricta, cylindrica,
recta, rarius, incurvata, apice late rotundata vel obtusata, edenticulata, indivisa, 16.8–73.5 × 5.2–11.5 µm,
brunneo-fusca, pellucida, solitaria, raro fasiculata. Conidia acrogena, transverse 1–3-septata, constricta,
cylindrica, obclavata, fusiformia, obpyriformia, recta, interdum incurvata, 22.5–90.3 × 3.4–14.7 µm,
brunneo-fusca, pellucida, solitaria vel catenulata. Caespituli orbiculares, irregulares, angulati, haud dense
velutini vel pulveracei, epiphylli pro more secus nervos dispositi, in ramulis convexi, compacti, atrobrunnei vel nigrescenti-olivacei, saepe confluentes. Maculae epiphyllae, plerumque subinconspicuae.
Fusicladium lini Sorauer, Z. Pflanzenkrankh. 5: 104 (1895).
Type: on leaves and stems of Linum usitatissimum, Belgium, Ardoye, Nijpels, 1894.
Lit.: LINDAU (1907: 784), VASSILJEVSKY & KARAKULIN (1937: 207).
Original description (translation based on SORAUER’s, l.c., and LINDAU´s (1907) descriptions in
German): Leaf spots oval to oblong, brown. Caespituli on the leaf spots, 0.75–1 mm long, almost
black. Conidiophores fasciculate, dense, geniculate–sinuous, more or less greenish brown at the
base, hyaline towards the apex, ca. 30 µm long, 3 µm wide. Conidia terminal, solitary, ovoid to
oblong, almost hyaline, 8 mm long, 4 µm wide, also with some longer conidia, 14–18 µm long.
Fusicladium lonicerae Vasyag., Izv. Akad. Nauk Kazahsk. SSR, Ser. Biol. 1(13):
101 (1957).
Holotype: on Lonicera tatarica, Kazakhstan, Karaganda, botanical garden, 4 Aug. 1951, M.P.
Vasyagina (AA).
Original description (VASYAGINA l.c.): Maculis rotundatis, albo-fuscis, solitariis, confluentibus.
Conidiophoris fasciculatis, hypophyllis, rectis vel curvatis, fuscis, apice hyalini, numerosis
denticulatis, 30–44 × 3 µm. Conidiis ovatis vel piroideis, uniseptatis vel unicellularibus, rarius 2
septatis, albo-fuscis, 12.8–19 × 5–6.5 µm.
Fusicladium stuckertii (Speg.) M.B. Ellis, in herb. (IMI).
Fusicladium gardeniae F.X. Chao & P.K. Chi, in Chi, Fungal Diseases of Cultivated
Medicinal Plants in Guangdong Province: 171 (1994).
≡ Napicladium stuckertii Speg., Anales Mus. Nac. Buenos Aires 8: 87 (1902).
≡ Sporhelminthium stuckertii (Speg.) Speg., Physis (Buenos Aires) 4(17): 292 (1918).
Type: on living cladodes of Baccachis trimera, Argentina, Córdoba.
Original description (SACCARDO 1906): Superficiale, late effusum, olivaceum; hyphis repentibus matrici
arctiuscule adnatis, araneosulis, dense ramuloso-intricatis atque anastomosantibus, torulosis, articulis a globoso
subcuboideis 5–10 µ diam., olivaceis; hyphis fertilibus hinc indeerectis, gracilibus, simplicinus, 3–5-septatis,
50–200 × 5–6, olivascentibus, apicem versus pallidioribus, parce septulatis, superne noduloso–geniculatis;
conidiis cylindraceis v. subfusoideis, continuis v. 1–3-septatis, levibus, 15–30 ×5–6, acro-pleurogenis, chlorinis.
Type: on leaves of Gardenia jasminoides, China, Prov. Guangdong, Qujiang, Oct. 1988, leg. F.X.
Chao No. 024.
Ill.: CHI (l.c.: Fig. 178).
Original description (CHI l.c.): Maculae orbiculares, flavo-brunneae, margine distinctae, 0.4–0.8
mm diam., annulatae, leviter depressae. Caespituli epiphylli, nigri; stromata subcuticulares, 30–220
µm diam.; conidiophora spanofasciculata, simplicia, sursum 0–1 geniculata, 1–4 septata, 13–24 × 3–
Type: on stems of Phaseolus vulgaris, Germany, Westphalia.
Original description (SACCARDO 1886: 347): Caespitulis sparsis aut densis, ex cinereo-viridibus;
hyphis erectis, simplicibus, parce septatis, apice obtusis, viridibus et fasciculato–conjunctis; conidiis
oblongo–fusiformibus, dilute viridibus.
Fusicladium fraxini var. phillyreae Trotter, in Pamp., Nuovo Giorn. Bot. Ital. 31:
233 (1924).
Original material is missing. This variety is very probably not conspecific with Spilocaea phillyreae.
Fusicladium tenue Bonord., Abh. Mykol. 2: 93 (1870).
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Schlechtendalia 9 (2003)
Fusicladium theae Hara, Tea J. 14: 16 (1919); Mycologia 12: 330 (1920).
Type: on leaves of Camellia thea (= Thea sinensis), Japan.
Original description (SACCARDO 1931: 803): Acervulis amphigenis, vellutinis, nigris; conidiophoris
filiformibus rectis v. curvis basi incrassatis continuis v. 3-septatis infra brunnescentibus supra pallide
coloratis et incurvatulis, 40–70 × 4–5 µm; conidiis terminalibus, cylindraceis v. ovato oblongis sub
medio uniseptatis non v. parum constrictis, apice obtusis, basi subacutatis, rectis v. curvis, hyalinis v.
flavescentibus, 15–28 × 5–6 µm.
Fusicladium vanillae Zimm., Centralbl. Bakteriol., 2 Abth., 8: 480 (1902).
Type: on leaves of Vanilla sp., Java, Buitenzorg.
Original description (SACCARDO 1906: 580): Hyphis sterilibus foliorum superficiei adpressis, rarius
liberis, brunneolis; conidiophoris erectis, rectiusculis, apice acutis, continuis, brunneolis, 25–30 µm
longis; conidiis ovato–oblongis, basi acutiusculis, apice rotundatis, bicellularibus, 8 × 4 µm.
Spilocaea concentrica Schwein., Trans. Amer. Philos. Soc., Ser. 2, 4(2): 297 (1832).
Type: In cortice Peponum putridorum, Bethl. Optime aut evoluta mense Octobri prope Philadelphia
in talibus.
Notes: Listed by SACCARDO (1886: 761) under doubtful and excluded species.
Spilocaea epiphylla Fr., Syst. Mycol. 3: 504 (1832).
Type: Plures tales formationes in foliis Pyri, Mali etc. misit Levieux, omnes e Gallia occid.
Original description (Fries): Maculis epidermide bullata circumscissa secedente minutis sparsis nigris.
Epidermidis valde relaxatae bullae ½ unc. circiter latae, haud rumpuntur, sed in ambitu integrae in
squamam aeque latam integram solvuntur. Tum sub hac conspicitur macula interrupta s. plures maculae
sparsae nigrae, folio arctae adnatae, nec pulverulentae, quae non sine difficultate a matrice separatae
characteres datos monstrant.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
113
Fusicladium alopecuri Sawada, Rep. Gov. Res. Inst. Formosa 85: 93 (1943), nom. inval.
Fusicladium anethi Nevod., Griby rossii (Russian fungi) IV, No. 191 (1917).
Type: on leaves of Peucedanum graveolens (= Anethum graveolens), Georgia, Prov. Tiflis, Distr.
Gori, near Skra, in horto Pridonov, 23 Jul. 1912, G.S. Nevodovskij, Griby ross. 191 (e.g., B, IMI
1423, K, LE).
= Passalora punctum (Delacr.) S. Petzoldt, in Arx, Plant Pathogenic Fungi: 288 (1987).
Lit.: CROUS & BRAUN (2003: 343).
Fusicladium angelicae Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 1891: 87
(1891).
Types: on leaves of Archangelica atropurpurea (= Angelica atropurpurea), USA, Wisconsin, Racine,
Sept. 1890, J.J. Davis (NY; Ellis & Everh, N. Am. F. 2790, e.g. M, NY).
≡ Passalora angelicae (Ellis & Everh.) U. Braun, Nova Hedwigia 55(1–2): 214 (1992).
Lit.: CROUS & BRAUN (2003: 437).
Fusicladium aronici Sacc., Michelia 2: 171 (1880).
Numerous collections (HBG, JE, M) have been examined.
= Fusicladiella melaena (Fuckel) S. Hughes, Mycol. Pap. 49: 21 (1952).
Fusicladium ascyrinum Ellis & Everh., J. Mycol. 4: 53 (1888).
Holotype: on leaves of Ascyrum hypericoides, USA, Louisiana, Natchitoches, 26 Sept. 1886, A.B.
Langlois (NY).
This is a hyphomycete of unclear affinity.
Fusicladium bambusicola (Sawada) Deighton, in herb.
Spilocaea opuntiae Rabenh., Flora 33: 625 (1850).
Type: “Auf unreif abgefallenen Früchten der indischen Feige, auf Capri.“
Original description (RABENHORST l.c.): Maculis aureo-fuscescentibus saepius obliteratis rugosoconstatis, spor. subconglobatis ovoideis pallide roseis demum expallentibus ab epidermide tectis,
episporio crassiusculo ruguloso.
Spilocaea scirpi Link, in Willd., Sp. pl., vol. 6(2): 87 (1825).
Type: Habitat in caulibus Scirporum siccis in Europa, nec non Aegypto.
Original diagnosis (Link): Maculas format in Scirporum majorum caulibus frequentissimas aggregatas
interdum confluentes, ita ut caulis inde saepe quasi marmoratus appareat. Maculae istae planae sunt
supra caulem minime eminentes. Sporidia minutissima intra caulem recondita seriatim juncta. Sp.
acervis oblongis effusis, epidermide non rumpente, sporidiis minutis fuscis.
Notes: Two samples from M (Botanische Staatssammlung München), deposited under Spilocaea
scirpi, have been examined, but no traces of Spilocaea-like structures have been found.
12.
Excluded species
≡
≡
Cercosporidium bambusicola Sawada, Taiwan Agric. Res. Inst. Rep. 87: 77 (1944), as
‘bambusicolum’, nom. inval.
Pseudospiropes bambusicola Goh & W.H. Hsieh, in Hsieh & Goh, Cercospora and similar
fungi from Taiwan: 147 (1990).
Fusicladium bicolor C. Massal., Nuovo Giorn. Bot. Ital. 21: 170 (1889) and Atti
Accad. Agric. Art. Comm. Verona, 3. Ser., 65: 115 (1889).
Isotype: on leaves of Chaerophyllum cf. hirsutum, Italy, prov. Verona, Mt. Lobia, 31 Aug. 1887, C.
Massalongo (HBG).
≡ Fusicladiella bicolor (C. Massal.) U. Braun, Schlechtendalia 5: 38 (2000).
Fusicladium butleri Syd., Ann. Mycol. 14: 260 (1916).
Holotype: on Jasminum arborescens, India, U.P., Orai, Bandlekhand, 27 Feb. 1907, E.J. Butler, No.
1710 (S).
≡ Pseudocercospora butleri (Syd.) U. Braun, Schlechtendalia 5: 42 (2000).
Lit.: CROUS & BRAUN (2003: 91).
Fusicladium alopecuri Ellis & Everh., J. Mycol. 4: 53 (1888).
Fusicladium caricae Sacc., Atti Congr. Bot. Palermo 1902: 58 (1902).
Holotype: on leaves of Setaria polystachya (= Alopecurus geniculatus), USA, Columbia, Montana,
20 May 1887, B.T. Galloway (NY).
≡ Cladosporium alopecuri (Ellis & Everh.) U. Braun, Schlechtendalia 5: 32 (2000).
Type: on leaves of Carica papaya, Paraguay, Guarapi, Feb. 1881, B. Balansa 2739 (LPS). Other
authentic material (Balansa 3855) at B.
= Asperisporium caricae (Speg.) Maubl., Lavoura 16: 212 (1913).
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Schlechtendalia 9 (2003)
Fusicladium caruanianum Sacc., Ann. Mycol. 11: 20 (1913), basionym
Fusicladium cecropiae (Stev.) Toro, Sci. Surv. Porto Rico & Virgin Islands 8: 224 (1932).
Pseudocercospora cecropiae (Stev.) Deighton, Mycol. Pap. 140: 70 (1976).
Fusicladium chlorinum Ellis & Kellerm., in herb.
=
Ramularia asteris var. latispora U. Braun, A Monograph of Cercosporella, Ramularia and allied genera, Vol. 2: 98 (1998).
Fusicladium cynanchi Reichert, Bot. Jahrb. Syst. 56: 720 (1921).
Holotype: on leaves of Cynanchum acutum, Egypt, near Damiettam, 20 Mar. 1921, C. Ehrenberg (B).
= Cercospora punctiformis Sacc. & Roum., Rev. Mycol. (Toulouse) 3: 29 (1881).
Lit.: CROUS & BRAUN (2003: 343).
Fusicladium depressum (Berk. & Broome) Roum., Fungi gall. exs. 86 (1879).
≡ Passalora depressa (Berk. & Broome) Sacc., Nuovo Giorn. Bot. Ital. 8: 187 (1876).
Lit.: CROUS & BRAUN (2003: 157).
Fusicladium depressum f. petroselini Sacc., Rev. Mycol. (Toulouse) 19: 53 (1897).
= Passalora punctum (Delacr.) S. Petzoldt, in Arx, Plant Pathogenic Fungi: 222 (1987).
Lit.: CROUS & BRAUN (2003: 343).
115
Fig. 58
≡ Pseudocladosporium caruanianum (Sacc.) U. Braun comb. nov.
Holotype: on dead leaves of Magnolia grandiflora, Malta, Balzan, leg. Caruano Gatto (PAD).
Lit.: SACCARDO (1931: 801).
On dead leaves, saprobic, lesions lacking. Colonies hypophyllous, effuse, loose to dense, forming
small to large patches, confluent, velutinous, brown, sooty. Mycelium internal and external, superficial, hyphae creeping, often aggregated, forming ropes or aggregations of swollen hyphal
cells, sparingly branched, 2–6 µm wide, septate, often with constrictions and swellings, pale
olivaceous to medium dark brow or olivaceous-brown. Conidiophores solitary, arising from swollen hyphal cells or from creeping hyphae, lateral, occasionally terminal, erect, straight and
subcylindrical to geniculate–sinuous, unbranched, 10–30 × 2–4 µm, (0–)1–3(–4)-septate, pale to
medium dark brown, smooth, wall thin or slightly thickened. Conidiogenous cells integrated, terminal, 10–20 µm long, conidiogenous loci denticle-like, mostly with several denticles, tips truncate, wall unthickened, not darkened or occasionally slightly darkened–refractive. Conidia catenate, often in branched chains, ellipsoid–ovoid, fusiform, subcylindrical, 5–18 × 2–4 µm,
0–1-septate, pale olivaceous or olivaceous-brown, yellowish brown, smooth to faintly rough-walled,
apex of primary conidia obtuse, rounded, hila (1–3, rarely up to 5) short obconically truncate,
unthickened, not darkened.
Notes: This species fits well into the concept of Pseudocladosporium U. Braun (BRAUN 1998). It
is distinguished from Pseudocladosporium hachijoense (Matsush.) U. Braun by having well-developed, longer, septate conidiophores and usually aseptate conidia. Pseudocladosporium
brevicatenatum (U. Braun & Feiler) U. Braun differs from P. caruanianum in having (0–)1–2septate conidia. The long, septate conidiophores of the latter species resemble those of
Pseudocladosporium sp. (BRAUN 1998), described from Japan, but the conidia in the latter fungus
are septate.
Together with Anungitea B. Sutton, Fusicladium, Polyscytalum Riess and similar genera,
Pseudocladosporium belongs to a group of acroblastosporic hyphomycetes (Acroblastosporae, KIFFER
& MORELET 1999). The taxonomy within this assemblage of genera is unsettled and uncertain. Monographic studies and molecular data are necessary to get a system of well-defined natural genera.
≡
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
A
B
C
Fig. 58: Pseudocladosporium caruanianum. A – conidia, B – conidiophores, C – mycelium, scale =
10 µm, U. Braun del.
Fusicladium depressum var. platysporum (Ellis & Holw.) Davis, Paras. fungi
Wisconsin: 113 (1942).
≡ Passalora platyspora (Ellis & Holw.) U. Braun, in Braun & Rogerson, Sydowia 47: 145 (1995).
Lit.: CROUS & BRAUN (2003: 328).
Fusicladium depressum var. sii (Ellis & Everh.) Davis, Paras. fungi Wisconsin: 113
(1942).
≡ Passalora sii (Ellis & Everh.) U. Braun, Nova Hedwigia 55: 214 (1992).
Lit.: CROUS & BRAUN (2003: 377).
Fusicladium depressum var. tommasiniae C. Massal., Atti Reale Ist. Veneto Sci. Lett.
Arti 59(2): 685 (1900).
=
Passalora sp., status unclear.
Fusicladium destruens Peck, Rep. (Annual) New York State Mus. Nat. Hist. 43: 30
(1890).
Material examined: on leaves of Avena sativa, Canada, London, Jul. 1907, J. Dearness, Barthol., F.
Columb. 2431 (HBG).
= Cladosporium macrocarpum Preuss, in Sturm, Deutschl. Fl. 3(6): 27–28 (1848).
116
Schlechtendalia 9 (2003)
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
117
Fusicladium dubiosum Speg., Anales Soc. Ci. Argent. 22: 211 (1886).
Fusicladium livistoniae P. Karst., Hedwigia 30: 302 (1891).
Syntypes: on leaves of Digitaria sp., Pl. du Paraguay 3517, Guarapi, Dez. 1882, Balansa (B, PC).
= Pyricularia grisea Sacc., Michelia 2: 20 (1880).
Holotype: on dead petioles of Livistona chinensis, Russia, Karelia, Vyborg, Liimatta, Sept. 1891, A.
Thesleff (H 4252).
Notes: This species does not belong in Fusicladium, status unclear (BRAUN 2000).
Fusicladium elasticae Koord., Bot. Unters. Java: 231 (1907).
The type at B consists of a drawing and some notes, but without any material. Based on the drawing,
this species can be excluded from Fusicladium, but its status and generic affinity are unclear.
Fusicladium euonymi-japonici Hori (TAI 1979), nom. nud.
Fusicladium fagopyri Oudem., Verh. Kon. Ned. Akad. Wetensch., Aft. Natuurk.,
Tweede Sect., 1897: 88 (1897).
Type: on leaves of Fagopyrum esculentum, Netherlands, Amsterdam, Jun. 1897, Oudemans (L
66363).
The type consists of various saprobic hyphomycetes, incl. Alternaria and Cladosporium. Some conidia agreeing with the original description of this species belong to Cladosporium herbarum s.lat.
Fusicladium fici Achundov, Novosti Sist. Nizsh. Rast. 16: 32 (1979), nom. illeg.,
homonym.
Fusicladium fuliginosum Kalchbr. & Cooke, Grevillea 1884: 24 (1884).
Syntype: on leaves of an unknown host plant, Africa, Port. Natal, Luanda, J.M. Wood (B).
Host plant unknown. This is a hyphomycete of unknown affinity.
Fusicladium gnaphaliatum Bonar, Mycologia 57: 392 (1965).
Fusicladium macrosporium Bonord., Hedwigia 3(5): 74 (1864).
= Monotospora ovata Sacc., Syll. fung. 4: 299 (1886).
Fusicladium maculicola (Ellis & Kellerm.) OndÍej, „eská Mykol. 27(4): 237 (1973).
Lectotype: on leaves of Phragmites australis, USA, Manhattan, Kansas, Jun. 1887, W.A. Kellerman
No. 934 (NY).
≡ Scolecotrichum maculicola Ellis & Kellerm., J. Mycol. 3: 103 (1887), as ‘maculicolum’.
≡ Passalora maculicola (Ellis & Kellerm.) U. Braun, Schlechtendalia 5: 39 (2000).
Fusicladium minutulum Sacc., Nuovo Giorn. Bot. Ital. 27: 85 (1920).
Isotype: on leaves of Vitis californica, USA, Sants Pars, Chryon., Sept. 1916, J.R. Weir (M).
≡ Asperisporium minutulum (Sacc.) Deighton, in Ellis, More Dematiaceous Hyphomycetes: 243
(1976).
Fusicladium peucedani Syd. & P. Syd., Ann. Mycol. 5: 340 (1907), nom. illeg., homonym of F. peucedani Ellis & Holw., 1895.
Types: on leaves of Peucedanum decursivum, Japan, Tokio, N. Nambu (B, S).
= Passalora depressa (Berk. & Broome) Sacc., Nuovo Giorn. Bot. Ital. 8: 187 (1876).
Lit.: CROUS & BRAUN (2003: 157).
Holotype: on leaves of Gnaphaliatum stramineum (= G. chilense), USA, California, Lake Merced,
San Francisco Co., 9 Sept. 1934, L. Bonar (UC 532308); paratypes: on G. stramineum, USA, California, San Francisco, Golden Gate Park, 16 Oct. 1925, L. Bonar (UC 257214); USA, California,
Humboldt Co., Hors-Linto Creek, 4 Sept. 1938, L. Bonar (UC 640690).
≡ Asperisporium gnaphaliatum (Bonar) U. Braun, Schlechtendalia 5: 32 (2000).
Fusicladium pongamiae Syd., Ann. Mycol. 11: 328 (1913).
Fusicladium hariotianum Sacc., Ann. Mycol. 6: 560 (1908).
Holotype: on Phellinus ferrea, on trunks of Alnus sp., USA, California, Humboldt Co., Van Duzen
River, 31 Mar. 1931, H.E. Parks 2726 (UC 1272179); isotype: California Fungi 1251 (PC, UC 568840).
≡ Porophilomyces poricola (Bonar) U. Braun, Schlechtendalia 5: 42 (2000).
Holoype: on leaves of Atropis borreri (= Glyceria borreri), France, Paris, Eatihou, Sept. 1907, leg. P.
Hariot (PC).
Original description (SACCARDO 1913: 1376): Caespitulis punctiformibus, nigricantibus, dense seriatis,
epiphyllis, superficialibus, 125–130 µm diam.; hyphis fertilibus dense fasciculatis, paliformibus,
simplicibus, continuis, atro-olivaceis, 70 × 5.5–6 µm, apice obtusulis; conidiis tereti–oblongis, basi
truncatis, apice rotundatis, 1-septatis, non constrictis, 30 × 8 µm, olivaceis.
= Cladosporium sp.
Notes: Long, brown, septate conidiophores with somewhat thickened walls and large solitary conidia, 16–30 × 8–11 µm, 0–3-sepate, olivaceous-brown, with somewhat thickened, echinulate walls.
This is a species of Cladosporium morphologically close to C. phlei (C.T. Greg.) G.A. de Vries.
≡
Asperisporium pongamiae (Syd.) Deighton, in Ellis, More Dematiaceous Hyphomycetes: 241
(1976).
Fusicladium poricola Bonar, Mycologia 57: 393 (1965).
Fusicladium praecox Niessl, in Rabenh., Fungi eur., Ed. Nov., Ser. II, No. 1166 (1868)
and Hedwigia 7: 124 (1868).
Syntypes: on leaves of Tragopogon orientalis, Czech Republic, pr. Bistenz ad Brunnam Moraviae,
May, G. de Niessl, Rabenh., F. eur. 1166 (e.g., B, HBG, HAL, LE, M).
≡ Cladosporium praecox (Niessl) U. Braun, Schlechtendalia 5: 34 (2000).
Fusicladium heterosporum Höhn., Ann. Mycol. 3: 337 (1905).
Fusicladium punctiforme G. Winter, in Rabenh., Fungi eur., Fasc. 16 (32), No. 3582
(1886), non Passalora punctiformis G.H. Otth, 1868.
Types: on leaves of Epilobium parviflorum, Austria, Wiener Wald, near Winten, 7 Jun. 1905, v.
Höhnel (B) and Kab. & Bub., F. imp. exs. 293 (e.g., BPI 424286, FH, K.)
≡ Passalora heterospora (Höhn.) Höhn., Centralbl. Bakteriol., 2. Abth., 60: 1 (1923).
Lit.: CROUS & BRAUN (2003: 455).
Syntypes: on leaves of Pimpinella integerrima (= Zizia integerrima), USA, Missouri, near Perryville,
Aestate 1885, C.H. Demetrio, Rabenh., F. eur. 3582 (e.g., HAL, M).
= Passalora platyspora (Ellis et Holw.) U. Braun, in Braun & Rogerson, Sydowia 47: 145 (1995).
Lit.: CROUS & BRAUN (2003: 328).
118
Schlechtendalia 9 (2003)
Fusicladium rhamni Fuckel, in herb.
Fusicladium robiniae Shear, Bull. Torrey Bot. Club. 29: 452 (1902).
Syntype: on leaves of Robinia pseudacacia, USA, Maryland, Glen Sligo, 3 May 1899, C.L. Shear,
Barthol., F. Columb. 1619 (HBG).
≡ Passalora robiniae (Shear) S. Hughes, Canad. J. Bot. 31: 572 (1953).
Lit.: CROUS & BRAUN (2003: 468).
Fusicladium ruthenicum Petr., Ann. Mycol. 19: 78 (1921).
Syntypes: Petr., F. polon. 638 (e.g., K, W).
= Passalora galii (Ellis et Holw.) Arx, Proc. Kon. Ned. Akad. Wetensch. C 86(1): 45 (1983).
Lit.: CROUS & BRAUN (2003: 193).
Fusicladium schnablianum Allesch., Fungi bavar. exs. 397 (1894).
Lectotype: on leaves of Carduus personata, Germany, Bavaria, Oberammergau, 1894, Allescher
(M), selected here; isolectotype: on Carduus personata, Germany, Bavaria, Oberammergau, Aug.
1894, Allescher, Allesch. & Schn., F. bavar. 397 (B).
= Fusicladiella melaena (Fuckel) S. Hughes, Mycol. Pap. 49: 21 (1952).
Fusicladium sorghi Pass., Hedwigia 16: 122 (1877).
Isotype: on leaves of Sorghum halepense, Italy, Pavia, Estate 1893, Briosi & Cavara, F. paras. 240
(HAL).
= Hadrotrichum sorghi Ferraris & C. Massal., Ann. Mycol. 10: 297 (1912).
Fusicladium statices Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 1894: 378
(1894).
Holotype: on leaves of Limonium vulgare (= Statice limonium), USA, New Jersey, Cape May, 13
Sept. 1894, A. Commons (NY).
= Cladosporium cf. herbarum (Pers.: Fr.) Link.
Fusicladium transversum Sacc., Ann. Mycol. 3: 170 (1905).
Syntype: on dead leaves of Ophiopogon japonicum, Italy, Padova, botanical garden, Feb. 1905, P.A.
Saccardo, Mycoth. ital. 1738 (M).
= Cladosporium sp.
Spilocaea proteae (Marasas et al.) Arx, Genera of Fungi Sporulating in Pure Culture,
ed. 3: 280 (1981).
≡
13.
Batcheloromyces proteae Marasas et al., J. S. African Bot. 41(1): 43 (1975).
References
Beside references cited in this work, additional papers, checklists and monographs are included that
have been used as sources of data on host range and distribution.
ACHUNDOV, T.M. 1979: Mikoflora Nakhichevanskoj ASSR. ‘Elm’ Publishing House, Baku.
ADERHOLD, R. 1896: Die Fusicladien unserer Obstbäume. I. Teil. Landwirthschaftliche Jahrbücher
25: 875–914.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
119
ADERHOLD , R. 1897: Revision der Species Venturia chlorospora, inaequalis und ditricha autorum.
Hedwigia 36: 67–83.
ADERHOLD , R. 1900: Die Fusicladien unserer Obstbäume. II. Teil. Landwirthschaftliche Jahrbücher
29: 541–588.
ADERHOLD , R. 1902: Über Venturia crataegi n. spec. Berichte der Deutschen Botanischen Gesellschaft 1902: 195–200.
Anonymous 1979: List of Plant Diseases in Taiwan. The Plant Protection Society, Republic of China.
ARX, J.A. von 1952: Studies on Venturia and related genera. Tijdschrift over Plantenziekten 58:
260–266.
ARX, J.A. von 1954: Revision einiger Gattungen der Ascomyceten. Acta Botanica Neerlandica 3: 83–84.
ARX, J.A. von 1957a: Schurft op Pyracantha. Tijdschrift over Plantenziekten 63: 198–199.
ARX, J.A. von 1957b: Ueber Fusicladium saliciperdum (Allesch. et Tubeuf) Lind. Tijdschrift over
Plantenziekten 63: 232–236.
ARX, J.A. von 1987: Plant Pathogenic Fungi. Beihefte zur Nova Hedwigia 87: 1–288.
ATANASOV, D. & PETROV, D. 1930: Spis’k na konstatiranite v B’lgarija prichiniteli na bolesti po
rastenijata. Sofja.
BACCARNI, P. 1906: Funghi dell´Eritrea. Annali di Botanica (Rome) 4: 269–277.
BALDACCI, E. & CIFERRI, R. 1937: Un nuovo genere di micete parassita del pioppo Pollaccia radiosa
(Lib.) Baldacci e Ciferri, Revisione dei G. Stigmella e Stigmina. I. Pollaccia radiosa (Lib.)
Baldacci e Ciferri. Atti dell’Istituto Botanico “Giovanni Briosi“ e Laboratorio Crittogamica
Italiano della Reale Università di Pavia, Ser. 4, 10: 55–72.
BARR , M.E. 1968: The Venturiaceae in North America. Canadian Journal of Botany 46: 799–864.
BATISTA, A.C. 1957: Novos generos e especies de fungos imperfeitos. Revista de Biologia. Lisbon 1:
97–112.
BENSANDE, M. & KEITT, G.W. 1928: Comparative studies of certain Cladosporium diseases of stone
fruits. Phytopathology 18(4): 313–329.
BILGRAMI, K.S., JAMALUDDIN, S. & RIZWI, M.A. 1991: Fungi of India. List and references (second
revised and enlarged and brought up to date edition). New Delhi.
BONORDEN , H.F. 1851: Handbuch der allgemeinen Mykologie. Stuttgart.
BONTEA , V. 1985: Ciuperci parazite Õi saprofite din România. Vol. I. Editura Academiei Republicii
Socialiste România, BucureÕti.
BONTEA , V. 1986: Ciuperci parazite Õi saprofite din România, Vol. II. Editura Academiei Republicii
Socialiste România, BucureÕti.
BRANDENBURGER, W. 1985: Parasitische Pilze an Gefäßpflanzen in Europa. Fischer Verlag, Stuttgart,
New York.
BRAUN, A. 1853: Über einige neue oder weniger bekannte Pflanzenkrankheiten, welche durch Pilze
erzeugt werden. Verhandlungen des Vereins zur Beförderung des Gartenbaues in den Königlich
Preussischen Staaten 1: 165–191.
BRAUN, U. 1992: Taxonomic notes on some species of the Cercospora-complex. Nova Hedwigia
55(1–2): 211–221.
BRAUN, U. 1998: A monograph of Cercosporella, Ramularia and allied genera (Phytopathogenic
Hyphomycetes) Vol. 2. IHW-Verlag, Eching.
BRAUN, U. 2000: Miscellaneous notes on some micromycetes. Schlechtendalia 5: 31–56.
BRAUN, U., CROUS, P.W., DUGAN, F., GROENEWALD, J.Z. & HOOG, G.S. de (2003): Phylogeny and
taxonomy of Cladosporium-like hyphomycetes, including Davidiella gen. nov., the teleomorph
of Cladosporium s. str. Mycological Progress 2(1): 3–18.
BRAUN, U. & M ELNIK, V.A. 1997: Cercosporoid fungi from Russia and adjacent countries. Trudy
Botanicheskogo Instituta im V.L. Komarova (St. Petersburg) 20: 1–130.
BRAUN, U. & M OUCHACCA, J. 2000: Reassessment of Cercospora byrsonimatis and Ramularia
ligustrina. Mycological Research 104(8): 1009–1011.
BRAUN, U., RITSCHEL, A. & SCHUBERT, K. 2002: Proposal to conserve the generic name Fusicladium
against Spilocaea (Hyphomycetes). Taxon 51: 557.
120
Schlechtendalia 9 (2003)
BRIDSON, G.D.R. & SMITH, E.R. 1991: Botanico-Periodicum-Huntianum/Supplementum. Carnegie
Mellon University. Pittsburgh, PA. Allen Press Inc., KS.
BRUMITT, R.K. & POWELL, C.E. 1992: Authors of Plant Names. Royal Botanic Gardens, Kew.
BUTIN, H. 1992: Pollaccia catenospora sp. nov. associated with leaf spots of willow. Mycological
Research 96(8): 658–660.
CASTAGNE, J.L.M. 1845: Catalogue des plantes qui croissent naturellement aux environs de Marseille. Aix.
CAVARA, F. 1888: Appunti di patologia vegetale. Alcuni funghi parassiti di piante coltivate. Atti dell’
Istituto Botanico dell’Università di Pavia, Ser. 2, 1: 425–438.
CHARKEVICH, S.S. 1978: Flora i rastitel’nost’ Ussurijskovo zapovednika. Moskva.
CHI, P.K. 1994: Fungal Diseases of Cultivated Medicinal Plants in Guangdong Province. Guangdong
Academy Press.
CORDA, A.C.J. 1829: Deutschlands Flora, Abtheilung III. Die Pilze Deutschlands, Band 2, Heft 7. Nürnberg.
C ROUS , P.W., APTROOT , A., K ANG , J.C., B RAUN , U. & W INGFIELD , M.J. 2000: The genus
Mycosphaerella and its anamorphs. Studies in Mycology 45: 107–121.
CROUS , P.W. & BRAUN , U. 2003: Mycosphaerella and its anamorphs: 1. Names published in
Cercospora and Passalora. CBS Biodiversity Series 1: 1–571.
CROUS, P.W., KANG, J.C. & BRAUN, U. 2001: A phylogenetic redefinition of anamorph genera in
Mycosphaerella based on ITS rDNA sequences and morphology. Mycologia 93: 1081–1101.
CROUS, P.W., PHILLIPS, A.J.L. & BAXTER, A.P. 2000: Phytopathogenic fungi from South Africa.
Department of Plant Pathology, University of Stellenbosch.
CRUCHET , D. 1925: Recherches mycologiques á Montagny et aux environs d’Yverdon. Bulletin de la
Société Vaudoise des Sciences Naturelles 55: 155–177.
DAHAL, G., AMATYA, P. & MANANDHAR, H. 1992: Plant diseases in Nepal. Review of Plant Pathology
71: 799–806.
DANCE, B.W. 1961: Leaf and shoot blight of poplars (Section Tacamahaca Spach) caused by Venturia
populina (Vuill.) Fabric. Canadian Journal of Botany 39: 875–890.
DA SILVA, M. & MINTER, D.W. 1995: Fungi from Brazil recorded by Batista and co-workers.
Mycological Papers 169: 1–585.
DAVID, J.C. 1997: A contribution to the systematics of Cladosporium. Revision of the fungi previously
referred to Heterosporium. Mycological Papers 172: 1–157.
DAVIS, J.J. 1922: Notes on Parasitic Fungi. Transactions of the Wisconsin Academy of Sciences,
Arts and Letters 20: 389–432.
DEIGHTON, F.C. 1967: Studies on Cercospora and allied genera II. (Passalora, Cercosporidium and
some species of Fusicladium on Euphorbia). Mycological Papers 112: 1–80.
DEIGHTON, F.C. 1990: Observation on Phaeoisariopsis. Mycological Research 94(8): 1096–1102.
DEIGHTON F.C. & PIROZYNSKI, K.A. 1965: African species of Uncinula; some species of Fusicladiella;
various hyphomycetes, mainly tropical. Mycological Papers 101: 1–43.
DENNIS, R.W.G. 1986: Fungi of the Hebrides. Royal Botanic Gardens, Kew.
DIAS DE SOUSA, M.R. de & LUCAS, M.T: Fungi Lusitanae XXV. Agronomia Lusitana 37: 95–103.
DIAS DE SOUSA, M.R. de, LUCAS, M.T. & LOPES, M.C. 1981: Fungi Lusitanae XXXVIII. Agronomia
Lusitana 41: 77–92.
DIEDICKE, H. 1910: Aufzählungen der in der Umgebung Erfurts beobachteten Micromyceten. Erfurt.
DINGLEY, J.M. 1969: Records of plant diseases in New Zealand. Bulletin, New Zealand Department
of Scientific and Industrial Research 192: 1–298.
EL-BUNI, A.M. & RATTAN, S.S. 1981: Check-list of Libyan Fungi. Al Faateh University, Faculty of
Science, Department of Botany, Supplement to Flora of Libya. Tripoli.
ELLIS, M.B. 1971: Dematiaceous hyphomycetes. CMI, Kew.
ELLIS, M.B. 1976: More dematiaceous hyphomycetes. CMI, Kew.
ELLIS, M.B. & ELLIS, J.P. 1997: Microfungi on land plants. An identification handbook. New enlarged
edition. The Richmond Publishing Co., Lfd., Slough.
ERSHAD, D. 1995: Fungi of Iran. Agricultural Research, Education and Extension Organization (10). Tehran.
FARR, D.F., BILLS, G.F., CHAMURIS, G.P. & R OSSMAN, A.Y. 1989: Fungi on plants and plant products
in the United States. APS Press, St. Paul, MN.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
121
FERRARIS, T. 1912: Hyphales, Tuberculariaceae – Stilbaceae. Flora Italica Cryptogama Pars I: Fungi, Fasc. 6: 195–534.
FISHER, E.E. 1961: Venturia carpophila sp. nov., the ascigerous state of the apricot freckle fungus.
Transactions of the British Mycological Society 44(3): 337–342.
FRIES, E.M. 1819: Novitiae florae svecicae 5: 61–80.
FRIES, E.M. 1825: Systema orbis vegetabilis. Lundae.
FRIES, E.M. 1832: Systema mycologicum 3: 261–524.
FUNK, A. 1989a: Pollaccia borealis sp. nov. associated with a purple-brown leaf spot of aspen.
Canadian Journal of Botany 67: 776–778.
FUNK, A. 1989b: Observations on an aspen leaf spot disease and associated fungus, Pollaccia borealis.
Canadian Journal of Plant Pathology 11: 353–356.
GINNS, J.H. 1986: Compendium of plant diseases and decay fungi in Canada 1960–1980. Research
Branch, Agriculture Canada, Publications 1813. Ottawa.
GOLOVINA, N.P. 1964: Karakulinia gen. nov. Novosti Sistematiki Nizshikh Rastenii 1: 210–214.
GONZÁLES FRAGOSO, D.R. 1927: Estudio sistemático de los Hifales de la Flora Española. Memorias.
Real Academia de Ciencias Exactas, Físicas y Naturales de Madrid, Ser. 2a, 6: 1–377.
GOTTWALD, T.R. 1982: Taxonomy of the pecan scab fungus Cladosporium caryigenum. Mycologia
74(3): 382–390.
GULYAMOVA, M.G., KUCHMI, N.P., RAMAZANOVA, S.S., SAGDULLAEVA, M.SH. & KIRGIZBAEVA,
K H .M. 1990: Flora Gribov Uzbekistana. Tom 7. Sumchatye Griby. ‘Fan’ Publishing
House,Tashkent.
HARVEY, I.C. & BRAITHWAITE , A.F. 1982: Records of fungal plant diseases. New Zealand Journal of
Agricultural Research 25(3): 437–441.
HAWKSWORTH, D.L., SUTTON, B.C. & AINSWORTH, G.C. 1983: Ainsworth & Bisby’s Dictionary of
Fungi, 7th ed. Kew.
HÖHNEL , F. von 1919: Fünfte vorläufige Mitteilung mycologischer Ergebnisse (Nr. 399–500). Berichte der Deutschen Botanischen Gesellschaft 37: 153–161.
HÖHNEL , F. von 1923: Studien über Hyphomyzeten. Centralblatt für Bakteriologie, Parasitenkunde
und Infektionskrankheiten. 2. Abteilung 60: 1–26.
HOLMGREN, P.K., HOLMGREN, N.H. & BARNETT, L.C. 1990: Index Herbariorum, Part 1: The Herbaria
of the World, 8th ed. New York Botanical Garden, New York.
HUGHES, S.J. 1953: Some foliicolous hyphomycetes. Canadian Journal of Botany 31: 565–576.
HUGHES, S.J. 1958: Revisiones hyphomycetum aliquot cum appendice de nominibus rejiciendis.
Canadian Journal of Botany 36(6): 724–836.
HUGHES, S.J. & PIROZYNSKI, K.A. 1972: Diccocum Corda. Canadian Journal of Botany 50: 2521–2534.
ISHII, H., UDAGAWA, H., NISHIMOTO, S., TSUDA, T. & NAKASHIMA, H. 1992: Scab resistance in pear
species and cultivars. Acta Phytopathologica et Entomologica Hungarica 27(1–4): 293–298.
ISHII, H., WATANABE, H. & TANABE, K. 1997: Physiological races of Venturia nashicola on pears.
IOBC wprs Bulletin, Bulletin OILB srop 20(9): 130–133.
ISHII, H. & YANASE, H. 2000: Venturia nashicola, the scab fungus of Japanese and Chinese pears: a
species distinct from V. pirina. Mycological Research 104: 755–759.
JÄRVA, L. & PARMASTO, E. 1980: Eesti seente koondnimestik (List of Estonian fungi with host index
and bibliography). Scripta Mycologica 7: 1–331.
JÄRVA, L., PARMASTO, I. & VAASMA, M. 1998: Eesti seente koondnimestik 1. Täiendusköide (1975–
1990) [List of Estonian fungi with host index and bibliography, Supplement 1 (1975–1990)].
Scripta Mycologica 12: 18–111.
KANCHAVELI, K.G., KUKHALEISHVILI, L.K., RUKHALZE, T.A., CHHAIDZE, R.I., GULMAGARASHVILI,
V.K H., M ELIYA, M.P., MURVANISHVILI , I.K., N AHUTSRISHVILI, I.G., I NASHVILI , T S .N. &
CHIKOVANI, N.V. 1986: Flora sporovykh rastenij Gruzii (konspekt). Tbilisi.
KASANEN, R., HANTULA, J. & KURKELO, T. 2001: The occurrence of an undescribed species of
Venturia in blighted shoots of Populus tremula. Mycological Research 105(3): 338–343.
KHAN, S.N. & MISRA, B.M. 1989: Pollaccia blight of polars in India. European Journal of Forest
Pathology 19: 379–381.
122
Schlechtendalia 9 (2003)
KIFFER, E. & MORELET, M. 1999: The Deuteromycetes. Mitosporic Fungi, Classification and Generic
Key. Enfield.
KIRK, P.M., CANNON, P.F., DAVID, J.C. & STALPERS, J.A. 2001: Dictionary of the fungi, 9th ed. CABI
Publishing, Egham.
KORBONSKAYA, JA.I. 1990: Griby Tadzhikistana. ‘Donish’ Publishing House, Dushanbe.
KOSHKELOVA, E.N. 1977: Mikromitsety yuzhnogo Turkmenistana. T. 1 ‘Ylym’ Publishing House,
Ashkhabad.
KUHNHOLTZ-LORDAT, G. & BLANCHET, G. 1948: Flore des Environs immédiats de Montpellier. T. 2.
Les végétaux vasculaires et leurs parasites cryptogames. Paris.
KUYPER, J. 1912: Een Fusicladium-ziekte op Hevea. Bullettin van het Departement van Landbouw
in Suriname 28: 3–10.
LAUNDON, G.F. 1970: Records of fungal plant diseases in New Zealand. New Zealand Journal of
Botany 8: 51–66.
LAWRENCE, G.H.M., BUCHHEIM, A.F.G., DANIELS, G.S. & DOLEZAL, H. 1968: Botanico-PeriodicumHuntianum. Hunt Botanical Library, Pittsburgh, PA.
LIND, J. 1905: Über einige neue und bekannte Pilze. Annales Mycologici 3: 427–432.
LIND, J. 1913: Danish fungi as represented in the herbarium of E. Rostrup. Copenhagen.
LIND, J. 1928: The micromycetes of Svalbard. Skrifter om Svalbard og Nordishavet 13: 1–61.
LIND, J. 1934: Studies on the geographical distribution of arctic circumpolar micromycetes. Kongelige
Danske Videnskabernes Selskabs Naturvidenskabelige og Mathematiske Afhandlinger 11(2): 1–152.
LINDAU, G. 1907: Dr. L. Rabenhorst’s Kryptogamen-Flora von Deutschland, Oesterreich und der
Schweiz. Zweite Auflage. Erster Band: Pilze. Die Pilze Deutschlands, Oesterreichs und der
Schweiz. VIII. Abteilung: Fungi imperfecti: Hyphomycetes (erste Hälfte), Mucedinaceae,
Dematiaceae (Phaeosporae und Phaeodidymae). Leipzig.
LINFORD, M.B. 1926: Black-Leaf of peas caused by Fusicladium pisicola n. sp. Phytopathology 16:
549–558.
LINK, H.F. 1825: in WILLDENOW, C.L., Species plantarum, Ed. 4, 6(2). Berlin.
LIU, Z.N., CHEN, S.F. & SHAO, Y.H. 1981: Fungus diseases of the North East trees. Science Press, China.
MAMLUK, O.F., ABU GHARBIEH, W.I., SHAW, C.G., AL-MUSA, A. & AL-B ANNA, L. 1984: A checklist
of plant diseases in Jordan. Faculty of Agriculture, University of Jordan, Amman.
MARHOLD, K. & HINDÁK, F. (eds.) 1998: Checklist of non-vascular and vascular plants of Slovakia.
VEDA, Bratislava.
MATHUR, R.S. 1977: Check-list of Afghani fungi and plant diseases. University of Baghdad, Natural
History Research Centre, Publication No. 32: 1–64.
MCCLAIN, R.L. 1925: Scab of Christmas Berry, Photinia arbutifolia Lindl., due to Fusicladium
photinicola n. sp. Phytopathology 15: 178–182.
MCKENZIE, E.H.C., O’SULLIVAN, P.J. & W ILKIE, J.P. 1992: A list of type specimens of New Zealand
fungi. Mycotaxon 43: 77–156.
MELNIK, V.A. & POPUSCHOI, I.S. 1992: Nesovershennye griby na drevesnych i kustarnikovych
porodach. Kishinev.
MENDEZ, M.A.S., SILVA, V.L. da, DIANESE, J.C., FERREIRA, M.A.S.V., SANTOS, C.E.N. dos, N ETO,
E.G., URBEN, A.F. & CASTRO, C. 1998: Fungos em plantas no Brasil. EMBRAPA. Brasília, D.F.
MENON, R. 1956: Studies on Venturiaceae on rosaceous plants. Phytopathologische Zeitschrift 27(2):
117–146.
MØLLER, F.H. 1958: Fungi of the Faeroes. Part 2. Copenhagen.
MORELET, M. 1978: Sur deux reclassements fongiques. Bulletin, Société des Sciences Naturelles et
d’Archéologie de Toulon et du Var 34: 12.
MORELET, M. 1985: Les Venturia des Peupliers de la section Leuce. 1. Taxonomie. Cryptogamie,
Mycologie 6: 101–117.
MORELET, M. 1993: Note préliminaire sur quatre ascomycètes pathogènes. Annales de la Société des
Sciences Naturelles et d’Archéologie de Toulon et du Var 45(3): 217–220.
MORELET, M. & SIGAUD, P. 1996: Observations on a poplar leaf infection occurring in North-East
China: The Grey Spot Disease. Cryptogamie, Mycologie 17(1): 11–20.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
123
MUJICA, R.F. & VERGARA, C.C. (revisada y actualizada por: E. Oehrens B.) 1980: Flora Fungosa
Chilena. Santiago de Chile (2nd edition). Universidad de Chile, Facultad de Agronomica, Ciencias
Agricolas, No. 5, Santiago de Chile.
MÜLLER, E. & ARX, J.A. von 1950: Einige Aspekte zur Systematik pseudosphaerialer Ascomyceten.
Bericht der Schweizerischen Botanischen Gesellschaft 60: 329–397.
MÜLLER, E. & ARX, J.A. von 1962: Die Gattungen der didymosporen Pyrenomyceten. Beiträge zur
Kryptogamenflora der Schweiz. 11(2): 1–922.
MUSKETT, A.E. & MALONE, J.P. 1985: Catalogue of Irish fungi – VI. Deuteromycotina. Proceedings of
the Royal Irish Academy. Section B, Biological, Geological and Chemical Sciences 85: 133–200.
NATTRASS, R.M. 1937: A first list of Cyprus fungi. Department of Agriculture, The Government of
Cyprus, Nicosia.
NÜESCH, J. 1960: Beitrag zur Kenntnis der weidenbewohnenden Venturiaceae. Phytopathologische
Zeitschrift 39: 329–360.
ONDÌEJ, M. 1971: Houby rodu Fusicladium Bonorden, tvoÍici konidie v Íetizcich (Hyphomycetes,
Fungi imperfecti). „eská Mykologie 25(3): 165–172.
ONDÌEJ, M. 1972: Ein Beitrag zur Kenntnis der parasitischen imperfekten Pilze der Gattung Pollaccia
E. Bald. et Cif. an Pappeln (Populus spp.). European Journal of Forest Pathology 2: 140–146.
ONDÌEJ, M. 1973: Nové a málo známé houby rodu Fusicladium Bonorden na topolech a vrb•. „eská
Mykologie 27(4): 236–240.
ONDÌEJ, M. 1984: Pollaccia spiraeae (Karakulin) OndÍej. „eská Mykologie 38: 46–48.
OSIPYAN, L.L. 1971: Novye vidy gifal’nykh gribov v Armyanskoj SSR. Mikologiya i Fitopatologiya
5(1): 87–90.
OSIPYAN, L.L. 1975: Gifal’nye Griby. Mikoflora Armjanskoj SSR. T. 3. Erevan.
PANDOTRA, V.R. 1997: Illustrated fungi of North India with special reference to J & K state. International Book Distributors, Dehra Dun.
PARTRIDGE, E.C. & MORGAN-JONES, G. 2003: Notes on Hyphomycetes. XC. Fusicladosporium, a
new genus for cladosporium-like anamorphs of Venturia, and the pecan scab-inducing fungus.
Mycotaxon 85: 357–370.
PASCOE, I.G. & SUTTON, B.C. 1990: Protoventuria parahebicola sp. nov. (Venturiaceae), the teleomorph
of Fusicladium veronicae on Parahebe perfoliata. Australian Systematic Botany 3: 281–285.
PASHCHENKO, A.YA., GAPONENKO, N.I., RAMAZANOVA, S.S., SAGDULLAEVA, M.SH., KIRGIZBAEVA,
KH.M. & MELNIK, V.A. 1978: Herbarium of fungi and collection of pure cultures (Institute of
Mikrobiology of the Academy of Sciences of Uzbek SSR). Tashkent. (in Russian).
PICBAUER, R. 1927: Addenda ad floram „echoslovakiae mycologicam Pars 3. Sborník Vysoké Òkoly
Zem•d•lské v Brn• D 7: 3–25.
PICBAUER, R. 1931: Addenda ad floram „echoslovakiae mycologicam Pars 5. Sborník Vysoké Òkoly
Zem•d•lské v Brn• D 18: 1–30.
PICBAUER, R. 1932: Addenda ad floram „echoslovakiae mycologicam Pars 6. Práce Moravské
pÍírodov•decké spole…nosti. 7(4): 1–17.
PICBAUER, R. 1937: Addenda ad floram „echoslovakiae mycologicam Pars 8. Verhandlungen des
Naturforschenden Vereines in Brünn 69: 29–45.
PLAKIDAS, A.G. 1942: Venturia acerina, the perfect state of Cladosporium humile. Mycologia 34: 27–37.
PÕLDMAA, P. 1967: Fitopatogennye Mikromicety Severnoj Estonii (= Phytopathogenic micromycetes
of the North Estonia). Tallin.
POSPELOV, A.G., ZAPROMETOV, N.G. & DOMASHEVA, A.A. 1957: Gribnaja Flora Kirgizskoj SSR,
Vyp. 1., Inst. Bot. Akad. Nauk Kirgizskoj SSR, Frunze.
P RILLIEUX , É.E. 1892: Observation sur le Napicladium tremulae, forme conidienne du
Didymosphaeria populina. Bulletin de la Société Mycologique de France 8: 26.
PYLDMAA, P. 1967: Fitopatogennye Mikromicety Severnoj Estonii. Tallin.
RAABE , R.D. & GARDENER, M.W. 1972: Scab on Pyracantha. Phytopathology. 62: 914–916.
RÉVAY, A. 1998: Review of the hyphomycetes of Hungary. Studia Botanica Hungarica 27–28: 5–74.
RITSCHEL, A. 2001: Taxonomische Revision der Gattungen Pollaccia und Spilocaea (Hyphomycetes,
Venturia-Anamorphen), Diplom-Arbeit, Martin-Luther-Universität Halle: 1–88.
124
Schlechtendalia 9 (2003)
ROSTRUP, E. 1883: Fortsatte Undersögelser over snyltesvampes Angreb pan Skovtroerus (med systen
Trosnit) (Fortgesetzte Untersuchungen über Angriffe von Schmarotzerpilzen an Waldbäumen.
Mit 17 Holzschnitten.). Tidsskrift for Skovbrug 6: 199–300.
ROSTRUP, O. 1935: Bidrag Til Danmarks Svampeflora 2. Dansk Botanisk Arkiv Udgivet af Dansk
Botanisk Forening 8(8): 46–57.
RULAMORT, M. de 1986: Remarques taxonomiques et nomenclaturales sur quelques micromycètes.
Bulletin de la Société Botanique du Centre-Ouest, N.S. 17: 191.
SACCARDO, P.A. 1878: Fungi Veneti novi vel critici vel Mycologiae Venetae addenti. Series VIII.
Michelia 1(2): 239–273.
SACCARDO, P.A. 1884: Sylloge Fungorum vol. 3. Padova.
SACCARDO, P.A. 1886: Sylloge Fungorum vol. 4. Padova.
SACCARDO, P.A. 1892: Sylloge Fungorum vol. 10. Padova.
SACCARDO, P.A. 1895: Sylloge Fungorum vol. 11. Padova.
SACCARDO, P.A. 1897: Sylloge Fungorum vol. 12 (Sydow, P. ed.). Borntraeger, Berlin.
SACCARDO, P.A. 1899: Sylloge Fungorum vol. 14 (Saccardo, P.A. & Sydow, P. eds.). Padova.
SACCARDO, P.A. 1902: Sylloge Fungorum vol. 16 (Saccardo, P.A. & Sydow, P. eds.). Padova.
SACCARDO, P.A. 1906: Sylloge Fungorum vol. 18 (Saccardo, P.A. & Saccardo, D. eds.). Padova.
SACCARDO, P.A. 1910: Sylloge Fungorum vol. 19 (Saccardo, P.A. & Traverso, G.B. eds.). Padova.
SACCARDO, P.A. 1913: Sylloge Fungorum vol. 22 (Saccardo, P.A. & Trotter, A. eds.). Padova.
SACCARDO, P.A. 1931: Sylloge Fungorum vol. 25 (Trotter, A. ed.). Avellino.
SAGDULLAEVA, M.Sh., KIRGIZBAEVA, KH.M., RAMAZANOVA, S.S., GULYAMOVA, M. & FAJZIEVA,
F.K H. 1990: Flora Gribov Usbekistana. T. 6. Gifal’nye Griby (Dematiaceae). ‘Fan’ Publishing
House, Tashkent.
SAMUELS, G.J. & SIVANESAN, A. 1975: Venturia asperata sp. nov. and its Fusicladium state on Apple
Leaves. New Zealand Journal of Botany 13: 645–652.
SCHEUER, C. 2003: Dupla Fungorum, Supplementum (2003), verteilt vom Institut für Botanik der
Universität Graz (GZU). Fritschiana 40: 1–51.
SCHNABEL, G., S CHNABEL, E.L. & JONES, A.L. 1999: Characterisation of ribosomal DNA from
Venturia inaequalis and its phylogenetic relationship to rDNA from other tree-fruit Venturia
species. Phytopathology 89: 100–108.
SCHOLLER, M., BRAUN, U. & RUHL, G. 2003: Fusicladium levieri, a new fungal parasite of Persimmon
in Indiana. Proceedings of the Indiana Academy of Sciences (in press).
SCHUBERT, K. 2001: Taxonomische Revision der Gattung Fusicladium (Hyphomycetes, VenturiaAnamorphen), Diplom-Arbeit, Martin-Luther-Universität Halle: 1–136.
SCHUBERT, K. & BRAUN, U. 2002: Fusicladium. IMI Descriptions of Fungi and Bacteria 152, no.
1511–1520.
SCHWEIZER, H. 1958: Beiträge zur Biologie des Kirschen- und Pfirsichschorferregers (Fusicladium
cerasi (Rabenh.) Sacc., Venturia cerasi Aderh. und Cladosporium carpophilum Thüm.). Phytopathologische Zeitschrift 33: 55–98.
SERVAZZI, O. 1939: Ricerce sulla cosi detta “Defogliazione primaverile dei Pioppi“. Bollettino del
Laboratorio Sperimentale e Osservatorio di Fitopatologia 15: 49–152.
SHIRAI, M. & HARA, K. 1927: A list of Japanese fungi hitherto known, Ed. 3.
SHVARTSMAN, S.R., VASYAGINA, M.P., BYZOVA, Z.M. & FILIMONOVA, N.M. 1975: Nesovershennye
griby – Fungi imperfecti (Deuteromycetes). 2. Monilial’nye – Moniliales. T. 8(2). Flora sporovykh
rastenij Kazakhstana. ‘Nauka’ Publishing House, Alma-Ata.
SIVANESAN, A. 1977: The taxonomy and pathology of Venturia species. Bibliotheca Mycologica 59: 1–139.
SIVANESAN, A. 1984a: The Bitunicate Ascomycetes and their anamorphs. Cramer Verlag, Vaduz.
SIVANESAN, A. 1984b: Acantharia, Gibberia and their anamorphs. Transactions of the British
Mycological Society 82(3): 507–529.
SORAUER, P.C.M. 1895: Die in Deutschland aufgetretenen Krankheitserscheinungen. E. Öl- und
Gemüsepflanzen. Zeitschrift für Pflanzenkrankheiten 5: 103–105.
STAHEL, G. 1917: De Zuid-Amerikaansche Hevea-Bladziekte veroozaakt door Melanopsammopsis
ulei nov. gen. Bullettin van het Departement van Landbouw in Suriname 34: 1–111.
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
125
STRASSER, P. 1907: Vierter Nachtrag zur Pilzflora des Sonntagberges (N.-Ö.), 1904. Verhandlungen
der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 57: 299–320.
SUBRAMANIAN, C.V. 1971: Hyphomycetes: an account of Indian species, except Cercosporae. New
Delhi.
SUTTON, B.C. 1970: Forest microfungi. IV. A leaf spot of Populus caused by Cladosporium subsessile.
Canadian Journal of Botany 48: 471–477.
SUTTON, B.C. & PASCOE, I. 1988: Fusicladium veronicae (Batista), comb. nov., causing brown leaf
blight of Parahebe species. Australian Systematic Botany 1: 79–86.
TAI, F.L. 1979: Sylloge Fungorum Sinicorum. Science Press, Academia Sinica, Peking.
TUBEUF, C. von 1902: Das Triebsterben der Weiden (Fusicladium saliciperdum Tub. syn. Septogloeum
saliciperdum Allesch. et Tub.). Arbeiten aus der Biologischen Reichsanstalt für Land- und Forstwirtschaft 2: 567–570.
ULVINEN, T., OHENOJA, E., AHTI, T. & ALANKO, P. 1981: A check-list of fungi (incl. lichens) of the
Koillismaa (Kuusamo) biological province, N.E. Finland. Oulanka Reports 2: 1–64.
VASSILJEVSKY, N.I. & KARAKULIN, B.P. 1937: Parazitnye nesovershennye griby. Ch. I. Gifomitsety.
Izdatel’stvo Akademii Nauk SSR, Leningrad.
VASYAGINA, M.P. 1957: Noye vidy gifomicetov Kazakhstana. Izvestiya Akademiya Nauk Kazakhskoi
S S R, Seriya Biologicheskaya 1, 13: 100–103.
V ERKLEY, G.J.M. 1997: Ultrastructural evidence for two types of proliferation in a single
conidiogenous cell of Septoria chrysanthemella. Mycological Research 102: 368–372.
VIÉGAS, A.P. 1961: Índice de Fungos da Amérika do Sul. Inst. Agron., Campinas.
VIENNOT-BOURGIN, E. 1949: Les champignons parasites des plantes cultivées. vol. 1. Paris.
WAKEFIELD, E.M. & BISBY, G.R. 1941: List of hyphomycetes recorded for Britain. Transactions of
the British Mycological Society 25(1): 84–91.
WARCUP, J.H. & TALBOT, P.H.B. 1981: Host-pathogen index of plant diseases in South Australia.
Deptartment of Plant Pathology, Waite Agriculture Research Institute, University Adelaide.
WU, W.P. & SUTTON, B.C. 1995: Further observations on Pollaccia mandshurica, a pathogen of
Populus sp. in China. Mycological Research 99: 983–986.
14.
Acknowledgements:
We are grateful to the directors and curators of the herbaria listed in chapter 2 for loaning type
material and other collections during the course of these monographic studies, and we thank D.
Triebel (Botanische Staatssammlung, Munich, Germany) for making it possible to carry out molecular examinations of Fusicladium spp. The colleagues of “Interdisziplinäres Wissenschaftliches Zentrum
für Materialwissenschaften” of the University Halle supported this project by providing possibilitiesto
use the ESEM technique for SEM studies, for which we are much obliged. We are especially grateful
to C.F. Hill (National Plant Pest Reference Laboratory, Ministry of Agriculture & Forestry, Auckland, New Zealand) who checked the whole text of the manuscript.
Addresses of the authors:
K. Schubert and Dr. U. Braun, Martin-Luther-Universität, FB Biologie, Intitut für Geobotanik und
und Botanischer Garten, Neuwerk 21, D-06099 Halle, Germany
(e-mail: schubert@botanik.uni-halle.de, braun@botanik.uni-halle.de)
A. Ritschel, Eberhard-Karls-Universität, Spezielle Botanik/Mykologie, Auf der Morgenstelle 1, D72076 Tübingen, Germany
(e-mail: anja.ritschel@uni-tuebingen.de).
126
15.
15.1.
Schlechtendalia 9 (2003)
Index
Index of fungal names
Acantharia 8
acerina Venturia 57
aconiti Fusicladium 109
ahmadii Fusicladium 1, 14, 17, 18
ahmadii Spilocaea 17, 80
alopecuri Cladosporium 112
alopecuri Fusicladium 112, 113
Alternaria 116
amelanchieris Spilocaea 77, 80
americana Pollaccia 87, 89
amygdali Fusicladium 29
anethi Fusicladium 113
angelicae Fusicladium 113
angelicae Passalora 113
Anungitea 114
Apiosporina 8
aplectri Fusicladium 109
aronici Fusicladium 113
ascyrinum Fusicladium 113
asiaticum Fusicladium 64, 65
asperata Venturia 5, 18, 20
asperatum Fusicladium 1, 13, 18, 19, 20
Asperisporium 10
asteris var. latispora Ramularia 114
asteroma Cladosporium 85
asteroma var. macrosporum Cladosporium
85
asteroma var. microsporum Cladosporium
85, 89
asteroma Fusicladium 85
asteroma Napicladium 43, 85
asteroma var. microsperma Napicladium
89
balsamiferae Pollaccia 43
bambusicola Cercosporidium 113
bambusicola Fusicladium 113
bambusicola Pseudospiropes 113
bambusicolum Cercosporidium 113
Basiascum 3, 8
betulae Asteroma 22
betulae Fusicladium 11, 20, 21, 22
bicolor Fusicladiella 113
bicolor Fusicladium 113
borealis Pollaccia 91, 92
borealis Venturia 91, 92
brevicatenatum Pseudocladosporium 114
brevipes Cladosporium 99
brevipes Fusicladium 12, 22, 23, 74
butleri Fusicladium 113
butleri Pseudocercospora 113
butyrospermi Fusicladium 109
byrsonimatis Fusicladium 1, 12, 23, 24
byrsonimatis Pseudocercospora 23
byrsonimatis Ramalia 23
caricae Asperisporium 113
caricae Fusicladium 113
caricinum Fusicladium 6, 7, 11, 24, 25
carpineum Fusicladium 11, 26, 27
carpini Fusicladium 26, 109
carpophila Venturia 5, 29
carpophilum Cladosporium 4, 26
carpophilum Clasterosporium 30
carpophilum Coryneum 30
carpophilum Fusicladium 4, 13, 26, 28,
30, 34, 35
carpophilum Fusicladosporium 4, 9, 26
carpophilum Megacladosporium 26
caruanianum Fusicladium 114
caruanianum Pseudocladosporium 1, 114,
115
caryigenum Cladosporium 4, 5, 41
caryigenum var. carpineum Cladosporium
26
caryigenum Fusicladium 41
catenospora Pollaccia 7, 30
catenosporum Fusicladium 1, 15, 30, 31
caulicola Fusicladium 1, 6, 11, 31, 32
cecropiae Fusicladium 114
cecropiae Pseudocercospora 114
cephalanthi Fusicladium 110
cerasi Acrosporium 33
cerasi Cladosporium 33
cerasi Fusicladiopsis 33
cerasi Fusicladium 4, 7, 13, 33, 34, 35
cerasi Karakulinia 9, 33
cerasi Megacladosporium 33
cerasi Venturia 5, 33
Cercostigmina 7
chaetomium Cladosporium 47
chanousii Fusicladium 110
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
chlorinum Fusicladium 114
chlorospora Venturia 94, 108, 109
Cladophialophora 5
cladosporioides Cladosporium 5
Cladosporium 3, 4, 5, 10, 20, 30, 43, 57,
65, 96, 105, 109, 116, 118
collinsii Apiosporina 105
collinsii Dimerosporium 105
collinsii Sphaeria 105
concentrica Spilocaea 3, 112
consors Fusicladium 11, 36
convolvularum Fusicladium 6, 11, 37, 38
crataegi Actinonema 76
crataegi f. sorbi ariae Actinonema 76
crataegi f. sorbi-torminalis Actinonema 76
crataegi var. arachnoideum f. sorbitorminalis Actinonema 76
crataegi Asteroma 76
crataegi var. pomi Asteroma 76
crataegi var. sorbi Asteroma 76
crataegi Capillaria 76
crataegi Fusicladium 13, 37, 39
crataegi Megacladosporium 37
crataegi Phlyctidium 76
crataegi Spilocaea 76
crataegi Venturia 37, 80
Cycloconium 3, 8
cynanchi Fusicladium 114
dearnessiana Acrotheca 82
dearnessianum Fusicladium 82
Denticularia 10
dentritica Passalora 76
dentritica var. orbiculata Passalora 76
dentriticum Cladosporium 76
dentriticum var. heteromeles Cladosporium
76
dentriticum Fusicladium 3, 4, 67, 76
dentriticum f. microsperma Fusicladium
77
dentriticum var. eriobotryae Fusicladium
77
dentriticum var. opuli Fusicladium 76, 81
dentriticum var. orbiculatum Fusicladium
76
dentriticum var. pyracanthae Fusicladium
76
dentriticum var. soraueri Fusicladium 76
dentriticum var. sorbinum Fusicladium 77
127
dentriticum var. sorbinum f. fruticola
Fusicladium 77
dentriticum var. sorbi-torminalis Fusicladium 77
depressa Passalora 72, 114, 117
depressum Fusicladium 114
depressum f. petroselini Fusicladium 114
depressum var. platysporum Fusicladium
115
depressum var. sii Fusicladium 115
depressum var. tommasiniae Fusicladium
115
destruens Fusicladium 115
diedickeanum Fusicladium 12, 40, 41
diospyri Fusicladium 61
ditricha Venturia 5, 20, 22, 97
dothidea Botryosphaeria 5
dubiosum Fusicladium 116
echinata Acantharia 104, 105
effusum Cladosporium 41
effusum Fusicladium 4, 6, 12, 26, 37, 41,
42, 43
effusum var. carpineum Fusicladium 26,
43
effusum Fusicladosporium 41
elasticae Fusicladium 116
elegans Fusicladium 1, 14, 43, 44
elegans Pollaccia 4, 43, 45, 87
ephedrae Fusicladium 110
ephedrae Venturia 110
epiphylla Spilocaea 3, 112
eriobotryae Basiascum 8, 77, 80
eriobotryae Fusicladium 77
eriobotryae Spilocaea 77, 80
euonymi-japonici Fusicladium 116
euphorbiae Fusicladiopsis 45
euphorbiae Fusicladium 12, 45, 46, 51
euphorbiae Karakulinia 45
fagopyri Fusicladium 116
fasciculata Passalora 47
fasciculatum Cercosporidium 47
fasciculatum Fusicladium 11, 49
fasciculatum var. fasciculatum Fusicladium 47, 48, 49
fasciculatum var. didymum Fusicladium
49, 50
fasciculatum Scolecotrichum 47
fautreyi Fusicladium 12, 51
128
fici Fusicladium 110, 116
fraxini Actinonema 52
fraxini Fusicladium 6, 12, 52, 53
fraxini var. phillyreae Fusicladium 53, 74,
110
fraxini Scolecotrichum 52
fraxini Spilocaea 52
fraxini Venturia 52
fuliginosum Fusicladium 116
fuscescens Fusicladium 82, 85
Fusicladiella 10
Fusicladiopsis 4, 8
Fusicladium 1, 2, 3, 4, 5, 6, 7, 8, 10, 11,
15, 31, 36, 37, 43, 45, 53, 55, 57, 62,
68, 71, 94, 96, 100, 105, 108, 109, 114,
116, 117
Fusicladium-state of Acantharia echinata
12, 104
Fusicladium-state of Apiosporina collinsii
12, 105, 106
Fusicladium-state of Apiosporina morbosa
13, 106, 107
Fusicladosporium 4, 9
galii Passalora 118
gardeniae Fusicladium 110
gnaphaliatum Asperisporium 116
gnaphaliatum Fusicladium 116
granulosum Fusicladium 52
grayana Phaeoisariopsis 54
grayiana Isariopsis 54
grayiana Phaeoisariopsis 54
grayianum Fusicladium 14, 54
grisea Pyricularia 116
hachijoense Pseudocladosporium 114
hariotianum Fusicladium 116
herbarum Cladosporium 116, 118
heterospora Passalora 116
heterosporum Fusicladium 116
heveae Fusicladium 12, 55, 56
heveae Passalora 55
heveae Scolecotrichum 55
hippophaës Fusicladium 111
Hormocladium 4, 8, 62
humile Cladosporium 57
humile Fusicladium 11, 57, 58
humile Fusicladosporium 57
inaequalis Venturia 5, 77, 80
Schlechtendalia 9 (2003)
junci Fusicladium 1, 6, 12, 57, 59
kaki Fusicladium 8, 61, 62
Karakulinia 4, 9
lageniforme Fusicladium 87
lalandi Fusicladium 77, 80
lathyrinum Dicoccum 60
lathyrinum Fusicladium 12, 60
lathyrmum Dicoccum 60
lethifera Pollaccia 87, 89
lethiferum Cladosporium 87, 89
levieri Cladosporium 61
levieri Fusicladium 6, 11, 61, 62
levieri Phaeoramularia 61
levieri Ragnhildiana 61
lini Fusicladium 111
livistoniae Fusicladium 117
lonicerae Fusicladium 111
macrocarpum Cladosporium 115
macrosporium Fusicladium 55, 117
macrosporum Fusicladium 55
macularis Venturia 87
maculicola Cladosporium 91
maculicola Fusicladium 90, 117
maculicola Passalora 117
maculicola Phaeoramularia 91, 100
maculicola Scolecotrichum 117
maculicola Torula 90
maculicolum Scolecotrichum 117
mali Asteroma 76
mali Coniosporum 77
mali Fumago 76
mandshurica Mycosphaerella 64
mandshurica Pollaccia 62
mandshurica Venturia 62
mandshuricum Fusicladium 1, 14, 62, 63
martianoffianum Cladosporium 64, 65
martianoffianum Fusicladium 6, 14, 57,
64, 65
Megacladosporium 4, 8
melaena Fusicladiella 113, 118
melanconioides Fusicladium 77
minutulum Asperisporium 117
minutulum Fusicladium 117
morbosa Apiosporina 106
moreletii Venturia 87, 89
Mycosphaerella 7, 8
Napicladium 8
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
nashicola Fusicladium 1, 14, 65, 66
nashicola Venturia 5, 6, 65, 67
nebulosa Spilocaea 6, 67
nebulosum Dicoccum 67
nebulosum Fusicladium 1, 12, 67, 68
obducens Fusicladium 6, 13, 68, 69
Oidium 8
oleaginea Spilocaea 6, 70
oleagineum Cycloconium 8, 70
oleagineum Fusicladium 1, 12, 70, 71
oleaginum Cycloconium 70
opuntiae Spilocaea 3, 112
orbiculatum Cladosporium 76
orbiculatum Fusicladium 76
ovata Monotospora 117
parahebicola Protoventuria 100
Passalora 10, 115
perexigua Didymoshaeria 36
peucedani Asperisporium 72
peucedani Fusicladium 11, 72, 73, 117
peucedani Pollaccia 72
Phaeoramularia 62
phillyreae Cycloconium 73
phillyreae Fusicladium 1, 12, 73, 74
phillyreae Spilocaea 53, 73, 110
phlei Cladosporium 116
photinicola Fusicladium 77
photinicola Spilocaea 77, 80
pirina Venturia 82
pirinum var. amelanchieris Fusicladium 76
pirinum var. pyracanthae Fusicladium 76
pirinum Megacladosporium 82
pisicola Fusicladium 7, 12, 74, 75
platyspora Passalora 115, 117
Pollaccia 1, 2, 3, 4, 5, 6, 7, 8, 11, 15, 43,
72, 87, 89, 100
polymorphum Cladosporium 82
Polyscytalum 114
pomi Actinonema 76
pomi Fusicladium 3, 12, 13, 14, 18, 76,
78, 80, 81
pomi Passalora 82
pomi Spilocaea 3, 6, 8, 76, 80
pongamiae Asperisporium 117
pongamiae Fusicladium 117
populi Clasterosporium 87
populi Fusariella 85
129
populi Stigmina 87
populi-albae Pollaccia 89
populina Venturia 4, 5, 43, 45
populinum Coryneum 64
populinum Dicoccum 87
poricola Fusicladium 117
poricola Porophilomyces 117
porrigo Cercospora 82, 85
praecox Cladosporium 117
praecox Fusicladium 117
proteae Batcheloromyces 118
proteae Spilocaea 118
Protoventuria 8
pruni Fusicladium 29
Pseudocercospora 7, 8, 10
Pseudocladosporium 114
Pseudofusicladium 4
psoraleae Dicoccum 81
psoraleae Fusicladium 7, 12, 81
punctiforme Fusicladium 117
punctiformis Cercospora 114
punctiformis Passalora 117
punctum Passalora 113, 114
pyracanthae Fusicladium 76
pyracanthae Passalora 76
pyracanthae Spilocaea 76, 80
pyrina Passalora 82
pyrina Venturia 5, 6, 67, 82
pyrinum Arthrinium 82
pyrinum Fusicladium 82
pyrinum Fusidium 82
pyrorum Fusicladium 3, 6, 8, 13, 14, 65,
82, 83, 85
pyrorum f. carpophila Fusicladium 82
pyrorum var. cladophilum Fusicladium 82
pyrorum Helminthosporium 76, 82
pyrorum Megacladosporium 82
radiosa Pollaccia 4, 8, 45, 53, 85, 87, 90, 92
radiosa var. lethifera Pollaccia 87, 89
radiosa var. populi-albae Pollaccia 89, 90
radiosa var. radiosa Pollaccia 90
radiosa Stigmina 43, 85
radiosa Venturia 85
radiosum Fusicladium 3, 4, 14, 43, 45,
85, 87, 89
radiosum var. balsamiferae Fusicladium
43
130
radiosum var. lethiferum Fusicladium 1,
15, 87, 88, 89
radiosum var. populi-albae Fusicladium
1, 14, 89, 90
radiosum var. microsporum Fusicladium
85
radiosum var. radiosum Fusicladium 14,
85, 86, 89
radiosum Oidium 4, 85
Ramalia 4, 9
Ramularia 7, 10
ramulosa Pollaccia 85
ramulosum Cladosporium 85, 90, 94
ramulosum Fusicladium 85, 92, 94
rhamni Fusicladium 118
robiniae Fusicladium 118
robiniae Passalora 118
romellianum Fusicladium 6, 14, 57, 90,
91, 92, 100
ruthenicum Fusicladium 118
saliciperda Pollaccia 92
saliciperda Venturia 92, 94
saliciperdum Fusicladium 15, 92, 93, 108
saliciperdum Septogloeum 92, 94
salicis Fusicladium 30, 31
schnablianum Fusicladium 118
scillae Cladosporium 94, 96
scillae Fusicladium 6, 12, 94, 95
scirpi Spilocaea 3, 112
scribnerianum Cladosporium 96
scribnerianum Fusicladium 11, 96, 97
Septoria 7
sii Passalora 115
sinensis Pollaccia 62
soraueri Napicladium 8, 76
sorghi Hadrotrichum 118
sorghi Fusicladium 118
Spilocaea 1, 2, 3, 4, 5, 6, 7, 8, 11, 15, 53,
68, 71, 112
spiraeae Fusicladium 14, 97, 98
spiraeae Pollaccia 97
spiraeae Scolecotrichum 97
statices Fusicladium 118
stuckertii Fusicladium 111
stuckertii Napicladium 111
stuckertii Sporhelminthium 111
subsessile Cladosporium 99
Schlechtendalia 9 (2003)
subsessile Fusicladium 14, 99, 100
syringae Venturia 40
tenue Fusicladium 111
theae Fusicladium 112
transversum Fusicladium 118
tremulae Fusicladium 43, 45, 52, 53, 85,
87, 89, 94
tremulae Napicladium 43, 45, 85
tremulae Venturia 4, 5, 87, 92
tremulae var. grandidentatae Venturia 87,
89
tremulae var. populi-albae Venturia 89
tremulae var. tremulae Venturia 85
ulei Microcyclus 55
vanillae Fusicladium 112
venosum Scolecotrichum 76
Venturia 2, 3, 4, 5, 7, 8, 9, 10, 26, 30, 35,
37, 43, 53, 57, 64, 87, 109
veronicae Fusicladium 6, 15, 100, 101
veronicae Ramalia 4, 9, 100
virescens Fusicladium 3, 4, 8, 82
virgaureae Fusicladium 11, 102
viticis Fusicladium 15, 103
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
2.2.2.3.
131
Index of host genera
Acacia 30
Acer 11, 57
Aconitum 109
Alnus 22, 117
Alopecurus 49, 112
Amelanchier 12, 77, 78, 79, 80, 105
Ammophila 49
Amygdalus 29
Anethum 113
Angelica 11, 72, 113
Aplectrum 109
Archangelica 113
Aronia 13, 79, 82, 84
Arum 81
Ascyrum 113
Astragalus 12, 22, 23
Atropis 116
Avena 115
Baccachis 111
Berberis 22
Betula 11, 20, 22, 96, 97
Bromus 30
Butyrospermum 109
Byrsonima 12, 23
Calystegia 11, 37
Camellia 112
Carduus 118
Carex 11, 24, 25
Carica 113
Carya 12, 26, 41, 43
Carpinus 11, 26, 43, 109
Cephalantus 110
Chaenomeles 13, 84
Chaerophyllum 113
Cheirodendron 30
Cicuta 11, 72
Convolvulus 11, 37
Cotoneaster 13, 79, 80
Crataegus 13, 37, 40, 76, 80, 81
Cynanchum 114
Digitaria 116
Diospyros 11, 61, 62
Docynia 13, 79, 81
Ephedra 110
Epilobium 116
Eriobotrya 13, 77, 78, 79, 80, 84, 85
Euphorbia 11, 12, 45, 47, 49, 51
Fagopyrum 116
Ficus 110
Filipendula 30
Fraxinus 12, 52, 53, 67, 68
Gardenia 110
Gentiana 110
Glehnia 11, 72
Glyceria 116
Gnaphaliatum 116
Heteromeles 13, 77, 78, 79, 80
Hevea 12, 55
Hippophaë 111
Hordeum 30
Jasminum 113
Juglans 43
Juncus 12, 57, 59
Kageneckia 13, 79, 80, 81
Lathyrus 12, 60
Limonium 118
Linum 111
Livistonia 117
Lomatium 11, 72
Lonicera 111
Magnolia 114
Malus 3, 13, 18, 20, 76, 78, 79, 80, 82, 84,
85, 112
Metrosideros 30
Olea 12, 70, 71
Ophiopogon 118
Parahebe 15, 100, 102
Persica 26
Peucedanum 11, 72, 113, 117
Phaseolus 111
Phillyrea 12, 53, 73, 74
Photinia 13, 77, 79
Phragmites 117
Pimpinella 117
Pisum 12, 74, 76
Populus 4, 14, 15, 43, 45, 53, 57, 62, 64, 65,
85, 86, 87, 88, 89, 90, 91, 92, 94, 99, 100
Potentilla 30
Prunus 13, 26, 29, 30, 33, 34, 68, 70, 79,
80, 107, 108
Psoralea 12, 81, 82
Pyracantha 13, 40, 78, 79, 80, 81
132
Pyrus 3, 14, 17, 18, 65, 67, 76, 79, 80, 82,
84, 85, 112
Quercus 12, 105
Robinia 118
Rosa 30
Rubus 14, 54
Salix 15, 30, 31, 92, 93, 94, 108, 109
Scabiosa 36
Scilla 12, 94, 96
Scirpus 3, 112
Sedum 11, 31
Setaria 112
Solidago 11, 102, 103
Sorbus 14, 76, 77, 78, 79, 80, 85
Sorghum 118
Sphaenosciadium 11, 72
Spiraea 14, 97, 98
Statice 118
Succisa 11, 36
Syringa 12, 40, 41
Thea 112
Tragopogon 117
Vanilla 112
Viburnum 81
Vitex 15, 103, 104
Vitis 117
Zizia 117
Schlechtendalia 9 (2003)
SCHUBERT, RITSCHEL & BRAUN: A monograph of Fusicladium s.lat.
133