Content uploaded by J.Z. Groenewald
Author content
All content in this area was uploaded by J.Z. Groenewald
Content may be subject to copyright.
© 2012 Nationaal Herbarium Nederland & Centraalbureau voor Schimmelcultures
You are free to share - to copy, distribute and transmit the work, under the following conditions:
Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work).
Non-commercial: You may not use this work for commercial purposes.
No derivative works: You may not alter, transform, or build upon this work.
For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be
waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights.
Persoonia 28, 2012: 126–137
www.ingentaconnect.com/content/nhn/pimj http://dx.doi.org/10.3767/003158512X652624
RESEARCH ARTICLE
INTRODUCTION
In his revision of coelomycetous fungi with appendage bear-
ing conidia, Nag Raj (1993) mentioned that in early papers
dealing with these fungi, descriptions of novel taxa only rarely
referred to the fact that conidia had appendages. In later years,
these appendages were seen as taxonomically informative
in separating species, especially in genera such as Coniella
(van Niekerk et al. 2004), Phyllosticta (Wulandari et al. 2009,
Glienke et al. 2011, Wikee et al. 2011), Pestalotiopsis (Maha-
rachchikumbura et al. 2011), Seimatosporium (Barber et al.
2011), and Tiarosporella (Crous et al. 2006). Sutton & Sellar
(1966) categorised appendages by their structure and position,
whether mucilaginous or cellular, endo- or exogenous, simple
or branched, as well as their position on the conidium. Nag Raj
(1993) further elaborated on these concepts, and defined nine
appendage types (A–I), with three subdivisions for append-
age type A. He further highlighted the diversity that exists in
appendage morphology, namely in shape (filiform, attenuated
or podiform), position (apical, basal, lateral, or in combination),
in patterns of distribution on the conidium, in branching, in
integrity with the conidium, in sequence of development, and
in structural changes (nucleate or not).
Appendages usually relate to an ecological function linked
to spore dispersal, and the colonisation of new substrates or
niches. Appendages play an important role in spore attachment
to substrates to ensure that the conidium can germinate and
its hyphae can infect or colonise the substrate (Nag Raj 1993).
Conidial appendages are known to occur in coelomycetes in
diverse habitats ranging from terrestrial to aquatic. Appendages
have subsequently been used to support the combination or
separation of taxa into different genera. One such genus is
Dinemasporium, which forms the basis of the present study. It
is characterised by superficial, cupulate to discoid conidiomata
with brown setae, and phialidic conidiogenous cells that give
rise to hyaline, oblong to allantoid, aseptate conidia with an
appendage at each end.
The genus Dinemasporium has several acknowledged syno-
nyms, namely Dendrophoma, Pycnidiochaeta, and Amphitia-
rospora (Sutton 1980, Nag Raj 1993). Sutton (1980) placed
species with septate conidia in Pseudolachnea, while Nag Raj
(1993) separated Pseudolachnea (1-septate conidia) from
Pseudolachnella (multiseptate conidia). Saccardo (1884) esta-
blished the subgenus Stauronema for taxa with conidia that had
apical, basal as well as lateral appendages. Sutton (1980) later
elevated Stauronema to generic level, a proposal subsequently
accepted by Nag Raj (1993).
The aim of the present study is to investigate the taxonomic
value of appendages as defining feature at generic level in
coelomycetous fungi, by using Dinemasporium as a case study.
A further aim is to clarify the phylogenetic position of the genus
Dinemasporium, and to revisit its circumscription in relation
to its synonyms and closely allied genera, many of which are
distinguished by a combination of conidium septation and ap-
pendage morphology.
How important are conidial appendages?
P.W. Crous1, 2, 3, G.J.M. Verkley1, M. Christensen4, R.F. Castañeda-Ruiz5,
J.Z. Groenewald1
Key words
Chaetosphaeriaceae
Dinemasporium
ITS
LSU
Sordariomycetes
systematics
Abstract The genus Dinemasporium is used as a case study to evaluate the importance of conidial appendages
for generic level classification of coelomycetous fungi. Based on morphology and sequence data of the large subunit
nuclear ribosomal RNA gene (LSU, 28S) and the internal transcribed spacers and 5.8S rRNA gene of the nrDNA
operon, the genus Dinemasporium is circumscribed, and an epitype designated for D. strigosum, the type of the
genus. A further five species are introduced in Dinemasporium, namely D. pseudostrigosum (isolated from Triticum
aestivum, Germany and Stigmaphyllon sagraeanum, Cuba), D. americana (soil, USA), D. polygonum (Polygonum
sachalinense, Netherlands), D. pseudoindicum (soil, USA), and D. morbidum (human sputum, Netherlands and
hare dung, New Zealand). Brunneodinemasporium, based on B. brasiliense, is introduced to accommodate Dinema-
sporium-like species with tightly aggregated brown conidiogenous cells, and pale brown conidia. Dendrophoma
(= Amphitiarospora) is reinstated as distinct from Dinemasporium, and an epitype designated for D. cytisporoides,
characterised by its superficial, stipitate to cupulate conidiomata, and small conidia with two polar, tubular, exogenous
appendages. The genus Stauronema is reduced to synonymy under Dinemasporium. Pseudolachnea (1-septate
conidia) is supported as distinct from Dinemasporium (aseptate conidia), and P. fraxini introduced as a novel spe-
cies. Taxa in this generic complex differ by combination of morphological characters of conidiomata, setae, conidia
and appendages. Appendage morphology alone is rejected as informative at the generic level.
Article info Received: 27 April 2012; Accepted: 20 May 2012; Published: 4 June 2012.
1 CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht,
The Netherlands; corresponding author e-mail: p.crous@cbs.knaw.nl.
2 Microbiology, Department of Biology, Utrecht University, Padualaan 8, 3584
CH Utrecht, The Netherlands.
3 Wageningen University and Research Centre (WUR), Laboratory of Phyto-
pathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands.
4 Department of Botany, 430 Lincoln Drive, University of Wisconsin, Madison,
WI 53706-1381, USA.
5 Instituto de Investigaciones Fundamentales en Agricultura Tropical “Alejan-
dro de Humboldt”, Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba,
C.P. 17200.
127
P.W. Crous et al.: Dinemasporium
MATERIALS AND METHODS
Isolates
The majority of the strains used in the present study were
obtained from the culture collection of the CBS-KNAW Fungal
Biodiversity Centre (CBS) Utrecht, the Netherlands. Fresh
collections were made from debris of diverse hosts by placing
material in damp chambers for 1–2 d. Single conidial colonies
were established from sporulating conidiomata on Petri dishes
containing 2 % malt extract agar (MEA; Crous et al. 2009b)
as described earlier (Crous et al. 1991). Colonies were sub-
cultured onto potato-dextrose agar (PDA), oatmeal agar (OA),
MEA, and pine needle agar (PNA) (Smith et al. 1996), and
incubated at 25 °C under continuous near-ultraviolet light to pro-
mote sporulation. Reference strains were deposited at the CBS.
DNA isolation, amplification and analyses
Genomic DNA was extracted from fungal colonies growing on
MEA using the UltraCleanTM Microbial DNA Isolation Kit (MoBio
Laboratories, Inc., Solana Beach, CA, USA) according to the
manufacturer’s protocol. The primers V9G (de Hoog & Gerrits
van den Ende 1998) and LR5 (Vilgalys & Hester 1990) were
used to amplify part (ITS) of the nuclear rDNA operon spanning
the 3’ end of the 18S rRNA gene, the first internal transcribed
spacer (ITS1), the 5.8S rRNA gene, the second ITS region
and the 5’ end of the 28S rRNA gene. The primers ITS4 (White
et al. 1990) and LSU1Fd (Crous et al. 2009a) were used as
internal sequence primers to ensure good quality sequences
over the entire length of the amplicon. The sequence alignment
and subsequent phylogenetic analyses for all the above were
carried out using methods described by Crous et al. (2006).
Gaps longer than 10 bases were coded as single events for
the phylogenetic analyses; the remaining gaps were treated as
‘fifth state’ data. Sequences derived in this study were lodged
at GenBank, the alignment in TreeBASE (www.treebase.org/),
and taxonomic novelties in MycoBank (www.MycoBank.org;
Crous et al. 2004).
Morphology
Morphological descriptions are based on slide preparations
mounted in clear lactic acid from colonies sporulating on PNA.
Observations were made with a Zeiss V20 Discovery stereo-
microscope, and with a Zeiss Axio Imager 2 light microscope
using differential interference contrast (DIC) illumination and
an AxioCam MRc5 camera and software. Colony characters
and pigment production were noted after 1 mo of growth on
MEA, PDA and OA (Crous et al. 2009b) incubated at 25 °C.
Colony colours (surface and reverse) were rated according to
the colour charts of Rayner (1970).
RESULTS
Phylogeny
Amplicons of approximately 1 700 bases were obtained of the
ITS region (including the first approximately 900 bp of LSU)
for the isolates listed in Table 1. The LSU sequences were
used to obtain additional sequences from GenBank, which
were added to the alignment (Fig. 1) and the ITS to determine
species identification (Fig. 2; discussed in species notes where
applicable). The manually adjusted LSU alignment contained
42 sequences (including the outgroup sequence) and 798
characters including alignment gaps (available in TreeBASE)
were used in the phylogenetic analysis; 151 of these were parsi-
mony-informative, 136 were variable and parsimony-uninform-
ative, and 511 were constant. Neighbour-joining analyses using
three substitution models on the sequence alignment yielded
trees with identical topologies to one another and support the
same terminal clades as obtained from the parsimony analy-
sis. The parsimony analysis of the LSU alignment yielded 142
equally most parsimonious trees (TL = 741 steps; CI = 0.533;
RI = 0.662; RC = 0.353). The manually adjusted ITS alignment
contained 31 sequences (including the outgroup sequence) and
514 characters including alignment gaps (available in Tree-
BASE) were used in the phylogenetic analysis; 203 of these
were parsimony-informative, 60 were variable and parsimony-
uninformative, and 251 were constant. Neighbour-joining analy-
ses using three substitution models on the sequence alignment
yielded trees with identical topologies to one another and sup-
port the same terminal clades as obtained from the parsimony
analysis. The parsimony analysis of the ITS alignment yielded
3 equally most parsimonious trees (TL = 929 steps; CI = 0.553;
RI = 0.706; RC = 0.390). All the species treated below are sup-
ported as distinct in the ITS phylogeny (Fig. 2).
Species Strain accession no.1 Substrate Country Collector(s) GenBank accession no.2
ITS LSU
Brunneodinemasporium brasiliense CBS 112007; INIFAT CO2/51 Decaying leaf Brazil R.F. Castañeda-Ruiz JQ889272 JQ889288
Dendrophoma cytisporoides CBS 223.95 Branches and twigs of Netherlands A. Aptroot JQ889273 JQ889289
Rhododendron
Dinemasporium americana CBS 127127; RMF 7507 A1 horizon soil of tallgrass USA:Iowa D.E. Tuthill JQ889274 JQ889290
prairie
Dinemasporium decipiens CBS 592.73 Soil under Elaeis guineensis Suriname J.H. van Emde JQ889275 JQ889291
Dinemasporium morbidum CBS 129.66 Sputum of man Netherlands N.G.M. Orie & A. Kikstra JQ889280 JQ889296
CBS 995.97; ATCC 200690 Hare dung New Zealand D.P. Mahoney JQ889281 JQ889297
Dinemasporium polygonum CBS 516.95 Polygonum sachalinense Netherlands A. Aptroot JQ889276 JQ889292
Dinemasporium pseudoindicum CBS 127402; RMF 8631 A1 horizon soil of tallgrass USA:Kansas M. Christensen JQ889277 JQ889293
prairie
Dinemasporium pseudostrigosum CBS 717.85 Triticum aestivum Germany P. Reinecke JQ889278 JQ889294
CBS 825.91; INIFAT C91/88-1 Stigmaphyllon sagraeanum Cuba R.F. Castañeda JQ889279 JQ889295
Dinemasporium strigosum CBS 520.78 Hay Netherlands M. Nieuwstad JQ889282 JQ889298
CBS 828.84 Leaf spot on Secale cereale Germany M. Hossfeld JQ889283 JQ889299
CPC 18898 Phragmites australis Netherlands W. Quaedvlieg JQ889284 JQ889300
CPC 18957 Malus USA:Illinois J. Batzer JQ889285 –
CPC 19796 Wildenovia incurvata South Africa A. Wood JQ889286 –
Pseudolachnea fraxini CBS 113701; UPSC 1833 Fraxinus excelsior Sweden K. & L. Holm JQ889287 JQ889301
1 ATCC: American Type Culture Collection, Virginia, USA; CBS: CBS Fungal Biodiversity Centre, Utrecht, The Netherlands; CPC: Culture collection of P.W. Crous, housed at CBS; INIFAT: Alexander
Humboldt Institute for Basic Research in Tropical Agriculture, Ciudad de La Habana, Cuba; RMF: Martha Christensen Soil Fungus Collection; UPSC: Uppsala University Culture Collection of
Fungi, Botanical Museum University of Uppsala, Uppsala, Sweden.
2 ITS: Internal transcribed spacers 1 and 2 together with 5.8S nrDNA; LSU: partial 28S nrDNA.
Table 1 Collection details and GenBank accession numbers of isolates for which novel sequences were generated in this study.
128 Persoonia – Volume 28, 2012
Saccharomyces cerevisiae Z73326
Infundibulomyces cupulata EF113979
Thozetella nivea EU825200
Rattania setulifera HM171322
Striatosphaeria codinaeophora AF466088
Tainosphaeria crassiparies AF466089
Chaetosphaeria luquillensis AF466074
CBS 112007 Brunneodinemasporium brasiliense
CBS 592.73 Dinemasporium decipiens
Ellisembia brachypus DQ408563
CBS 113701 Pseudolachnea fraxini
CBS 825.91 Dinemasporium pseudostrigosum
Chaetosphaeria preussii AF178561
Melanopsammella gonytrichii AF466085
Melanopsammella vermicularioides AF466087
Melanochaeta hemipsila AY346292
Melanochaeta aotearoae AF466082
Pyrigemmula aurantiaca HM241692
Lecythothecium duriligni AF261071
CBS 223.95 Dendrophoma cytisporoides
CPC 18898 Dinemasporium strigosum
CBS 520.78 Dinemasporium strigosum
CBS 717.85 Dinemasporium pseudostrigosum
CBS 828.84 Dinemasporium strigosum
Chaetosphaeria tropicalis AF466080
Chaetosphaeria conirostris AF466066
Chaetosphaeria fuegiana EF063574
Chaetosphaeria callimorpha AF466062
Codinaeopsis gonytrichoides AF178556
CBS 127402 Dinemasporium pseudoindicum
Chaetosphaeria innumera AY017375
Chloridium lignicola AF178544
Chaetosphaeria capitata AF466061
Umbrinosphaeria caesariata AF261069
Chaetosphaeria ovoidea AF064641
CBS 127127 Dinemasporium americana
Zignoella pulviscula AF466090
CBS 516.95 Dinemasporium polygonum
Zignoella pulviscula AF466091
Chaetosphaeria ciliata GU180637
CBS 995.97 Dinemasporium morbidum
CBS 129.66 Dinemasporium morbidum
10 changes
100
98
95
53
86
93
58
89
100
85
65
99
80
57
100
74 100
100
55
Taxonomy
Brunneodinemasporium Crous & R.F. Castañeda, gen. nov.
— MycoBank MB800158
Type species. Brunneodinemasporium brasiliense Crous.
Etymology. Similar to Dinemasporium, but conidiogenous cells and
conidia brown in colour.
Conidiomata stromatic, scattered or aggregated, superficial,
dark brown to black, cupulate, unilocular, globose, setose; basal
stroma of textura angularis. Setae abundant, brown to black,
simple, septate, subulate to cylindrical, unbranched, smooth,
thick-walled, multi-septate, arising randomly throughout basal
stroma. Conidiophores lining the basal stroma in a dense layer,
brown, septate, unbranched, cylindrical, thin-walled, smooth.
Conidiogenous cells integrated, determinate, phialidic with
conspicuous periclinal thickening at an attenuated apex, brown,
smooth, subcylindrical to lageniform. Conidia hyaline to pale
brown, aseptate, thin-walled, smooth, fusiform, gently curved
or straight, apex obtuse to subobtusely rounded, base truncate,
eguttulate or guttulate, with a single, cellular, unbranched,
flexuous, with tubular appendage at each end, separated by a
septum; basal appendage excentric.
Fig. 1 The first of 142 equally most parsimonious trees obtained from a
heuristic search with 100 random taxon additions of the LSU sequence align-
ment. The scale bar shows 10 changes, and bootstrap support values from
1 000 replicates are shown at the nodes. The species treated in this study
are located in the coloured blocks. Branches present in the strict consensus
tree are thickened and the tree was rooted to a sequence of Saccharomyces
cerevisiae (GenBank accession Z73326).
129
P.W. Crous et al.: Dinemasporium
Cephalotheca sulfurea AB278194
Pyrigemmula aurantiaca HM241692
CBS 592.73 Dinemasporium decipiens
CBS 113701 Pseudolachnea fraxini
CBS 112007 Brunneodinemasporium brasiliense
Thozetella havanensis EF029184
CBS 129.66 Dinemasporium morbidum
CBS 995.97 Dinemasporium morbidum
CBS 516.95 Dinemasporium polygonum
CBS 223.95 Dendrophoma cytisporoides
Codinaeopsis gonytrichoides AF178556
Thozetella acerosa AY330996
CBS 127402 Dinemasporium pseudoindicum
Chaetosphaeria hebetiseta AF178549
Chaetosphaeria callimorpha AF178555
Menispora tortuosa AF178558
Menispora manitobaensis EU488738
Menispora ciliata EU488737
Zignoella pulviscula AF178543
Dictyochaeta fertilis AF178540
Dictyochaeta simplex EF029193
Thozetella falcata AY331004
Thozetella nivea EU825201
CBS 825.91 Dinemasporium pseudostrigosum
CBS 717.85 Dinemasporium pseudostrigosum
CBS 127127 Dinemasporium americana
CBS 520.78 Dinemasporium strigosum
CBS 828.84 Dinemasporium strigosum
CPC 18957 Dinemasporium strigosum
CPC 18898 Dinemasporium strigosum
CPC 19796 Dinemasporium strigosum
25 changes
100
91
100
87
88
52
100
100
100
100
52
99
69
67
54
100
99
86
100
Pyrigemmula
Dinemasporium
Pseudolachnea
Thozetella
Brunneodinemasporium
Dendrophoma
Chaetosphaeria
Codinaeopsis
Dictyochaeta
Menispora
Zignoella
Brunneodinemasporium brasiliense Crous & R.F. Casta ñeda,
sp. nov. — MycoBank MB800159; Fig. 3
Etymology. Named after the country from where this fungus was collected,
Brazil.
Conidiomata stromatic, scattered or aggregated, superficial,
dark brown to black, cupulate, unilocular, globose, up to 600
μm diam, setose with a central buff conidial mass on PNA;
basal stroma of textura angularis. Setae abundant, brown to
black, simple, septate, subulate to cylindrical, with obtuse apex,
unbranched, smooth, thick-walled, multi-septate, 200–1300 μm
long, 12–15 μm diam at base, 5–10 μm diam at obtuse apex,
arising from basal stroma. Conidiophores lining the basal stroma
in a dense layer, brown, 4–6-septate, unbranched, cylindrical,
thin-walled, smooth, 40–70 × 3 – 4 μm. Conidiogenous cells
integrated, determinate, phialidic with clearly visible periclinal
thickening at apex 1.5–2 µm diam, brown, smooth, subcylin-
drical to lageniform, 10–16 × 3–4 μm. Conidia hyaline to pale
brown, aseptate, thin-walled, smooth, fusiform, gently curved
or straight, apex obtuse to subobtusely rounded, base truncate,
eguttulate or guttulate, (17–)18–19(– 20) × (2–)2.5 – 3 μm, with
a single, unbranched, flexuous, tubular appendage at each
end, 5 –7 μm long, apparently separated by a septum; basal
appendage excentric.
Culture characteristics — Colonies flat, spreading, with sparse
to moderate aerial mycelium. On MEA smoke grey (surface), with
even, lobate margins, surrounded by a zone of diffuse, red pig-
ment; reverse olivaceous grey in centre, outer region red due to
diffuse pigment; on OA iron grey with even, lobate margin; on
PDA surface isabelline, in outer region smoke grey, in reverse
grey olivaceous.
Fig. 2 The first of three equally most parsimonious trees obtained from a heuristic search with 100 random taxon additions of the ITS sequence alignment.
The scale bar shows 25 changes, and bootstrap support values from 1 000 replicates are shown at the nodes. Genera are indicated in the coloured blocks and
the names of genera treated in this study are written in red. Ex-type strains are printed in bold. Branches present in the strict consensus tree are thickened
and the tree was rooted to a sequence of Cephalotheca sulfurea (GenBank accession AB278194).
130 Persoonia – Volume 28, 2012
Specimen examined. Brazil, Corcorvado, on decaying leaf, 12 Oct. 2002,
R.F. Castañeda-Ruiz (holotype CBS H-20948, culture ex-type CBS 112007
= INIFAT CO2/51).
Notes — Brunneodinemasporium has randomly distributed
setae throughout the basal stroma, which differs from Dinema-
sporium, which has a densely aggregated layer of brown
conidiogenous cells with prominent periclinal thickening and
apical taper, and conidia that appear pale brown, and have
setae that separate from the conidia by a septum (though the
latter is not clear under the light microscope, but differs from
Dinemasporium s.str.).
Dendrophoma Sacc., Michelia 2: 4. 1880.
= Amphitiarospora Agnihothr., Sydowia 16: 75. 1962 (1963).
Type species. Dendrophoma cytosporoides Sacc.
Conidiomata stromatic, scattered to gregarious, superficial,
stipitate, globose and closed, becoming cupulate, unilocular,
dark brown to black; basal excipulum brown, of dense textura
intricata. Setae arising from the outer elements of excipulum, or
restricted to base of conidioma, sparse, subulate to subcylindri-
cal, apex blunt to acutely rounded, straight to curved, trans-
versely septate, dark brown, thick-walled, smooth. Conidio-
phores arising from conidiomatal cavity, septate, branched,
hyaline. Conidiogenous cells discrete, or integrated, terminal
and lateral, lageniform to subcylindrical, hyaline, thin-walled,
smooth. Conidia naviculate to botuliform, aseptate, hyaline, thin-
walled, smooth, with an unbranched cellular appendage at
each end.
Dendrophoma cytosporoides Sacc., Michelia 2: 4. 1880;
Fig. 4
Basionym. Phoma cytosporoides Sacc. (as ‘cytisporoides’), Michelia 1,
5: 522. 1879.
≡ Dinemasporium cytosporoides (Sacc.) B. Sutton, Trans. Brit. Mycol.
Soc. 48: 613. 1965.
= Amphitiarospora neottiosporoides Agnihothr., Sydowia 16: 75. 1962
(1963).
Conidiomata stromatic, scattered to gregarious, superficial,
stipitate, at first globose and closed, then opening to become
cupulate, up to 300 µm diam, unilocular, dark brown to black;
basal excipulum brown, of dense textura intricata. Setae aris-
ing from the outer elements of excipulum, or restricted to base
of conidioma, sparse, subulate to subcylindrical, apex blunt to
acutely rounded, straight to curved, 2–8-septate, dark brown,
thick-walled, smooth, 100–300 × 4–6 µm, apex 1–1.5 µm
wide. Conidiophores arising from conidiomatal cavity, septate,
branched, hyaline, up to 40 µm long. Conidiogenous cells
discrete or integrated, terminal and lateral, lageniform to sub-
cylindrical, frequently in terminal whorls, hyaline, thin-walled,
smooth, 6–15 × 1.5 µm. Conidia naviculate to botuliform, asep-
tate, hyaline, thin-walled, smooth, (3.5–)4 –5 (–6) × (1–)1.5 µm,
with an unbranched appendage at each end, 0.5–1 µm long.
Culture characteristics — Colonies flat, spreading, with
sparse aerial mycelium and even, lobate margins. On OA sur-
face grey olivaceous, reverse olivaceous grey; on PDA surface
brick to cinnamon, reverse dark brick.
Specimens examined. France, Rouen, L’ Abbee Letendre, on Ulmus
sp., designated as lectotype of Phoma cytosporoides by Sutton (1965),
IMI 111879 ex Herb. PAD. – netherlands, Utrecht, Soest, near hospital,
on Rhododendron, coll. A. Aptroot, isol. G. Verkley, 11 Nov. 1994 (epitype
designated here CBS H-12004, culture ex-epitype CBS 223.95).
Fig. 3 Brunneodinemasporium brasiliense (CBS 112007). a, b. Acervular conidiomata with setae; c. bases of setae; d. obtuse apices of setae; e, f. tightly
aggregated layer of conidiogenous cells; g. squash of conidiogenous cells giving rise to conidia; h, i. conidia. — Scale bars: b = 20 µm, all others = 10 µm;
c applies to d; e applies to f, g.
cb da
g h ie f
131
P.W. Crous et al.: Dinemasporium
Notes — The characteristic features that separate Dendro-
phoma from Dinemasporium are conidiomata that are superficial
and stipitate, becoming cupulate. The conidiogenous cells of
Dendrophoma form a dense layer that gives rise to conidia
with short, tubular exogenous appendages at each end, also
differing from Dinemasporium s.str.
Based on an examination of the holotype specimen, Sutton
(1977, 1980) regarded Amphitiarospora as synonymous with
Dinemasporium. The original description (conidia 4– 5 × 1.5– 2
µm, setae 1–2 µm; Agnihothrudu 1962), indicated that A. neot-
tio sporoides is synonymous with D. cytosporoides.
Dinemasporium Lév., Ann. Sci. Nat., Bot., sér. 3, 5: 274. 1846.
= Dinemasporium (Lév.) subg. Stauronema Sacc., Syll. Fung. 3: 686.
1884.
≡ Stauronema (Sacc.) Syd., P. Syd. & E.J. Butler, Ann. Mycol. 14: 217.
1916.
= Pycnidiochaeta Sousa da Câmara, Agron. Lusit. 12: 109. 1950.
Type species. Dinemasporium strigosum (Pers.: Fr.) Sacc.
Conidiomata stromatic, cupulate, often discoid, superficial (also
on SNA in culture), unilocular, setose, black, with basal stroma
of textura angularis. Setae arise from the basal stroma and/
or from excipular margin, unbranched, subulate to cylindrical,
straight or curved, brown to dark brown, smooth or verruculose,
with obtuse to acute apices. Conidiophores lining the inner
cavity, mostly branched, septate, hyaline (at times brown at
base), smooth, invested in mucus. Conidiogenous cells discrete
or integrated, lageniform, subcylindrical or cylindrical, hyaline,
smooth, with visible periclinal thickening. Conidia fusiform, navi-
culate or allantoid, aseptate, hyaline to pale brown, smooth,
with a single, unbranched, filiform, cellular appendage at each
end (not separated from body via septa); lateral appendages
present or absent.
Dinemasporium americana Crous & Tuthill, sp. nov. — Myco-
Bank MB800160; Fig. 5
Etymology. Named after its country of origin, Unites States of America.
Conidiomata stromatic, scattered or aggregated, superficial,
dark brown to black, cupulate, unilocular, globose, up to 350
μm diam, setose with a central buff to rosy buff conidial mass
on PNA; basal stroma of textura angularis, layer 20–30 µm
thick. Setae brown to black, simple, septate, subulate with
acute apex, unbranched, smooth, thick-walled, up to 7-septate,
50–200 × 4–10 μm, 1–1.5 µm wide at acute apex, arising from
basal stroma or lateral from excipulum. Conidiophores lining the
basal stroma, prominently septate (–8), branched, cylindrical,
thin-walled, smooth, base pale brown, apex hyaline, up to 70 µm
long. Conidiogenous cells determinate, phialidic with periclinal
thickening, hyaline, smooth, subcylindrical, 7–10 × 2–2.5 µm.
Conidia hyaline, aseptate, thin-walled, smooth, naviculate to
fusiform or ellipsoid, gently curved or straight, apex obtuse to
subobtusely rounded, base truncate, eguttulate or guttulate,
(9–)12–13(–16) × (2.5 –) 3 μm, with a single, unbranched, fle-
xuous, tubular appendage at each end, (11–)12–14(–16) µm;
basal appendage excentric.
Culture characteristics — Colonies spreading, erumpent,
with moderate aerial mycelium. On MEA buff (surface), reverse
cinnamon; on OA and PDA buff (surface and reverse).
Specimen examined. USA, Iowa, Cherokee County, Steele Prairie, A1
horizon soil, 1985, D.E. Tuthill, T93N R40W S16 (holotype CBS H-20949,
dried culture of RMF 7507 = CBS 127127).
Notes — Isolate RMF 7507 was formerly treated as D. stri-
gosum, but is specifically distinct. Although the conidiomata
produced are similar in general morphology to those of D. strigo-
sum, they differ by having shorter setae, larger conidia, longer
appendages (setae 60–400 μm long, conidia (9–)10–12(–13)
μm long, appendages 6–9 µm long in D. strigosum).
Dinemasporium morbidum Crous, sp. nov. — MycoBank
MB800161; Fig. 6
Etymology. Name reflects on the unpleasant substrata from which the
fungus was isolated.
Conidiomata stromatic, scattered or aggregated, superficial,
dark brown to black, cupulate, unilocular, globose, up to 250 μm
diam, setose with a central buff conidial mass on PNA; basal
stroma of textura angularis, layer 20–30 µm thick. Setae brown
to black, simple, septate, subulate with acute apex, unbranched,
smooth, thick-walled, up to 9-septate, 50–330 × 5 –10 μm,
1–1.5 µm wide at acute apex, arising from basal stroma or
lateral from excipulum. Conidiophores lining the basal stroma,
hyaline, septate, sparingly branched, cylindrical, thin-walled,
smooth, 30–40 µm long. Conidiogenous cells determinate,
phialidic with periclinal thickening, hyaline, smooth, subcylindri-
Fig. 4 Dendrophoma cytosporoides (CBS 223.95). a. Stalked acervuli forming in culture; b–d, f. conidiogenous cells; e. setae; g, h. conidia. — Scale bars
= 10 µm, except e = 20 µm.
cb da
g he f
132 Persoonia – Volume 28, 2012
cal to lageniform, 8–13 × 2–2.5 µm. Conidia hyaline, aseptate,
thin-walled, smooth, naviculate to fusiform or ellipsoid, gently
curved or straight, apex obtuse to subobtusely rounded, base
truncate, eguttulate or guttulate, (10–)12–14(–15) × (2.5–)3
μm, with a single, unbranched, flexuous, tubular appendage
at each end, (6–)7–10(–12) µm; basal appendage excentric.
Culture characteristics — Colonies spreading, erumpent,
with moderate aerial mycelium. On MEA, OA and PDA buff (sur-
face), reverse buff with patches of cinnamon; sporulation rosy
buff.
Specimens examined. netherlands, from human sputum, 19 Nov. 1965,
N.G.M. Orie and A. Kikstra, No. A 29/9 (holotype CBS H-12014, culture
ex-type 129.66). – new zealand, South Island, Hobson’s Nature trail near
Highway 13, Arthur’s Pass, on hare dung, Mar. 1995, D.P. Mahoney, CBS
995.97 = ATCC 200690.
Notes — Dinemasporium morbidum has longer conidia and
appendages than D. strigosum, which has conidia (9–)10–12
(–13) μm long, with appendages 6–9 µm long.
Fig. 5 Dinemasporium americana (CBS 127127). a, b. Acervular conidiomata forming in culture; c–e. setae and conidiogenous cells; f. conidiogenous cells;
g, h. conidia. — Scale bars = 10 µm; e applies to f.
cb da
g he f
Fig. 6 Dinemasporium morbidum (CBS 129.66). a, b. Acervular conidiomata forming in culture; c. setae. d, e. conidiogenous cells; f, g. bases and apices of
setae; h, i. conidia. — Scale bars = 10 µm, except a = 20 µm.
cb da
g h ie f
133
P.W. Crous et al.: Dinemasporium
Dinemasporium polygonum Crous & Verkley, sp. nov. —
MycoBank MB800162; Fig. 7
Etymology. Named after the host genus from which it was isolated,
Polygonum.
Conidiomata stromatic, scattered or aggregated, superficial,
dark brown to black, cupulate, unilocular, globose, up to 250
μm diam, setose with a central buff to rosy buff conidial mass
on PNA; basal stroma of textura angularis, layer 20–30 µm
thick. Setae brown to black, simple, septate, subulate with
acute apex, unbranched, smooth, thick-walled, up to 9-septate,
100–300 × 5– 8 μm, 1–1.5 µm wide at acute apex, arising
from basal stroma or lateral from excipulum. Conidiophores
lining the basal stroma, hyaline, septate, sparingly branched,
cylindrical, thin-walled, smooth, 30–60 µm long. Conidiog-
enous cells determinate, phialidic with periclinal thickening,
hyaline, smooth, subcylindrical to lageniform, 7–16 × 2–2.5
µm. Conidia hyaline, aseptate, thin-walled, smooth, naviculate
to fusiform or ellipsoid, gently curved or straight, apex obtuse
to subobtusely rounded, base truncate, eguttulate or guttulate,
(9–)10–12(–13) × (2–)2.5 (– 3) μm, with a single, unbranched,
flexuous, tubular appendage at each end, (9–)10–12(–13) µm;
basal appendage excentric.
Culture characteristics — Colonies spreading, erumpent, with
moderate aerial mycelium. On MEA, OA and PDA buff (surface
and reverse).
Specimen examined. netherlands, Soest, de Stompert, on Polygonum
sachalinense, 22 Feb. 1995, coll. A. Aptroot, isol. G.J.M. Verkley (holotype
CBS H-20950, culture ex-type CBS 516.95).
Notes — Dinemasporium polygonum has conidia with longer
appendages than D. strigosum though shorter than in D. ameri-
canum (6–9 µm long in D. strigosum, 11–16 µm in D. ameri-
canum).
Dinemasporium pseudoindicum Crous & M. Chr., sp. nov.
— MycoBank MB800163; Fig. 8
Etymology. Named after its morphological similarity to Stauronema
indicum.
Conidiomata stromatic, scattered or aggregated, superficial,
dark brown to black, cupulate, unilocular, globose to oval,
irregular in outline, 300 – 400 μm diam, setose with a central
cream conidial mass on PNA; basal stroma of textura angularis,
layer 20–30 µm thick. Setae brown, simple, septate, subulate
with acute to obtuse apex, unbranched, smooth, thick-walled,
Fig. 7 Dinemasporium polygonum (CBS 516.95). a, b. Acervular conidiomata forming in culture; c. setae; d, e. conidiogenous cells; f, g. conidia. — Scale
bars = 10 µm, except a, b = 20 µm.
cb
da
g
e
f
Fig. 8 Dinemasporium pseudoindicum (CBS 127402). a, b. Acervular conidiomata forming in culture; c, d. setae; e. conidiogenous cells; f– h. conidia, showing
apical, lateral and basal appendages. — Scale bars = 10 µm, except a = 20 µm, applies also to b.
cb da
g he f
134 Persoonia – Volume 28, 2012
up to 4-septate, 100–400 × 4 – 8 μm, 1–1.5 µm wide at acute
apex, or 4 µm when obtuse at apex, arising from basal stroma
or lateral from excipulum. Conidiophores lining the basal
stroma, hyaline, but pigmented in basal part, septate, sparingly
branched, cylindrical, thin-walled, smooth, 20–40 µm long,
invested in mucus. Conidiogenous cells determinate, phialidic
with periclinal thickening, rarely indeterminate and proliferating
percurrently at apex, hyaline, smooth, subcylindrical to lageni-
form, 5–15 × 2–2.5 µm. Conidia hyaline, aseptate, thin-walled,
smooth, naviculate to fusiform or ellipsoid, gently curved or
straight, apex obtuse to subobtusely rounded, base truncate,
eguttulate or guttulate, (9–)10–12(–13) × (3–)3.5 (– 4) μm, with
a single, unbranched, flexuous, tubular appendage at each
end; basal appendage excentric; apical and basal appendages
9–22 µm long; lateral appendages 2, inserted 4–6 µm below
apex, 10–17 µm long.
Culture characteristics — Colonies spreading, erumpent,
with moderate aerial mycelium on OA, but with sparse aerial
mycelium on PDA and MEA. On MEA rosy buff (surface), re-
verse cinnamon with patches of buff; on OA buff; on PDA grey
olivaceous in centre, buff in outer region, and buff in reverse.
Specimen examined. USA, Kansas, near Manhattan, Konza Prairie
Research Natural Area, long-term ecological research site, A1 horizon soil,
tallgrass prairie, June 1987, M. Christensen (holotype CBS H-20951, culture
ex-type RMF 8631 = CBS 127402).
Notes — Morphologically similar to Stauronema indicum, but
distinct in that it has larger conidia and appendages of different
length, and at different positions in relation to the conidial apex
(conidia 7–13 × 2–3 µm, terminal appendages 6.5–17 µm,
lateral appendages 8–20 µm long, but 3–4 µm below apex;
Nag Raj 1993).
Dinemasporium pseudostrigosum Crous, sp. nov. — Myco-
Bank MB800164; Fig. 9
Etymology. Named after its morphological similarity to D. strigosum.
Conidiomata stromatic, scattered or aggregated, superficial,
dark brown to black, cupulate, unilocular, globose, up to 250
μm diam, setose with a central buff to rosy buff conidial mass
on PNA; basal stroma of textura angularis, layer 20–30 µm
thick. Setae brown to black, simple, septate, subulate with
acute apex, unbranched, smooth, thick-walled, up to 7-septate,
100–250 × 5– 8 μm, 1–1.5 µm wide at acute apex, arising
from basal stroma or lateral from excipulum. Conidiophores
lining the basal stroma, hyaline, septate, sparingly branched,
cylindrical, thin-walled, smooth, 30–40 µm long. Conidiogenous
cells determinate, phialidic with periclinal thickening, hyaline,
smooth, subcylindrical to lageniform, 10–15 × 2–2.5 µm.
Conidia hyaline, aseptate, thin-walled, smooth, naviculate to
fusiform or ellipsoid, gently curved or straight, apex obtuse to
subobtusely rounded, base truncate, eguttulate or guttulate,
(10–)12–13(–14) × (2.5–)3 μm, with a single, unbranched,
flexuous, tubular appendage at each end, (11–)13–15(–17)
µm; basal appendage excentric.
Culture characteristics — Colonies spreading, erumpent,
with moderate aerial mycelium. On MEA buff (surface), reverse
cinnamon in middle, buff in outer region; on OA buff with profuse
sporulation (peach in colour); on PDA dirty white to buff (surface
and reverse).
Specimens examined. cuBa, Granma, on Stigmaphyllon sagraeanum,
14 Mar 1991, R.F. Castañeda, INIFAT C91/88-1 = CBS 825.91. – Germany,
Monheim, on Triticum aestivum, Sept. 1985, P. Reinecke, (holotype CBS
H-12018, culture ex-type CBS 717.85).
Notes — Morphologically D. pseudostrigosum is similar to
D. strigosum, but distinct in that it has larger conidia, and longer
appendages.
Dinemasporium strigosum (Pers.: Fr.) Sacc., Michelia 2:
281. 1881; Fig. 10
Basionym. Peziza strigosa Pers.: Fr., Syst. Mycol. 2: 103. 1823.
≡ Polynema strigosa (Pers.: Fr.) Fuckel, Jahrb. Nassauischen Vereins
Naturk. 23– 24: 367. 1870.
= Vermicularia graminum Lib., Pl. Crypt. Arduenna, fasc. (Liège) 4: no.
348. 1837.
≡ Excipula graminum (Lib.) Corda, Icon. Fungorum (Prague) 3: 29. 1839.
≡ Dinemasporium graminum (Lib.) Lév., Ann. Sci. Nat., Bot., ser. 3, 5:
274. 1846.
= Excipula betulae Fuckel, Jahrb. Nassauischen Vereins Naturk. 15: 64.
1860.
≡ Polynema betulae (Fuckel) Fuckel, Jahrb. Nassauischen Vereins
Naturk. 23– 24: 367. 1870.
≡ Dinemasporium betulae (Fuckel) Sacc., Syll. Fung. 3: 686. 1884.
= Dinemasporium longisetum Syd., Ann. Mycol. 34, 4/5: 400. 1936.
= Ellisiellina biciliata Sousa da Câmara, Agron. Lusit. 11: 72. 1949.
= Pycnidiochaeta biciliata Sousa da Câmara, Agron. Lusit. 12, 1: 109.
1950.
For additional synonyms see Sutton (1980) and Nag Raj (1993).
Conidiomata stromatic, scattered or aggregated, superficial,
dark brown to black, cupulate, unilocular, globose, up to 250 μm
Fig. 9 Dinemasporium pseudostrigosum (CBS 717.85). a, b. Acervular conidiomata forming in culture; c, d. setae; e, f. conidiogenous cells; g, h. conidia. —
Scale bars = 10 µm, except a = 20 µm, applies also to b.
cb
d
a
g he f
135
P.W. Crous et al.: Dinemasporium
diam, setose with a central buff to rosy buff conidial mass on
PNA; basal stroma of textura angularis, layer 20–30 µm thick.
Setae brown to black, simple, septate, subulate with acute
apex, unbranched, smooth, thick-walled, up to 9-septate,
60–400 × 7–10 μm, 1–1.5 µm wide at acute apex, arising
from basal stroma or lateral from excipulum. Conidiophores
lining the basal stroma, hyaline, septate, sparingly branched,
cylindrical, thin-walled, smooth, 30–40 µm long. Conidiogenous
cells determinate, phialidic with periclinal thickening, hyaline,
smooth, subcylindrical to lageniform, 12–15 × 2–2.5 µm.
Conidia hyaline, aseptate, thin-walled, smooth, naviculate to
fusiform or ellipsoid, gently curved or straight, apex obtuse to
subobtusely rounded, base truncate, eguttulate or guttulate,
(9–)10–12(–13) × (2–)2.5 (– 3) μm, with a single, unbranched,
flexuous, tubular appendage at each end, (6–)7–8(–9) µm;
basal appendage excentric.
Culture characteristics — Colonies spreading, erumpent,
with moderate aerial mycelium. On MEA buff (surface), rosy
buff (reverse); on OA buff with profuse sporulation (peach in
colour); on PDA buff (surface and reverse).
Specimens examined. BelGium, Ardenne, on dried grass blades, Libert,
Pl. Crypt. Arduennae, 1837, syntype of D. graminum, K(M) 175982. – Ger-
many, Baden-Werttemberg, Eberbach, on rotton wood of Betula sp., Fuckle’s
Fungi Rhenani exs 205, syntype of D. betulae, K(M) 175979; Brandenburg,
Freienwalde near Oder, on leaves of Populus tremula, H. Sydow Mycotheca
Germanica exs 2978, 30 Sept. 1934, syntype of D. longisetum, K(M) 175980;
Monheim, on Secale cereale, May 1984, M. Hossfeld, (epitype designated
here CBS H-20952, culture ex-epitype CBS 828.84). – netherlands, Beem-
ster, from hay, M. Nieuwstad, Oct. 1978, specimen CBS H-12008, culture
CBS 520.78; Wageningen, on Phragmites australis, 15 Nov. 2010, W.
Quaedvlieg, CPC 18898, 18899. – PortuGal, Ribatejo, on leaves of Dactylis
hispanica, Garcia Cabral 307, 30 Aug. 1950, holotype of Ellisiellina biciliata
LISE 41853; Serra do Geres, on culms of Agrostis castellana, 9 July 1948,
M.R. de J. Dias, holotype of Pycnidiochaeta biciliata, LISE 50037. – south
aFrica, Western Cape Province, Bracken Nature Reserve, on Wildenovia
incurvata, 18 Aug. 2011, A. Wood, CPC 19796, 19797. – sweden, on grass
blades, E. Fries, Scler. Suecicae exs 136, isotype of Peziza strigosa K(M)
175981.– USA: Illinois, Rockford, from Malus, J. Batzer, Sept. 2000, speci-
men CBS H-19787, culture CBS 18957.
Notes — Dinemasporium strigosum and its purported syno-
nyms have been treated by several authors (Webster 1955,
Sutton 1980, Nag Raj 1993, Yamaguchi et al. 2005, Duan et
al. 2007). The synonyms proposed by Sutton (1980) and Nag
Raj (1993) were accepted based on the examination of the type
material cited above.
Although D. strigosum was linked to a sexual state described
as Phomatospora dinemasporium (Webster 1955), the generic
placement was questioned by Rappaz (1992), as other spe-
cies of Phomatospora have been linked to hyphomycetes with
Fusarium-like conidia. No sexual states were encountered or
induced in the present study.
Pseudolachnea Ranoj., Ann. Mycol. 8: 593. 1910 [non Pseudo-
lachnea Velen. 1934].
= Dinemasporiella Bubák & Kabát, Hedwigia 52: 358. 1912; non Dinema-
sporiella Speg., Annales Mus. Nac. Buenos Aires 20: 366. 1910 (nom. dub.).
≡ Dinemasporiopsis Bubák & Kabát, nom. nov., apud Died., Krypt.-Fl.
Mark Brandenburg 9: 750. 1915.
= Chaetopatella Hino & Katum., J. Jap. Bot. 33: 238. 1958.
Type species. Pseudolachnea hispidula (Schrad.) B. Sutton (= P. bubakii
Ranoj.).
Conidiomata stromatic, scattered to gregarious, cupulate, super-
ficial, unilocular, setose, dark brown to black; basal stroma of
textura angularis. Setae divergent, subulate with blunt or acute
apices, unbranched, septate, thick-walled, smooth, dark brown.
Conidiophores arising from the uppermost cells of the basal
stroma and the inner cells of the excipulum, branched, septate,
brown at base, hyaline in upper part, smooth, invested in a thin
layer of mucus. Conidiogenous cells discrete, phialidic with
periclinal thickening, cylindrical, hyaline, smooth. Macroconidia
fusiform with obtuse ends, 1-septate, hyaline, smooth, guttulate,
bearing an unbranched, short, cellular, filiform appendage at
each end; appendages not delimited by septa, basal append-
age excentric. Microconidia intermixed with macroconidia in
same conidioma, fusiform, curved with rounded ends, aseptate,
smooth, guttulate, lacking appendages.
Pseudolachnea fraxini Crous, sp. nov. — MycoBank
MB800165; Fig. 11
Etymology. Named after the host from which it was collected, Fraxinus.
Conidiomata on OA stromatic, scattered to gregarious, cu-
pulate, superficial, unilocular, setose, dark brown to black,
up to 300 µm diam; basal stroma of textura angularis. Setae
divergent, subulate with blunt or acute apices, unbranched,
septate only at the base, thick-walled, smooth, dark brown,
40–170 × 4–7 µm. Conidiophores arising from the uppermost
Fig. 10 Dinemasporium strigosum (CBS 828.84). a–c. Acervular conidiomata forming in vivo; d, e. setae; f, g. conidiogenous cells; h, i. conidia. — Scale
bars = 10 µm, except a = 20 µm, applies also to b, c; d applies to e; f applies to g.
cb da
g h i
e
f
136 Persoonia – Volume 28, 2012
cells of the basal stroma and the inner cells of the excipulum,
sparsely branched and septate at the base, up to 20 µm long,
brown at base, hyaline in upper part, smooth, invested in a
thin layer of mucus. Conidiogenous cells discrete, phialidic
with periclinal thickening, cylindrical, hyaline, smooth, 6–12 ×
2–3 µm. Macroconidia fusiform with obtuse ends, 1-septate,
hyaline, smooth, guttulate, (12–)15–16(–18) × 2(–2.5) µm,
bearing an unbranched, short, cellular, filiform appendage at
each end, 1.5–2 µm long; appendages not delimited by septa,
basal appendage excentric. Microconidia common, intermixed
with macroconidia in same conidioma, fusiform, curved with
rounded ends, aseptate, smooth, guttulate, (6–)7–9 × (1–)1.5
µm, lacking appendages.
Culture characteristics — Colonies flat, spreading, with
sparse aerial mycelium and even, lobate margins. On MEA
dirty white (surface), reverse cinnamon; on OA surface honey
to isabelline, reverse honey; on PDA dirty white (surface and
reverse).
Specimen examined. sweden, Uppland, Dalby par., Jerusalem, on Fraxi-
nus excelsior, coll. K. & L. Holm, isol. O. Constantinescu, 31 Mar. 1986 (holo-
type CBS H-20953, culture ex-type CBS 113701).
Notes — The ex-type culture of P. fraxini (CBS 113701) was
originally identified as P. hispidula. Pseudolachnea fraxini differs
from P. hispidula by having much smaller conidiomata, shorter
setae, larger conidia (conidiomata 600–1200 µm diam, setae
80–220 µm long, conidia 11–14 µm long in P. hispidula; Nag
Raj 1993), and forms primarily microconidia in culture, an aspect
not yet observed in P. hispidula.
DISCUSSION
The present study examined the genus Dinemasporium with
the aim of evaluating the importance of conidial appendages as
taxonomic characters at generic level among coelomycetous
taxa. Dinemasporium has several acknowledged synonyms,
including Dendrophoma, Pycnidiochaeta, and Amphitiarospora.
Species with septate conidia were allocated to other genera
(Pseudolachnea, Pseudolachnella), while those with apical,
basal and lateral setae, were placed in Stauronema (Sutton
1980, Nag Raj 1993).
Based on these findings, a new genus, Brunneodinemasporium
is introduced to accommodate a Dinemasporium-like species
with tightly aggregated brown conidiogenous cells, and pale
brown conidia. Furthermore, the genus Dendrophoma (= Am-
phitiarospora) is reinstated as distinct from Dinemasporium.
Dendrophoma (which is based on D. cytosporoides) has super-
ficial, stipitate to cupulate conidiomata, and small coni dia with
two polar, tubular, exogenous appendages, which separates
it from Dinemasporium. The genus Stauronema is reduced
to synonymy under Dinemasporium, and lateral conidial ap-
pendages rejected as a useful single character for generic
separation. The genus Pseudolachnea (1-septate coni dia) is
supported as distinct from Dinemasporium (aseptate conidia),
though no cultures were available to determine if Pseudolach-
nella (multi-septate conidia) needs to be reduced to synonymy
with Pseudolachnea (Sutton 1980), or retained as separate
genus (Nag Raj 1993).
The genus Dinemasporium is a phylogenetically well-defined
genus in the Chaetosphaeriaceae. Dinemasporium is circum-
scribed based on D. strigosum, which is also epitypified in this
study. Five novel species are described from a collection of
cultures formerly treated as D. strigosum. Taxa in this complex
appear to differ by a combination of morphological features of
the setae, conidia and conidial appendages.
Other than revising taxa in the Dinemasporium complex, a
major aim of the present study was to determine the relevance
of appendages as delimiting feature at generic level in coelo-
mycetous fungi. Conidial appendages have been used exten-
sively to support the separation of species (e.g. Coniella, Phyllo-
sticta, van Niekerk et al. 2004, Glienke et al. 2011), but have
also been used to separate genera of coelomycetous fungi
(e.g. Discosia, Pestalotiopsis, Seimatosporium, Tiarosporella,
Crous et al. 2006, Barber et al. 2011, Maharachchikumbura et
al. 2011, Tanaka et al. 2011).
Results from the present study revealed that conidial append-
ages in coelomycetes are not informative as generic characters
when looked at in isolation, as recently reported for the Seima-
tosporium complex (Barber et al. 2011). Although appendage
morphology appeared to be highly informative at the species
level (mucilaginous or cellular, endo- or exogenous, simple
or branched, as well as their position on the conidium body),
they appeared to be less informative when used in isolation as
generic feature. We can conclude that genera separated chiefly
on the basis of appendage morphology, e.g. Dinemasporium
Fig. 11 Pseudolachnea fraxini (CBS 113701). a, b. Acervular conidiomata forming in culture; c, d. setae; e, f. conidiogenous cells; g. h. macro- and micro-
conidia. — Scale bars = 10 µm, except a = 20 µm, applies also to b.
cb da
g he f
137
P.W. Crous et al.: Dinemasporium
and Stauronema, or Seimatosporium and Vermisporium (Barber
et al. 2011), represent only two genera, namely Dinemasporium
and Seimatosporium. This finding could also have implications
for other more commonly encountered genera, e.g. Pestalotiop-
sis and Pestalotia, which are also separated solely on the basis
of apical appendage morphology. Pestalotia (1839) predates
Pestalotiopsis (1949), which will have serious implications for
names in Pestalotiopsis, which is presently the more commonly
used genus.
Acknowledgements We thank the technical staff, Arien van Iperen (cul-
tures), Marjan Vermaas (photo plates), and Mieke Starink-Willemse (DNA
isolation, amplification and sequencing) for their invaluable assistance.
REFERENCES
Agnihothrudu V. 1962. Notes on fungi from North-east India XIV – A new
genus of Discellaceae from Assam. Sydowia 16: 73–76.
Barber PA, Crous PW, Groenewald JZ, Pascoe IG, Keane P. 2011. Reas-
sessing Vermisporium (Amphisphaeriaceae), a genus of foliar pathogens
of eucalypts. Persoonia 27: 90–118.
Crous PW, Gams W, Stalpers JA, Robert V, Stegehuis G. 2004. MycoBank:
an online initiative to launch mycology into the 21st century. Studies in
Mycology 50: 19– 22.
Crous PW, Schoch CL, Hyde KD, Wood AR, Gueidan C, et al. 2009a. Phylo-
genetic lineages in the Capnodiales. Studies in Mycology 64: 17–47.
Crous PW, Slippers B, Wingfield MJ, Rheeder J, Marasas WFO, et al. 2006.
Phylogenetic lineages in the Botryosphaeriaceae. Studies in Mycology
55: 235– 253.
Crous PW, Verkley GJM, Groenewald JZ, Samson RA (eds). 2009b. Fungal
Biodiversity. CBS Laboratory Manual Series 1. Centraalbureau voor Schim-
melcultures, Utrecht.
Crous PW, Wingfield MJ, Park RF. 1991. Mycosphaerella nubilosa a synonym
of M. molleriana. Mycological Research 95: 628– 632.
Duan J, Wu W, Liu XZ. 2007. Dinemasporium (coelomycetes). Fungal Diver-
sity 26: 205– 218.
Glienke C, Pereira OL, Stringari D, Fabris J, Kava-Cordeiro V, et al. 2011.
Endophytic and pathogenic Phyllosticta species, with reference to those
associated with Citrus Black Spot. Persoonia 26: 47–56.
Hoog GS de, Gerrits van den Ende AHG. 1998. Molecular diagnostics of
clinical strains of filamentous Basidiomycetes. Mycoses 41: 183–189.
Maharachchikumbura SSN, Guo LD, Chukeatirote E, Bahkali AH, Hyde KD.
2011. Pestalotiopsis – morphology, phylogeny, biochemistry and diversity.
Fungal Diversity 50: 167–187.
Nag Raj TR. 1993. Coelomycetous anamorphs with appendage-bearing
conidia. Mycologue Publications, Waterloo, Ontario.
Niekerk JM van, Groenewald JZ, Verkley GJM, Fourie PH, Wingfield MJ,
Crous PW. 2004. Systematic reappraisal of Coniella and Pilidiella, with
specific reference to species occurring on Eucalyptus and Vitis in South
Africa. Mycological Research 108: 283– 303.
Rappaz F. 1992. Phomatospora berkeleyi, P. arenaria and their Sporothrix
anamorphs. Mycotaxon 45: 323– 330.
Rayner RW. 1970. A mycological colour chart. Commonwealth Mycological
Institute, Kew.
Saccardo PA. 1884. Sylloge fungorum 3: 1–860
Smith H, Wingfield MJ, Crous PW, Coutinho TA. 1996. Sphaeropsis sapinea
and Botryosphaeria dothidea endophytic in Pinus spp. and Eucalyptus spp.
in South Africa. South African Journal of Botany 62: 86–88.
Sutton BC. 1965. Typification of Dendrophoma and a reassessment of D. ob-
scurans. Transactions of the British Mycological Society 48: 611–616.
Sutton BC. 1977. Coelomycetes VI. Nomenclature of generic names pro-
posed for Coelomycetes. Mycological Papers 141: 1–253.
Sutton BC. 1980. The Coelomycetes: Fungi imperfecti with pycnidia, acervuli,
and stromata. Commonwealth Mycological Institute, Kew, Surrey.
Sutton BC, Sellar PW. 1966. Toxosporiopsis n. gen., an unusual member of
melanconiales. Canadian Journal of Botany 44: 1505–1513.
Tanaka K, Endo M, Hirayama K, Okane I, Hosoya T, Sato T. 2011. Phylogeny
of Discosia and Seimatosporium, and introduction of Adisciso and Immer-
sidiscosia genera nova. Persoonia 26: 85– 98.
Vilgalys R, Hester M. 1990. Rapid genetic identification and mapping of en-
zymatically amplified ribosomal DNA from several Cryptococcus species.
Journal of Bacteriology 172: 4238– 4246.
Webster J. 1955. Graminicolous pyrenomycetes V. Conidial state of Lepto-
sphaeria michotii, L. microspica, Pleospora vagans and the perfect states
of Dinemasporium graminum. Transitions of the British Mycological Society
38: 347–365.
White TJ, Bruns T, Lee J, Taylor SB. 1990. Amplification and direct sequenc-
ing of fungal ribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand
DH, Sninsky JJ, White TJ (eds), PCR protocols: a guide to methods and
applications: 315– 322. Academic Press, San Diego, California, USA.
Wikee S, Udayanga D, Crous PW, Chukeatirote E, McKenzie EHC, et al.
2011. Phyllosticta – an overview of current status of species recognition.
Fungal Diversity 51: 43– 61.
Wulandari NF, To-anun C, Hyde KD, Duong LM, Gruyter J de, et al. 2009.
Phyllosticta citriasiana sp. nov., the cause of Citrus tan spot of Citrus
maxima in Asia. Fungal Diversity 34: 23–39.
Yamaguchi Y, Masuma R, Tomoda H, Omura S. 2005. A new species of
Dinemasporium from sugar cane on Irabujima island, Japan. Myco science
46: 367–369.