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New insights into the systematics and phylogeny of the genus
Jattaea and similar fungi of the Calosphaeriales
Martina Réblová
#Kevin D. Hyde 2011
Abstract The phylogenetic relationship of Jattaea and
Wegelina, two morphologically related genera of the
Calosphaeriales, is analyzed in two molecular analyses
using the Internal transcribed spacer region of the DNA
operon, and the nuclear large and small subunits of
ribosomal DNA combined with the second largest subunit
of RNA polymerase for a multigene analysis. For each data
set maximum likelihood, maximum parsimony and Bayesian
analyses were performed. Type and herbarium material of two
lectotype species Jattaea algeriensis and Wegelina discreta
were examined and correlated with results from in vitro
studies. The phylogenetic analyses show that both species are
congeneric. The name Jattaea is accepted to include species
with hyaline, allantoid to suballantoid ascospores in clavate,
stipitate asci without apical annulus borne on individual cells
on ascogenous hyphae and with phialophora-like anamorphs
produced in axenic culture. Twelve species are accepted in
Jattaea, the concept of the genus is outlined and a key to its
species is provided. The genus Weg e l i n a is reduced to its
synonymy. Molecular data also show that septation of
ascospores, a diagnostic feature used to separate calosphaeria-
like fungi into the genus Phragmocalosphaeria, does not
appear to be relevant in distinguishing genera in the
Calosphaeriales. Therefore, Jattaea includes species with
both one-celled and septate ascospores and Phragmocalos-
phaeria is accepted as its generic synonym. The genus
Jattaea and its phialophora-like anamorphs is compared with
other genera of the Calosphaeriales and the phylogenetically
related Togniniaceae. The revision of four species of
Ceratostomella, i.e., C. echinella,C.leptorrhyncha,C. mali
and C. microspora, revealed their striking morphological
similarities with the Calosphaeriaceae and the To gn in i a c e a e .
These are identified as species of Jattaea,Tog n i n i a and
Togniniella and the respective new combinations are
proposed.
Keywords Calosphaeriaceae .Conidiogenesis .
Phragmocalosphaeria .Phialophora-like anamorph .
Multigene phylogeny .Wegelina
Introduction
The Calosphaeriales is a phylogenetically and morphologi-
cally well-established order of perithecial ascomycetous
fungi with allantoid to suballantoid ascospores and charac-
teristic ascogenous hyphae, ascogenous cells and centrum,
considered unique among the ascomycetes. These fungi are
classified in two families, the Calosphaeriaceae and the
monotypic Pleurostomataceae (Réblová et al. 2004). This
group of fungi was for the first time recognized by Munk
(1957) as the family Calosphaeriaceae of the broadly
perceived Sphaeriales.Barr(1983) recognized the order
Calosphaeriales; however, no anamorphs were yet linked to
teleomorphs of the accepted genera. The concepts and
history of the order and the family were outlined by Barr
(1983,1985) and Barr et al. (1993). The distinctness of the
morphologically similar Togninia Berl. and its Phaeoacre-
monium W. Gams, Crous & M.J. Wingf. anamorphs from
Calosphaeria was later corroborated by molecular DNA and
morphological data and in vitro studies leading to the
exclusion of Togn in i a from the Calosphaeriales and the
description of the Togn i n i a c e ae (Réblová et al. 2004).
After the stromatic formerly calosphaeriaceous family
Graphostromataceae, associated with nodulisporium-like
anamorphs, was excluded in the Xylariales (Barr et al.
1993), the Calosphaeriales encompassed nine genera, eight
M. Réblová (*)
Department of Taxonomy, Institute of Botany,
Academy of Sciences,
CZ-252 43 Průhonice, Czech Republic
e-mail: reblova@ibot.cas.cz
Fungal Diversity (2011) 49:167–198
DOI 10.1007/s13225-011-0099-8
Received: 14 January 2011 / Accepted: 9 March 2011 / Published online: 22 April 2011
nonstromatic and one stromatic (Kirk et al. 2008). The
correct placement for only four of them could be confirmed
with molecular data, i.e., Calosphaeria Tul. & C. Tul.,
Jattaea Berl., Pleurostoma Tul. & C. Tul. and Togniniella
Réblová, L. Mostert, W. Gams & Crous (Damm et al. 2008;
Réblová et al. 2004; Vijaykrishna et al. 2004). Other genera
like Enchnoa Fr., Kacosphaeria Speg., Pragmocalosphaeria
Petr., Sulcatistroma A.W. Ramaley and Wegeli na Berl. await
to be studied with molecular tools.
The calosphaerialean taxa are relatively little known and
until recently, none of the species were studied in vitro.
Their perithecia are inconspicuous on the host, usually
collected ‘accidentally’together with other fungi on dying
branches and twigs still attached to the trees or shrubs or
they grow on decaying wood and bark in the litter.
Perithecia often develop solitarily or in circinate groups
on wood beneath the loosened periderm or in soft, decaying
wood. Recently, they were also isolated from wood of
branches and trunks of fruit trees showing canker symptoms
(Damm et al. 2008; Trouillas et al. 2010), a substratum
domain generally believed to be restricted to the morpho-
logically similar Togniniaceae (e.g. Mostert et al. 2006).
Berlese (1900) described Jattaea and Wegelina as
morphologically similar genera with hyaline, allantoid,
one-celled ascospores, clavate asci, which he distinguished
primarily by characters of perithecia and length of the neck;
papillate to short-beaked in the former taxon vs.cylindrical
elongate necks in the latter. Jattaea algeriensis Berl. and
Wegelina discreta Berl. were designated as the respective
lectotypes by Clements and Shear (1931). In both genera,
the perithecia are nonstromatic, with different degree of
immersion in the natural substratum, solitary, scattered or
gregarious, sometimes in small irregular or valsoid groups,
but they do not form large circinate configurations typical
of Calosphaeria. Although Jattaea was always considered
a separate, though closely related taxon to Calosphaeria,
Wegelina was subsequently reduced to synonymy of
Scoptria Nitschke (Barr 1985)orCalosphaeria (Barr
et al. 1993), but it was later reinstated and emended by
Barr (1998) to include species with hyaline to dull brown,
allantoid, suballantoid to oblong, one-celled or delicately
septate ascospores, asci with an amyloid to inamyloid
apical ring and perithecia with conspicuous, sometimes
sulcate necks. One new species and two new combinations
were introduced in the genus (Barr 1998). Wegelina sensu
Barr (1998) included a heterogeneous collection of various
species now placed in several genera. Based on molecular
and morphological data, Wegelina species with dull brown,
suballantoid ascospores and perithecia with sulcate necks,
i.e., W. grumsiniana (Kirschst.) V. Kumm., T. Richt. &
Schwik (Kummer et al. 2005) and W. polyporina M.E. Barr
(Barr 1998), were transferred to Ceratostomella and listed
in the synonymy of C. rostrata (Tode : Fr.) Sacc. (Réblová
2006). Wegelina species with hyaline, one-septate, sub-
allantoid to oblong ascospores in clavate, stipitate asci with
an inamyloid apical ring were transferred to Barbatosphaeria
Réblová (Réblová 2007). Wegelina subdenudata (Peck) M.E.
Barr with ellipsoidal pigmented ascospores does not fit the
present concept of Jattaea and is excluded from the genus
(still under revision). Wegelina discreta and other morpho-
logically related species, which follow Berlese’sgeneric
treatment (1900), have hyaline, allantoid to suballantoid,
one-celled ascospores and clavate, stipitate asci without an
apical annulus. These species are considered in this study.
Petrak (1923) introduced a monotypic genus Phragmoca-
losphaeria Petr., typified with P. piskorsii Petr., to accom-
modate calosphaeria-like fungi with septate ascospores.
Three Jattaea species, i.e., J. ceanothina (Peck) M. E. Barr,
J. leucospermi Marinc., M. J. Wingf. & Crous and J. villosa
(Nitschke) L. Mostert, also have septate ascospores. The
revision of herbarium material of other fungi and search in
the literature revealed that there are four other species, which
conform to the description of Phragmocalosphaeria and
would require re-evaluation, i.e., Calosphaeria polyblasta
Romell & Sacc., Calospora allantospora Ellis & Everh.,
Calospora suecica Rehm and Romellia cornina (Peck) M. E.
Barr. Four lignicolous species of Ceratostomella,whichdo
not match the generic concept of that genus (Réblová 2006),
i.e.,C.echinellaEllis & Everh., C.leptorrhyncha (Durieu &
Mont.) Sacc., C. mali Ellis & Everh. and C. microspora Ellis
& Everh., are remarkably similar in morphological characters
of perithecia, asci, ascospores and centrum to the members of
the Calosphaeriaceae and Togniniaceae, which are associated
with phialidic anamorphs in vitro. These species are revised
here.
To evaluate relationships of Jattaea,Phragmocalosphae-
ria,Wegelina, and the four Ceratostomella species resem-
bling the Calosphaeriales, the type and other herbarium
material of their species was examined and freshly collected
specimens were cultivated and studied in vitro. In order to
study the phylogenetic relationships of the taxa in question,
the DNA sequences were analyzed in two separate phyloge-
nies: the Internal transcribed spacer region of the DNA operon
(ITS; including ITS1, 5.8S and ITS2) and nuclear large and
small subunits ribosomal DNA (nc28S and nc18S rDNA)
combined with the second largest subunit of RNA polymerase
(RPB2) for a multigene analysis.
Materials and methods
Morphological observations of herbarium material
and fungal strains
The herbarium specimens examined and cultures studied
are listed under each species treated. Dried specimens were
168 Fungal Diversity (2011) 49:167–198
rehydrated in water; material was examined with an
Olympus SZX12 dissecting microscope and centrum
material (including asci, ascospores and paraphyses) was
mounted in Melzer’s reagent or 90% lactic acid. Hand
sections of the perithecium wall were studied. When
present, conidiophores, conidiogenous cells and conidia
were examined as microscope slide preparations mounted
in water, Melzer’s reagent or 90% lactic acid. All measure-
ments were made in Melzer’s reagent. Means ± standard
errors (s.e.) based on 25 measurements are given for
dimensions of asci, ascospores, conidia and conidiogenous
cells. Images were captured using differential interference
(DIC) or phase contrast (PC) microscopy using an Olympus
DP70 Camera operated by Imaging Software Cell* on an
Olympus BX51 compound microscope. Images were
processed with Adobe Photoshop CS4 Extended.
Cultures were grown on potato-carrot agar (PCA, Gams
et al. 1998). Colonies were examined after 7, 21 and 30 d at
25°C in the dark. Cultures are maintained at CBS-KNAW
Fungal Biodiversity Center, Utrecht, The Netherlands and
ICMP, International Collection of Microorganisms and
Plants, Landcare Research, Auckland, New Zealand and
in the Institute of Botany, Academy of Sciences, Průhonice,
Czech Republic.
DNA extraction, amplification and sequencing
DNA was isolated with an UltraClean Microbial DNA Kit
(MoBio Laboratories, Inc., Canada) using mycelium removed
from a PCA culture following the manufacturer’sprotocolfor
filamentous fungi. All PCR experiments were carried out using
a PTC-200 thermal cycler (MJ Research Inc., Watertown, MA,
U.S.A.). PCR reactions containing 2–4 mM MgSO
4
were
performed using Platinum Taq DNA polymerase High
Fidelity (Invitrogen, Carlsbad, CA, U.S.A.) in 25 μL volume
reactions. PCR conditions were as follows: for nc18S 2 min
at 94°C; 35 cycles of 30 s at 94°C, 30 s at 55°C and 150–
300 s at 68°C; for ITS and nc28S 2 min at 94°C; 35 cycles of
30 s at 94°C, 30 s at 55–60°C, and 165–270 s at 68°C; all
amplifications were concluded by incubation for 10 min at
68°C. Amplicons were purified using the UltraClean PCR
Clean-up Kit (MoBio Laboratories, Inc., Canada) following
the manufacturer’s directions. All nucleotide sequences were
obtained by the dideoxy chain-terminating method using
automated DNA sequencers ABI PRISM 3100 or ABI
PRISM 3130xl (Applied Biosystems, Foster City, CA, U.S.
A.). For PCR amplifications, the following primers were
used: for nc18S, NSSU131–NS24 (Kauff and Lutzoni 2002;
White et al. 1990); for nc28S, ITS5/NS5/LR0R–LR8 (White
et al. 1990; Vilgalys, unpublished: www.botany.duke.edu/
fungi/mycolab); for ITS NS5/ITS5–ITS4 (White et al. 1990);
for RPB2 fRPB2-5F–fRPB2-7cR (Liu et al. 1999). For
sequencing reactions, the following primers were used: for
nc18S, NSSU131, SR11R, SR7, SR7R, NSSU897R,
NSSU1088, NSSU1088R, NS6, NS24 (Gargas and Taylor
1992; Kauff and Lutzoni 2002; Spatafora et al. 1995;White
et al. 1990; Vilgalys, unpublished: as above); for nc28S
LR0R, LR3R, LR6, LR7, LR16, LR5 (Rehner and Samuels
1994; Vilgalys and Hester 1990; Vilgalys and Sun 1994), JS7
and JS8 (Landvik 1996); for ITS, ITS5 and ITS4 (White et al.
1990); for RPB2 fRPB2-5F, fRPB2-7cR, RPB2-980R and
RPB2-1014F (Reeb et al. 2004).
Sequences were edited using Sequencher 4.9 software
(Gene Codes Corp., Ann Arbor, MI, U.S.A.).
Sequence alignment
Accession numbers and isolate information for new ITS,
nc18S, nc28S rDNA and RPB2 sequences derived from
DNA extract of ascospore and/or conidial isolates are
collated in Table 1. The new sequences were aligned with
data retrieved from GenBank, mostly from studies pub-
lished by Spatafora et al. (2007) and Zhang et al. (2007).
Other DNA data of members of the Calosphaeriales were
published by Damm et al. (2008), Mostert et al. (2006),
Réblová (2006), Réblová and Mostert (2007), Réblová and
Štěpánek (2009) and Vijaykrishna et al. (2004). Accession
numbers of analysed sequences are given in the phylograms
(Figs. 1and 2).
All sequences were manually aligned in BioEdit 7.0.9.0
(Hall 1999) and two alignments for combined sets of
nc28S-nc18S-RPB2 and ITS (ITS1-nc5.8S-ITS2) were
constructed. Predicted models of the secondary structure
of the nc28S and nc18S molecules of Saccharomyces
cerevisiae Meyen ex E.C. Hansen (Gutell 1993; Gutell et
al. 1993; Wuyts et al. 2002) were used to improve decisions
on homologous characters. For the ITS alignment we used
the predicted models of the secondary structure designed
for species of Chaetosphaeria from the family Chaetos-
phaeriaceae (Réblová and Winka 2000). We analyzed the
first two thirds of the nc28S region, the whole nc18S
region, the whole ITS region and 5–7 domains of the RPB2
gene. All characters in the ITS alignment were included in
the analysis. Bases 1–74 were excluded from analyses of
the nc28S and nc18S alignments, and bases 1–40 of the
RPB2 alignment, because of the incompleteness of the 5′-
end of the majority of the available sequences of all three
genes. The lengths of the alignments (see Results) were
determined after introduction of gaps. The ITS alignment
and the combined alignment of all three genes are available
in TreeBase (no. 11155).
Phylogenetic analyses
Phylogenetic relationships of two newly investigated strains
of Wegelina discreta were examined using nc28S, nc18S
Fungal Diversity (2011) 49:167–198 169
rDNA and RPB2 sequences of species from 20 orders or
families of the Sordariomycetes and nc5.8S, ITS1 and ITS2
rDNA sequences of representatives of the Calosphaeriales
and Togniniaceae. For all analyses rooting was accom-
plished by the outgroup method (Nixon and Carpenter
1993). Togninia fraxinopennsylvanica (T.E. Hinds) Georg
Hausner, Eyjólfsd. & J. Reid, T. leptorrhyncha (Durieu &
Mont.) Réblová and T. minima (Tul. & C. Tul.) Berl. in the
family Togniniaceae were used as outgroup in the ITS
phylogeny, and two outgroup taxa, S. cerevisiae and
Vanderwaltozyma polyspora (van der Walt) Kurtzman in
the family Saccharomycetaceae (Saccharomycetes), were
used in the three-gene phylogeny.
Maximum likelihood, maximum parsimony and Bayesian
analyses were used to estimate the phylogenetic relationships
of W. discreta and other related taxa in the Calosphaeriales.
The three genes in the combined analysis (nc18S-nc28S-
RPB2) were tested for heterogeneity between data partitions
before combining them in a total-evidence analysis. We used
the partition homogeneity/incongruence-length difference
test implemented in PAUP (Swofford 2002)todetermineif
different partitions of the data have significantly different
signals. Because combining data with value P>0.01 generally
improves phylogenetic accuracy (Cunningham 1997)and
our data did not show significant heterogeneity (P=0.01), the
sequences were combined for the further analyses.
Maximum likelihood analysis (ML) was performed with
RAxML-HPC (Stamatakis et al. 2005) using a GTRMIX
model of evolution, which is a combination of GTRGAMMA
and GTRCAT (a RAxML-specific alternative model, in
which the alignment sites are pooled into a pre-specified
number of rate categories) models. The GTRCAT model is
used for the heuristic search and the best tree is then
optimized and the likelihood values are calculated under the
GTR+G model.
Table 1 Sources and accession numbers of isolates sequenced for this study
Teleomorph Source* Locality Substrate/Host GenBank accession numbers**
ITS LSU SSU RPB2
Barbatosphaeria
barbirostris
CBS 127690 Czech republic Wood of
Betula verruculosa
HQ878592 HQ878596 HQ878602
Jattaea algeriensis CBS 120871 South Africa Wood of
Prunus salicina
EU367446
a
U367457
a
EU367462
a
HQ878603
Jattaea discreta CBS 127681 France Wood of
Crataegus sp.
HQ878587 HQ878593 HQ878597
Jattaea discreta J. F. 06100 Italy Wood of
Quercus cerris
HQ878588 HQ878594
Jattaea
mookgoponga
CBS 120867 South Africa Wood of
Prunus persica
var. nucipersica
HQ878589 EU367458
a
EU367463
a
HQ878604
Natantiella
ligneola
CBS 123470 Belgium Wood of
Betula pendula
FJ617556
d
HQ878598 HQ878605
Pleurostoma
ootheca
CBS 115329 Thailand Wood HQ878590 AY249089
e
AY761074
e
HQ878606
Pleurostomophora
richardsiae
CBS 270.33 Sweden Wood AY179948
e
AY249089
e
AY761066
e
HQ878607
Tectonidula
hippocrepida
ICMP 17630 New Zealand Wood of
Nothofagus sp.
FJ617557
d
HQ878599 HQ878608
Togninia
fraxinopennsylvanica
CBS 128920 Czech Republic Wood of a
deciduous tree
HQ878591 HQ878595 HQ878600 HQ878609
Togninia minima CBS 111015 South Africa Stem of
Vitis vinifera
AF266647
b
AY761082
b
AY761068
b
HQ878610
Togninia vibratilis CBS 117115 France Wood of
Fagus sylvatica
DQ649065
c
HQ878601 HQ878611
* CBS = CBS-KNAW, Fungal Biodiversity Center, Utrecht, the Netherlands; ICMP = Landcare Research, Auckland, New Zealand. The culture J.
F. 06100 is no longer available
** Published sequences:
a
Damm et al. (2008),
b
Mostert et al. (2006),
c
Réblová and Mostert (2007),
d
Réblová and Štěpánek (2009),
e
Vijaykrishna et al.
(2004)
Fig. 1 Phylogram inferred from maximum likelihood analysis of a
combined data set (nc28S-nc18S-RPB2) using a GTRMIX model of
evolution. Maximum likelihood bootstrap (ML BP), maximum
parsimony bootstrap (MP BP) and Bayesian posterior probability
(PP) are indicated at the nodes. Thickened branches indicate posterior
probability values = 1.0 PP, 100% MP and 100% ML bootstrap
support. The GenBank accession numbers given in the tree after the
names are those of nc28S/nc18S/RPB2 genes. Missing sequences are
indicated by “–”
1 Fungal Diversity (2011) 49:167–19870
Fungal Diversity (2011) 49:167–198 171
0.2
Jattaea discreta HQ878594/-/-
Chaetosphaerella phaeostroma AY346274/-/AY780172
Coniochaeta ostrea DQ470959/DQ471007/DQ470909
Lentomitella crinigera AY761086/-/-
Togniniella microspora AY761076/AY761073/GU180660
Saccharomyces cerevisiae J01355/J01353/M15693
Calosphaeria pulchella AY761075/AY761071/GU180661
Lentomitella cirrhosa AY761085/-/-
Valsa ambiens AF362564/DQ862056/DQ862025
Pleurostomophora repens AY761078/AY761067/-
Apiognomonia errabunda AF408334/NG013199/DQ862014
Peethambara spirostriata AY489724/AY489692/EF692516
Hypocrea americana AY544649/AY544693/AF545561
Jattaea algeriensis EU367457/EU367462/HQ878603
Lulworthia grandispora DQ522856/DQ522855/DQ518181
Calosphaeria africana EU367455/EU367460/-
Pleurostoma ootheca AY249089/AY761074/HQ878606
Petriella setifera AY281100/U43908/DQ368640
Anthostomella torosa DQ836902/DQ836895/DQ836885
Diaporthe phaseolorum U47830/L36985/AY641036
Natantiella ligneola FJ617556/HQ878598/HQ878605
Jattaea leucospermi EU552127/-/-
Papulosa amerospora DQ470950/DQ470998/DQ470901
Lindra thalassiae DQ470947/DQ470994/DQ470897
Microascus trigonosporus DQ470958/DQ471006/AF107792
Ceratostomella pyrenaica DQ076323/-/-
Graphostroma platystoma DQ836906/DQ836900/DQ836893
Gelasinospora tetrasperma DQ470980/DQ471032/DQ470932
Reticulascus clavatus GU180622/GU180643/-
Jattaea discreta HQ878593/HQ878597/-
Tectonidula hippocrepida FJ617557/HQ878599/HQ878608
Leucostoma niveum AF408367/DQ862050/DQ862019
Lasiosphaeria ovina AF064643/AY083799/AY600292
Camarops microspora AY083821/DQ471036/DQ470937
Pseudonectria rousseliana U17416/AF543767/DQ522459
Jattaea mookgoponga EU367458/EU367463/HQ878604
Lignincola laevis U46890/AF050487/DQ836886
Coniochaeta discoidea AY346297/AJ875179/AY780191
Melanospora zamiae AY046579/AY046578/DQ368634
Vanderwaltozyma polyspora AY048169/X83825/EF599484
Sordaria fimicola AY780079/X69851/DQ368647
Cylindrotrichum setosum GU180617/GU180652/-
Melanospora tiffanii AY015630/AY01561/AY015637
Jobellisia luteola AY346286/-/-
Xylaria hypoxylon AY544648/AY544692/DQ470878
Camarops ustulinoides DQ470941/DQ470989/DQ470882
Barbatosphaeria barbirostris HQ878592/HQ878596/HQ878602
Corollospora maritima FJ176901/U46871/DQ368632
Ceratosphaeria lampadophora AY761084/AY761088/-
Magnaporthe grisea AB026819/DQ493955/-
Kylindria peruamazonensis GU180638/GU180609/GU180656
Togninia leptorrhyncha AY761081/AY761069/-
Glomerella cingulata DQ286199/M55640/DQ858455
Halosphaeria appendiculata AY090892/U46872/-
Chaetosphaeria ciliata GU180637/GU180614/GU180659
Jobellisia fraterna AY346285/-/-
Togninia vibratilis DQ649065/HQ878601/HQ878611
Melanconis stilbostoma DQ862043/DQ862054/DQ836890
Gibellulopsis nigrescens GU180613/GU180648/GU180664
Pleurostomophora richardsiae AY249089/AY761066/HQ878607
Ophiostoma piliferum AY281094/AJ243295/DQ470905
Ophiostoma stenoceras DQ836904/DQ836897/DQ836891
Chaetosphaeria curvispora GU180636/AY502933/GU180655
Bertia moriformis AY695261/-/AY780151
Plectosporium tabacinum GU180612/GU180647/GU180663
Monilochaetes infuscans GU180645/GU180619/GU180658
Gnomonia gnomon AF408361/DQ471019/DQ470922
Togninia minima AY761082/AY761068/HQ878610
Ceratostomella cuspidata FJ617558/-/-
Togninia fraxinopennsylvanica HQ878595 /HQ878600/HQ878609
84/-/1.0
90/99/0.9
99/99/1.0
98/99/1.0
80/-/1.0
100/90/1.0
100/81/1.0
78/79/0.8
98/93/1.0
98/73/1.0
100/-/1.0
61/-/0.8
100/93/0.9
78/55/0.9
84/79/0.9
86/100/0.9
59/85/0.8
83/82/1.0
96/82/1.0
98/67/1.0
92/-/1.0
100/99/1.0
99/85/1.0
100/78/1.0
98/95/1.0 100/86/1.0
95/90/1.0
90/-/1.0
99/100/1.0
61/55/1.0
61/72/0.8
92/65/1.0
97/68/1.0
98/60/1.0
72/79/0.9
98/83/1.0
99/99/1.0
100/99/1.0
100/95/1.0
Xylariales
Diaporthales
Jobellisiaceae
Calosphaeriales
Togniniaceae
Magnaporthaceae
Ophiostomatales
Papulosaceae
Coniochaetales
Sordariales
Chaetosphaeriales
Boliniales
Microascales
Hypocreales
Coronophorales
Melanosporales
Plectosphaerellaceae
Glomerellales
Lulworthiales
Saccharomycetales
52/61/1.0
-/-/0.8
calosphaeria-like fungi with
Ramichloridium, Sporothrix
anamorphs
calosphaeria-like fungi with
Nodulisporium anamorph
Maximum parsimony (MP) analyses were conducted
with PAUP 4.0b10 (Swofford 2002). A heuristic search was
performed with the stepwise-addition option with 1,000
random taxon addition replicates and TBR branch swap-
ping. All characters were unordered and given equal
weight. Gaps were treated as missing data. Branch support
was estimated on the recovered topologies by performing
1,000 bootstrap replicates with a full heuristic search,
consisting of ten random-addition replicates for each
bootstrap replicate.
Bayesian analysis (BAY) was performed in a likelihood
framework as implemented by MrBayes v3.0b4 software
package to reconstruct phylogenetic trees (Huelsenbeck and
Ronquist 2001). The program MrModeltest2 v. 2.3.
(Nylander 2008) was used to infer the appropriate substitu-
tion model, which would best fit the model of DNA
evolution for our sequence data sets. Multiple Bayesian
searches using Metropolis-coupled Markov chain Monte
Carlo sampling were conducted. One cold and three heated
Markov chains were used in the analysis. Bayesian analysis
was run for 5 million generations, with trees sampled every
1,000 generations. The first 20,000 trees, which represented
the burn-in phase of the analysis, were discarded. To estimate
posterior probabilities (PP) of recovered branches (Larget
Fig. 2 Phylogram inferred from maximum likelihood analysis of ITS
sequences using a GTRMIX model of evolution. Maximum likelihood
bootstrap (ML BP), maximum parsimony bootstrap (MP BP) and
Bayesian posterior probability (PP) are indicated at the nodes.
Thickened branches indicate posterior probability values= 1.0 PP,
100% MP and 100% ML bootstrap support. The strain numbers are
given in front of each name. The GenBank accession numbers given
in the tree after the names are those of ITS (ITS1-5.8S-ITS2)
1 Fungal Diversity (2011) 49:167–19872
0.07
CBS 120867 Jattaea mookgoponga HQ878589 ex-type
SS07 Calosphaeria pulchella HM237297
CBS 113726 Togniniella microspora EU367452 ex-type
CBS 270.33 Pleurostomophora richardsiae AY179948
STEU 6401 Jattaea mookgoponga EU367450
CBS 128920 Togninia fraxinopennsylvanica HQ878591
STEU 6399 Jattaea algeriensis EU367447
LM06 Calosphaeria pulchella HM237298
STEU 6182 Calosphaeria africana EU367444 ex-type
CBS 115329 Pleurostoma ootheca HQ878590
J. F. 06100 Jattaea discreta HQ878588
CBS 120871 Jattaea algeriensis EU367446 ex-type
HNDC06 Jattaea algeriensis GU361954
SM05 Calosphaeria pulchella HM237300
AMH9590 Togninia leptorrhyncha AY179945
CBS 119343 Jattaea leucospermi EU552127 ex-type
YNDC23 Jattaea algeriensis GU361945
CBS 631.94 Togninia minima AF266647
CBS 115999 Calosphaeria pulchella EU367451 ex-epitype
CBS 294.39 Pleurostomophora repens AF083195
CBS 127681 Jattaea discreta HQ878587 ex-epitype
CBS 113648 Togniniella microspora EU367453
DC04 Calosphaeria pulchella HM237299
YNDC19 Jattaea algeriensis GU361941
STEU 6400 Jattaea algeriensis EU367448
STEU 6181 Calosphaeria africana EU367445
50/97/0
99/97/0.7
98/92/1.0
76/60/0.9
100/71/1.0
95/96/1.0
100/99/0.9
62/74/71
99/100/0.8
Calosphaeriaceae Pleurostomataceae Togniniaceae
Jattaea
Calosphaeria
Togniniella
Pleurostoma
Calosphaeriales
and Simon 1999) 50% majority rule consensus trees were
created from the remaining trees using PAUP.
Results
In the first analysis, a combination of the nc28S, nc18S and
RPB2 sequences was assessed for 70 taxa representing 69
species in 20 ascomycetous orders or families of the
Sordariomycetes. The alignment consisted of 4,237 total
characters: 2,238 constant, 346 not parsimony-informative
and 1,653 parsimony-informative. A maximum parsimony
heuristic search produced twenty-four MPTs with a length of
12,561 steps (CI=0.282, RI=0.479, HI=0.717). The ML tree
isshowninFig.1. For the BAY analysis, the GTR+I+G
substitution model was inferred. In all three analyses,
the Calosphaeriales are shown as a well-supported group
(61% bootstrap support from MP/72% bootstrap support
from ML/0.8 posterior probability from BAY) and include
two strongly supported families, the Calosphaeriaceae (100/
100/1.0) and the Pleurostomataceae (100/100/1.0). Two
strains of Jattaea discreta are grouped with Jattaea algeriensis
(97/68/1.0) and other Jattaea species in a monophyletic
subclade (98/60/1.0) within the Calosphaeriaceae.Jattaea
leucospermi, representing the morphological pattern with
septate suballantoid ascospores, groups in a monophyletic
subclade with a strain of J. mookgoponga (92/65/1.0) among
other Jattaea species.
The second data set comprised an analysis of the ITS
region (ITS1-5.8S-ITS2). The alignment included 26
isolates representing 12 species in 3 families, the
Calosphaeriaceae,Pleurostomataceae and Togniniaceae,
and consisted of 603 total characters: 350 constant, 32 not
parsimony-informative and 221 parsimony-informative. A
maximum parsimony (MP) heuristic search produced two
most parsimonious trees (MPTs) with a length of 524 steps
(CI=0.730, RI=0.874, HI=0.269). The ML tree is shown
in Fig. 2. For the BAY analysis, the GTR+I+G substitution
model was inferred. In ML, MP and BAY analyses, the
two strains of Jattaea discreta grouped as a sister to Jattaea
algeriensis (without branch support) in a monophyletic well-
supported clade of the genus Jattaea (76/60/0.9). The
grouping of J. leucospermi with J. mookgoponga as its
clademate (98/92/1.0) within Jattaea is also confirmed in the
ITS phylogeny.
Taxonomy
Jattaea Berl., Icon. Fung. 3: 6. 1900.
=Wegelina Berl., Icon. Fung. 3: 8. 1900.
=Phragmocalosphaeria Petr., Annls. Mycol. 21: 109. 1923.
Perithecia nonstromatic, arranged singly, in small
irregular clusters or in loose valsoid configurations, usually
on wood beneath the periderm, sometimes associated with
dead ascomata of Diaporthales or Diatrypaceae,or
superficial on bark or decorticated decaying wood; venter
globose, subglobose to conical, dark brown to black,
sometimes covered with hyphal tomentum, with central
papilla or cylindrical, short to elongated neck; ostiolum
periphysate. Perithecial wall leathery to fragile, two-
layered; outer layer of pigmented cells of textura angularis
or prismatica; inner layer of subhyaline to hyaline,
flattened, elongated, thin-walled cells. Ascogenous hyphae
persistent, terminated in growth, discrete, short-branched,
aseptate, each branch sequentially and simultaneously
producing several lateral and terminal dehiscent cells, from
each of which an ascus arises as an outgrowth. Paraphyses
persistent, abundant, septate, hyaline, apically free, tape-
ring, longer than the asci. Asci clavate to oblong-clavate,
stipitate, obtuse to broadly rounded at the thickened apex,
tapering towards the base, withnodistinctdischarge
mechanism, floating freely within the centrum at maturity,
often with a short appendix remaining at the base after ascal
dehiscence from the ascogenous hyphae, 8-spored. Asco-
spores allantoid, suballantoid to oblong, hyaline, one-celled
or one- to several-septate, smooth, 2–3-seriate or fasciculate
in the ascus.
Anamorph: The anamorph does not regularly occur on
substratum near the perithecia. When the ascospores are
isolated in axenic culture, a phialophora-like anamorph is
produced. Anamorphs were formed in vitro by Jattaea
algeriensis,J. discreta,J. leucospermi and J. mookgoponga.
Typus:Jattaea algeriensis Berl.
Notes: Perithecia of Jattaea species are scarcely visible
on the surface, usually as tips of erumpent minute necks
often converging, but protruding separately. A tendency of
the perithecia to form small clusters, circular or valsoid
configurations beneath the periderm was observed in
several collections of J. discreta; but such formations occur
more frequently in Jattaea species with septate ascospores,
viz.J. allantospora,J. ceanothina and J. leucospermi. The
Jattaea species tend to be associated with rosaceous hosts
among other woody plants.
The length of the ascus stipe can vary significantly
within a single collection. Upon ascus maturation the
stipeelongatesandthesizeofthewholeascusincreases
accordingly; the size of the sporiferous part remains
constant. Thus, in one collection short-stipitate oblong
asci can occur together with long-stipitate clavate asci
clearly divided into a sporiferous part and elongated
tapering stipe. Therefore, it is important to measure asci
in pars sporifera because of the dynamic nature of the
stipe and its elasticity.
Fungal Diversity (2011) 49:167–198 173
Key to accepted Jattaea species
1. Ascospores aseptate 2
1. Ascospores septate 8
2. Perithecia papillate 3
2. Perithecia with cylindrical, elongated, sometimes flexu-
ous neck 4
3. Asci oblong-clavate, (21−)25–35(−48)×4.5–6μm, in pars
sporifera 19–32 μm long; ascospores suballantoid, 5–
7.5× 1.5–2μmJ. algeriensis
3. Asci clavate, (30−)33–40×7–8μm, in pars sporifera 21–
26 μm long; ascospores oblong, suballantoid to subcylin-
drical, 6–8×1.5(−2) μmJ. microtheca
4. Perithecial venter covered by a tomentum, neck glabrous,
dark brown to black 5
4. Perithecia glabrous, dark brown to black 6
5. Tomentum golden-brown; asci oblong-clavate, 18–25×
4–6μm; ascospores cylindric-oblong, 3–6×1.5–
2μmJ. stachybotryoides
5. Tomentum golden-yellow to golden-orange; asci
clavate, 30–35× 6–7μm, in pars sporifera
20–25 μm long; ascospores suballantoid, (7−)8–9×
(1.5−)2 μmJ. aurea
6. Asci in pars sporifera (12−)14–16(−18) μm long;
ascospores suballantoid to oblong, 5–6(−7)×1.5(−2)
μm; apex of perithecial neck red-brown in transmitted
light J. echinella
6. Asci longer in pars sporifera, apex of perithecial neck dark,
opaque in transmitted light 7
7. Asci clavate, 6.5–8μm wide, in pars sporifera 18–25
(−31) μm long; ascospores (6−)6.5–8×1.5–2μm;
perithecia usually in small irregular or valsoid
groups J. discreta
7. Asci oblong-clavate, (4.5−)5–5.5(−6) μm wide, in pars
sporifera (21−)23–27(−30) μm long; ascospores 6–7×
(1.5−)2 μm; perithecia usually scattered J. tumidula
8. Asci 12–15 μm wide, clavate; ascospores 3(−3.5) μm
wide, 3–7-septate J. ceanothina
8. Asci narrower 9
9. Asci 7–11 μm wide 10
9. Asci 5–7μm wide, 30–70 μm long, in pars sporifera
20–47(−55) μm; ascospores 0–3-septate, 8–11(−12)×
(1−)1.5–2μmJ. villosa
10. Ascospores (10−)12–14×2–2.5 μm, 1–3-septate; asci
48–68×(6−)7(−9) μmJ. allantospora
10. Ascospores shorter 11
11. Ascospores 1–3-septate, (8−)9–10.5(−11)× (1.5−)2 μm;
asci 34–36×7–8μmJ. leucospermi
11. Ascospores 1-septate, (8−)9–10.5× 2–2.5 μm; asci (36−)
40–50×(8−)9.5–11(−12) μmJ. pleurostoma
Jattaea algeriensis Berl., Icon. Fung. 3: 7. 1900.
(Figs. 3–5)
=Jattaea prunicola Damm & Crous, Persoonia 20: 45.
2008.
Perithecia scattered, solitary on wood beneath the thin
periderm; venter 200–300 μm diam, 190–220 μmhigh,
subglobose, dark brown to black, glabrous, slightly com-
pressed horizontally, papillate with sparse brown hairs in the
upper half attached to the periderm; ostiolum periphysate.
Perithecial wall leathery to fragile, two-layered, 24–28 μm
thick; outer layer of brown polyhedral cells of textura
angularis; inner layer of subhyaline to hyaline, flattened,
elongated, thin-walled cells. Cells on ascogenous hyphae
ellipsoidal to ovoidal, 2.5–3×3–4μm. Paraphyses 2.5–5μm
wide near the base. Asci (21−)25–35(−48)×4.5–6μm
(mean ± s.e.=25.4±0.6×5.4±0.1), in pars sporifera 19–
32 μm long, short-stipitate, oblong-clavate, broadly rounded
at the thickened apex, tapering towards the base, without a
distinct discharge mechanism, floating freely within the
centrum at maturity, with a short appendix remaining at the
base after ascal dehiscence from ascogenous hyphae, 8-
spored. Ascospores 5–7.5×1.5–2μm(mean±s.e.=7.1±
0.2×1.7±0.1), suballantoid, hyaline, aseptate, smooth, 2–3-
seriate in the ascus.
Anamorph: phialophora-like (Damm et al. 2008: 46,
Fig. 6)
Known distribution: Algeria, South Africa.
Habitat: Decayed herbaceous stem.
Material and cultures examined: ALGERIA, La Calle:
on decayed canes of Rubus fruticosus L. lying in the
brushwood, No. 43, MC 8938 (PC, holotype). SOUTH
AFRICA, Western Cape Province: Franschhoek, isolated
from brown necrosis in wood of Prunus salicina Lindl., 10
June 2004, U. Damm (living culture CBS 120871, ex-type
strain of J. prunicola).
Notes: The paraphyses were seen only as fragments in
the holotype collection. Damm et al. (2008) isolated J.
algeriensis (as prunicola) from wood of Prunus salicina
displaying necrotic symptoms; the perithecia and the
phialophora-like anamorph developed in axenic culture.
The anamorph formed cylindrical, elongate-ampulliform to
ampulliform phialides on aerial mycelium, while hyphae
bearing adelophialides were predominantly submerged in
the agar medium; conidia are suballantoid to oblong,
hyaline.
Jattaea algeriensis is similar to J. microtheca in having
papillate perithecia bearing sparse inconspicuous hyphae,
but it differs from it by shorter, narrower, oblong-clavate
asci with a short stipe. The asci of J. microtheca are
wider (7–8μm), distinctly clavate and stipitate, tapering
from the sporiferous part towards the base. For further
illustration of the teleomorph and anamorph refer to
Damm et al. (2008).
The type specimen of J. algeriensis (PC, MC 8938) is
the same as the type specimen of Sphaeria myriospora
1 Fungal Diversity (2011) 49:167–19874
Durieu & Mont. = Coronophora myriospora (Durieu &
Mont.) Sacc.
Jattaea allantospora (Ellis & Everh.) Réblová, comb.
nov. (Figs. 6–13)
MycoBank MB519576
≡Calospora allantospora Ellis & Everh., J.
Mycol. 9: 223. 1903 (basionym).
=Phragmocalosphaeria piskorsii Petr., Annls. Mycol.
21: 110. 1923.
Perithecia aggregated in groups of 6–10(−20) individu-
als on wood beneath the periderm; venter subglobose, dark
brown to black, glabrous, (350−)400–480 μmdiam,
440–500 μm high; necks central, cylindrical, elongate, ca
100 μm wide, up to 500 μm long, straight or slightly
flexuous, converging, at first decumbent to the substratum
then upright, piercing the periderm separately in a fissure in
the periderm; ostiolum periphysate. Perithecial wall lea-
thery to fragile, two-layered, 37–50 μm thick; outer layer of
brown polyhedral cells of textura prismatica; inner layer of
subhyaline to hyaline, flattened, elongated, thin-walled
cells. Cells on ascogenous hyphae ovoidal, 3–4.5×3–
5.5 μm. Paraphyses were seen only as ca 2μm wide
fragments. Asci 48–68×(6−)7(−9) μm (mean ± s.e.= 56.7 ±
2.8×7.1±0.2), in pars sporifera 37–48(−55) μm long
(mean ± s.e.=47.4±1.7), short-stipitate, oblong to oblong-
clavate, obtuse at the thickened apex, tapering towards the
base from the sporiferous portion, without a distinct
discharge mechanism, floating freely within the centrum
at maturity, with a short appendix remaining at the base
after ascal dehiscence from ascogenous hyphae, 8-spored.
Ascospores (10−)12–14× 2 μm (mean ± s.e.=12.3±0.6 × 2),
suballantoid to oblong, slightly curved, hyaline, 1–3-
septate, smooth, 2-seriate in the ascus.
Anamorph: Unknown
Material examined: CANADA, Ontario: London, on
decaying wood of Acer saccharinum L., 27 October 1903,
J. Dearness, Ellis & Everhart’s Fungi Columbiani no. 1912
(NY, isotype of Calospora allantospora).
Notes: The extreme case of stipe elongation can be seen
in J. allantospora, the variability in the stipe length is
conspicuous (Fig. 9). The majority of the asci, which are
attached to the ascogenous hyphae, are perfectly oblong
with a short stipe. However, some asci still attached can be
seen with already prolonging stipe as the process of stipe
prolongation has already started. The asci, which are
floating freely in the centrum, become distinctly clavate
with a prolonged stipe.
The type material of Phragmocalosphaeria piskorsii
[CZECH REPUBLIC, Moravia: Bělkovice-Lašťany,
decayed twig of Prunus spinosa (L.) Walt., leg. F. Petrak]
is not located in the W herbarium in Vienna (A. Igersheim,
Curator of Cryptogam Collections, pers. comm.). Petrak did
not specify the day or year of his collection. According to
the protologue (Petrak 1923), the species description
matches well that of J. allantospora. Therefore, P. piskorsii
is reduced to its synonymy.
Jattaea aurea Réblová & Fournier, sp. nov. (Figs. 14–27)
MycoBank MB 519577
Perithecia solitaria vel gregaria, immersa vel superficialia,
ventribus globosis usque subglobosis, 250–300 μmdiam,
Figs. 3–5Jattaea algeriensis.3,4. Asci. 5. Ascospores. 3–5from MC 8938 (PC, holotype). Scale bars: 3–5=10 μm
Fungal Diversity (2011) 49:167–198 175
240–300 μm altis, tomento aureo flavido et collis longis,
nigris, centralibus, rectis vel leviter flexuosis praedita. Paries
perithecii coriaceus, 25–32 μm crassus, bistratosus. Hyphae
ascogenae discretae. Paraphyses septatae, ad septa modice
constrictae, 4–4.5 μm latae prope basim, sursum ad 2–2.5 μm
angustatae, ascos superantes. Asci unitunicati, oblongo-
clavati, 30–35×6–7μm (in medio ± s.e.=32.3±0.6×6.1±
0.1), in parte sporifera 20–25 μm longi (in medio ± s.e.=23.2±
0.6), longit.:latit. 5.3, exhyphis ascogenis oriundi. Ascosporae
suballantoideae, (7−)8–9×(1.5−)2 μm (in medio ± s.e.= 8.3±
0.2×1.9±0.05), hyalinae, continuae, leves, in parte superiore
asci 2–3-seriate.
Etymology:aureus (L) golden, referring to the colour of
the golden-yellow tomentum covering the perithecial
venter.
Perithecia solitary or aggregated in groups, superficial
rarely semiimmersed in wood and bark; venter 250–
300 μm diam, 240–300 μm high, globose to subglobose,
covered by a golden-yellow to golden-orange tomentum;
necks black, central, elongate, 70–85 μm wide, 200–
Figs. 6–13 Jattaea allantospora.6. Longitudinal section of the perithecial wall. 7–11. Asci (arrow on Fig. 9indicate the length of the stipe). 12,13.
Ascospores. 6–13 from Ellis & Everhart’s Fungi Columbiani no. 1912 (NY, isotype). Scale bars: 6=50 μm; 7–11 = 10 μm; 12, 13=5 μm
1 Fungal Diversity (2011) 49:167–19876
Figs. 14–27 Jattaea aurea.14–16. Perithecia on the natural substra-
tum. 17. Longitudinal section of the perithecial wall. 18. Paraphyses.
19. Asci attached to the cells on ascogenous hyphae. 20–22.
Ascogenous hyphae and cells. 23–25. Asci with ascospores. 26,27.
Ascospores. 17–27 from PRM 917687 (holotype). Scale bars: 14–16=
500 μm; 17= 50 μm; 18–25 = 10 μm; 26, 27 =5 μm
Fungal Diversity (2011) 49:167–198 177
500 μm long, straight or slightly flexuous, upright
sometimes decumbent, tapering at the apex; ostiolum
periphysate. Perithecial wall leathery, two-layered, 25–
32 μm thick; outer layer of pale brown to red-brown
polyhedral cells of textura prismatica, externally with 5–
7.5 μm thick layer of reddish-brown cells and short hyphae
tightly interwoven, forming a thin stromatic layer around
the aggregated perithecia, inner layer of subhyaline to
hyaline, flattened, elongated, thin-walled cells. Cells of
ascogenous hyphae subglobose to ovoid, 4–5.5× 2.5–
3.5 μm. Paraphyses persistent, abundant, unbranched,
septate, hyaline, apically free, 4–4.5 μm wide near the
base, tapering to 2–2.5 μm, longer than the asci. Asci 30–
35×6–7μm (mean ± s.e.=32.3±0.6× 6.1 ± 0.1), in pars
sporifera 20–25 μm long (mean ± s.e.= 23.2 ± 0.6), short-
stipitate, oblong-clavate, obtuse at the apex, tapering
towards the base from the sporiferous portion; ascal wall
thickened only along the sides, discontinuous at the apex,
without a distinct discharge mechanism; asci floating freely
within the centrum at maturity, with a short appendix
remaining at the base after ascal dehiscence from ascoge-
nous hyphae, 8-spored. Ascospores (7−)8–9×(1.5−)2 μm
(mean ± s.e.=8.3±0.2×1.9 ± 0.05), suballantoid, hyaline,
aseptate, smooth, 2–3-seriate in the ascus.
Anamorph: Unknown.
Material examined: FRANCE, Pyrénées Atlantiques:
Ariège, Camarade, Les Moulins, 21 April 2002, on bark
of a dead stem of Coriaria myrtifolia L., J. Fournier J.F.
02068 (PRM 917687, holotype).
Notes: Cultivation attempts were not successful and the
ascospores did not germinate (water agar, water agar with
0.5% bovine serum albumin, PCA). The golden-yellow
tomentum on the exterior of the perithecial venter is a
conspicuous character and together with the size of asci and
ascospores distinguishes J. aurea from other Jattaea
species. The morphologically similar Jattaea stachybo-
tryoides (Romero and Samuels 1991) has a golden-brown
tomentum covering the perithecial venter and differs from
J. aurea in having shorter and narrower asci (18–25×4–
6μm) and shorter ascospores (3–6×1.5–2μm). Jattaea
algeriensis and J. microtheca also possess a tomentum, but
this is brown, macroscopically inconspicuous and formed
of short, septate flexuous hairs and the perithecia are
regularly papillate to short-beaked. Two other species from
the family Calosphaeriaceae,Calosphaeria africana
Damm & Crous and Calosphaeria ludens Berl., also have
a perithecial venter covered with a tomentum and glabrous
and black necks (Berlese 1900; Damm et al. 2008).
Jattaea ceanothina (Peck) M.E. Barr, Mycologia 77:
559. 1985. (Figs. 28–36)
≡Sphaeria (Obtectae)ceanothina Peck, Ann. Rep.
New York State Mus. 29: 62. (for 1875) 1878.
≡Physalospora caenothina (Peck) Sacc., Syll.
Fung. 1: 441. 1882.
=Valsa cornina Peck, Ann. Rep. New York State Mus.
38: 102–103. (for 1884) 1885.
≡Engizostoma corninum (Peck) Kuntze, Revis.
Gen. Pl. (Leipzig) 3: 473. 1898.
≡Romellia cornina (Peck) M.E. Barr, Mycologia
77: 560. 1985.
≡Jattaea cornina (Peck) M.E. Barr, J.D. Rogers &
Y.M. Yu, Mycotaxon 48: 534. 1993.
Perithecia solitary or forming irregular or loose valsoid
groups of 3–4 or aggregated in larger circular groups of 6–
8 individuals on wood beneath the periderm, or on exposed
wood on the margins of loosened periderm; venter (200−)
290–410 μm diam, (220−)300–400 μm high, globose to
subglobose, dark brown to black, glabrous; necks or short
beaks central, cylindrical, elongate, (90−)100–120 μm
wide, up to 200–400 μm long, straight, rounded at the
apex, converging radially with aggregated perithecia,
piercing the periderm in a circular fissure; ostiolum
periphysate. Perithecial wall leathery, two-layered, 25–
39 μm thick; outer layer of pale brown to red-brown
polyhedral cells of textura prismatica; inner layer of
subhyaline to hyaline, flattened, elongated, thin-walled
cells. Cells on ascogenous hyphae ellipsoidal to ovoidal,
3–3.5×5–6μm. Paraphyses were seen only as ca 2–3μm
wide fragments. Asci (54−)60–70×(11−)12–14(−17) μm
(mean ± s.e.=64.5±1.4×13.1 ± 0.3), in pars sporifera (27−)
33–45 μm long (mean ± s.e.=38.2±0.9), long-stipitate,
clavate, obtuse at the thickened apex, wall ca 1.5–2μm
thick, tapering towards the base from the sporiferous
portion, without a distinct discharge mechanism, floating
freely within the centrum at maturity, with a short appendix
remaining at the base after ascal dehiscence from ascoge-
nous hyphae, 8-spored. Ascospores (12−)13–16(−17)×3–
3.5 μm (mean ± s.e.=14.8±0.3×3±0.02), cylindrical-
allantoid to oblong, slightly curved, hyaline, 3–7-septate,
not constricted at the septa, smooth, broadly rounded at the
ends, arranged in a fascicle or 3-seriate in the upper part of
the ascus.
Anamorph: Unknown
Material examined: U.S.A., New York: Center
(Karner), Albany Co., on decaying wood of Ceanothus
americanus L., May 1875, C. H. Peck (NYS 701,
holotype of Sphaeria ceanothina); ibid., Kenwood,
Albany County, Side Hill, south of Susq. R. R. Bridge
No. 2 near Albany, on decaying wood beneath the
periderm of twigs of Cornus racemosa Lam., April 1884,
C. H. Peck (NYS 879, holotype of Valsa cornina); ibid.,
Bethlehem, C. H. Peck (NYS 3470).
Notes: The type material of V. cornina (NYS 879)
contains two packets with locality information different
from that given in the prologue. The protologue gives
1 8 Fungal Diversity (2011) 49:167–1987
Albany as the locality whereas one packet gives Kenwood
and the other gives Bethlehem as the locality. The locality
of the specimen labelled Kenwood is near the village
Kenwood in the town of Bethlehem on the Albany town
border. The Bethlehem packet contains a portion of the
same collection. Barr (1985) discussed the authenticity of
the holotype material of V. cornina. She concluded that
Peck was referring in the protologue to this collection (NYS
879, divided in two portions) made on the bank of the small
river separating the two towns.
Jattaea discreta (Berl.) Réblová, comb. nov.(Figs.37–
48,49–57)
MycoBank MB519578
≡Wegelina discreta Berl., Icon. Fung. 3: 8. 1900
(basionym).
Figs. 28–36 Jattaea ceanothina.28–30. Perithecia on the natural
substratum. 31. Longitudinal section of the perithecial wall. 32–34.
Asci containing ascospores. 35,36. Ascospores (arrows indicate
mucilaginous cushions at each end). 28–32,34–36 from NYS 879
(holotype of Valsa cornina); 33 from NYS 3470. Scale bars: 28–30=
500 μm; 31= 50 μm; 32, 33, 36 = 10 μm; 34 =20 μm; 35=5 μm
Fungal Diversity (2011) 49:167–198 1 97
Figs. 37–48 Jattaea discreta.37–40. Perithecia on the natural
substratum. 41. Longitudinal section of the perithecial wall. 42–47.
Asci with ascospores and in 45 with paraphyses. 48. Ascospores. 37–
41,46–48 from (J.F. 06100); 42–44 from NY (lectotype of W.
discreta); 45 from PAD 2817 (holotype of W. saccardoana). Scale
bars: 37–40= 500 μm; 41 = 25 μm; 42–48 =10 μm
1 Fungal Diversity (2011) 49:167–19880
=Wegelina saccardoana (as Saccardiana) Berl., Icon.
Fung. 3: 9. 1900.
Perithecia aggregated in small groups or in irregular
loose formations, superficial to semiimmersed in wood
beneath the peeled off periderm, often associated with dead
ascomata of other fungi; venter 230–400 μm diam, 250–
400 μm high, globose to subglobose, dark brown, glabrous;
neck central, elongate, 80–100 μm wide, 300–1,200 μm
long, straight or slightly flexuous, rounded to slightly
tapering at the apex; when perithecia are aggregated necks
converging radially, perithecia and necks at first decumbent
to the substratum necks then upright, not united in a disc at
the top and piercing the periderm separately in a common
point; ostiolum periphysate. Perithecial wall leathery, two-
layered, 32–40 μm thick; outer layer of brown polyhedral
cells of textura prismatica to angularis; inner layer of
subhyaline to hyaline, flattened, elongated, thin-walled
cells. Cells on ascogenous hyphae ellipsoidal to obpyri-
form, 2.5–3.5×4–5μm. Paraphyses persistent, abundant,
unbranched, septate, hyaline, cylindrical, apically free, 4.5–
6μm wide near the base, tapering to 2–3μm, longer than
the asci. Asci (30−)35–45(−50)×6.5–8μm (mean ± s.e.=
40±1.2×6.8±0.1), in pars sporifera 18–25(−31) μm long
(mean ± s.e.=22.3±0.5), long-stipitate, clavate, obtuse at
the thickened apex, tapering towards the base from the
sporiferous portion, without a distinct discharge mecha-
nism, floating freely within the centrum at maturity, with a
short appendix remaining at the base after ascal dehiscence
from ascogenous hyphae, 8-spored. Ascospores (6−)6.5–8×
1.5–2μm (mean ± s.e.=7.4±0.1×1.5 ± 0.05), suballantoid
to oblong, hyaline, aseptate, smooth, 2–3-seriate or
arranged in a fascicle in the upper part of the ascus.
Cultural characteristics: Colonies in vitro slow-growing,
after 21 d on PCA at 25°C 1.4–1.7 mm diam, felty or
cottony in texture, with entire margin; aerial mycelium
medium to sparse; colony surface subhyaline, pale brown to
pale olivaceous, whitish towards the margin, reverse grey-
ish in the centre, whitish to inconspicuous towards the
margin. Vegetative mycelium hyaline, subhyaline to pale
yellow-brown, 1.5–2.5 μm wide, septate, smooth. Con-
idiophores arising from aerial or submerged hyphae,
variable in length, 4–20 μm long, 2–2.5 μm wide, erect,
simple or branched, hyaline or subhyaline, one- to several-
septate, often with numerous sterile, inflated, subhyaline
cells. Phialides 4–10×2.5–3.5 μm (mean ± s.e.= 6.6 ± 0.5 ×
2.9±0.1), tapering to 1–1.5 μm just below the collarette,
Figs. 49–57 Phialophora-like anamorph of Jattaea discreta.49,52–56. Aerial mycelium with phialides and conspicuous collarette. 50,51. Aerial
mycelium with adelophialides. 49–57 from living strain (ex-epitype, CBS 127681). Scale bars: 49–57=10 μm
Fungal Diversity (2011) 49:167–198 181
terminal or intercalary, monophialidic, smooth, hyaline,
elongate-ampulliform, often attenuated at the base or
subcylindrical, frequently pigmented in the apical region
below the collarette; adelophialides cylindrical or ampulli-
form, neck 2.5–3×1.5 μm. Phialides developing a terminal,
funnel-shaped collarette, 2.5–4μm long, 1.5–2μm wide,
with a distinct periclinal thickening. Conidia 3.5–4.5(−5) ×
1–1.3 μm (mean ± s.e.=4.1±0.2×1.2±0.03), hyaline,
oblong-ellipsoidal or suballantoid, one-celled, smooth,
aggregated in round, slimy heads at the phialide tip.
Chlamydospores not observed.
Anamorph: phialophora-like
Material examined: FRANCE, Pyrénées Atlantiques:
Ariège, Rimont, Las Muros, on dead corticated branch of
Crataegus sp. still attached to the trunk, 9 May 2006, J.
Fournier J. F. 06100. ITALY, Veneto: Belluno, on decaying
wood of a branch of Acer pseudoplatanus L., autumn 1879,
Saccardo Mycotheca Veneta no. 1450 as Calosphaeria
wahlenbergii Nitschke (NY, lectotype of W. discre t a ,
designated here); ibid., Montello forest, on dead stem of Galium
sp., as Calosphaeria minima in Saccardo’s herbarium (PAD
2817, holotype of W. saccardoana); Lazio: Viterbo province,
Farnese, Nature Reserve Selva di Lamone, on decayed wood
of Quercus cerris L., 19 March 2010, M. Réblová & W.
Gams, M. R. 3139 (PRM 917688, epitype, designated here,
living culture CBS 127681). U.S.A., New Jersey: Gloucester
County, Newfield, on decaying wood of Vaccinium sp., 6
December 1878 (NY as Sphaeria microtheca).
Notes: Among Jattaea species, J. discreta is comparable
with J. echinella, but the latter taxon differs from J. discreta
in having shorter ascospores 5–6(−7) μm, and shorter asci in
a(12−)14–16(−18) μmlongpars sporifera, and smaller
perithecia usually scattered to gregarious in the decaying
wood with longer tapering necks with the reddish brown
apex.
A fungus close to J. discreta was collected in New
Zealand (NEW ZEALAND, Westlands: Ross ca 28 km SW
of Hokitika, on Ruatapu Ross Road 6, Totara Walley Road,
on decaying bark of a branch, 12 April 2005, M. Réblová
NZ 767, PDD 99960). It is similar to J. discreta in having
similar shape and size of ascospores, 6–7.5× 1.5(−2) μm
and asci, 30–36×5.5–6.5 μm (in pars sporifera 17–23 μm),
the perithecia grow solitarily or aggregated along the
margins of partly decayed and peeled off bark. The fungus
differs from J. discreta in morphology of the perithecial
wall of the venter. The wall is thicker, 43–52 μm wide; the
cells of the outer wall near the interior are brown brick-like
of textura prismatica becoming dark reddish-brown of
textura angularis towards the exterior. Neck is black,
elongated, central, glabrous and opaque. The morphology
of the perithecial wall resembles the wall of Jattaea aurea
and J. stachybotryoides.Jattaea aurea differs from the new
zealandian collection in having the perithecial wall of the
venter brown, it does not become red or reddish-brown
towards the exterior, but possesses a thin reddish layer of
cells or short hyphae tightly attached to the outer layer.
Jattaea stachybotryoides differs from the new zealandian
collection in shorter asci (18–25×4–6μm) and shorter
cylindric-oblong ascospores (3–6×1.5–2μm) (Romero and
Samuels 1991). The new zealandian collection (PDD
99960) is a further link between J. discreta and J.
stachybotryoides and other Jattaea species with similar
structure of the perithecial wall.
Jattaea echinella (Ellis & Everh.) Réblová, comb. nov.
(Figs. 58–65)
MycoBank MB519579
≡Ceratostomella echinella Ellis & Everh., North
Amer. Pyren. p. 195. 1892 (basionym).
Perithecia scattered, solitary to gregarious or arranged
in groups, semiimmersed to immersed in wood only
necks protruding, often beneath the periderm; venter
230–250 μm diam, 240–260 μm high, globose to
subglobose, dark brown to black, glabrous; neck dark
brown to black, red-brown at the apex, central, elongate,
80–90 μm wide, 350–3,000 μm long, straight or slightly
flexuous, tapering upwards the apex; when perithecia are
aggregated necks tightly converging radially, at first
decumbent to the substratum then upright, not united in
a disc at the top and piercing the periderm separately in
a narrow fissure; ostiolum periphysate. Perithecial wall
leathery to fragile, two-layered, 25–35 μm thick; outer
layer of red-brown polyhedral cells of textura prismatica,
with a thin layer of reddish pigment on the surface; inner
layer of subhyaline to hyaline, flattened, elongated, thin-
walled cells. Cells on ascogenous hyphae ellipsoidal, 2–
3×4–5μm. Paraphyses persistent, abundant, unbranched,
septate, hyaline, cylindrical, apically free, 3.5–6μmwide
near the base, tapering to 2–2.5 μm, longer than the asci.
Asci (31−)35–42(−50)×5–7μm (mean ± s.e.=38.3±0.6×
5.6±0.1), in pars sporifera (12−)14–16(−18) μm long
(mean ± s.e.= 14.9±0.2), long-stipitate, clavate, obtuse at
the thickened apex, tapering towards the base from the
sporiferous portion, without a distinct discharge mecha-
nism, floating freely within the centrum at maturity, with a
short appendix remaining at the base after ascal dehis-
cence from ascogenous hyphae, 8-spored. Ascospores 5–6
(−7)×1.5(−2) μm (mean ± s.e.=5.7±0.1×1.5±0.02), sub-
allantoid to oblong, hyaline, aseptate, smooth, 2–3-seriate
or arranged in fascicles in the sporiferous part of the ascus.
Anamorph: Unknown
Material examined: CANADA, Belis Corners: on bark of
Acer rubrum L., 22 April 1953, D. E. Wells (DAOM 35410);
Ontario: Merivale, on bark of Acer sp., 12 September 1952,
S. J. Hughes (DAOM 28793); Ontario: London, on decaying
wood of Acer sp., 15 October 1889, J. Dearness (NY 1003).
1 Fungal Diversity (2011) 49:167–19882
Figs. 58–65 Jattaea echinella.58,59. Perithecia on the natural
substratum. 60. Longitudinal section of the perithecial wall. 61–64.
Asci with ascospores and paraphyses. 65. Ascospores. 58–59 from
DAOM 28793; 60 from NY (holotype); 61–65 from DAOM 35410.
Scale bars 58, 59=1,000 μm; 60 = 25 μm; 61–65 = 10 μm
Fungal Diversity (2011) 49:167–198 183
U.S.A., Ohio: on the inner side of cast-off bark of Acer
rubrum, 11 February 1890, Morgan 928 (NY, holotype);
ibid., on inner side of the loosened bark of a trunk of Acer
sp., November 1880, J. B. Ellis, North American Fungi no.
580 (NY); New Jersey: Newfield, on loosened bark of Acer
rubrum, 6 November 1902, J. B. Ellis, Ellis & Everhart
Fungi Columbiani no. 1712 (NY).
Notes:Jattaea echinella is well distinguishable by
elongated often flexuous neck; the apex is reddish brown
in the transmitted light.The reddish colour is also conspicuous
when material is fresh or well hydrated. The species can be
compared with J. discreta or J. tumidula,fromwhichit
differs in having shorter asci in pars sporifera,viz. (12−)14–
16(−18) μmlongvs.18–25(−31) μmlonginJ. discreta and
(21−)23–27(−30) μmlonginJ. tumidula.
Jattaea leucospermi Marinc., M.J. Wingf. & Crous,
CBS Biodiv. Ser. 7: 54. 2008.
Anamorph: phialophora-like (Marincowitz et al. 2008:54)
Material examined: SOUTH AFRICA, Western Cape
Province: Kogelberg Nature Reserve, decayed bark of
Leucospermum conocarpodendron subsp. viride Rourke, 3
November 2000, S. Marincowitz S. L. 359 (PREM 59517,
holotype; living culture CBS 119343, ex-type strain).
Notes: For detailed description and illustration refer to
Marincowitz et al. (2008:54–55). The fungus is characterized
by the eutypelloid configuration of perithecia and hyaline to
subhyaline 1–3-septate ascospores, pale yellow in mass. The
phialophora-like anamorph was obtained in axenic culture
from ascospore isolates. Based on the molecular analysis, J.
leucospermi is closely related to J. mookgoponga, which is
known only as asexually reproducing species.
Jattaea microtheca (Cooke & Ellis) Berl., Icon. Fung. 3:
7. 1900. (Figs. 66–74)
≡Sphaeria (Obtectae)microtheca Cooke & Ellis,
Grevillea 5: 51. 1876.
≡Calosphaeria microtheca (Cooke & Ellis) Sacc.,
Syll. Fung. 1: 97. 1882.
≡Physalospora microtheca (Cooke & Ellis) Sacc.,
Syll. Fung. 1: 445. 1882.
≡Valsa (Calosphaeria)microtheca (Cooke &
Ellis) Cooke, Grevillea 14: 52. 1885.
≡Circinostoma microthecum (Cooke & Ellis)
House, Bull. New York State Mus. 266: 41. 1925.
Perithecia solitary or aggregated in irregular groups,
superficial to semiimmersed in wood, often erumpent through
wooden fibres; venter 200–250 μm diam, 250–280 μmhigh,
subglobose, ovoid to conical, dark brown to black, covered
with tomentum of brown, septate, flexuous hairs, ca 1.5 μm
wide, 1.5–2.5 μm long, papillate or with a short central neck;
ostiolum periphysate. Perithecial wall leathery to fragile,
two-layered, 30–40 μm thick; outer layer of gray brown
polyhedral cells of textura angularis; inner layer of subhya-
line to hyaline, flattened, elongated, thin-walled cells. Cells
on ascogenous hyphae ellipsoidal to subglobose, 3.5–4.5×3–
4.5 μm. Paraphyses persistent, abundant, unbranched,
septate, hyaline, cylindrical, apically free, 3–3.5 μmwide
near the base, tapering to 2 μm, longer than the asci. Asci
(30−)33–40×7–8μm (mean ± s.e.= 37.5±1.0×7 ± 0.1), in
pars sporifera 21–26 μm long (mean ± s.e.=22.8±0.5), long
stipitate, clavate, obtuse at the thickened apex, tapering towards
the base from the sporiferous portion, without a distinct
discharge mechanism, floating freely within the centrum at
maturity, with a short appendix remaining at the base after ascal
dehiscence from ascogenous hyphae, 8-spored. Ascospores 6–
8×1.5(−2) μm (mean ± s.e.=6.9±0.2×1.5±0.03), oblong,
suballantoid to subcylindrical, hyaline, aseptate, smooth, 2–3-
seriate to fasciculate in the upper part of the ascus.
Anamorph: Unknown
Material examined: U.S.A., New Jersey: Newfield, on a
twig of Andromeda sp., J. B. Ellis, (NY, presumed isotype
of Sphaeria microtheca); ibid., November 1880, J. B. Ellis,
North American Fungi no. 580 (NY).
Notes: Perithecia occur usually in softer decaying wood,
often erumpent through wooden fibres. Barr (1985)
reported the species to be fairly common on various
substrates in North America.
Two exsiccate collections of Sphaeria microtheca
deposited in the NY herbarium in New York were
examined; J. B. Ellis: North American Fungi no. 580, on
the inner surface of loosened bark of maple logs, November
1880. One exsiccate contained the correct substratum and
perithecia of the fungus described here as J. microtheca.
The other exsiccate collection contained two envelopes
with different specimens. One envelope contained a
wooden stem mounted on the paper and labelled “on
Andromeda”; only empty perithecia could be observed. The
second envelope contained a piece of maple bark with
perithecia of Capronia sp. and a calosphaerialean fungus
identified as J. echinella (this study).
Another collection (U.S.A., New Jersey: Newfield, on
decaying wood of Quercus sp., 22 March 1877, as Sphaeria
microtheca, J. B. Ellis 2484, NY) with asci and ascospores
similar to Jattaea microtheca was examined. This fungus
differs from J. microtheca in having the perithecial wall
thicker, 50–62 μm; outer wall consisting of layers of brown
cells of textura prismatica that become reddish-brown of
textura angularis towards the exterior, tomentum was not
observed. The fungus formed globose to subglobose
perithecia, usually arranged in rows with papilla or short
glabrous neck. This collection is not accepted as a
representative of J. microtheca
Jattaea mookgoponga Damm & Crous, Persoonia 20:
47. 2008.
1 Fungal Diversity (2011) 49:167–19884
Anamorph: phialophora-like (Damm et al. 2008: 47, fig. 7)
Culture examined: SOUTH AFRICA, Limpopo Prov-
ince: Mookgopong, from central necrosis in wood of
Prunus persica var. nucipersica (Suckow.) C.K. Schneid.,
31 August 2004, U. Damm (living culture CBS 120867;
ex-type strain).
Notes:Jattaea mookgoponga was isolated from necrotic
wood in axenic culture, but perithecia, asci and ascospores
Figs. 66–74 Jattaea microtheca.66. Longitudinal section of the
perithecial wall. 67. perithecium. 68. Hairs of the tomentum covering
perithecial venter. 69–72. Asci with ascospores still attached to the cells of
the ascogenous hyphae. 73. Paraphyses. 74. Ascospores. 66–68,71,72,
74 from North American Fungi no. 580 (NY); 69,70,73 from NY
(isotype). Scale bars: 66= 25 μm; 67 =250 μm; 68–71= 10 μm; 72, 73=
20 μm; 74= 5 μm
Fungal Diversity (2011) 49:167–198 185
did not develop in vitro, instead the phialophora-like
anamorph was produced (Damm et al. 2008). Attempts to
produce the teleomorph were not successful. Based on
molecular sequence data of ITS and nc28S rDNA, the
species was confirmed to belong to the family Calosphaer-
iaceae and described under the holomorph name as a new
Jattaea species. In the present ITS and the three-gene
phylogenies J. mookgoponga is phylogenetically closely
related to J. leucospermi.
For full description of the phialophora-like anamorph
refer to Damm et al. (2008).
Jattaea pleurostoma (Starbäck) Berl., Icon. Fung. 3: 8.
1900. (Figs. 75–84)
≡Massalongiella pleurostoma Starbäck, Ascom.
Oeland., p. 5, 1889.
Perithecia solitary or in irregular to small circular groups
of 3–5, on wood beneath the periderm, dark brown to
black, glabrous, venter 300–400 μm diam, 380–450 μm
high, globose to subglobose, sometimes laterally pressed,
dark brown to black; neck central, elongate, 95–110 μm
wide, 250–1,000 μm long, straight or slightly flexuous,
broadly rounded at the glabrous apex, often decumbent to
the substratum or upright, piercing the periderm separately;
ostiolum periphysate. Perithecial wall leathery, two-
layered, 33–45 μm thick; outer layer of brown, polyhedral
cells of textura prismatica; inner layer of subhyaline to
hyaline, flattened, elongated, thin-walled cells. Cells on
ascogenous hyphae ellipsoidal to obovoidal, 6–7×3–3.5 μm.
Paraphyses persistent, abundant, unbranched, septate, hya-
line, cylindrical, apically free, 2–3.5 μm wide near the base,
tapering to ca 2μm, longer than the asci. Asci (36−)40–50×
(8−)9.5–11(−12) μm (mean ± s.e.= 44.1±0.7× 10.4±0.1),
in pars sporifera 24–33 μm long (mean ± s.e.=26.9 ± 0.5),
long-stipitate, clavate, obtuse at the ca 0.3–0.5 μm thick
apex, tapering towards the base from the sporiferous portion,
without a distinct discharge mechanism, floating freely
within the centrum at maturity, with a short appendix
remaining at the base after ascal dehiscence from ascogenous
hyphae, 8-spored. Ascospores (8−)9–10.5× 2–2.5 μm
(mean ± s.e.= 9.6 ± 1.2 × 1.9±0.06), suballantoid, hyaline,
1-septate, smooth, arranged 3–4-seriate or in a fascicle in
the upper part of the ascus.
Anamorph: Unknown
Material examined: SWEDEN, Öland: Torslunda parish,
Färjestaden, on decaying twig of Rosa sp., 3 June 1888, K.
Starbäck (UPS 411626, F-129302, slide from the holotype
collection); Uppland: Dalby parish, ca 200 m E of
Jerusalem, roadside, on dead stem of Rosa canina L., 28
March 1980, K. Holm and L. Holm 1989b (UPS 411625, F-
129301); ibid., Dalby parish, Jerusalem, outside the SE
corner of the garden, on decaying twig of R. canina,2
November 1982, K. Holm and L. Holm 2770e (UPS
411624, F-129300); ibid., Gryta parish, ca 500 m ESE of
Husby, on decaying twig of R. canina, K. Holm and L.
Holm 2347a (UPS 411623, F-129999).
Notes: In the original collection the perithecia have long
necks, often decumbent to the substratum. In the other three
specimens examined the length of the neck can vary from long
upright to decumbent, and some perithecia in the collection
UPS 411623 were also seen papillate to short-necked.
Jattaea stachybotryoides Romero & Samuels, Sydowia
43: 234. 1991.
Anamorph: Unknown
Notes: For discussion, description and illustration refer
to Romero and Samuels (1991: 234, pl. 2 G–L, pl. 4 O–S).
Jattaea stachybotryoides was collected on bark and
wood of Eucalyptus viminalis Labill. and it is known only
from the type locality in Argentina. The fungus is
characteristic by golden-brown tomentum covering the
perithecial venter and growing away from aggregated
perithecia; hyphae form a loose stromatic layer and release
a yellow pigment in 3% KOH. The neck is brown,
elongated, glabrous and usually perpendicular to the
substrate; asci are 18–25×4–6μm, oblong-clavate, apex
thickened and a short-bristle like appendage is attached at
base after ascal dehiscence from ascogenous hyphae;
paraphyses are persistent, abundant, longer than the asci,
septate, tapering to 3–6μm at the tip; ascospores are 3–6×
1.5–2μm, cylindric-oblong, hyaline, one-celled, arranged
2-seriately in the ascus (Romero and Samuels 1991).
Jattaea tumidula (Sacc.) Réblová, comb. nov.
(Figs. 85,86–96)
MycoBank MB519580
≡Calosphaeria tumidula Sacc., Atti Soc. Veneto-
Trent. Sci. Nat. Padova 4: 77–100 (Fungi Ven. novi Ser. 4:
20) (basionym) 1875. Fungi Italici Autographice delineati.
Fascs 9–12, Tab. 474. 1878.
≡Togninia minima var. tumidula (Sacc.) Berl.,
Icon. Fung. 3: 11. 1900.
=Ceratostomella mali Ellis & Everh., Proc. Acad. Nat.
Sci. Philad. 42: 225. 1890.
Perithecia scattered, solitary, immersed in wood beneath
the loosened bark, only necks protruding; venter 230–300 μm
diam, 220–280 μm high, globose to subglobose, dark brown
to black, glabrous; necks central, elongate, 80–90 μm wide,
150–300 μm long, straight or slightly flexuous; ostiolum
periphysate. Perithecial wall leathery, two-layered, 25–32
(−45) μm thick; of brown polyhedral cells of textura
angularis on the exterior, becoming textura prismatica
towards the interior. Cells on ascogenous hyphae ellipsoidal
to subglobose, 2.5–3×4–5μm. Paraphyses persistent, abun-
dant, unbranched, septate, hyaline, cylindrical, apically free,
4–5μm wide near the base, tapering to 2–2.5 μm, longer
1 Fungal Diversity (2011) 49:167–19886
Figs. 75–84 Jattaea pleurostoma.75–77. Perithecia on the natural
substratum. 78. Longitudinal section of the perithecial wall. 79–82.
Asci and in 81,82 with paraphyses. 83. Ascogenous cells on
ascogenous hyphae. 84. Ascospores. 75–78,80,83 from UPS
411624, F-129300; 79,81,82,84 from UPS 411623, F-129999.
Scale bars: 75–77= 500 μm; 78 = 50 μm; 79 =25 μm; 80–83=20 μm;
84= 10 μm
Fungal Diversity (2011) 49:167–198 187
than the asci. Asci 35–45× (4.5−)5–5.5(−6) μm(mean±s.e.=
39.3± 0.6×5.3± 0.1), in pars sporifera (21−)23–27(−30) μm
long (mean ± s.e.=25.3±0.4), long-stipitate, oblong-clavate,
obtuse at the thickened apex, tapering towards the base from
the sporiferous portion, without a distinct discharge mecha-
nism, floating freely within the centrum at maturity, with a
short appendix remaining at the base after dehiscence from the
ascogenous hyphae, 8-spored. Ascospores 6–8×(1.5−)2 μm
(mean ± s.e.=6.9±0.1×1.9±0.03), suballantoid to oblong,
hyaline, aseptate, smooth, 2–3-seriate in the ascus.
Anamorph: Unknown
Material examined: ITALY, Treviso, Cansigio, branch of
Fagus sylvatica, Saccardo (PAD, holotype of Calosphaeria
tumidula). U.S.A., New Jersey: Gloucester County, New-
field, on inner surface of loose hanging bark of partly dead
apple tree, December 1889 (NY, holotype of Ceratosto-
mella mali); ibid., Newfield, on apple tree bark, December
1889 (NY, as Sphaeria microtheca).
Notes:Jattaea tumidula is similar to J. discreta and J.
microtheca in the shape and length of asci, but it differs from
them in having narrower asci; the asci of J. discreta are also
slightly shorter in pars sporifera,18–25(−31) μm. Jattaea
microtheca differs further from J.tumidula by perithecia
covered with macroscopically inconspicuous brown tomen-
tum. An illustration based on type material of Calosphaeria
tumidula was recently published in Mostert et al. (2006:97,
fig. 47). An illustration of C. tumidula (Fig. 85) based on the
type material is reproduced from Saccardo (1878).
Jattaea villosa (Nitschke) L. Mostert, Stud. Mycol. 54:
101. 2006.
≡Calosphaeria villosa Nitschke, Pyrenom. Germ.
p. 98. 1867.
≡Togninia villosa (Nitschke) Berl., Icon. Fung. 3:
10. 1900.
Note: For full synonymy, discussion, description and
illustration refer to Mostert et al. (2006: 101, fig. 49).
Jattaea villosa is characterized by allantoid to suballan-
toid, septate, hyaline ascospores [8–11(−12)×(1−)1.5–
2μm] in oblong-clavate, long-stipitate asci [30–70×5–
7μm, in pars sporifera 20–47(−55) μm]; perithecia solitary
or in small loose groups developing beneath the periderm;
they are papillate, rarely with a short neck. The septation of
ascospores is delayed, one-celled ascospores occur usually
in the mature asci and after they are released the second and
third septa are developed. The perithecia are on the natural
substratum associated with subglobose to flattened, subepi-
dermal spermogonia, for details see Mostert et al. (2006).
Jattaea villosa differs from other Jattaea species with
septate ascospores in having a perithecial venter covered
with sparse, pale yellowish-brown tomentum and narrower
asci, 5–7(−10) μm wide.
Species excluded from Jattaea and Wegelina
Accepted names are printed in bold; many excluded species
are not reclassified.
Jattaea berlesiana Sacc. & Traverso, Annls. Mycol. 1:
432. 1903. (Figs. 97–99)
Material examined: ITALY, Sardinia: Sassari, on dead
wood of trunk and branch of Cistus salviaefolius L., A. N.
Berlese no. 10 (PAD, holotype); ibid., on dead wood of
Cistus sp., A. N. Berlese no. 225 (PAD).
Notes: In the type collection A. N. Berlese no. 10 (PAD),
no perithecia were present, instead a card with a drawing
and a description was enclosed. This description matched
well the protologue and my observations of the fungus in
Fig. 85 Jattaea tumidula. Illustration of Calosphaeria tumidula
(Saccardo, Fungi Italici Autographice Delineati, Fascs 9–12, Tab.
474. 1878)
1 8 Fungal Diversity (2011) 49:167–1988
the collection A. N. Berlese no. 225 (PAD). This specimen
contains mature perithecia of Jattaea berlesiana with asci
and ascospores. Perithecia form small circular groups
beneath the periderm. Asci are 40–46 ×(8−)9–10 μm,
oblong, rounded at the base, without a stipe and bristle-
like appendix, obtuse at the thin-walled apex; apical
annulus and paraphyses were not observed; ascospores are
11–12×3–4μm, ellipsoidal, one-celled, sometimes slightly
curved, flattened at one side, hyaline. Saccardo (in Traverso
1903) noted the occurrence of a Cytospora sp. anamorph
associated with perithecia on the host. This fungus should
better be classified in the genus Valsa Fr.
Jattaea brevirostris (Ellis) Berl., Icon. Fung. 3: 7. 1900.
Notes:Berlese(1900) based his new species on
Calosphaeria microtheca var. brevirostris Ellis (Saccardo
Figs. 86–96 Jattaea tumidula.86. Longitudinal section of the
perithecial wall. 87. Perithecium, the long neck is depicted only as a
fragment. 88. Paraphyses. 89–94. Asci containing ascospores. 95.
Ascospores. 86–95 from NY (holotype of C. mali). Scale bars: 86=
25 μm; 85= 100 μm; 88–94 = 10 μm; 95, 96 =5 μm
Fungal Diversity (2011) 49:167–198 1 98
1891: Tab. 449) and illustrated it with globose perithecia
without tomentum. However, Ellis and Everhart (1892)
and Cash (1952) did not mention such a variety and
neither Barr (1985) nor I could locate any material of this
fungus.
Jattaea curvicolla (Peck) M. E. Barr, Mycologia 77:
558. 1985. (Figs. 100–103)
≡Sphaeria (Caulicolae)curvicolla Peck, Ann.
Rep. New York State Mus. Nat. Hist. 31: 50. (for 1877)
1879.
≡Gnomoniella curvicolla (Peck) Sacc., Syll.
Fung. 1: 417. 1882.
≡Eutypa curvicolla (Peck) M. E. Barr, Mycol.
Mem. 7: 20. 1978.
Material examined: U.S.A., New York: Karner (Center),
Albany Co., on dead stems of Polygonella articulata (L.)
Meisn., October 1877, C. H. Peck (NY, holotype).
Notes: Peck (1879) described the perithecia of S.
curvicolla as black, hemispherical with short, subcylin-
drical, slightly curved ostiolum, 300–400 μm wide, asci
are oblong containing hyaline, oblong ascospores, 6–9×
3μm. Barr (1985) based her observations on the type
material and described and illustrated the fungus with
ascomata 315–490 μm diam, slightly depressed, with
papilla or short beak often slightly curved, asci are in pars
sporifera 32.5–39.5× 4.5–5.5 μm and contain allantoid,
hyaline ascospores, 5.5–7×2–3μm.
According to my observations, the type material contains
ca 35 longer (9–13 cm long) and ca 35 shorter (1–5 cm)
twigs of Polygonella articulata. They are covered with
dark, nonstromatic subglobose perithecia that externally
match the original macroscopic description of S. curvicolla
by Peck (1879), asci are oblong 75–95×(18−)20–25 μm, in
pars sporifera 65–82 μm long and contain hyaline,
ellipsoidal ascospores, (15−)17–22×7–9μm. Every twig
was searched for any deviation in perithecia that might
belong to Jattaea and several randomly selected perithecia
were examined microscopically, but the ascospores and asci
illustrated and described by Peck (1879) and Barr (1985)
could not be found. The same conflict between the type
material of S. curvicolla and observations made by Barr and
Peck were reported earlier by Pirozynski and Morgan-Jones
(1968).
Probably the type collection contained two fungi, one of
which is a species of Jattaea; it might be in a minority or its
perithecia could already be depleted. Therefore, for the time
being, the fungus in the type collection of S. curvicolla is
not accepted in Jattaea until it is recollected and its generic
status revealed based on examination of fresh material. The
fungus, whose perithecia are abundantly present in the
holotype, should be accommodated in the genus Botryos-
phaeria Ces. & De Not. of the Dothideomycetes.
Jattaea faginea Moesz, Annls Mus. Nat. Hung., Pars
Bot. 36: 129. 1944. (Figs. 104–107)
=Togninia vibratilis (Fr.: Fr.) Réblová & L. Mostert,
Mycol. Res. 111: 301. 2007.
Anamorph.Phaeoacremonium sp.
Material examined: HUNGARY, Mátrakása: on decayed
wood of Fagus sylvatica, 31 May 1939, J. Moesz (BP
12460, holotype of J. faginea).
Notes: For the full synonymy, discussion, description
and illustration refer to Réblová and Mostert (2007: 301,
fig. 3a–f).
Jattaea herbicola (Ellis & Everh.) Berl., Icon. Fung. 3:
8. 1900.
≡Calosphaeria herbicola Ellis & Everh., North
Amer. Pyren. p. 511. 1892.
=Togninia leptorrhyncha (Durieu & Mont.) Réblová,
comb.nov. (Figs. 108–112)
MycoBank MB 519581
≡Sphaeria leptorrhyncha Durieu & Mont.,
Explor. Scient. Algérie 1: 510. 1849 (basionym).
≡Ceratostomella leptorrhyncha (Durieu & Mont.)
Sacc., Syll. Fung. 1: 412. 1882.
=Togninia novaezealandiae Georg Hausner, Eyjólfsd. &
J. Reid, Can. J. Bot. 70: 729. 1992.
Material examined: ALGERIA, La Calle: on bark of
Chamaerops humilis L. (Arecales), 19 November 1840
(PC, holotype of Sphaeria leptorrhyncha). ITALY, on wood
of Castanea sativa Mill. (PAD, as Calosphaeria wahlen-
bergii). U.S.A., New Jersey: Newfield, on decaying stems
of Lactuca canadensis L., 20 January 1891 (NY, holotype
of Calosphaeria herbicola); Massachusetts: Baptist Hill,
Conway, on Acer saccharum Marshall, 18 March 1973, M.
E. Barr Bigelow 6035 (NY, as Calosphaeria microtheca);
New York: Warren Co., Camp Dippikill, Cam/Ridge Trails,
on dead branches of Acer spicatum Lamarck, 17 September
1993, S. M. Huhndorf 641 (NY, as Jattaea microtheca).
UKRAINE, Carpathian Mts.: Boržava Massif, Pilipec,
decayed bark of Fagus sylvatica, 17 July 1998, M. Réblová
M. R. 1281 (PRA).
Jattaea spermatozoides Berl., Icon. Fung. 3: 7. 1900.
(Figs. 113–115)
Material examined: ALGERIA, La Calle: as Sphaeria
tetragona Duby on stem of Smilax aspera subsp. maur-
itanica Aschers. & Graebn., Duby (PC, two identical
specimens, not type).
Notes:Berlese(1900) based his new species on
material of Eutypa tetragona (Duby) Sacc. f. simplex
Durieu & Mont., Expl. Sci. Alg. 1: 460. 1848. The fungus
was originally described with immersed, globose perithe-
cia with erumpent thickened ostiolum with 3–4outer
ridges with clavate, long-stipitate asci containing allan-
1 Fungal Diversity (2011) 49:167–19890
Figs. 97–107 97–99.Jattaea berlesiana.97,98. Asci. 99. Asco-
spores. 100–103.Jattaea curvicola.100,101. Asci. 102. Longitudinal
section of the perithecial wall. 103. Ascospores. 104–107.Togninia
vibratilis.104,105,107. Asci with ascospores. 106. Perithecia on the
natural substratum. 97–99 from PAD (holotype); 100–103 from NY
(holotype); 104–107 from BP12460 (holotype of Jattaea faginea).
Scale bars: 97–100, 103–105, 107=10 μm; 102=25 μm; 106 =
500 μm
Fungal Diversity (2011) 49:167–198 191
toid, hyaline ascospores (Berlese 1900). In the PC
herbarium are deposited two specimens labelled Sphaeria
tetragona Duby, but none represents a holotype. Both
collections contain a fungus, which is identical to J.
spermatozoides described by Berlese (1900).
The ascomata are surrounded by a thin, dark, marginal
stromatic zone in the wood, the asci are unitunicate,
clavate, cylindrical in pars sporifera with a long tapering
stipe and contain allantoid, one-celled, hyaline ascospores,
which are yellowish in mass and arranged in a fascicle in
the middle of the sporiferous part beneath the ascal apex.
Both collections represent a species of Eutypa sp.
Jattaea tympanoides (M. E. Barr) M.E. Barr, J.D. Rogers
& Y.M. Ju, Mycotaxon 48: 534 1993.
≡Romellia tympanoides M. E. Barr, Mycologia
77: 561. 1985.
≡Conidiotheca tympanoides (M. E. Barr)
Réblová & L. Mostert, Mycol. Res. 111: 305. 2007.
Material examined: U.S.A., Massachusetts: Baptist Hill,
Conway, on decaying twigs of Prunus pennsylvanica L., 21
April 1969, M. E. Barr 5275 (NY, holotype).
Notes: For the full synonymy, discussion, description
and illustration refer to Réblová and Mostert (2007: 305,
fig 3g–t).
Wegelina barbirostris (Dufour) M. E. Barr, Cryptog.
Bryol.-Lichénol. 19: 172. 1998.
≡Barbatosphaeria barbirostris (Dufour) Réblová,
Mycologia 99: 727. 2007.
Anamorphs:Ramichloridium and Sporothrix type
Material examined: ITALY, on decayed wood, Fries’
herbarium collection no. 540, L. Dufour (UPS 59201, F-
004917, holotype).
Notes: For the full synonymy, discussion, description
and illustration refer to Réblová (2007: 727, figs 3, 4).
Wegelina cryptomeriae Sawada, Bull. Gov. Forest Exp.
Sta. Tokyo 45: 51. 1950.
≡Pleuronaema cryptomeriae Sawada (nom. inval.,
Art. 36)
Notes:Pleuronaema cryptomeriae is an unpublished
name. The name W. cryptomeriae was validly published
by Sawada (1950: 51, fig. 10), it contained a Latin
diagnosis and syntype designation, and should be cited
as the basionym for any future combinations (Art. 33.7a).
The fungus was described from leaves of Cryptomeria
japonica D. Don having pycnidia associated with perithecia,
which are immersed and covered with epidermis, asci
cylindrical, obtuse at the base and ascospores hyaline, one-
celled. The genus Pleuronaema Höhn. is a synonym of the
coelomycete genus Cytospora Ehrenb., the anamorph of
Va l s a .NotypematerialofW. cryptomeriae is available.
Wegelina grumsiniana (Kirschst.) V. Kumm., T. Richt. &
Schwik, Z. Mykol. 71: 232. 2005.
=Ceratostomella rostrata (Tode : Fr.) Sacc., Syll. Fung.
1:409. 1882.
Material examined: GERMANY, Bredowen Forest:
Orthavelland, in hymenium of the decayed basidioma of
Fomes fomentarius (L.) J. Kickx on Betula sp., 6 October
1916, W. Kirschstein (B 700006552, holotype).
Notes: For full synonymy, discussion, description and
illustration refer to Réblová (2006: 80, figs 16–18).
Wegelina polyporina M.E. Barr, Cryptogamie, Bryol.
Lichénol. 19: 170. 1998.
=Ceratostomella rostrata (Tode : Fr.) Sacc., Syll. Fung.
1: 409. 1882.
Material examined: SWEDEN, Närke: Glanshammar
Parish, Skölv, in hymenium of Fomes fomentarius,20
May 1993, K. G. Nilsson (DAOM 227786, isotype);
Notes: For full synonymy, discussion, description
and illustration refer to Réblová (2006: 80, figs 16–
18).
Wegelina sepulta (Berk. & Broome) Berl., Icon. Fung. 3:
8. 1900.
≡Sphaeria sepulta Berk. & Broome, J. Linn. Soc.,
Bot. 10: 387. 1868.
=Sphaeria scoparia Schwein., Schr. Naturf. Ges.
Leipzig 1: 37. 1822.
≡Eutypella scoparia (Schwein. : Fr.) Ellis &
Everh, North Amer. Pyren. p. 495. 1892.
≡Peroneutypa scoparia (Schwein. : Fr.) Carmarán
& A. I. Romero, Fung. Divers. 23: 84. 2006.
Notes: According to Barr et al. (1993), W.sepulta is a
synonym of Eutypella scoparia, now recognized as
Peroneutypa scoparia of the Diatrypaceae.
Wegelina subdenudata (Peck) M. E. Barr, Cryptogamie,
Bryol. Lichénol. 19: 172. 1998. (Figs. 116,117)
≡Sphaeria (Ceratostomae)subdenudata Peck,
Ann. Rep. New York State Mus. 32: 52 (for 1878) 1880.
≡Ceratostoma subdenudatum (Peck) Sacc., Syll.
Fung. 9: 481. 1891.
≡Ceratostomella subdenudata (Peck) M.E. Barr,
New York State Mus. Bull. 459: 44. 1986.
Material examined: U.S.A., New York: Griffins, Catskill
Mts., Delaware Co., on decaying wood, September 1877,
C. H. Peck (NY, holotype).
Notes: The species differs from Jattaea and other genera
of the Calosphaeriales in having robust perithecia with
long sulcate necks, oblong-cylindrical asci with short stipe,
distinct apical annulus and ascospores, which are hyaline,
becoming light dull brown, one-celled but with 1–5
pseudosepta at maturity. The species is not accepted in
1 Fungal Diversity (2011) 49:167–19892
Jattaea; it is morphologically related to species classified in
Lentomitella Höhn. (Réblová 2006).
Species accepted in Togniniella
Togniniella microspora (Ellis & Everh.) Réblová, comb.
nov. (Figs. 118–120)
MycoBank MB 519582
≡Ceratostomella microspora Ellis & Everh., Proc.
Acad. Nat. Sci. Philadelphia 45: 444. 1894 (basionym).
=Togniniella acerosa Réblová, L. Mostert, W. Gams &
Crous, Stud. Mycol. 50: 545. 2004.
Anamorph:Phaeocrella acerosa Réblová, L. Mostert,
W. Gams & Crous
Material and cultures examined: NEW ZEALAND,
Tasman Province: Nelson Lake National Park, Lake Rotoiti
near St. Arnaud, Lakehead track ca 1.5 km SE of NP
Headquarters, on decayed wood of a trunk of Nothofagus
sp., 22 February 2003, M. Réblová M.R. 2593 (PDD
81431, holotype of T. acerosa; living culture ICMP 15113,
ex-type strain); ibid., Westlands: Harihari, Saltwater
Forest, Poerua River valley, on decaying wood, 12 March
2003, M. Réblová M. R. 2811 (PDD 81432, living culture
ICMP 15149). U.S.A., New York: Alcove, on rotten beech
log, September 1893, C. L. Shear 174 (00911861 NY,
holotype of C. microspora).
Notes:Togniniella microspora is the only species of the
genus. It is characterized by minute allantoid ascospores in
clavate, stipitate asci arranged in spicate formation on
ascogenous hyphae. For further discussion, description and
illustration of the holomorph refer to Réblová et al. (2004:
545, figs 41–60).
Species of uncertain status, morphologically similar
to Jattaea
Calosphaeria microtheca (Cooke & Ellis) Sacc. var.
rosmarini Rolland, Bull. Soc. Mycol. Fr., p. 2. 1896.
Notes: The var. rosmarini was described from decayed
wood of rosemary in France with stromatic, black, minute
perithecia growing beneath the periderm, long to short
necks, asci are 20×5 μm, clavate, short-stipitate containing
allantoid, hyaline 5×1 μm ascospores arranged biseriately.
The type variety Jattaea microtheca var. microtheca differs
from var. rosmarini in having longer and wider asci, (30−)
33–40×7–8μm and longer and wider ascospores, 6–8×1.5
(−2) μm. The type material of var. rosmarini was not
located in the PC herbarium (Bart Buyck, Curator, pers.
comm.). Calosphaeria microtheca var. rosmarini is not
accepted in the genus Jattaea, because the protologue does
not contain sufficient information for the taxon classification.
Calosphaeria microtheca (Cooke & Ellis) Sacc. f.
salicis-albae Fautrey, Rev. Mycol. Toulouse, p. 53. 1897.
Notes: The forma salicis-albae was described from a
branch of Salix alba L. in France. Perithecia were described
as minute, aggregated in the bark and with a long neck, asci
are 20×5 μm, pyriform, short-stipitate containing hyaline,
allantoid 5×1 μm ascospores. The type material of f.
salicis-albae was not located in the PC herbarium (Bart
Buyck, Curator, pers. comm.). The fungus is not accepted
in the genus Jattaea, because the protologue does not
contain information sufficient for its classification.
Calosphaeria polyblasta Romell & Sacc., Grevillea 21:
65. 1893. Fig. 121
≡Cesatiella polyblasta (Romell & Sacc.) Höhn.,
Annls Mycol. 3(2): 188. 1905.
Notes: According to the protologue and the original
illustration of Calosphaeria polyblasta (Saccardo 1893a,b),
the fungus is similar to Jattaea species with septate
ascospores. Perithecia are described as papillate, subglo-
bose, black, arranged in circular configurations beneath the
periderm with asci 42–50×9–10 μm, long-stipitate, clavate,
obtuse at the apex containing eight ascospores, which are
12–22×3 μm (average 15×3), oblong, curved, 4-septate,
hyaline, arranged in fascicles or 2-seriate in the ascus. The
fungus was collected on a decaying branch of Salix caprea
L. Calosphaeria polyblasta is similar to J. ceanothina in
the morphology of the ascospores, their size and septation,
but it differs from the latter by shorter and narrower asci.
The type material of C. polyblasta is not available. In
order to accept this species in Jattaea, fresh material needs
to be recollected. The illustration based on the type material
is reproduced from Saccardo (1893b).
Calosphaeria polyblasta was combined under the
genus Cesatiella Sacc. in the Hypocreaceae by von
Höhnel (1905). Upon examination of the type species
Cesatiella australis Sacc. & Speg., Rossman et al. (1999)
concluded that Cesatiella is similar to Leiosphaerella
Höhn. and belongs in the Hyponectriaceae in the
Xylariales.Petrak(1923) regarded Calosphaeria poly-
blasta as another species of Phragmocalosphaeria,buthe
invalidly proposed the combination without citing the
basionym.
Calospora suecica Rehm,inVleugel,SvenskBot.
Tidskr. 5: 327. 1911.
Notes: According to the protologue, the fungus is morpho-
logically similar to Jattaea allantospora in characters of 3–5-
septate ascospores, 10–12.5×2–2.5 μm; clavate short-stipitate
asci 35–40× 5.5–7μm and persistent paraphyses. The
perithecia are described as black, coriaceous, 400 μmdiam
with short convergent necks protruding and arranged in small
clusters or solitary.
Fungal Diversity (2011) 49:167–198 193
1 Fungal Diversity (2011) 49:167–19894
The type material is not available. The protologue does
not contain sufficient information for its classification.
Discussion
The revision of the type, other herbarium material and in
vitro studies of Wegelina discreta and its comparison to
Jattaea algeriensis and other species classified in Jattaea
revealed that Jattaea and Wegelina are morphologically
closely related. Two phylogenetic analyses, based on
molecular DNA data of ITS and nc28S, nc18S rDNA and
RPB2 of J. algeriensis,W. discreta and other members of the
Calosphaeriales confirm that these species are congeneric.
Berlese erected both genera Jattaea (Berlese, Icon. Fung. 3:
6. 1900) and We ge l i n a (ibid. 3: 8. 1900), in the same
publication. The generic name Jattaea is accepted with
Wegelina as its synonym to include species with hyaline,
allantoid to suballantoid ascospores in clavate, stipitate asci
without apical annulus borne on individual cells on ascoge-
nous hyphae. The associated anamorphs are referred to as
phialophora-like. Twelve species are classified in Jattaea.
The molecular data and cultivation studies support the
placement of species with one-celled and one- to several-
septate ascospores in Jattaea. In the ITS and three-gene
phylogenies, Jattaea leucospermi represents the phenotypic
pattern with septate allantoid to suballantoid ascospores.
This pattern was formerly considered a diagnostic feature of
the monotypic genus Phragmocalosphaeria (Petrak 1923).
Molecular data confirm that ascospore septation is not a
significant character suitable to distinguish genera in the
Calosphaeriales,andPhragmocalosphaeria is accepted as
a synonym of Jattaea. The arrangement of perithecia on the
host, morphology of asci, particularly the ontogeny of
ascogenous hyphae and morphology of phialides, adelo-
phialides and conidia of the anamorphs are important
characters at genus level.
Several researchers have noted that the Calosphaeriales
represent a polyphyletic group of phenotypically similar
taxa that may comprise at least two other phylogenetic
lineages (Barr et al. 1993; Samuels and Candoussau 1996;
Barr 1998). The exclusion of Graphostroma of the Graph-
ostromataceae in the Xylariales from the Calosphaeriales
was a first step in the phylogenetic tidying of the
Calosphaeriales (Barr et al. 1993). A revision of some
Ceratostomella (inc. sedis) species that do not match the
generic concept around the lectotype species C. rostrata
(Tode : Fr.) Sacc. (Réblová 2006), revealed that they are
morphologically similar to the Calosphaeriales in having
perithecia growing between cortex and wood, arranged
separately or in circular formations with converging necks,
hyaline allantoid to suballantoid ascospores, densely
branching cylindrical-clavate ascogenous hyphae and ellip-
soidal ascogenous cells arranged in characteristic patterns,
but they are associated with two modes of conidiogenesis.
Five Ceratostomella species with a calosphaeria-like
phenotype and three former Calosphaeria species, associated
with holoblastic denticulate conidiogenesis, conidiophores of
Ramichloridium- or Sporothrix-types, were classified in three
distinct holomorph genera, i.e., Barbatosphaeria Réblová,
Natantiella Réblová and Tectonidula Réblová (Réblová
2007; Réblová and Štěpánek 2009). These three genera
(inc. sedis) are phylogenetically related to each other and
represent a second putative phylogenetic lineage of
calosphaeria-like fungi (Fig. 1) with closest relatives in the
Ophiostomatales. The revision of type and other herbarium
material of four further species, i.e., Ceratostomella echi-
nella,C.leptorrhyncha,C. mali and C. microspora revealed
their striking morphological similarities with calosphaeria-
like fungi with phialidic conidiogenesis. These species are
classified in Jattaea,Togninia and Togniniella and are
presented in this study.
Jattaea is closely related to Calosphaeria and Togni-
niella in the Calosphaeriaceae, order Calosphaeriales.In
Jattaea and Calosphaeria the ascogenous hyphae are
cylindrical, clavate to ellipsoidal and seem to have
terminated growth (Jattaea: Figs. 20,21 and 22,63,71,
Figs. 108–120 108–112.Togninia leptorrhyncha.108. Paraphyses.
109,111. Asci containing ascospores. 11 0. Asci attached to the
ascogenous hyphae. 112. Ascospores. 113 –115.Jattaea spermato-
zoides.113. Ascus. 114 . Ascospores. 115 . Asci with paraphyses in the
hymenium. 116,117.Wegelina subdenudata.116,117 . Asci and
ascospores. 118–120.Togniniella microspora.118. Longitudinal
section of the perithecial wall. 11 9,120. Asci with ascospores
attached to the cells on ascogenous hyphae. 108–112 from NY
(holotype of Calosphaeria herbicola); 113 –115 from PC; 11 6,11 7
from NY (holotype); 118–120 from NY 00911861 (holotype of
Ceratostomella microspora). Scale bars: 108= 20 μm; 109–117, 119,
120= 10 μm; 118=50 μm
R
Fig. 121 Calosphaeria polyblasta. Illustration of C. polyblasta
(Saccardo, Fungi novi Europaei et Asiatici. Grevillea 21: Plate 184.
1893.) Scale bars: a=500 μm; b= 10 μm; c = 5 μm
Fungal Diversity (2011) 49:167–198 195
80,83;Calosphaeria: Réblová et al. 2004: figs. 3, 9–11).
The cells growing on ascogenous hyphae are often arranged
in a terminal cluster; croziers occur among the cells. In
Togniniella ascogenous hyphae often proliferate and elon-
gate (Fig. 119; Réblová et al. 2004: figs. 28, 31, 32), giving
the attached asci a spicate arrangement. The ascospores of
Togniniella are generally narrower; in T. microspora they
do not exceed 1 μm in width, while in Jattaea and
Calosphaeria ascospores are regularly 1.5–3μm wide.
The delimitation of Jattaea from Calosphaeria is rather
narrow and it is predominantly based on the arrangement of
the perithecia on the natural substratum and the morphology
of asci, ascospores and anamorphs. However, the difference
between the two genera is more remarkable when the primary
and secondary structures of ITS rDNA sequences of their
species are compared. The ITS1 and ITS2 sequences of
Calosphaeria are generally longer (195–201 bp) and their
ITS1 region forms different and more sophisticated secondary
structure motifs than the ITS1 region of Jattaea species (169–
181 bp) (Réblová, in prep.). The perithecia of Calosphaeria
species have a globose to subglobose venter with elongated
neck and grow in dense circinate or valsoid configurations
beneath the periderm often in few vertical levels with
converging necks, while the perithecia of Jattaea and
Togniniella grow usually solitarily, scattered or in small
irregular to loose valsoid groups beneath the periderm.
The known anamorphs associated with Jattaea species
have phialidic conidiogenesis and are referred to as
phialophora-like (Damm et al. 2008; present study). Their
conidiophores are semimacronematous, phialides and ade-
lophialides are usually subhyaline to lightly pigmented with
a flaring collarette with periclinal thickening and produce
hyaline, allantoid to suballantoid conidia. The phialophora-
like anamorphs of Jattaea are morphologically very similar
to the Calosphaeriophora Réblová, L. Mostert, W. Gams &
Crous anamorphs of Calosphaeria (Réblová et al. 2004)in
having phialides with a pigmented apical region and deep,
flaring collarettes and hyaline one-celled conidia. The
phialides of Calosphaeriophora are described as hyaline,
while those of the phialophora-like anamorph of Jattaea are
subhyaline or with a pale yellow-brown pigment. The other
two anamorphs linked with the Calosphaeriales are the
Phaeocrella Réblová, L. Mostert, W. Gams & Crous
anamorph of Togniniella and the Pleurostomophora Vijaykr.,
L. Mostert, Jeewon, W. Gams, K.D. Hyde & Crous
anamorph of Pleurostoma (Réblová et al. 2004; Vijaykrishna
et al. 2004).
Jattaea resembles the genus Togninia of the Togninia-
ceae in suballantoid one-celled, hyaline ascospores, clavate
asci without apical annulus and solitarily growing perithe-
cia. The anamorphs of Togninia exclusively belong to the
dematiaceous hyphomycete genus Phaeoacremonium,a
genus of approximately 35 species introduced to include
fungi that are intermediate between Acremonium Link : Fr.
and Phialophora Medlar (Crous et al. 1996; Dupont et al.
2000; Groenewald et al. 2001; Mostert et al. 2003,2004,
2006).
Damm et al. (2008) described ascus dehiscence and
ascospore release in the Calosphaeriaceae on the example
of Calosphaeria africana Damm & Crous. The authors
concluded that the ascus of C. africana possesses two
functionally different walls and the process should be
compared to a fissitunicate ascus dehiscence of the
Dothideomycetes (e.g. Schoch et al. 2007). This is an
interesting phenomenon and it was observed for the first
time in calosphaerialean fungi. Damm et al. (2008)
interpreted the ascal apex of Jattaea algeriensis (as
prunicola) as having an apical ring. According to our
observations, the ascal apex of all four genera classified in
the Calosphaeriales and accepted in this study, does not
contain an apical annulus (Réblová et al. 2004; Vijaykrishna
et al. 2004); instead, the ascus wall at the apex and extending
to the sides towards the stipe is conspicuously thickened. In
some species, like J. algeriensis (Damm et al. 2008,fig.5g,
h) and J. aurea (Figs. 23,24, present study), the wall is
disrupted centrally at the apex and mature asci contain some
kind of central pore through which the ascospores are
released. In asci, which are collapsed and have shrunken
walls after ascospore release, the ascal apex with thickened
walls approaching each other may mimic a ring.
Species of Calosphaeria and Jattaea have been deter-
mined to function as plant pathogens associated with
disease of Prunus. Trouillas et al. (2010) isolated Calo-
sphaeria pulchella from diseased branches of sweet cherry
(Prunus avium L.) in California. The fungus was isolated
from cankers and vascular necroses of the wood, perithecia
were also formed beneath the periderm. Pathogenesis of
four isolates of C. pulchella was determined by branch
inoculations. The ITS sequences of the four pathogenic
strains grouped in a strongly supported clade (100/99/0.9)
with the other strain of C. pulchella, which formed
perithecia on dead branches of P. avium in France. Also,
the three latest species additions to the family Calosphaer-
iaceae were obtained by isolation from wood necroses
close to old pruning wounds (Damm et al. 2008):
Calosphaeria africana was isolated from V-shaped necrosis
in wood of Prunus armeniaca L., Jattaea algeriensis (as
prunicola) from brown necrosis in wood of Prunus salicina
Lindl., and J. mookgoponga from reddish brown central
necrosis in wood of Prunus persica var. nucipersica.
Acknowledgement This study was financially supported by the
Project of the National Foundation of the Czech Republic (GA206/09/
0547) and Institutional Research Concept No. AV0Z60050516 (Institute
of Botany of the ASCR). Prof. Walter Gams is thanked for careful reading
of the manuscript and providing many suggestions. Drs Paul Kirk and
Scott Redhead are thanked for help with solving the nomenclatural
1 Fungal Diversity (2011) 49:167–19896
problem of Wegelina cryptomeriae and supplying the literature. I am
grateful for the help of Dr Václav Štěpánek, who undertook the DNA
sequencing of ascospore isolates of Jattaea discreta, other members of
the Calosphaeriales and provided other sequences newly presented in
this study. The curators of herbaria (B, BP, CBS-H, DAOM, NY, NYS,
PAD, PC, PDD, PREM, UPS) are acknowledged for loans of the type
and other herbarium material and the curators of the CBS culture
collection for supplying reference strains.
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