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Pseudosclerococcum golindoi gen. et sp. nov., a new taxon with apothecial ascomata and a Chalara-like anamorph within the Sclerococcales (Eurotiomycetes)

Authors:
  • King Juan Carlos University / Naturhistoriska Riksmuseet

Abstract and Figures

Sclerococcales encompasses a heterogeneous group of fungi, with most of the species included in the genus Sclerococcum (= Dactylospora). Species of Sclerococcum are characterized by having apothecial ascomata with asci covered by an external hemiamyloid gelatin and a thick euamyloid apical cap, while lacking an inner amyloid wall thickening. Asexual morphs, known for few species, are sporodochial. In this study, we describe Pseudosclerococcum golindoi as a new genus and species sister to Sclerococcum in a multigene phylogeny (nuITS, nuLSU, nuSSU, mtSSU). The fungus produces ascomata similar to those of Sclerococcum, but differs in having cylindrical asci embedded in an overall hemiamyloid gelatin with a fissitunicate dehiscence. Unlike Sclerococcum, Pseudosclerococcum golindoi produces a Chalara-like asexual morph. A possible symbiotic association of P. golindoi with Ascocoryne cylichnium is discussed. The presence of a hemiamyloid gelatin on lateral wall of asci, so far largely overlooked, is reported for some Sclerococcum species. Based on ascal characters and interpretation of the phylogenetic analyses, 14 names assigned to saprotrophic species, previously placed in Dactylospora, are combined in Sclerococcum.
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ORIGINAL ARTICLE
Pseudosclerococcum golindoi gen. et sp. nov., a new taxon
with apothecial ascomata and a Chalara-like anamorph
within the Sclerococcales (Eurotiomycetes)
Ibai Olariaga
1,2
&Joxepo Teres
2
&Joaquín Martín
2
&María Prieto
1
&Hans-Otto Baral
3
Received: 14 February 2019 /Revised: 10 April 2019 /Accepted: 10 May 2019
#German Mycological Society and Springer-Verlag GmbH Germany, part of Springer Nature 2019
Abstract
Sclerococcales encompasses a heterogeneous group of fungi, with most of the species included in the genus Sclerococcum (=
Dactylospora). Species of Sclerococcum are characterized by having apothecial ascomata with asci covered by an external
hemiamyloid gelatin and a thick euamyloid apical cap, while lacking an inner amyloid wall thickening. Asexual morphs, known
for few species, are sporodochial. In this study, we describe Pseudosclerococcum golindoi as a new genus and species sister to
Sclerococcum in a multigene phylogeny (nuITS, nuLSU, nuSSU, mtSSU). The fungus produces ascomata similar to those of
Sclerococcum, but differs in having cylindrical asci embedded in an overall hemiamyloid gelatin with a fissitunicate dehiscence.
Unlike Sclerococcum,Pseudosclerococcum golindoi produces a Chalara-like asexual morph. A possible symbiotic association
of P. golindoi with Ascocoryne cylichnium is discussed. The presence of a hemiamyloid gelatin on lateral wall of asci, so far
largely overlooked, is reported for some Sclerococcum species. Based on ascal characters and interpretation of the phylogenetic
analyses, 14 names assigned to saprotrophic species, previously placed in Dactylospora, are combined in Sclerococcum.
Keywords Bitunicate asci .Dactylospora .Dactylosporaceae .Fissitunicate .Sclerococcum
Introduction
Sclerococcales encompasses a heterogeneous group of fungi that
was recovered as a supported monophyletic group in recent phy-
logenetic studies (Chen et al. 2015; Réblová et al. 2016;Yuetal.
2018).Accordingtoourestimation,partlybasedonDiederich
et al. (2018), Sclerococcales comprises ca. 80 species, including
biotrophic species on lichenized, non-lichenized fungi or leafy
liverworts, wood saprobes and species isolated from the intestinal
flora of beetles (Vargas-Asensio et al. 2014). Most species are
terrestrial, but marine and freshwater species have also been de-
scribed (Jones et al. 1999;Pangetal.2014). Known sexual
morphs are apothecial, with asci generally considered unitunicate
(Hafellner 1979; but see Wood et al. 2016), whereas asexual
morphs consist of sporodochia or effuse colonies with (semi-)
macronematous conidiophores with phialides and catenate co-
nidia (Réblová et al. 2016).
The core group of the Sclerococcales is constituted by the
genus Sclerococcum Fr. : Fr. The long-established concept of
Sclerococcum comprised anamorphic hyphomycetous fungi pro-
ducing blackish sporodochia with multicellular conidia growing
on lichenized fungi (Hawksworth 1975,1979; Diederich et al.
2013). On the other hand, Dactylospora Körb. was a commonly
used genus name for teleomorphic fungi, being well character-
ized by black apothecial ascomata with asci covered with an
external, amyloid apical gelatinous cap instead of an inner amy-
loid wall thickening (tholus, Hafellner 1979). Dactylospora,the
phylogenetic placement of which has been clarified recently
(Pino-Bodas et al. 2017; Diederich et al. 2018), has been merged
with Sclerococcum based on the close phylogenetic relationship
of their type species (Diederich et al. 2018). The monophyly of
Dactylospora has been questioned, since several authors (Pang
Section Editor: Gerhard Rambold
*Ibai Olariaga
ibai.olariaga@urjc.es
1
Department of Biology and Geology, Physics and Inorganic
Chemistry, Rey Juan Carlos University, C/ Tulipán s/n,
28933 Móstoles, Madrid, Spain
2
Aranzadi Society of Sciences, Mycology section, Zorroagagaina 11.
P.C. 200014, Donostia-San Sebastián, Basque Country, Spain
3
Tübingen, Germany
Mycological Progress (2019) 18:895905
https://doi.org/10.1007/s11557-019-01500-7
et al. 2014;Chenetal.2015; Réblová et al. 2016;Yuetal.2018)
recovered Dactylospora as polyphyletic with four species nested
with Sclerococcum, while sequences of Dactylospora imperfecta
(Ellis) Hafellner and D. lobariella (Nyl.) Hafellner, obtained
from cultures, nested in the Chaetothyriales close to Capronia
Sacc. Pino-Bodas et al. (2017) provided for the first time se-
quences of the type species of Dactylospora,D. parasitica
(Flörke) Zopf., and found it was close to Sclerococcum sphaerale
(Ach. : Fr.) Fr., type of Sclerococcum. Diederich et al. (2018)
reached the same conclusion after sequencing two more speci-
mens of D. parasitica, and further showed that at least the
nuLSU sequence of D. lobariella published by Schoch et al.
(2009) was a contaminant. Thus, these authors merged
Scleroccocum and Dactylospora, but transferred only
lichenicolous species of Dactylospora to the older genus name
Sclerococcum. Although molecular data are not available for the
vast majority of species of Sclerococcum, the fact that all species
with asci covered by an amyloid gelatin (Hafellner 1979)forma
monophyletic group substantiates the idea of Sclerococcum be-
ing monophyletic (Pino-Bodas et al. 2017; Diederich et al. 2018).
Although the Sclerococcum-Dactylospora lineage was recov-
ered as a distinct unnamed lineage in the broad multigene phy-
logenies by Schoch et al. (2009) and Chen et al. (2015),
Sclerococcales was first formally recognized by Réblová et al.
(2016). Basal to the core group formed by Sclerococcum, asec-
ond lineage within the Sclerococcales includes two dematiaceous
phialidic fungi (Rhopalophora clavispora (W. Gams) Réblová,
Unter. & W. Gams and Fusichalara minuta Hol.-Jech.) and a
third lineage comprising several strains isolated from digestive
tracts of beetles, basal to the rest. Yu et al. (2018) described the
new aquatic hyphomycete Cylindroconidiis aquaticus X.D. Yu
& H. Zhang and found it to be closely related to R. clavispora
and F
. minuta in the second clade. In this scenario, more hypho-
mycetes are likely to belong to the Sclerococcaceae.
During our surveys in the northern Iberian Peninsula (Basque
Country) in 2006 and 2014, we collected a fungus characterized
by black apothecial ascomata reminiscent of those of
Sclerococcum (formerly Dactylospora), but differing in that asci
were cylindrical and lacked an amyloid gelatinous apical cap.
The asexual morph obtained in culture is of the Chalara type
(dematiaceous hyphomycete) and thus different to the
sporodochial asexual morphs characteristic of Sclerococcum.
Preliminary blast searches and phylogenetic analyses placed
our fungus within the Sclerococcales. Thus, this study investi-
gates the identity and phylogenetic position of this fungus.
Materials and methods
Morphological study
The description of Pseudosclerococcum golindoi is based on
two fresh specimens made in the type locality and one more
dried specimen. Additional material of Sclerococcum
mediterraneum,S. pseudourceolatum,S. stygium and
S. parasiticum was examined for comparison. The macro-
and microscopic description was made from fresh material,
and was completed later by observing material soaked in wa-
ter or in KOH 5%. Structures where measured from fresh
material or from rehydrated material in H
2
O. Only mature
spores discharged from asci without applying external pres-
sure on the cover slip were measured. Spore statistics were
calculated for each collection based on measures of 25 spores
(excluding ornamentation). Abbreviations of statistics refer-
ring to ascospores are L
m
=meanlength;W
m
=meanwidth
and Q
m
=L
m
/W
m
. Mounting reagents used were water, Congo
red in sodium dodecyl sulphate, Lugols solution (IKI) and
KOH 5%. The symbol * refers to living structures and to
dead material. RR and RB refer to types of hemiamyloid re-
action as defined by Baral (2009). Material is deposited in
ARAN-Fungi and UPS herbaria (Thiers 2014). Geographical
coordinates are provided in decimal degrees (WGS 84). The
substrate was identified based on the wood anatomy.
Culture observations
AdicaryoticcultureofPseudosclerococcum golindoi was ob-
tained from shed ascospores on 2% malt extract agar (MEA;
2% malt extract, 2% agar-agar). The plate was sealed with
laboratory film and incubated at room temperature. The cul-
ture was deposited at Westerdijk Fungal Biodiversity Institute,
Utrecht (the Netherlands, CBS).
DNA extraction, PCR amplification and sequencing
DNA was extracted from fresh specimens or two living cul-
tures using DNeasy Plant Mini Kit (Qiagen) according to the
manufacturers instructions. The nuITS was amplified
employing ITS1, ITS4 and ITS5 primers (White et al. 1990),
while primers LR0R (Rehner and Samuels 1994) and LR5
(Vilgalys and Hester 1990)wereusedtoamplifythenuLSU
rDNA region. The nuSSU rDNA region was amplified using
NS1, NS3, NS4 and NS8 primers (White et al. 1990). The
mtSSU region was amplified with mtSSU1 and mtSSU3R
(Zoller et al. 1999) primers. PCR products were subsequently
purified using Exo-sap-IT (USB Corporation, Santa Clara,
CA, USA) and sequenced at Macrogen (www.macrogen.
com), using the same primers as for amplification.
Novel sequences generated in this study (Table 1) were
subjected to BLAST searches to screen for contaminants.
Sequences were edited and assembled using Sequencher v.
4.7 (Gene Codes Corporation Ann Arbor, Michigan, USA)
and were deposited in GenBank (Table 1). ITS sequences
obtained from ascomata and from the culture of P. golindoi
were checked to be identical.
896 Mycol Progress (2019) 18:895905
Blast queries retrieved members of the Eurotiomycetes as
closest matches to sequences of Pseudosclerococcum
golindoi. To explore its phylogenetic affinities preliminarily,
we included its nuLSU sequence in the Eurotiomycetes 7-loci
alignment assembled by Chen et al. (2015), available in
TreeBase (S17149). This alignment was subjected to a maxi-
mum likelihood (ML) analysis using the RAxML HPC2 on
XSEDEtool (Stamatakis 2014) in the CIPRES Science
Gateway (Miller et al. 2010), starting from a random tree. A
GTR-GAMMA model with four rate categories was selected.
For branch confidence, 1000 ML bootstrap replicates were
conducted using rapid bootstrapping. Since P. golindoi nested
in a well-supported clade together with Sclerococcum se-
quences (Dactylospora mangrovei,D. haliotrepha and
Sclerococcum sphaerale), we assembled four alignments
(nuSSU, nuLSU, nuITS, mtSSU) with a more comprehensive
sampling of Sclerococcales. Nucleotide sequences were
aligned manually using AliView (Larsson 2014). Based on
the multigene phylogeny of the Eurotiomycetes by Chen
et al. (2015), Caliciopsis orientalis,Onygena corvina,
Placidium lacinulatum and Trichocoma paradoxa were se-
lected as outgroup. Ambiguous regions of the nuITS and
mtSSU genes were excluded. Each locus was subjected to a
ML analysis, as described above, to assess gene congruence.
A supported clade for one marker was considered to be in
conflict when contradicted with significant support by another
(bootstrap support > 70%). As no conflict was detected, all the
markers were concatenated into a single alignment with four
Table 1 GenBank accession numbers of the species used in the phylogenetic analyses. Newly generated sequences are marked in bold
isolate SSU LSU ITS mtSSU
Caliciopsis orientalis CBS 138.64 DQ471039 DQ470987 KP881690 FJ190654
Cylindroconidiis aquaticus MFLUCC 110294 MH236580 MH236579 MH236576
Fusichalara minuta CBS 709.88 KX537773 KX537758 KX537754 KX537762
Onygena corvina CBS 281.48 FJ358352 AB075355 FJ225792
Placidium lacinulatum AFTOL-ID 2236/2287 EF689847 EF643762 FJ225689
Pseudosclerococcum golindoi ARAN-Fungi 6619 (holotype) MK759887 MK759890 MK759885 MK759897
Rhopalophora clavispora CBS 281.75 KX537771 KX537756 KX537752 KX537760
Sclerococcum ahtii RP23 (H) KY661659 KY661686 KY661630
Sclerococcum ahtii RP127 (H) ––KY661618
Sclerococcum ahtii RP182 (H) ––KY661622 KY661687
Sclerococcum deminutum RP235 (H) ––KY661629
Sclerococcum glaucomarioides RPB275 (LE 261065) KY661660 KY661632 KY661683
Sclerococcum haliotrephum ATCC:MYA-3590 FJ176802 FJ176855 KJ766382
Sclerococcum lobariellum Diederich 18109 MH698499 MH698503
Sclerococcum lobariellum Diederich 17708 MH698498 MH698502
Sclerococcum lobariellum ARAN-Fungi 10091 MK759891 MK759898
Sclerococcum mangrovei CBS 110.444 FJ176836 FJ176890 KJ766383
Sclerococcum parasiticum ARAN-Fungi 2724 MK759888 MK759892 MK759899
Sclerococcum parasiticum RP422 (LE 260868) KY661666 KY661646 KY661690
Sclerococcum parasiticum RP423 (LE 261336) ––KY661629 KY661691
Sclerococcum parasiticum F283586(S) MK759894 MK759901
Sclerococcum parasiticum F283587(S) MK759895 MK759902
Sclerococcum parasiticum ARAN-Fungi A3044025 MK759893 MK759900
Sclerococcum sp. RP391 (H) KY661664 KY661689
Sclerococcum sphaerale Diederich 17279 JX081672 JX081677
Sclerococcum sphaerale Ertz 17425 (BR) JX081674 JX081676
Sclerococcum stygium ARAN-Fungi 3395 MK759889 MK759896 MK759903
Sclerococcum stygium ARAN-Fungi 823 ––MK759886 MK759904
Sclerococcum stygium BHI-F312 (FH) ––MF161218
Sclerococcum vrijmoediae NTOU 4002 KC692152 KC692153 NR_138396
Trichocoma paradoxa CBS 78893-IFO 30659 FJ358354 FJ358290 FJ225782
Beetle-associated isolate INBio 4503Q KM242358 KM242358
Beetle-associated isolate INBio 4513 L KM242300 KM242300
Mycol Progress (2019) 18:895905 897
partitions: nrITS, nrLSU, nrSSU and mtSSU. This dataset was
subjected to ML and Bayesian analyses. The ML analysis was
performed as explained above. The Bayesian analysis was
carried out in MrBayes 3.2.3 (Ronquist et al. 2012;MB),with
four parallel runs of eight Metropolis-coupled Markov Monte
Carlo (MCMCMC) chains for 30 million generations, starting
from a random tree, and sampling one tree every 1000th gen-
eration. The substitution models were sampled across the
GTR space by the MCMCMC analysis (Ronquist et al.
2012). Stationarity was assumed when log-likelihood values
of the chains reached the same stable equilibrium and when
average standard deviation of split frequencies fell below
0.01. A burn-in sample of 60,000 trees was discarded. To
assess branch confidence, a 50% majority rule consensus tree
was computed with the remaining 60,004 trees using the
SUMT command of MrBayes.
Results
The four-locus alignment contained 2916 aligned characters
(395 nuITS, 839 nrLSU, 996 nrSSU, 686 mtSSU) after ex-
cluding ambiguous regions. The MB analysis reached an av-
erage standard deviation of split frequencies of 0.01 after
1,375,000 generations. The single best tree resulting from
the maximum likelihood analysis is shown in Fig. 1, with
ML bootstrap values (ML-BP) and Bayesian Posterior
Probabilities (PP) by nodes.
Nearly all deep nodes of the tree were highly sup-
ported, including the Sclerococcales (ML-BP 100%/PP
1). Within the Sclerococcales, species of Sclerococcum,
with either sporodochial anamorphs or apothecial
ascomata with asci covered with an amyloid gelatinous
apical cap, are encompassed in a highly supported clade
(ML-BP 96%/PP 1, Fig. 1). Sclerococcum comprises
two strongly supported clades: (a) a clade containing
the lichenicolous species S. ahtii,S. deminutum and
S. glaucomarioides, and (b) a clade emcompassing
lichenicolous, marine and wood-inhabiting species, in-
cluding S. sphaerale,typeofSclerococcum and
S. parasiticum, type of Dactylospora. Five specimens
of S. parasiticum form a clade only supported in the
Bayesian analysis (ML-BP 59%/PP 0.97), while speci-
men RP423, also identified as S. parasiticum,isstrong-
ly supported to S. sphaerale.Pseudosclerococcum
golindoi, characterized by a Chalara-like anamorph
and apothecial ascomata without an amyloid apical
cap, is sister to Sclerococcum. Three dematiaceous
phialidic fungi, Rhopalophora clavispora,Fusichalara
minuta and Cylindroconidiis aquaticus, form another
highly supported clade (ML-BP 100%/PP 1) sister to
the Sclerococcum-Pseudosclerococcum clade.
Taxonomy
Pseudosclerococcum Olariaga, Teres, J.M. Martín, M. Prieto
&Baralgen. nov.
Mycobank: MB 829762.
Etymology: referring to the macroscopic similarity to the
sexual morph of species of Sclerococcum and its close phylo-
genetic relationship.
Saprobic or possibly associated with other fungi, terrestrial.
Sexual morph producing apothecial ascomata, disc-shaped, ses-
sile to substipitate, black. Ectal excipulum composed of thin-
walled, isodiametric to vertically elongated, mainly polygonal
cells, forming a textura angularis. Paraphyses distantly septate,
not constricted at septa, unbranched, apical cells claviform. Asci
cylidrical, thin-walled to slightly thick-walled, apically with a
thicker wall, without an apical apparatus, fissitunicate, with a
basal crozier, inamyloid, covered with an IKI red and KOH/
IKI blue gelatin. Ascospores ellipsoid to slightly obovate, 1-sep-
tate, hyaline to brown; spore wall covered with hemisphaerical
warts. Asexual morph Chalara-like. Conidiophores simple,
brown, erect, cylindrical, septate. Conidia in short chains, ellip-
soid to claviform, with a truncate basal end, not septate, thin-
walled, smooth, pale brown to brown.
Pseudosclerococcum differs from Sclerococcum in its cy-
lindrical asci devoid of an amyloid gelatinous cap, a
fissitunicate dehiscence type and a Chalara-like anamorph.
Typ e s p e cies:Pseudosclerococcum golindoi Olariaga,
Teres, J.M. Martín, M. Prieto & Baral.
Pseudosclerococcum golindoi Olariaga, Teres, J.M.
Martín, M. Prieto & Baral sp. nov. (Figs. 2,3,4and 5).
MycoBank: MB 829827.
Etymology: named after José Luis Albizu Golindo,first
co-collector of this species and as homage to a life devoted to
collect and study fungi.
Holotype: Spain, Basque Country, Gipuzkoa, Aia, Mindi
erreka, 43.246226, 2.154849, 117 m asl., on very rotten,
decorticated log of Platanus hispanica lying on the ground,
in riparian forest with Alnus glutinosa,Corylus avellana,
Fraxinus excelsior and planted Platanus hispanica trees, adest
Ascocoryne cylichnium,leg.J.L.Teres,J.Martín,J.M.
Lekuona, P.M. Pasaban, J.I. López Amiano & I. Olariaga,
10 Oct 2014, ARAN-Fungi 6619. Isotype: UPS. Ex-type cul-
ture: CBS 143732.
Other specimens examined: Spain, Basque Country,
Gipuzkoa, Aia, Altzolarats erreka, 43.231422, 2.201852,
127 m asl., on very rotten, decorticated log of Ulmus glabra,
lying on the ground, adest Tomentella crinalis and Ascocoryne
cylichnium, 3 Dec 2006, leg. J.L.Teres, J.L. Albizu & I.
Olariaga, ARAN-Fungi 6618. Same locality and log as the
type, leg. J.L. Teres & J. Martín, 03 Oct 2014, ARAN-Fungi
265; leg. J.L. Teres, 21 Dec 2014, ARAN-Fungi 11725; leg.
J.L. Teres, 28 Dec 2014, ARAN-Fungi 11726 (topotypes).
898 Mycol Progress (2019) 18:895905
Sexual morph apothecial, scattered or densely gregarious,
usually (sub-)sessile. Apothecia, black, shiny, initially cup-
shaped, then plane and finally pulvinate, rarely umbilicate in
the centre, 1.53.5 mm diam. Margin slightly to strongly ele-
vated and distinct in young ascomata, obtuse, regular, finely
rough. Stipe sometimes present, black, glabrous, 00.5 × 0.4
0.5 mm. Apothecial outside black, dull. Epithecium formed
by dark-reddish brown exudates around paraphysis heads,
pigmentation amorphous-resinous, sometimes rather indis-
tinct, not dissolved in KOH. Hymenium below terminal cells,
hyaline to pale reddish brown, 120150 μm thick. Paraphyses
distantly septate (more densely above), slightly constricted at
septa at the lower part, unbranched, 22.5 μm diam, apical
cells usually with pale reddish brown exudate, claviform, 3
4μm diam., without visible cytoplasmic contents, IKI.Asci
cylindrical, slightly thick-walled in the upper part (dead state),
developing a 38μm thick apical wall thickening (tholus)
during ascus maturation; tholus compressed in living asci,
sometimes with a pore-like structure, bitunicate, fissitunicate,
with endotunica strongly elongating during discharge, with a
basal crozier, 8-spored, *120141 × 810.5 μm, 67.5 μm
wide, lateral walls 12μmthick,IKI, KOH/IKI,hyaline;
congophobous gelatin covering the ascus wall hemiamyloid
(IKI red, type RR, KOH/IKI blue). Ascospores ellipsoid to
slightly obovate, rarely slightly curved, homopolar or some-
what heteropolar, thin-walled, 1-septate, hyaline to very pale
sexual morph
sporodochial
asexual morph
phialidic asexual
morph
Fig. 1 Best tree resulting from
the maximum likelihood
analysis of the nuITS-nuLSU-
nuSSU-mtSSU regions of the
Sclerococcales. Maximum
Likelihood bootstrap values
(ML-BP) are shown above
nodes and Bayesian posterior
probabilities (PP) below nodes.
Thickened branches are nodes
with high support in both analyses
(ML-BP 75; PP 95).
Lichenicolous species are marked
in green
Mycol Progress (2019) 18:895905 899
brown inside living asci, brown when overmature, *with a
large guttule and several smaller ones per cell, with a large
guttule per cell, spore wall covered with hemisphaerical warts,
0.30.5 μm high, *(9)1012(13.5) × (4)4.55.5(6) μm
(L
m
=10.311.9, W
m
=4.95.3, Q
m
=2.12.3), (8)912 ×
45μm(L
m
=1010.3, W
m
=4.64.7, Q
m
= 2.2), obliquely
uniseriate in the ascus. Subhymenium of dense, dark reddish-
brown textura intricata. Medullary excipulum dark reddish-
brown to black-brown, 130500 μm thick, composed of two
sharply delimited zones: a) inner part composed of subglobose
to cylindrical, thick-walled cells surrounded by a dark reddish
brown matter, 1.53μm broad, forming a dense textura
intricata and b) outer part composed of subglobose to polyg-
onal or elongated, slightly thick-walled cells with dark reddish
brown pigment at septum edges, 1233 × 1015 μm, forming
atextura angularis. Ectal excipulum bright reddish-brown,
120350 μm thick, with isodiametric to elongated, polygonal
cells, arranged in vertical, vaguely parallel rows, slightly
thick-walled, with dark reddish brown pigment at septum
edges, 1545 × 827 μm, forming a textura angularis.
Outermost cells covered with an amorphous, almost crystal-
loid matter, dark reddish brown. Anchoring hyphae some-
times present near the apothecial base, sparse, brown,
encrusted, septate, thick-walled, 23.5 μm thick. Apothecial
sections losing reddish colour by turning dull brown in KOH,
not releasing dissolved pigment into the medium.
Asexual morph observed in culture. Colonies on MEA 10
15 mm diam in 60 days at 20 °C, superficial, effuse, convex,
hairy, glaucous green, sometimes whitish in the centre.
Reverse blackish green. Margin regular and distinct.
Vegetative hyphae cylindrical, septate, pale greenish brown
to dark brown, smooth, sometimes covered with dark brown
resinous exudates, 2.54μm broad. Conidiophores scattered,
simple, brown, erect, straight, rarely with a sinuous wall,
cylindrical, septate, smooth, sometimes with brown resinous
exudates, thin-walled to slightly thick-walled, 59132 × 3
Fig. 2 Pseudosclerococcum
golindoi. ARAN-Fungi 6618. a
In situ ascomata. ARAN-Fungi
6619 (holotype). bIn situ
ascomata, cclose-up of ascomata,
dascomata co-occurring with
Ascocoryne cylichnium,e
ascomata growing on ascomata of
A. cylichnium,fspot of type
locality where the holotype was
found. All fresh
900 Mycol Progress (2019) 18:895905
5.5 μm. Conidia in short chains, ellipsoid to shortly clavate,
truncate at the basal end with minute inconspicuous marginal
frill, non-septate, thin-walled, smooth, pale brown, with a few
inconspicuous minute guttules, *6.58(8.5) × 34(4.5) μm
(L
m
=7.5,W
m
=3.7,Q
m
=2).
Ecology: on fallen and decorticated logs of angiosperms,
highly decayed, lying on the ground, fruiting on the upper side
or on the underside of the log. Found in the vicinity of
Ascocoryne cylichnium (Tul.) Korf. Both localities from
where P. golindoi is known are narrow ravines along streams,
with well-preserved riparian forests with ecological continuity
and abundant dead wood. Due the mild oceanic climate with
abundant precipitation, both sites host several ferns, bryo-
phytes and lichens of tropical affinity and considered tertiary
relicts, such as Dumortiera hirsuta,Hookeria lucens,
Osmunda regalis,Stegnogramma pozoi, Vandenboschia
speciosa and Woodwardia radicans (Blanco et al. 1998;
Fernández-Palacios et al. 2011;Frahm2012).
Comments
Our attempts to key out the sexual and asexual morphs using
the existing monographs (e.g. Hafellner 1979;NagRajand
Kendrick 1975) and recent literature were unsuccesful.
Pseudosclerococcum golindoi is a unique species due to its
black apothecia, cylindrical asci devoid of an amyloid apical
cap or an amyloid tholus and fissitunicate dehiscence by
means of a strong elongation of the endotunica (Fig. 3f), in
combination with a Chalara-like anamorph.
Concerning the sexual morph, several characters of
Pseudosclerococcum golindoi are reminiscent of Catinella
olivacea (Batsch. : Fr.) Boud. (Dothideomycetes) such as the
sessile darkapothecial ascomata and brown warted ascospores
uniseriately arranged within cylindrical asci. Nevertheless,
C. olivacea differs in having olive green to greenish black
apothecia with a vertically furrowed margin, aseptate spores,
asci without a hemiamyloid gelatin, lacking an apical wall
Fig. 3 Pseudosclerococcum
golindoi. ARAN-Fungi 6619
(holotype), except for j (ARAN-
Fungi 6618). a*Ascospores,
arrow indicates overmature
spores, bhymenium showing
*paraphyses with a brown
resinous exudate around terminal
cells, c*young ascus, d*ascus
with a slightly thick-walled apex,
containing submature ascospores
with incipient ornamentation, e
*ascus with a pore-like structure
at the apex containing mature
ascospores with clear
ornamentation, fempty asci in
which the endotunica has
extruded out of the ectotunica, g
ascus in IKI showing a red
(hemiamyloid) reaction of the
surrounding gelatin, hascus in
KOH/IKI showing a blue reaction
induced by KOH-pretreatment, i
submature ascus with a thick-
walled tholus, containing
immature ascospores still without
ornamentation, jsubmature
ascus with a thick-walled tholus
showing a subapical constriction
with ascoplasm extension, k
*ascus base. Scale bars = 10 μm
Mycol Progress (2019) 18:895905 901
thickening, rupturing by a prominent bivalvate split and a
forcible spore discharge instead of by a strong elongation of
the endotunica (Greif et al. 2007; Baral ined.). Likewise, other
species of Catinella differ also in having aseptate spores and
asci without a hemiamyloid gelatin (Seaver 1946;Gamundí
1981). Catinella olivacea nests in the Dothideomycetes (Greif
et al. 2007)whereasP. golindoi clusters in the Eurotiomycetes
(Fig. 1), and thus, both genera are not closely related.
Our phylogenetic analyses show that P. golindoi is sister
to the Sclerococcum-Dactylospora lineage (Fig. 1).
Ascomata of both have a very similar macroscopic appear-
ance but ascus characters separate them clearly. First, asci
in P. golindoi are covered entirely with a hemiamyloid
gelatin (type RR), while the asci of Sclerococcum have
been described as being covered with an amyloid gelati-
nous apical cap in the literature (Hafellner 1979; Sarrión
et al. 2002; Ihlen et al. 2004; Pino-Bodas et al. 2017).
Interestingly, we observed a hemiamyloid (type RB) reac-
tion of the gelatin on the lateral walls of asci in some
species of Sclerococcum as well (S. bloxamii,S. stygium
and several undescribed species, personal observations by
Baral and Olariaga), while the gelatin on lateral walls of
asci was clearly euamyloid in other species
(S. mediterraneum,S. pseudourceolatum). The
hemiamyloid reaction of some Sclerococcum species ap-
pears to have been largely overlooked so far (e.g. in
Hafellner 1979) and very few authors described it (Joshi
et al. 2010,hemiamyloid, (I blue turning red)). Our ob-
servations suggest that this reaction might be of taxonomic
value in Sclerococcum and it should be further studied.
Second, the dehiscence mechanism is fissitunicate in
P. golindoi as the endoascus elongates and extrudes out
of the exoascus shortly before spore release (Fig. 3f). In
contrast, asci in Sclerococcum are non-fissitunicate and lack
an endoascal thickening and elongation during discharge
(Bellemère and Hafellner 1982). Third, ascal shape differs
between P. golindoi and Pseudosclerococcum. Asci are typ-
ically clavate and up to 60 μmlonginSclerococcum (with
exception of the marine species D. haliotrepha and
D. vrijmoediae), whereas those of P. golindoi are
Fig. 4 Pseudosclerococcum
golindoi. ARAN-Fungi 6619
(holotype). a*Apothecial median
section in water, showing
upraised margin, b*close-up of
ectal excipulum, cectal
excipulum in water, showing an
opaque crystalloid matter on the
surface, dectal excipulum losing
red tones in KOH, eouter part of
medullary excipulum in water, f)
inner part of medullary
excipulum in KOH. Scale bars=
50 μm(a,b), 10 μm(c,d,e,f)
902 Mycol Progress (2019) 18:895905
cylindrical and considerably longer. Further evidence for a
separation between Sclerococcum and P. g o l i n d o i is found
in their asexual morphs. The asexual morph in
Sclerococcum, only known in a few lichenicolous species,
is sporodochial (Diederich et al. 2018) and much unlike the
phialidic hyphomycetous Chalara-like asexual morph in
P. golindoi. Also, ecological adaptations appear to be dif-
ferent between P. golindoi and Sclerococcum. Species of
Sclerococcum are typically drought-tolerant and grow on
exposed, periodically dry substrates whereas P. golindoi
was found on the lower part of damp wood lying on the
forest ground.
The asexual morph of P. golindoi keys out in Chalara
(Nag Raj and Kendrick 1975), due to the absence of
aleuriospores, sterile setae, cylindrical phialides and the shape
of the conidiophores. Within Chalara, it stands alone on ac-
count of its long conidiophores, hyphae with brown resinous
exudates and pale brown claviform conidia with a truncate
base (Nag Raj and Kendrick 1975;McKenzieetal.2002).
Chalara breviclavata Nag Raj & W.B. Kendr. conforms to
P. golindoi in having conidia with a truncate base with a
minute marginal frill, but its hyaline conidia are considerably
larger (1020 × 3.54.5 μm) and its colonies lack green col-
our (Nag Raj and Kendrick 1975)unlikeinP. golindoi.The
only available sequence of C. breviclavata, not a type se-
quence, nests within the Leotiomycetes (Cai et al. 2009).
Chalara brunnipes Nag Raj & W.B. Kendr. produces also
claviform conidia with a truncate base like those of
P. golindoi, but its conidia are shorter, hyaline, the
conidiophores are usually asperulate and have a broadly con-
ical base (Nag Raj and Kendrick 1975). Chalara selaginellae
Farr shares with P. golindoi pale brown conidia, but they
differ in being 1-septate (Nag Raj and Kendrick 1975).
Interestingly, a non-type strain identified as Fusichalara
minuta Hol.-Jech, another species with a Chalara-like asex-
ual morph (Réblová et al. 2016), is part of a lineage basal to
P. golindoi in the Sclerococcales (Fig. 1). Fusichalara
minuta, like other species of Fusichalara, differs morpholog-
ically from P
. golindoi and species conventionally assigned to
Chalara in having phialides with an apical thickening in the
wall (Hughes and Nag Raj 1973;NagRajandKendrick
1975). Judging from this, it is conceivable that more
Chalara-like anamorphic fungi are discovered in the future
as belonging to the Sclerococcales.
Based on all the evidence explained above, broaden-
ing the circumscription of Sclerococcum to encompass a
species with fissitunicate asci devoid of a euamyloid
gelatinous cap and a different Chalara-like asexual
morph, would make Sclerococcum a heterogeneous ge-
nus. On the contrary, morphological, molecular and eco-
logical characters support the distinctiveness of
P. golindoi andwethusproposeanewgenustoappro-
priately accommodate it. So far, P. golindoi has been
found only on two logs, always in close co-occurrence
with Ascocoryne cylichnium.SincemostSclerococcum
species are symbionts growing on other fungi or on
bryophytes, it cannot be ruled out that P. golindoi has
symbiotic interactions with A. cylichnium.
Fig. 5 Pseudosclerococcum
golindoi. Ex-holotype culture
(CBS 143732). aSix-month-old
culture, b*hyphae, c*phialides,
d*conidia, arrow indicating an
inconspicuous marginal frill.
Scale bars = 10 μm
Mycol Progress (2019) 18:895905 903
Nomenclatural novelties
Diederich et al. (2018) transferred various lichenicolous spe-
cies of Dactylospora to Sclerococcum, but most species asso-
ciated with non-lichenized fungi or growing on dead plant
parts were not combined in Sclerococcum. Our analyses
(Fig. 1) suggest that species with clavate to cylindrical asci
covered with an amyloid, gelatinous apical cap, including
species growing on wood, nest in the monophyletic genus
Sclerococcum. Accordingly, we propose the following new
combinations:
Sclerococcum arthrobotryinum (Hafellner) Olariaga,
Teres, J.M. Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829763.
Basionym. Dactylospora arthrobotryina Hafellner, Beih
Nova Hedwigia 62:97 (1979).
Sclerococcum bloxamii (Berk.) Olariaga, Teres, J.M.
Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829764.
Basionym. Patellaria bloxamii Berk. & W. Phillips, Man
Brit Discomyc:361 (1887). [as bloxami].
Dactylospora bloxamii (Berk.) Hafellner, Beih Nova
Hedwigia 62:103 (1979).
Sclerococcum caledonicum (Hafellner) Olariaga, Teres,
J.M. Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829765.
Basionym.DactylosporacaledonicaHafellner, Beih Nova
Hedwigia 62:105 (1979).
Sclerococcum canariense (Kohlm. & Volkm.-Kohlm.)
Olariaga, Teres, J.M. Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829766.
Basionym. Dactylospora canariensis Kohlm. & Volkm.-
Kohlm., Mycotaxon 67:248 (1998).
Sclerococcum epimyces (Tobisch) Olariaga, Teres, J.M.
Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829767.
Basionym. Karschia epimyces Tobisch, Öst bot Z 83:127
(1934).
Dactylospora epimyces (Tobisch) Hafellner, Beih Nova
Hedwigia 62:107 (1979).
Sclerococcum imperfectum (Ellis) Olariaga, Teres, J.M.
Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829768.
Basionym. Patellaria imperfecta Ellis, Bull Torrey Bot
Club 8(2):74 (1881).
Dactylospora imperfecta (Ellis) Hafellner, Beih Nova
Hedwigia 62:114 (1979).
Sclerococcum inopinum (Döbbeler & W.R. Buck)
Olariaga, Teres, J.M. Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829769.
Basionym. Dactylospora inopina Döbbeler & W.R. Buck,
Nova Hedwigia 105(12):88 (2017).
Sclerococcum luridum (Hafellner) Olariaga, Teres, J.M.
Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829770.
Basionym. Dactylospora lurida Hafellner, Beih Nova
Hedwigia 62:120 (1979).
Sclerococcum mediterraneum (Sarrión & Hafellner)
Olariaga, Teres, J.M. Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829772.
Basionym.Dactylospora mediterranea Sarrión &
Hafellner in Sarrión, Hafellner & Burgaz, Lichenologist
34(5):363 (2002).
Sclerococcum pseudourceolatum (Sarrión & Hafellner)
Olariaga, Teres, J.M. Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829812.
Basionym. Dactylospora pseudourceolata Sarrión &
Hafellner in Sarrión, Hafellner & Burgaz, Lichenologist
34(5):365 (2002).
Sclerococcum rubiginosum (Hafellner) Olariaga, Teres,
J.M. Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829813.
Basionym. Dactylospora rubiginosa Hafellner, Beih Nova
Hedwigia 62:127 (1979).
Sclerococcum stipitatum (Hafellner) Olariaga, Teres, J.M.
Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829814.
Basionym. Dactylospora stipitata Hafellner, Beih Nova
Hedwigia 62:137 (1979).
Sclerococcum stygium (Berk. & M.A. Curtis) Olariaga,
Teres, J.M. Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829815.
Basionym. Patellaria stygia Berk. & M.A. Curtis,
Grevillea 4(29):2 (1875).
Dactylospora stygia (Berk. & M.A. Curtis) Hafellner,
Beih Nova Hedwigia 62:137 (1979).
Sclerococcum verruculosum (Hafellner) Olariaga, Teres,
J.M. Martín, M. Prieto & Baral, comb. nov.
MycoBank: MB 829816.
Basionym. Dactylospora verruculosa Hafellner, Beih
Nova Hedwigia 62:151 (1979).
Acknowledgments This paper is dedicated to José Luis Albizu Mayoz
Golindofor his friendship and a life devoted to study fungi in the
Basque Country. Bejondaizula! We wish to thank S. Pérez Ortega for
lending us relevant literature on Sclerococcum and R. Pino-Bodas for
being willing to share unpublished sequences with us.
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Mycol Progress (2019) 18:895905 905
... Based on the results of this preliminary phylogenetic analysis, we constructed a four-locus data set (sensu Olariaga et al. 2019) of Sclerococcales and related taxa within Eurotiomycetes to investigate the exact placement of Polydiscidium. We selected 31 isolates of Sclerococcales and 18 taxa in closely related orders (Coryneliales, Eurotiales, Onygenales, Mycocaliciales, Pyrenulales, Verrucariales) as the ingroup, and Geoglossum nigritum (Geoglossomycetes) as the outgroup taxon (details in TABLE 1). ...
... The species most closely related to S. martynii in our data set is S. stygium, which is easily differentiated micromorphologically by its considerably larger asci (50-75 × 6-10 µm) and larger ascospores (15-20 × 4-6 µm) (Butler 1940 [as Karschia stygia]). Olariaga et al. (2019) noted that some Sclerococcum species have an easily overlooked hemiamyloid gelatinous hymenium, but this was not observed in S. martynii. Other Sclerococcum species with known teleomorphs can be differentiated by ascomatal structure, ascospore morphology, and habitat. ...
... The genus was only known from the asexual S. sphaerale until new taxa sharing the lichenicolous ecology began to be added in the late 20th century (e.g., Diederich 1990). Diederich et al. (2013) revealed a relationship of Sclerococcum to marine species of Dactylospora based on a multilocus phylogenetic analysis, which later broadened to lichenicolous, terrestriallignicolous, and fungicolous Dactylospora species, resulting in formal combinations (Diederich et al. 2013(Diederich et al. , 2018Olariaga et al. 2019). Currently, Sclerococcum is mostly characterized by small (<1 mm) asexual or sexual lichenicolous species, or sexual saprotrophs on wood in marine or terrestrial environments. ...
Article
Full-text available
Polydiscidium is an enigmatic, monotypic, and rarely reported genus of Ascomycota of uncertain placement. The morphologically unique Polydiscidium martynii grows on dead wood and forms compound ascomata composed of thick, black, gelatinous somatic tissue that branches out from a common base. Multiple apothecia are located on the branches, mostly toward the tips, and are composed of 8-spored asci and paraphyses embedded in a gelatinous matrix that turns blue in Melzer's reagent. The species was previously known from only three collections from Guyana (holotype), Trinidad, and the Democratic Republic of the Congo and no sequences exist. Due to its peculiar morphology, taxonomic affinities of Polydiscidium have been debated, with different authors having placed it in Helotiaceae, Leotiaceae, or Leotiomycetes incertae sedis. Recent collections of this species resulting from long-term field work in Guyana and Cameroon led us to revisit the morphology and phylogenetic position of this fungus. Newly generated sequences of P. martynii were added to an Ascomycota-wide six-locus data set. The resulting phylogeny showed Polydiscidium to be a member of order Sclerococcales (Eurotiomycetes). Next, a four-locus (18S, ITS, 28S, mtSSU) phylogenetic reconstruction revealed that Polydiscidium is congeneric with Sclerococcum. A new combination is proposed for this species, Sclerococcum martynii. Micromorphological features, including the gelatinous hymenium composed of asci with amyloid gel cap and septate brown ascospores, are in agreement with Sclerococcum. New combinations are proposed for two additional species: Sclerococcum chiangraiensis and S. fusiformis. Finally, Dactylosporales is considered a later synonym of Sclerococcales.
... Dactylosporaceae accommodates ecologically and morphologically diverse genera, and was reinstated by Diederich et al. (2018) to replace Sclerococcaceae (Réblová et al., 2016). For example, the freshwater genus Cylindroconidiis has holoblastic conidiogenous cells (Yu et al., 2018), while the terrestrial genera Pseudosclerococcum and Rhopalophora are apothecial ascomycetes and dematiaceous phialidic hyphomycetes, respectively (Réblová et al., 2016;Olariaga et al., 2019). The terrestrial and marine genus Sclerococcum (= Dactylospora) has loose sporodochia with catenate conidia or apothecia-like ascomata often growing on lichens or decaying wood (Hawksworth, 1975;Jones et al., 1999;Pang et al., 2014;Pino-Bodas et al., 2017). ...
... The qualities of raw sequences generated in this study were checked with Finch TV version 1.4.0. Based on nucleotide BLAST 2 and previous publications (Raja et al., 2008;Koukol and Kolárová, 2010;Réblová et al., 2012Réblová et al., , 2016Diederich et al., 2013;Pang et al., 2014;Boonmee et al., 2016;Su et al., 2016;Yang et al., 2016;Hernández-Restrepo et al., 2017;Pino-Bodas et al., 2017;Yu et al., 2018;Dayarathne et al., 2019;Ekanayaka et al., 2019;Olariaga et al., 2019), related sequences together with newly generated ones were selected for constructing a phylogenetic tree. All sequences used in this study are listed in Table 1. ...
... Pseudobactrodesmium nests well within the family Dactylosporaceae in our phylogenetic tree of the combined sequence dataset (Figure 1). The unique combination of morphological characters of Pseudobactrodesmium stands apart from other existing genera in this family (Hawksworth, 1975;Ellis, 1976;Jones et al., 1999;Réblová et al., 2016;Yu et al., 2018;Olariaga et al., 2019). Pseudobactrodesmium is morphologically similar to a few dematiaceous hyphomycetous genera with long or short, septate conidia, e.g., Bactrodesmiastrum (Holubová-Jechová, 1984), Bactrodesmium (Ellis, 1971(Ellis, , 1976, Digitodesmium (Boonmee et al., 2016), Distoseptispora , Scolecostigmina ...
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During our ongoing surveys of fungi on submerged wood in the Greater Mekong Subregion, we collected two new species similar to Bactrodesmium longisporum. Pseudobactrodesmium gen. nov. is introduced to accommodate the new species, P. aquaticum, P. chiangmaiensis and B. longisporum is transferred to this genus. Fasciculate conidiophores, enteroblastic conidiogenous cells and subulate to fusiform, phragmoseptate conidia with a tapering apical cell and sheath characterize the genus. Pseudobactrodesmium aquaticum has longer conidia than P. chiangmaiensis. The placement of Pseudobactrodesmium in Dactylosporaceae (Eurotiomycetes) is a novel finding based on analyses of combined LSU, SSU, ITS and RPB2 sequence data. Our study reveals that Pseudobactrodesmium is likely to be a speciose genus with different species in streams around the world.
... Although the Sclerococcum-Dactylospora lineage was recovered as a distinct lineage in the broad multigene phylogenies by Schoch et al. (2009) andChen et al. (2015), Sclerococcomycetidae was first formally recognized by Réblová et al. (2017) including in it the single order Sclerococcales. Later, Dactylospora was merged with Sclerococcum based on the close phylogenetic relationship of their type species (Diederich et al. 2018;Olariaga et al. 2019). Members of this subclass are characterized by apothecial ascomata with unitunicate, non-amyloid asci, covered with an amyloid or hemiamyloid gelatinous c a p . ...
... They are non-lichenized, terrestrial, marine, bryophytic, corticolous, lignicolous, lichenicolous or are associated with beetles as a part of their intestinal microbiota. Olariaga et al. (2019) also pointed out the existence of several dematiaceous phialidic fungi within this clade, several strains isolated from digestive tracts of beetles and a new aquatic hyphomycete. Thus, more hyphomycetes are likely to belong to the Sclerococcomycetidae. ...
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The class Eurotiomycetes (Ascomycota, Pezizomycotina) comprises important fungi used for medical, agricultural, industrial and scientific purposes. Eurotiomycetes is a morphologically and ecologically diverse monophyletic group. Within the Eurotiomycetes, different ascoma morphologies are found including cleistothecia and perithecia but also apothecia or stromatic forms. Mazaediate representatives (with a distinct structure in which loose masses of ascospores accumulate to be passively disseminated) have evolved independently several times. Here we describe a new mazaediate species belonging to the Eurotiomycetes. The multigene phylogeny produced (7 gene regions: nuLSU, nuSSU, 5.8S nuITS, mtSSU, RPB1 , RPB2 and MCM7 ) placed the new species in a lineage sister to Eurotiomycetidae. Based on the evolutionary relationships and morphology, a new subclass, a new order, family and genus are described to place the new species: Cryptocalicium blascoi . This calicioid species occurs on the inner side of loose bark strips of Cupressaceae ( Cupressus , Juniperus ). Morphologically, C. blascoi is characterized by having minute apothecioid stalked ascomata producing mazaedia, clavate bitunicate asci with hemiamyloid reaction, presence of hamathecium and an apothecial external surface with dark violet granules that becomes turquoise green in KOH. The ancestral state reconstruction analyses support a common ancestor with open ascomata for all deep nodes in Eurotiomycetes and the evolution of closed ascomata (cleistothecioid in Eurotiomycetidae and perithecioid in Chaetothyriomycetidae) from apothecioid ancestors. The appropriateness of the description of a new subclass for this fungus is also discussed.
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This article is the 13th contribution in the Fungal Diversity Notes series, wherein 125 taxa from four phyla, ten classes, 31 orders, 69 families, 92 genera and three genera incertae sedis are treated, demonstrating worldwide and geographic distribution. Fungal taxa described and illustrated in the present study include three new genera, 69 new species, one new combination, one reference specimen and 51 new records on new hosts and new geographical distributions. Three new genera, Cylindrotorula (Torulaceae), Scolecoleotia (Leotiales genus incertae sedis) and Xenovaginatispora (Lindomycetaceae) are introduced based on distinct phylogenetic lineages and unique morphologies. Newly described species are Aspergillus lannaensis, Cercophora dulciaquae, Cladophialophora aquatica, Coprinellus punjabensis, Cortinarius alutarius, C. mammillatus, C. quercofocculosus, Coryneum fagi, Cruentomycena uttarakhandina, Cryptocoryneum rosae, Cyathus uniperidiolus, Cylindrotorula indica, Diaporthe chamaeropicola, Didymella azollae, Diplodia alanphillipsii, Dothiora coronicola, Efbula rodriguezarmasiae, Erysiphe salicicola, Fusarium queenslandicum, Geastrum gorgonicum, G. hansagiense, Helicosporium sexualis, Helminthosporium chiangraiensis, Hongkongmyces kokensis, Hydrophilomyces hydraenae, Hygrocybe boertmannii, Hyphoderma australosetigerum, Hyphodontia yunnanensis, Khaleijomyces umikazeana, Laboulbenia divisa, Laboulbenia triarthronis, Laccaria populina, Lactarius pallidozonarius, Lepidosphaeria strobelii, Longipedicellata megafusiformis, Lophiotrema lincangensis, Marasmius benghalensis, M. jinfoshanensis, M. subtropicus, Mariannaea camelliae, Melanographium smilaxii, Microbotryum polycnemoides, Mimeomyces digitatus, Minutisphaera thailandensis, Mortierella solitaria, Mucor harpali, Nigrograna jinghongensis, Odontia huanrenensis, O. parvispina, Paraconiothyrium ajrekarii, Parafuscosporella niloticus, Phaeocytostroma yomensis, Phaeoisaria synnematicus, Phanerochaete hainanensis, Pleopunctum thailandicum, Pleurotheciella dimorphospora, Pseudochaetosphaeronema chiangraiense, Pseudodactylaria albicolonia, Rhexoacrodictys nigrospora, Russula paravioleipes, Scolecoleotia eriocamporesi, Seriascoma honghense, Synandromyces makranczyi, Thyridaria aureobrunnea, Torula lancangjiangensis, Tubeufa longihelicospora, Wicklowia fusiformispora, Xenovaginatispora phichaiensis and Xylaria apiospora. One new combination, Pseudobactrodesmium stilboideus is proposed. A reference specimen of Comoclathris permunda is designated. New host or distribution records are provided for Acrocalymma fci, Aliquandostipite khaoyaiensis, Camarosporidiella laburni, Canalisporium caribense, Chaetoscutula juniperi, Chlorophyllum demangei, C. globosum, C. hortense, Cladophialophora abundans, Dendryphion hydei, Diaporthe foeniculina, D. pseudophoenicicola, D. pyracanthae, Dictyosporium pandanicola, Dyfrolomyces distoseptatus, Ernakulamia tanakae, Eutypa favovirens, E. lata, Favolus septatus, Fusarium atrovinosum, F. clavum, Helicosporium luteosporum, Hermatomyces nabanheensis, Hermatomyces sphaericoides, Longipedicellata aquatica, Lophiostoma caudata, L. clematidisvitalbae, Lophiotrema hydei, L. neoarundinaria, Marasmiellus palmivorus, Megacapitula villosa, Micropsalliota globocystis, M. gracilis, Montagnula thailandica, Neohelicosporium irregulare, N. parisporum, Paradictyoarthrinium difractum, Phaeoisaria aquatica, Poaceascoma taiwanense, Saproamanita manicata, Spegazzinia camelliae, Submersispora variabilis, Thyronectria caudata, T. mackenziei, Tubeufa chiangmaiensis, T. roseohelicospora, Vaginatispora nypae, Wicklowia submersa, Xanthagaricus necopinatus and Xylaria haemorrhoidalis. The data presented herein are based on morphological examination of fresh specimens, coupled with analysis of phylogenetic sequence data to better integrate taxa into appropriate taxonomic ranks and infer their evolutionary relationships.
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The newly discovered systematic placement of Bactrodesmium abruptum, the lectotype species of the genus, prompted a re-valuation of the traditionally broadly conceived genus Bactrodesmium. Fresh material, axenic cultures and new DNA sequence data of five gene regions of six species, i.e. B. abruptum, B. diversum, B. leptopus, B. obovatum, B. pallidum and B. spilomeum, were studied. Bactrodesmium is a strongly resolved lineage in the Savoryellales (Sordariomycetes), supported by Bayesian and Maximum Likelihood methods. The genus Bactrodesmium is emended and delimited to hyphomycetes characterised by sporodochial conidiomata, mononematous often fasciculate conidiophores, holoblastic conidiogenesis and acrogenous, solitary, dry, pigmented, transversely or rarely longitudinally septate conidia. The conidia are seceding rhexolytically, exhibiting multiple secession patterns. An identification key to 35 species accepted in Bactrodesmium is given, providing the most important diagnostic characters. Novel DNA sequence data of B. longisporum and B. stilboideum confirmed their placement in the Sclerococcales (Eurotiomycetes). For other Bactrodesmium, molecular data are available for B. cubense and B. gabretae, which position them in the Dothideomycetes and Leotiomycetes, respectively. All four species are excluded from Bactrodesmium and segregated into new genera, Aphanodesmium, Gamsomyces and Kaseifertia. Classification of other 20 species and varieties not recognised in the genus is discussed. Based on new collections of Dematiosporium aquaticum, the type species of Dematiosporium, the genus is emended to accommodate monodictys-like freshwater lignicolous fungi of the Savoryellales characterised by effuse colonies, holoblastic conidiogenous cells and dictyosporous, pigmented conidia with a pore in each cell. Study of additional new collections, cultures and DNA sequence data revealed several unknown species, which are proposed as taxonomic novelties in the Savoryellales and closely related Pleurotheciales. Ascotaiwania latericolla, Helicoascotaiwania lacustris and Pleurotheciella erumpens are described from terrestrial, lentic and lotic habitats from New Zealand and France, respectively. New combinations are proposed for Helicoascotaiwania farinosa and Neoascotaiwania fusiformis. Relationships and systematics of the Savoryellales are discussed in the light of recent phylogenies and morphological patterns newly linked with the order through cultural studies.
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Neotropical mountain forests are characterized by having hyperdiverse and unusual fungi inhabiting lichens. The great majority of these lichenicolous fungi (i.e., detectable by light microscopy) remain undescribed and their phylogenetic relationships are mostly unknown. This study focuses on lichenicolous fungi inhabiting the genus Lobariella ( Peltigerales ), one of the most important lichen hosts in the Andean cloud forests. Based on molecular and morphological data, three new genera are introduced: Lawreyella gen. nov. ( Cordieritidaceae , for Unguiculariopsis lobariella ), Neobaryopsis gen. nov. ( Cordycipitaceae ), and Pseudodidymocyrtis gen. nov. ( Didymosphaeriaceae ). Nine additional new species are described ( Abrothallus subhalei sp. nov., Atronectria lobariellae sp. nov., Corticifraga microspora sp. nov., Epithamnolia rugosopycnidiata sp. nov., Lichenotubeufia cryptica sp. nov., Neobaryopsis andensis sp. nov., Pseudodidymocyrtis lobariellae sp. nov., Rhagadostomella hypolobariella sp. nov., and Xylaria lichenicola sp. nov.). Phylogenetic placements of 13 lichenicolous species are reported here for Abrothallus , Arthonia , Globonectria , Lawreyella , Monodictys , Neobaryopsis , Pseudodidymocyrtis , Sclerococcum , Trichonectria and Xylaria . The name Sclerococcum ricasoliae comb. nov. is reestablished for the neotropical populations formerly named S. lobariellum ( Sclerococcales ). A key to sexual and asexual states of 40 species of lobariellicolous ascomycetous fungi is provided. Teleomorph-anamorph connections were established for several species using molecular methods and/or visual observations in nature. Additionally, we found that the anamorphic species Cornutispora ophiurospora inhabiting Lobariella was often accompanied by ascomata of Spirographa . Results of phylogenetic analyses, including newly generated sequences of several Cornutispora and Spirographa species inhabiting various host lichens, support the conclusion that Cornutispora is a synonym of Spirographa . Our Maximum Likelihood inference based on multiple loci show that all studied Spirographa (including Cornutispora ) belong to a new lineage within Ostropales . Based on these highly supported phylogenetic placements and the distinct character states of their conidiomata, in comparison with other Lecanoromycetes , a new family is proposed – Spirographaceae fam. nov. This new lineage includes broadly distributed mycoparasites, inhabiting various lichen and fungal hosts, and representing an early diversification event preceding the lichen-forming clade of Fissurinaceae , Gomphillaceae and Graphidaceae . Two lichenicolous species, Asteroglobulus giselae and Pleoscutula arsenii , were found to be nested within the Spirographa clade, and their teleomorph-anamorph connections were confirmed based on genotypic and phenotypic data. This phylogenetic result is corroborated by their highly similar ascomata anatomy. Together these results strongly indicate that both species are congeneric with Spirographa . As a result, four new species ( S. aggregata sp. nov., S. galligena sp. nov., S. maroneae sp. nov., and S. parmotrematis sp. nov.) and 15 new combinations are proposed ( Spirographa ascaridiella comb. nov., S. arsenii comb. nov., S. ciliata comb. nov., S. giselae comb. nov., S. herteliana comb. nov., S. hypotrachynae comb. nov., S. intermedia comb. nov., S. lichenicola comb. nov., S. limaciformis comb. nov., S. ophiurospora comb. nov., S. pittii comb. nov., S. pyramidalis comb. nov., S. triangularis comb. nov., S. tricupulata comb. nov., and S. vermiformis comb. nov.). Species of the genus Spirographa , as outlined here, are strongly host-specific, mainly at the generic level of their host. Some host genera can harbour more than one Spirographa species.
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Lichenicolous fungi represent a highly specialized and successful group of organisms that live exclusively on lichens, most commonly as host-specific parasites, but also as broad-spectrum pathogens, saprotrophs or commensals. We present here the most recent update to the classification of lichenicolous fungi in the Ascomycota and Basidiomycota to genus level, arranged phylogenetically according to published classifications. For each genus, all known lichenicolous taxa (obligately lichenicolous taxa, lichenicolous lichens, and facultatively lichenicolous taxa) are listed, along with information about types, synonyms, pertinent literature and whether or not molecular data are available for any of the listed species. The number of accepted lichenicolous fungi is now 2319, with 2000 obligately lichenicolous species, subspecies or varieties, 257 lichenicolous lichens and 62 facultatively lichenicolous taxa. These species are found in 10 different classes of Fungi (Ascomycota and Basidiomycota), 55 orders, 115 families and 397 genera. The 2319 total taxa is an increase from the 1559 total species reported in the last published catalogue in 2003, and a larger number than the approximately 1800 reported in the most recent online checklist (www.lichenicolous.net) posted in January 2018. Of the total number of taxa, 2219 (96%) are ascomycetes and 100 (4%) are basidiomycetes. Of the 397 genera containing lichenicolous species, c. 50% (198) are entirely lichenicolous. In addition, six families (Abrothallaceae, Adelococcaceae, Cyphobasidiaceae, Obryzaceae, Polycoccaceae, Sarcopyreniaceae) and two orders (Abrothallales, Cyphobasidiales) are entirely lichenicolous. Sequence information is available for lichenicolous species in 128 (32%) of the 397 genera containing lichenicolous species, and in 56 (28%) of the 198 entirely lichenicolous genera. Many species are known from only one host lichen, but it is likely that broader host ecologies will be discovered as new sequence information is obtained from ongoing microbiome studies. Phaeopyxis Rambold & Triebel is considered as a new synonym of Bachmanniomyces D.Hawksw., resulting in five new combinations B. australis (Rambold & Triebel) Diederich & Pino-Bodas (≡ P. australis), B. carniolicus (Arnold) Diederich & Pino-Bodas (≡ Biatora carniolica), B. muscigenae (Alstrup & E.S.Hansen) Diederich & Pino-Bodas (≡ P. muscigenae), B. punctum (A.Massal.) Diederich & Pino-Bodas (≡ Nesolechia punctum) and B. varius (Coppins, Rambold & Triebel) Diederich & Pino-Bodas (≡ P. varia). As a consequence of a phylogenetic analysis including new sequences, Dactylospora Körb. is regarded as a new synonym of Sclerococcum Fr.: Fr., resulting in one new name (S. acarosporicola Ertz & Diederich) and 46 new combinations. Sclerococcaceae Réblová, Unter. & W.Gams is considered as a new synonym of Dactylosporaceae Bellem. & Hafellner. The new Sclerococcum ophthalmizae Coppins is described. Sclerophyton occidentale Herre is lectotypified on the lichenicolous fungus present in the type specimen and becomes a younger synonym of Sclerococcum parasiticum. A replacement name is Arthonia polydactylonis Diederich & Ertz (≡ A. ceracea). Further new combinations are Abrothallus lobariae (Diederich & Etayo) Diederich & Ertz (≡ Phoma lobariae), A. psoromatis (Zhurb. & U. Braun) Diederich & Zhurb. (≡ P. psoromatis), Asteroglobulus pyramidalis (Etayo) Diederich (≡ Cornutispora pyramidalis), Didymocyrtis grumantiana (Zhurb. & Diederich) Zhurb. & Diederich (≡ Phoma grumantiana), Epithamnolia atrolazulina (Etayo) Diederich (≡ Hainesia atrolazulina), Gyalolechia epiplacynthium (Etayo) Diederich (≡ Fulgensia epiplacynthium), Nesolechia doerfeltii (Alstrup & P.Scholz) Diederich (≡ Phacopsis doerfeltii), N. falcispora (Triebel & Rambold) Diederich (≡ P. falcispora), N. oxyspora var. fusca (Triebel & Rambold) Diederich (≡ P. oxyspora var. fusca), Preussia peltigerae (Brackel) Diederich (≡ Sporormiella peltigerae), Scutula curvispora (D.Hawksw. & Miadl.) Diederich (≡ Libertiella curvispora), S. didymospora (D.Hawksw. & Miadl.) Diederich (≡ L. didymospora), Stigmidium haesitans (Nyl.) Diederich (≡ Verrucaria haesitans), and S. parvum (Henssen) Diederich (≡ Pharcidia parvum). © 2018 by The American Bryological and Lichenological Society, Inc.
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During our ongoing investigations of freshwater fungi in Thailand, Cylindroconidiis aquaticus gen. et sp. nov. is described in family Sclerococcaceae (Eurotiomycetes), based on morphological characters and analyses of LSU, SSU and ITS sequence data. Its closest relatives are Rhopalophora and Fusichalara minuta. The new genus is characterized by mononematous, unbranched conidiophores, polyblastic, terminal or intercalary conidiogenous cells, and cylindrical,1-septate conidia. It is the first report of freshwater fungi encountered in Sclerococcaceae.
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Though most of the lichenicolous fungi belong to the Ascomycetes, their phylogenetic placement based on molecular data is lacking for numerous species. In this study the phylogenetic placement of 19 species of lichenicolous fungi was determined using four loci (LSU rDNA, SSU rDNA, ITS rDNA and mtSSU). The phylogenetic analyses revealed that the studied lichenicolous fungi are widespread across the phylogeny of Lecanoromycetes. One species is placed in Acarosporales, Sarcogyne sphaerospora; five species in Dactylosporaceae, Dactylospora ahtii, D. deminuta, D. glaucoides, D. parasitica and Dactylospora sp.; four species belong to Lecanorales, Lichenosticta alcicorniaria, Epicladonia simplex, E. stenospora and Scutula epiblastematica. The genus Epicladonia is polyphyletic and the type E. sandstedei belongs to Leotiomycetes. Phaeopyxis punctum and Bachmanniomyces uncialicola form a well supported clade in the Ostropomycetidae. Epigloea soleiformis is related to Arthrorhaphis and Anzina. Four species are placed in Ostropales, Corticifraga peltigerae, Cryptodiscus epicladonia, C. galaninae and C. cladoniicola comb. nov. (= Lettauia cladoniicola). Three new species are described, Dactylospora ahtii, Cryptodiscus epicladonia and C. galaninae. © 2017 Naturalis Biodiversity Center & Westerdijk Fungal Biodiversity Institute.
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Species belonging to the Coryneliaceae and parasitizing Podocarpaceae hosts were collected from different locations in South Africa and studied morphologically by light microscopy and molecularly by obtaining partial nrDNA (ITS-1/5.8S/ITS-2, 18S and 28S) gene sequences. The position of the Coryneliaceae within the Eurotiomycetidae was not confirmed and a new subclass, Coryneliomycetidae, was introduced. While Eurotiomycetidae usually form cleistothecia/gymnothecia with evanescent, unitunicate asci, and Chaetothyriomycetidae mostly perithecia with bitunicate/fissitunicate to evanescent asci, Coryneliomycetidae form pseudothecial mazaedial ascomata, initially with double-walled asci with the outer layer deliquescing, resulting in passive ascospore release. The Coryneliomycetidae thus occupies a unique position in the Eurotiomycetes. Furthermore, epitypes were designated for Corynelia uberata, the type species of Corynelia (type genus of the family, order and subclass), Lagenulopsis bispora, the type species of Lagenulopsis, and Tripospora tripos the type species of Tripospora, with Lagenulopsis and Tripospora confirmed as belonging to the Coryneliaceae. Corynelia uberata resolved into three clades, one on Afrocarpus (= Podocarpus) falcatus and A. gracilior, and two clades occurring on P. latifolius, herein described as C. africana and C. fructigena. Morphologically these three species are not readily distinguishable, although they differ in spore dimensions, ascomata shape, ornamentation and DNA phylogeny. It is likely that several more species from other parts of the world are currently erroneously placed in C. uberata. © 2015-2016 Naturalis Biodiversity Center & Centraalbureau voor Schimmelcultures.
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1. The species of Karschia s. lat. are taxonomically treated. 2. In a general part technical problems, the history of the genus, the characters used for the delimitation of the genera are discussed, a new ascus-type characteristic for Dactylospora is described and annotations to biology and distribution patterns are given. Essential results seem to be that all the species of the lecanoralean genera treated live together or close to algae and that the lichenicolous and parasitic species of Karschia s. lat. are specific to at least groups of hosts. 3. Keys to the genera with apothecial or pseudothecial fruiting bodies as well as to all species and varieties of Karschia s. lat. are given. 4. Sixty-eight species and varieties belonging to 20 different genera and 3 species, the generic position of which is not clear at this time, are treated. Four genera, 1 subgenus and rnany infrageneric taxa are recognized as synonyms. Six genera, 9 species and 5 varieties are described as new and 30 new combinations are proposed. 5. The species of Karschia s. lat. belong to the following 19 genera: Buellia, Epilichen, Rhizocarpon, Rinodina, Dactylospora, Buelliella, Colensoniella, Cycloschizon, Dothidea, Eutryblidiella, Gibbera, Heterosphaeriopsis, Karschia s. str., Poetschia, Pseudodiscus, Rhizodiscina, Rhizogene, Schrakia, and Stratisporella. 6. Line drawings, all are originals of the author, are added to many descriptions of accepted taxa. 7. Twenty-five not seen, doubtful or to be exduded taxa are listed. 8. Both an index of the hosts and an index of the species as well as infraspecific taxa are given.
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We present a 6-gene, 420-species maximum-likelihood phylogeny of Ascomycota, the largest phylum of Fungi. This analysis is the most taxonomically complete to date with species sampled from all 15 currently circumscribed classes. A number of superclass-level nodes that have previously evaded resolution and were unnamed in classifications of the Fungi are resolved for the first time. Based on the 6-gene phylogeny we conducted a phylogenetic informativeness analysis of all 6 genes and a series of ancestral character state reconstructions that focused on morphology of sporocarps, ascus dehiscence, and evolution of nutritional modes and ecologies. A gene-by-gene assessment of phylogenetic informativeness yielded higher levels of informativeness for protein genes (RPB1, RPB2, and TEF1) as compared with the ribosomal genes, which have been the standard bearer in fungal systematics. Our reconstruction of sporocarp characters is consistent with 2 origins for multicellular sexual reproductive structures in Ascomycota, once in the common ancestor of Pezizomycotina and once in the common ancestor of Neolectomycetes. This first report of dual origins of ascomycete sporocarps highlights the complicated nature of assessing homology of morphological traits across Fungi. Furthermore, ancestral reconstruction supports an open sporocarp with an exposed hymenium (apothecium) as the primitive morphology for Pezizomycotina with multiple derivations of the partially (perithecia) or completely enclosed (cleistothecia) sporocarps. Ascus dehiscence is most informative at the class level within Pezizomycotina with most superclass nodes reconstructed equivocally. Character-state reconstructions support a terrestrial, saprobic ecology as ancestral. In contrast to previous studies, these analyses support multiple origins of lichenization events with the loss of lichenization as less frequent and limited to terminal, closely related species.
Book
We began with the intention of monographing Chalara and very similar fungi (Excioconidium, etc.). We soon extended the scope of our study to encompass those dimorphic imperfect genera with Chalara-like phialides (Thielaviopsis, Chalaropsis, Stilbochalam, Hughesiella), then to cover two other genera with Chalara-like phialides but having characteristic ancillary sterile structures (Chaetochalara, Sporoschisma). Finally we considered several other genera with phialides having more or less cylindrical collarettes and deep-seated conidiogenous loci (Bloxamia, Endosporostilbe, Ascoconidium, Sporendocladia) or whose descriptions raised the suspicion that they had such attributes (Endoconidium, Columnophora, Milowia). To have extended the study further would have meant including phialidic genera which, although often having relatively deep collarettes, were otherwise not particularly morphologically comparable to Chalara (Catenularia, Phialophora, Phialocephala, Sporoschismopsis). Our study, which began as an attempt to revise one genus, thus finished up by considering fifteen existing generic names (five reduced to synonymy, one to the status of nomen dubium) and adding one additional generic name (first published elsewhere). Readers will find in the taxonomic part of this book a compilation of descriptions and illustrations of species of Sporoschisma and Fusichalara. Some may consider these redundant, especially since we have added nothing significantly new to what is already known about them: our purpose in including them is solely to bring together all taxonomic data relating to Chalara and other closely allied genera in a single source-book.
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A new genus Rhopalophora is described for Phialophora clavispora, a lignicolous species formerly placed in Phialophora section Catenulatae that possesses pigmented conidiophores, phialides with a single conidiogenous locus that occasionally appear as schizophialides, and clavate, aseptate conidia arranged in chains or sometimes in heads. Sexual morphs are not known for this taxon. Phylogenetic analysis of DNA sequences from five loci (nucSSU, ITS, nucLSU, mitSSU, rpb1 and rpb2) of this and related fungi supports the introduction of a new family, Sclerococcaceae, for which we establish the order Sclerococcales. This order belongs to the new subclass Sclerococcomycetidae, a strongly supported clade within the Eurotiomycetes that is basal to a lineage containing the Chaetothyriomycetidae, Coryneliomycetidae and Eurotiomycetidae. Rhopalophora clavispora fits in this new family and is closely related to an isolate of Fusichalara minuta. The Sclerococcales also encompass marine, lignicolous species of Dactylospora, two species of the lichenicolous genus Sclerococcum, and a lineage comprised of strains from the digestive tracts of Neotropical wood-inhabiting beetles. We confirm that Dactylospora is polyphyletic; the phylogenetic placement of D. parasitica, the generic type, remains unknown.
Article
The Euantennariaceae and the Metacapnodiaceae are two new families of sooty moulds which have hyphomycetous conidial states, distinguishing them from the Capnodlaceae sensu stricto in which the conidial states are pyenidial. In the Euantennariaceae, all species investigated have the Dothidea centrum development while in the Metacapnodiaceae, centrum development is of the Pleospora type. The Pleospora centrum of the Metacapnodiaceae differs from the centrum of Pleospora herbarum only in one minor detail; in the Metacapnodiaceae, the pseudoparaphyses remain relatively short and do not reach the base of the centrum locule.