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

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.

Full text

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) 18:895–905
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, Lugol’s 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
manufacturer’s 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) 18:895–905

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
XSEDE’tool (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 11–0294 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) 18:895–905 897

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) 18:895–905

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, 1.5–3.5 mm diam. Margin slightly to strongly ele-
vated and distinct in young ascomata, obtuse, regular, finely
rough. Stipe sometimes present, black, glabrous, 0–0.5 × 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, 120–150 μm thick. Paraphyses
distantly septate (more densely above), slightly constricted at
septa at the lower part, unbranched, 2–2.5 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 †3–8μm 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, *120–141 × 8–10.5 μm, †6–7.5 μm
wide, lateral walls †1–2μmthick,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) 18:895–905 899

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,
0.3–0.5 μm high, *(9–)10–12(−13.5) × (4–)4.5–5.5(−6) μm
(L
m
=10.3–11.9, W
m
=4.9–5.3, Q
m
=2.1–2.3), †(8–)9–12 ×
4–5μm(L
m
=10–10.3, W
m
=4.6–4.7, 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, 130–500 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, 1.5–3μm 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, 12–33 × 10–15 μm, forming
atextura angularis. Ectal excipulum bright reddish-brown,
120–350 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, 15–45 × 8–27 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, 2–3.5 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, 2.5–4μm 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, 59–132 × 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) 18:895–905

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, *6.5–8(−8.5) × 3–4(−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, f†empty asci in
which the endotunica has
extruded out of the ectotunica, g
†ascus in IKI showing a red
(hemiamyloid) reaction of the
surrounding gelatin, h†ascus 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, j†submature
ascus with a thick-walled tholus
showing a subapical constriction
with ascoplasm extension, k
*ascus base. Scale bars = 10 μm
Mycol Progress (2019) 18:895–905 901

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, c†ectal
excipulum in water, showing an
opaque crystalloid matter on the
surface, d†ectal excipulum losing
red tones in KOH, e†outer 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) 18:895–905

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 (10–20 × 3.5–4.5 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) 18:895–905 903

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(1–2):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
‘Golindo’for 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.
References
Baral H-O (2009) Iodine reaction in Ascomycetes: why is Lugol's solu-
tion superior to Melzer's reagent?. https://in-vivo-veritas.de/articles/
iodine-reaction-in-ascomycetes-why-is-lugols-solution-superior-to-
melzers-reagent/. Accessed 2 Apr 2019
904 Mycol Progress (2019) 18:895–905

Bellemère A, Hafellner J (1982) L’ultrastructure des asques de genre
Dactylospora (Discomycetes) et son interet taxonomique.
Cryptogam Mycol 3:71–93
Blanco E, Casado MA, Costa M, Escribano R, García M, Génova M,
Gómez A, Gómez F, Moreno JC,Morla C, Regato P, Sainz H (1998)
Los bosques ibéricos: una interpretación geobotánica. Editorial
Planeta, Barcelona
Cai L, Wu W-P, Hyde KD (2009) Phylogenetic relationships of Chalara
and allied species inferred from ribosomal DNA sequences. Mycol
Progress 8:133–143
Chen K-H, Miadlikowska J, Arnold AE, U’RenJM,GayaE,GueidanC,
Lutzoni F (2015) Phylogenetic analyses of eurotiomycetous endo-
phytes reveal their close affinities to Chaetothyriales, Eurotiales, and
a new order –Phaeomoniellales. Mol Phylogenet Evol 85:117–130
Diederich P, Ertz D, Lawrey JD, Sikaroodi M, Untereiner WA (2013)
Molecular data place the hyphomycetous lichenicolous genus
Sclerococcum close to Dactylospora (Eurotiomycetes) and
S. parmeliae in Cladophialophora (Chaetothyriales). Fungal
Divers 58:61–72
Diederich P, Lawrey JD, Ertz D (2018) The 2018 classification and
checklistof lichenicolous fungi, with 2000 non-lichenized, obligate-
ly lichenicolous taxa. Bryologist 121(3):340–425
Fernández-Palacios JM, de Nascimento L, Otto R, Delgado JD, García-
del-Rey E, Arévalo JR, Whittaker RJ (2011) A reconstruction of
Palaeo-Macaronesia, with particular reference to the long-term bio-
geography of the Atlantic island laurel forests. J Biogeogr 38(2):
226–246
Frahm JP (2012) The phytogeography of European bryophytes. Bot
Serbica 36(1):23–36
Gamundí IJ (1981) On Lagerheima Sacc. and Bulgariella Karst. Sydowia
34:82–93
Greif MD, Gibas CF, Tsuneda A, Currah R (2007) Ascoma development
and phylogeny of an apothecioid dothideomycete, Catinella
olivacea. Am J Bot 94(11):1890–1899
Hafellner J (1979) Karschia. Revision einer Sammelgattung an der
grenze von lichenisierten und nichtlichenisierten Ascomyceten.
Beih Nova Hedwigia 62:1–248
Hawksworth DL (1975) A revision of lichenicolous fungi accepted by
Keissler in Coniothecium. Trans Br Mycol Soc 65:219–238
Hawksworth DL (1979) The lichenicolous hyphomycetes. Bull Br Mus
Nat Hist Bot Ser 6:183–300
Hughes SJ, Nag Raj TR (1973) New Zealand Fungi. 20. Fusichalara gen.
nov. N Z J Bot 11:661–671
Ihlen G, Holien H, Tønsberg T (2004) Two new species of Dactylospora
(Dactylosporaceae, Lecanorales), with a key to the known species in
Scandinavia. Bryologist 107(3):357–362
Jones EBG, Abdel-WahabMA ASA, Hsieh SY (1999) Dactylospora
mangrovei sp. nov. (Discomycetes, ascomycota) from mangrove
wood. Mycoscience 40:317–320
Joshi Y, Knudsen K, Want XY, Hur J-S (2010) Dactylospora
glaucomarioides (Ascomycetes, Dactylosporaceae): a
Lichenicolous Fungus New to South Korea. Mycobiology 39(4):
321–322
Larsson A (2014) AliView: a fast and lightweight alignment viewer and
editor for large data sets. Bioinformatics 30:3276–3278
McKenzie EHC, Pinnoi A, Wong MKM, Hyde KD, Jones EBG (2002)
Two new hyaline Chalara species, and a key to species described
since 1975. Fungal Divers 11:129–139
Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES science
gateway for inference of large phylogenetic trees. In: Proceedings of
the gateway computing environments workshop (GCE), New
Orleans, LA, pp 1–8
Nag Raj TR, Kendrick B (1975) A monograph of Chalara and allied
genera. Wilfrid Laurier University Press, Waterloo
Pang KL, Guo S-Y, Alias SA, Hafellner J, EBG Johnes (2014) A new
species of marine Dactylospora and its phylogenetic affinities with-
in the Eurotiomycetes, Ascomycota. Bot Mar 57:315–321
Pino-Bodas R, Zhurbenko MP, Stenroos S (2017) Phylogenetic place-
ment within Lecanoromycetes of lichenicolous fungi associated with
Cladonia and some other genera. Persoonia 39:91–117
Réblová M, Untereiner WA, Štěpánek V, Gams W (2016) [2017]
Disentangling Phialophora section Catenulatae: disposition of taxa
with pigmented conidiophores and recognition of a new subclass,
Sclerococcomycetidae (Eurotiomycetes). Mycol Prog 16:27–46.
https://doi.org/10.1007/s11557-016-1248-y
Rehner S, Samuels GJ (1994) Taxonomy and phylogeny of Gliocladium
analyzed from nuclear large subunits ribosomal DNA sequences.
Mycol Res 98:625–634
Ronquist F, Teslenko M, van der Mark P, Ayres D, Darling A, Hohna S,
Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2:
efficient Bayesian phylogenetic inference and model choice across a
large model space. Syst Biol 61:539–554
Sarrión FJ, Hafellner J, Burgaz AR (2002) Three new species of the genus
Dactylospora in Spain. Lichenologist 34(5):361–368
Schoch CL, Sung GH, López-Giráldez F et al (2009) The Ascomycota
tree of life: a phylum-wide phylogeny clarifies the origin and evo-
lution of fundamental reproductive and ecological traits. Syst Biol
58:224–239
Seaver FJ (1946) Photographs and descriptions of cup-fungi: XLI.
Catinella nigro-olivacea. Mycologia 38(4):473–476
Stamatakis A (2014) RAxMLVersion 8: a tool for phylogenetic analysis
and post-analysis of large phylogenies. Bioinformatics 30:1312–
1313
Thiers B (2014) (continuously updated): index Herbariorum: a global
directory of public herbaria and associated staff, New York
Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/
ih/ Last accession: 25/01/2018
Vargas-Asensio G, Pinto-Tomas A, Rivera B, Hernandez M, Hernandez
C, Soto-Montero S, Murillo C, Sherman DH, Tamayo-Castillo G
(2014) Uncovering the cultivable microbial diversity of Costa
Rican beetles and its ability to break down plant cell wall compo-
nents. PLoS One 9(11):e113303
Vilgalys R, Hester M (1990) Rapid genetic identification and mapping of
enzymatically amplified ribosomal DNA from several Cryptococcus
species. J Bacteriol 172:4238–4246
White TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct
sequencing of fungal ribosomal RNA genes for phylogenetics. In:
Innis MA, Gelfand DH, Sninsky JJ, White TJ (eds) PCR protocols: a
guide to methods and applications. Academic Press, New York, pp
315–322
Wood AR, Damm U, van der Linde EJ, Groenewald JZ, Cheewangkoon
R, Crous PW (2016) Finding the missing link: resolving the
Coryneliomycetidae within Eurotiomycetes. Persoonia 37:37–56
Yu X-D, Dong W, Bhat DJ, Boonmee S, Zhang D, Zhang H (2018)
Cylindroconidiis aquaticus gen. et sp. nov., a new lineage of aquatic
hyphomycetes in Sclerococcaceae (Eurotiomycetes). Phytotaxa
372(1):079–087
Zoller S, Scheidegger C, Sperisen C (1999) PCR primers for the ampli-
fication of mitochondrial small subunit ribosomal DNA of lichen
forming ascomycetes. Lichenologist 31:511–516
Publisher’snoteSpringer Nature remains neutral with regard to
jurisdictional claims in published maps and institutional affiliations.
Mycol Progress (2019) 18:895–905 905