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ORIGINAL RESEARCH
Taxonomy and the evolutionary history of Micropeltidaceae
Xiang-Yu Zeng
1,2,3
•Hai-Xia Wu
1
•Sinang Hongsanan
3,4,5
•Rajesh Jeewon
6
•Ting-Chi Wen
2
•
Sajeewa S. N. Maharachchikumbura
7
•Putarak Chomnunti
8
•Kevin D. Hyde
3,8,9
Received: 25 December 2018 / Accepted: 31 May 2019
School of Science 2019
Abstract
Micropeltidaceae species are flyspeck fungi which have been subjected to few systematic studies. We re-examined 27 genera
which were accepted in the Micropeltidaceae and re-described them based on herbaria materials and protologues. Based on
morphologyand phylogenetic investigations, we transfer Micropeltidaceae to a new order, Micropeltidales (Lecanoromycetes).
Genera with bluish or greenish upper walls (Dictyopeltella,Dictyothyriella,Dictyothyrina,Dictyothyrium,Haplopeltheca,
Micropeltis,Scolecopeltidium and Stomiopeltopsis) are accepted in the new taxonomic concept for Micropeltidaceae. A
molecular clock approach estimated the divergence time of the Micropeltidaceae crown group at 130 (165–104) Mya,which also
supports its rank as an order (diverging from 220–100 Mya). The evolutionary histories between Micropeltidaceae species and
host plants are interpreted by cophylogenetic analyses calibrated by their divergence times. The result indicates that the
diversification of Angiospermae (130–80 Mya) fosters the formation of genera of Micropeltidaceae mainly via cospeciation
events, and this codivergent period would be an important reference when establishing generic boundaries of epifoliar fungi.
Keywords Codivergence Cophylogeny Dating Lecanoromycetes Micropeltidales
Introduction
The family Micropeltidaceae comprises ‘flyspeck’ fungi typi-
fied by shiny, black, round to ovoid, sclerotium-like bodies
lacking a visible mycelial mat (Yang et al. 2010;Lietal.2011).
This family was accomodated in the Microthyriales, Doth-
ideomycetes based on its morphological similarities to Asteri-
naceae, Microthyriaceae and Phaeothecoidiellaceae. Species of
Micropeltidaceae are, however, not clearly distinct from related
families based on morphology. The phylogenetic placement of
Micropeltidaceae within the Lecanoromycetes has been sup-
ported based on DNA sequence data available from only two
species in GenBank (Hongsanan and Hyde 2017). The taxo-
nomic boundaries of species within the Micropeltidaceae have
remained unclear for more than half a century and there has not
Electronic supplementary material The online version of this
article (https://doi.org/10.1007/s13225-019-00431-8) con-
tains supplementary material, which is available to autho-
rized users.
&Hai-Xia Wu
aileen2008haixia@gmail.com
1
International Fungal Research and Development Centre, Key
Laboratory of Resource Insect Cultivation & Utilization State
Forestry Administration, The Research Institute of Resource
Insects, Chinese Academy of Forestry, Kunming 650224,
China
2
The Engineering Research Center of the Utilization for
Characteristic Bio-Pharmaceutical Resources in Southwest,
Guizhou University, Guiyang 550025, China
3
Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
4
Shenzhen Key Laboratory of Microbial Genetic Engineering,
College of Life Sciences and Oceanography, Shenzhen
University, Shenzhen 518000, China
5
Shenzhen Key Laboratory of Laser Engineering, College of
Optoelectronic Engineering, Shenzhen University,
Shenzhen 518000, China
6
Department of Health Sciences, Faculty of Science,
University of Mauritius, Reduit 80837, Mauritius
7
Department of Crop Sciences, College of Agricultural and
Marine Sciences, Sultan Qaboos Universty, PO Box 8,
123 Al Khoud, Oman
8
School of Science, Mae Fah Luang University,
Chiang Rai 57100, Thailand
9
Key Laboratory for Plant Diversity and Biogeography of East
Asia, Kunming Institute of Botany, Chinese Academy of
Sciences, 132 Lanhei Road, Kunming 650201, China
123
Fungal Diversity
https://doi.org/10.1007/s13225-019-00431-8(0123456789().,-volV)(0123456789().,-volV)
been any major taxonomic revision since Batista (1959). It is
therefore important to resolve the taxonomy of Micropelti-
daceae, as well as other related ‘grey’ families for a better
understanding of their evolution and taxonomic circumscription.
Historical overview
Micropeltidaceae can be traced back to Pyrenomycetes Sch-
wein., when Montagne (1842) introduced the genus Micro-
peltis. Saccardo (1883) included taxa that are characterised by
superficial, black, dimidiate, flattened, membranous or car-
bonaceous perithecia in the new family Microthyriaceae. von
Ho
¨hnel (1910) suggested that those fungi with non-radiating
upper walls should be excluded from Microthyriaceae. Theis-
sen (1913) agreed with von Ho
¨hnel (1910) and introduced a
new order Hemisphaeriales Theiss., along with a new family
Hemisphaeriaceae Theiss., to accommodate taxa that are
characterised by meandering upper walls. Arnaud (1918)sug-
gested to group Microthyriaceae and Hemisphaeriaceae under
the new order Microthyriales G. Arnaud. Clements and Shear
(1931) synonymised Hemisphaeriaceae under the new family
Micropeltidaceae, with 29 genera and 186 species. However,
the family name Micropeltidaceae was not formally accepted
until the publication of the monograph by Batista (1959), which
included 45 genera in the family. Kirk et al. (2008)included27
genera and Lumbsch and Huhndorf (2010) included 24 genera
in Micropeltidaceae. Wu et al. (2011) transferred Byssopeltis,
Caudella and Scolecopeltidium from Microthyriaceae to
Micropeltidaceae based on the structure of the upper wall. Hyde
et al. (2013) revised the taxonomy of Micropeltidaceae and
accepted 12 genera. Hongsanan et al. (2014) transferred
Anariste to Micropeltidaceae from Asterinaceae. Recently, the
outlines provided by Wijayawardene et al. (2014,2018)
included 13 and 14 genera in Micropeltidaceae, respectively.
The phylogenetic study of Micropeltidaceae can be traced
back to Wu et al. (2011), who provided the first sequence data
(LSU and SSU sequences of Micropeltis zingiberaceicola H.X.
Wu & K.D. Hyde) of the modern Micropeltidaceae. Abarca
et al. (2011) investigated the phylogenetic relationships of the
asexual genera, Heliocephala and Holubovaniella,andcon-
cluded that Stomiopeltis betulae J.P. Ellis is the closest relative
sexual genus. Hongsanan et al. (2017a)excludedChaetothyr-
ina from Micropeltidaceae, and placed it into Phaeothecoi-
diellaceae based on molecular data and disease symptoms.
However, all previous phylogenetic studies were based on a
putative fact that Micropeltidaceae is a member of Doth-
ideomycetes. Hongsanan and Hyde (2017) concluded that
Micropeltidaceae should be transferred from Dothideomycetes
to Lecanoromycetes based on the phylogeny, but its placement
within Lecanoromycetes still remains unclear.
Taxonomic review
Members of Micropeltidaceae are characterised by flat-
tened, blue-green upper walls with pseudoparenchymatous
hyphae, and a central ostiole, clavate to cylindrical or
elongate asci that develop inclined towards the center of
ascomata and hyaline ascospores with uni- or multi-
transversal septa (Fig. 1). Clements and Shear (1931)
divided the family into three subfamilies viz.
Fig. 1 Characters of
Micropeltidaceae.aColonies
of Micropeltis asiatica (MFLU
16-0082) on leaf surface. bThe
upper wall of Dictyothyriella
garciniae (MFLU 17-1056). c, d
Ascomata of Micropeltis
asiatica (MFLU 16-0082) and
Dictyothyriella garciniae
(MFLU 17-1056) on leaf
surface. e, f The ascus and
ascospore of Micropeltis
asiatica (MFLU 16-0083)
Fungal Diversity
123
Dictyopeltineae Theissen, Haplopeltineae Theissen and
Plochmopeltineae Theissen based on colour and structure
of upper walls. Batista (1959) divided Micropeltidaceae
into four subfamilies based on colour of upper walls:
Dictyopeltoideae Batista (species with dark green or dark
blue upper walls with meandering reticulate hyphae);
Haplopeltoideae Batista (species with brown or black
upper walls with pseudo-parenchyma); Gymnopeltoideae
Batista (species with red upper walls with pseudo-par-
enchyma); and Stomiopeltoideae Batista (species with
brown upper walls with free hyphae). Batista (1959) also
provided keys to genera based on the size of ascospores,
presence of ostiole, setae or paraphyses, as well as keys to
species for each genus in Micropeltidaceae based on the
size of ascospores.
Internal hyphae are completely absent in Micropelti-
daceae. The hyphal filaments are delicate, forming a weft
or film, usually hyaline, and are interpreted as elements of
the ascending stroma. Ascomata are flattened, circular,
with a papillate ostiole, surrounded by a hyaline film or not,
with whole or crenellate margins. The upper walls consist
of closely jointed, meandrous cells, and its blue-green
coloration might be caused by some co-inhabiting algae or
bacteria. Asci are 8-spored, clavate, ellipsoid, globose or
cylindrical, and radially arranged from the periphery
towards the center. Paraphyses are presented as sterile fil-
aments between asci. These filaments are simple or bran-
ched, tubular or not, with or without septa, shorter or longer
than the asci, sometimes tapering at the apex, and hyaline
and rarely coloured (Batista 1959). The presence or
absence of paraphyses is a main taxonomic criterion for
generic differentiation (Wu et al. 2011; Hyde et al. 2013).
Ascospores are uni- or multi-cellular, constricted or not at
septa, hyaline, rarely chlorinate or brown. Filiform spores
always have a length–width ratio of 10:1 (Batista 1959).
Species of Micropeltidaceae differs from Microthyri-
aceae in having bluish or greenish upper walls with
meandrous cells and mostly multi-septate ascospores,
while Microthyriaceae has brownish upper walls with
radially arranged cells and 1-septate ascospores. Although
some genera have uniseptate ascospores (e.g. Dictyopel-
tella), they were identified as members of Micropeltidaceae
based on the arrangement of upper wall cells. The taxo-
nomic concept of genera in Micropeltidaceae is still
unclear due to lack of studies and insufficient phylogenetic
evidence to support classification.
Ecology
Species of Micropeltidaceae are highly diverse in tropical
regions and can be found during cold seasons when asci
and ascospores are mature (Batista 1959). They are mostly
epiphyllous and distributed on a wide range of plant fam-
ilies and records indicate that they are not host-specific
(Batista 1959; Acosta 1995; Reynolds and Gilbert
2005,2006). Similar to other sooty blotch and flyspeck
fungi, species of Micropeltidaceae cause no damage to the
underlying hosts. They neither produce haustoria to obtain
nutrients from the living organisms, nor utilize the con-
stituent elements from the dead bodies (Batista 1959).
Species of Micropeltidaceae are entirely superficial and
commensal with their host plants. They use mineral and
organic substances resulting from transpiration residues
and epidermal secretions (e.g. potassium, calcium salts and
phosphatides) to satisfy their growth, even in very small
quantities, and the exchange of gases between host plants
and the environment also benefits these fungi (Batista
1959).
The sooty blotch and flyspeck fungal group are gener-
ally difficult to culture, as they either rarely sporulate, or
grow extraordinarily slow and thus can be easily contam-
inated by saprobes, and this has resulted in a poor under-
standing of the group (Batzer et al. 2005; Yang et al. 2010;
Gleason et al. 2011; Zeng et al. 2018a). Numerous attempts
have been made to explore ways to cultivate Micropelti-
daceae species under artificial laboratory environments, but
without success (Batista 1959).
Life cycle
The asexual state of Micropeltidaceae has not been repor-
ted so far. For the sexual state, Batista (1959) concluded
that species of Micropeltidaceae are entirely free of fila-
mentous metagenic forms. After ascospores germinate, the
germinating tube will be fixed and dilated as a ‘appresso-
ria-like’ structure, from which the ascomata will develop.
The development of ascomata begins with process of
pseudo-parenchyma formation, giving rise to the develop-
ment of the upper wall to protect asci. The basal wall,
where the hyphae contact the leaf epidermis, absorbs
metabolic products until the marginal film is formed. The
thickness of the basal ascomata is considerably reduced
when mature.
Divergence time estimations
The key to reconciling different species concepts is to
identify a common element that implies a single, more
general, concept of species (de Queiroz 2007), and recently
Hyde et al. (2017) provided critical insights into how
divergence time estimates could be used in the taxonomic
ranking of fungi. Time is a common currency that allows
biologists to compare historical events among different
Fungal Diversity
123
groups, while molecular clocks provide the means to
convert molecular change into geological time that can
integrate fungal evolution with the dates of plate tectonic
movements, paleoecology, and the evolution of plants and
animals (Zuckerkandl and Pauling 1965; Sarich and Wil-
son 1967; Taylor and Berbee 2006; Berbee and Taylor
2010). Fossils with geological information provides abso-
lute time estimations for evolutionary events and can be
used to calibrate molecular clocks (Hyde et al. 2017;
Berbee and Taylor 2010). The ages of different lineages
can be then estimated by applying molecular clocks to
phylogenies, to better understand fungal evolution.
The first study that applied a molecular clock to fungal
phylogeny was carried out by Simon et al. (1993), to
investigtate if the radiation of arbuscular mycorrhizal fungi
coincided with the appearance of green plants on land.
Vijaykrishna et al. (2006) studied the direction of the
evolution of aquatic ascomycetes and showed that they had
evolved from land saprobes, while Li et al. (2005) used the
molecular clock to postulate how nematode trapping
methods had evolved in fungi. As suggested by Hennig
(1966) and implemented by Avise and Johns (1999),
divergence time estimated by using molecular clock
approach has now become a criterion to delimitate higher
ranks of fungal taxa in systematic studies (Hongsanan et al.
2017b; Hyde et al. 2017; Liu et al. 2017; Zhao et al. 2017).
It provides more objective evidence to justify the bound-
aries of higher fungal rank, which compensates the draw-
backs in morphology and phylogeny.
Hongsanan et al. (2016) estimated the divergence times
for some epifoliar fungal groups, but not Micropeltidaceae.
This study investigates the evolution of the Micropelti-
daceae based on divergence time estimates and provide
new insights into their taxonomy.
Cophylogenetic analyses
The long-term evolutionary dynamics encompass several
speciation events. The joint speciation of two or more
lineages that are ecologically associated is referred to as
‘cospeciation’, and the association can be parasitic, mutu-
alistic, symbiotic and other relationships (Page 2003). The
idea of cospeciation is derived from the pioneering studies
on avian parasites (Fahrenholz 1913), in which the
hypothesis that they proposed is known as the Fahrenholz
rule: ‘parasite phylogeny mirrors that of its host’. Host taxa
have since been used as a taxonomic criterion for the
classification of parasites due to lack of DNA sequences in
the past (Savile 1979; Hoberg et al. 1997; de Vienne et al.
2013). In Hafner and Nadler (1988), the first cophyloge-
netic analysis that compared independent phylogenies for
rodents and their ectoparasites was published.
Cophylogeny mostly tries to determine the significant
match between host and symbiont trees, as well as the best
explanation for the differences between the two. To better
interpret the evolutionary history, it is also important to
estimate the age of nodes in host and symbiont phylogenies
(de Vienne et al. 2007). A study by Zhang et al. (2015)
demonstrated that both historical and contemporary events
have played important roles in the phylogeography and
evolution of the Ophiocordyceps sinensis–ghost moth
parasitism on the Tibetan Plateau and results of cophylo-
genetic analyses accompany with estimated divergence
time raised the possibility of the occurrence of cospeciation
events. Such an approach has also been applied to study
parasitic fungal-plant associations (McTaggart et al. 2016),
but not for epiphytes.
Generally, cophylogenetic studies concentrate on host-
specific parasites, while multi-host parasitic species may
result from situations such as cryptic parasitic species,
recent host switching or failure to speciate (Banks and
Paterson 2005). Lice are believed to be host-specific and
show a high degree of codivergence with their hosts, but it
is relatively common that they infest multiple host species
(Banks and Paterson 2005). Therefore, it is with caution
that a conclusion on host-specificity should be made based
only on the statistical data, whereas identifying evolu-
tionary patterns may explain the true ecological traits of
parasites.
Aim of the study
The aim of this study was to clarify the taxonomic
boundaries of Micropeltidaceae, solve the phylogenetic
placement of Micropeltidaceae, estimate the divergence
time of Micropeltidaceae and investigate the evolutionary
history of Micropeltidaceae. We also crushed the ascomata
and established the microbial community on the surface of
leaves to verify if the blue-green coloration is caused by
some algae or bacteria that co-inhabiting with Micropelti-
daceae species.
Materials and methods
Morphological studies
Type specimens of Dictyopeltella,Dictyothyriella,Dic-
tyothyrina,Dictyothyrium,Haplopeltheca,Stomiopeltop-
sis,Stigmatodothis,Stomiopeltis,Dictyopeltis,Mitopeltis,
Stomiotheca,Thyriodictyella,Hansfordiopsis,Stig-
matophragmia and Polypedia were obtained from GZU, K,
LPS, NY, PDD, S and URM. Fresh collections with black
spots were collected in Thailand and China, and returned to
Fungal Diversity
123
the laboratory in paper envelopes. The samples were pro-
cessed and examined following the methods described by
Chomnunti et al. (2014). Photographs of ascomata were
taken using a compound stereomicroscope (Zeiss Discov-
ery.V8 with camera AxioCam ERc 5S). Sections were
made using a stereomicroscope (Motic) and mounted in
water or rehydrated 5% KOH solution. Photomicrographs
of fungal structures were taken with a light microscope
(Nikon Eclipse Ni–U) fitted with a digital camera (Canon
DS126311 EOS 600D) and a scanning electron microscope
(SEM). Small pieces of leaves containing several clean
ascomata were cut and kept in 1.5 ml microcentrifuge
tubes with silica gel inside for later DNA extraction (Chase
and Hills 1991). Holotypes are deposited at Mae Fah Luang
University (MFLU) Herbarium and The Research Institute
of Resource Insects, Chinese Academy of Forestry (IFRD),
isotypes are deposited at Herbarium of Cryptogams Kun-
ming Institute of Botany, Academia Sinica (HKAS). Index
Fungorum and Facesoffungi numbers are registered
(Jayasiri et al. 2015; Index Fungorum 2018). New taxa are
established based on recommendations outlined by Jeewon
and Hyde (2016).
DNA extraction, amplification and sequencing
Total genomic fungal DNAs were extracted following the
protocol described by Zeng et al. (2018b). The partial large
subunit (LSU) rDNA was amplified with the primer pair
LROR and LR5 (Vilgalys and Hester 1990). The 5.8S
rDNA along with the internal transcribed spacer (ITS) was
amplified with the primer pair ITS1 and ITS4 (White et al.
1990). The partial small subunit (SSU) rDNA was ampli-
fied with the primer pair NS1 and NS4 (White et al. 1990).
The partial translation elongation factor 1-alpha (TEF)
pcDNA was amplified with the primer pair EF1-983F and
EF1-2218R (Rehner and Buckley 2005). The partial second
largest subunit of RNA polymerase II (RPB2) pcDNA was
amplified with two primer pairs fRPB2-5F and fRPB2-7cR,
and fRPB2-7cF and fRPB2-11aR (Liu et al. 1999).
Total genomic plant DNAs were extracted using Plant
gDNA maxi kit (BioMIGA, USA) according to the man-
ufacturer’s instructions. The partial ribulose-1,5-bisphos-
phate carboxylase/oxygenase (rbcL) pcDNA was amplified
with primer pair SI_For and SI_Rev developed by Kress
et al. (2009). The trnH-psbA intergenic spacer was ampli-
fied with primer pair psbAF (Sang et al. 1997) and trnH2
(Tate and Simpson 2003). The 5.8S rDNA along with the
internal transcribed spacer (ITS) was amplified with the
primer pair ITS1 and ITS4 (White et al. 1990).
PCR reactions were employed in 20 lL reaction mixture
containing 10 lL29Bench Top Taq Master Mix (Bio-
miga, AT1201, China), 7 lL ddH
2
O, 1 lL forward and
reward primers (10 lM/lL) and 1 lL DNA template.
Amplifications were performed in a T100
TM
Thermal
Cycler (BIO-RAD), which was programmed for an initial
denaturation at 95 C for 3 min followed by 34 cycles of
1 min at 95 C, 50 s at 49 C(rbcL, fRPB2-7cF and
fRPB2-11aR for RPB2), 50 C(trnH-psbA), 51 C (LSU
and SSU), 53 C (5.8S, fRPB2-5F and fRPB2-7cR for
RPB2), 55 C (TEF) and extension at 72 C for 1 min, and
a final extension at 72 C for 5 min. PCR products were
sequenced by using appropriate PCR primers used in
amplification reactions by Sangon Biotech (Shanghai) Co.,
Ltd.
Phylogenetic analyses
Sequences of each gene generated from forward and
reverse primers were assembled with BioEdit v.7.2.5 (Hall
2013) to obtain consensus sequences. Representative
sequences of each class in Pezizomycotina and each order
in Lecanoromycetes were downloaded from GenBank
(Supplementary Table 1). Each gene dataset was aligned
separately by Mafft v7.187 (Katoh and Standley 2013) and
then manually aligned where necessary. Phylogenetic trees
based on LSU, SSU, 5.8S, TEF, RPB2 and combined LSU,
SSU, 5.8S, TEF and RPB2 datasets were performed using
maximum likelihood (ML) and Bayesian inference (BI) at
the CIPRES webportal (Miller et al. 2010).
Maximum likelihood analysis was performed using
‘RAxML-HPC BlackBox’ tool (Stamatakis 2014). One
thousand non-parametric bootstrap iterations were
employed with a general time reversible (GTR) model and
a discrete gamma distribution, plus estimating the propor-
tion of invariable sites (Stamatakis et al. 2008). Bayesian
inference (Larget and Simon 1999) was performed using
the ‘MrBayes on XSEDE’ tool (Ronquist et al. 2012) based
on the nucleotide substitution models which were deter-
mined by the Akaike Information Criteria (AIC) in
MrModeltest 2.3 (Nylander 2008). The Markov Chain
Monte Carlo (MCMC) algorithm of four chains started
from a random tree topology with two parallel runs. Trees
were sampled every 1000 generations, and the run was
stopped automatically when the average standard deviation
of split frequencies fell below 0.01. A 50% majority rule
consensus tree was summarised after discarding the first
25% samples.
The resulting trees were visualised in FigTree v1.4.3
(Rambaut 2012).
Fossil calibrations and divergence time
estimations
Four fossils were selected to constrain the node age with an
additional secondary calibration (Samarakoon et al. 2019):
the fossil Anzia electra (Rikkinen and Poinar 2002) was
Fungal Diversity
123
Table 1 Genera accepted in Micropeltidaceae
Clements and Shear (1931) Batista (1959) Lumbsch and Huhndorf (2010) Wijayawardene et al. (2014) This study
Aphysa Amazonotheca ?Armata Anariste Dictyopeltella
Chaetopeltopsis Bonaria Bonaria Caudella Dictyothyriella
Chaetoplaca Chaetopeltopsis Chaetothyrina Chaetothyrina Dictyothyrina
Clypeolum Chaetoplaca Clypeolina Dictyopeltella Dictyothyrium
Dictyopeltis Chaetothyrina Cyclopeltis Haplopeltheca Haplopeltheca
Dictyothyrina Ciferriotheca Dictyopeltella Heliocephala Micropeltis
Dictyothyrium Clypeolinopsis Dictyopeltis Holubovaniella Scolecopeltidium
Eremotheca Clypeolum Dictyostomiopelta Micropeltis Stomiopeltopsis
Eremothecella Dictyopeltella Dictyothyriella Scolecopeltidium
Griggsia Dictyopeltis Dictyothyrina Sirothyriella
Haplopeltis Dictyostomiopelta Dictyothyrium Stomiopeltis
Metathyriella Dictyothyrina Hansfordiopsis Stomiopeltopsis
Micropeltella Dictyothyrium Haplopeltheca Stomiotheca
Micropeltis Didymopeltis Mendoziopeltis
Microthyriella Didymothyriella Micropeltis
Mitopeltis Eremotheca ?Mitopeltis
Moesziella Gymnopeltis Muricopeltis
Phaeaspis Hansfordiopsis Polypedia
Phragmothyriella Haplopeltis Stigmatodothis
Plochmopeltis Kerniomyces Stigmatophragmia
Polyclypeolum Mendoziopeltis Stomiopeltis
Protopeltis Metathyiella Stomiopeltopsis
Saccardinula Micropeltella Stomiotheca
Schizothyrium Micropeltidium Thyriodictyella
Scolecopeltis Micropeltis
Scolecopeltium Microthyriella
Stigmatophragmia Muricopeltis
Stomiopeltella Neopeltella
Stomiopeltis Oswaldoa
Parapeltella
Plochmopeltidella
Polyclypeolina
Polyclypeolum
Protopeltis
Schizopeltis
Schizothyrina
Scolecopeltella
Scolecopeltidium
Scolecopeltis
Setopeltis
Spegazziniella
Stomiopeltella
Stomiopeltis
Stomiotheca
Sydowiellina
Fungal Diversity
123
used to calibrate the divergence of Anzia and Cano-
parmelia (normal distribution, mean = 35, SD = 50). The
fossil Aspergillus collembolorum (Do
¨rfelt and Schmidt
2005) was used to calibrate the crown group of Aspergillus
(normal distribution, mean = 35, SD = 50). The fossil
Calicium sp. (Rikkinen 2003) was used to calibrate the
genus Calicium (normal distribution, mean = 35, SD =
50). The fossil Chaenotheca sp. (Rikkinen 2003) was used
to calibrate the genus Chaenotheca crown group (normal
distribution, mean = 35, SD = 50). The tree ‘rootHeight’
was used to calibrate the Pezizomycotina by applying a
truncated normal distribution (mean = 400, SD = 150)
with an upper hard bound (truncation) set to 400 Mya
(Beimforde et al. 2014).
Subsequent divergence time analysis was carried out
using BEAST v1.8.4 (Drummond et al. 2012). Aligned
sequence data of each LSU, SSU, 5.8S, TEF and RPB2
dataset was partitioned and loaded to prepare an XML file
constructed with BEAUTI v1.8.4. Clock and substitution
models were set to be unlinked (independently estimated
for each gene partition), while the tree prior parameters
were set to be linked across partitions (concatenation). The
best fit substitution model for each gene was tested from
eleven substitution schemes by using ‘jModelTest2 on
XSEDE’ tool (Darriba et al. 2012) at the CIPRES web
portal (Miller et al. 2010), and determined by the Bayesian
information criterion (BIC). An uncorrelated relaxed clock
model (Drummond et al. 2006) with a lognormal distri-
bution of rates was applied for each partition. The Yule
process (Yule 1925; Gernhard 2008) was used to model the
speciation of nodes in the topology, with uniform priors on
probability of splits and extinctions.
After the analyses, Tracer v.1.7 (Rambaut et al. 2018)
was used to check if the effective sample sizes (ESS) were
higher than 200, after removing a proportion of states as
burn-in. The log files and tree files generated by BEAST
were combined in LogCombiner 1.8.4. The maximum
clade creditability (MCC) tree with median node heights
was summarised by using TreeAnnotator v1.8.4, and then
visualised and modified using the Interactive Tree Of Life
(iTOL) (https://itol.embl.de) online tool (Letunic and Bork
2019).
Cophylogenetic analyses
Independent phylogenetic analyses of targeted fungal
group and their host plants were carried out using BEAST
v1.8.4 with the same steps in the molecular clock analyses.
The node ages of the fungal dataset and plant dataset were
calibrated by the estimated divergence times resulting from
this study and previous studies (Clarke et al. 2011;
Magallo
´n et al. 2015) respectively.
Distance-based analysis of cophylogeny was performed
using ParaFit, implemented in Copycat 2.04 (Meierkolthoff
et al. 2007). The global fit between the fungal and host
trees was computed and statistical significance tested by
randomizing individual fungus–plant associations. ParaFit
was also used to assess the contribution of each individual
fungus–plant association (‘link’) to this global congruence.
The permutation tests of significance were calculated using
999 permutations.
Event-based analysis was performed using Jane 4
(Conow et al. 2010). Jane assigns costs to five events that
are used to describe host–parasite cophylogeny, namely
cospeciation, duplication, failure to diverge, host switch
and sorting (or loss). We used the default eventcost
scheme (cospeciation = 0, duplication = 1, duplication and
host switch = 2, sorting = 1, failure to diverge = 1). The
Vertex-based cost model method was implemented; the
genetic algorithm was set to 100 generations with a pop-
ulation size of 300, and the statistical significance of
reconstructions was evaluated with 1000 random tip map-
ping permutations.
Microbial diversity analyses
The specimen Micropeltis asiatica (MFLU 16-0083) was
selected for the surface diversity analyses of bacteria. The
leaf sample was processed as follows: A clean leaf that
contains the fungi was mounted in the sterilised phosphate
buffer saline (PBS) solution (replace by new PBS solution
before each step) and centrifuged for 20 min at 180 rpm
twice and ultrasonication for 10 min (160 W, 30 s/30 s).
The supernatants from the above three steps were collected
and filtered with a 0.2 lm filter membrane and preserved at
-80 C. The processed sample was sent to Shanghai
Majorbio Bio-pharm Technology Co., Ltd, and the data
was analysed on the free online platform of Majorbio
I-Sanger Cloud Platform (www.i-sanger.com).
Results
The combined LSU, SSU, 5.8S, TEF and RPB2 dataset
(Supplementary Table 1) consists of 204 taxa and 4289
unambiguously aligned sites (LSU: 942, SSU: 1007, 5.8S:
164, TEF: 892, RPB2: 1284). The best fit substitution
model of each gene is TrNef ?I?G (5.8S), TPM1uf ?
I?G (TEF), TVM ?I?G (RPB2), SYM ?I?G (SSU)
and GTR ?I?G (LSU). The family Micropeltidaceae is
represented by eight species comprising 12 strains. They
form four distinct clades at the generic level viz. Dic-
tyothyriella,Scolecopeltidium,Stomiopeltopsis and the
type Micropeltis, which are also distinguished by
morphology.
Fungal Diversity
123
Fig. 2 The best scoring
RAxML tree based on
combined LSU, SSU, 5.8S, TEF
and RPB2 sequence data of
Pezizomycotina. Maximum
likelihood bootstrap support
values (BS) greater than 50%
and Bayesian posterior
probabilities (PP) greater than
0.9 are shown above the
branches. Clades with strong
support (BS C70, PP C0.95)
are indicate in bold.
Micropeltidales species and
other similar taxa are
highlighted in green and brown
colour respectively. New
sequences obtained from this
study are in green bold
Fungal Diversity
123
Phylogenetic analyses of Pezizomycotina revealed that
Micropeltidaceae clusters with Cyanodermella (Sticti-
daceae, Ostropales,) as a sister clade of Graphidales,
Gyalectales, Ostropales and Thellenellales in class Leca-
noromycetes, forming a distinct lineage that segregates
from Microthyriaceae (Microthyriales) in Dothideomycetes
(Fig. 2). On the other hand, Micropeltidaceae species
phylogenetically separate from those similar taxa with
brown upper walls (Asterinaceae, Microthyriaceae and
Phaeothecoidiellaceae) (Fig. 2). Therefore, Micropelti-
daceae is excluded from Microthyriales and placed in the
newly introduced Micropeltidales ord. nov.
Divergence time estimations used the same dataset as
the phylogenetic analyses. The MCC tree summarised from
finalised 1731 trees demonstrated that the Micropeltidales
crown group diverged at 130 (165–104) Mya (Fig. 3),
which is also within the period that major fungal orders
diverged (220–100 Mya) (Liu et al. 2017).
The distance-based approach ParaFit illustrated that the
global congruence between Micropeltidaceae species and
host plant trees is not significant (P= 0.22). Scenarios
reconstructed by the event-based approach Jane 4 resulted
in a single solution, which consists three cospeciation
events, one duplication, two duplication and host switches,
five losses and two failures to diverge. However, the
Fig. 2 continued
Fungal Diversity
123
program has overestimated the probability of cospeciation
when verified by the divergence time of fungi and host
plants. Therefore, we interpret the co-evolutionary histories
based on the cophylogenetic relationships calibrated by
their divergence times (Fig. 4). The tanglegram demon-
strated that the most recent common ancestors of Micro-
peltis,Dictyothyriella,Stomiopeltopsis and their host
plants appeared in the same period when the Angiospermae
started to diversify, which indicates that cospeciations
mainly drove the formation of genera in Micropeltidaceae.
This codivergent period could be applied as a new refer-
ence for delimiting epifoliar fungal genera.
The crushed ascomata did not exhibit any structure of
algae or bacteria, but the bacterial diversity analyses gave a
positive result. Totally 105,168 raw reads were obtained
from the result of bacterial diversity analyses. After fil-
tering, 52,584 high-quality sequences were retained and
clustered into 33 Operational Taxonomic Units (OTUs)
(Supplementary Table 2). Of the 33 OTUs, 90.6% belon-
ged to Cyanobacteria, 2.67% belonged to Sphingomon-
adales, 2.33% belonged to Burkholderiales, 1.10%
belonged to Rickettsiales, and the remaining 3.27%
belonged to other groups (Fig. 5). The parameter of Chao 1
Index is 63, the Shannon Index is 0.56, and the Good’s
coverage is 1.
Taxonomy
Micropeltidales X.Y. Zeng, H.X. Wu & K.D. Hyde, ord.
nov.
Fig. 2 continued
Fungal Diversity
123
Index Fungorum number: IF 555493; Facesoffungi
number: FoF 05125
Epiphytic on living leaves, epiphyllous or hypophyllous,
visible as black, solitary dots on host surface. Hyphae
superficial, hyaline, difficult to detect. Sexual morph:
Ascomata superficial, solitary, mostly gregarious, black,
flattened, circular, dimidiate, with a central ostiole. Upper
wall greenish or bluish in water, reddish or brownish in
KOH solution, membranous, comprising overlapping, net-
like or puzzle-like, compact or sparse, pseudoparenchy-
matous hyphae, lacking a basal plate. Hamathecium com-
prising asci inclined from the base towards the ostiole.
Paraphyses present or absent. Asci 8-spored, bitunicate,
cylindrical to clavate, hyaline. Ascospores overlapping,
2-seriate, cylindrical to subclavate, aseptate or uni- to
multi-septate, constricted at septa when mature, hyaline,
head cell larger than others, middle cell globose, tail cell
longer than others. Asexual morph: Undetermined.
Notes: Micropeltidales is introduced to accommodate
the type family Micropeltidaceae. Its placement in Leca-
noromycetes is supported by both phylogeny and estimated
divergence time.
Type family:Micropeltidaceae Clem. & Shear [as
‘Micropeltaceae’], Gen. fung., Edn 2 (Minneapolis): 100
(1931)
Cenozoic Mesozoic Paleozoic
Mya 0
100
200
300
400
20
40
60
80
120
140
160
180
220
240
260
280
320
340
360
380
420
Tubeufia chiangmaiensis
Dothidea sambuci
Xylaria hypoxylon
Natipusilla decorospora
Heliocephala zimbabweensis
Cyphelium inquinans
Calicium viride
Pyronema domesticum
Dendrographa decolorans
Lecanora hybocarpa
Catolechia wahlenbergii
Houjia yanglingensis
Rhizina undulata
Letrouitia domingensis
Teloschistes flavicans
Zeloasperisporium hyphopodioides
Sclerophora sanguinea
Aspergillus nidulans
Acrospermum adeanum
Chlorociboria aeruginosa
Varicellaria rhodocarpa
Eleutherascus lectardii
Peltigera degenii
Cyanodermella asteris
Spirosphaera beverwijkiana
Stomiopeltis betulae
Chaenotheca phaeocephala
Brachiosphaera tropicalis
Aureobasidium pullulans
Usnea antarctica
Umbilicaria vellea
Coccocarpia erythroxyli
Thelenella antarctica
Lindra thalassiae
Roccellographa cretacea
Lepra amara
Melanomma pulvis-pyrius
Myriangium duriaei
Chaetomium globosum
Phaeocryptopus nudus
Roccella fuciformis
Sympoventuria capensis
Acarosporina microspora
Exophiala dermatitidis
Stomiopeltopsis antidesmatis
Gyalecta friesii
Staurothele frustulenta
Lecidea laboriosa
Condylospora vietnamensis
Lophiotrema nucula
Ionaspis odora
Chaenotheca brachypoda
Ochroconis dracaenae
Loxospora cismonica
Chaenotheca xyloxena
Sclerophora farinacea
Micropeltis amomi
Tubeufia cerea
Phaeothecoidiella missouriensis
Phyllobaeis imbricata
Orbilia vinosa
Botrytis cinerea
Cladosporium cladosporioides
Solenopsora candicans
Anisomeridium ubianum
Chaetothyriothecium elegans
Aliquandostipite khaoyaiensis
Lembosia xyliae
Cladonia caroliniana
Canoparmelia caroliniana
Leptosphaeria doliolum
Tumidispora shoreae
Dissoconium aciculare
Potebniamyces pyri
Megalospora tuberculosa
Caliciopsis pinea
Gregorella humida
Caliciopsis orientalis
Acrospermum gramineum
Gnomonia gnomon
Gyromitra californica
Dermatocarpon miniatum
Lichenoconium erodens
Microthyrium buxicola
Capnodium coffeae
Elsinoe veneta
Trichoglossum hirsutum
Trichocoma paradoxa
Myriangium hispanicum
Arthroderma redellii
Diploschistes cinereocaesius
Phaeotrichum benjaminii
Porpidia albocaerulescens
Aspicilia caesiocinerea
Placynthiella uliginosa
Microcalicium arenarium
Macrophomina phaseolina
Diatrype disciformis
Penicillium viridicatum
Rhizocarpon geographicum
Lichenoconium usneae
Sordaria fimicola
Coccomyces dentatus
Physcia aipolia
Orbilia auricolor
Trapelia placodioides
Lecidea fuscoatra
Disciotis venosa
Absconditella sphagnorum
Lasiosphaeria ovina
Dictyothyriella garciniae
Melaspilea costaricensis
Massaria inquinans
Schaereria fuscocinerea
Pyxine subcinerea
Venturia inaequalis
Chaetothyrina guttulata
Cordyceps militaris
Asterina cestricola
Dermea acerina
Mollisia cinerea
Apiospora bambusae
Sclerophora coniophaea
Chaetothyrina musarum
Anzia colpodes
Stictis populorum
Lecanactis abietina
Lachnum virgineum
Clonostachys rosea
Ascobolus crenulatus
Stemphylium vesicarium
Baeomyces placophyllus
Capronia pilosella
Lulworthia fucicola
Geoglossum nigritum
Speiropsis pedatospora
Caloscypha fulgens
Onygena corvina
Asterina melastomatis
Micropeltis sp.
Buellia frigida
Scolecopeltidium wangtianshuiense
Endocarpon pallidulum
Cladoriella rubrigena
Trichodelitschia bisporula
Microthyrium propagulensis
Lobothallia radiosa
Peltula auriculata
Lindra obtusa
Ramularia endophylla
Umbilicaria mammulata
Chaenotheca trichialis
Cyanodermella oleoligni
Dyfrolomyces rhizophorae
Micropeltis asiatica
Phlyctis argena
Melaspilea lentiginosa
Natipusilla naponensis
Bulgaria inquinans
Aleuria aurantia
Cladoriella eucalypti
Cudoniella clavus
Calicium salicinum
Simonyella variegata
Acanthostigma lignicola
Placopsis perrugosa
Arthrorhaphis citrinella
Phacidium lacerum
Stomiopeltis versicolor
Schaereria dolodes
Dendrographa minor
Brigantiaea fuscolutea
Aspergillus fumigatus
Sarcoscypha coccinea
Teratosphaeria fibrillosa
Megalotremis verrucosa
Spirosphaera minuta
Microthyrium microscopicum
Mazzantia napelli
Botryosphaeria dothidea
Loramyces macrosporus
Barriopsis fusca
Meria laricis
Parmelia saxatilis
Sagiolechia protuberans
Zeloasperisporium ficusicola
Jahnula aquatica
Melaspilea diplasiospora
Lecophagus sp.
Scolecopeltidium menglaense
Lasallia pustulata
Trichoderma viride
Fissurina insidiosa
Toensbergia leucococca
Lobaria scrobiculata
Peltula umbilicata
Sporastatia polyspora
Micropeltis dendrophthoes
Hamatispora phuquocensis
Loxospora ochrophaea
Classes
Lecanoromycetes
Eurotiomycetes
Coniocybomycetes
Lichinomycetes
Dothideomycetes
Arthoniomycetes
Sordariomycetes
Leotiomycetes
Geoglossomycetes
Pezizomycetes
Orbiliomycetes
Orders
Lecideales
Sporastatiales
Peltigerales
Lecanorales
Teloschistales
Caliciales
Leprocaulales
Baeomycetales
Sarrameanales
Rhizocarpales
Pertusariales
Schaereriales
Umbilicariales
Micropeltidales
Gyalectales
Graphidales
Ostropales
Thellenellales
Fig. 3 The MCC tree with divergence times estimates for main
groups of the Pezizomycotina obtained from a Bayesian approach
(BEAST) using four fossil constraints. Classes of Pezizomycotina and
orders of Lecanoromycetes are differentiate with different colours.
Nodes with PP lesser than 0.9 are marked with blue circles in
proportion to their PP values. Assignments of the fossil constraints are
marked with blue stars
Fungal Diversity
123
Generic type:Micropeltis Mont., Annls Sci. Nat., Bot.,
se
´r. 2 17: 122 (1842)
Micropeltidaceae Clem. & Shear [as ‘Micropeltaceae’],
Gen. fung., Edn 2 (Minneapolis): 100 (1931)
Index Fungorum number: IF 81006; Facesoffungi
number: FoF 05126
Epiphytic on living leaves, epiphyllous or hypophyllous,
visible as black, solitary dots on host surface. Hyphae
superficial, hyaline, difficult to detect. Sexual morph:
Ascomata superficial, solitary, mostly gregarious, black,
flattened, circular, dimidiate, with a central ostiole. Upper
wall greenish or bluish in water, reddish or brownish in
KOH solution, membranous, comprising overlapping, net-
like or puzzle-like, compact or sparse, pseudoparenchy-
matous hyphae, lacking a basal plate. Hamathecium com-
prising asci inclined from the base towards the ostiole.
Paraphyses present or absent. Asci 8-spored, bitunicate,
cylindrical to clavate, hyaline. Ascospores overlapping,
2-seriate, cylindrical to subclavate, tapering at the end,
aseptate or uni- to multi-septate, constricted at septa when
mature, hyaline, head cell larger than others, middle cell
globose, tail cell longer than others. Asexual morph:
Undetermined.
Notes: Micropeltidaceae was established by Clements
and Shear (1931) with 29 genera and 186 species that has
been accommodated in Microthyriales. A monograph was
provided by Batista (1959) with keys to genera and species.
Descriptions of Micropeltidaceae are generally very simple
or incomplete, which results in different interpretations by
different researchers (Batista 1959). Recently, Micropelti-
daceae was placed in Lecanoromycetes based on phy-
logeny (Hongsanan and Hyde 2017). However, the DNA
sequence data and phylogenetic findings for this family are
limited due to few sequences being available in GenBank.
In this study, we accept eight genera in Micropeltidaceae
(Table 1).
Type genus:Micropeltis Mont., Annls Sci. Nat., Bot.,
se
´r. 2 17: 122 (1842)
Key to genera of Micropeltidaceae
1. Paraphyses absent or inconspicuous……………..…2
1. Paraphyses present……………………………..……4
2. Ascospore 2-septate………………..…Dictyothyrium
2. Ascospore more than 2-septate…………………...…3
3. Ascospores 3–4-septate…………...…Stomiopeltopsis
3. Ascospores 5–6-septate…………….…Haplopeltheca
Early Angiospermae
150 140 130 120 110 100 90 80 70 60 50 40 30 20 10
Cenozoic
0 (Mya)
Mesozoic
Semecarpus sp. (Sapindales)
Glochidion sp. (Malpighiales)
Smilax sp. (Liliales)
Amomum villosum (Zingiberales)
136Mya
119Mya
117Mya
Syzygium jambos (Myrtales)
Garcinia sp. (Malpighiales)
Antidesma sp. (Malpighiales)
116Mya
103Mya
Micropeltis asiatica
Dictyothyriella garciniae
Dictyothyriella garciniae
Stomiopeltopsis antidesmatis
Micropeltis asiatica
Micropeltis sp.
Micropeltis amomi
39Mya
41Mya
81Mya
130Mya
114Mya
92Mya
Scolecopeltidium wangtianshuiense
Scolecopeltidium menglaense
Fig. 4 The tanglegram shows
the evolutionary histories
between fungi (black) and
plants (green). White circles,
black circles, arrows and broken
lines indicate cospeciation,
duplication, host shift and
failure to diverge events
respectively
Fungal Diversity
123
4. Ascospores aseptate………………...…Dictyothyrina
4. Ascospores septate………………………………..…5
5. Ascospores 1-septate………………..…Dictyopeltella
5. Ascospores more than 1 septum……………………6
6. Ascospores 2–4-septate………………Dictyothyriella
6. Ascospores more than 4 septa………………………7
7. Ascospores 4–6-septate………………..…Micropeltis
7. Ascospores more than 6 septa…….Scolecopeltidium
Dictyopeltella Bat. & I.H. Lima, Publicac¸o
˜es Inst. Micol.
Recife 56: 242 (1959)
Index Fungorum number: IF 1533; Facesoffungi num-
ber: FoF 05127
Epiphytic on living leaves, hypophyllous, visible as
black dots. Hyphae superficial, thin, hyaline, hard to detect.
Ascomata superficial, solitary, subdense, gregarious, black,
flattened, circular, dimidiate, with a central ostiole. Upper
wall blackish green, membranous, comprising overlapping,
compact, pseudoparenchymatous hyphae with textura epi-
dermoidea, lacking a basal plate. Hamathecium comprising
asci inclined from the base towards the ostiole. Paraphyses
filiform, thin, evanescent. Asci 8-spored, bitunicate,
cylindrical to ovoid. Ascospores overlapping, 2-seriate,
fusiform to subclavate, 1-septate, constricted at the septum,
hyaline.
Notes:Dictyopeltella was introduced by Batista (1959)
to distinguish species of Dictyopeltis that lack paraphyses.
The type species D. domingensis is similar to members of
Micropeltis in having blue-green upper walls, however, the
presence of paraphyses and the septation of ascospores
makes it different. Therefore, we maintain Dictyopeltella
as a member of Micropeltidaceae.
Type species:Dictyopeltella domingensis (Petr. & Cif.)
Bat. & I.H. Lima, Publicac¸o
˜es Inst. Micol. Recife 56: 243
(1959)
:Dictyopeltis domingensis Petr. & Cif., Annls mycol.
30(3–4): 179 (1932)
Index Fungorum number: IF 296628; Facesoffungi
number: FoF 05128, Fig. 6
Colonies hypophyllous. Sexual morph:Ascomata up to
270 lm in diam., subdense, with an irregular central osti-
ole. Upper wall blackish green, comprising net-like
hyphae. Asci 66–82 913–17 lm(
x=75915 lm,
n = 10), 8-spored, bitunicate, cylindrical to ovoid, with
Fig. 5 Percent of community
abundance that co-inhabiting
with Micropeltis asiatica
(MFLU 16-0083) on the leaf
surface. a, b Squash mount of
ascoma. cCommunity barplot
analysis at the order level
Fungal Diversity
123
short pedicel, ocular chamber not observed. Ascospores
18–23 94.5–6.5 lm(
x=2095.5 lm, n = 20), over-
lapping, 2-seriate, fusiform to subclavate, 1-septate, con-
stricted at the septum, upper cell oval to ovoid, lower cell
tapering at the end, hyaline. Asexual morph:
Undetermined.
Material examined: DOMINICAN REPUBLIC, Penin-
sula de Samana
´, Prov. de Samana
´,Sa
´nchez, old clearings,
300 m, on leaves of Renealmia aromatic (Aubl.) Griseb.,
30 April 1930, E. L. Ekman (GZU 000282267, holotype).
Dictyothyriella Speg., Boln Acad. nac. Cienc. Co
´rdoba
27(4): 384 (1924)
Index Fungorum number: IF 1538; Facesoffungi num-
ber: FoF 05129
Epiphytic on living leaves, epiphyllous, visible as black
dots. Hyphae superficial, hyaline, hard to detect. Ascomata
superficial, solitary, gregarious, subdense, bluish green,
flattened, circular, dimidiate, papillate, with a raised, cen-
tral ostiole. Upper wall bluish green, membranous, com-
prising overlapping, compact, pseudoparenchymatous
hyphae with textura epidermoidea, lacking a basal plate.
Hamathecium comprising asci inclined from the base
towards the ostiole. Paraphyses hyaline, embedded in a
gelatinous matrix. Asci 8-spored, bitunicate, clavate to
fusiform, short pedicel or absent, ocular chamber present.
Ascospores overlapping, 2-seriate, cylindrical to clavate,
2–3-septate, slightly constricted at septa, rounded at both
Fig. 6 Dictyopeltella
domingensis (GZU 000282267)
aThe material packet. bThe
specimen. cColonies on leaf
surface. dAscomata on host
surface. eCross section of
ascoma on leaf. fSquash mount
of ascoma. gThe upper wall. h–
jAsci. kPseudoparaphyses. l,
mAscospores. Scale bars
d= 100 lm, e, f =50lm,
g=20lm, h–m =10lm
Fungal Diversity
123
end cells, head cell bigger than others, smooth-walled,
hyaline.
Notes:Dictyothyriella was established by Spegazzini
(1924) with a single species D. bauhiniae (Rehm) Theiss.
& Syd. The genus was first placed in Microthyriaceae, but
Batista (1959) re-described the type species and transferred
it to Micropeltis. Species of Dictyothyriella and Micro-
peltis share the bluish green upper walls and filiform
paraphyses, but they can be distinguished by the septation
of ascospores. Therefore, we accept this genus as a member
of Micropeltidaceae.
Type species:Dictyothyriella bauhiniae (Rehm) Theiss.
& Syd., Annls mycol. 15(6): 430 (1918)
:Micropeltis bauhiniae Rehm, Leafl. of Philipp. Bot.
6: 1945 (1913)
Index Fungorum number: IF 215255; Facesoffungi
number: FoF 05130, Fig. 7
Colonies epiphyllous. Sexual morph:Ascomata up to
350 lm in diam., subdense, grouped, papillate, with a
raised, central ostiole. Upper wall bluish green, comprising
puzzle-like, lobed hyphae. Asci 40–60 97–14.5 lm
(
x=49912 lm, n = 15), 8-spored, bitunicate, clavate to
fusiform, short pedicel or absent, ocular chamber present.
Ascospores 14–20 93.5–7 lm(
x=1694.5 lm,
n = 20), overlapping, 2-seriate, cylindrical to clavate, 2–3-
septate, slightly constricted at septa, rounded at both ends,
head cell bigger than others, smooth-walled, hyaline.
Asexual morph: Undetermined.
Material examined: PHILIPPINES, P. I., Los Ban
˜os, on
leaves of Bauhinia sp., 20 May 1913, C. F. Baker, No.
1170 (S F8816, holotype).
Dictyothyriella garciniae X.Y. Zeng, H.X. Wu & K.D.
Hyde, sp. nov.
Index Fungorum number: IF 555492; Facesoffungi
number: FoF 05131, Fig. 8
Etymology: Refer to the host genus Garcinia.
Holotype: MFLU 17-1056
Fig. 7 Dictyothyriella
bauhiniae (S F8816) aThe
specimen. bColonies on leaf
surface. cSquash mount of
ascoma. dThe upper wall e, f
Cross section of ascoma. g–i
Asci mounted in Melzer
reagent. jPseudoparaphyses
mounted in Melzer reagent. k, l
Ascospores mounted in Melzer
reagent. Scale bars b= 500 lm,
c–e =20lm, f–j =10lm, k,
l=5lm
Fungal Diversity
123
Colonies epiphyllous, visible as black dots. Hyphae
superficial, hyaline, hard to detect. Sexual morph:Asco-
mata up to 680 lm in diam., superficial, solitary, gregari-
ous, subdense, black, flattened, circular, dimidiate, with a
raised, central ostiole. Upper wall blackish green, mem-
branous, comprising overlapping, net-like, branched,
compact, pseudoparenchymatous hyphae with textura epi-
dermoidea, lacking a basal plate. Hamathecium comprising
asci inclined from the base towards the ostiole. Paraphyses
filiform, thin, hyaline. Asci 41–53 912–15 lm
(
x=48914 lm, n = 5), 8-spored, bitunicate, ellipsoid to
subfusiform, ocular chamber not observed. Ascospores 23–
29 95lm(
x=2695lm, n = 3), overlapping,
2-seriate, cylindrical to subclavate, 2–4-septate, slightly
constricted at the septum, head cell a little larger, bottom
cell a little longer, hyaline. Asexual morph:
Undetermined.
Material examined: THAILAND, Trat, Amphoe Ko
Chang, Yuttha Navi Ko Chang Memorial, on living leaves
of Garcinia sp. (Clusiaceae), 27 April 2017, Xiang-Yu
Zeng (MFLU 17-1056, holotype; HKAS 101762, isotype);
THAILAND, Trat, Amphoe Ko Chang, Yuttha Navi Ko
Chang Memorial, on living leaves of Syzygium jambos
(Myrtaceae), 27 April 2017, Xiang-Yu Zeng (MFLU
17-1051).
Fig. 8 Dictyothyriella
garciniae (MFLU 17-1056)
aThe host leaf. bThe ascoma
on leaf surface. cSquash mount
of ascoma. dThe upper wall.
eParaphyses. f–g Asci.
hAscospores. Scale bars
c= 100 lm, d=20lm, e–
h=10lm
Fungal Diversity
123
Notes: This new species should be a member of Dic-
tyothyriella, due to its greenish upper walls, filiform para-
physes, and 2–4-septate ascospores. It is most similar to
Dictyothyriella harmsiana (Henn.) F. Stevens & Manter and
Dictyothyriella inaequiseptata Cif. in terms of ascospore size.
However, D. harmsiana has smaller ascomata (200–240 lm)
and lack paraphyses (Hennings 1897), while D. inaequisep-
tata has hypophyllous ascomata and larger asci (50–70 9
20–35 lm) (Ciferri 1956). Therefore, a new species, Dic-
tyothyriella garciniae sp. nov., is introduced. This species is
sister to Scolecopeltidium and Stomiopeltopsis, but phyloge-
netically distinct as a new species (Fig. 2).
Dictyothyrina Theiss., Annls mycol. 11(5): 469 (1913)
Index Fungorum number: IF 1539; Facesoffungi num-
ber: FoF 05132
Epiphytic on living leaves, hypophyllous, visible as
black dots. Hyphae superficial, hyaline, hard to detect.
Ascomata superficial, solitary, gregarious, subdense, black,
flattened, circular, dimidiate, with a circular central ostiole.
Upper wall blackish green, membranous, comprising
overlapping, compact, pseudoparenchymatous hyphae with
textura intricata, lacking a basal plate. Hamathecium
comprising asci inclined from the base towards the ostiole.
Paraphyses filiform, hyaline, embedded in a gelatinous
matrix. Asci 8-spored, bitunicate, cylindrical, short pedicel
or absent. Ascospores overlapping, 2-seriate, cylindrical to
ellipsoid, aseptate, hyaline.
Notes:Dictyothyrina was placed in Hemisphaeriaceae
(Hemisphaeriales) when introduced by Theissen (1913).
This is the only genus with aseptate ascospores in Micro-
peltidaceae, and the arrangement of its upper wall is
slightly different from Micropeltis. We accept this genus in
Micropeltidaceae due to its greenish, pseudoparenchyma-
tous upper wall. Sequence data is needed to confirm its
placement.
Type species:Dictyothyrina fecunda (Sacc.) Theiss.,
Annls mycol. 11(5): 469 (1914)
:Muyocopron fecundum Sacc., Bolm Soc. broteriana,
Coimbra, se
´r. 1 11: 62 (1893)
Fig. 9 Dictyothyrina
atrocyanea (S F9937) a, b The
specimen and descriptions.
cAscomata on leaf surface.
dSquash mount of ascoma.
eThe upper wall. fParaphyses.
gParaphyses mounted in
Melzer reagent. h, i Asci. j, k
Unicellular ascospores.
lAscospores in Melzer reagent.
Scale bars c= 500 lm,
d= 100 lm, e=50lm, h,
i=20lm, f, g, j–l =10lm
Fungal Diversity
123
Dictyothyrina atrocyanea (Starba
¨ck) Theiss., Annls mycol.
11(5): 469 (1914)
:Muyocopron fecundum var. atrocyaneum Starba
¨ck,
Bih. K. svenska VetenskAkad. Handl., Afd. 3 25(no. 1): 24
(1899)
Index Fungorum number: IF 215102; Facesoffungi
number: FoF 05133, Fig. 9
Colonies hypophyllous. Sexual morph:Ascomata up to
270 lm in diam., subdense, gregarious, with a circular
central ostiole. Upper wall blackish green, comprising
meandering, lobed hyphae. Asci 66–78 99–11(-14) lm
(
x=70911 lm, n = 5), 8-spored, bitunicate, cylindrical,
short pedicel or absent, ocular chamber not observed. As-
cospores10–14 93–5 lm(
x=1194lm, n = 15),
overlapping, 2-seriate, cylindrical to ellipsoid, aseptate,
verrucose, hyaline. Asexual morph: Undetermined.
Material examined: BRAZIL, Cuyaba, Matto Grosso, on
leaves of Hexalobi brasiliensis, 13 January 1894, G.
A. Malme (S F9937, holotype).
Notes: Information of the type species of Dictyothyrina
is scarce, and we were unable to locate the type specimen.
Therefore, instead another species in S herbarium was
examined. This specimen is characterised by pseudo-
parenchymatous upper walls and aseptate ascospores,
which is quite similar to Muyocopronales species. How-
ever, species of Muyocopronales have brown upper walls
rather than greenish, and they do not commonly occur on
leaves (Mapook et al. 2016). As upper walls of flyspeck
fungi in Dothideomycetes are all brown colour, we con-
sider the greenish, pseudoparenchymatous upper wall as
the typical features to accommodate the species in
Micropeltidaceae.
Dictyothyrium Theiss., O
¨st. bot. Z. 62: 277 (1912)
Index Fungorum number: IF 1540; Facesoffungi num-
ber: FoF 05134
Epiphytic on living leaves, epiphyllous, visible as black
dots. Hyphae superficial, hyaline, hard to detect. Ascomata
superficial, solitary, gregarious, subdense, black, flattened,
circular, dimidiate, papillate, with a raised, central ostiole.
Upper wall blackish green, membranous, comprising
overlapping, compact, pseudoparenchymatous hyphae with
textura epidermoidea, lacking a basal plate. Hamathecium
comprising asci inclined from the base towards the ostiole.
Paraphyses absent. Asci 8-spored, bitunicate, cylindrical to
fusoid, with short pedicel. Ascospores overlapping, 2-se-
riate, cylindrical to long clavate, 2-septate, slightly con-
stricted at septa, hyaline.
Notes:Dictyothyrium was placed in Hemisphaeriaceae
(Hemisphaeriales) when introduced by Theissen (1912).
The holotype specimen differs from protologue in position
of colony, septation of ascospores and present of guttulas in
ascospores. By considering the characters of the herbarium,
it is distinct to Micropeltis by the presence of paraphyses
and the number of septa in ascospores. We keep this genus
in Micropeltidaceae due to its greenish upper wall.
Type species:Dictyothyrium chalybeum (Rehm) Theiss.
[as ‘chalybaeum’], O
¨st. bot. Z. 62: 277 (1912)
:Clypeolum chalybeum Rehm, Hedwigia 37: 322
(1898)
Index Fungorum number: IF 203967; Facesoffungi
number: FoF 05135, Fig. 10
Colonies epiphyllous. Sexual morph:Ascomata up to
635 lm in diam., subdense, gregarious, papillate, with a
raised, central ostiole. Upper wall blackish green, com-
prising net-like hyphae. Asci 73–128 915–23 lm
(
x= 100 919.5 lm, n = 15), 8-spored, bitunicate, cylin-
drical to fusoid, with short pedicel, ocular chamber not
observed. Ascospores 35–53 98–11 lm
(
x= 40.8 99.5 lm, n = 20), overlapping, 2-seriate,
cylindrical to long clavate, 2-septate, constricted at septa,
hyaline. Asexual morph: Undetermined.
Material examined: BRAZIL, on leaves of Araceae, 3
May 1886, R. Laihallia (S F12647, holotype).
Haplopeltheca Bat., J.L. Bezerra & Cavalc., Publicac¸o
˜es
Inst. Micol. Recife 388: 5 (1963)
=Phaeostomiopeltis Bat. & Cavalc., in Batista, Bezerra,
Cavalcante, Maia & Silva, Publicac
¸o
˜es Inst. Micol. Recife
392: 9 (1963)
Index Fungorum number: IF 2233; Facesoffungi num-
ber: FoF 05136
Epiphytic on living leaves, epiphyllous, visible as black
dots. Hyphae superficial, hyaline, hard to detect. Ascomata
superficial, solitary, gregarious, scattered, black, flattened,
circular, dimidiate, papillate, with a central ostiole. Upper
wall blackish green, membranous, comprising overlapping,
compact, pseudoparenchymatous hyphae with textura epi-
dermoidea, lacking a basal plate. Hamathecium comprising
asci inclined from the base towards the ostiole. Paraphyses
absent. Asci 8-spored, bitunicate, clavate to ovoid, with
short pedicel. Ascospores overlapping, 2-seriate, cylindri-
cal, 5–6-septate, constricted at septa, head cell longer than
others, tapering at bottom, hyaline.
Notes:Haplopeltheca was introduced by Batista et al.
(1963b). Its blackish green upper wall and transverse septa
in ascospores is typical to Micropeltidaceae. Haplopeltheca
differs from other genera in Micropeltidaceae by lack of
paraphyses, 5–6-septate ascospores. We keep this genus as
a member of Micropeltidaceae due to its greenish, pseu-
doparenchymatous upper wall.
Type species:Haplopeltheca hyleensis Bat., J.L. Bez-
erra & Cavalc., Publicac¸o
˜es. Instituto de Micologia da
Universidade do Recife & Instituto Nacional de Pesquis
388: 7 (1963)
Fungal Diversity
123
Index Fungorum number: IF 331693; Facesoffungi
number: FoF 05137, Fig. 11
Colonies epiphyllous. Sexual morph:Ascomata up to
390 lm in diam., scattered, papillate, with a raised, ‘X’-
shaped, central ostiole when mature. Upper wall blackish
green, comprising net-like hyphae. Asci 43–60 914–
23 lm(
x=54919 lm, n = 15), 8-spored, bitunicate,
clavate to ovoid, with short pedicel, ocular chamber not
observed. Ascospores 24–40 95–8 lm
(
x= 29.5 96.6 lm, n = 20), overlapping, 2-seriate,
cylindrical, 5–6-septate, constricted at septa, head cell
longer than others, tapering at bottom, hyaline. Asexual
morph: Undetermined.
Material examined: BRAZIL, on leaves of an uniden-
tified plant, collection date unknown, A. C. Batista (URM
15859, holotype).
Micropeltis Mont., in Sagra, Annls Sci. Nat., Bot., se
´r. 2
17: 122 (1842)
=Dictyothyriella Rehm, in Theissen, Brote
´ria, se
´r. bot.
12: 92 (1914)
=Hormopeltis Speg., Anal. Mus. nac. Hist. nat. B. Aires
23: 84 (1912)
=Micropeltella Syd. & P. Syd., Annls mycol. 11(5): 404
(1913)
=Micropeltidium Speg., Boln Acad. nac. Cienc. Co
´r-
doba 23(3-4): 574 (1919) [1918]
=Ophiopeltis J.V. Almeida & Sousa da Ca
ˆmara, Revta
agron., Lisb. 1: 175 (1903)
=Parapeltella Speg., Boln Acad. nac. Cienc. Co
´rdoba
23(3-4): 506 (1919) [1918]
=Scolecopeltella Speg., Boln Acad. nac. Cienc. Co
´r-
doba 26(2-4): 354 (1921)
Fig. 10 Dictyothyrium
chalybeum (S F12647) aThe
specimen. bAscoma on leaf
surface. dThe upper wall
mounted in 3% KOH. eCross
section of ascoma. f–h Asci.
iAsci mounted in blue cotton
reagent. jAscospores. Scale
bars:b= 200 lm, c,
e=50lm, i, j =20lm, d, f–
h=10lm
Fungal Diversity
123
=Scolecopeltis Speg., Boln Acad. nac. Cienc. Co
´rdoba
11(4): 574 (1889)
=Scolecopeltopsis Ho
¨hn., Sber. Akad. Wiss. Wien,
Math.-naturw. Kl., Abt. 1 118: 316 [42 repr.] (1909)
=Theciopeltis F. Stevens & Manter, Bot. Gaz. 79(3):
285 (1925)
Index Fungorum number: IF 3180; Facesoffungi num-
ber: FoF 05138
Epiphytic on living leaves, hypophyllous, visible as
black dots. Hyphae superficial, hyaline, hard to detect.
Ascomata superficial, solitary, gregarious, subdense, black,
flattened, circular, dimidiate, with a central ostiole. Upper
wall bluish green or blackish green, membranous, com-
prising overlapping, compact, pseudoparenchymatous
hyphae with textura epidermoidea, lacking a basal plate.
Hamathecium comprising asci inclined from the base
towards the ostiole. Paraphyses filiform, thin, hyaline,
evanescent. Asci 8-spored, bitunicate, ovoid to widely
fusiform. Ascospores overlapping, 2-seriate, clavate to
fusiform, 4–6-septate, constricted at septa, hyaline, head
cell longer and larger, tapering at bottom.
Notes:Micropeltis was introduced by Montagne (1842)
as a member of Pyrenomycetes. The isotype from K is in
very poor condition. Nevertheless, Wu et al. (2011) has
illustrated the isotype from P and designated a portion of
the specimen as the lectotype. Therefore, we re-described
Micropeltis applanata based on the examined material and
descriptions from Wu et al. (2011).
Type species:Micropeltis applanata Mont., Annls Sci.
Nat., Bot., se
´r. 2 17: 122 (1842)
Index Fungorum number: IF 220119; Facesoffungi
number: FoF 05139
Colonies hypophyllous. Sexual morph:Ascomata up to
330 lm in diam., subdense, gregarious, with a circular
central ostiole. Upper wall bluish green or blackish green,
comprising puzzle-like hyphae. Asci 65–95 915–25 lm
(
x= 78.4 920.3 lm, n = 15), 6–8-spored, bitunicate,
obclavate to widely fusiform, with a short pedicel, ocular
Fig. 11 Haplopeltheca
hyleensis (URM 15859) aThe
specimen. b, c Colonies on leaf
surface. dSquash mount of
ascoma. eSection of ascoma. f–
hAsci mounted in Melzer
reagent. i–k Ascospores
mounted in Melzer reagent.
Scale bars:c= 200 lm,
d=50lm, e–h =10lm, i–
k=5lm
Fungal Diversity
123
chamber not obvious (Wu et al. 2011). Ascospores 30.5–
35 95–8 lm(
x= 32.8 95.7 lm, n = 20), overlapping,
2-seriate, (4-)5(-8) septate, constricted at septa, long cla-
vate, hyaline, smooth-walled (Wu et al. 2011). Asexual
morph: Undetermined.
Material examined: CUBA, on leaves of an unidentified
plant, collection date unknown, ex herb C. Montage (ex
herb. Berkeley) (K 164010 isotype).
Micropeltis asiatica X.Y. Zeng, H.X. Wu & K.D. Hyde,
nom. nov.
:Micropeltis zingiberaceicola H.X. Wu & K.D. Hyde
[as ‘zingiberacicola’], in Wu, Schoch, Boonmee, Bahkali,
Chomnunti & Hyde, Fungal Diversity 51(1): 211 (2011)
Index Fungorum number: IF 555494; Facesoffungi
number: FoF 05140, Fig. 12
Etymology:asiatica= Asian; in reference to its location,
China and Thailand.
Colonies mostly epiphyllous, sometimes hypophyllous,
visible as black dots. Hyphae superficial, hyaline, hard to
detect. Sexual morph:Ascomata up to 600 lm in diam.,
superficial, solitary, gregarious, subdense, black, flattened,
circular, dimidiate, with a central ostiole. Upper wall
blackish green, membranous, comprising overlapping,
puzzle-like, lobed, compact, pseudoparenchymatous
hyphae with textura epidermoidea, lacking a basal plate.
Hamathecium comprising asci inclined from the base
towards the ostiole. Paraphyses filiform, thin, hyaline. Asci
55–60 911–16 lm(
x=58914 lm, n = 5), 8-spored,
bitunicate, clavate to subfusiform, ocular chamber present.
Ascospores 30–42 94.5–6.5 lm(
x=3696lm,
n = 15), overlapping, 2-seriate, cylindrical, 4–6-septate,
constricted at the septum when mature, head cell larger,
middle cell globose when mature, bottom cell longer,
tapering at the ends, hyaline. Asexual morph:
Undetermined.
Material examined: THAILAND, Krabi, Khao Hang
Nak, on living leaves of Semecarpus sp. (Anacardiaceae),
17 December 2015, Xiang-Yu Zeng (MFLU 16-0083;
HKAS 101758); THAILAND, Krabi, Khao Hang Nak, on
living leaves of Glochidion sp. (Phyllanthaceae), 17
December 2015, Xiang-Yu Zeng (MFLU 16-0082);
THAILAND, Chiang Mai, Mae Taeng, Pa Pae, Bahn Pa
Deng, 128 Moo 3, Mushroom Research Centre, on living
Fig. 12 Micropeltis asiatica
(MFLU 16-0083) aHost leaves.
b, c Ascomata on leaf surface.
dSquash mount of ascoma.
eAn overview of ascoma
(SEM). fHyphae on leaf surface
(SEM). gParaphyses. h, i Asci.
j, k Ascospores. Scale bars
b= 200 lm, c, d = 100 lm,
e=20lm, f–k =10lm
Fungal Diversity
123
leaves of Xanthophyllum sp. (Polygalaceae), 18 July 2015,
Xiang-Yu Zeng (MFLU 16-0073).
Notes: The three new collections (MFLU 16-0073,
MFLU 16-0082 and MFLU 16-0083) are identical to Mi-
cropeltis zingiberaceicola H.X. Wu & K.D. Hyde in both
morphological features and DNA sequences. Sequence of
IFRDCC 2264 is 99% similar to the LSU and SSU
sequence of MFLU 16-0082 (820/821 nucleotides, no gaps;
1184/1199 nucleotides, 3/1199 gaps) and MFLU 16-0083
(817/818 nucleotides, no gaps; 1188/1198 nucleotides,
2/1198 gaps). Comparing MFLU 16-0082 and MFLU
16-0083, there are 99% similarity in ITS-LSU (1323/1328
nucleotides, no gaps), SSU (1501/1508 nucleotides, no
gaps), and TEF (749/754 nucleotides, no gaps) sequence,
while the major differences happens in RPB2 sequence
with 98% similarity (1899/1931 nucleotides, no gaps). The
difference among these sequences does not reach the
threshold for establishing new taxa (Jeewon and Hyde
2016). However, M. zingiberaceicola H.X. Wu & K.D.
Hyde should not be a valid name, as M. zingiberaceicola
Henn. had been published earlier in 1897. Therefore, Mi-
cropeltis asiatica nom. nov. is introduced here.
Micropeltis amomi X.Y. Zeng, H.X. Wu & K.D. Hyde, sp.
nov.
Index Fungorum number: IF 555495; Facesoffungi
number: FoF 05210, Fig. 13
Etymology: Refer to the host genus Amomum.
Holotype: MFLU 16-0081
Colonies epiphyllous, visible as black dots. Hyphae
superficial, hyaline, hard to detect. Sexual morph:Asco-
mata up to 400 lm in diam., superficial, solitary, gregari-
ous, subdense, black, flattened, circular, dimidiate, with a
central ostiole. Upper wall blackish green, membranous,
comprising overlapping, puzzle-like, lobed, compact,
pseudoparenchymatous hyphae with textura epidermoidea,
with hyaline, membranous margin, lacking a basal plate.
Hamathecium comprising asci inclined from the base
towards the ostiole. Paraphyses absent. Asci 42–56 918–
20 lm(
x=53919 lm, n = 5), 8-spored, bitunicate,
clavate, ocular chamber present. Ascospores 20–30 95–
6lm(
x=2495lm, n = 10), overlapping, 2-seriate,
cylindrical, 4–6-septate, constricted at the septum, head
cell larger, tapering at the ends. Asexual morph:
Undetermined.
Fig. 13 Micropeltis amomi
(MFLU 16-0081) aHost leaves.
bColonies on leaf surface.
cThe ascoma on leaf surface
(SEM). dHyphae (SEM).
eSquash mount of ascoma.
f. Hyphae under margin g, h
Asci. i, j Ascospores. Scale bars
b, e = 200 lm, f=50lm, c, d,
g, h =20lm, i, j =10lm
Fungal Diversity
123
Material examined: THAILAND, Krabi, Khao Hang
Nak, on living leaves of Amomum villosum (Zingiber-
aceae), 17 December 2015, Xiang-Yu Zeng (MFLU
16-0081, holotype; HKAS 101756, isotype).
Notes:Micropeltis amomi is most similar to Micropeltis
gravataensis Bat. & A.F. Vital in ascospore size. However,
M. gravataensis has paraphyses, larger asci, hypophyllous
colonies, and larger ascomata. Phylogenetically, the new
species is related with Micropeltis sp. MFLU 16-0090 and
M. dendrophthoes. There are 98% (792/810 nucleotides, no
gaps) similarity in LSU region, and 99% (429/433
nucleotides, no gaps) similarity in SSU region between the
new species and M. dendrophthoes. The new collection has
membranous margins, smaller ascospores, asci and asco-
mata, and the tail cell of ascospores in M. dendrophthoes is
much longer than other cells. Therefore, we introduce it as
a new species in Micropeltis.
Micropeltis sp. Figure 14
Colonies epiphyllous, visible as black dots. Hyphae
superficial, hyaline, hard to detect. Sexual morph:Asco-
mata superficial, solitary, gregarious, subdense, black,
flattened, circular, dimidiate, up to 450 lm in diam., with a
central ostiole. Upper wall blackish green, membranous,
comprising overlapping, puzzle-like, compact,
pseudoparenchymatous hyphae with textura epidermoidea,
with hyaline, membranous margin, lacking a basal plate.
Hamathecium comprising asci inclined from the base
towards the ostiole. Paraphyses absent. Asci not observed.
Ascospores not observed. Asexual morph: Undetermined.
Material examined: THAILAND, Chiang Mai, Mae
Taeng, Pa Pae, Bahn Pa Deng, 128 Moo 3, Mushroom
Research Centre, on living leaves of Smilax sp. (Smila-
caceae), 22 December 2015, Xiang-Yu Zeng (MFLU
16-0090, holotype; HKAS 101759, isotype).
Notes: Although there are a lot of ascomata available in
the new collection, we could not find structures of any asci
or ascospores. However, the sequence data shows it to be a
member of Micropeltis, which clusters with the newly
introduced species Micropeltis amomi within the clade that
represents Micropeltis. The comparison of the ITS regions
between the two related collections reveals a difference of
5% in the nucleotides that supports them to be different as
recommended by Jeewon and Hyde (2016). We therefore
could only identify it to the genus level.
Scolecopeltidium F. Stevens & Manter, Bot. Gaz. 79(3):
282 (1925)
Index Fungorum number: IF 4951; Facesoffungi num-
ber: FoF 05141
Fig. 14 Micropeltis sp. (MFLU
16-0090) aHost leaves. b, c
Colonies on leaf surface. dThe
ascoma (SEM). eHyphae
(SEM). fThe upper wall. Scale
bars b= 200 lm, c= 100 lm,
d–f =20lm
Fungal Diversity
123
Epiphytic on living leaves, hypophyllous, visible as
black dots. Hyphae superficial, hyaline, hard to detect.
Ascomata superficial, solitary, gregarious, subdense, black,
flattened, circular, dimidiate, with a circular central ostiole.
Upper wall blackish green, membranous, comprising
overlapping, compact, pseudoparenchymatous hyphae with
textura intricata, lacking a basal plate. Hamathecium
comprising asci inclined from the base towards the ostiole.
Paraphyses filiform, hyaline, branched. Asci 8-spored, long
ovate. Ascospores overlapping, 2-seriate, cylindrical, con-
tinuous, chain structure, more than 6 septa, constricted at
septa, hyaline, head cell longer and larger than others.
Notes:Scolecopeltidium was introduced in Microthyri-
aceae by Stevens and Manter (1925). The descriptions were
re-written based on the Wu and Hyde (2013), who reap-
praised the genus and concluded that this genus is typified
by ascospores with more than 5 septa. However, we found
that ascospores of the two specimens examined by Wu
et al. (2011) and Wu and Hyde (2013) both have more than
15 septa. Ascospores of Scolecopeltidium have the largest
size and more septa among other genera in
Micropeltidaceae. We conclude that ascospores in Scole-
copeltidium should have more than 6 septa and can reach
15 septa when mature. However, sequence data is needed
to confirm its placement.
Type species:Scolecopeltidium salacensis (Racib.) F.
Stevens & Manter, Bot. Gaz. 79(3): 282 (1925)
:Scolecopeltis salacensis Racib., Parasit. Alg. Pilze
Java’s (Jakarta) 3: 35 (1900)
Scolecopeltidium menglaense H.X. Wu, X.Y. Zeng &
K.D. Hyde, sp. nov.
Index Fungorum number: IF 555496; Facesoffungi
number: FoF 05142, Fig. 15
Etymology: in reference to its association (Latin -ensis)
with Meng La = menglaense.
Holotype: IFRD 9303
Colonies epiphyllous, visible as black dots. Hyphae
superficial, hyaline, hard to detect. Sexual morph:Asco-
mata up to 340 lm in diam., superficial, solitary, gregari-
ous, black, flattened, circular, with a papillate central
ostiole. Upper wall greenish, comprising overlapping, net-
like, compact, pseudoparenchymatous hyphae with textura
Fig. 15 Scolecopeltidium
menglaense (IFRD 9303)
aHost leaves. bAscoma on leaf
surface. cSquash mount of
ascoma. d, e Ascus with
paraphyses. f, g Ascospores. h, i
Ascospores mounted in cotton
blue reagent. Scale bars
b= 200 lm, c= 100 lm,
d=20lm, e=10lm, f–
i=5lm
Fungal Diversity
123
epidermoidea, with hyaline, membranous margin, lacking a
basal plate. Hamathecium comprising asci inclined from
the base towards the ostiole. Paraphyses present, about
1.5 lm wide. Asci 45–51 916–23 lm
(
x= 47.5 918.5 lm, n = 15), 8-spored, bitunicate, cla-
vate, ocular chamber present. Ascospores 24–38 96–
9lm(
x=3297.5 lm, n = 20), overlapping, 2-seriate,
cylindrical, 6–7-septate, constricted at the septum, head
cell a little larger. Asexual morph: Undetermined.
Material examined: CHINA, Xi Shuang Ban Na, Meng
La city, on living leaves of Amomum villosum
Fig. 16 Scolecopeltidium
wangtianshuense (IFRD 9302)
aHost leaves. b–d Colonies on
leaf surface. e, f Squash mount
of ascoma mounted in cotton
blue reagent. g, i, j Asci. h, k
Asci mounted in cotton blue
reagent. l–p Ascospores
mounted in cotton blue reagent.
Scale bars b= 400 lm,
c= 200 lm, d= 100 lm,
e=50lm, f=25lm, g–
o=10lm, p=5lm
Fungal Diversity
123
(Zingiberaceae), 6 March 2013, Hai-Xia Wu (IFRD 9303,
holotype; HKAS 102285, isotype).
Notes: This new species is placed in Scolecopeltidium
based on its greenish upper walls and 7-septate ascospores.
There is no record of Scolecopeltidium species that found
from the host genus Amomum.Scolecopeltidium men-
glaense is most similar to S. cassiae Bat. in ascospore size,
but differs in the septation of ascospores and the size of
asci. In phylogenetic tree, the new species clusters within
the family Micropeltidaceae as a sister to S. wangtian-
shuiense (the new species below) with high bootstrap value
(100% ML and 1.00 BYPP). Scolecopeltidium menglaense
is 96% (796/831 nucleotides, 1/831 gaps) similar to S.
wangtianshuiense in LSU region, and has smaller ascomata
and less septation of ascospores. Therefore, we introduce it
as a new species based on morphology and phylogeny.
Scolecopeltidium wangtianshuiense H.X. Wu, X.Y. Zeng
& K.D. Hyde, sp. nov.
Index Fungorum number: IF 555497; Facesoffungi
number: FoF 05143, Fig. 16
Etymology: in reference to its association (Latin -ensis)
with Wang Tian Shui = wangtianshuiense.
Holotype: IFRD 9302
Colonies epiphyllous, visible as black dots. Hyphae
superficial, hyaline, hard to detect. Sexual morph:Asco-
mata up to 460 lm in diam., superficial, solitary, gregari-
ous, black, flattened, circular, with a papillate central
ostiole. Upper wall greenish, comprising overlapping, net-
like, compact, pseudoparenchymatous hyphae with textura
epidermoidea, with hyaline, membranous margin, lacking a
basal plate. Hamathecium comprising asci inclined from
the base towards the ostiole. Paraphyses filiform. Asci 49–
83.5 915.6–18.6 lm(
x= 69.5 916.5 lm, n = 15), 8-
spored, bitunicate, clavate, ocular chamber present. As-
cospores 25.4–44.2 94.3–6.1 lm(
x= 36.5 95lm,
n = 20), overlapping, 2-seriate, cylindrical, 8-septate,
constricted at the septum, head cell a little larger. Asexual
morph: Undetermined.
Material examined: CHINA, Xi Shuang Ban Na, Wang
Tian Shu, on living leaves of Amomum villosum
Fig. 17 Stomiopeltopsis
linacearum (URM 18108)
aThe specimen. b, c Colonies
on leaf surface. dSquash mount
of ascoma. eThe upper wall.
fCross section of ascoma. g–i
Asci mounted in Melzer
reagent. j, k Ascospores
mounted in Melzer reagent.
Scale bars c, d = 200 lm, e–
k=10lm
Fungal Diversity
123
(Zingiberaceae), 6 March 2016, Hai-Xia Wu (IFRD 9302,
holotype; HKAS 102284, isotype).
Notes: By having greenish upper walls and 8-septate
ascospores, this new collection belongs to Scolecopeltid-
ium. It is most similar to Scolecopeltidium garciniae
(Henn.) F. Stevens & Manter. in terms of ascospore size,
but differs in the presence of paraphyses and the size of
ascomata. Phylogenetically, this new collection clusters
with S. menglaense as discussed above. Therefore, we
introduce it as a new species based on morphology and
phylogeny.
Stomiopeltopsis Bat. & Cavalc., Publicac¸o
˜es Inst. Micol.
Recife 392: 30 (1963)
Index Fungorum number: IF 5275; Facesoffungi num-
ber: FoF 05144
Epiphytic on living leaves, epiphyllous, visible as black
dots. Hyphae superficial, hyaline, hard to detect. Ascomata
superficial, black, solitary, subdense, gregarious, flattened,
circular, dimidiate, papillate, with a circular central ostiole.
Upper wall blackish green, membranous, comprising
overlapping, compact, pseudoparenchymatous hyphae with
textura intricata, lacking a basal plate. Hamathecium
comprising asci inclined from the base towards the ostiole.
Paraphyses absent. Asci 8-spored, bitunicate, cylindrical to
fusoid. Ascospores, cylindrical, 3–4-septate, constricted at
septa, hyaline, head cell longer and larger than others.
Notes:Stomiopeltopsis was introduced by Batista et al.
(1963a) as a monotypic genus. Parvosympodium nolinae
A.W. Ramaley has been reported to be the anamorph of
Stomiopeltopsis nolinae (Ramaley 2002). Like other gen-
era, the greenish upper wall makes Stomiopeltopsis a
member of Micropeltidaceae.
Type species:Stomiopeltopsis linacearum Bat. &
Cavalc., Publicac¸o
˜es. Instituto de Micologia da Universi-
dade do Recife & Instituto Nacional de Pesquis 392: 31
(1963)
Index Fungorum number: IF 339762; Facesoffungi
number: FoF 05145, Fig. 17
Colonies epiphyllous. Sexual morph: Ascomata up to
1 mm in diam., subdense, gregarious, papillate, with a
raised, circular central ostiole. Upper wall blackish green,
comprising net-like hyphae. Asci 36–59 98–14 lm
(
x= 46.2 910.3 lm, n = 15), 8-spored, bitunicate, cylin-
drical to fusoid. Ascospores 18–21 95–7 lm
(
x= 19.7 95.6 lm, n = 20), overlapping, 2-seriate,
cylindrical, 3–4-septate, constricted at septa, hyaline, head
cell longer and larger than others. Asexual morph:
Undetermined.
Material examined: BRAZIL, on leaves of Linaceae,
collection date unknown, A. C. Batista (URM 18108,
holotype).
Stomiopeltopsis antidesmatis X.Y. Zeng, H.X. Wu & K.D.
Hyde, sp. nov.
Index Fungorum number: IF 555498; Facesoffungi
number: FoF 05146, Fig. 18
Etymology: Refer to the host genus Antidesma.
Holotype: MFLU 17-1055
Colonies epiphyllous, visible as black dots. Hyphae
superficial, hyaline, hard to detect. Sexual morph:Asco-
mata up to 450 lm in diam., superficial, solitary, subdense,
black, flattened, circular, dimidiate, with a raised, central
ostiole. Upper wall blackish green, membranous, com-
prising overlapping, net-like, branched, sparse, pseudo-
parenchymatous hyphae with textura epidermoidea,
lacking a basal plate. Hamathecium comprising asci
inclined from the base towards the ostiole. Paraphyses
absent. Asci 46–47 913 lm(
x=46913 lm, n = 3),
8-spored, bitunicate, clavate to subfusiform, with short
pedicel, ocular chamber present. Ascospores 18 94lm
(n = 4), overlapping, 2-seriate, cylindrical to subclavate,
3–4-septate, slightly constricted at the septum, head cell a
little larger, bottom cell a little longer, tapering at bottom.
Asexual morph: Undetermined.
Material examined: THAILAND, Trat, Amphoe Ko
Chang, Yuttha Navi Ko Chang Memorial, on living leaves
of Antidesma sp. (Phyllanthaceae), 27 April 2017, Xiang-
Yu Zeng (MFLU 17-1055, holotype; HKAS 101761,
isotype).
Notes: There are very few fruiting bodies found from
this new collection, and the fungus seems not completely
mature. Morphologically, this new collection is most sim-
ilar to the type species of Stomiopeltopsis. However, the
type has larger ascomata and thinner ascospores.
Stomiopeltopsis antidesmatis is the first sequence data of
Stomiopeltopsis, and constitutes an independent lineage
that close to Dictyothyriella. There are 95% (871/921
nucleotides, 9/921 gaps) similarity in LSU region, 84%
(319/382 nucleotides, 25/382 gaps) similarity in ITS region
and 88% (825/939 nucleotides, 2/939 gaps) similarity in
TEF region between Stomiopeltopsis antidesmatis and
Dictyothyriella garciniae, which are similar to the differ-
ences between Stomiopeltopsis antidesmatis and Micro-
peltis asiatica. Therefore, Stomiopeltopsis antidesmatis sp.
nov. is introduced.
Genera transferred to Armatellaceae (Meliolales)
Armata W. Yamam., Sci. Rep. Hyogo Univ. Agric. 3 (2):
89 (1958)
Index Fungorum number: IF 308; Facesoffungi number:
FoF 05147
Colonies hypophyllous, scattered. Hyphae superficial,
brown, undulate, closely reticulate, with hyphopodia.
Hyphopodia alternate to irregular, 1-septate, without
phialides and hyphal setae. Head cell irregular, more or
Fungal Diversity
123
less angular. Perithecia up to 300 lm in diam., superficial,
globose, black. Paraphyses numerous, filiform, single,
hyaline. Asci 8-spored, ellipsoid. Ascospores ellipsoid,
rounded at both ends, continuous, olive-brown or brown.
Notes:Armata was introduced by Yamamoto (1958)asa
member of Meliolales and comprises a single species Ar-
mata macrospora (W. Yamam.) W. Yamam. that found on
leaves of Phoebe formosana. It had been placed in
Micropeltidaceae by Lumbsch and Huhndorf (2010). The
genus is characterised by superficial, branched hyphae with
hyphopodia, superficial, globose, dark perithecia without
setae, 8-spored asci contains ellipsoid, brown ascospores.
The character of ascospores is not clear in the protologue,
but seems to resemble a chain-like structure. As we were
not able to locate the type specimen, we transfer it back to
Armatellaceae (Meliolales) according to the description.
Type species:Armata macrospora (W. Yamam.) W.
Yamam., Sci. Rep. Hyogo Univ. Agric. 3 (2): 89 (1958)
:Armatella macrospora W. Yamam., Sci. Rep. Hyogo
Univ. Agric. 3 (1): 21 (1957)
Genera transferred to Asterinales incertae sedis
Caudella Syd. & P. Syd., Annls mycol. 14(1-2): 90 (1916)
Index Fungorum number: IF 852; Facesoffungi number:
FoF 05211
Epiphytic on living leaves. Colonies scattered to sub-
dense. Hyphae superficial, brown, reticulate, with alternate
sparse hyphopodia. Thyriothecia hypophyllous, dense,
circular, flattened, black, with a central ostiole. Upper wall
brown, comprising radial arrangement of irregularly lobed
cells, tightly fitting together like a jigsaw puzzle. Pseu-
doparaphyses filiform, thin. Asci 8-spored, bitunicate,
cylindrical to clavate. Ascospores fusoid to subclavate,
1-septate, not constricted at the septum, round at apex,
lower cell longer, tapering at the end, hyaline (Wu et al.
2011).
Notes:Caudella was introduced by Sydow and Sydow
(1916) as a member of Microthyriaceae and is typified by
Caudella oligotricha Syd. & P. Syd. that found on leaves of
Flacourtiaceae. Wu et al. (2011) transferred it to Micro-
peltidaceae. However, it must have been placed in
Fig. 18 Stomiopeltopsis
antidesmatis (MFLU 17-1055)
aHost leaves. bAscomata on
host surface. cSquash mount of
ascoma. dThe upper wall. eThe
hamathecium. f, g Asci with
ascospores. Scale bars
b= 200 lm, c= 100 lm, d–
g=20lm
Fungal Diversity
123
Micropeltidaceae by mistake due to its brown thyriothecia,
hyphopodia on hyphae and 2-celled ascospores. We
therefore transfer it to Asterinales based on characters
defined by Hongsanan et al. (2014).
Type species:Caudella oligotricha Syd. & P. Syd.,
Annls mycol. 14(1-2): 90 (1916)
Genera transferred to Capnodiales incertae sedis
Anariste Syd., Annls mycol. 25 (1-2): 76 (1927)
Index Fungorum number: IF 189; Facesoffungi number:
FoF 05148
Colonies hypophyllous, scattered. Hyphae superficial,
brown, reticulate, with hyphopodia. Ascomata solitary,
flattened, circular, black, with a circular central ostiole.
Upper wall dark brown, composed of radial arrangement of
hyphae. Paraphyses hyaline, thin. Asci 8-spored, bituni-
cate, cylindrical to clavate, ocular chamber present. As-
cospores clavate, 1-septate, slightly constricted at the
septum, upper cell globose to oval, lower cell tapering at
the end.
Notes:Anariste was first introduced by Sydow (1927)as
a member of Microthyriaceae and is typified by a single
species Anariste poliothea Syd. which were found on
leaves of Phoebe neurophylla. von Arx and Mu
¨ller (1975)
and Hosagoudar et al. (2001) placed the genus in the family
Asterinaceae, while Hongsanan et al. (2014) transfer this
genus from Asterinaceae to Micropeltidaceae due to its
unusual ascospores. There is no sequence data for this
genus so far, but its brown upper walls is distinguishable
from the hyaline hyphae and greenish upper walls in
Micropeltidaceae. We therefore transfer it to Capnodiales
according to its morphology.
Type species:Anariste poliothea Syd., Annls mycol. 25
(1-2): 76 (1927)
:Asterina poliothea (Syd.) F. Stevens, Illinois Biol.
Monogr. 17 (2): 64 (1939)
Stigmatodothis Syd. & P. Syd., Philipp. J. Sci., C, Bot.
9(2): 173 (1914)
Index Fungorum number: IF 5250; Facesoffungi num-
ber: FoF 05149
Epiphytic on living leaves, visible as black dots. Hyphae
superficial, brown, branched. Thyriothecia hypophyllous,
amphigenous, scattered to dense, circular, black, with a
central circular ostiole. Upper wall dark brown, comprising
radial arrangement, meandrous, compact hyphae, lacking a
Fig. 19 Stigmatodothis
palawanensis (S F6523) aThe
specimen. bColonies on leaf
surface. cSquash mount of
thyriothecium. d, e Cross
section of thyriothecium. f–i
Asci. j–l Ascospores. Scale bars
b= 200 lm, c=50lm,
d=20lm, e–i =10lm, j–
l=5lm
Fungal Diversity
123
basal plate. Asci 8-spored, bitunicate, ellipsoid to clavate.
Ascospores overlapping, 2–3 series, cylindrical, 3-septate,
mostly not constricted at septa, hyaline.
Notes:Stigmatodothis was introduced by Sydow and
Sydow (1914) and is typified by Stigmatodothis palawa-
nensis Syd. & P. Syd. that found on leaves of Dendrobium
sp. von Arx and Mu
¨ller (1975) reviewed Stigmatodothis
and described S. palawanensis as the type species. We
transfer this genus from Micropeltidaceae to Capnodiales
due to its brown upper wall and hyaline, muti-septate
ascospores.
Type species:Stigmatodothis palawanensis Syd. & P.
Syd., Philipp. J. Sci., C, Bot. 9(2): 173 (1914)
Index Fungorum number: IF 159709; Facesoffungi
number: FoF 05150, Fig. 19
Colonies hypophyllous, amphigenous, visible as black
dots. Hyphae superficial, brown, branched. Sexual morph:
Thyriothecia up to 180 lm in diam., dense, grouped, cir-
cular, black, with a central circular ostiole. Upper wall dark
brown, comprising radial arrangement, meandrous, com-
pact hyphae, lacking a basal plate. Asci 23–37 911–
20 lm(
x=30914 lm, n = 15), 8-spored, bitunicate,
ellipsoid to clavate, orcular chamber not observed. As-
cospores 7–14 93–4.5 lm(
x=1194lm, n = 20),
overlapping, 2–3 series, cylindrical, 3-septate, constricted
at the middle septum, hyaline. Asexual morph:
Undetermined.
Fig. 20 Stomiopeltis aspersa
(K 164017) aThe specimen.
bColonies on leaf surface.
cSquash mount of
thyriothecium. dThe upper
wall. e, g–j Asci mounted in
Melzer reagent. fAsci. k–n
Ascospores mounted in Melzer
reagent. Scale bars b= 200 lm,
c=50lm, d=10lm, e–
n=5lm
Fungal Diversity
123
Material examined: PHILIPPINES, Palawan, Taytay; on
leaves of Dendrobii sp., 12 May 1913, Merrill E. D. No.
8909 (S F6523, holotype).
Stomiopeltis Theiss., Brote
´ria, se
´r. bot. 12: 85 (1914)
=Akaropeltella M.L. Farr, Mycologia 64(2): 252 (1972)
=Akaropeltis Bat. & J.L. Bezerra, Publicac¸o
˜es Inst.
Micol. Recife 313: 5 (1961)
=Akaropeltopsis Bat. & Peres, Atas Inst. Micol. Univ.
Recife 3: 45 (1966)
=Clypeolina Speg., Hong. Exo
´t.: 27 (1923)
=Clypeolinopsis Bat., Publicac¸o
˜es Inst. Micol. Recife
56: 315 (1959)
=Clypeolopsis F. Stevens & Manter, Bot. Gaz. 79(3):
287 (1925)
=Diplocarponella Bat., Anais Soc. Biol. Pernambuco
15(1): 141 (1957)
=Leptopeltina Speg., Boln Acad. nac. Cienc. Co
´rdoba
27(4): 397 (1924)
=Sirothyriella Ho
¨hn., Sber. Akad. Wiss. Wien, Math.-
naturw. Kl., Abt. 1 119: 451 (1910)
=Stomiopeltella Theiss., Brote
´ria, se
´r. bot. 12: 86
(1914)
=Verlandea Bat. & Cif., Beih. Sydowia 1: 338 (1957)
[1956]
Index Fungorum number: IF 5274; Facesoffungi num-
ber: FoF 05151
Epiphytic on living leaves, visible as black dots. Hyphae
superficial, brown, reticulate. Thyriothecia hypophyllous,
dense to subdense, flattened, circular, brown, with an
irregular central ostiole. Upper wall brown, comprising
meandrous, compact hyphae that anastomosing at central
ostiole, lacking a basal plate. Paraphyses filiform, dense.
Asci 8-spored, bitunicate, cylindrical to clavate. Ascospores
overlapping, 2–3 series, cylindrical, 1-septate, slightly
constricted at the septum or not, hyaline.
Notes:Stomiopeltis was introduced by Theissen (1914)
to accommodate a single species Stomiopeltis aspersa
(Berk.) Theiss., which was transferred from Calothyrium
(Microthyriaceae) to Micropeltidaceae. The genus is dis-
tinguished from other genera in the subfamily Plochmo-
peltineae by the presence of paraphyses. The type specimen
from K herbarium is in poor condition, and the specimens
from GZU herbarium differs from the type in texture of
upper wall. Nevertheless, the brown upper walls as well as
phylogenetic analyses both illustrate that Stomiopeltis is a
member of Capnodiales.
Type species:Stomiopeltis aspersa (Berk.) Theiss.,
Brote
´ria, se
´r. bot. 12: 85 (1914)
:Asterina aspersa Berk., Hooker’s J. Bot. Kew Gard.
Misc. 6: 210 (1854)
=Calothyrium aspersum (Berk.) Theiss., O
¨st. bot. Z.
62(6): 219 (1912)
Index Fungorum number: IF 221479; Facesoffungi
number: FoF 05152, Fig. 20
Colonies hypophyllous, scattered, visible as black dots.
Hyphae superficial, brown, reticulate. Sexual morph:
Thyriothecia up to 176 lm in diam., subdense, flattened,
circular, brown, easily removed from the host surface, with
a central ostiole. Upper wall brown, comprising mean-
drous, compact hyphae, lacking a basal plate. Pseudopa-
raphyses filiform. Asci 30–43 99–12 lm
(
x=36910 lm, n = 15), 8-spored, ellipsoid, with short
pedicel, orcular chamber present. Ascospores 8–11 92.5–
3lm(
x=1093lm, n = 20), overlapping, 2–3 series,
cylindrical, 1-septate, not constricted at the septum, hya-
line, upper cell shorter and broader than lower cell.
Asexual morph: Undetermined.
Material examined: INDIA, Khasia, on leaves of Laurus
sp., collection date unknown, Berkeley (K 164017,
holotype).
Genera transferred to Dothideomycetes incertae sedis
Dictyopeltis Theiss., Annls mycol. 11(5): 469 (1913)
Index Fungorum number: IF 1534; Facesoffungi num-
ber: FoF 05153
Epiphytic on living leaves, visible as black dots. Hyphae
superficial, brown. Ascomata solitary, flattened, black,
ostiolate, forming above a layer of radiate, membranous
plant cells. Mycelial pellicle brown, comprising radiate,
meandrous, compact hyphae that anastomosing at central
ostiole. Peridium light brown, with a layer of compressed
cell. Pseudoparaphyses filiform. Asci 8-spored, bitunicate,
clavate to ellipsoid. Ascospores cylindrical, 3-septate,
hyaline.
Notes:Dictyopeltis was introduced in Hemisphaeriaceae
by Theissen (1913). The specimen is different from the
protologue in the presence of ostiole and paraphyses, sep-
tation of ascospores, and the size of both asci and ascos-
pores. It is difficult to place Dictyopeltis in a certain order
without any reliable sequence data. However, we exclude
this genus from Micropeltidaceae due to its brownish upper
wall.
Type species:Dictyopeltis vulgaris (Racib.) Theiss.,
Annls mycol. 11(5): 469 (1913)
:Clypeolum vulgare Racib., Bull. int. Acad. Sci. Lett.
Cracovie, Cl. sci. math. nat. Se
´r. B, sci. nat. 3: 384 (1909)
Index Fungorum number: IF 191844; Facesoffungi
number: FoF 05154, Fig. 21
Colonies epiphyllous, visible as black dots. Hyphae
superficial, brown. Sexual morph: Ascomata up to 300 lm
in diam., solitary, flattened, black, ostiolate, forming above
a layer of radiate, membranous plant cells. Mycelial pel-
licle brown, comprising radiate, meandrous, compact
hyphae that anastomosing at central ostiole. Peridium light
brown, with a layer of compressed cell. Pseudoparaphyses
Fungal Diversity
123
filiform. Asci 36.5–40 98.5–10 lm(
x=3899.5 lm,
n = 5), 8-spored, bitunicate, clavate to ellipsoid, orcular
chamber not observed. Ascospores 16.5–22.5 93–5 lm
(
x= 18.5 94lm, n = 10), cylindrical, 3-septate, hyaline.
Asexual morph: Undetermined.
Material examined: INDONESIA, Java, Tjampea, on
leaves of Hydnocarpus sp., collection date unknown, Leg.
M. Raciborski (ZTMyc 56239, holotype).
Stomiotheca Bat., Publicac¸o
˜es Inst. Micol. Recife 56: 453
(1959)
Index Fungorum number: IF 20614; Facesoffungi
number: FoF 05155
Epiphytic on living leaves. Hyphae superficial, brown,
septate, flexuous to substraight, twisted, closely reticulate.
Thyriothecia hypophyllous, solitary or grouped, gregarious,
dense, brown, circular, flattened, forming beneath a layer
of interwoven hyphae, with an irregular central ostiole.
Upper wall light brown, comprising radiate, meanderous,
compact hyphae that anastomosing at central ostiole,
lacking a basal plate. Pseudoparaphyses branched, thin,
dense, numerous. Asci 8-spored, bitunicate, cylindrical to
subclavate, with long pedicel, ocular chamber present.
Ascospores overlapping, 2–4 seriate, clavate, 2-septate,
slightly constricted at the top septum, hyaline, head cell
oval to obovoid, 2-celled tail filiform, tapering at the end,
not constricted at the bottom septum.
Notes:Stomiotheca was introduced by Batista (1959)
and maintained its placement in Micropeltidaceae till now.
The type specimen from NY differs from the protologue in
septation of hyphae, guttulas in ascospores and colour of
ascospores. Nevertheless, we exclude this genus from
Micropeltidaceae based on its brown upper wall, but cannot
place it in a certain order without sequence data.
Type species:Stomiotheca amazonensis Bat., Pub-
licac¸o
˜es Inst. Micol. Recife 56: 457 (1959)
Index Fungorum number: IF 306526; Facesoffungi
number: FoF 05156, Fig. 22
Colonies hypophyllous, scattered. Hyphae superficial,
brown, septate, flexuous to substraight, twisted, closely
reticulate. Sexual morph:Thyriothecia up to 660 lmin
diam., solitary or grouped, gregarious, dense, brown, cir-
cular, flattened, forming beneath a layer of interwoven
hyphae, with an irregular central ostiole. Upper wall light
brown, comprising radiate, meanderous, compact hyphae
that anastomosing at central ostiole, lacking a basal plate.
Pseudoparaphyses branched, thin, dense, numerous. Asci
(123-)134–160(-165) 923–31 lm(
x= 144 925 lm,
n = 10), 8-spored, bitunicate, cylindrical to subclavate,
with long pedicel, ocular chamber present. Ascospores 60–
Fig. 21 Dictyopeltis vulgaris
(ZTMyc 56239) aThe material
packet. bThe specimen.
cAscomata on leaf substrate.
dSquash mount of ascoma in
overview. eThe upper wall.
fThe basal wall.
gPseudoparaphyses. hAsci. i, j
Ascospores. Scale bars d,
e=50lm, f–h =20lm, i,
j=10lm
Fungal Diversity
123
80 98–10 lm(
x=71910 lm, n = 20), overlapping,
2–4 seriate, clavate, 2-septate, slightly constricted at the
top septum, hyaline, head cell oval to obovoid, 2-celled tail
filiform, tapering at the end, not constricted at the bottom
septum. Asexual morph: Unknown.
Material examined: BRAZIL, Amazonas, Colonia Pedro
Borges, on dried leaves of an unidentified plant, 22 August
1923, J. R. Weir (NY 914557, isotype)
Genera transferred to Microthyriales incertae sedis
Heliocephala V. Rao, K.A. Reddy & de Hoog, Persoonia
12(3): 239 (1984)
=Holubovaniella R.F. Castan
˜eda, Deuteromycotina de
Cuba, Hyphomycetes (La Habana) 3: 14 (1985)
Index Fungorum number: IF 11097; Facesoffungi
number: FoF 05157
Colonies effuse, light to dark grey. Conidiophores
macronematous, mononematous, brown, bearing terminal
conidiogenous cells which are monoblastic, ampulliform or
ovoidal in dense, drepanoid arrangement. Conidia
Fig. 22 Stomiotheca amazonensis (NY 914557) aThe material
packet and specimen. bThyriothecia on leaf substrate. cClose up of
thyriothecium. dSquash mount of thyriothecium in overview. eThe
upper wall covered by aerial hyphae. fSeptate hyphae. gParaphyses.
h–k Asci mounted in 5% KOH. l, m Ascospores mounted in 5%
KOH. Scale bars b= 400 lm, c, d = 200 lm, e=50lm, f, h–
k=20lm, g, l, m =10lm
Fungal Diversity
123
produced in radial, compact heads, dry, obclavate, 2-sep-
tate, rostrate, sometimes bearing heads of second or third
order.
Notes:Heliocephala was introduced by Rao et al. (1984)
as an asexual genus of Micropeltidaceae (Microthyriales).
Abarca et al. (2011) reported that Holubovaniella is a
synonym of Heliocephala and is related to Stomiopeltis
betulae. It is difficult to compare the morphology between
asexual fungi and sexual fungi, but we transfer it to
Microthyriales based on the present phylogeny.
Type species:Heliocephala proliferans V. Rao, K.A.
Reddy & de Hoog, Persoonia 12(3): 239 (1984)
Mitopeltis Speg., Boln Acad. nac. Cienc. Co
´rdoba 25: 93
(1921)
Index Fungorum number: IF 3218; Facesoffungi num-
ber: FoF 05158
Epiphytic on living leaves, visible as black dots. Hyphae
superficial, brown, septate, reticulate. Thyriothecia
amphigenous, circular, flattened, black, with a central
ostiole. Upper wall brown, membranous, comprising cells
of radial and parallel arrangements. Paraphyses absent.
Asci 8-spored, bitunicate, clavate, with short pedicel. As-
cospores 2-seriate, cylindrical, 3-septate, slightly con-
stricted at the septum, hyaline.
Notes:Mitopeltis was introduced with a single species
M. chilensis Speg. by Spegazzini (1921). The holotype
specimen from LPS was in poor condition, and only few
ascomata are available. Nevertheless, stellate lobulated
ascomata and texture of upper walls are quite different
from species of Micropeltidaceae, according to the original
descriptions and illustrations. Therefore, we transfer Mi-
topeltis to Microthyriales due to its radial structure of upper
wall.
Type species:Mitopeltis chilensis Speg., Boln Acad.
nac. Cienc. Co
´rdoba 25: 93 (1921)
Index Fungorum number: IF 158806; Facesoffungi
number: FoF 05159, Fig. 23
Colonies amphigenous, visible as black dots. Hyphae
superficial, brown, septate, reticulate. Sexual morph:
Thyriothecia up to 190 lm in diam., epiphytic, circular,
flattened, black, with a central ostiole. Upper wall brown,
membranous, comprising cells of radial and parallel
arrangements. Paraphyses absent. Asci
40–50 915–18 lm (from the protologue), 8-spored,
bitunicate, clavate, with short pedicel, orcular chamber not
observed. Ascospores 22–24 94–5 lm (from the proto-
logue), 2-seriate, cylindrical, 3-septate, slightly constricted
at the septum, hyaline. Asexual morph: Undetermined.
Material examined: CHILE, Los Perales, on stem of
Jubaea spectabilis, 26 April 1917, C. Spegazzini (LPS
1053, holotype)
Fig. 23 Mitopeltis chilensis
(LPS 1053) aThe specimen.
bThyriothecium on leaf
surface. cSuperfical hyphae on
leaf surface. dSquash mount of
thyriothecium. eHyphae. fThe
drawing the upper wall and
hyphae. gThe drawing of asci
and ascospores. Scale bars b,
c= 100 lm, d, e =20lm, f,
g=10lm
Fungal Diversity
123
Thyriodictyella Cif., Atti Ist. bot. Univ. Lab. crittog. Pavia,
Ser. 4 19: 129 (1962)
Index Fungorum number: IF 5467; Facesoffungi num-
ber: FoF 05160
Epiphytic on living leaves, visible as black dots. Hyphae
superficial, brown, flexuous, anastomosing at central osti-
ole. Thyriothecia hypophyllous, dense to subdense, flat-
tened, circular, papillate, with an irregular central ostiole.
Upper wall dark brown, comprising radial arrangement of
irregularly lobed hyphae that anastomosing at central
ostiole. Asci 8-spored, bitunicate, cylindrical, lack pseu-
doparaphyses. Ascospores overlapping, 2-seriate, cylindri-
cal to fusoid, 1-septate, very thick, not constricted at the
septum, hyaline.
Notes:Thyriodictyella was introduced by Ciferri (1961)
with a single species T. microsperma (Syd. & P. Syd.) Cif.
which was found on leaves of Uncaria guianensis. The
placement of the genus has not changed since introduced as
a member of Micropeltidaceae. However, the characters of
this genus are more similar to Microthyriaceae by having
radiate arrangement of upper wall, circular central ostiole,
brown hyphae without hyphopodia, and hyaline, 1-septate
ascospores. Therefore, we exclude it from
Micropeltidaceae.
Type species:Thyriodictyella microsperma (Syd. & P.
Syd.) Cif., Atti Ist. bot. Univ. Lab. crittog. Pavia, Ser. 4 19:
129 (1962)
:Micropeltella microsperma Syd. & P. Syd., Annls
mycol. 14(1–2): 89 (1916)
=Dictyothyriella microsperma (Syd. & P. Syd.) F.
Stevens & Manter, Bot. Gaz. 79(3): 273 (1925)
Index Fungorum number: IF 340080; Facesoffungi
number: FoF 05161, Fig. 24
Colonies hypophyllous, visible as black dots. Hyphae
superficial, brown, flexuous, anastomosing at central osti-
ole. Sexual morph:Thyriothecia 160–190 lmin
diam. 923–63 lm high, dense to subdense, flattened,
circular, papillate, with an irregular central ostiole. Upper
wall dark brown, comprising radial arrangement of irreg-
ularly lobed hyphae that anastomosing at central ostiole.
Asci 55–75 99–12 lm(
x= 65.8 910.6 lm, n = 15), 6–
8-spored, bitunicate, cylindrical, lack pseudoparaphyses.
Ascospores 15–21 94–5 lm(
x= 17.5 94lm, n = 20),
overlapping, 2-seriate, cylindrical to fusoid, 1-septate, very
Fig. 24 Thyriodictyella
microsperma (S F8633-8634)
aThe specimen. bColonies on
leaf surface. cSquash mount of
thyriothecium. dCross section
of thyriothecium. eThe upper
wall. f–i Asci. j–m Ascospores.
Scale bars b= 200 lm, c,
e=20lm, d, f–i =10lm, j–
m=5lm
Fungal Diversity
123
thick, not constricted at the septum, hyaline. Asexual
morph: Undetermined.
Material examined: BRAZIL, Rio, Seringal Auristella,
on dried leaves of an unidentified plant, 8 June 1911, J.
Gmel (S F8633-8634, holotype).
Doubtful genera
Bonaria Bat., Publicac¸o
˜es Inst. Micol. Recife 56: 438
(1959)
Index Fungorum number: IF 624; Facesoffungi number:
FoF 05164
Colonies epiphyllous, dense, confluent. Hyphae super-
ficial, brown, reticulate, without setae or hyphopodia. As-
comata circular, brown, ostiolate, lack basal stroma. Asci
8-spored, bitunicate, lack paraphyses. Ascospores fusiform
to clavate,1-septate, hyaline.
Notes:Bonaria was introduced as a member of Micro-
peltidaceae, and is typified by B. lithocarpi (V.A.M. Mill.
& Bonar) Bat. that was found on leaves of Lithocarpus
densiflorus. We could not locate any specimens of this
genus. However, its brown upper walls described in the
protologue makes it distinct from member of the Micro-
peltidaceae. Hence, we treat it as a doubtful genus as no
specimen from Bonaria has been re-examined.
Type species:Bonaria lithocarpi (V.A.M. Mill. &
Bonar) Bat., Publicac¸o
˜es Inst. Micol. Recife 56: 439 (1959)
:Protopeltis lithocarpi V.A.M. Mill. & Bonar,
University of Calif. Publ. Bot. 19: 412 (1941)
Clypeolina (Theiss.) Theiss., in Theissen & Sydow, Annls
mycol. 15(6): 419 (1917)
=Clypeolella sect. Clypeolina Theiss., Centbl. Bakt.
ParasitKde, Abt. II 34(8–9): 234 (1912)
Index Fungorum number: IF 1117; Facesoffungi num-
ber: FoF 05165
Subiculum composed of dark brown hyphae, irregularly
branched, densely compact, without hyphopodia. Thyrio-
thecia inverted, flattened-conical, circular, brown. Asci
8-spored, lack paraphyses. Ascospores oblong to ellipsoid,
gray-brown, rounded at both sides, constricted at the sep-
tum, upper cell slightly wider but smaller.
Notes:Clypeolina was first introduced as a member of
Microthyriaceae (Theissen and Sydow 1917). Lumbsch
and Huhndorf (2010) placed the genus in Micropeltidaceae.
Its dark brown subiculum with branched hyphae and brown
2-celled ascospores make it different from Micropelti-
daceae. However, none of the specimens of this genus was
obtained. Therefore, we treat it as a doubtful genus.
Type species:Clypeolina apus (Theiss.) Theiss., in
Theissen & Sydow, Annls mycol. 15(6): 419 (1917)
:Clypeolella apus Theiss., Centbl. Bakt. ParasitKde,
Abt. II 34(8–9): 234 (1912)
Cyclopeltis Petr., Sydowia 7(5-6): 370 (1953)
Index Fungorum number: IF 1374; Facesoffungi num-
ber: FoF 05166
Ascomata amphigenous, scattered, circular, black, mar-
gin lobate. Hypostroma dark brown, in epidermal pseudo-
parenchyma, Asci widely ovoid or ellipsoid, 8-spored.
Ascospores oblong, ovoid or ellipsoid, hyaline at young
state, become olive at mature state.
Notes:Cyclopeltis was placed in Polystomellaceae when
introduced by Petrak (1953). Lumbsch and Huhndorf
(2010) placed the genus in Micropeltidaceae. Although we
could not locate any specimens of this genus, we exclude it
from Micropeltidaceae due to its coloured ascospores.
Type species:Cyclopeltis orbicularis Petr., Sydowia
7(5-6): 370 (1953)
Dictyostomiopelta Vie
´gas, Bragantia 4(1-6): 53 (1944)
Index Fungorum number: IF 1536; Facesoffungi num-
ber: FoF 05167
Hyphae septate, branched, hard to detect. Ascomata
punctiform, flattened, circular, with more or less sharp
margin. Asci 8-spored, globose to clavate, thickened at the
apex, with short pedicel. Ascospores oblong, muriform,
constricted at septa, hyaline, smooth.
Notes:Dictyostomiopelta was introduced by Vie
´gas
(1944) as a member of Hemisphaeriaceae (Microthyriales)
and is typified by a single species Dictyostomiopelta
manihoticola Vie
´gas that found on stems of Manihot
utilissima. We were not able to locate the type specimen,
but we exclude this genus from Micropeltidacceae based
on its muriform ascospores.
Type species:Dictyostomiopelta manihoticola Vie
´gas
[as ‘manigoticola’], Bragantia 4(1-6): 54 (1944)
Hansfordiopsis Bat., Publicac¸o
˜es Inst. Micol. Recife 56:
407 (1959)
Index Fungorum number: IF 2221; Facesoffungi num-
ber: FoF 05162
Lack free hyphae. Thyriothecia epiphyllous, flattened,
orbicular, brown, with irregular dehiscence and clear
margins. Upper wall pseudo-parenchyma, comprising
reticulate hyphae. Asci 8-spored, unitunicate, lack para-
physes. Ascospores clavate or subglobose, 1-septate, con-
stricted at the septum, dark brown.
Notes: The genus Hansfordiopsis was introduced by
Batista (1959) with a single species Hansfordiopsis
phaeospora (Hansf.) Bat., which was synonymised under
Microthyriella phaeospora Hansf. (Batista 1959). How-
ever, the taxon found from the holotype is Enterographa
bella, a lichenised Arthoniales. A technical error must have
been done by the staff of the herbarium, like mislabeling of
the specimen. An epitype should be collected from the type
locality for further studies. Therefore, we treat Hansfor-
diopsis as a doubtful genus.
Fungal Diversity
123
Type species:Hansfordiopsis phaeospora (Hansf.) Bat.,
Publicac¸o
˜es Inst. Micol. Recife 56: 407 (1959)
:Microthyriella phaeospora Hansf., Proc. Linn. Soc.
N.S.W. 79(3-4): 112 (1954)
Enterographa bella R. Sant., Symb. bot. upsal. 13(no. 1):
106 (1952) Fig. 25
Colonies epiphyllous, developing on a thin sheath.
Sexual morph:Thyriothecia superficial, flattened, linear to
branched, brown, solitary, poorly develop at the basal.
Upper wall brown, composed of textura globulosa inner
cells and textura angularis outer cells, with hyaline margin.
Pseudoparaphyses thin, dense. Asci 42–48(-59) 920–
26 lm(
x=47923 lm, n = 5), 8-spored, bitunicate,
subglobose to ellipsoid, orcular chamber not obvious. As-
cospores 20–25 93–5 lm(
x=2494lm, n = 10),
overlapping, 2–3-seriate, fusiform, 7-septate, not con-
stricted at septa, hyaline, top third and fourth cells bigger
than the others, tapering at both ends, with hyaline sheath.
Asexual morph: Unknown.
Fig. 25 Enterographa bella (PDD 6788) aThe material packet.
bSpecimens. c, d Colonies on leaf substrate. e, f Squash mount of
ascoma. gThe margin of ascomata. hCross section of ascomata.
iPseudoparaphyses. j–l Asci. m–o Ascospores. Scale bars
e= 200 lm, f, g = 100 lm, h=50lm, i–l =20lm, m–o =10lm
Fungal Diversity
123
Material examined: NEW ZEALAND, Kapiti Island, on
leaves of Knightia excelsa, November 1936, H. H. Allan
(PDD 6788)
Mendoziopeltis Bat., Publicac¸o
˜es Inst. Micol. Recife 56:
434 (1959)
Index Fungorum number: IF 3120; Facesoffungi num-
ber: FoF 05168
Hyphae superficial, brown, septate, reticulate, without
setae or hyphopodia. Ascomata flattened, brown, with a
central pseudo-ostiole and fimbriated margins. Upper wall
brown, comprising cells of hyphal network that anasto-
mosing at central ostiole, closely reticulate, constricted at
septa. Asci 3-spored, unitunicate, ellipsoid to subglobose,
lack paraphyses. Ascospores cylindrical, muriform, 7–9
transverse septa, 7–12 vertical septa, constricted at septa,
hyaline at young state, become light brown when mature,
with a basal hyaline appendage.
Notes:Mendoziopeltis was introduced in the monograph
of Micropeltidaceae by Batista (1959). We could not locate
the type specimen of this genus. Nevertheless, muriform
ascospores and texture of upper wall are quite different
from species of Micropeltidaceae, according to the original
descriptions and illustrations. Therefore, we exclude
Mendoziopeltis from Micropeltidaceae.
Type species:Mendoziopeltis byrsonimae Bat. & Nas-
cim. ex Bat., Publicac¸o
˜es Inst. Micol. Recife 56: 434
(1959)
Muricopeltis Vie
´gas, Bragantia 4(1-6): 55 (1944)
Index Fungorum number: IF 3291; Facesoffungi num-
ber: FoF 05169
Ascomata hypophyllous, black, grouped, ostiolate, with
hyaline margin. Upper wall comprising brown meandering
hyphae. Asci globose, 8-spored, hyaline, lack paraphyses.
Ascospores clavate, muriform, brown, smooth.
Notes:Muricopeltis was introduced as a member of
Hemisphaeriaceae (Microthyriales) by Vie
´gas (1944) and
is typified by a single species Muricopeltis brasiliensis
Vie
´gas that found on leaves of Piper sp. We were unable to
locate the type specimen, and descriptions were re-written
based on the protologue. Nevertheless, we exclude this
genus from Micropeltidacceae based on its brown, muri-
form ascospores.
Type species:Muricopeltis brasiliensis Vie
´gas, Bra-
gantia 4(1-6): 54 (1944)
Stigmatophragmia Tehon & G.L. Stout, Mycologia 21(4):
180 (1929)
Index Fungorum number: IF 5255; Facesoffungi num-
ber: FoF 05170
Epiphytic on living leaves. Ascomata epiphyllous,
widely scattered, flattened, circular, brown, opening by a
somewhat umbonate, carbonaceous, with a circular ostiole.
Upper wall brown, comprising a layer of pseudo-
parenchymatous membranous cell. Paraphyses filiform,
unbranched. Asci 8-spored, bitunicate, cylindrical, with
short pedicel. Ascospores cylindrical, 3-septate, constricted
at the septa, tapering at both ends, hyaline.
Notes:Stigmatophragmia was introduced as a mono-
typic genus in Stigmateaceae (Hemisphaeriales), which is
represented by Stigmatophragmia sassafrasicola Tehon &
G.L. Stout that were found on leaves of Sassafras vari-
ifolium (Tehon and Stout 1929). The specimen from ILLS
is in poor condition. Descriptions were made based on the
protologue. We excluded it from Micropeltidaceae due to
its brown upper walls and ascospores. Fresh collections are
needed for future molecular analyses.
Type species:Stigmatophragmia sassafrasicola Tehon
& G.L. Stout, Mycologia 21(4): 181 (1929)
Index Fungorum number: IF 183497; Facesoffungi
number: FoF 05171, Fig. 26
Colonies epiphyllous. Ascomata diaphyllous, circular or
somewhat angular, up to 225 lm in diam., tan to brown,
with a distinct, unraised, purplish margin. Upper wall
brown, comprising a layer of pseudoparenchymatous
membranous cell. Paraphyses filiform, unbranched, abun-
dant. Asci 65–80 910–15 lm (from the protologue),
8-spored, bitunicate, cylindrical, with short pedicel. As-
cospores 14–17 93–4 lm (from the protologue), cylin-
drical, 3-septate, constricted at the septa, tapering at both
ends, hyaline.
Material examined: AMERICA, Illinois, Champaign
County, Seymour, on leaves of Sassafras variifolium,15
October 1925, L. R. Tehon (ILLS 21698, holotype).
Polypedia Bat. & Peres, Atti Ist. bot. Univ. Lab. crittog.
Pavia, Ser. 4 16: 118 (1959)
Index Fungorum number: IF 4318; Facesoffungi num-
ber: FoF 05172
Epiphytic on living leaves, superficial hyphae absent.
Ascomata epiphyllous, scattered to subdense, flattened,
orbicular, black, opening by irregularly radiating fissures
when wet, intimately mixed with conidiomata. Upper wall
comprising two different tissue types, reddish brown tex-
tura prismatica in the outer wall and olive brown textura
angularis in the inner wall. Asci 8-spored, bitunicate,
cylindrical, clavate. Ascospores cylindrical to ellipsoid,
1-septate, hyaline, slightly verrucose. Conidiomata solitary
or some gregarious, circular. Conidia, aseptate, smooth,
ellipsoidal-globose to ovoid, wall with guttulate.
Notes:Polypedia was introduced with a single species
P. leopoldinensis Bat. & Peres by Batista and Peres (1959).
Ina
´cio and Cannon (2008) synonymised the type species to
be a member of Peltistroma. The specimen from URM is in
poor condition. However, the dark brown upper wall also
makes us believe that the genus is not a member of
Fungal Diversity
123
Micropeltidaceae. Fresh collections and sequences are
needed to establish their taxonomic affinities.
Type species: Peltistroma leopoldinense (Bat. & Peres)
Ina
´cio & P.F. Cannon, CBS Diversity Ser. (Utrecht) 8: 54
(2008)
=Polypedia leopoldinensis Bat. & Peres, Atti Ist. bot.
Univ. Lab. crittog. Pavia, Ser. 4 16: 119 (1959)
Index Fungorum number: IF 540722; Facesoffungi
number: FoF 05173, Fig. 27
Sexual morph: Ascomata epiphyllous, scattered to
subdense, flattened, orbicular, up to 2 mm in diam, black,
opening by irregularly radiating fissures when wet, inti-
mately mixed with conidiomata. Upper wall comprising
two different tissue types, reddish brown textura prismat-
ica in the outer wall and olive brown textura angularis in
the inner wall. Asci 48–67 910–15 lm (from the proto-
logue), 8-spored, bitunicate, cylindrical, clavate. As-
cospores 13–16 94–6 lm (from the protologue),
cylindrical to ellipsoid, aseptate, hyaline. Asexual morph:
Undetermined.
Material examined: BRAZIL, on unidentified coria-
ceous leaves, 9 July 1958, A.C. Batista (URM 14047,
holotype).
Discussion
Previous descriptions for Micropeltidaceae included 24
genera with bluish green upper walls with meandrous cells
(Lumbsch and Huhndorf 2010; Hyde et al. 2013). How-
ever, many of those genera (Bonaria,Caudella,Clype-
olina,Cyclopeltis,Chaetothyrina,Dictyopeltis,
Hansfordiopsis,Mendoziopeltis,Mitopeltis,Muricopeltis,
Stigmatodothis.Stigmatophragmia,Stomiopeltis,Stomio-
theca,Thyriodictyella and Polypedia) have brown and
similar structures of the upper walls as in Micropeltidaceae.
The morphological features of Micropeltidaceae are very
similar to Asterinaceae, Microthyriaceae and Phaeothe-
coidiellaceae. Asterinaceae is typified by superficial
hyphae with 1-celled hyphopodia, thyriothecia with a
central fissured dehisce, and brown ascospores (Hongsanan
Fig. 26 Stigmatophragmia
sassafrasicola (ILLS 21698
Notes: Erroneously as ILLS
20103 in the protologue) aThe
label of specimen. bThe
specimen. cFruit body on leaf
surface. d, e Hyphae. fThe
original drawing of an ascus
with spores and paraphyses by
Tehon and Stout (1929). g, h
The conidia and conidiole.
Scale bars c= 200 lm, d, e, g,
h=20lm
Fungal Diversity
123
et al. 2014). Microthyriaceae is typified by brown upper
walls with radial cells, a darkened central ostiole, and lack
hyphopodia on hyphae (Wu et al. 2011). Phaeothecoidiel-
laceae is a newly introduced flyspeck family typified by
brown upper walls with interwoven cells, setae and invis-
ible superficial hyphae (Hongsanan et al. 2017a). The
upper walls and form of ascospores appear to be important
characters to distinguish these similar groups. Asteri-
naceae, Microthyriaceae and Phaeothecoidiellaceae that
have scutate and brown upper walls are quite different from
Micropeltidaceae. Micropeltidaceae are characterised by
membranous, greenish upper walls. Phylogenies generated
herein also provide evidences to justify that these charac-
ters are phylogenetically significant in familial circum-
scription. Asterinaceae (characterised by brown, dimidiate
upper walls), Microthyriaceae (characterised by brown,
radial upper walls) and Phaeothecoidiellaceae (charac-
terised by brown, interwoven upper walls) are segregated
clades in Dothideomycetes and are distinct to Micropelti-
daceae (characterised by greenish, membranous upper
walls) in Lecanoromycetes (Fig. 1). Similar phylogenetic
scenarios have been reported previously where families
have been differentiated based on ascomatal features
among the Sordariomycetes and Dothideomycetes (Cai
et al. 2006; Kodsueb et al. 2006; Zhang et al. 2009; Hon-
gsanan et al. 2015). The upper wall of Micropeltidaceae is
more like a mycelial pellicle covering the leaf surface and
not a radial structure. Therefore, ascomata is a more suit-
able term for the fruiting bodies of Micropeltidaceae rather
than thyriothecia. The dominant bacterial community,
Fig. 27 Polypedia
leopoldinensis (URM 14047)
aThe specimen. b, c Colonies
on leaf surface, intimately
mixed with conidiomata. d, e
Squash mount of ascoma and
conidiomata. fCross section of
conidiomata. g–l Conidia. Scale
bars c= 200 lm, d= 100 lm,
e, f =50lm, g–l =5lm
Fungal Diversity
123
which inhabited the surface of Micropeltis asiatica on
Semecarpus sp., are Cyanobacteria. This may explain the
cause of blue-green coloured upper walls in this family and
gives a clue as to their nutritional mode. Most relatives in
nearby taxa of this class are lichenised (Wedin et al. 2005b;
Miadlikowska et al. 2006,2014). Identification of these
symbionts provides a foundation for future studies on
determining roles of different symbionts.
In this study, we accept eight genera with hyaline
hyphae and bluish or greenish upper walls to be the
members of Micropeltidaceae, and accommodate them in
the new order Micropeltidales. Of the remaining genera,
we transfer one genus to Asterinales, one to Meliolales,
three to Capnodiales, two to Dothideomycetes incertae
sedis, three to Microthyriales, and treat nine as doubtful
genera. We also introduce five new species based on
morphology and the sequence data.
Phylogenetically, the modern Micropeltidaceae is a
sister clade of Graphidales, Gyalectales, Ostropales and
Thellenellales in Lecanoromycetes, and morphologically
differs in having flattened ascomata rather than cup-like
structures (Wedin et al. 2005a). In our dataset, we select
strains to represent orders of Lecanoromycetes by follow-
ing several previous studies (Wedin et al. 2005b; Miad-
likowska et al. 2006,2014; Hofstetter et al. 2007; Pino-
Bodas et al. 2017; Kraichak et al. 2018). The placement of
Cyanodermella is doubtful, as the genus does not cluster
with members of Ostropales, but is sister to the Micro-
peltidales clade. The sexual morph of Cyanodermella has
not been described, and we could not compare their mor-
phological features with Micropeltidaceae species. There-
fore, we do not include Cyanodermella in Micropeltidales.
Although the sample size for the cophylogenetic study is
relatively small, the congruent divergence times and
topologies between genera of Micropeltidaceae and host
plants indicate convincing coevolutionary events during
the Cretaceous period when the diversity of Angiospermae
has massively increased (130–80 Mya) (Clarke et al. 2011;
Magallo
´n et al. 2015). This result indicates that the diver-
sification of Angiospermae might foster the formation of
genera of Micropeltidaceae, and it would be an important
reference for establishing the generic boundaries of epifo-
liar fungi. Coevolutionary dynamics of hosts and sym-
bionts probably do not favour long-term cospeciation,
while host shifts appear to be more common in shaping the
diversity of symbiotic fungal species (de Vienne et al.
2013; Millanes et al. 2014; McTaggart et al. 2016; Navaud
et al. 2018). After the diversification period of
Angiospermae, there are no significant coevolutionary
events between Micropeltidaceae species and their host
plants, indicating an independent evolutionary tendency of
Microepltidaceae. We suspect that epifoliar fungi may
have different strategies when co-inhabiting with plants
during different periods of time.
In addition, we noticed that the clade representing
Arthoniomycetes clusters inside Dothideomycetes, and this
is congruent to results reported by Ertz and Diederich
(2015). Asterinales, Cladoriellales, Eremithallales, Jahnu-
lales, Lichenoconiales, Microthyriales, Natipusillales,
Phaeotrichales, Venturiales and Zeloasperisporiales are
distinct from the major Dothideomycetes group in our
phylogenetic analyses. These orders have low bootstrap
support with their sister clades in previous systematic
studies (Schoch et al. 2009; Hyde et al. 2013), and both of
the studies did not include Arthoniomycetes data in their
phylogenetic trees. On the other hand, the Sordariomycetes
clade is sister to Dothideomycetes in our phylogenetic
analyses supported by 1.00 PP but low bootstrap, while it
usually cluster with Leotiomycetes in other phylogenetic
studies. This difference might result from the inclusion of
the above orders and the different length of the RPB2 gene
used in the analyses. More evidences are needed to clarify
the placement of Arthoniomycetes and the above orders.
Acknowledgement This study is supported by the Fundamental
Research Funds for the Central Non-profit Research Institution of
CAF (grant no. CAFYBB2019QB005), the National Natural Science
Foundation of China (grant no. 31300019), the Science and Tech-
nology Foundation of Guizhou Province (No. [2017]5788), the
Impact of climate change on fungal diversity and biogeography in the
Greater Mekong Subregion (RDG6130001) and The future of spe-
cialist fungi in a changing climate: baseline data for generalist and
specialist fungi associated with ants, Rhododendron species and
Dracaena species (grant no: DBG6080013). Dr. R Jeewon thanks the
University of Mauritius for research support.
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