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Checklist of Laboulbeniomycetes from Belgium and the Netherlands Checklist of thallus-forming Laboulbeniomycetes from Belgium and the Netherlands, including Hesperomyces halyziae and Laboulbenia quarantenae spp. nov.

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In this paper we present an updated checklist of thallus-forming Laboulbeniomycetes (Ascomycota, Pezi-zomycotina), that is, the orders Herpomycetales and Laboulbeniales, from Belgium and the Netherlands. Two species are newly described based on morphology, molecular data (ITS, LSU ribosomal DNA) and ecology (host association). These are Hesperomyces halyziae on Halyzia sedecimguttata (Coleoptera, Coc-cinellidae) from both countries and Laboulbenia quarantenae on Bembidion biguttatum (Coleoptera, Car-abidae) from Belgium. In addition, nine new country records are presented. For Belgium: Laboulbenia aubryi on Amara aranea (Coleoptera, Carabidae) and Rhachomyces spinosus on Syntomus foveatus (Co-leoptera, Carabidae). For the Netherlands: Chitonomyces melanurus on Laccophilus minutus (Coleoptera, Dytiscidae), Euphoriomyces agathidii on Agathidium laevigatum (Coleoptera, Leiodidae), Laboulbenia fas-ciculata on Omophron limbatum (Coleoptera, Carabidae), Laboulbenia metableti on Syntomus foveatus and S. truncatellus (Coleoptera, Carabidae), Laboulbenia pseudomasei on Pterostichus melanarius (Coleoptera, Carabidae), Rhachomyces canariensis on Trechus obtusus (Coleoptera, Carabidae), and Stigmatomyces hydrel-liae on Hydrellia albilabris (Diptera, Ephydridae). Finally, an identification key to 140 species of thallus-forming Laboulbeniomycetes in Belgium and the Netherlands is provided. Based on the combined data, we are able to identify mutual gaps that need to be filled as well as weigh the impact of chosen strategies (fieldwork, museum collections) and techniques in these neighboring countries. The aim of this work is to serve as a reference for studying Laboulbeniomycetes fungi in Europe.
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Checklist of Laboulbeniomycetes from Belgium and the Netherlands 23
Checklist of thallus-forming Laboulbeniomycetes from
Belgium and the Netherlands, including Hesperomyces
halyziae and Laboulbenia quarantenae spp. nov.
Danny Haelewaters1,2,3, André De Kesel4
1 Purdue University, West Lafayette, Indiana, United States of America 2 University of South Bohemia, České
Budějovice, Czech Republic 3 Ghent University, Ghent, Belgium 4 Meise Botanic Garden, Meise, Belgium
Corresponding author: Danny Haelewaters (danny.haelewaters@gmail.com)
Academic editor: Cecile Gueidan|Received19 May 2020|Accepted 7 July 2020 | Published 30 July2020
Citation: Haelewaters D, De Kesel A (2020) Checklist of thallus-forming Laboulbeniomycetes from Belgium and
the Netherlands, including Hesperomyces halyziae and Laboulbenia quarantenae spp. nov.. MycoKeys 71: 23–86.
https://doi.org/10.3897/mycokeys.71.53421
Abstract
In this paper we present an updated checklist of thallus-forming Laboulbeniomycetes (Ascomycota, Pezi-
zomycotina), that is, the orders Herpomycetales and Laboulbeniales, from Belgium and the Netherlands.
Two species are newly described based on morphology, molecular data (ITS, LSU ribosomal DNA) and
ecology (host association). ese are Hesperomyces halyziae on Halyzia sedecimguttata (Coleoptera, Coc-
cinellidae) from both countries and Laboulbenia quarantenae on Bembidion biguttatum (Coleoptera, Car-
abidae) from Belgium. In addition, nine new country records are presented. For Belgium: Laboulbenia
aubryi on Amara aranea (Coleoptera, Carabidae) and Rhachomyces spinosus on Syntomus foveatus (Co-
leoptera, Carabidae). For the Netherlands: Chitonomyces melanurus on Laccophilus minutus (Coleoptera,
Dytiscidae), Euphoriomyces agathidii on Agathidium laevigatum (Coleoptera, Leiodidae), Laboulbenia fas-
ciculata on Omophron limbatum (Coleoptera, Carabidae), Laboulbenia metableti on Syntomus foveatus and
S. truncatellus (Coleoptera, Carabidae), Laboulbenia pseudomasei on Pterostichus melanarius (Coleoptera,
Carabidae), Rhachomyces canariensis on Trechus obtusus (Coleoptera, Carabidae), and Stigmatomyces hydrel-
liae on Hydrellia albilabris (Diptera, Ephydridae). Finally, an identication key to 140 species of thallus-
forming Laboulbeniomycetes in Belgium and the Netherlands is provided. Based on the combined data,
we are able to identify mutual gaps that need to be lled as well as weigh the impact of chosen strategies
(eldwork, museum collections) and techniques in these neighboring countries. e aim of this work is to
serve as a reference for studying Laboulbeniomycetes fungi in Europe.
Keywords
2 new taxa, arthropod-associated fungi, Ascomycota, Herpomycetales, integrative taxonomy, key, La-
boulbeniales
Copyright D. Haelewaters, A. De Kesel. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
MycoKeys 71: 23–86 (2020)
doi: 10.3897/mycokeys.71.53421
http://mycokeys.pensoft.net
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Launched to accelerate biodiversity research
RESEARCH ARTICLE
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
24
Introduction
Herpomycetales and Laboulbeniales are two orders within the class Laboulbeniomycet-
es (Ascomycota, Pezizomycotina), consisting of arthropod-associated biotrophs. Both
orders are unique among related fungi in that they do not form hyphae; instead, thalli
are produced by mitotic divisions from a two-celled ascospore. Herpomycetales was
recently described and includes a single genus, Herpomyces axt., with 27 described
species–all associated with cockroaches (Blattodea) (Haelewaters et al. 2019b; Gutier-
rez et al. 2020). e Laboulbeniales order, on the other hand, successfully radiated on
a wide range of hosts. Representatives of this order can be found in three arthropod
subphyla, including mites and harvestmen (in subphylum Chelicerata), millipedes (in
subphylum Myriapoda), and many orders of true insects (in subphylum Hexapoda).
e vast majority of about 2,325 described species (Kirk 2019) are known from beetles
(order Coleoptera), hence the common name once introduced for the group, “beetle
hangers” (Cooke 1892). e early taxonomic history of these fungi is fraught with
confusion (Blackwell et al. 2020), but the incorporation of sequence data has led to
a conclusive placement of these fungi within Ascomycota (Blackwell 1994; Weir and
Blackwell 2001; Schoch et al. 2009).
Early studies on Laboulbeniales (including Herpomyces at that time) in Belgium
and the Netherlands are scarce. In Belgium, Collart (1945, 1947) and Rammeloo
(1986) made noteworthy contributions, followed by multiple publications by De Ke-
sel and colleagues (1989–present). e Laboulbeniomycetes from Belgium were for
the rst time summarized by De Kesel and Rammeloo (1992), who reported 1 species
of Herpomyces and 47 species of Laboulbeniales. De Kesel et al. (2020) provided an up-
dated – and illustrated – Catalogue of the Laboulbeniomycetes of Belgium, with a total of
115 species (3 Herpomycetales, 112 Laboulbeniales) from 222 host species. For more
details regarding the study of Herpomycetales and Laboulbeniales in Belgium, we refer
to De Kesel and Rammeloo (1992) and De Kesel et al. (2020). In the Netherlands,
thus far, no eort has been made to publish a checklist.
e study of Laboulbeniales in the Netherlands started during a meeting of the
Dutch Entomological Society in 1906, triggered by a question from Dr. Johannes P.
Lotsy, then director of the “Rijksherbarium” (Leiden). In response, Prof. Dr. De Meijere
remembered that he once observed an infected Drosophila funebris (Fabricius, 1787) y,
collected at the ARTIS Amsterdam Royal Zoo in 1904, but had not thought it worthy
of mention at the time. Recent infected material of D. funebris from nature reserve De
Kaaistoep has thus far always been associated with Stigmatomyces entomophilus (Peck)
axt. (Haelewaters et al. 2015b) and hence it is likely that S. entomophilus represents the
very rst report of Laboulbeniales from the Netherlands. e rst published account was
a developmental study of Stigmatomyces baeri H. Karst. by Boedijn (1923). e fungus
was found on an atypical host – Fannia canicularis (Linnaeus, 1761); this y is the only
reported host for Fanniomyces ceratophorus (Whisler) T. Majewski, which is morphologi-
cally dierent from Boedijns (1923) drawings. We agree with axter (1931) that the
fungus was probably correctly identied by Boedijn, but perhaps the host was not.
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 25
Next, in the 1930s, only two species of Laboulbeniales were reported in the Neth-
erlands: Laboulbenia cristata axt. from Paederus riparius (Linnaeus, 1758) (Kossen
1936, 1938) and Laboulbenia agellata Peyr. from Platynus spp. (Zaneveld 1938). It
was not until Abraham Middelhoek (1906–1968) that the number of reported species
of Laboulbeniales in the Netherlands would increase by 25 (Middelhoek 1941, 1942,
1943a, b, c, d, 1945, 1947a, b, 1949). Middelhoek was rst an artist who, among
other things, made stained glass windows. Only after World War II, he studied biol-
ogy and raised an interest in fungi, particularly the Laboulbeniales. After Middelhoek,
Laboulbeniales were forgotten about in the Netherlands except for a single paper by
Meijer (1975), who proposed to use Laboulbeniales fungi as “biological tags” to trace
migration patterns. Since 2012, Haelewaters and colleagues have published several
papers dealing with Laboulbeniales in the Netherlands, which together have more than
doubled the number of reported species in this country (De Kesel and Gerstmans
2012; Haelewaters 2012, 2013; Haelewaters et al. 2012a, b, 2014, 2015a, b, 2020; De
Kesel et al. 2013; Haelewaters and De Kesel 2013; De Kesel and Haelewaters 2014,
2019; Haelewaters and van Wielink 2016). To date, 79 species of Laboulbeniales are
reported from the Netherlands.
In this contribution we compile all available data from Belgium and the Neth-
erlands. Keeping in mind that both countries show some geographical dierences,
especially due to specic soils and increasing altitude in the southern part of Belgium,
we think a combined checklist makes sense at this point. is is mainly because the
sampling eort for Laboulbeniomycetes in the southern part of Belgium has been
much lower compared to the northern and central areas of the country (De Kesel et al.
2020). As a result, the bulk of Belgian and Dutch records come from biogeographically
comparable regions. e here presented checklist is useful to illustrate where mutual
gaps need to be lled and what the impact has been of the chosen strategies (eldwork,
museum collections) and trapping techniques. In combination with the recently pub-
lished Belgian catalogue (De Kesel et al. 2020) presenting illustrations and identica-
tion keys to 115 taxa, this checklist will serve as a reference for mycologists, students,
and scholars studying Laboulbeniomycetes fungi. In addition, this work is an appro-
priate starting point for an updated checklist of thallus-forming Laboulbeniomycetes
from Europe–an ongoing project that needs to be updated, three decades after the
massive undertaking of Santamaría et al. (1991).
Materials and methods
Specimen collection and morphological study
Insects were collected in Belgium and the Netherlands using pitfall traps and on an
illuminated white screen at night. Specimens were preserved in 96–99% ethanol until
they were screened for presence of thalli of Laboulbeniomycetes at 20–50× magnica-
tion. alli were removed from the host at the foot and mounted in Amann solution
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
26
following the methods in De Kesel et al. (2020). Drawings and measurements were
made using a BX51 light microscope (Olympus, Tokyo, Japan) with drawing tube,
digital camera, and AnalySIS software (Soft Imaging System GmbH, Münster, Ger-
many); or an an Olympus BH2 bright eld compound microscope with SC30 camera
and cellSens 1.18 imaging software.
Infected hosts found in Belgium and the Netherlands are preserved at Meise Bo-
tanic Garden (BR) and the Brabant Museum of Nature, Tilburg (NNKN), respective-
ly. Microscope slides of Laboulbeniales are deposited at BR, FH, GENT, and NMBT
(iers continuously updated).
DNA extraction, PCR amplification, sequencing
ree thalli of Laboulbenia quarantenae sp. nov. were used for DNA isolation using the
REPLI-g Single Cell Kit (Qiagen, Stanford, California) with modications (Haele-
waters et al. 2019b). e DNA extract was stored at -20 °C until PCR amplication.
Recent studies found that even though the internal transcribed spacer (ITS) region is a
good marker for species delimitation in Laboulbeniomycetes, it is dicult to amplify
in this group. Instead, the large subunit (LSU) of the ribosomal RNA gene has been
put forward as a secondary barcode because it is easy to amplify and provides high
discriminative resolution at species-level (e.g., Haelewaters et al. 2018; Sundberg et
al. 2018b; Walker et al. 2018; Liu et al. 2020). e partial LSU was amplied using
primers LIC15R (Miadlikowska et al. 2002) and LR6 (Vilgalys and Hester 1990). Se-
quencing was outsourced to Macrogen Europe (Amsterdam, the Netherlands) with the
same PCR primers and an additional reverse primer, LR3 (Vilgalys and Hester 1990).
Resulting forward and both reverse sequence reads were assembled and edited with
Sequencher version 5.2.3 (Gene Codes Corporation, Ann Arbor, Michigan).
For Hesperomyces halyziae, molecular work had been done previously (Haele-
waters et al. 2018). DNA was extracted using the Extract-N-Amp Plant PCR Kit
(Sigma-Aldrich, St. Louis, Missouri) (methods in Haelewaters et al. 2015c). Seven
thalli were placed in a 1.5 mL tube with 40 µL of Extraction Solution and steri-
lized sand. e tube was then placed in a FastPrep FP120 Cell Disrupter (ermo
Fisher Scientic, Waltham, Massachusetts) to mechanically crush fungal material
at 5.5m/s for 20 sec, and then on a heating block to incubate at 95 °C for 10 min.
Finally, a total of 120 µL Dilution Solution was added to the mixture. Because
we needed to dene “H. virescens sensu stricto”, additional extractions from single
Hesperomyces thalli removed from Chilocorus stigma (Say, 1835) were performed us-
ing the REPLI-g Single Cell Kit with modications. Amplication of the ITS was
done using primers ITS1f (Gardes and Bruns 1993) and ITS4 (White et al. 1990)
as well as Hesperomyces-specic primers ITShespL and ITShespR (Haelewaters et al.
2019b). Purication and sequencing (same primers) of these PCR products were
outsourced to Genewiz (Plaineld, New Jersey).
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 27
Phylogenetic analyses
Methods for both datasets – ITS for Hesperomyces, LSU for Laboulbenia – were largely
identical. Sequences were downloaded from NCBI GenBank (https://www.ncbi.nlm.
nih.gov/genbank/) and supplemented with sequences that were generated during this
study. Sequences were aligned using MUSCLE version 3.7 (Edgar 2004), which is
available on the CIPRES Science Gateway V. 3.3 (Miller et al. 2010). After alignment
of the ITS dataset, partial SSU and partial LSU were removed by looking for the motifs
5’-ATCATTA-3’ (3’ end of SSU) and 5’-TGACCT-3’ (5’ start of LSU), and deleting
downstream and upstream sequence data, respectively (Baral et al. 2018). For the LSU
dataset, we unsuccessfully searched for the 5’-TGACCT-3’ motif. We then looked for
the motif following 5’-TGACCT-3’ in a Hesperomyces sequence (GenBank acc. no.
MG757513), which is 5’-CGGAT-3’, found this motif in the Laboulbenia dataset, and
then realized that the 5’ start of LSU in Laboulbenia includes one nucleotide substi-
tution compared to the conventional motif: 5’-TGGCCT-3’. We deleted the down-
stream sequence data to remove partial ITS. Next, ambiguously aligned regions and
uninformative positions were removed using the command line version of trimAl v1.2
(Capella-Gutiérrez et al. 2009) with gap threshold = 0.6 and minimal coverage = 0.5.
Models of nucleotide substitution were selected by considering the Akaike Informa-
tion Criterion corrected for small samples (AICc) with ModelFinder Plus (Kalyaana-
moorthy et al. 2017). Maximum likelihood (ML) was inferred for each dataset under
the selected model with IQ-TREE (Nguyen et al. 2015; Chernomor et al. 2016).
Ultrafast bootstrap (BS) analysis with 1000 replicates estimated branch support in the
ML trees (Hoang et al. 2018).
Bayesian analyses were done using a Markov chain Monte Carlo (MCMC) coales-
cent approach implemented in BEAST 1.8.4 (Drummond et al. 2012), with a strict
clock assuming a constant rate of evolution across the tree, a Yule Speciation tree prior
(Yule 1925; Gernhard 2008), and the nucleotide substitution model as selected by
jModelTest 2.1 (Darriba et al. 2012) under the AICc criterion. For each dataset, four
runs were performed from a random starting tree for 10 million generations with a
sampling frequency of 1000. All settings were entered in BEAUti 1.8.4 to generate an
XML le, which was run in BEAST on the CIPRES Science Gateway (Miller et al.
2010). Resulting log les were entered in Tracer version 1.6 (Rambaut et al. 2014) to
check MCMC trace plots for convergence and to assess eective sample sizes (ESS). A
standard 10% burn-in was used resulting in overall ESS values of well above 200 for
all sampled parameters. After removal of 10% burn-in, trees les were combined in
LogCombiner 1.8.4. TreeAnnotator 1.8.4 was used to generate consensus trees with
0% burn-in and to infer the Maximum Clade Credibility tree with highest product of
individual clade posterior probabilities (pp) for both datasets.
Trees with ML BS and Bayesian pp were visualized in FigTree version 1.4.3 (http://
tree.bio.ed.ac.uk/software/gtree/) and edited in Adobe Illustrator 2020 version 24.1.1
(San Jose, California).
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
28
Checklist
For the checklist of thallus-forming Laboulbeniomycetes from Belgium and the Neth-
erlands, we used De Kesel et al. (2020) for Belgium and all available published papers
(since 1938 up to 2020) for the Netherlands. Laboulbeniomycetes and their hosts
are listed alphabetically, starting with Herpomycetales, followed by Laboulbeniales.
Fungal species are numbered throughout (1–140), authority and reference to the pro-
tologue are presented. For each fungus, hosts are presented alphabetically, with classi-
cation (order, family) and country in which the association has been reported: “Be” for
Belgium, “Nl” for the Netherlands. No detailed collection information is shown except
for new country records. In several instances, taxonomic notes are provided. Hosts are
according to Vorst (2010) and Beccaloni et al. (2014). Names of fungi correspond to
Index Fungorum (2020).
Identification key
e key to species of Laboulbeniomycetes in Belgium and the Netherlands is based on
diagnostic characters referring to morphology and/or host taxa. It requires microscope
equipment and morphological study as described in Benjamin (1971), Huldén (1983),
Majewski (1994), Santamaría (1998), and De Kesel et al. (2020). Terminology follows
Tavares (1985), Santamaría (1998, 2003), and De Kesel et al. (2020).
Results
e ITS dataset consisted of 31 Hesperomyces sequences (Table 1) and 724 characters,
of which 462 were constant and 198 were parsimony-informative. e selected nu-
cleotide substitution model under AICc was TVM+F+G4 (-lnL = 2790.545, ModelF-
inder Plus) and TVM+G (-lnL = 2786.8769, jModelTest 2). e Hesperomyces virescens
sensu lato (Haelewaters et al. 2018) clade has maximum support from both ML and
Bayesian analyses (Figure 1). Each of the nine clades within H. virescens s.l. consists
of isolates from thalli removed from a single host species, except for the Adalia clade,
which includes isolates from both A. bipunctata and A. decempunctata. One of the
clades consists of isolates from Chilocorus stigma, the host on which H. virescens was
originally described (axter 1891). is clade, representative of Hesperomyces virescens
sensu stricto, receives maximum support. e single isolate of Hesperomyces halyziae,
from Halyzia sedecimguttata, is placed as sister to H. virescens s.l. from Harmonia axy-
ridis (Pallas, 1773) (pp = 0.8).
e LSU dataset consisted of 24 Laboulbenia sequences (Table 2) and 682 char-
acters, of which 558 were constant and 63 were parsimony-informative. e selected
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 29
Table 1. Hesperomyces sequences used in phylogenetic analysis of the ITS dataset. Asterisks (*) indicate
sequences that were generated during the course of this study.
Species Host Isolate GenBank (ITS) Reference
Hesperomyces coleomegillae Coleomegilla maculata 632A KF192888 Goldmann et al. (2013)
Coleomegilla maculate 635D KF192906 Goldmann et al. (2013)
Hesperomyces halyziae Halyzia sedecimguttata D. Haelew. 955b MG757813 Haelewaters et al. (2018)
Hesperomyces virescens s.s. Chilocorus stigma D. Haelew. 1444a MT373697* is paper
Chilocorus stigma D. Haelew. 1444b MT373698* is paper
Hesperomyces virescens s.l. Adalia bipunctata D. Haelew. 1193g MG757817 Haelewaters et al. (2018)
Adalia bipunctata D. Haelew. 1231a MG757821 Haelewaters et al. (2018)
Adalia bipunctata D. Haelew. 1232a MG757822 Haelewaters et al. (2018)
Adalia decempunctata D. Haelew. 1248b MG757823 Haelewaters et al. (2018)
Azya orbigera D. Haelew. 928g MG745343 Haelewaters et al. (2018)
Cheilomenes propinqua D. Haelew. 655c MG757804 Haelewaters et al. (2018)
Cheilomenes propinqua D. Haelew. 659b MG757805 Haelewaters et al. (2018)
Cheilomenes propinqua D. Haelew. 1259a MG757828 Haelewaters et al. (2018)
Cycloneda sanguinea D. Haelew. 924a MG757808 Haelewaters et al. (2018)
Cycloneda sanguinea D. Haelew. 1374a MG757831 Haelewaters et al. (2018)
Harmonia axyridis 352B KF192916 Goldmann et al. (2013)
Harmonia axyridis D. Haelew. 361a MG757801 Haelewaters et al. (2018)
Harmonia axyridis D. Haelew. 486c KT800044 Haelewaters et al. (2015c)
Harmonia axyridis D. Haelew. 669a MG757807 Haelewaters et al. (2018)
Harmonia axyridis D. Haelew. 1188g MG438317 Haelewaters et al. (2019b)
Harmonia axyridis D. Haelew. 1268d MG757830 Haelewaters et al. (2018)
Harmonia axyridis DH1 KF192920 Goldmann et al. (2013)
Harmonia axyridis LT1 KF192910 Goldmann et al. (2013)
Harmonia axyridis MT001 KT800048 Haelewaters et al. (2015c)
Olla v-nigrum D. Haelew. 954e MG757812 Haelewaters et al. (2018)
Olla v-nigrum D. Haelew. 1200h MG757819 Haelewaters et al. (2018)
Olla v-nigrum JP353b MG757799 Haelewaters et al. (2018)
Olla v-nigrum JP354b MG757800 Haelewaters et al. (2018)
Psyllobora vigintimaculata D. Haelew. 1250b MG757825 Haelewaters et al. (2018)
Psyllobora vigintimaculata D. Haelew. 1250c MG757826 Haelewaters et al. (2018)
Psyllobora vigintimaculata D. Haelew. 1251b MG757827 Haelewaters et al. (2018)
nucleotide substitution model under AICc was TN+F+G4 (-lnL = 1876.681, ModelF-
inder Plus) and TrN+G (-lnL = 1872.4616, jModelTest 2). Our phylogenetic analyses
show nine distinct species, which are all supported. e relationships among species
are unresolved in dierent places, but this is not unsurprising because of extremely lim-
ited taxon sampling. Laboulbenia quarantenae holds an unresolved position in the tree
but is clearly separated from both L. agellata and the morphologically similar L.vul-
garis, conrming its status as a separate species. Laboulbenia vulgaris isolates E10T2
and E11T6, which originated from Bembidion tetracolum, are placed among isolates of
the same species removed from Ocys harpaloides. Interestingly, and in accordance with
De Weggheleire (2019) and Haelewaters et al. (2019a), L. agellata falls apart in three
species. However, only ten isolates are included, originating from six host species, none
of which were reported in the protologue (Peyritsch 1873). As a result, it is too early to
make taxonomic decisions within this problematic taxon.
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
30
Figure 1. Maximum clade creditability tree of Hesperomyces isolates reconstructed from an ITS dataset,
with H. coleomegillae as outgroup. e topology is the result of Bayesian inference performed with BEAST.
For each node, ML BS (≥ 65) and Bayesian pp (≥ 0.7) are presented above/below the branch leading to
that node. Hesperomyces virescens sensu lato is highlighted with light gray shading, isolates are color-coded
by host; H. virescens sensu stricto and H. halyziae sp. nov. are highlighted with dark gray shading.
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 31
Taxonomy
Hesperomyces halyziae Haelew. & De Kesel, sp. nov.
MycoBank No: 835489
Figure 3
Etymology. Referring to the host genus, Halyzia.
Diagnosis. Morphologically very similar to other taxa within H. virescens sensu
lato, but forming a distinct species supported by ITS data. e ITS sequence shares
95.8–97.9% identity with H. virescens s.l. from Harmonia axyridis, and 96.5–95.4%
with H. virescens s.l. from Adalia bipunctata/A. decempunctata. Unique molecular syna-
pomorphies in the ITS at positions 478, 517, 652.
Types. Holotype: e Netherlands, Noord Brabant Province, Tilburg, nature reserve
De Kaaistoep, 51.5333333N, 5.0166667E, 11 Aug. 2015, leg. H. Spijkers & P. van
Wielink, on female Halyzia sedecimguttata (Linnaeus, 1758) (Coleoptera, Coccinellidae)
(NNKN), slide D. Haelew. 955a (FH, 4 juvenile and 3 mature thalli, left elytron),
reported as Hesperomyces virescens in Haelewaters and van Wielink (2016). Paratypes:
Belgium, Province Vlaams-Brabant, Meise, Domein van Bouchout, 50.927925N,
4.333069E, 28 Mar. 2019, leg. C. Gerstmans, on H. sedecimguttata (BR, CG437–
CG440), slides BR5020212155379V, BR5020212156406V, BR5020212157434V,
and BR5020212158462V; reported as Hesperomyces virescens sensu lato in De Kesel et
al. (2020). Ibid., 1 Apr. 2019, leg. C. Gerstmans, on H. sedecimguttata (BR, CG441–
Table 2. Laboulbenia sequences used in phylogenetic analysis of the LSU dataset. Asterisks (*) indicate
sequences that were generated during the course of this study.
Species Host Isolate GenBank (LSU) Reference
Laboulbenia bruchii Neolema adunata D. Haelew. 1346b MN394843 Haelewaters et al. (2019a)
Laboulbenia collae Paranchus albipes D. Haelew. 1456a MN394844 Haelewaters et al. (2019a)
Paranchus albipes D. Haelew. 1456b MN394845 Haelewaters et al. (2019a)
Paranchus albipes D. Haelew. 1461b MN397131 Haelewaters et al. (2019a)
Laboulbenia quarantenae Bembidion biguttatum, ADK6448 E13T12 MT371368* is paper
Laboulbenia agellata Agonum emarginatum, ADK6428 E13T1 MT703825* is paper
Agonum micans, ADK6332 D. Haelew. 1457a MN394851 Haelewaters et al. (2019a)
Agonum micans, ADK6332 D. Haelew. 1457b MN394852 Haelewaters et al. (2019a)
Agonum micans, ADK6332 D. Haelew. 1457c MN394853 Haelewaters et al. (2019a)
Agonum nigrum, ADK6445 E13T11 MT703826* is paper
Limodromus assimilis, ADK6329-1 D. Haelew. 1454a MN394849 Haelewaters et al. (2019a)
Limodromus assimilis, ADK6329-1 D. Haelew. 1454b MN394850 Haelewaters et al. (2019a)
Limodromus assimilis, ADK6329-2 D. Haelew. 1458a MN394854 Haelewaters et al. (2019a)
Loricera pilicornis H85-1 KY350538 Sundberg et al. (2018a)
Oxypselaphus obscurus, ADK6374 E11T11 MT703824* is paper
Laboulbenia pedicellata Dyschirius globosus H84-1 KY350537 Sundberg et al. (2018a)
Laboulbenia systenae Disonycha procera D. Haelew. 1342b MN394858 Haelewaters et al. (2019a)
Laboulbenia vulgaris Bembidion tetracolum, ADK6420 E10T2 MT703822* is paper
Bembidion tetracolum, ADK5557 E11E6 MT703823* is paper
Ocys harpaloides, ADK6330-1 D. Haelew. 1455a MN397135 Haelewaters et al. (2019a)
Ocys harpaloides, ADK6330-1 D. Haelew. 1455b MN397136 Haelewaters et al. (2019a)
Ocys harpaloides, ADK6330-2 D. Haelew. 1459a MN397137 Haelewaters et al. (2019a)
Ocys harpaloides, ADK6330-3 D. Haelew. 1460a MN397138 Haelewaters et al. (2019a)
Ocys harpaloides, ADK6353-1 E0T6 MT703821* is paper
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
32
442), slides BR5020212159490V and BR5020212160236V; reported as Hesperomyces
virescens sensu lato in De Kesel et al. (2020).
Description. allus 335–453 µm long from foot to perithecial apex; colored
yellow except for a somewhat darker region right above the foot. Cell I obtriangular,
2.0–2.5× longer than broad, broadening distally, with very oblique septum I–II.
Cell II longer than broad, 23–28 × 16–21 µm, subtrapezoidal in section. Cell III
always smaller than cell II, 14–20 × 14–19 µm, with inated dorsal cell wall. Pri-
mary appendage consisting of 4 superposed cells, 61–67 µm long; in the same axis
as cells I and III, separated from the latter by the constricted primary septum; its
basal cell somewhat longer than broad, longer than each of the remaining cells of the
appendage; second to fourth cells carrying a single antherium externally, the fourth
cell also carrying a second upwardly directed antherium. Antheridia ask-shaped,
Figure 2. Maximum clade creditability tree of Laboulbenia isolates reconstructed from an LSU dataset,
with L. bruchii as outgroup. e topology is the result of Bayesian inference performed with BEAST. For
each node, ML BS (≥ 65) and Bayesian pp (≥ 0.7) are presented above/below the branch leading to that
node. Isolates are color-coded by host; L. quarantenae sp. nov. is highlighted with gray shading.
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 33
Figure 3. Hesperomyces halyziae Haelew. & De Kesel from Halyzia sedecimguttata A mature thallus from
slide D. Haelew. 955a, holotype B mature thallus from slide BR5020212156406V. Scale bar: 100 µm.
with slightly (dorsally and/or basally) curved eerent necks, the upper antheridium
carrying at its dorsal side a pointed process, which represents the original ascospore
apex. Cell VI with subparallel margins to broadening distally, 33–70 × 23–33 µm.
Perithecium 194–291 × 62–86µm (not including basal cells), symmetric or with
the anterior margin convex and the posterior one almost straight or concave; broad-
est near the upper third, then gradually tapering towards the apex; apex complex
with 2 short lower lobes, 2 upper (terminal) lobes, and 2 prominent lips surround-
ing the ostiole; lower lobes tapering to a rounded tip, the ventral lobe outwardly
directed; terminal lobes unicellular, elongated, 29–42 µm in length, curved up-
wards and outwardly; ostiole with two lips, 25–29 µm in length, one lip triangular,
the other slightly shorter, blunt or rounded, basally carrying the remainder of the
trichogyne. Ascospores 70–85 µm long, with conspicuous slime sheath only sur-
rounding the larger cell.
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
34
Material sequenced. e Netherlands, Noord Brabant Province, Tilburg, nature
reserve De Kaaistoep, 51.5333333N, 5.0166667E, 11 Aug. 2015, leg. H. Spijkers & P.
van Wielink, on female Halyzia sedecimguttata (Coleoptera, Coccinellidae) (NNKN),
isolate D. Haelew. 955b (7 thalli, elytra, ITS: MG757813).
Hosts and distribution. On Halyzia sedecimguttata from Belgium and the Neth-
erlands. Previously reported as H. virescens (Haelewaters and van Wielink 2016, Haele-
waters et al. 2017) and H. virescens sensu lato (De Kesel et al. 2020). One unveried
record is available from France (Justamond 2019).
Notes. Supported by multi-locus phylogenetic analyses and sequence-based species
delimitation methods, Haelewaters et al. (2018) showed that H. virescens axt. is a com-
plex of multiple species, segregated by host. e authors proposed to “restrict H. virescens
sensu stricto to those thalli found on Chilocorus stigma, the host species on which the
fungus was originally described” (axter 1891). Here, we included two isolates from C.
stigma (Say, 1835), and found the clade representative of H. virescens sensu stricto. Based
on this analysis and previous work (Haelewaters et al. 2018), we can start describing the
individual clades as distinct species. A monographic work with formal descriptions for
the seven other species within H. virescens s.l. is in preparation, but in the light of this
checklist we decided to describe H. halyziae, which was only known from a single col-
lection in the Netherlands until we recently collected it in Belgium (Mar.–Apr. 2019).
Haelewaters and van Wielink (2016) reported an infected specimen of Halyzia se-
decimguttata from nature reserve De Kaaistoep in the Netherlands. In 1997–2015, 476
individuals of H. sedecimguttata were collected on a lighted white sheet and screened
for presence of Laboulbeniales, only resulting in one individual (parasite prevalence
0.2%). In Belgium, a population of infected H. sedecimguttata was found at the Meise
Botanic Garden. Specimens were collected in spring 2019 while they were leaving their
overwintering place–deep cracks in the woodwork of a small forest chapel. Screening
of 46 specimens of H. sedecimguttata revealed nine infected ones (parasite prevalence
19.5%). is ladybird species seems to overwinter singly or in small congregations
in narrow overwintering places, including in leaf litter, under foliage on stone walls,
on trunks and branches (Majerus and Williams 1989). is congregation behavior
is benecial for transmission of the fungus and is also observed in Harmonia axyridis
(Haelewaters et al. 2017).
Morphologically, H. halyziae is very similar to what we have thus far accepted as
H. virescens. Within the Kingdom Fungi, there is an incredible diversity that cannot be
perceived through morphology. Cryptic species are being uncovered in Agaricomycetes
(e.g., Stefani et al. 2014; Sánchez-García et al. 2016), Lecanoromycetes (e.g., Singh
et al. 2015), Leotiomyces (e.g., Grünig et al. 2008), Pucciniomycetes (Bennett et al.
2011), Ustilaginomycetes (e.g., Li et al. 2017), and other major clades. And while the
Laboulbeniales has been the subject of a large-scale study to estimate the global species
richness of the group (Weir and Hammond 1997), cryptic diversity was not part of the
equation. In other words, the number of estimated species of Laboulbeniales, between
15,000 and 75,000, is likely to be corrected to include cryptic species. We note that
the recognition of H. halyziae is only possible through molecular data and host associa-
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 35
tion. Our current understanding is that, within this species complex, there is a strict
parasite-host association, with one parasite found only on one host. We think that
this host specicity exists at the genus level, given the Adalia clade (Figure 1), which
includes isolates from thalli removed from two host species within the same genus.
Laboulbenia quarantenae De Kesel & Haelew., sp. nov.
MycoBank No: 835490
Figure 4
Diagnosis. Morphologically similar to Laboulbenia vulgaris Peyr., but the insertion
cell is attached to the lower fth of the posterior margin of the perithecial wall and the
outer appendage is composed of 4–6(–8) branches resulting from successive dichoto-
mies starting at the suprabasal cell, which is poorly pigmented or nearly hyaline. e
LSU sequence shares 89.7–98.0% identity with other sequenced taxa of Laboulbenia,
97.4% with L. agellata from Agonum nigrum, 97.5–98.0% with L. agellata from
Limodromus assimilis, 97.0–98.0% with L. agellata from Agonum emarginatum/A. mi-
cans/Loricera pilicornis/Oxypselaphus obscurus, and 97.0–97.7% with L. vulgaris from
Bembidion tetracolum/Ocys harpaloides. Unique molecular synapomorphies in the LSU
at positions 503, 545.
Types. Holotype: Belgium, Province Vlaams Brabant, Meise, Domein van Bou-
chout, 50.9267056N, 4.3220028E, 30 m a.s.l., 26 Apr. 2019, leg. A. De Kesel, rivu-
let-associated grassland, on Bembidion (Philochtus) biguttatum (Fabricius, 1779) (Co-
leoptera, Carabidae), ADK6448 (BR), slide BR5020212163329V (1 mature thallus,
prothorax). I: ibid., slides BR5020212162292V (2 mature thalli, right mesofe-
mur), BR5020212161264V (6 mature thalli, right protibia), BR5020212166412V (5
immature thalli, mesothorax), BR5020212165385V (1 mature thallus, right protibia),
and BR5020212164357V (1 mature thallus, right mesofemur). Paratype: Belgium,
Province Vlaams-Brabant, Meise, Domein van Bouchout, 50.92745N, 4.323917E,
32m a.s.l., 30 Apr. 2020, leg. A. De Kesel, rivulet-associated grassland, on B. (P. ) bigut-
tatum, ADK6523 (BR), slide BR5020195033527V (2 mature thalli, mesosternum).
Etymology. From quarantena, which was used in 14th–15th century Venetian lan-
guage for a forty-day isolation period. e new species was described during the 2020
quarantine period imposed to curb the spread of the COVID-19 virus.
Description. allus 300–465 µm long from foot to perithecial tip; colored hya-
line at the lower receptacular cells (I and II) and the inner appendage, otherwise pig-
mented light to dark brown; especially the upper receptacular cells (III, IV and V),
cell VI, and the perithecium darkening with age. Cell I elongated, usually straight,
56–107× 22–33 µm; sometimes bent and then wider at the upper end. Cell II slen-
der, mostly with parallel margins, longer than cell I, 73–160 × 29–40 µm, anterior
margin shorter than posterior. Cells III and VI side by side, with septum II–III always
much shorter than septum II–VI. Cell III with a narrow base, 29–43 µm long, wid-
ening upwards and then 22–29 µm wide at the apex. Cell VI more or less rectangu-
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
36
Figure 4. A–I Laboulbenia quarantena De Kesel & Haelew. from Bembidion biguttatum, specimen
ADK6448: A mature thallus from prothorax, slide BR5020212163329V, holotype B mature thallus from
prothorax with less pigmented perithecium C mature thallus from the right mesofemur D–F mature thal-
li from the right protibia G immature thallus from the prothorax H mature thallus from the right mesofe-
mur I ascospores J–K laboulbenia vulgaris Peyr: J mature thallus from prothorax of Bembidion tetracolum,
specimen ADK5557 K mature thallus from mesothorax of Ocys harpaloides, specimen ADK6353. One of
the diagnostic characteristics of the new species–the positioning of the insertion cell–is shown in a mature
thallus of L. quarantenae (E) and one of L. vulgaris (J). Scale bar: 100 µm.
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 37
lar, 30–34× 23–30 µm. Cell IV more or less rectangular, slightly broader than long,
20–32 × 25–30µm. Cell V small, triangular, situated in the inner-upper corner of
cell IV, 9–14 × 7–14 µm, as pigmented as surrounding cells. Insertion cell brownish
black, attened, barely marking a constriction on the posterior margin of the thallus,
attached to the lower fth of the posterior margin of the perithecial wall, 18–25 µm
wide and 90–128 µm from the perithecial tip. Inner appendage hyaline, composed of
2–4(–6) short branches, rarely exceeding the perithecial tip, 88–150 µm long, result-
ing from successive dichotomies starting at the basal cell, the latter 9–14 × 6–12 µm.
Antheridia short, ask-shaped, few in number, usually on the young inner appendage
and arising laterally from its suprabasal cell. Outer appendage up to 250–335 µm
long, extending beyond the perithecial tip, often entirely light brown, composed of
4–6(–8) branches, resulting from successive dichotomies starting at the suprabasal cell;
the basal cell longer than broad, 23–32 × 15–21 µm, almost entirely hyaline. Perithe-
cium ellipsoid, venter only very slightly asymmetrical, anterior and posterior margins
almost equally convex, 109–157 × 43–64 µm, length/width ratio 1.9–2.5, widest in
the middle; perithecial tip asymmetrical, with prominent and rounded posterior mar-
gin; preostiolar spots black, in older thalli merging into a pre-apical ring, always with
distinctly paler zone under the posterior spot. Ascospores two-celled, hyaline, 59–65 ×
4.2–5.5 µm, with slime sheath.
Material sequenced. Belgium, Province Vlaams Brabant, Meise, Domein van
Bouchout, 50.9267056N, 4.3220028E, 30 m a.s.l., 26 Apr. 2019, leg. A. De Ke-
sel, rivulet associated grassland, on Bembidion biguttatum (Coleoptera, Carabidae),
ADK6448 (BR), isolate E13T12 (3 mature thalli, prothorax, LSU: MT371368).
Hosts and distribution. us far only known on Bembidion biguttatum from the
type locality in Belgium. Reported as Laboulbenia sp. nov. in De Weggheleire (2019).
Notes. Morphologically, L. quarantenae mostly resembles L. vulgaris Peyr., but it
diers from it by the very low position of the insertion cell (regardless of the origin
of the thallus), the successive dichotomous branching of the outer appendage, the
poorly pigmented to nearly hyaline basal cell of the outer appendage, and the slender
habitus. Although these characters may vary to some extent, eventually resulting in
specimens that are morphologically close to L. vulgaris, our LSU phylogeny (Figure 2)
shows that sequences of typical L. vulgaris obtained from Carabidae known to host L.
vulgarisBembidion tetracolum Say, 1823 and Ocys harpaloides (Audinet-Serville, 1821)
(Santamaría et al. 1991; Majewski 1994; Haelewaters et al. 2019a; De Kesel et al.
2020)–fall in a monophyletic clade separated from L. quarantenae. e two isolates of
L. vulgaris from B. tetracolum were collected in Belgium (isolate E10T2) and Latvia
(isolate E11T6), from populations that are 1,550 km apart, but they were placed together
among isolates from O. harpaloides (all from Belgium). Laboulbeniaquarantenae, on
the other hand, was collected between <1 and 21 km distance from where hosts of L.
vulgaris were collected.
Phylogenetically, L. quarantenae may be more closely related to L. agellata than
to L. vulgaris. Laboulb quarantenae and L. agellata (sensu lato) were retrieved as sister
taxa in our phylogeny, although no statistical support was retrieved for this sister rela-
tionship. Whereas species boundaries are evident based on our phylogeny, it goes with-
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
38
out saying that both taxon sampling and sequence data need to be greatly expanded
upon to resolve relationships among species of Laboulbenia. e new species is appar-
ently very rare and was never found in combination with L. vulgaris, the more common
parasite from Bembidion biguttatum in Belgium (De Kesel 1998; De Kesel et al. 2020).
In Europe, many species of Laboulbenia have been reported on Bembidion Latreille,
1802 (Santamaría et al. 1991). Of those, L. pedicellata axt. and L. vulgaris Peyr. are
among the most reported ones. Bembidion biguttatum belongs to subfamily Trechinae.
To our knowledge, this species is infected by either L. murmanica Huldén (S. San-
tamaría pers. comm.), L. pedicellata (Scheloske 1969; Majewski 1994), or L. vulgaris
(Majewski 1994; De Kesel et al. 2020). Based on the position of its insertion cell as well
as the morphology of both the outer appendage and the androstichum (cells II, IV, and
V), L. quarantenae is fundamentally dierent from these three species. e outer ap-
pendage of L. quarantenae is reminiscent of the one from L. agellata, which, however,
is a more robust species reported from 80 genera of Carabidae belonging to Anthiinae,
Brachininae, Elaphrinae, Harpalinae, Loricerinae, Nebriinae, and Patrobinae (but not
Trechinae) (Santamaría et al. 1991; Santamaría 1998; Haelewaters et al. 2019a).
Bembidion biguttatum, the host for L. quarantenae, belongs to the subgenus
Philochtus. Representatives of Laboulbenia reported from Bembidion subgenus Philoch-
tus are few and include two species only: L. pedicellata and L. vulgaris. Two thalli of
Laboulbenia “sp. similar to L. vulgaris” from Bembidion bruxellense Wesmael, 1835 [as
B. rupestre (Linnaeus, 1767) are illustrated in Majewski (1994: Pl. 53, Figs 1, 2). eir
morphology comes close to L. quarantenae but cell V is much larger and the insertion
cell is not situated low enough along the posterior margin of the perithecial wall. Also
L. parvula is reported on subgenus Philochtus in Santamaría et al. (1991), but this spe-
cies is much smaller (180–190 µm total length) compared to L. quarantenae, it has a
deeply pigmented basal cell of the outer appendage, the inner and outer appendage
each carry 4–8 very slender branches, and its perithecial tip is rather squarish.
As we explore patterns of speciation of taxa in both Herpomycetales and Laboul-
beniales using integrative taxonomy, we can start linking some of these patterns to
morphological or life history traits. One candidate trait is the haustorium–a rhizoidal
structure that penetrates the host’s integument to make contact with the haemocoel,
increasing surface area for nutrient uptake and providing holdfast. We hypothesize
that – due to the invasive nature of their haustorium – Herpomycetales and hausto-
rial Laboulbeniales, such as species of Hesperomyces, maintain close interactions with
their hosts, possibly involving adaptations to the hosts’ defense systems and leading to
escape-and-radiate coevolution (Ehrlich and Raven 1964). ese developments result
in an evolutionary arms race, with specialization and leading to speciation (One Host
One Parasite model, Figure 1). While all 27 species of Herpomyces form multiple haus-
toria, not all Laboulbeniales penetrate their host. Recently, Tragust et al. (2016) pre-
sented evidence for four species of Laboulbeniales to be supercially attached to their
host, and also L. agellata and L. vulgaris do not seem to perforate their hosts. ere
are no strict developmental barriers for non-penetrating species and their ascospores
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 39
may develop on multiple arthropods given that they co-occur in a given microhabitat,
resulting in parasite species with more than one host (e.g., L. vulgaris in Figure 2), in
contrast to the host-specic species of Hesperomyces. Undoubtedly, other factors come
into play; more studies of speciation and species limits, specicity, host shifting, and
transmission patterns are needed to test said hypothesis.
Alphabetical checklist of thallus-forming Laboulbeniomycetes in Belgium and the
Netherlands
Herpomycetales
1. Herpomyces ectobiae axt., Proc. Am. Acad. Arts Sci. 38(2): 20 (1902) [1903]
Blattella germanica (Linnaeus, 1767) (Blattodea, Ectobiidae) ........................... Be
2. Herpomyces periplanetae axt., Proc. Am. Acad. Arts Sci. 38(2): 13 (1902)[1903]
Blatta orientalis Linnaeus, 1758 (Blattodea, Blattidae) ..................................... Be
Periplaneta americana (Linnaeus, 1758) (Blattodea, Blattidae) ......................... Be
3. Herpomyces stylopygae Speg., Anal. Mus. Nac. Hist. Nat. B. Aires 29: 551 (1917)
Blatta orientalis Linnaeus, 1758 (Blattodea, Blattidae) ..................................... Be
Laboulbeniales
4. Aphanandromyces audisioi W. Rossi, Mycologia 74: 522 (1982)
Brachypterus urticae (Fabricius, 1792) (Coleoptera, Kateretidae) ....................... Be
5. Asaphomyces tubanticus (Middelh. & Boelens) Scheloske, Parasitol. Schriftenr.
19: 92 (1969)
Catops fuliginosus Erichson, 1837 (Coleoptera, Leiodidae) ............................... Nl
Catops fuscus (Panzer, 1794) ....................................................................... Be, Nl
Catops longulus Kellner, 1846 ........................................................................... Be
Catops nigricans (Spence, 1813) ................................................................ Be, Nla
Catops sp. ......................................................................................................... Be
Choleva sp. (Coleoptera, Leiodidae) ................................................................. Nl
a Fungus as Barbariella tubantica Middelh. & Boelens ex Middelh. in Middelhoek (1949).
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
40
6. Bordea denotata Haelew. & De Kesel, Nova Hedwig. 98: 114 (2014)
Bibloporus bicolor (Denny, 1825) (Coleoptera, Staphylinidae) .......................... Nl
7. Botryandromyces heteroceri (axt.) I.I. Tav. & T. Majewski, Mycotaxon 3:
195(1976)
Heterocerus fenestratus (unberg, 1784) (Coleoptera, Heteroceridae) .............. Be
Heterocerus exuosus Stephens, 1828 ................................................................. Be
Heterocerus hispidulus Kiesenwetter, 1843......................................................... Be
Heterocerus obsoletus Curtis, 1828 ..................................................................... Nl
8. Cantharomyces denigratus axt., Mem. Am. Acad. Arts Sci. 16: 27 (1931)
Dryops luridus (Erichson, 1847) (Coleoptera, Dryopidae) ................................ Be
9. Cantharomyces elongatus Haelew. & De Kesel, Mycotaxon 123: 468 (2013)
Syntomium aeneum (Müller, 1821) (Coleoptera, Staphylinidae) ....................... Nl
10. Cantharomyces italicus Speg., Anal. Mus. Nac. Hist. Nat. B. Aires 27: 42 (1915)
Dryops luridus (Erichson, 1847) (Coleoptera, Dryopidae) ................................ Be
11. Cantharomyces orientalis Speg., Anal. Mus. Nac. Hist. Nat. B. Aires 27: 43 (1915)
Carpelimus corticinus (Gravenhorst, 1806) (Coleoptera, Staphylinidae) ..... Be, Nla
Carpelimus foveolatus (Sahlberg, 1832) ............................................................. Be
Carpelimus sp. .................................................................................................. Be
Diglotta mersa (Haliday, 1837) (Coleoptera, Staphylinidae) ............................. Be
a Host as Troglophloeus corticinus (Gravenhorst, 1806), fungus as Cantharomyces thax-
teri Maire in Middelhoek (1949).
12. Cantharomyces platystethi axt., Proc. Am. Acad. Arts Sci. 35: 415 (1900)
Platystethus sp. (Coleoptera, Staphylinidae) ...................................................... Be
13. Cantharomyces robustus T. Majewski, Acta Mycol. 23: 99 (1990) [1987]
Carpelimus bilineatus Stephens, 1834 (Coleoptera, Staphylinidae) ................... Be
Carpelimus corticinus (Gravenhorst, 1806) ....................................................... Be
Carpelimus rivularis (Motschulsky, 1860) ................................................... Be, Nl
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 41
Carpelimus sp. .................................................................................................. Be
Gnypeta rubrior Tottenham, 1939 (Coleoptera, Staphylinidae) ........................ Be
14. Chaetarthriomyces crassiappendicatus Scheloske
Chaetarthria seminulum (Herbst, 1797) (Coleoptera, Hydrophilidae) .............. Nl
15. Chitonomyces aculeifer Speg., Anal. Mus. Nac. Hist. Nat. B. Aires 27: 44 (1915)
Graptodytes pictus (Fabricius, 1787) (Coleoptera, Dytiscidae) ........................... Be
Haliplus sp. (Coleoptera, Haliplidae) ............................................................... Be
16. Chitonomyces bidessarius (axt.) axt., Mem. Am. Acad. Arts Sci. 12:
292(1902)
Hygrotus impressopunctatus (Schaller, 1783) (Coleoptera, Dytiscidae) ............... Nl
17. Chitonomyces italicus Speg., Anal. Mus. Nac. Hist. Nat. B. Aires 27: 46 (1915)
Laccophilus hyalinus (De Geer, 1774) (Coleoptera, Dytiscidae) ........................ Be
18. Chitonomyces melanurus Peyr., Sitzber. Akad. Wiss. Wien Math.-Naturw. Kl.
68: 250 (1873)
Laccophilus hyalinus (De Geer, 1774) (Coleoptera, Dytiscidae) ........................ Be
Laccophilus minutus (Linnaeus, 1758) ............................................................. Nla
a New record: Utrecht Province, Soest, Soesterveen, 17 Oct. 1924, leg. F.C. Drescher, on
Laccophilus minutus [as Laccophilus obscurus (Panzer, 1795)] (Naturalis Biodiver-
sity Center), slide D. Haelew. 075a (BR-MYCO, 5 thalli, margin of left elytron).
19. Chitonomyces paradoxus (Peyr.) axt., Mem. Am. Acad. Arts Sci. 12:
287(1902)
Laccophilus hyalinus (De Geer, 1774) (Coleoptera, Dytiscidae) ........................ Be
Laccophilus minutus (Linnaeus, 1758) .............................................................. Nl
20. Compsomyces lestevae axt., Proc. Am. Acad. Arts Sci. 35: 439 (1900)
Lesteva longoelytrata (Goeze, 1777) (Coleoptera, Staphylinidae) ....................... Be
Lesteva pubescens Mannerheim, 1830 ............................................................... Be
Lesteva sicula subsp. heeri Fauvel, 1871....................................................... Be, Nl
Lesteva sp. ........................................................................................................ Be
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
42
21. Coreomyces arcuatus axt., Mem. Am. Acad. Arts Sci. 16: 324 (1931)
Sigara striata (Linnaeus, 1758) (Hemiptera, Corixidae).................................... Be
22. Corethromyces henrotii Balazuc [as ‘henroti’], Bull. Mens. Soc. Linn. Lyon 42:
283 (1973)
Choleva cisteloides (Frölich, 1799) (Coleoptera, Leiodidae) ............................... Be
Choleva fagniezi Jeannel, 1922 ......................................................................... Nl
Choleva jeanneli Britten, 1922.......................................................................... Nl
Choleva oblonga Latreille, 1708 ........................................................................ Nl
23. Corethromyces stilici axt., Proc. Am. Acad. Arts Sci. 37: 42 (1901)
Rugilus (Rugilus) rupes Germar, 1836 (Coleoptera, Staphylinidae) ........... Be, Nla
Rugilus (Rugilus) similis (Erichson, 1839) ......................................................... Be
Rugilus sp. ........................................................................................................ Be
a Host as Stilicus rupes (Germar, 1836) in Middelhoek (1943a, 1945).
24. Cryptandromyces bibloplecti T. Majewski, Acta Mycol. 25: 43 (1990)
Pselaphinae gen et sp. indet. (Coleoptera, Staphylinidae) ................................. Be
25. Cryptandromyces elegans (Maire) W. Rossi & D. Castaldo, Pl. Biosystems 138:
264 (2004)
Brachygluta fossulata (Reichenbach, 1816) (Coleoptera, Staphylinidae) ............ Nl
Brachygluta xanthoptera Reichenbach, 1816 ..................................................... Be
26. Cryptandromyces euplecti Santam., Nova Hedwig. 72: 384 (2001)
Euplectus sanguineus Denny, 1825 (Coleoptera, Staphylinidae) ........................ Be
27. Dimorphomyces myrmedoniae axt., Proc. Am. Acad. Arts Sci. 36: 409
(1900)[1901]
Gnypeta rubrior Tottenham, 1939 (Coleoptera, Staphylinidae) ......................... Be
28. Diphymyces kaaistoepi Haelew. & De Kesel, Sterbeeckia 35: 63 (2019)
Choleva cisteloides (Frölich, 1799) (Coleoptera, Leiodidae) ............................... Be
Choleva fagniezi Jeannel, 1922 ......................................................................... Nl
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 43
29. Distolomyces forculae (T. Majewski) I.I. Tav., Mycol. Mem. 9: 207 (1985)
Forcula auricularia Linnaeus, 1758 (Dermaptera, Forculidae) ................ Be, Nl
30. Ecteinomyces trichopterophilus axt., Proc. Am. Acad. Arts Sci. 38: 26
(1902)[1903]
Acrotrichis fascicularis (Herbst, 1793) (Coleoptera, Ptiliidae) ............................ Be
Acrotrichis grandicollis (Mannerheim, 1844) ..................................................... Nl
Acrotrichis intermedia (Gillmeister, 1845) ......................................................... Be
Acrotrichis sp. ................................................................................................... Be
31. Eucantharomyces stammeri Scheloske, Parasitol. Schriftenr. 19: 108 (1969)
Calathus melanocephalus (Linnaeus, 1758) (Coleoptera, Carabidae) ................. Be
32. Euphoriomyces agathidii (Maire) I.I. Tav., Mycol. Mem. 9: 218 (1985)
Agathidium laevigatum Erichson, 1845 (Coleoptera, Leiodidae) ...................... Nla
a New record: Noord Brabant Province, Tilburg, nature reserve De Kaaistoep, 51.540672
N 5.013867 E, 3–17 Jun. 2000, leg. Working Group Insects of the Royal Dutch
Natural History Association (KNNV), pitfall trap, ±2.5 m S of Quercus robur #2,
on Agathidium laevigatum (NNKN), slides D. Haelew. 1064a (FH, 1 submature
and 2 mature thalli, tip of left elytron) and D. Haelew. 1064b (NMBT, 1 juvenile
and 2 mature thalli, tip of right elytron).
33. Euzodiomyces lathrobii axt., Proc. Am. Acad. Arts Sci. 35: 449 (1900)
Lathrobium brunnipes (Fabricius, 1793) (Coleoptera, Staphylinidae) ................ Be
Lathrobium elongatum (Linnaeus, 1767) .................................................... Be, Nl
Lathrobium geminum Kraatz, 1857 ............................................................ Be, Nl
Lathrobium laevipenne Heer, 1839 ................................................................... Nl
Lathrobium sp. ................................................................................................. Be
Lobrathium multipunctum (Gravenhorst, 1802) (Coleoptera, Staphylinidae) .... Be
Patrobus atrorufus (Stroem, 1768) (Coleoptera, Carabidae) .............................. Be
Pterostichus strenuus (Panzer, 1796) (Coleoptera, Carabidae) ............................ Be
34. Fanniomyces burdigalensis Balazuc, Revue Mycol. 43: 402 (1979)
Copromyza stercoraria (Meigen, 1830) (Diptera, Sphaeroceridae) .....................Bea
Crumomyia pedestris (Meigen, 1830) (Diptera, Sphaeroceridae) .......................Bea
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
44
a Fungus as Stigmatomyces burdigalensis (Balazuc) A. Weir & W. Rossi in De Kesel et
al. (2020).
35. Fanniomyces ceratophorus (Whisler) T. Majewski, Acta Mycol. 8: 230 (1972)
Fannia canicularis (Linnaeus, 1761) (Diptera, Fanniidae) ............................... Nla
a Fungus described as Stigmatomyces ceratophorus Whisler, and later recombined in Fan-
niomyces T. Majewski by Majewski (1972), based on the branching pattern of the
primary appendage. Weir and Rossi (1995), in turn, found no valid rationale to
maintain Fanniomyces as a separate genus and considered it a junior synonym of
Stigmatomyces, stating that “the structure of the antheridial appendage is particu-
larly variable”. However, based on an SSU–LSU ribosomal DNA dataset, Haele-
waters et al. (in press) found that 1) Stigmatomyces as currently circumscribed is
paraphyletic and 2) Fanniomyces is supported as a stand-alone genus with two
species, F. burdigalensis and F. ceratophorus.
36. Haplomyces texanus axt., Proc. Am. Acad. Arts Sci. 28: 160 (1893)
Bledius gallicus (Gravenhorst, 1806) (Coleoptera, Staphylinidae) .................... Nla
a Host as Bledius fracticornis (Paykull, 1790) in Middelhoek (1943a).
37. Helodiomyces elegans F. Picard, Bull. Soc. Mycol. Fr. 29: 557 (1913)
Dryops anglicanus Edwards, 1909 (Coleoptera, Dryopidae) .............................. Nl
Dryops auriculatus (Georoy, 1785) ................................................................. Nl
Dryops luridus (Erichson, 1847) ................................................................. Be, Nl
38. Hesperomyces coccinelloides axt., Mem. Am. Acad. Arts Sci. 16: 110 (1931)
Stethorus punctillum (Weise, 1891) (Coleoptera, Coccinellidae) ....................... Be
39. Hesperomyces halyziae Haelew. & De Kesel, sp. nov.
Halyzia sedecimguttata (Linnaeus, 1758) (Coleoptera, Coccinellidae) .......Bea, Nlb
a Fungus as Hesperomyces virescens axt. sensu lato in De Kesel et al. (2020).
b Fungus as Hesperomyces virescens axt. in Haelewaters and van Wielink (2016) and
Haelewaters et al. (2017).
40. Hesperomyces virescens axt., Proc. Am. Acad. Arts Sci. 25: 264 (1891), sensulato
Harmonia axyridis (Pallas, 1773) (Coleoptera, Coccinellidae) .................... Be, Nl
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 45
Tytthaspis sedecimpunctata (Linnaeus, 1761) (Coleoptera, Coccinellidae) ......... Be
41. Hydraeomyces halipli (axt.) axt., Mem. Am. Acad. Arts Sci. 12: 294 (1902)
Haliplus avicollis Sturm, 1834 (Coleoptera, Haliplidae) ................................. Nl
Haliplus immaculatus Gerhardt, 1877 .............................................................. Be
Haliplus lineatocollis (Marsham, 1802) ............................................................. Be
Haliplus lineolatus Mannerheim, 1844 ............................................................. Be
Haliplus rucollis (De Geer, 1774) ............................................................. Be, Nl
Haliplus sp. ...................................................................................................... Be
42. Hydrophilomyces cf. gracilis T. Majewski, Acta Mycol. 10: 272 (1974)
Cercyon marinus omson, 1853 (Coleoptera, Hydrophilidae) ........................ Be
Cercyon sp. ....................................................................................................... Be
43. Hydrophilomyces cf. hamatus T. Majewski, Acta Mycol. 10: 274 (1974)
Cercyon marinus omson, 1853 (Coleoptera, Hydrophilidae) ........................ Be
44. Idiomyces peyritschii axt., Proc. Am. Acad. Arts Sci. 28: 162 (1893)
Deleaster dichrous Gravenhorst, 1802 (Coleoptera, Staphylinidae) .............. Be, Nl
45. Kainomyces rehmanii T. Majewski, Polish Bot. Stud. 1: 121 (1990)
Acrotrichis dispar (Matthews, 1865) (Coleoptera, Ptiliidae) .............................. Nl
Acrotrichis sp. ................................................................................................... Be
46. Laboulbenia acupalpi Speg., Anal. Mus. Nac. Hist. Nat. B. Aires 26: 458 (1915)
Acupalpus parvulus (Sturm, 1825) (Coleoptera, Carabidae) .............................. Nl
47. Laboulbenia anoplogenii axt., Proc. Am. Acad. Arts Sci. 35: 156 (1899)
[1899–1900]
Stenolophus mixtus (Herbst, 1784) (Coleoptera, Carabidae)........................ Be, Nl
Stenolophus teutonus (Schrank, 1781) ............................................................... Be
48. Laboulbenia argutoris Cépède & F. Picard, Bull. Biol. Fr. Belg. 42: 260 (1909)
Pterostichus diligens (Sturm, 1824) (Coleoptera, Carabidae) ............................. Be
Pterostichus strenuus (Panzer, 1796) ............................................................ Be, Nl
Pterostichus vernalis (Panzer, 1796) ................................................................... Nl
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
46
49. Laboulbenia atlantica axt., Mem. Am. Acad. Arts Sci. 12: 336 (1902)
Lobrathium multipunctum (Gravenhorst, 1802) (Coleoptera, Staphylinidae) .... Be
50. Laboulbenia aubryi Balazuc, Revue Mycol. 43: 393 (1979)
Amara aenea (De Geer, 1774) (Coleoptera, Carabidae) ....................................Bea
a New record: Belgium, Province Vlaams Brabant, Meise, Domein van Bouchout,
50.9274389 N 4.323925 E, ca. 25 m a.s.l., 6 Apr. 2020, leg. A. De Kesel, wet mead-
ow, on Amara aenea, ADK6520 (BR), slides ADK6520a (BR-MYCO, 1 mature thal-
lus, elytra) and ADK6520b (BR-MYCO, 2 immature and 4 mature thalli, elytra).
51. Laboulbenia barbara Middelh. & Boelens, Ned. Kruidk. Arch. 53: 99 (1943a)
Philonthus punctus (Gravenhorst, 1802) (Coleoptera, Staphylinidae) ................ Nl
52. Laboulbenia benjaminii Balazuc ex Santam., Fl. Mycol. Iber. 4: 45 (1998)
Badister bullatus (Schrank, 1798) (Coleoptera, Carabidae) ........................ Be, Nla
Badister lacertosus Sturm, 1815 ......................................................................... Be
Badister sodalis (Duftschmid, 1812) .......................................................... Be, Nlb
Badister unipustulatus Bonelli, 1813 ................................................................. Be
a Host as Badister bipustulatus (Fabricius, 1792), fungus as Laboulbenia polyphaga
axt. in Middelhoek (1949) and Meijer (1975).
b Fungus as Laboulbenia polyphaga axt. in Meijer (1975).
53. Laboulbenia calathi T. Majewski, Polish Bot. Stud. 7: 89 (1994)
Calathus erratus (Sahlberg, 1827) (Coleoptera, Carabidae) ............................... Be
Calathus fuscipes (Goeze, 1777) ........................................................................ Nl
Calathus melanocephalus (Linnaeus, 1758) ................................................. Be, Nl
54. Laboulbenia clivinalis axt., Proc. Am. Acad. Arts Sci. 35: 155 (1899)
[1899–1900]
Clivina collaris (Herbst, 1784) (Coleoptera, Carabidae) ................................... Be
Clivina fossor (Linnaeus, 1758) .................................................................. Be, Nl
55. Laboulbenia collae T. Majewski, Polish Bot. Stud. 7: 104 (1994)
Agonum micans (Nicolai, 1822) (Coleoptera, Carabidae) ................................. Be
Paranchus albipes (Fabricius, 1796) (Coleoptera, Carabidae) ...................... Be, Nl
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 47
56. Laboulbenia coneglianensis Speg., Redia 10: 47 (1914)
Harpalus anis (Schrank, 1781) (Coleoptera, Carabidae) ......................... Be, Nla
Harpalus atratus Latreille, 1804 ........................................................................ Be
Harpalus attenuatus Stephens, 1828 ................................................................. Be
Harpalus griseus (Panzer, 1796) ................................................................. Be, Nlb
Harpalus rupes (De Geer, 1774) ..................................................................... Be
Harpalus tardus (Panzer, 1796) ................................................................... Be, Nl
Harpalus sp. ..................................................................................................... Be
Ophonus rubarbis (Fabricius, 1792) (Coleoptera, Carabidae) .......................... Be
Parophonus maculicornis (Duftschmid, 1812) (Coleoptera, Carabidae) ............ Nlc
a Host as Harpalus aeneus (Fabricius, 1775), fungus as Laboulbenia elongata axt. in
Middelhoek (1949).
b Host as Pseudophonus griseus (Panzer, 1796), fungus as Laboulbenia elongata axt. in
Middelhoek (1949).
c Fungus as Laboulbenia melanaria axt. in Haelewaters et al. (2012a).
57. Laboulbenia cristata axt., Proc. Am. Acad. Arts Sci. 29: 174 (1893)
Paederus fuscipes Curtis, 1826 (Coleoptera, Staphylinidae) ............................... Nl
Paederus littoralis Gravenhorst, 1802 ................................................................ Be
Paederus riparius (Linnaeus, 1758) ............................................................. Be, Nl
Paederus sp. ...................................................................................................... Be
58. Laboulbenia dubia axt., Proc. Am. Acad. Arts Sci. 38: 35 (1902) [1903]
Philonthus cognatus Stephens, 1832 (Coleoptera, Staphylinidae) ...................... Be
59. Laboulbenia egens Speg., Anal. Soc. Cient. Argent. 85: 323 (1918)
Elaphropus parvulus (Dejean, 1831) (Coleoptera, Carabidae) ........................... Be
Paratachys micros (Fischer von Waldheim, 1828) (Coleoptera, Carabidae) ........ Be
60. Laboulbenia elaphri Speg., Anal. Mus. Nac. B. Aires 26: 64 (1915)
Elaphrus cupreus Duftschmid, 1812 (Coleoptera, Carabidae) ........................... Be
Elaphrus riparius (Linnaeus, 1758) ................................................................... Be
61. Laboulbenia eubradycelli Huldén, Karstenia 25: 4 (1985)
Bradycellus harpalinus (Audinet-Serville, 1821) (Coleoptera, Carabidae) .... Be, Nl
Bradycellus rucollis (Stephens, 1828) ............................................................... Be
Bradycellus verbasci (Duftschmid, 1812) ..................................................... Be, Nl
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
48
Trichocellus placidus (Gyllenhal, 1827) (Coleoptera, Carabidae) ....................... Be
62. Laboulbenia fasciculata Peyr., Sitzber. Akad. Wiss. Wien Math.-naturw. Kl.
68: 248 (1873)
Nebria brevicollis (Fabricius, 1792) (Coleoptera, Carabidae) ............................. Be
Omophron limbatum (Fabricius, 1777) (Coleoptera, Carabidae) ................ Be, Nla
Patrobus atrorufus (Stroem, 1768) (Coleoptera, Carabidae) .............................. Be
Pterostichus nigrita (Paykull, 1790) (Coleoptera, Carabidae) ............................. Be
a New record: No locality, no date, on Omophron limbatum (Naturalis Biodiversity
Center), slide D. Haelew. 074a (BR-MYCO, 3 thalli, left metatibia).
63. Laboulbenia fennica Huldén, Karstenia 23: 54 (1983)
Gyrinus marinus Gyllenhal, 1808 (Coleoptera, Gyrinidae) ............................... Nl
Gyrinus substriatus Stephens, 1829 ............................................................. Be, Nl
64. Laboulbenia lifera axt., Proc. Am. Acad. Arts Sci. 28: 165 (1893)
Harpalus anis (Schrank, 1781) (Coleoptera, Carabidae) ............................... Nla
a Host as Harpalus aeneus (Fabricius, 1775) in Middelhoek (1949). e microscope
slide from the collection of W.J. Kossen was reported to be in very poor condi-
tion; as a result, no illustrations could be made (Middelhoek 1949). For the
time being, we retain the identication of the fungus. Laboulbenia lifera was
described on a species of Anisodactylus Dejean, 1829 (Coleoptera, Carabidae) in
the USA, and it is possible that European records of L. lifera belong in L. agel-
lata (Majewski 1994, Haelewaters et al. 2019a). e species is not included in
the identication key.
65. Laboulbenia agellata Peyr., Sitzber. Akad. Wiss. Wien Math.-naturw. Kl. 68:
247 (1873), sensu lato
Agonum emarginatum (Gyllenhal, 1827) (Coleoptera, Carabidae) .................... Be
Acupalpus avicollis (Sturm, 1825) .................................................................. Nla
Agonum fuliginosum (Panzer, 1809) ........................................................... Be, Nlb
Agonum marginatum (Linnaeus, 1758) ....................................................... Be, Nl
Agonum micans (Nicolai, 1822) ........................................................................ Be
Agonum moestum (Duftschmid, 1812) ...................................................... Be, Nla
Agonum muelleri (Herbst, 1784) ................................................................ Be, Nl
Agonum nigrum Dejean, 1828 .......................................................................... Be
Agonum thoreyi Dejean, 1828 ..................................................................... Be, Nl
Agonum viduum (Panzer, 1796) ........................................................................ Nl
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 49
Agonum viridicupreum Goeze, 1777 ................................................................. Be
Anchomenus dorsalis (Pontoppidan, 1763) (Coleoptera, Carabidae) ................. Nlc
Anisodactylus binotatus (Fabricius, 1787) (Coleoptera, Carabidae) .................... Be
Laemostenus terricola (Herbst, 1784) (Coleoptera, Carabidae) .......................... Be
Limodromus assimilis (Paykull, 1790) (Coleoptera, Carabidae) .................. Be, Nld
Loricera pilicornis (Fabricius, 1775) (Coleoptera, Carabidae) ............................ Be
Nebria brevicollis (Fabricius, 1792) (Coleoptera, Carabidae) ............................. Be
Oxypselaphus obscurus (Herbst, 1784) (Coleoptera, Carabidae)......................... Be
Paranchus albipes (Fabricius, 1796) (Coleoptera, Carabidae) ..................... Be, Nle
Parophonus maculicornis (Duftschmid, 1812) ................................................... Be
Pterostichus vernalis (Panzer, 1796) ................................................................... Be
Trichotichnus laevicollis (Duftschmid, 1812) (Coleoptera, Carabidae)............... Be
a Fungus as Laboulbenia elongata axt. in Middelhoek (1949).
b Host as Europhilus fuliginosus (Panzer, 1809), fungus as Laboulbenia elongata axt.
in Middelhoek (1949).
c Host as Platynus dorsalis (Pontoppidan, 1763) in Zaneveld (1938), as Agonum dorsale
(Pontoppidan, 1763) in Meijer (1975).
d Host as Platynus assimilis (Paykull, 1790) in Zaneveld (1938).
e Host as Platynus rucornis (Goeze, 1777) in Zaneveld (1938).
66. Laboulbenia giardii Cépède & F. Picard, Bull. Sci. Fr. Belg. 42: 258 (1908)
Dicheirotrichus gustavii Crotch, 1871 (Coleoptera, Carabidae) .................. Be, Nla
Dicheirotrichus obsoletus (Dejean, 1829) ........................................................... Be
a Host as Dicheirotrichus pubescens (Paykull, 1790) in Meijer (1975).
67. Laboulbenia gyrinicola Speg., Redia 10: 34 (1914)
Gyrinus marinus Gyllenhal, 1808 (Coleoptera, Gyrinidae) ......................... Be, Nl
Gyrinus natator (Linnaeus, 1758) ..................................................................... Be
Gyrinus substriatus Stephens, 1829 ................................................................... Nl
68. Laboulbenia hyalopoda De Kesel, Sterbeeckia 18: 17 (1998)
Paradromius linearis (Olivier, 1795) (Coleoptera, Carabidae) ........................... Be
69. Laboulbenia inata axt., Proc. Am. Acad. Arts Sci. 27: 41 (1892)
Acupalpus dubius Schilsky, 1888 (Coleoptera, Carabidae) ........................... Be, Nl
Acupalpus exiguus Dejean, 1829 ....................................................................... Be
Acupalpus parvulus (Sturm, 1825) .................................................................... Nl
Stenolophus mixtus (Herbst, 1784) (Coleoptera, Carabidae).............................. Be
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
50
70. Laboulbenia kajanensis Huldén, Karstenia 23: 56 (1983)
Pterostichus diligens (Sturm, 1824) (Coleoptera, Carabidae) ............................. Be
Pterostichus strenuus (Panzer, 1796) .................................................................. Be
71. Laboulbenia lecoareri (Balazuc) Huldén, Karstenia 25: 6 (1985)
Trechoblemus micros (Herbst, 1784) (Coleoptera, Carabidae) ........................... Be
72. Laboulbenia leisti J. Siemaszko & Siemaszko, Polsk. Pism. Entomol. 6: 203
(1928) [1927]
Agonum muelleri (Herbst, 1784) (Coleoptera, Carabidae) ................................ Be
Leistus ferrugineus (Linnaeus, 1758) (Coleoptera, Carabidae) ..................... Be, Nl
73. Laboulbenia lichtensteinii F. Picard, Bull. Sci. Fr. Belg. 50: 449 (1917)
[1916–1917]
Cillenus lateralis Samouelle, 1819 (Coleoptera, Carabidae) .............................. Nl
74. Laboulbenia littoralis De Kesel & Haelew., Mycologia 106: 408 (2014)
Caus xantholoma (Gravenhorst, 1806) (Coleoptera, Staphylinidae) .......... Be, Nl
75. Laboulbenia luxurians Peyr., Sitzber. Akad. Wiss. Wien Math.-naturw. Kl. 68:
248 (1873)
Bembidion dentellum (unberg, 1787) (Coleoptera, Carabidae) ..................... Nl
76. Laboulbenia metableti Scheloske, Parasitol. Schriftenr. 19: 124 (1969)
Syntomus foveatus (Georoy, 1785) (Coleoptera, Carabidae) ..................... Be, Nla
Syntomus truncatellus (Linnaeus, 1760) ..................................................... Be, Nla
a New records: Noord-Holland Province, Zuid-Kennemerland National Park,
31 Oct. 2016, leg. M. Boeken, pitfall trap, on Syntomus truncatellus, slide D.
Haelew. 1236b (GENT, 2 juvenile thalli, pronotum). Ibid., 5 Jun. 2017, leg. M.
Boeken, pitfall trap, on Syntomus truncatellus, slide D. Haelew. 1378a (GENT,
2 mature thalli, posterior margin of right elytron). Ibid., 5 Jun. 2017, leg. M.
Boeken, pitfall trap, on Syntomus foveatus, slide D. Haelew. 1387a (GENT, 1
mature thallus, left elytron). Ibid., 5 Jun. 2017, leg. M. Boeken, pitfall trap, on
Syntomus foveatus, slides D. Haelew. 1391a (FH, 5 mature thalli, right elytron),
D. Haelew. 1391b (FH, 1 mature thallus, left metatrochanter), and D. Haelew.
1391c (FH, 1 submature and 2 mature thalli, mesocoxae). Ibid., 17 Jul. 2017,
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 51
leg. M. Boeken, pitfall trap, on Syntomus truncatellus, slide D. Haelew. 1379a
(GENT, 2juvenile thalli, left elytron).
77. Laboulbenia murmanica Huldén, Karstenia 23: 57 (1983)
Bembidion assimile Gyllenhal, 1810 (Coleoptera, Carabidae) ........................... Be
78. Laboulbenia notiophili Cépède & F. Picard, Bull. Biol. Fr. Belg. 42: 259 (1909)
Demetrias atricapillus (Linnaeus, 1758) (Coleoptera, Carabidae) ...................... Be
Demetrias imperialis (Germar, 1824) ................................................................ Be
Demetrias monostigma Leach, 1819 .................................................................. Be
Notiophilus biguttatus (Fabricius, 1779) (Coleoptera, Carabidae) ............... Be, Nl
Notiophilus rupes Curtis, 1829 ........................................................................ Be
Notiophilus substriatus Waterhouse, 1833 ......................................................... Nl
Notiophilus sp. .................................................................................................. Be
Paradromius linearis (Olivier, 1795) (Coleoptera, Carabidae) .................... Be, Nla
Philorhizus melanocephalus (Dejean, 1825) (Coleoptera, Carabidae)................. Nl
a Fungus as Laboulbenia casnoniae axt. in Haelewaters et al. (2012a).
79. Laboulbenia ophoni axt., Proc. Am. Acad. Arts Sci. 35: 190 (1899)
[1899–1900]
Harpalus rubripes (Duftschmid, 1812) (Coleoptera, Carabidae) ....................... Be
Ophonus rubarbis (Fabricius, 1792) (Coleoptera, Carabidae) .......................... Be
80. Laboulbenia pedicellata axt., Proc. Am. Acad. Arts Sci. 29: 109 (1893)
Bembidion aeneum Germar, 1824 (Coleoptera, Carabidae) ........................ Be, Nl
Bembidion articulatum (Panzer, 1796) .............................................................. Nl
Bembidion biguttatum (Fabricius, 1779) ........................................................... Nl
Bembidion gilvipes Sturm, 1825........................................................................ Be
Bembidion guttula (Fabricius, 1792) ........................................................... Be, Nl
Bembidion iricolor Bedel, 1879 ................................................................... Be, Nl
Bembidion lunulatum (Georoy, 1785) ...................................................... Be, Nl
Bembidion minimum (Fabricius, 1792) ...................................................... Be, Nl
Bembidion normannum Dejean, 1831 ........................................................ Be, Nl
Bembidion obtusum Audinet-Serville, 1821 ...................................................... Be
Bembidion quadrimaculatum (Linnaeus, 1760) .......................................... Be, Nl
Bembidion ustulatum (Linnaeus, 1758) ............................................................ Nl
Bembidion varium (Olivier, 1795) .............................................................. Be, Nl
Dyschirius globosus (Herbst, 1784) (Coleoptera, Carabidae) .............................. Nl
Dyschirius salinus Schaum, 1843 ...................................................................... Nl
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
52
Dyschirius thoracicus (P. Rossi, 1790) ............................................................... Nla
Dyschirius tristis Stephens, 1827 ....................................................................... Be
Dyschirius sp. ................................................................................................... Nl
Pogonus chalceus (Marsham, 1802) (Coleoptera, Carabidae) ....................... Be, Nl
a Host as Dyschirius arenosus Stephens, 1827 in Middelhoek (1943a).
81. Laboulbenia philonthi axt., Proc. Am. Acad. Arts Sci. 28: 174 (1893)
Philonthus micans (Gravenhorst, 1802) (Coleoptera, Staphylinidae) ................. Nl
Philonthus rubripennis Stephens, 1832 (Coleoptera, Staphylinidae) .................. Be
Philonthus sp. ................................................................................................... Be
82. Laboulbenia pseudomasei axt., Proc. Am. Acad. Arts Sci. 35: 196 (1899)
Loricera pilicornis (Fabricius, 1775) (Coleoptera, Carabidae) ............................ Be
Nebria brevicollis (Fabricius, 1792) (Coleoptera, Carabidae) ............................. Be
Pterostichus anthracinus (Panzer, 1795) (Coleoptera, Carabidae) ....................... Be
Pterostichus melanarius (Illiger, 1798) .............................................................. Nla
Pterostichus minor (Gyllenhal, 1827) ................................................................ Be
Pterostichus nigrita (Paykull, 1790) ................................................................... Be
Pterostichus strenuus (Panzer, 1796) .................................................................. Be
Stomis pumicatus (Panzer, 1796) (Coleoptera, Carabidae) ................................. Be
a New record: Drenthe Province, Oude Willem, 52.897438 N 6.323432 E, 2 Jun.
2014, leg. A.J. Dees, on Pterostichus melanarius (NNKN), slides D. Haelew. 1013a
(FH, 1 juvenile thallus, right elytron) and D. Haelew. 1013b (FH, 1 submature
thallus, prosternum).
83. Laboulbenia quarantenae De Kesel & Haelew, sp. nov.
Bembidion (Philochtus) biguttatum (Fabricius, 1779) (Coleoptera, Carabidae) ... Be
84. Laboulbenia rougetii Mont. & C.P. Robin, in Robin, Histoire Naturelle des
végétaux parasites qui croissent sur l’homme et sur les animaux vivants (Paris):
622 (1853)
Brachinus crepitans (Linnaeus, 1758) (Coleoptera, Carabidae) .......................... Be
85. Laboulbenia slackensis Cépède & F. Picard, Compt. Rend. Assoc. Franç.
Avancem. Sci. 35: 775 (1907)
Pogonus chalceus (Marsham, 1802) (Coleoptera, Carabidae) ....................... Be, Nl
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 53
86. Laboulbenia stilicicola Speg., Redia 10: 41 (1914)
Rugilus orbiculatus (Paykull, 1789) (Coleoptera, Staphylinidae) ................ Be, Nla
Rugilus rupes Germar, 1836 (Coleoptera, Staphylinidae) ......................... Be, Nlb
a Host as Stilicus orbiculatus (Paykull, 1789), fungus as Laboulbenia subterranea axt.
in Middelhoek (1943a, 1947a).
b Host as Stilicus rupes Germar, 1836, fungus as Laboulbenia subterranea axt. in
Middelhoek (1943a, 1945).
87. Laboulbenia thaxteri Cépède & F. Picard, Bull. Biol. Fr. Belg. 42: 260 (1909)
Asaphidion avipes (Linnaeus, 1760) (Coleoptera, Carabidae) .......................... Be
88. Laboulbenia vulgaris Peyr., Sitzber. Akad. Wiss. Wien Math.-naturw. Kl. 68:
248 (1873)
Asaphidion avipes (Linnaeus, 1760) (Coleoptera, Carabidae) .......................... Nl
Bembidion assimile Gyllenhal, 1810 (Coleoptera, Carabidae) ........................... Nl
Bembidion biguttatum (Fabricius, 1779) ..................................................... Be, Nl
Bembidion bruxellense Wesmael, 1835 ............................................................. Nla
Bembidion dentellum (unberg, 1787) ..................................................... Be, Nl
Bembidion elongatum Dejean, 1831 ................................................................. Be
Bembidion femoratum Sturm, 1825 ............................................................ Be, Nl
Bembidion iricolor Bedel, 1879 ......................................................................... Nl
Bembidion mannerheimi Sahlberg, 1827 ........................................................... Be
Bembidion minimum (Fabricius, 1792) ............................................................ Nl
Bembidion normannum Dejean, 1831 .............................................................. Nl
Bembidion pallidipenne (Illiger, 1802) .............................................................. Nl
Bembidion properans (Stephens, 1828) ....................................................... Be, Nl
Bembidion stephensii Crotch, 1866 ................................................................... Be
Bembidion testaceum (Duftschmid, 1812) ........................................................ Nl
Bembidion tetracolum Say, 1823 ................................................................. Be, Nl
Bembidion tibiale (Duftschmid, 1812) ............................................................. Be
Bembidion ustulatum (Linnaeus, 1758) ............................................................ Nl
Bembidion sp.................................................................................................... Be
Dyschirius globosus (Herbst, 1784) (Coleoptera, Carabidae) .............................. Nl
Dyschirius salinus Schaum, 1843 ...................................................................... Nl
Ocys harpaloides (Audinet-Serville, 1821) (Coleoptera, Carabidae) ................... Be
Trechus quadristriatus (Schrank, 1781) (Coleoptera, Carabidae) ....................... Be
Trechus rubens (Fabricius, 1792) ....................................................................... Be
a Host as Bembidion rupestre (Linnaeus, 1767) in Meijer (1975).
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
54
89. Mimeomyces zeelandicus Middelh. & Boelens, Ned. Kruidk. Arch. 53: 102 (1943)
Heterothops binotatus (Gravenhorst, 1802) (Coleoptera, Staphylinidae) ............ Nl
90. Misgomyces dyschirii axt., Proc. Am. Acad. Arts Sci. 35: 443 (1900)
Dyschirius aeneus (Dejean, 1825) (Coleoptera, Carabidae) ......................... Be, Nl
Dyschirius globosus (Herbst, 1784) .............................................................. Be, Nl
Dyschirius intermedius Putzeys, 1846 ................................................................ Be
Dyschirius politus (Dejean, 1825) ..................................................................... Nl
Dyschirius salinus Schaum, 1843 ...................................................................... Nl
Dyschirius tristis Stephens, 1827 ................................................................ Be, Nla
a Host as Dyschirius luedersi Wagner, 1915 in Middelhoek (1943a).
91. Monoicomyces bolitocharae T. Majewski, Polish Bot. Stud. 7: 193 (1994)
Bolitochara obliqua Erichson, 1837 (Coleoptera, Staphylinidae) ....................... Be
92. Monoicomyces britannicus axt., Proc. Am. Acad. Arts Sci. 35: 412 (1900)
Acrotona fungi (Gravenhorst, 1806) (Coleoptera, Staphylinidae) ......................Bea
Acrotona orbata (Erichson, 1837) .................................................................... Beb
Acrotona pseudotenera (Cameron, 1933) ........................................................... Nl
Atheta sp. (Coleoptera, Staphylinidae).............................................................. Be
a Host as Atheta (Mocyta) fungi (Gravenhorst, 1806) in De Kesel et al. (2020).
b Host as Atheta (Mocyta) orbata (Erichson, 1837) in De Kesel et al. (2020).
93. Monoicomyces californicus (axt.) axt., Mem. Am. Acad. Arts Sci. 16: 38 (1931)
Anotylus sculpturatus (Gravenhorst, 1806) (Coleoptera, Staphylinidae) ...... Be, Nla
a Host as Oxytelus sculpturatus Gravenhorst, 1806 in Middelhoek (1943a).
94. Monoicomyces fragilis Scheloske, Parasitol. Schriftenr. 19: 138 (1969)
Ocalea picata (Stephens, 1832) (Coleoptera, Staphylinidae) ............................. Be
95. Monoicomyces homalotae axt., Proc. Am. Acad. Arts Sci. 35: 412 (1900)
Atheta aeneicollis (Sharp, 1869) (Coleoptera, Staphylinidae) ............................. Nl
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 55
Atheta amicula (Stephens, 1832) ...................................................................... Nl
Atheta crassicornis (Fabricius, 1792) .................................................................. Nl
Atheta gagatina (Baudi, 1848) .......................................................................... Nl
Atheta longicornis (Gravenhorst, 1802) ............................................................. Be
Atheta triangulum (Kraatz, 1856) ............................................................... Be, Nl
Atheta vestita (Gravenhorst, 1806) ................................................................... Be
Atheta xanthopus (omson, 1856) .................................................................. Nl
Atheta sp. ......................................................................................................... Be
96. Monoicomyces invisibilis axt., Proc. Am. Acad. Arts Sci. 36: 414 (1900)[1901]
Anotylus sculpturatus (Gravenhorst, 1806) (Coleoptera, Staphylinidae) ............. Be
Anotylus sp. ...................................................................................................... Be
Oxytelus laqueatus (Marsham, 1802) (Coleoptera, Staphylinidae) ..................... Be
Oxytelus sp. ...................................................................................................... Be
Platystethus arenarius (Georoy, 1785) (Coleoptera, Staphylinidae) .................. Be
97. Monoicomyces matthiatis T. Majewski, Acta Mycol. 25: 49 (1990) [1989]
Platystethus cf. arenarius (Georoy, 1785) (Coleoptera, Staphylinidae) ............. Be
98. Monoicomyces myllaenae Santam., Nova Hedwig. 82: 358 (2006)
Myllaena elongata (Matthews, 1838) (Coleoptera, Staphylinidae) ..................... Nl
99. Monoicomyces nigrescens axt., Proc. Am. Acad. Arts Sci. 35: 412 (1900)
Atheta atramentaria (Gyllenhal, 1810) (Coleoptera, Staphylinidae) .................. Nl
Atheta sp. ......................................................................................................... Be
Brundinia marina (Mulsant & Rey, 1853) (Coleoptera, Staphylinidae) ............Bea
Dilacra luteipes (Erichson, 1837) (Coleoptera, Staphylinidae) ......................... Nlb
a Host as Atheta (Actophylla) marina (Mulsant & Rey, 1853) in De Kesel et al. (2020).
b Host as Atheta luteipes (Erichson, 1837) in Middelhoek (1943a).
100. Peyritschiella biformis (axt.) I.I. Tav., Mycol. Mem. 9: 270 (1985)
Philonthus umbratilis (Gravenhorst, 1802) (Coleoptera, Staphylinidae) ............ Be
101. Peyritschiella dubia (axt.) I.I. Tav., Mycol. Mem. 9: 270 (1985)
Philonthus politus (Linnaeus, 1758) (Coleoptera, Staphylinidae) ...................... Be
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
56
102. Peyritschiella furcifera (axt.) I.I. Tav., Mycol. Mem. 9: 270 (1985)
Philonthus albipes (Gravenhorst, 1802) (Coleoptera, Staphylinidae) ................ Nla
Philonthus rectangulus Sharp, 1874 .................................................................. Nla
a Fungus as Dichomyces furciferus axt. in Middelhoek (1943a).
103. Peyritschiella heinemanniana De Kesel, Belg. J. Bot. 131: 177 (1999) [1998]
Xantholinus longiventris Heer, 1839 (Coleoptera, Staphylinidae) ...................... Be
104. Peyritschiella princeps (axt.) I.I. Tav., Mycol. Mem. 9: 270 (1985)
Bisnius cephalotes (Gravenhorst, 1802) (Coleoptera, Staphylinidae) ........... Be, Nla
Bisnius sordidus (Gravenhorst, 1802) ......................................................... Be, Nlb
Bisnius subuliformis (Gravenhorst, 1802) ..........................................................Nl
Philonthus politus (Linnaeus, 1758) (Coleoptera, Staphylinidae) ...................... Be
Philonthus varians (Paykull, 1789) .................................................................. Nlc
Philonthus sp. ................................................................................................... Be
a Host as Philonthus cephalotes (Gravenhorst, 1802), fungus as Dichomyces vulgatus
axt. in Middelhoek (1943a, 1947a).
b Host as Philonthus sordidus (Gravenhorst, 1802), fungus as Dichomyces princeps axt.
in Middelhoek (1941, 1943a, 1943b, 1943c), fungus also as Dichomyces vulgatus
axt. (variety sensu axter 1908: 252) in Middelhoek (1943a, 1943b).
c Fungus as Dichomyces princeps axt. in Middelhoek (1941).
105. Peyritschiella protea axt., Proc. Am. Acad. Arts Sci. 35: 427 (1900)
Anotylus insecatus Gravenhorst, 1806 (Coleoptera, Staphylinidae) .................... Be
Anotylus rugosus (Fabricius, 1775) ............................................................. Be, Nla
Anotylus sp. ...................................................................................................... Be
Staphylinidae sp. indet. (Coleoptera, Staphylinidae) ........................................ Be
a Host as Oxytelus rugosus (Fabricius, 1775) in Middelhoek (1943a, 1947a).
106. Peyritschiella vulgata (axt.) I.I. Tav., Mycol. Mem. 9: 271 (1985)
Philonthus albipes (Gravenhorst, 1802) (Coleoptera, Staphylinidae) ................ Nla
a Fungus as Dichomyces vulgatus axt. in Middelhoek (1943b, 1943c).
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 57
107. Phaulomyces simplocariae De Kesel, Mycotaxon 50: 192 (1994)
Simplocaria semistriata Fabricius, 1794 (Coleoptera, Byrrhidae) ....................... Be
108. Rhachomyces canariensis axt., Proc. Am. Acad. Arts Sci. 35: 436 (1900)
Trechus obtusus Erichson, 1837 (Coleoptera, Carabidae) ........................... Be, Nla
Trechus quadristriatus (Schrank, 1781) ............................................................. Be
Trechus sp. ........................................................................................................ Be
a New record: Noord-Holland Province, Zuid-Kennemerland National Park, 17 Oct.
2016, leg. M. Boeken, pitfall trap, on Trechus obtusus Erichson, 1837 (Coleoptera,
Carabidae), slides D. Haelew. 1242a (GENT, 9 thalli, right margin of pronotum)
and D. Haelew. 1242b (GENT, 3 juvenile thalli, elytra). Ibid., 5 Jun. 2017, leg.
M. Boeken, pitfall trap, on Trechus obtusus, slide D. Haelew. 1388a (GENT, 1
submature thallus, tip of left elytron).
109. Rhachomyces furcatus (axt.) axt., Proc. Am. Acad. Arts Sci. 30: 467
(1895) [1894]
Othius punctulatus (Goeze, 1777) (Coleoptera, Staphylinidae) ......................... Be
Othius subuliformis Stephens, 1833 ........................................................... Bea, Nl
a Host as Othius myrmecophilus Kiesenwetter, 1843 in De Kesel et al. (2020).
110. Rhachomyces lasiophorus (axt.) axt., Proc. Am. Acad. Arts Sci. 30: 468
(1895) [1894]
Acupalpus dubius Schilsky, 1888 (Coleoptera, Carabidae) ................................. Be
Acupalpus exiguus Dejean, 1829 ................................................................. Be, Nl
Anthracus consputus (Duftschmid, 1812) (Coleoptera, Carabidae) .................... Nl
111. Rhachomyces philonthinus axt., Proc. Am. Acad. Arts Sci. 35: 435 (1900)
Bisnius metarius (Gravenhorst, 1802) (Coleoptera, Staphylinidae)........... Be, Nla
Philonthus cruentatus (Gmelin, 1790) (Coleoptera, Staphylinidae) .................. Nlb
Philonthus fumarius (Gravenhorst, 1806) ......................................................... Be
Philonthus marginatus (Müller, 1764) ......................................................... Be, Nl
Philonthus rectangulus Sharp, 1874 ................................................................... Be
Philonthus varians (Paykull, 1789) ............................................................ Be, Nlb
Philonthus sp. ................................................................................................... Be
a Host as Philonthus metarius (Gravenhorst, 1802) in Middelhoek (1943a, 1943d).
b Fungus as Rhachomycesphilonthi’ axt. in Middelhoek (1943b).
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
58
112. Rhachomyces pilosellus (C.P. Robin) axt., Proc. Am. Acad. Arts Sci. 30: 467
(1895) [1894]
Lathrobium fulvipenne (Gravenhorst, 1806) (Coleoptera, Staphylinidae) .......... Be
Lathrobium geminum Kraatz, 1857 .................................................................. Be
113. Rhachomyces spinosus Santam. & A.D. Cuesta-Segura, Nova Hedwig. 110:
362 (2020)
Syntomus foveatus (Georoy, 1785) (Coleoptera, Carabidae) ............................Bea
a Fungus as Rhachomyces sciakyi W. Rossi in De Kesel et al. (2020)
114. Rhachomyces tenenbaumii J. Siemaszko & Siemaszko, Polsk. Pism. Entomol.
6: 205 (1928)
alassophilus longicornis (Sturm, 1825) (Coleoptera, Carabidae) ..................... Be
115. Rhadinomyces cristatus axt., Proc. Am. Acad. Arts Sci. 28: 180 (1893)
Lathrobium brunnipes (Fabricius, 1793) (Coleoptera, Staphylinidae) ................ Be
Lathrobium castaneipenne Kolenati, 1846 ......................................................... Be
Lathrobium elongatum (Linnaeus, 1767) .......................................................... Be
Lathrobium fulvipenne (Gravenhorst, 1806) ..................................................... Be
Lathrobium geminum Kraatz, 1857 .................................................................. Be
Lathrobium sp. ................................................................................................. Be
116. Rhadinomyces pallidus axt., Proc. Am. Acad. Arts Sci. 28: 180 (1893)
Lathrobium elongatum (Linnaeus, 1767) (Coleoptera, Staphylinidae) ............... Nl
117. Rhynchophoromyces anacaenae Scheloske, Parasitol. Schriftenr. 19:
143(1969)
Anacaena lutescens (Stephens, 1829) (Coleoptera, Hydrophilidae) .................... Be
118. Rickia dendroiuli W. Rossi, Rev. Mycol. 41: 531 (1977)
Julida sp. indet. ................................................................................................ Be
119. Rickia laboulbenioides De Kesel, Sterbeeckia 32: 6 (2013)
Cylindroiulus latestriatus (Curtis, 1845) (Julida, Julidae) ............................. Be, Nl
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 59
Cylindroiulus punctatus Leach, 1814 ................................................................. Be
Julida sp. indet. ................................................................................................ Be
120. Rickia peyerimhoi Maire, Bull. Sci. Fr. Belg. 7: 290 (1916)
Scaphisoma sp. (Coleoptera, Staphylinidae) ...................................................... Be
121. Rickia proteini T. Majewski, Acta Mycol. 19: 191 (1985)
Proteinus sp. (Coleoptera, Staphylinidae) ......................................................... Be
122. Rickia wasmannii Cavara, Malpighia 13: 182 (1899)
Myrmica ruginodis Nylander, 1846 (Hymenoptera, Formicidae) ...................... Nl
Myrmica sabuleti Meinert, 1861 (Hymenoptera, Formicidae).................... Be, Nla
Myrmica scabrinodis Nylander, 1846 ................................................................ Nl
a Host as Myrmica scabrinodis Nylander, 1846 in Haelewaters (2012).
123. Siemaszkoa fennica Huldén, Karstenia 23: 63 (1983)
Ptenidium formicetorum Kraatz, 1851 (Coleoptera, Ptiliidae) ........................... Nl
124. Siemaszkoa ptenidii (Scheloske) I.I. Tav. & T. Majewski, Mycotaxon 3: 204(1976)
Ptenidium sp. (Coleoptera, Ptiliidae) ................................................................ Be
125. Stichomyces conosomatis axt., Proc. Am. Acad. Arts Sci. 37: 38 (1901)
Sepedophilus marshami (Stephens, 1832) (Coleoptera, Staphylinidae) ............... Be
Sepedophilus nigripennis (Stephens, 1832) .................................................. Be, Nl
Sepedophilus pedicularius (Gravenhorst, 1802) .................................................. Be
Sepedophilus testaceus (Fabricius, 1792) ............................................................. Nl
Sepedophilus sp. ................................................................................................ Be
126. Stigmatomyces baeri H. Karst., Chemismus P.-Zelle: 78 (1869)
• “Fannia canicularis” (Linnaeus, 1761) (Diptera, Fanniidae) ............................ Nla
a Host as Homalomyia canicularis (Linnaeus, 1761) in Boedijn (1923). e host iden-
tication may have been incorrect; Fannia canicularis is typically associated with
Fanniomyces ceratophorus Whisler, whereas S. baeri is typically found on Musca
domestica Linnaeus, 1758 (Diptera, Muscidae).
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
60
127. Stigmatomyces crassicollis axt., Proc. Am. Acad. Arts Sci. 52: 661 (1917)
Leptocera caenosa (Rondani, 1880) (Diptera, Sphaeroceridae) ........................... Be
Leptocera fontinalis (Fallén, 1826) .................................................................... Be
Leptocera lutosoidea (Duda, 1938) .................................................................... Be
Opacifrons humida (Haliday, 1836) (Diptera, Sphaeroceridae) ......................... Be
Spelobia rulabris (Stenhammar, 1855) (Diptera, Sphaeroceridae) ................... Be
Sphaeroceridae sp. indet. (Diptera) .................................................................. Be
128. Stigmatomyces divergatus axt., Mem. Am. Acad. Arts Sci. 16: 122 (1931)
Apteromyia claviventris (Strobl, 1909) (Diptera, Sphaeroceridae) ...................... Be
Spelobia parapusio (Dahl, 1909) (Diptera, Sphaeroceridae)............................... Be
Spelobia sp. ...................................................................................................... Be
129. Stigmatomyces entomophilus (Peck) axt., Proc. Am. Acad. Arts Sci. 36: 398
(1900) [1901]
Drosophila funebris (Fabricius, 1787) (Diptera, Drosophilidae) ........................ Nl
130. Stigmatomyces hydrelliae axt., Proc. Am. Acad. Arts Sci. 36: 404 (1900) [1901]
Hydrellia albilabris (Meigen, 1830) (Diptera, Ephydridae) .............................. Nla
a New record: Noord-Brabant Province, Udenhout, nature reserve De Brand, 51.631777
N 5.132998 E, 14–21 Jun. 1990, leg. Working Group Insects of the Royal Dutch
Natural History Association (KNNV), malaise trap (van Zuijlen et al. 1996), on
Hydrellia albilabris (Meigen, 1830) (Diptera, Ephydridae), slide WR1746 (will
be deposited at FI, Herbarium Universitatis Florentinae, Florence, Italy), det. W.
Rossi, comm. J.W.A. van Zuijlen.
131. Stigmatomyces limosinae axt., Proc. Am. Acad. Arts Sci. 36: 406 (1900) [1901]
Spelobia clunipes (Meigen, 1830) (Diptera, Sphaeroceridae) ............................. Be
Spelobia talparum (Richards, 1927) .................................................................. Nl
132. Stigmatomyces majewskii H.L. Dainat, Manier & Balazuc, Bull. Trimest.
Soc. Mycol. Fr. 90: 171 (1974)
Drosophila subobscura Collin, 1936 (Diptera, Drosophilidae) ........................... Nl
133. Stigmatomyces minilimosinae T. Majewski, Polish Bot. Stud. 1: 122 (1990)
Minilimosina parvula (Stenhammar, 1855) (Diptera, Sphaeroceridae) .............. Be
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 61
134. Stigmatomyces platensis Speg., Anal. Mus. Nac. Hist. Nat. B. Aires 29:
676(1917)
Paralimosina fucata (Rondani, 1880) (Diptera, Sphaeroceridae) ....................... Be
Paralimosina subcribrata (Rohacek, 1977) ........................................................ Be
135. Symplectromyces vulgaris (axt.) axt., Mem. Am. Acad. Arts Sci. 13: 315
(1908)
Philonthus sp. (Coleoptera, Staphylinidae) ....................................................... Be
Quedius curtipennis Bernhauer, 1908 (Coleoptera, Staphylinidae) .................... Be
Quedius fuliginosus (Gravenhorst, 1802) ........................................................... Be
Quedius fumatus (Stephens, 1833).................................................................... Be
Quedius lateralis (Gravenhorst, 1802) ............................................................... Nl
Quedius levicollis (Brullé, 1832) .......................................................................Bea
Quedius maurorufus (Gravenhorst, 1806) ......................................................... Nl
Quedius mesomelinus (Marsham, 1802) ...................................................... Be, Nl
Quedius sp. ...................................................................................................... Be
a Host as Quedius tristis (Gravenhorst, 1802) in De Kesel et al. (2020).
136. Teratomyces actobii axt. Proc. Am. Acad. Arts Sci. 29: 98 (1894)
Gabrius nigritulus (Gravenhorst, 1802) (Coleoptera, Staphylinidae) ................. Be
Gabrius sp. ....................................................................................................... Be
137. Teratomyces philonthi axt., Proc. Am. Acad. Arts Sci. 35: 432 (1901)
Gabrius nigritulus (Gravenhorst, 1802) (Coleoptera, Staphylinidae) ................. Be
Gabrius trossulus (Nordmann, 1837) ............................................................... Nla
Gabrius sp. ....................................................................................................... Be
Quedius nitipennis (Stephens, 1833) (Coleoptera, Staphylinidae) ..................... Be
Quedius sp. ...................................................................................................... Be
a Host as Philonthus trossulus Nordmann, 1837 in Middelhoek (1943a).
138. Troglomyces manfrediae S. Colla [as ‘manfredii’], Nuovo G. Bot. Ital. 39:
451(1932)
Julida sp. indet. ................................................................................................ Be
139. Troglomyces triandrus Santam. & Engho, Organ. Divers. Evol. 15: 253 (2015)
Archiboreoiulus pallidus (Brade-Birks, 1920) (Julida, Blaniulidae) ..................... Be
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
62
140. Zodiomyces vorticellarius axt., Proc. Am. Acad. Arts Sci. 25: 263 (1891)
Helochares punctatus (Sharp, 1869) (Coleoptera, Hydrophilidae) ...................... Nl
Helochares sp. ................................................................................................... Be
Doubtful records and combinations
Laboulbenia elegans axt. on Harpalus anis (Schrank, 1781) (Coleoptera, Carabi-
dae) [as Harpalus aeneus (Fabricius, 1775)] (Middelhoek 1949). is material could
not be veried since the Middelhoek collection is currently untraceable, but it likely
represents L. coneglianensis. Laboulbenia coneglianensis is reported from species of Har-
palus Latreille, 1802 and Ophonus Dejean, 1821 in Europe, whereas L. elegans is thus
far only conrmed from New England, USA (axter 1890, 1896).
Laboulbenia agellata [as Laboulbenia elongata axt.] on Calathus erratus (Sahl-
berg, 1827) (Coleoptera, Carabidae) (Middelhoek 1947b). e material is incom-
plete and impossible to verify. Given the host, it is doubtful that this report represents
L.agellata. Possibly it is L. calathi T. Majewski, which is already known from the
Netherlands (Haelewaters et al. 2012b).
Laboulbenia agellata on Pterostichus nigrita (Paykull, 1790) (Coleoptera, Carabi-
dae) (Meijer 1975). is record possibly represents L. pseudomasei axt. but we can-
not verify because the material of Meijer is untraceable. Pterostichus nigrita is routinely
reported as host to L. pseudomasei, not L. agellata (axter 1899; Scheloske 1969;
Majewski 1994; Santamaría 1998; De Kesel et al. 2020). Both species are easily distin-
guished with morphological characters.
Laboulbenia pedicellata on Trechus quadristriatus (Schrank, 1781) (Coleoptera, Car-
abidae) (Meijer 1975). is would be the only worldwide record of L. pedicellata on a
species of Trechus Clairville, 1806 and thus is likely incorrect. Laboulbenia pedicellata is
generally reported on species of Bembidion Latreille, 1802 sensu lato (Coleoptera, Car-
abidae) and Dyschirius Bonelli, 1810 (Coleoptera, Carabidae) (Haelewaters et al. 2019a).
Discussion
New species and new records
In this paper, we describe two new species of Laboulbeniales based on the combination
of molecular data, morphology, and ecology (host association). ese are Hesperomyces
halyziae on Halyzia sedecimguttata in Belgium and the Netherlands, and Laboulbenia
quarantenae on Bembidion biguttatum in Belgium. Additionally, Laboulbenia aubryi
and Rhachomyces spinosus are newly reported from Belgium. Seven previously described
species of Laboulbeniales are reported for the rst time from the Netherlands: Chi-
tonomyces melanurus, Euphoriomyces agathidii, Laboulbenia fasciculata, Laboulbenia me-
tableti, Laboulbenia pseudomasei, Rhachomyces canariensis, and Stigmatomyces hydrelliae.
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 63
e report of L. aubryi from Belgium is only the third one from Europe. Laboul-
benia aubryi was thus far only recorded from India, Nepal, Poland, and Spain (type).
Reported hosts are species in Amara Bonelli, 1810 (= Bradytus Stephens, 1827, = Lei-
ronotus Ganglbauer, 1892) (Santamaría et al. 1991; Santamaría 1998; Majewski 1999),
a diverse genus that is only exceptionally reported with Laboulbeniales (Santamaría
et al. 1991). Scheloske (1969) mentioned L. agellata on Amara plebeja (Gyllenhal,
1810), but considered it an accidental host (“Zufallswirt”). Moreover, based on its
simple outer appendage, L. aubryi can easily be separated from L. agellata. e clos-
est related species, morphologically speaking, is L. argutoris Cépède & F. Picard, but
L.aubryi can be separated from it by the insertion cell that is free from the perithecium
wall and by the structure of its inner appendage (Santamaría 1998).
Rhachomyces spinosus was recently described from Spain (Santamaría et al. 2020).
e most characteristic feature of this species is the spinous process on the second
cell of the primary appendage, absent in similar species R. lavagnei (F. Picard) W.
Rossi and R. sciakyi W. Rossi. e reported host for R. spinosus in both Belgium and
Spain is Syntomus foveatus (Coleoptera, Carabidae). Rhachomyces lavagnei is found on
Microlestes spp. and R. sciakyi on Pseudomesolestes sp. All these hosts are placed in the
subtribe Dromiusina (Harpalinae, Lebiini); it is possible that these species of Rha-
chomyces have a high degree of host specicity, which will only come to light as more
material will be collected.
Chitonomyces melanurus is easily recognized from other congeneric species by the
apically hooked, dark brown to blackish basal cell of its primary appendage. Nine
species of Chitonomyces Peyr. occur in Europe, all of them occupying a specic posi-
tion of the host integument. Chitonomyces melanurus grows almost exclusively on the
upper margin of the left elytron of Laccophilus Leach, 1815 water beetles (Coleoptera,
Dytiscidae). It has thus far has been reported in Europe from Austria (type), Belgium,
Croatia, Finland, France, Germany, Hungary, Italy, Poland, Spain, Ukraine, United
Kingdom; also found in Asia and Africa (Bánhegyi 1960; Huldén 1983; Santamaría et
al. 1991; Majewski 1994; De Kesel and Werbrouck 2008; Rossi 2018).
e Dutch report of E. agathidii is found on Agathidium laevigatum, the host spe-
cies from which the type was described (Maire 1920). Euphoriomyces agathidii is thus
far found on members of Agathidium Panzer, 1796, Amphicyllis Erichson, 1845, and
Cyrtusa Erichson, 1842 (Coleoptera, Leiodidae) in Bulgaria, Germany, Italy, Morocco
(type), Poland, South Korea, Spain, and Sweden (Huldén 1983; Majewski 1994; Lee
et al. 2007; Rossi et al. 2018). Our material is consistent with E. agathidii, with two
mature perithecia at one side and a third, immature perithecium at the other side of
the receptacular axis.
Laboulbenia fasciculata is recognized by the receptacular cell V, which proliferates
upwards in a series of 4–8 superposed cells V’ gradually decreasing in size. Each of these
cells V’ gives rise to a small trapezoidal cell that carries an appendage consisting of cells
separated by dark and constricted septa. is species is very widespread, with reports
across Europe, in Africa, Asia, and North and South America. Hosts are members of
Carabidae, often Chlaenius Bonelli, 1810 (subfamily Harpalinae) and Patrobus Dejean,
1821 (subfamily Trechinae), but also several other genera in subfamilies Cicindelinae,
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
64
Brachininae, Harpalinae, Nebriinae, Omophroninae, Patrobinae, and Trechinae (San-
tamaría et al. 1991). e reports on Omophron spp. are sometimes considered a form
of L. fasciculata but this is not accepted by all (Spegazzini 1914; Majewski 1994; but
Santamaría 1998).
e status of L. metableti as a separate species has been disputed. Formally syn-
onymized with L. notiophili by Rossi and Santamaría (2006), De Kesel et al. (2020)
reinstated L. metableti as a separate species based on characteristics of the appendage
system. is species has a European distribution, with reports in Andorra, Austria,
Belgium, Finland, Germany (type), Hungary, Italy, Poland, Russia, and the United
Kingdom (reviewed in Rossi and Santamaría 2006). Hosts are species of Syntomus
Hope, 1838 (= Metabletus Schmidt-Goebel, 1846) (Coleoptera, Carabidae, Harpali-
nae, Lebiini). We propose using molecular characters to resolve the debate given the
taxonomic confusion of species of Laboulbenia on European hosts in the Lebiini tribe:
L. baetica Balazuc, L. blanchardii Cépède, L. cymindicola Speg., L. metableti, L. notio-
phili, and L. pulchella Speg.
Laboulbenia pseudomasei is recognized by cell V that has an internal convex margin
and is separated from the perithecium (Villarreal et al. 2010). Cell V sometimes pro-
liferates into a simple or divided branch that grows upwards between the perithecium
and insertion cell (Majewski 1994; Santamaría 1998). Rossi and Weir (1997) illus-
trated that L. pseudomasei can be morphologically highly variable even on a single host
insect. Also in the newly reported material from the Netherlands, L. pseudomasei was
variable, with the thallus from the right elytron without proliferation of cell V, and the
thallus from the prosternum with proliferation of cell V. e geographic distribution
of L. pseudomasei is problematic; many old records are unillustrated and the specimens
are not preserved (Rossi and Weir 1997).
Rhachomyces canariensis was described from Tenerife (axter 1900) and has since
been reported from several countries in Europe and North Africa, Madeira, and the
Canary Islands, always associated with species of Trechus Clairville, 1806 (Coleoptera,
Carabidae) (Arndt and Santamaría 2004). Majewski (1994) noted the variability of
this species and Tavares (1985) suggested material from large geographic distances to
the type locality be segregated into a separate taxon.
e only species of Laboulbeniales found on Hydrellia Robineau-Desvoidy,
1830 ies (Diptera, Ephydridae) is S. hydrelliae. axter (1901) described it from
Kittery Point in Maine, USA (axter 1901) and it has since then been reported in
Finland, France, Italy, Poland, Portugal, Russia, the United Kingdom (Santamaría
and Rossi 1993, Weir and Rossi 1995), and New Zealand (Hughes et al. 2004). e
new report from the Netherlands is the rst one on the European continent in 25
years. Stigmatomyces hydrelliae is recognized by its straight appendage with sterile
basal cell and stout antheridia, the spiralled cell walls of the perithecium, and the
rounded perithecial apex with one of the lip cells forming a slender, bluntly pointed
projection. Hughes et al. (2004) noted that S. hydrelliae thalli from New Zealand are
dierent in their perithecial wall cells not being spiralled and lacking apical projec-
tions at the perithecial apex.
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 65
Checklist
e current list of thallus-forming Laboulbeniomycetes from Belgium and the Neth-
erlands includes 140 species. Sixty-three species have been found in both countries. A
total of 118 species are found in Belgium, and 85 species in the Netherlands. Of the
140 species in the checklist, 55 have not (yet) been reported from the Netherlands,
and 22 species have not (yet) been reported from Belgium. Laboulbeniales research in
both Belgium and the Netherlands has also resulted in the discovery of new taxa; over
the years, 16 species have been described based on material from Belgium and/or the
Netherlands (Table 3). It is remarkable that we keep nding undescribed species in two
of the most urbanized countries in the world. e reason for this can be found in the
fact that Laboulbeniomycetes are severely understudied; only a handful of researchers
work on these fungi. In addition, some of the most recently described species are the
result of previously unavailable molecular data, long-term study of humid habitats,
and focus on unexplored niches.
is checklist is based on fungal records obtained from at least 283 host species
(including only those identied to species level). To increase the number of thallus-
forming Laboulbeniomycetes known from Belgium and the Netherlands, future re-
search should focus on screening Acari (with Rickia), Blattodea (Herpomyces–especially
in the Netherlands), Coleoptera (many genera), Corixidae (Coreomyces), Diplopoda
(Diplopodomyces, Troglomyces), Diptera (Stigmatomyces), Hebridae (Tavaresiella,
Triceromyces), and Mallophaga (Trenomyces). Within Coleoptera, the beetles, aquatic
and semi-aquatic hosts, such as Dytiscidae (Chitonomyces), Hydraenidae (Autoicomyces,
Hydrophilomyces, ripomyces), and Hydrophilidae (Chaetarthriomyces, Eusynaptomy-
ces) deserve special attention. More genera of Laboulbeniales that are currently not yet
Table 3. Species of Laboulbeniales described based on type material from Belgium (Be) and the Neth-
erlands (Nl).
Country Laboulbeniales species Host species Host classication Reference
Nl Asaphomyces tubanticus [as
Barbariella tubantica]
Catops nigricans Coleoptera, Leiodidae Middelhoek (1949)
Nl Cantharomyces elongatus Syntomium aeneum Coleoptera, Staphylinidae Haelewaters and De Kesel (2013)
Nl Bordea denotata Bibloporus bicolor Coleoptera, Staphylinidae Haelewaters et al. (2014)
Be Cryptandromyces euplecti Euplectus sanguineus Coleoptera, Staphylinidae Santamaría (2001)
Be, NL Diphymyces kaaistoepi Choleva cisteloides, C. fagniezi Coleoptera, Leiodidae De Kesel and Haelewaters (2019)
Be, NL Hesperomyces halyziae Halyzia sedecimguttata Coleoptera, Coccinellidae is paper
Nl Laboulbenia barbara Philonthus punctus Coleoptera, Staphylinidae Middelhoek (1943a)
Be Laboulbenia quarantenae Bembidion biguttatum Coleoptera, Carabidae is paper
Be Laboulbenia elaphri Elaphrus cupreus Coleoptera, Carabidae Spegazzini (1915)
Be Laboulbenia hyalopoda Paradromius linearis Coleoptera, Carabidae De Kesel (1998)
Be, NL Laboulbenia littoralis Caus xantholoma Coleoptera, Staphylinidae De Kesel and Haelewaters (2014)
Nl Mimeomyces zeelandicus Heterothops binotatus Coleoptera, Staphylinidae Middelhoek (1943a)
Be Peyritschiella heinemanniana Xantholinus longiventris Coleoptera, Staphylinidae De Kesel (1999)
Be Phaulomyces simplocariae Simplocaria semistriata Coleoptera, Byrrhidae De Kesel (1994)
Be, NL Rickia laboulbenioides Cylindroiulus latestriatus Julida, Julidae De Kesel et al. (2013)
Be Troglomyces triandrus Archiboreoiulus pallidus Julida, Blaniulidae Engho and Santamaría (2015)
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
66
recorded from either Belgium or the Netherlands, could be discovered on Anthicidae
(Dioicomyces), Ptiliidae (Siemaszkoa), Silvanidae (Cucujomyces), Staphylinidae (Amor-
phomyces, Diplomyces, Dipodomyces, Haplomyces, Mimeomyces, Sphaleromyces), and Ten-
ebrionidae (Dimeromyces).
As Laboulbeniomycetes research progresses, lesser known host groups will need to
be incorporated into our studies. is will eventually require intensied collaborations
with specialist entomologists, as well as screening museum insect collections and the
use of dierent collecting methods. at dierent sampling techniques have an im-
pact on Laboulbeniales studies may be illustrated by our work with Rickia wasmannii
Cavara. Based on pitfall trapping, Haelewaters et al. (2015a) reported R. wasmannii
from three host species: Myrmica sabuleti Meinert, 1861 (parasite prevalence 38%),
M.scabrinodis Nylander, 1846 (11%), and M. ruginodis Nylander, 1846 (0.55%). Di-
rect sampling from a M. scabrinodis nest at the same locality in the Netherlands, how-
ever, resulted in a 100% prevalence (De Kesel et al. 2016).
Finally, undersampled habitats have been cited repeatedly as one of the main sources
to nd undescribed fungi (e.g., Blackwell 2011, Hawksworth and Lücking 2017, Wijaya-
wardene et al. 2020). is is especially true for the Laboulbeniomycetes. Sampling from
dung, fresh and brackish water, animal nests, caves, carcasses, and rotting plant debris
has greatly contributed to discoveries in this eld of research, not only adding to num-
bers of described species but also building on our understanding of the ecology of these
minute fungi. For example, Piegler et al. (2016) sampled ants and their associates from
ant nests and, for the rst time since its description (Cavara 1899), R. wasmannii was
observed on hosts other than Myrmica, including inquiline mites and a y larva. A sur-
vey of Laboulbeniales from coprophilic beetles on Galloway dung in Belgium resulted in
two reports of species that until then had only been found in Poland, thus representing a
large geographical range expansion (De Kesel 2010). And signal craysh traps in nature
reserve ‘De Kaaistoep’ have thus far revealed an undescribed species of Diphymyces (De
Kesel and Haelewaters 2019) and more material is awaiting detailed study.
Key to Laboulbeniomycetes from Belgium and the Netherlands
1 Dioecious; on Blattodea (cockroaches) ............................... 50 (Herpomyces)
alli mostly monoecious; on other host groups ......................................... 2
2 Perithecial wall cells numerous, subequal, always ≥ 6 cells per vertical row ....3
Perithecial wall with < 6 cells per vertical row ............................................7
3 Receptacle uniseriate, composed of numerous superposed cells ................... 5
Receptacle multiseriate, often massive ......................................................... 4
4 Receptacle turbinate, with apical depression holding numerous sterile ap-
pendages, stalked perithecia and antheridial branchlets; on Hydrophilidae ...
............................................................................Zodiomyces vorticellarius
Receptacle not turbinate, bearing numerous perithecia and appendages later-
ally; on Carabidae and Staphylinidae ......................Euzodiomyces lathrobii
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 67
5 Perithecium with an apical, darkened rostrum; receptacle with 4–5 cells; on
Ptiliidae ....................................................................Kainomyces rehmanii
Perithecium without a rostrum; receptacle with > 5 cells .............................6
6 Perithecium long-necked, without lobes or ne appendages on the perithecial
wall; on Hydrophilidae .............................. Rhynchophoromyces anacaenae
Perithecium without long neck, ostiolum with 4 ne ligulae, lower wall bear-
ing slender ramied appendices; on Dryopidae ...........Helodiomyces elegans
7 Antheridia simple, ask shaped; release of spermatia through small necks ....8
Antheridia grouped into a compound structure with wall ......................... 44
8 Sterile appendages unicellular with black basal septum; antheridia small, al-
ways with black basal septum; receptacle formed by 3 vertical tiers of cells
(not always clear), at least one tier partly or entirely anking the perithe-
cium ...........................................................................................52 (Rickia)
Not this combination .................................................................................9
9 Suprabasal cell of the receptacle (cell II) produces multi-celled secondary ap-
pendages; the latter supporting a perithecium (with cell VI) at their base .....
..................................................................................Compsomyces lestevae
Cell II not producing secondary appendages ............................................. 10
10 Perithecial wall with an elongated accessory cell along its outer venter; unicel-
lular outgrowths are formed above the foot; on Cercyon (Hydrophilidae) .... 11
Perithecium without accessory cell; no such outgrowths above the foot ....12
11 Lower receptacular cells isodiametric; perithecium neck more or les
straight ............................................................Hydrophilomyces cf. gracilis
Lower receptacular cells attened; perithecium neck strongly curved ............
..................................................................... Hydrophilomyces cf. hamatus
12 Cell VII and basal cells of the perithecium clearly visible in mature perithe-
cia ............................................................................................................. 13
Cell VII and basal cells of the perithecium not visible in mature perithecia ....40
13 Receptacle produces longitudinal septa, leading to a suprabasal complex with
numerous secondary appendices ...............................................................14
Receptacle stays a series of superposed cells, rarely forming longitudinal septa,
not forming a suprabasal complex or secondary appendices ......................20
14 Receptacle composed of a series of superposed cells (4–5 or more), each form-
ing on one side a basal cell with numerous, fairly large, pigmented and mul-
ticellular appendages; thalli usually with only one perithecium .....................
........................................................................................ 56 (Rhachomyces)
Not with these features .............................................................................15
15 allus hyaline; appendages not in bunches; On Cholevinae (Leiodidae)......
.............................................................................. Asaphomyces tubanticus
allus moderately to deeply pigmented in some parts; appendages appear in
bunches on the receptacle .........................................................................16
16 Receptacle asymmetrical ........................................................................... 17
Receptacle mostly symmetrical ..................................................................18
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
68
17 Antheridia in lateral series on fertile appendages; dorsal and ventral cell of the
triangular receptacle supporting a series of appendages and their basal cells;
perithecium stalked by elongated cells VI and VII ...... Idiomyces peyritschii
Antheridia never organized in lateral series; appendages not in series; recepta-
cle 5-celled; cells VI and VII relatively short ..................... 62 (Laboulbenia)
18 Appendages with pointed-curved tips, darkened septa; antheridia terminal,
ask shaped, not forming ramications with age ................19 (Teratomyces)
Appendages with rounded tips, with series of intercalary antheridia, the latter
ramifying into new appendages with age ..............Symplectromyces vulgaris
19 Cells I and II from receptacle becoming brown with age; basal cells of ap-
pendages with laterally aligned antheridia/septa ........ Teratomyces philonthi
Cell I hyaline, contrasting with a deep blackened cell II; basal cells of append-
ages without such laterally aligned septa ........................ Teratomyces actobii
20 Primary appendage bicellular, both cells separated by a dark constricted sep-
tum; antheridium below the primary appendage; on aquatic Coleoptera .... 21
Primary appendage more developed .......................................................... 22
21 All 4 vertical tiers of the perithecial wall have 4 cells each .............................
.....................................................................................109 (Chitonomyces)
Only 2 vertical tiers of the perithecial wall have 4 cells, the others have 6 cells;
on Haliplidae ............................................................ Hydraeomyces halipli
22 Receptacle composed of ≥ 4 cells .............................................................23
Receptacle composed of ≤ 3 cells ..............................................................26
23 Primary receptacle composed of a chain of cells (≥ 3) ................................24
Primary receptacle composed of cells I and II, entire receptacle with ve
cells .................................................................................. 62 (Laboulbenia)
24 Perithecium with obtuse apex and inconspicuous neck ..............................
................................................................................Misgomyces dyschirii
Perithecium with long neck and dierentiated venter ...............................25
25 Antheridia sessile, develop as corner cells of the primary appendage; Recepta-
cle cells attened, broadening upwards .......Ecteinomyces trichopterophilus
Antheridia not sessile but formed on lateral branchlets; receptacle cells elon-
gate .................................................................. Botryandromyces heteroceri
26 Cell III attened and entirely appressed against the perithecium; on Julida ...
........................................................................................113 (Troglomyces)
Cell III dierent; on Hexapoda ................................................................. 27
27 On Coleoptera ..........................................................................................29
Not on Coleoptera .................................................................................... 28
28 Basal cell of appendage dark; perithecial apex with outgrowths; on Forcula
(Dermaptera, Forculidae) .......................................Distolomyces forculae
On Diptera ....................................... 114 (Fanniomyces & Stigmatomyces)
29 Primary appendage easily breaking o at its strongly narrowed basal cell; on
Kateretidae........................................................ Aphanandromyces audisioi
Primary appendage persistent ....................................................................30
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 69
30 Receptacle cells (I, II, III) more or less superposed .................................... 31
Receptacle cells not superposed (cell I and III touching) ........................... 39
31 Distal cell of primary appendage is a simple antheridium, with eerent neck
and spine ........................................................................... Bordea denotata
Primary appendage without such a single and terminal antheridium .........32
32 Antheridial structures are born on corner cells of appendage axis cells .......33
Antheridial branches not born from corner cells .......................................36
33 Basal (m, n, n’) and stalk cells (VI, VII) of the perithecium small (together
<25 µm long); on Hydrophilidae ....Chaetarthriomyces crassiappendicatus
Basal (m, n, n’) and stalk cells (VI, VII) ≥ 25 µm long; on Staphylinidae .... 34
34 Cell III mostly without antheridial branches, with or without perithecium;
cells VI and VII of similar length .......................... Stichomyces conosomatis
Cell III always with antheridial branches, never with perithecium; cell VI
much taller than cell VII ...........................................................................35
35 allus forms perithecia and corner cells on one side (anteri-
or) ...........................................................................Rhadinomyces pallidus
allus forms perithecia and corner cells on both sides (anterior and poste-
rior) ........................................................................ Rhadinomyces cristatus
36 Primary appendage simple, composed of numerous similar superposed
cells .........................................................................124 (Cryptandromyces)
Primary appendage branched .................................................................... 37
37 Cell VI adnate to cell II; exclusively on Cholevinae (Leiodidae) ....................
................................................................................ Diphymyces kaaistoepi
Cell VI supported by cell II; mostly on Staphylinidae, rarely on Cholevinae
(Leiodidae) ...............................................................................................38
38 Cell I tall and elongated, cell II attened............... Mimeomyces zeelandicus
Cell I very short, cell II not attened (isodiametric) .... 126 (Corethromyces)
39 Perithecial tip with prominent ostiolar lips and lobes; appendage short, with
sessile lateral antheridia on each cell; fresh thalli often greenish-yellow; on
Coccinellidae ................................................................127 (Hesperomyces)
Perithecial tip without such lobes; appendage long, with lateral antheridia on
few cells; not on Coccinellidae ................................ 124 (Cryptandromyces)
40 Receptacle between foot and cell VI with ≥ 3 cells. ...................................41
Receptacle between foot and cell VI with 2 cells; foot entirely black .............
...........................................................................Phaulomyces simplocariae
41 Receptacle a series of superposed cells, many of which laterally producing
perithecia and/or appendages ................................ Euphoriomyces agathidii
Receptacle a series of superposed cells without lateral cells bearing perithecia
and appendages ......................................................................................... 42
42 Receptacle with attened and nely appendiculate cells above cell III; Foot
entirely black; on Corixidae (Hemiptera) .................... Coreomyces arcuatus
Receptacle without such attened cells above cell III; foot with a small black-
ish dot; on Ptiliidae ...................................................................................43
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
70
43 e appendage is a prolongation of the receptacle axis, the perithecium is
lateral ........................................................................... Siemaszkoa ptenidii
e perithecium is often terminal and in continuation with the receptacular
axis ............................................................................... Siemaszkoa fennica
44 Cell I laterally extending and supporting a series of cells derived from cell II;
thallus dioecious ..........................................Dimorphomyces myrmedoniae
Cell I not laterally extending; thallus monoecious ..................................... 45
45 Primary receptacle composed of a chain of ≥ 3 cells .... Misgomyces dyschirii
Primary receptacle not a chain of cells ....................................................... 46
46 Primary appendage fertile, with a compound antheridium ........................47
Primary appendage sterile or absent .......................................................... 49
47 Compound antheridium with eerent neck and tall cell on the outer side; on
Carabidae .........................................................Eucantharomyces stammeri
Compound antheridium dierent, never with eerent neck; on Staphylinidae
and Dryopidae (= Parnidae) ......................................................................48
48 Primary appendage is entirely transformed into a compound antheridium,
with spine .................................................................... Haplomyces texanus
Compound antheridium is an intercalary structure of the primary append-
age ............................................................................. 129 (Cantharomyces)
49 Receptacle formed by 3 horizontal tiers of cells; antheridia compound, sessile,
often on the median series; sterile appendages unicellular ....134 (Peyritschiella)
Receptacle dierently organized; sterile appendages multicellular; antheridial
structure stalked, large .................................................140 (Monoicomyces)
50 Secondary receptacle (female thallus) without concentrically organized cells .
....................................................................................Herpomyces ectobiae
Secondary receptacle a series of concentrically organized and attened cells ...
.................................................................................................................. 51
51 Secondary receptacle ≤ 80 µm long, with rounded apex and partially dark-
ened cells ................................................................. Herpomyces stylopygae
Secondary receptacle ≥ 100 µm long, with pointed apex and without dark
pigmentations ...................................................... Herpomyces periplanetae
52 Perithecium almost entirely embedded in the receptacle ....Rickia peyerimhoi
Anterior part of the perithecium free ........................................................53
53 On Diplopoda .......................................................................................... 54
On other arthropods ................................................................................. 55
54 Dorsal margin of the perithecium free in its upper third; anterior series of
receptacle consisting of 2(–3) cells ...........................Rickia laboulbenioides
Dorsal margin of the perithecium only free at the apex; Anterior series of
receptacle consisting of > 2 cells ....................................... Rickia dendroiuli
55 Cell I 12–18 µm long; on Staphylinidae ...............................Rickia proteini
Cell I 60–90 µm long; on Myrmica (Hymenoptera, Formicidae) ..................
....................................................................................... Rickia wasmannii
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 71
56 Primary appendage hyaline, 3-celled, dierent from other appendages; On
Syntomus (Carabidae) ................................................ Rhachomyces spinosus
Primary appendage pigmented, identical to secondary appendages ...........57
57 Receptaculum between cells I and VI usually with < 6 cells, sterile appendages
very long; on Lathrobium (Staphylinidae) ................Rhachomyces pilosellus
Receptaculum between cells I and VI composed of ≥ 6 cells; sterile append-
ages do not exceed perithecial apex ...........................................................58
58 Cells of the B-appendages of unequal length .............................................59
Cells of the B-appendages of similar to equal length .................................60
59 B-appendages elongate, slender, tapering upwards; On Philonthus (Staphyli-
nidae) ................................................................Rhachomyces philonthinus
B-appendages short, stout, width broad rounded apex; On alassophilus
(Carabidae) ........................................................ Rhachomyces tenenbaumii
60 Cell VI elongate and situated in the median to subapical part of the (second-
ary) receptacle; On Othius (Staphylinidae) .................Rhachomyces furcatus
Cell VI short, distally on the (secondary) receptacle; on Carabidae ........... 61
61 Perithecial apex with black spots; terminal cell of the B-appendages widest in
the middle; on Acupalpus (Carabidae) .................. Rhachomyces lasiophorus
Perithecial apex hyaline; terminal cell of B-appendages cylindrical, usually
proliferating; on Trechus (Carabidae) .....................Rhachomyces canariensis
62 Insertion cell absent .................................................................................. 63
Insertion cell present ................................................................................. 65
63 Appendages with large basal cells and dark septa; on Carabidae ....................
.............................................................................. Laboulbenia fasciculata
Appendages liform, with ne basal cells and dark septal on Gyrinidae ....64
64 Perithecium with two hyaline apical outgrowths, one straight one hooked ....
............................................................................... Laboulbenia gyrinicola
Both perithecial outgrowths with black spots, irregularly shaped ...................
................................................................................... Laboulbenia fennica
65 On Carabidae ........................................................................................... 66
On Staphylinidae ....................................................................................103
66 Insertion cell free ...................................................................................... 67
Insertion cell attached to the posterior margin of the perithecium (not free) ....72
67 Foot almost hyaline with only a small black dot ......Laboulbenia hyalopoda
Foot entirely black ....................................................................................68
68 Cell V as tall as cell IV ..............................................Laboulbenia clivinalis
Cell V smaller than cell IV ........................................................................ 69
69 Outer appendage not branched .................................................................70
Outer appendage branched ................................. Laboulbenia pseudomasei
70 Inner appendage hardly branched, with a single antheridium........................
................................................................................. Laboulbenia lecoareri
Inner appendage branched, with multiple antheridia ................................71
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
72
71 Lower 4–5 cells of outer appendage deeply pigmented in their middle; osti-
olar papillae not conspicuous; on Syntomus (Carabidae) ................................
................................................................................ Laboulbenia metableti
Lower 4–5 cells of outer appendage evenly pigmented; ostiolar papillae con-
spicuous; on Amara (Carabidae) .................................. Laboulbenia aubryi
72 Cell V as tall as cell IV, or almost so .......................................................... 73
Cell V smaller than cell IV ........................................................................ 78
73 Outer wall of perithecium with knobs ............................ Laboulbenia egens
Outer wall of the perithecium without knobs ...........................................74
74 Outer appendage without dark septum, growing beyond the perithecium ....
.................................................................................... Laboulbenia ophoni
Outer appendage with at least one dark septum, not growing beyond the
perithecium ..............................................................................................75
75 Cells IV and V attened, broader than long; On Cillenus (Carabidae) ..........
.......................................................................... Laboulbenia lichtensteinii
Cells IV and V isodiametric or longer than broad ..................................... 76
76 allus and receptaculum poorly pigmented (yellow-amber); basal cell of out-
er appendage inated; on Pogonus (Staphylinidae) .... Laboulbenia slackensis
allus and receptaculum strongly pigmented; basal cell of outer appendage
not so inated ........................................................................................... 77
77 Cell III attened and oblique; posterior margin of cell IV longer than the one
from cell III; insertion cell extremely at and opaque ....Laboulbenia luxurians
Cell III not attened; posterior margin of cell IV equal or shorter than the one
from cell III; insertion cell well-formed and black ....Laboulbenia pedicellata
78 Outer appendage not growing beyond the perithecium ................................
............................................................................ Laboulbenia murmanica
Outer appendage growing beyond the perithecium ................................... 79
79 Outer appendage branched ....................................................................... 80
Outer appendage not branched .................................................................90
80 Cell IV very long, often with a conspicuous dorso-apical bump, sometimes
divided ......................................................................................................81
Cell IV not so long, never divided, without dorso-apical bump ................82
81 Outer appendage with > 2 branches; on Stenolophus (Carabidae) ..................
............................................................................ Laboulbenia anoplogenii
Outer appendage consisting of 2 branches; on Acupalpus (Carabidae) ...........
................................................................................. Laboulbenia acupalpi
82 Insertion cell on or above the middle of the perithecium; inner appendage less
developed than outer appendage ...............................................................83
Insertion cell below the middle of the perithecium ...................................84
83 allus and receptaculum pale; septa from basal cells of outer appendage not
darkened; on Paranchus albipes (Carabidae).................... Laboulbenia collae
allus and receptaculum strongly pigmented; septa from basal cells of outer
appendage darkened .................................................. Laboulbenia vulgaris
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 73
84 Outer side of the base of outer appendage strongly darkened .................... 85
Outer side of the base of outer appendage not or only very slightly darkened
................................................................................................................. 87
85 allus pale brown; appendages numerous, with tapering and pointed apices;
on Dicheirotrichus (Carabidae) ..................................... Laboulbenia giardii
allus deep brown; appendages not so numerous, not tapering, with round-
ed apices ...................................................................................................86
86 Septum II/III clearly shorter than septum II/VI; cell V clearer than surround-
ing structures; on Harpalus and Ophonus (Carabidae) ...................................
........................................................................ Laboulbenia coneglianensis
Septum II/III nearly as long as septum II/VI; cell V not much paler than sur-
rounding structures; on Brachinus (Carabidae) ............Laboulbenia rougetii
87 allus often bent, anterior side of the thallus concave ............................89
allus not so bent, anterior side of the thallus fairly straight ....................88
88 Insertion cell near the base of the perithecium; outer appendage often com-
posed of 4–6(–8) branches, resulting from successive dichotomies starting at
the suprabasal cell ...............................................Laboulbenia quarantenae
Insertion cell not so deep; outer appendage branched once or twice, not as
dichotomies ............................................ Laboulbenia agellata sensu lato
89 Cell V quite small, less than half the length of cell IV; perithecium very slen-
der, subcylindrical (not a stable feature); on Harpalus (Carabidae) ................
........................................................................ Laboulbenia coneglianensis
Cell V longer, usually more than half the length of cell IV; perithecium more
ovate; on Elaphrus (Carabidae) .....................................Laboulbenia elaphri
90 Insertion cell located at or above the middle of the perithecium; adaxial side
of the perithecium half free .......................................................................91
Insertion cell located well below the middle of the perithecium; adaxial side
of the perithecium more than half free ......................................................95
91 Basal cells of outer appendage with darkened septa ................................... 93
Basal cells of outer appendage with normal septa ...................................... 92
92 Cell VI broader than long ..................................... Laboulbenia benjaminii
Cell VI longer than broad .........................................Laboulbenia argutoris
93 Basal cell of outer appendage inated; inner appendage growing beyond the
perithecium ................................................................. Laboulbenia inata
Basal cell of outer appendage normal; inner appendage never beyond perithe-
cium .........................................................................................................94
94 Inner appendage composed of a single antheridium supported by one basal
cell; on Asaphidion (Carabidae) .................................. Laboulbenia thaxteri
Inner appendage with ≥ 2 cells each supporting one or more antheridia .......
.................................................................................. Laboulbenia vulgaris
95 Outer appendage rotated relative to the perithecium; on Pterostichus diligens
(Carabidae) ............................................................ Laboulbenia kajanensis
Outer appendage not rotated .................................................................... 96
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
74
96 Inner appendage growing far beyond the perithecium ...............................97
Inner appendage not or hardly beyond the perithecium ............................ 98
97 Cell V clearly paler than surrounding cells and perithecium (III, IV
andVI) ..................................................................................................... 98
Cell V not paler than its surrounding cells .......................Laboulbenia leisti
98 Cell IV (and cell III) evenly and deeply pigmented ................................. 100
Cell IV (and cell III) hyaline or pigmented, their outer margins distinctly
more pigmented than inner margins .........................................................99
99 Cell VI longer than broad; thallus ≥ 230 µm long; on Calathus (Carabidae) ...
.......................................................................................Laboulbenia calathi
Cell VI isodiametric; thallus smaller; on Demetrias, Notiophilus and Paradro-
mius (Carabidae) .................................... Laboulbenia notiophili sensu lato
100 Inner appendage hardly branched, with a single antheridium; cell IV longer
than broad ................................................................ Laboulbenia lecoareri
Inner appendage branched, with multiple antheridia; cell IV isodiametrical ...
.................................................................................................................. 101
101 Cell V minute; upper margin of cell IV 4–6× the width of cell V ............102
Cell V larger; upper margin of cell IV only 1–2× the width of cell V ............
............................................................................Laboulbenia eubradycelli
102 Lower 4–5 cells of outer appendage deeply pigmented in their middle; lower
3–4 cells of both branches of the inner appendage each producing a short
straight branch .........................................................Laboulbenia metableti
Lower 4–5 cells of outer appendage evenly pigmented; inner appendage dif-
ferently constructed ............................... Laboulbenia notiophili sensu lato
103 Cell V as long as cell IV, or almost so ......................................................106
Cell V smaller than cell IV ...................................................................... 104
104 Insertion cell free from the perithecium; outer appendage branched .............
......................................................................................Laboulbenia dubia
Insertion cell attached to the posterior margin of the perithecium (not free);
outer appendage not branched ................................................................105
105 Outer appendage with dark septa between basal cells; insertion cell near the
base of the perithecium ............................................ Laboulbenia stilicicola
Outer appendage without dark septa at the basal cells; insertion cell near the
middle of the perithecium.........................................Laboulbenia atlantica
106 Outer appendage with at least one dark septum at the basal and suprabasal
cells .........................................................................................................108
Outer appendage without dark septa at the basal cells ............................. 107
107 Outer appendage forming a tuft of branches, posterior margins of both its
basal and suprabasal cell entirely darkened ................. Laboulbenia barbara
Outer appendage simple or forked once; posterior margin of suprabasal cell
of outer appendage with black remains of primary appendage ......................
................................................................................... Laboulbenia cristata
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 75
108 Cell II hyaline; one black septum above the basal cell of the outer append-
age ............................................................................ Laboulbenia littoralis
Anterior part of cell II pigmented black; black septa between all basal cells of
inner and outer appendage ....................................... Laboulbenia philonthi
109 Perithecium with conspicuous outgrowths (spikes, thorns) .....................110
Perithecium without outgrowths; receptacle with outgrowth...................111
110 Perithecial outgrowth, arising from the apical-most wall cell .........................
............................................................................. Chitonomyces paradoxus
Perithecial outgrowth lateral, arising from sub-apical wall cell .......................
................................................................................Chitonomyces aculeifer
111 Suprabasal cell of the receptacle (Ia) attened ......Chitonomyces bidessarius
Suprabasal cell of the receptacle (Ia) isodiametric ....................................112
112 Receptacular outgrowth hyaline, straight or arcuate ....Chitonomyces italicus
Receptacular outgrowth black, straight, with conspicuously hooked apex .....
............................................................................ Chitonomyces melanurus
113 Primary appendage with 1 antheridium, always situated in the lowest cell ....
.............................................................................. Troglomyces manfrediae
Primary appendage with 3 antheridia situated in the third, fourth and fth
cell ............................................................................ Troglomyces triandrus
114 Appendage branched ..............................................................................115
Appendage an unbranched axis ............................................................... 116
115 Basal cell of appendage small, pigmented; appendage cells normal ................
.......................................................................... Fanniomyces burdigalensis
Basal cell of appendage elongate, not pigmented; appendage cells elongated .
.......................................................................... Fanniomyces ceratophorus
116 Cell VI shorter than cell III; appendage consisting isodiametric to elongated
cells .........................................................................................................117
Cells III and VI equally long; appendage with dark basal cell, consisting of
attened cells; on Sphaeroceridae (Diptera) ............................................120
117 Venter of perithecium without protuberances .........................................118
Venter of perithecium with protuberances; on Ephydridae (Diptera) ............
............................................................................ Stigmatomyces hydrelliae
118 Appendage arcuated or sigmoid; perithecial neck shorter than the venter; on
Musca (Diptera, Muscidae) ..........................................Stigmatomyces baeri
Appendage not arcuated; perithecial neck longer than venter; on Drosophila
(Diptera, Drosophilidae) .........................................................................119
119 Perithecial neck as long as venter; appendage hyaline, its axis composed of 4
cells; On Drosophila subg. Sophophora (Diptera, Drosophilidae) ...................
............................................................................ Stigmatomyces majewskii
Perithecial neck 2× as long as venter; appendage brown, its axis composed of
6 cells; On Drosophila subg. Drosophila (Diptera, Drosophilidae) .................
...................................................................... Stigmatomyces entomophilus
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
76
120 Venter of perithecium without protuberances .........................................121
Venter of perithecium with protuberances ..............................................122
121 Perithecial basal cells elongated, longer than the appendage ..........................
.............................................................................Stigmatomyces limosinae
Perithecial basal cells not elongated, never longer than the appendage ..........
........................................................................... Stigmatomyces crassicollis
122 Perithecial apex abruptly becoming conical; appendage not proliferating ......
.............................................................................. Stigmatomyces platensis
Perithecial apex gradually tapering; appendage proliferating distally ........123
123 Perithecial venter with numerous knobs, below the neck and also down-
wards ............................................................Stigmatomyces minilimosinae
Perithecial venter with 4 knobs, only below the neck ....................................
........................................................................... Stigmatomyces divergatus
124 Basal cells of the primary appendage dark brown on the outer side ...............
........................................................................... Cryptandromyces euplecti
Primary appendage entirely hyaline .........................................................125
125 Cell I supporting cell II; thallus > 125 µm long ....Cryptandromyces elegans
Cell I supporting cells II and III; thallus < 100 µm long ...............................
.......................................................................Cryptandromyces bibloplecti
126 Basal cell of the receptacle with a black upgrowth; on Rugilus (Staphylini-
dae)..............................................................................Corethromyces stilici
Basal cell of the receptacle normal, without a black upgrowth; on Choleva
(Leiodidae) ............................................................. Corethromyces henrotii
127 Upper and lower lobes of perithecium of equal length, not exceeding perithe-
cial tip..............................................................Hesperomyces coccinelloides
Upper lobes long, exceeding the perithecial tip; lower lobes short ...........128
128 On Halyzia (Coleoptera, Coccinellidae) ................... Hesperomyces halyziae
– On Harmonia, Tytthaspis (Coleoptera, Coccinellidae) ...................................
...............................................................Hesperomyces virescens sensu lato
129 Cell II of receptacle blackened ...........................Cantharomyces denigratus
Cell II never black, at most brownish or with a black spot ......................130
130 Basal cell of primary appendage supporting a 300–415 µm long unbranched
series of 7–11 elongate cells..................................Cantharomyces elongatus
Primary appendage not so long ............................................................... 131
131 Primary appendage not ramied ............................. Cantharomyces italicus
Primary appendage with ramications ....................................................132
132 Primary appendage ramied above the suprabasal cell; basal cell not spheri-
cal ...........................................................................................................133
Primary appendage ramied above the basal cell, the latter spherical .............
............................................................................. Cantharomyces robustus
133 Antheridium small, not reaching the upper and lower septa of the basal cell
of the appendage ................................................ Cantharomyces platystethi
Antheridium larger, reaching the lower (and upper) septum of the basal cell
of the appendage ..................................................Cantharomyces orientalis
Checklist of Laboulbeniomycetes from Belgium and the Netherlands 77
134 Sterile appendages blackish brown, without conspicuous black septa ............
.....................................................................Peyritschiella heinemanniana
Sterile appendages hardly pigmented, with black septa ............................135
135 Lower horizontal tier of cells pigmented black, septa between cells obscured ...
.................................................................................................................. 136
Lower horizontal tier of cells not as pigmented, septa between cells visible....138
136 Distal part of receptacle broad; upper tier composed of > 20 cells .................
.................................................................................Peyritschiella biformis
Upper tier of the receptacle with < 15 cells; lateral sides of median tier each
produce one pigmented outgrowths (some reach the upper tier) .............137
137 allus with 2 perithecia .......................................... Peyritschiella furcifera
allus with (2–)3–12 perithecia ............................... Peyritschiella vulgata
138 Median cell of both lowest horizontal tiers as long as neighboring cells .........
.................................................................................... Peyritschiella protea
Median cell of both lowest horizontal tiers much larger than neighboring
cells, the latter becoming smaller outwards. ............................................139
139 Perithecium usually with auricula; ird and second horizontal tier of similar
length ...........................................................................Peyritschiella dubia
Perithecium without auricula; third horizontal tier longer than second .........
................................................................................. Peyritschiella princeps
140 Secondary receptacula unicellular ............................................................141
Secondary receptacula multicellular ........................................................146
141 Secondary receptacula pigmented ...........................................................145
Secondary receptacula hyaline ................................................................. 142
142 Cell VI much longer than perithecium ................ Monoicomyces myllaenae
Cell VI shorter than perithecium ............................................................143
143 Perithecium asymmetrical and bent; antheridium usually without apical
branchlets ............................................................ Monoicomyces matthiatis
Perithecium symmetrical, straight; antheridium with several apical branch-
lets ..........................................................................................................144
144 Basal cell of primary appendage at least partially pigmented, not narrowing
towards the apex ..................................................Monoicomyces homalotae
Basal cell of primary appendage entirely hyaline, narrowing towards the
apex ................................................................... Monoicomyces britannicus
145 Antheridial and primary appendages very long, reaching beyond the perithe-
cial apex .....................................................................Monoicomyces fragilis
Antheridial and primary appendages shorter, not reaching beyond the peri-
thecial apex ...........................................................Monoicomyces nigrescens
146 Secondary receptaculum pigmented ..................Monoicomyces bolitocharae
Secondary receptaculum hyaline .............................................................147
147 Secondary and antheridial appendages blackish brown .................................
.......................................................................... Monoicomyces californicus
Secondary and antheridial appendages hyaline or hardly pigmented, never
blackish brown ......................................................Monoicomyces invisibilis
D. Haelewaters & A. De Kesel / MycoKeys 71: 23–86 (2020)
78
Acknowledgements
D. Haelewaters received funding from the Uyttenboogaart-Eliasen Foundation and
through a SYNTHESYS+ grant (no. BE-TAF-151), nanced by the Horizon 2020
Research Infrastructures Programme of the European Commission. We want to take
the opportunity to recognize friends and collaborators for providing and/or identify-
ing infected insects throughout the years: Mario Amal, Berend Aukema, Wim Baert,
Michiel Boeken, Peter Boer, Johan Bogaert, Emiel H.M. Bouvy, André Braeckman, Sil-
vano Canzoneri, Luc De Bruyn, Gerdien De Kock, Kris Decleer, Wouter Dekoninck,
Albert J. Dees, Patrice Deramaix, Konjev Desender, Shanice De Weggheleire, Didier
Drugmand, Marc Dufrène, Henrik Engho, Ron Felix, Cyrille Gerstmans, Guy Hae-
ghebaert, Frederik Hanssens, Bart Horvers, Ivan Hoste, Sophie Janssens, Rudy Joc-
qué, Peter-Jan Keizer, Ilse Kranen, Mark Lammers, Violet Middelman, Jinze Noordijk,
eo M.J. Peeters, C. Lidwien Raak-van den Berg, Ivo Raemakers, Jindřich Roháček,
Menno Schilthuizen, Edwin Schuller, Dieter Slos, Piet Stoelen, Henk Spijkers, Tom
Van den Neucker, Douwe van der Ploeg, Joke van Erkelens, Marck Van Kerckvoorde,
ierry Vercauteren, Paul S. van Wielink, Tom Werbrouck, Jan Willem A. van Zuijlen,
Oscar Vorst, and Remco Vos. We should also acknowledge the Emergency Temporary
Access Service developed by HathiTrust in response to the COVID-19 pandemic and
subsequent lockdown, which made it possible for us to access necessary literature. Fi-
nally, we thank Alejandra Gutierrez, Patricia Kaishian, and Walter Rossi for reviewing
the paper and providing valuable suggestions for improvement.
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... While we have been able to sequence other species of Laboulbenia Haelewaters & De Kesel 2020), attempts to sequence L. ephemerolacuna have thus far been unsuccessful. Many specimens of T. surinamensis were observed along Pipeline Road but only about a dozen were collected for this study. ...
... We expect the number of Laboulbeniales reported for the country to increase in coming years given research that is being done by students at the Universidad Autónoma de Chiriquí, among others. As has also been suggested for continued Laboulbeniomycetes research in Belgium and the Netherlands (Haelewaters & De Kesel 2020), future work of Laboulbeniales in Panama will need to focus on (i) lesser-known host groups (e.g., aquatic insects, coprophilic Coleoptera, Diptera, Ptiliidae, Staphylinidae) and (ii) undersampled habitats such as animal nests, caves, and rotting fruit and plant debris. ...
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... (Ascomycota: Laboulbeniales, Laboulbeniaceae) is an obligate ectoparasite infecting adult ladybirds of over thirty species (Haelewaters et al., 2017). Recent integrative taxonomic analyses have revealed that H. virescens is in fact made up of many different species, each specifically adapted to its individual host Haelewaters and De Kesel, 2020;Crous et al., 2021). H. virescens completes its entire lifecycle on its living host; sexual spores divide mitotically to produce yellowish, multicellular, three-dimensional structures called thalli on the outside of any part of the host's body. ...
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... For Laboulbeniales, the bacteria could perhaps directly or indirectly provide nutrients necessary for growth into mature thalli, a hypothesis that, we think, merits further investigation. This can be more important for those Laboulbeniales lacking a haustorium, for mining nutrients from the host's integument (Tragust et al. 2016, Haelewaters & De Kesel 2020, Haelewaters et al. 2022. ...
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... Two fungal species now have pandemic-inspired monikers. Mycologists have named the one found in Netherlands as Diadolocovida claustri and the one found in US is being called Lauboulbenia quarantanae [9,10]. These names arose out of frustration of trying to keep research alive in a pandemic affected world. ...
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Denmark, Lasiosphaeria deviata on pieces of wood and herbaceous debris. Dominican Republic, Calocybella goethei among grass on a lawn. France (Corsica), Inocybe corsica on wet ground. France (French Guiana), Trechispora patawaensis on decayed branch of unknown angiosperm tree and Trechispora subregularis on decayed log of unknown angiosperm tree. Germany, Paramicrothecium sambuci (incl. Paramicrothecium gen. nov.) on dead stems of Sambucus nigra. India, Aureobasidium microtermitis from the gut of a Microtermes sp. termite, Laccaria diospyricola on soil and Phylloporia tamilnadensis on branches of Catunaregam spinosa. Iran, Pythium serotinoosporum from soil under Prunus dulcis. Italy, Pluteus brunneovenosus on twigs of broadleaved trees on the ground. Japan, Heterophoma rehmanniae on leaves of Rehmannia glutinosa f. hueichingensis. Kazakhstan, Murispora kazachstanica from healthy roots of Triticum aestivum. Namibia, Caespitomonium euphorbiae (incl. 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Russia, Colpoma junipericola from needles of Juniperus sabina, Entoloma cinnamomeum on soil in grasslands, Entoloma verae on soil in grasslands, Hyphodermella pallidostraminea on a dry dead branch of Actinidia sp., Lepiota sayanensis on litter in a mixed forest, Papiliotrema horticola from Malus communis, Paramacroventuria ribis (incl. Paramacroventuria gen. nov.) from leaves of Ribes aureum and Paramyrothecium lathyri from leaves of Lathyrus tuberosus. South Africa, Harzia combreti from leaf litter of Combretum collinum ssp. sulvense, Penicillium xyleborini from Xyleborinus saxesenii, Phaeoisaria dalbergiae from bark of Dalbergia armata, Protocreopsis euphorbiae from leaf litter of Euphorbia ingens and Roigiella syzygii from twigs of Syzygium chordatum. Spain, Genea zamorana on sandy soil, Gymnopus nigrescens on Scleropodium touretii, Hesperomyces parexochomi on Parexochomus quadriplagiatus, Paraphoma variabilis from dung, Phaeococcomyces kinklidomatophilus from a blackened metal railing of an industrial warehouse and Tuber suaveolens in soil under Quercus faginea. Svalbard and Jan Mayen, Inocybe nivea associated with Salix polaris. Thailand, Biscogniauxia whalleyi on corticated wood. UK, Parasitella quercicola from Quercus robur. USA, Aspergillus arizonicus from indoor air in a hospital, Caeliomyces tampanus (incl. Caeliomyces gen. nov.) from office dust, Cippumomyces mortalis (incl. Cippumomyces gen. nov.) from a tombstone, Cylindrium desperesense from air in a store, Tetracoccosporium pseudoaerium from air sample in house, Toxicocladosporium glendoranum from air in a brick room, Toxicocladosporium losalamitosense from air in a classroom, Valsonectria portsmouthensis from air in men’s locker room and Varicosporellopsis americana from sludge in a water reservoir. Vietnam, Entoloma kovalenkoi on rotten wood, Fusarium chuoi inside seed of Musa itinerans, Micropsalliota albofelina on soil in tropical evergreen mixed forests and Phytophthora docyniae from soil and roots of Docynia indica. Morphological and culture characteristics are supported by DNA barcodes.
... Thalli removed from Pareochomus nigripennis and P. quadriplagitus are morphologically similar and form a single phylogenetic clade. This is in line with the hypothesis that the host specificity within the H. virescens complex exists at the generic level (Haelewaters et al. 2018, Haelewaters & De Kesel 2020 Notes -In our Maximum Likelihood tree using LSU, ITS, rpb2 and tub2, our four isolates represented an independent clade in the genus Heterophoma. The isolates were different from seven existing species of Heterophoma, i.e., H. nobilis, H. poolensis, H. adonidis, H. sylvatica, H. verbascicola, H. ver bascidensiflori and H. novaeverbascicola, based on shape and size of conidia, and NaOH reaction (see Table in Supplementary material; De Gruyter & Noordeloos 1992, De Gruyter et al. 1993, Chen et al. 2015, 2017, Hou et al. 2020a. ...
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Hesperomyces virescens (Ascomycota, Laboulbeniales), a fungal ectoparasite, is thus far reported on Harmonia axyridis from five continents: North and South America, Europe, Africa, and Asia. While it is known that He. virescens can cause mortality of Ha. axyridis under laboratory conditions, the role of biotic and abiotic factors in influencing the distribution of He. virescens in the field is unknown. We collected and screened 3,568 adult Ha. axyridis from 23 locations in seven countries in Central Europe between October and November 2018 to test the effect of selected host characters and climate and landscape variables on the infection probability with He. virescens. Mean parasite prevalence of He. virescens on Ha. axyridis was 17.9%, ranging among samples from 0 to 46.4%. Host sex, climate, and landscape composition did not have any significant effect on the infection probability of He. virescens on Ha. axyridis. Two color forms, f. conspicua and f. spectabilis, had a significantly lower parasite prevalence compared to the common Ha. axyridis f. novemdecimsignata.
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Since the publication of the Catalogue of the Laboulbeniomycetes of Belgium, eleven species were found that represent new records for Belgium. Some of these taxa are rare in Europe. In this paper we present brief descriptions, notes and illustrations of these taxa. Some corrections to the Catalogue are given.
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Laboulbeniomycetes is a class of fungi that have obligate associations with arthropod hosts, either for dispersal (order Pyxidiophorales) or as biotrophic parasites (orders Herpomycetales and Laboulbeniales). Here, we focus on Herpomy-cetales and Laboulbeniales, which include fungi that form thalli, 3-dimensional, multicellular units of 1000 s of cells. Based on recently published data regarding patterns of speciation, we present the One-Host-One-Parasite model (1H1P) for haustorial thallus-forming Laboulbeniomycetes. We hypothesize that taxa with haustoria, rhizoidal structures that make contact with the host's body cavity, have very strict host specificity. For taxa without haustoria, the micro-habitat-as selected by the host-governs host shifting, presence or absence of the fungus, abundance, effective host range, and geographic distribution. We make suggestions for future research including fluorescent labeling of waxy lipids and mass spectrometry. These techniques have the potential to generate the data necessary to evaluate the here proposed 1H1P hypothesis for Herpomycetales and Laboulbeniales.
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In this study, the thallus-forming Laboulbeniomycetes (Herpomycetales and Laboulbeniales) from Denmark are presented as an illustrated monograph. Sixteen species and one genus are newly described based on morphology and ecology (host association). The new genus is named Tanmaurkiella Santam. gen. nov. and includes two species: T. pselaphi Santam. gen. et sp. nov. (type species) and T. huggertii Santam. gen. et sp. nov., both on Pselaphus heisei Herbst, 1792 (Col. Staphylinidae Pselaphinae). The other 14 new species are Amorphomyces ventricosus Santam. sp. nov. on Myrmecocephalus concinnus (Erichson, 1839) (Col. Staphylinidae Aleocharinae), Cantharomyces papillatus Santam. sp. nov. on Bledius terebrans (Schiødte, 1866) (Col. Staphylinidae Oxytelinae), Cryptandromyces cryptophagi Santam. sp. nov. on Cryptophagus distinguendus Sturm, 1845 (Col. Cryptophagidae), Cryptandromyces danicus Santam. sp. nov. on Euconnus wetterhallii (Gyllenhal, 1813) (Col. Staphylinidae Scydmaeninae), Dimeromyces oculatus Santam. sp. nov. on Longitarsus luridus (Scopoli, 1763) (Col. Chrysomelidae), Euphoriomyces enghoffii Santam. sp. nov. on Leiodes rugosa Stephens, 1829 (Col. Leiodidae), Euphoriomyces smicri Santam. sp. nov. on Smicrus filicornis (Fairmaire & Laboulbène, 1855) (Col. Ptiliidae), Laboulbenia inexpectata Santam. sp. nov. on Acupalpus exiguus Dejean, 1829 (Col. Carabidae), Laboulbenia pygidicola Santam. sp. nov. on Syntomus truncatellus (Linnaeus, 1761) (Col. Carabidae), Monoicomyces brachiatus Santam. sp. nov. on Atheta sodalis (Erichson, 1837) and Ocyusa picina (Aubé, 1850) (Col. Staphylinidae Aleocharinae), Monoicomyces crassicaulis Santam. sp. nov. on Oxypoda elongatula Aubé, 1850 (Col. Staphylinidae Aleocharinae), Monoicomyces reboleirae Santam. sp. nov. on Gnypeta carbonaria (Mannerheim, 1830) (Col. Staphylinidae Aleocharinae), Monoicomyces validus Santam. sp. nov. on Atheta vestita (Gravenhorst, 1806), Aleochara grisea Kraatz, 1856, and Geostiba circellaris (Gravenhorst, 1806) (Col. Staphylinidae Aleocharinae), and Stigmatomyces thoracochaetae Santam. sp. nov. on Thoracochaeta brachystoma (Stenhammer, 1855) (Diptera Sphaeroceridae). The total number of laboulbeniaceous species from Denmark rises from 29 to 195, which means that 166 are here reported as new country records. Among these, Kainomyces isomali Thaxt. deserves special mention as it is a new European record. The same applies to nine species which are reported here for the first time after their original description. For this study, an intensive sampling programme has been realized, with 429 Danish localities screened including around 1900 collections with fungi. Two new synonymies are established: Laboulbenia acupalpi Speg. (Spegazzini 1915a) syn. nov. for Laboulbenia stenolophi Speg. (Spegazzini 1914), and Monoicomyces oxytelis Huldén (Huldén 1983) syn. nov. for Monoicomyces invisibilis Thaxt. (Thaxter 1900). The new combination Peyritschiella oxyteli (Cépède & F.Picard) Santam. comb. nov. is proposed for Rheophila oxyteli Cépède & F.Picard including neotypification, and delimitation of Peyritschiella protea Thaxt. is incorporated. Lectotypes for Laboulbenia polyphaga Thaxt. and Symplectromyces vulgaris (Thaxt.) Thaxt. are designated. Nineteen species are illustrated here with photographs for the first time. Three species: Eumonoicomyces papuanus Thaxt., Peyritschiella protea, and Stigmatomyces euconni F.Picard, which were reported from Denmark in the literature should be removed from the Danish Funga. We have examined the following types of Thaxter from FH (Farlow Herbarium, Harvard University Herbaria): Asaphomyces cholevae Thaxt., Dimorphomyces myrmedoniae Thaxt., Eumonoicomyces papuanus, Laboulbenia polyphaga, Peyritschiella protea, Rhadinomyces pallidus Thaxt., and Symplectromyces vulgaris. Ceratomyces pyrenaeus Santam. is newly recorded from USA, and this is also a new record from the American continent. Description of the genus Cryptandromyces Thaxt. has been emended to incorporate the new species here described. Morphology of the antheridium in Eumonoicomyces papuanus is studied, and the status of the genus Eumonoicomyces Thaxt. vis-à-vis Monoicomyces Thaxt. is discussed. Identification keys are provided for genera and species. In support of the additional aim of this work to serve as a reference for the study of Laboulbeniomycetes fungi in Europe, we include maps and the Appendix 1 for comparison of the known species in the ten most diverse, better studied, European countries.
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This paper describes and illustrates a new species of Laboulbeniales (Ascomycota, Laboulbeniomycetes) recovered from Mastoptera guimaraesi bat flies (Diptera, Streblidae) in Ecuador and Panama. Bat fly-associated Laboulbeniales are still unexplored in the Neotropics, with only four described species of Gloeandromyces and one species of Nycteromyces known. Morphological characteristics and phylogenetic analyses support placement of the new taxon in Gloeandromyces and its recognition as an undescribed species. Gloeandromyces hilleri sp. nov. is easily recognized by 2-3 longitudinal rows of undulations at its perithecial venter. Phylogenetic reconstructions of the large subunit (LSU) ribosomal DNA and the translation elongation factor 1α (TEF1) both resolve G. hilleri and G. nycteribiidarum as sister species. We discuss the utility of LSU and TEF1 as secondary barcodes in Laboulbeniomycetes taxonomy.
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Historically, thallus-forming Laboulbeniomycetes, including the orders Laboulbeniales and Herpomycetales, were set apart because of their distinctive morphology and ecology. Although some biologists correctly interpreted these arthropod ectoparasites as fungi, even ascomycetes, others thought they were worms, red algae, or members of taxa described especially for them. Speculation on the evolution of the group involving red algae, the morphology-based Floridean Hypothesis, persisted deep into the 20th century, in part because valid alternatives were not presented. Although the distinctive features of Laboulbeniales clearly set them apart from other fungi, the difficulty was in the absence of characters grouping them among the fungi. Thaxter considered the Laboulbeniales to be ascomycetes, but he avoided phylogenetic discussions involved in the Floridean Hypothesis all of his life. Eventually, developmental studies of the life history of Pyxidiophora species, hyphal perithecial ascomycetes with 2-celled ascospores, revealed characters connecting Laboulbeniales to other ascomycetes. The distinctive morphological features of Laboulbeniales (absence of mycelium, a thallus developed from 2-celled ascospores by cell divisions in several planes, arthropod parasitism) can be best understood by comparison with Pyxidiophora. The development of a 3-dimensional thallus composed of true parenchyma occurs not only in Laboulbeniales, but also in Pyxidiophora species. The life history of arthropod ectoparasitism of Laboulbeniales as well as mycoparasitism and phoretic dispersal by arthropods of Pyxidiophora species can be explained by Tranzschel's Law, originally applied to rust fungi. Molecular analyses including other arthropod-associated fungi have contributed to a better understanding of an enlarged class, Laboulbeniomycetes, which now includes a clade comprising Chantransiopsis, Tetrameronycha, and Subbaromyces. A two-locus phylogenetic tree highlights evolutionary and life history questions with regard to the placement of Herpomycetales as the first diverging lineage of the Laboulbeniomycetes. The sister group for all the Laboulbeniomycetes remains to be discovered.
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