Acremonium s. lat

Abstract

This chapter addresses the asexual stages of lichenicolous Hypocreales species with non-curved conidia. A total of 23 species are accepted, currently classified in 11 genera: Acremonium, Cylindromonium, Galloea, Globonectria, Nectriopsis, Paranectria, Patriciomyces, Pronectria, Sarocladium, Trichonectria, and Trichothecium. The sexual stages of 13 of these species are known. Acremonium vanderkolkii is newly described here for a European species with large, 7–11-septate conidia. Cephalosporiopsis epiparasitaster is treated as a new synonym of Cylindromonium lichenicola, and Cylindromonium lichenophilum as a new synonym of Trichonectria rubefaciens. A phylogenetic analysis suggests that many lichenicolous Trichonectria species are congeneric with lichenicolous Cylindromonium species, although sequences from the type species, T. hirta, are still lacking.

Full text

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Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
Acremonium s. lat.
by P. Diederich, D. Ertz, W. von Brackel, P. Pinault, J. Boers, J. D. Lawrey & D. Hawksworth
1 Conidia aseptate
2 Conidia subspherical, 10–13.5(–18.5) μm diam.; conidiogenous cells 50–55 × 4.5–6 μm; usually accompanied by
supercial perithecia of the sexual stage entirely covered by pinkish brown hairs; on Physcia and other corticolous
lichens ................................................................................................................................. Nectriopsis physciicola (60)
2’ Conidia ellipsoidal to subcylindrical; when subspherical, then < 4 μm diam.
3 Conidia 2.5–4 μm long, broadly ellipsoidal; on Lobariella
4 Colonies white with pale pinkish droplets; conidia (2–)2.5–4(–5) × 1.5–2.2(–2.5) μm; usually accompanied
by immersed to erumpent, orange perithecia of the sexual stage ........................... Globonectria cochensis (48)
4’ Colonies pinkish with orange droplets; conidia broader, (2–)2.5–3.8(–4.5) × 2–2.5(–3) μm; usually accompa-
nied by supercial, orange brown perithecia of the sexual stage ........................Trichonectria setadpressa (66)
3’ Conidia 4–25 μm long
4 Conidiophores irregularly branched; sexual stage unknown
5 Conidia 4–7 μm long
6 Conidia 4–6 × 2–3 μm
7 Conidiophores verruculose; conidia 4–6 × 2–3 μm; conidiogenous cells 15–60 × 2.5–3.5 μm; on
Leptogium s. lat. ................................................................................... Acremonium spegazzinii (66)
7’ Conidiophores smooth; conidia 5–6 × 2–3 μm; conidiogenous cells 17–35 × 2.5–3 μm; on Clado-
nia and Lepraria incana ................................................................ Acremonium a. spegazzinii (67)
6’ Conidia broader, 5–7 × 3–4.5 μm; conidiophores smooth; conidiogenous cells (15–)23–45 × 2–4 μm;
on Lepra and Pertusaria ...........................................................................Acremonium pertusariae (59)
5’ Conidia 7–10.5 × 4–6 μm; conidio phores verruculose in the lower part; conidiogenous cells 20–30(–35) ×
3–4.5 μm; on Physcia ....................................................................................  (69)
4’ Conidiophores usually unbranched
5 Conidia 4–8 μm long (but septate conidia in Cylindromonium lichenicola longer)
6 Conidiogenous cells 8–20 μm long
7 Colonies sporodochium-like, chalk-white, usually accompanied by supercial, pale to orange
brown perithecia of the sexual stage; conidiogenous cells 8–17 μm long; conidia ellipsoidal,
(3.5–)4–6.5 × 2–2.5(–3) μm; on Lobariella, Ricasolia and Sticta ............. Nectriopsis curtiseta (50)
7 Colonies euse; sexual stage unknown; conidiogenous cells 15–20 μm long; conidia ellipsoidal,
4–5.5 × 1.5–2.5 μm; on various lichens ............................................ Acremonium antarcticum (46)
6’ Conidiogenous cells 20–60 μm long
7 On the discoloured thalli of Dirinaria and Polyblastidium, often accompanied by supercial, pale
to medium orange perithecia of the sexual stage; conidiogenous cells 20–41 μm long, basally
1.5–2.5 μm, apically 1–1.5 μm; conidia (4–)4.5–7(–10.5) × (1.5–)2–3(–3.5) μm, aseptate
........................................................................................................ Cylindromonium dirinariae (51)
Diederich, P., D. Ertz, W. von Brackel, P. Pinault, J. Boers, J. D. Lawrey & D. Hawksworth. 2024. Acremonium s. lat. Flora of Lichenicolous Fungi 2: 40–77.
Abstract. This chapter addresses the asexual stages of lichenicolous Hypocreales species with non-curved conidia. A total of 23 species are
accepted, currently classied in 11 genera: Acremonium, Cylindromonium, Galloea, Globonectria, Nectriopsis, Paranectria, Patriciomyces,
Pronectria, Sarocladium, Trichonectria, and Trichothecium. The sexual stages of 13 of these species are known. Acremonium vanderkolkii is
newly described here for a European species with large, 7–11-septate conidia. Cephalosporiopsis epiparasitaster is treated as a new synonym
of Cylindromonium lichenicola, and Cylindromonium lichenophilum as a new synonym of Trichonectria rubefaciens. A phylogenetic analysis
suggests that many lichenicolous Trichonectria species are congeneric with lichenicolous Cylindromonium species, although sequences from the
type species, T. hirta, are still lacking.

41
Acremonium s. lat.
7 On the thalli of Melanelixia or Melanohalea, rarely accompanied by supercial, reddish peri-
thecia of the assumed sexual stage; conidiogenous cells 32–50 μm long, basally 2.5–3.5 μm,
apically 1–2 μm; conidia (4–)4.5–5.5(–6) × (1.5–)1.8–2.5 μm, aseptate
.......................................................................................................Acremonium bavaricum (46)
7’ On other hosts, often on decaying lichen thalli, facultative lichenicolous; sexual stage unknown
8 Conidiogenous cells 30–60 μm long, basally 2–4 μm, apically 1–1.5 μm; conidia varying from
aseptate, 5–8 × 1.5–2.8 μm, 1-septate, 6–13 × 2–3 μm to 2–3-septate, 14–18 × 3 μm
..................................................................................................Cylindromonium lichenicola (57)
8’ Conidiogenous cells 20–40(–65) μm long, basally 1.5–2(–2.5) μm, apically 0.5–1 μm; conidia
always aseptate, 4–7 × 1–3.5 μm; probably heterogeneous ................ Sarocladium strictum (68)
5’ Conidia (7–)8–27 μm long
6 Conidia subcylindrical, l/w ratio 3.5–10
7 Conidiogenous cells 40–80 μm long; conidia 11.5–27 × 2–3.5 μm, often with annular thickenings
at both ends; frequently accompanied by supercial, reddish brown perithecia of the sexual stage;
on the thallus of macrolichens, mainly of Parmeliaceae .................. Trichonectria rubefaciens (63)
7’ Conidiogenous cells 13–22 μm long; conidia broader, 19–25 × 3.5–5 μm, without annular thicken-
ings at both ends; usually accompanied by immersed to erumpent, orange to red perithecia of the
sexual stage; conidiophores immersed in the necrotised thallus of Hypogymnia
............................................................................................................ Trichonectria anisospora (45)
6’ Conidia ellipsoidal, l/w ratio 1.8–3
7 Conidiogenous cells 20–26 × 3–3.5 μm, smooth; conidia 8–12 × 4–5.5 μm; sexual stage unknown;
on Evernia..........................................................................................Cylindromonium everniae (54)
7’ Conidiogenous cells 26–50 μm long; often accompanied by immersed, reddish brown perithecia of
the sexual stage [identication without the sexual stage dicult]
8 Conidia 7.5–20 × 4–7 μm; conidiogenous cells 30–55 × 2.5–5 μm, smooth or slightly verrucu-
lose; on Dirina, Physcia and Physconia .............................................Pronectria echinulata (52)
8’ Conidia 8.5–10.5 × 4–5 μm; conidiogenous cells 26–50 × 3–4 μm, nely verrucose and wavy
in outline; on Anaptychia ................................................................... Pronectria santessonii (65)
1 Conidia (0–)1(–3)-septate; sexual stage unknown
2 Conidiogenesis retrogressive, not phialidic; conidia 13–25 × 6–11 μm; rst conidium born as a blowout from
the side of the conidiogenous cell apex; next conidium produced just below, but on the opposite side; succes-
sive conidia thus produced in a characteristic zigzag patterned chain; facultative lichenicolous
................................................................................................................................. Trichothecium roseum (62)
2’ Conidiogenesis phialidic
3 Conidia 0–1(–3)-septate, 1-septate conidia 6–13 × 2–3 μm, subcylindrical, both apices rounded; facultative
lichenicolous.........................................................................................................Cylindromonium lichenicola (57)
3’ Conidia (0–)1-septate, 17–20 × 5.5–6.5 μm, biconical, one or both apices attenuated; on Physcia
.............................................................................................................................. Patriciomyces valentinianus (70)
1 Conidia 3-septate
2 Conidiogenous cells little dierentiated, subcylindrical, developing laterally from aerial hyphae, 2‒4 × 1.2‒1.5 μm;
conidia narrowly claviform, more or less straight, 3-septate, 33‒45 × 2.5‒3.5 μm, apically tapering to 1.5‒2 μm;
sexual stage unknown; on Cladonia .......................................................................................Galloea cladoniicola (48)
2’ Conidiogenous cells 50–110 × 4–6 μm; conidia 25–30 × 5–8 μm; usually accompanied by supercial, reddish, hairy
perithecia of the sexual stage; on various corticolous lichens ..............................................Paranectria oropensis (58)
1’ Conidia 7‒11-septate, 40–75 μm long; on various corticolous lichens
2 Conidia 7-septate, 40–55 × 5.5–8 μm; conidiogenous cells 40–95 × 4–5 μm; usually accompanied by supercial,
reddish, hairy perithecia of the sexual stage ............................................................................. Trichonectria hirta (55)
2’ Conidia 5‒11-septate, 45‒75 × 2.5‒4 μm; conidiogenous cells shorter, 12‒36 × 3‒5 μm; rarely accompanied by red-
dish, hairy perithecia of the assumed sexual stage ........................................................Acremonium vanderkolkii (71)

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Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
Phylogeny (nuLSU) of Hypocreales (with three members of Glomerellales used as an outgroup) obtained using a RAxML analysis, with the po-
sition of several lichenicolous or facultative lichenicolous species indicated in bold. Internal branches in bold-face indicate ML-BS values ≥70.
Lichenicolous or facultative lichenicolous members of the genera Acremonium, Cylindromonium, Globonectria, Pronectria, Sarocladium, Tri-
chothecium and Trichonectria are highlighted in yellow. The names of sequences obtained from a type specimen of a lichenicolous fungus are
marked by ‘(type)’. The scale bar is proportional to the substitution rate. The length of the dashed line was reduced by 50% for edition reasons.
0.03
Scolecofusarium ciliatum
Geosmithia lavendula
Cyanonectria cyanostoma
Roselliniella atlantica
Hapsidospora flava
Lasionectriella rubioi
Trichonectria pertusariae
Trichonectria rubefaciens 2
Microcera physciae (type)
Cylindromonium everniae (type)
Hapsidospora globosa
Hydropisphaera peziza
Cosmospora coccinea
Pronectria robergei 1
Globonectria cochensis 1
Trichonectria anisospora
Xenoacremonium recifei
Trichonectria pyrenaica
Paracosmospora physciae
Nectria cinnabarina
Acremonium alternatum
Verrucostoma freycinetiae
Lasionectria lecanodes
Nectria aurantiaca
Trichonectria usneicola
Cylindromonium alloxyli
Blastophorum aquaticum
Lasionectria mantuana
Hapsidospora inversa
Trichonectria rectipila
Roselliniella euparmeliicola
Trichothecium roseum 1
Cylindromonium eugeniicola
Pseudocosmospora vilior
Stilbocrea macrostoma
Pronectria robergei 2
Thyronectria coryli
Emericellopsis glabra
Roumegueriella rufula
Ochronectria calami
Sarocladium kiliense
Waltergamsia hennebertii
Trichothecium roseum 2
Nectriopsis violacea
Trichonectria setadpressa 1
Gliocephalotrichum bulbilium
Pronectria loweniae (type)
Thyronectria berolinensis
Bulbithecium hyalosporum
Nectriopsis rexiana
Bulbithecium arxii
Hydropisphaera aurantiaca
Calonectria cylindrospora
Reticulascus tulasneorum
Persiciospora africana
Geosmithia putterillii
Acremonium brachypenium
Pronectria cf. echinulata
Selinia pulchra
Sarocladium bactrocephalum
Trichonectria setadpressa 2
Sarocladium strictum (type)
Globonectria cochensis 2
Stilbocrea walteri
Bryocentria metzgeriae
Trichonectria epimegalosporae (type)
Cylindromonium lichenicola 2
Pseudonectria rousseliana
Waltergamsia fusidioides
Stephanonectria keithii
Sarcopodium macalpinei
Acremonium egyptiacum
Protocreopsis pertusa
Acremonium acutatum
Trichonectria rubefaciens 1
Cylindromonium lichenophilum (type)
Microcera larvarum
Bionectria grammicospora
Cylindromonium dirinariae (type)
Cylindromonium lichenicola 1
Emericellopsis maritima
Gliomastix roseogrisea
Illosporium carneum
Acremonium psychrophilum
‘Acremonium psychrophilum’
Trichonectria puncteliae (type)
Outgroup (Glomerellales)
HYPOCREALES
Kylindria chinensis
Bionectriaceae
Nectriaceae

43
Acremonium s. lat.
Introduction
Asexual stages of Hypocreales with hyaline conidia pro-
duced from phialidic conidiogenous cells have commonly
been included in the genus Acremonium. The genus was re-
vised by Gams (1971) who treated 82 species, many of which
obtained in pure culture. Four lichenicolous species were re-
vised and accepted by Hawksworth (1979), viz. A. antarcti-
cum, A. lichenicola, A. rhabdosporum and A. spegazzinii.
Lowen (1995) described the new section Lichenoidea Low-
en for nine lichenicolous species, showed that Acremonium
rhabdosporum represents the asexual stage of Nectria rube-
faciens, obtained an Acremonium-like anamorph in a culture
of Pronectria oligospora, and presented a key to all known
lichenicolous species of Acremonium.
More recently, molecular data have allowed splitting the
heterogeneous genus Acremonium in a number of natural
genera, but sequences from many species are still missing.
We have therefore decided to treat the genus Acremonium
here broadly, including all asexual stages of lichenicol-
ous Hypocreales, except those with curved conidia (Mi-
crocera, Reconditella, Roselliniella, Roselliniopsis and
Scolecofusarium), which are treated separately. We have
also included the facultative lichenicolous Trichothecium
roseum with non-phialidic conidiogenous cells, as it might
supercially be mistaken for an Acremonium s. lat. species.
Phylogeny
The order Hypocreales is placed within the class Sordari-
omycetes and includes fungi with various lifestyles such as
algicolous, bryophilous, corticolous, endophytic, entomo-
genous, foliicolous, fungicolous, lichenicolous and sapro-
bic, while some are known to be plant and human patho-
gens and several are used in industrial applications (e. g.,
Rossman et al. 1999, Lombard et al. 2015, Maharachchi-
kumbura et al. 2016, Diederich et al. 2018, Hyde et al.
2020, Hou et al. 2023, Perera et al. 2023). The number of
families accepted in Hypocreales increased from 8 to 22
in the past few years but their delimitation is not always
clearly dened (Wijayawardene et al. 2022, Hou et al. 2023,
Perera et al. 2023). The genus Acremonium was a large and
highly polyphyletic group of asexual fungi, with members
distributed over dierent orders, a situation that resulted in
the recombination of many species into other genera us-
ing molecular results (e. g., Summerbell et al. 2011, Hou et
al. 2023). The lichenicolous Acremonium antarcticum for
instance was placed in the order Glomerellales (Summer-
bell et al. 2011), but still needs to be transferred to another
genus. Other lichenicolous fungi with Acremonium-like
asexual stages belong to the Hypocreales. They represent a
heterogeneous assemblage of taxa distributed mainly over
the families Bionectriaceae, Nectriaceae and Niessliaceae.
Their generic assignment is often dicult without molecu-
lar data because of their poor morphological dierentiation.
The genus Cylindromonium was described for taxa with
an Acremonium-like morphology following phylogenetic
results (Crous et al. 2019). In our phylogeny, species of
Cylindromonium and Trichonectria are intermixed within
a strongly supported clade belonging to the Nectriaceae, as
already shown in previous studies (e. g., Crous et al. 2023,
Ohmaki et al. 2023). Therefore, the circumscription of these
two genera is problematic. Connections between asexual/
sexual stages for Cylindromonium and Trichonectria have
been highlighted for some species (Ohmaki et al. 2023)
suggesting that both genera should be merged. However,
while the type species of Cylindromonium (C. eugeniicola)
is placed within this clade, the generic type of Trichonectria
(T. hirta) is not sequenced yet leaving some doubts about
the phylogenetic position of this latter genus. Within the
family Bionectriaceae, species of Acremonium-like taxa
are recovered as polyphyletic. A rst lineage represents
Acremonium s. str. as it includes the generic type, A. alter-
natum, while a second lineage includes two lichenicolous
Acremonium-like species, one representing probably the
asexual stage of Pronectria echinulata, and another named
A. psychrophilum. The position of Acremonium psychro-
philum is uncertain, with the nuLSU sequence (OQ055395)
published by Hou et al. (2023) being part of Acremonium
s. str., while the nuLSU sequence (OP883935) published by
Etayo et al. (2023) is part of the lineage including species of
Pronectria. The connection of the sexual and asexual stages
of lichenicolous Hypocreales was proved by molecular data
for Globonectria cochensis and Trichonectria setadpressa
(Flakus et al. 2019). The genera Sarocladium and Tricho-
thecium, though being respectively part of the Nectriaceae
and Bionectriaceae in our phylogeny, are included in the
Sarocladiaceae and the Myrotheciomycetaceae in Hou et
al. (2023). The latter authors also placed Cylindromonium
and Trichonectria in the family Niessliaceae, whereas these
genera were placed in the Nectriaceae in other studies (e. g.,
Crous et al. 2019, 2023, Ohmaki et al. 2023).
Genera included
 Link
Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Fre-
unde Berlin 3: 15 (1809). Type: A. alternatum Link
Molecular data: yes (T, L). Number of species: 5–0–0 (100).
Notes. The genus Acremonium, as revised by Gams (1971),
was a heterogeneous assemblage of asexual fungi mainly
belonging to Hypocreales, but also to Coniochaetales, Mi-
crosascales and Sordariales (e. g., Summerbell et al. 2011).
Phylogenetic results have allowed transferring many spe-
cies to other genera, while the status of other unsequenced
taxa, including ve lichenicolous species currently kept
in Acremonium, is not yet settled. Acremonium psychro-

44
Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
philum, a species isolated from a lichen thallus in Antarc-
tica, may be closely related to the type A. alternatum (see
above, under Phylogeny), but it is not known whether this
is a true lichenicolous fungus.
 Crous
in Crous et al., Persoonia 43: 313 (2019). Type: C. eugeniicola Crous
Molecular data: yes (T, L). Number of species: 4–0–1 (7).
Notes. This genus was described for species of Acremoni-
um with cylindrical conidia (Crous et al. 2019, 2021), most
being lichenicolous, and several with a known sexual stage
in the genus Trichonectria. Additional species currently
included in Trichonectria belong to the Cylindromonium
clade and may be transferred there, when molecular data of
the type of Trichonectria become available.
 Alstrup & Søchting
Graphis Scripta 21: 33 (2009). Type: G. cladoniicola Alstrup &
Søchting
Molecular data: no. Number of species: 1–0–0.
Notes. A poorly known monotypic genus, distinguished
by the abundant aerial hyphae, the reduced conidiophores,
and the long, narrowly claviform conidia.
 Etayo
Biblioth. Lichenol. 84: 47: (2002). Type: G. cochensis Etayo
Molecular data: yes (T, L). Number of species: 1–0–0.
Notes. A monotypic genus similar to Pronectria, char-
acterized by the immersed perithecia, 4-spored asci and
ascospores rapidly breaking into subpherical half-spores
(Flakus et al. 2019). Phylogenetically not related to
Pronectria (Bionectriaceae), but included in Nectriaceae.
 Maire
Ann. Mycol. 9: 323 (1911), nom. cons. Type: Nectriopsis violacea
(J. C. Schmidt) Maire, nom. sanct.
Molecular data: yes (T). Number of species: 23–0–0 (73).
Notes. This genus was accepted and revised by Samuels
(1988) for fungicolous (incl. lichenicolous) hypocrealean fun-
gi with mainly pale coloured, supercial, thin-walled perithe-
cia and 1-septate ascospores. Many additional lichenicolous
species have later been described in Nectriopsis, but none of
them has ever been included in a phylogenetic analysis.
 Sacc.
Michelia 1 (3): 317 (1878). Type: P. anis Sacc.
= Ciliomyces Höhn., Sitzungsber. Akad. Wiss. Wien, Math.-
Naturw., Kl., Abt. 1 115: 673 (1906). Type: Sphaeria oropen-
sis, i. e., P. oropensis (Ces.) D. Hawksw. & Piroz.
Molecular data: yes (T, L). Number of species: 4–0–0 (5).
Notes. A genus characterized by the supercial perithecia
and transversely septate to muriform ascospores with long
and thin attenuated ends (Rossman et al. 1999).
 D. Hawksw.
in Cole & Hawksworth, Mycotaxon 77: 327 (2001). Type: P. val-
entinianus D. Hawksw.
Molecular data: no. Number of species: 1–0–0.
Notes. A monotypic genus with an asexual stage having
narrowly biconical, 1-septate conidia, obviously belonging
to Hypocreales.
 Clem.
Gen. Fungi 78: 282 (1931). Type: P. lichenicola (Ces.) Clem.,
i. e., P. robergei (Mont. & Desm.) Lowen
Molecular data: yes (T, L). Number of species: 48–0–1.
Notes. This genus comprises mainly lichenicolous species
with yellow to orange or red coloured, immersed perithecia,
and hyaline, 1-septate ascospores (Rossman et al. 1999). Asex-
ual stages of several Pronectria are of the Acremonium type.
 W. Gams & D. Hawksw.
Kavaka 3: 57 (1976 [‘1975’]). Type: S. oryzae (Sawada) W. Gams
& D. Hawksw.
Molecular data: yes (T, L). Number of species: 0–0–1 (22).
Notes. The genus Sarocladium is used for a number of for-
mer Acremonium species phylogenetically related to the
type (Summerbell et al. 2011). A single species is faculta-
tive lichenicolous.
 Kirschst.
Verh. Bot. Vereins der Prov. Brandenburg 48: 60 (1907). Type: T.
aculeata Kirschst., i. e., T. hirta (A. Bloxam) Petch
Molecular data: yes (L). Number of species: 16–0–5 (26).
Notes. The genus Trichonectria includes species with light
to dark coloured perithecia and thick-walled setae. While
the generic type has long subcylindrical to vermiform,
11–21-septate ascospores (Rossman et al. 1999) and an
asexual stage with large, subcylindrical, 7-septate conidia,
most other species included in the genus have 1-septate as-
cospores and aseptate conidia, suggesting that they may not
be congeneric with the type. Unfortunately no molecular
data are available for the generic type. As several other spe-
cies described or combined in Trichonectria belong to the
Cylindromonium clade, it is possible that all of them, except
the type, will have to be transferred to Cylindromonium.

45
Acremonium s. lat.
 Link
Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Freun-
de Berlin 3: 18 (1809), nom. sanct., Fr., Syst. Mycol. 3: 426
(1832). Type: T. roseum (Pers.) Link
Molecular data: yes (T, L). Number of species: 0–0–1 (9).
Notes. This genus comprises a number of species phy-
logenetically close to the generic type, but with vari-
able morphologies, including species with phialoconidia,
sympodial blastoconidia, and retrogressive blastoconidia
(Summerbell et al. 2011). One species is facultative li-
chenicolous.
The species
As molecular data of many species included here are still missing,
some of them may need to be transferred to other genera in the
future. The species are subsequently treated here in the alphabeti-
cal order of their epithets.
Trichonectria anisospora (Lowen) van den Boom &
Diederich
in Sérusiaux et al., Lejeunia n. S. 173: 33 (2003); Nectriella an-
isospora Lowen, Mem. New York Bot. Gard. 49: 248 (1989);
Pronectria anisospora (Lowen) Lowen, Mycotaxon 39: 461
(1990). Type: USA, Maine, Sagadahoc Co., Bates-Morse
Mountain Coastal Research Area, in the corticolous thallus
of Hypogymnia physodes, 19 Sept. 1987, R. Lowen 358-87
(NY – holotype).
= Acremonium pedatum Lowen, Mem. New York Bot. Gard. 49:
248 (1989), nom. inval. (ICN Art. 40.1).
Ascomata orange to red, immersed, becoming super-
cial, setose. Asexual stage permanently immersed. Co-
nidiophores reduced to the conidiogenous cells, rarely
1-septate, with a short lower cell, arising more or less
perpendicularly from the mycelium. Conidiogenous cells
discrete, terminal, hyaline, thin-walled, smooth, subulate,
often bent in the middle, phialidic, collarette not enlarged,
13–22 μm long, basally 3–6.5 μm wide, apically 1.3–2
μm. Conidia solitary, subcylindrical, often narrower in
the middle, apically rounded, basally truncate, 0(–1)-sep-
tate, hyaline, smooth, thick-walled, (13–)19–25(–28) ×
3.5–5 μm.
Notes. The asexual stage of this species is distinguished by
the large cylindrical conidia that are often narrower in the
middle, and by the relatively short, usually aseptate con-
idiophores. Lowen (1989) obtained the asexual stage from
single-spore cultures, but did not observe it in vivo. Here
we report it for the rst time in nature. In both specimens
examined, conidiophores and conidia were seen only in
microscopical sections of the necrotised host thallus close
to immersed perithecia. Paracosmospora physciae, iso-
lated from Physcia tenella, has similar conidia, which are,
however, constantly 1-septate.
Ecology and hosts. In the thallus of Hypogymnia physodes,
more rarely H. tubulosa, killing the infected parts.
Distribution. Europe (Austria; Belgium; Czech Republic;
Estonia; France; Germany; Ireland; Italy; Latvia; Lithu-
ania; Luxembourg; Netherlands; Norway; Poland; Russia;
Spain; Switzerland; UK; Ukraine), Macaronesia (Madeira)
and North America (USA: Maine).
Asexual specimens examined (both on Hypogymnia physodes).
France: Puy-de-Dôme: Saint-Ours, SSW of Vulcania park,
45.8074°N, 2.9309°E, 930 m, 2020, Pinault (BR, in holotype of
Tremella pinaultii). Luxembourg: Lellingen, Bärel, 49.9912°N,
6.0280°E, 420 m, 2021, Diederich 19594 (BR).
References. Lowen 1989, Sérusiaux et al. 2003 [Alstrup et al.
2008, Aptroot et al. 2005, Berger & Zimmermann 2016, Brackel
2006, 2008, 2015, Brackel & Berger 2019, Brackel & Puntillo
2016, Brackel et al. 2018, Cezanne & Eichler 2015, Etayo 1998,
Gorczak et al. 2021, Hafellner 1996, Hafellner et al. 2008, Himel-
brant et al. 2020, Hitch 1998, John et al. 2011, Malíček & Palice
2013, Malíček et al. 2018, Motiejūnaitė et al. 2011, 2016, Roux
2020, Schiefelbein et al. 2012, 2017, Stepanchikova et al. 2020,
Zhurbenko 2017, Zhurbenko & Kobzeva 2014, Zimmermann &
Berger 2018; britishlichensociety.org.uk; www.verspreidingsat-
las.nl].
Trichonectria anisospora. A, Luxembourg, Diederich 19594, B,
France, Pinault. Conidiogenous cells and conidia immersed in the
necrotised thallus of Hypogymnia physodes (arrow: 1-septate conid-
ium). Photos: P. Diederich (A) and P. Pinault (B).Scale bars: 10 μm.
Trichonectria anisospora

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Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
Acremonium antarcticum (Speg.) D. Hawksw.
Bull. Brit. Mus. (Nat. Hist.), Bot. 6: 192 (1979); Sporotrichum ant-
arcticum Speg., Anales Mus. Nac. Buenos Aires, ser. 3, 13:
416 (1910 [1911]). Type: Antarctic Islands, South Orkney Is-
lands, on Gondwania cf. regalis (Vain.) Zahlbr., Jan. 1908, C.
Spegazzini (LPS 21677 – holotype).
Sexual stage unknown. Colonies supercial, euse. Conidi-
ophores erect, simple or loosely aggregated, unbranched or
with one branch at the base, hyaline, smooth. Conidiogenous
cells discrete, terminal, hyaline, thin-walled, smooth, narrow-
ly subulate, phialidic with a broad collarette, 15–20 μm long,
basally c. 2 μm wide, apically 1–1.5 μm. Conidia solitary, ad-
hering in a slimy mass, ellipsoidal, rounded at both apices,
aseptate, hyaline, smooth, 4–5.5(–6) × 1.5–2.5(–3) μm.
Notes. W. Gams (pers. comm. 1997) had doubts about the
conspecicity of some European specimens with the type.
Ecology and hosts. On the thallus of Gondwania cf. regalis,
Hypogymnia physodes, Lepraria incana, Parmelina tiliacea,
Physcia tenella, Rusavskia elegans and Xanthoria parietina.
Distribution. Europe (Belarus; Estonia; Germany; Luxem-
bourg; Poland; Russia) and Antarctic Islands (South Ork-
ney Islands).
Specimen examined. Luxembourg: NE of Kehlen, cimetière de
Schéimerech, on Parmelina tiliacea, 1997, Diederich 12931 (BR).
References. Hawksworth 1979 [Diederich 1989, Diederich et al. 1991,
Eichler et al. 2010, Kukwa & Flakus 2009, Kukwa et al. 2013, Suija
et al. 2009, 2010, Tsurykau et al. 2014, Zhurbenko & Kobzeva 2016].
Acremonium bavaricum Brackel
in Brackel et al., Mycosphere 3: 858 (2012). Type: Germany, Bavaria,
Oberpfalz, Kreis Regensburg, Karlswiesbachtal, 49°04’32”N,
12°18’51”E, 435 m, on Fraxinus excelsior, on Melanelixia
glabratula, 28 Oct. 2009, W. von Brackel (M – holotype).
Sexual stage reddish, supercial perithecia. Colonies su-
percial, euse. Conidiophores erect, mostly simple and
unbranched, rarely septate and branched, hyaline, smooth.
Conidiogenous cells discrete, terminal, hyaline, thin-
walled, smooth, narrowly subulate, phialidic, collarette not
enlarged, 32–50 μm long, basally 2.5–3.5 μm wide, apical-
ly 1–2 μm. Conidia solitary or assembled in small drops,
narrowly ellipsoidal, apically rounded, basally truncate,
aseptate, hyaline, smooth, (4–)4.5–5.5(–6) × (1.5–)1.8–2.5
μm (in vitro 2.5–3.5 × 1.5–2 μm).
Notes. This species has conidia similar in size to those of
Acremonium antarcticum and A. spegazzinii. The former
Acremonium antarcticum
Acremonium antarcticum, Luxembourg, Diederich 12931. Colonies
over the thallus of Parmelina tiliacea. Scale bar: 200 μm.
Acremonium antarcticum, South Orkney Islands, holotype (modied
from Hawksworth 1979). Conidiophores, conidiogenous cells and
conidia. Scale bars: 10 μm.
Acremonium bavaricum, Germany, holotype (modied from Brackel
et al. 2012). Colonies over the thallus of Melanelixia glabratula.
Scale bar: 500 μm.

47
Acremonium s. lat.
diers in the shorter conidiogenous cells, while the lat-
ter has verruculose conidiophores. Following Brackel et
al. (2012), two species of Hypocreales were known from
Melanelixia and Melanohalea, viz. Paranectria oropensis
and Pronectria septemseptata, but none of them has been
found associated to Acremonium bavaricum. Addition-
ally, a culture of P. septemseptata yielded a very dierent
asexual stage, with conidiogenous cells having up to 20
conidiogenous loculi and conidia becoming brown (Brack-
el et al. 2012). In specimen 17739 from Luxembourg, P.
septemseptata was growing on a neighbouring tree, but
was not associated to A. bavaricum. Instead, small red-
Acremonium bavaricum, Luxembourg, Diederich 17739. Colonies over the thallus of Melanelixia glabratula, showing immature red perithecia
(colour faded in herbarium). Scale bar: 200 μm.
Acremonium bavaricum, France, Pinault. Conidiophores, conidiogenous cells and conidia, in a mixture of ammoniacal congo red and phloxine
(A–C), or in water (D–F). Several conidiophores carry a drop (stained orange) lled with conidia (purple) that includes conidia. Photos: P.
Pinault. Scale bars: 10 μm.

48
Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
dish, supercial perithecia grew amongst the Acremonium
population, unfortunately immature, and these most likely
represent its sexual stage.
Ecology and hosts. On the thallus of Melanelixia glabratula,
M. subargentifera and Melanohalea exasperatula.
Distribution. Europe (France; Germany; Luxembourg).
Additional specimens examined. France: Puy-de-Dôme: Romag-
nat, Opme, 45.7100°N, 3.0835°E, on Melanelixia subargentifera,
2024, Pinault (herb. Pinault). Luxembourg: Strassen, Gaascht-
gronn, 49.6261°N, 6.0515°E, 300 m, on M. glabratula, 2014,
Diederich 17739 (BR).
References. Brackel et al. 2012 [Brackel 2019, 2023].
Galloea cladoniicola Alstrup & Søchting
Graphis Scripta 21: 33 (2009). Type: Denmark, East Jutland, 19
km WSW Vejle, Liegård Plantage, 55°37’45”N, 9°15’41”E,
on Cladonia cf. chlorophaea, 22 Aug. 2007, U. Søchting (C –
holotype; BG – isotype).
Sexual stage unknown. Colonies of dense, white aggrega-
tions of aerial hyphae and conidiophores. Mycelium im-
mersed, hyphae hyaline, 1.5‒2.5 μm diam., with long erect,
frequently branched aerial hyphae. Conidiophores reduced
to conidiogenous cells. Conidiogenous cells little dieren-
tiated, monophialidic, subcylindrical, developing laterally
from the aerial hyphae, 2‒4 × 1.2‒1.5 μm. Conidia dry,
hyaline, narrowly claviform, more or less straight, 3-sep-
tate, 33‒45 × 2.5‒3.5 μm, apically tapering to 1.5‒2 μm,
thin- and smooth-walled.
Notes. This is a remarkable species characterized by the
large, 3-septate conidia, born on very short and narrow
conidiogenous cells, emerging laterally or terminally from
long aerial hyphae.
Ecology and host. Over basidiomatal galls of Tremella cla-
doniae on the thallus of Cladonia cf. chlorophaea.
Distribution. Europe (Denmark), known only from the
type locality.
Reference. Alstrup & Søchting 2009.
Globonectria cochensis Etayo
Biblioth. Lichenol. 84: 47 (2002). Type: Colombia, Nariño, Pasto,
corregimiento El Encano, Reserva Natural Tunguragua, SE
lago La Cocha (Guamués), 2700 m, on Lobariella pallida,
29 July 1998, J. Etayo 15859, S. Churchill, J. Muñoz & B.
Ramírez (COL – holotype; herb. Etayo – isotype).
Ascomata orange, immersed to erumpent perithecia. Col-
onies of the asexual stage white sporodochium-like ag-
Galloea cladoniicola
Acremonium bavaricum, Germany, holotype (modied from Brackel
et al. 2012). Conidiophores, conidiogenous cells and conidia. Scale
bars: 10 μm.
Acremonium bavaricum
Globonectria cochensis

49
Acremonium s. lat.
Galloea cladoniicola, Denmark, holotype. A, Aerial hypha (arrow head) with perpendicular conidiogenous cells, and conidia attached to
conidiogenous cells (arrows). B, Conidia and conidiogenous cell (arrow). C, Aerial hyphae. D, Mycelium with two small branches, narrow
conidiogenous cells (arrows) and young conidia. E, Colonies over basidiomatal galls of Tremella cladoniae on a podetium of Cladonia cf.
chlorophaea. A–D, in ammoniacal Congo red. Scale bars: A–D = 10 μm, E = 200 μm.

50
Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
gregations of conidiophores on an arachnoid subiculum,
often with pale pinkish ‘droplets’ made of conidia. Con-
idiophores reduced to the conidiogenous cell. Conidiog-
enous cells discrete, terminal, hyaline, straight or slightly
curved, 15–30 μm long, basally 1–2 μm wide, tapering at
the apex. Conidia solitary, subspherical to broadly ellipsoi-
dal, not truncate, aseptate, hyaline, smooth, (2–)2.5–4(–5)
× 1.5–2.2(–2.5) μm.
Notes. The dierences with the very similar asexual
stage of Trichonectria setadpressa are given under that
species.
Ecology and hosts. On the thallus of Lobariella pallida and
L. crenulata.
Distribution. South America (Bolivia; Colombia).
References. Etayo 2002, Flakus et al. 2019.
Nectriopsis curtiseta Etayo
Opera Lilloana 50: 292 (2017). Type: Ecuador, Prov. Imbabura,
Otavalo, Lagunas de Mojanda, ladera entre Laguna grande y
Laguna Chiquita, 0°07’01”N, 78°16’16”W, 3700 m, bosque
de Polylepis orientado, on Sticta sp., 27 Febr. 2003, J. Etayo
25414 & Z. Palice (QCA – holotype; herb. Etayo – isotype).
Ascomata pale to orange brown, supercial perithecia. Col-
onies of the assumed asexual stage supercial, chalk-white,
sporodochium-like, 200–800 μm diam. Conidiophores re-
Asexual stage of Globonectria cochensis, Bolivia, on Lobariella crenulata, Flakus 22184 (A and E), on Lobariella sp., Etayo 33-20 (B–D) (re-
produced from Flakus et al. 2019). A–B, Colonies over the host thallus. C, Conidiophores and conidia in water. D–E, Conidia in LPCB. Scale
bars: A–B = 250 μm, C–E = 10 μm.
Nectriopsis curtiseta, Madeira, Diederich 19753. Assumed asexual
stage (white) and orange brown perithecia of the sexual stage over
an old apothecium and the thallus of Ricasolia sublaevis. Scale bar:
500 μm.
Nectriopsis curtiseta, Madeira, Diederich 19753. Conidiophores, con-
idiogenous cells and conidia, in phloxine. Scale bar: 10 μm.

51
Acremonium s. lat.
duced to the conidiogenous cell. Conidiogenous cells dis-
crete, terminal, hyaline, straight, 8–17 μm long, basally
1.5–3 μm wide, apically 1–1.5 μm. Conidia solitary, nar-
rowly ellipsoidal, apically rounded, basally truncate, asep-
tate, hyaline, smooth, (3.5–)4–6.5 × 2–2.5(–3) μm.
Notes. The sexual stage of this species was described from
Sticta by Etayo (2017); this author further described and il-
lustrated an unnamed Acremonium on Lobariella crenulata.
Flakus et al. (2019) observed the same Acremonium several
times on Lobariella apothecia in the presence of Nectriopsis
curtiseta ascomata and suggested that it may represent the
asexual stage of that species. The co-occurrence of both stag-
es in our recent Madeira specimen supports that hypothesis.
Ecology and hosts. On the thallus and apothecia of Lobari-
ella crenulata, L. pallida, Lobariella sp., Ricasolia pat-
inifera, R. sublaevis, Sticta sp. and S. weigelii, the asexual
stage mainly on the apothecial and thalline margins.
Distribution. Macaronesia (Madeira), Central America
(Guatemala) and South America (Bolivia; Ecuador; Peru).
Specimen examined. Madeira: Ribeiro Frio, 32.7337°N,
16.8870°W, 920 m, on Ricasolia sublaevis, 2023, Dieder-
ich 19753 (BR).
References. Etayo 2017, Flakus et al. 2019.
Cylindromonium dirinariae Ohmaki & Okane
in Ohmaki et al., Fungal Syst. Evol. 11: 4 (2023). Type: Japan, Ibaraki,
Tsukuba, Tennodai, Univ. of Tsukuba, 36˚06’08”N, 140˚06’24”E,
on bark of Zelkova serrata, on Dirinaria applanata, 1 Aug. 2020,
A. Ohmaki, I. Okane, K. Ohmachi, K. Miyazawa & K. Gibu, FAO
005 (TNS-L-131533 – holotype; ex-type culture NBRC 115852).
Ascomata supercial, perithecioid, pale to medium orange, with
short, sti setae. Mycelium consisting of hyaline, smooth, sep-
tate, branched, 1.5–2 μm diam. hyphae. Conidiophores solitary,
erect, unbranched or rarely branched near the base, 0–2-sep-
tate, hyaline, smooth, 45–51 × 2–3.5 μm. Conidiogenous cells
hyaline, smooth, subcylindrical, 20–41 μm long, 1.5–2.5 μm
wide at the base, 1–1.5 μm at the apex, phialidic, collarette not
enlarged. Conidia solitary, adhering in a slimy mass, hyaline,
smooth, aseptate, ellipsoidal, apically rounded, basally narrow-
ly truncate, (4–)4.5–7(–10.5) × (1.5–)2–3(–3.5) μm.
Notes. This is the rst known Cylindromonium species in
which both the sexual and asexual stages have been ob-
served in nature and included in a phylogenetic analysis.
The asexual stage strongly resembles that of the facultative
lichenicolous Sarocladium strictum, but is easily distin-
guished by the frequent presence of the sexual stage, su-
percial, orange perithecia with short setae.
Nectriopsis curtiseta
Cylindromonium dirinariae, Vietnam, Zhurbenko 19223. Sexual and asexual stages over the thallus of Polyblastidium sp. Scale bar: 200 μm.
Cylindromonium dirinariae

52
Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
Ecology and hosts. On the thallus and apothecial margin of
Dirinaria applanata, D. purpurascens, and Polyblastidium
sp., often decolorizing the host thallus.
Distribution. North America (USA: Florida) and Asia (Japan;
Vietnam).
Specimens examined. USA: Florida: Pasco Co., Zephyrhills,
Woodfern Ave., 28.246°N, 82.192°W, 35 m, on corticolous Di-
rinaria purpurascens, 2019, Common 10337C (BR) (asexual
stage); Pasco Co., Zephyrhills, along Henry Ave., 28.248°N,
82.179°W, 25 m, on corticolous D. purpurascens, 2013, Com-
mon 9591A (BR) (sexual stage). Vietnam: Northeast Region,
Cao Bằng Province, Nguyên Bình District, Phia Oắc-Phia Đén
National Park, 5.2 km SW of Tinh Tuc village, 22°36’28”N,
105°51’59”E, 1680 m, on Polyblastidium sp., 2019, Zhurbenko
19223 (BR) (sexual and asexual stages).
Reference. Ohmaki et al. 2023.
Pronectria echinulata Lowen
in Rossman et al., Stud. Mycol. 42: 58 (1999). Type: Ireland, Gort-
naskehy, Aug. 1985, on Salix, on Physcia aipolia, M. R. D.
Seaward (K IMI 105139 – holotype).
= Acremonium hypholomatis auct., non (Boedijn) D. Hawksw.
Ascomata reddish brown perithecia, immersed in the host
thallus or apothecia. Colonies of the assumed asexual stage
euse over the host thallus, whitish to pale pinkish. Co-
nidiophores erect, simple, basally septate. Conidiogenous
cells discrete, terminal, hyaline, relatively thick-walled,
smooth to slightly verruculose, narrowly subulate, phial-
idic, collarette not enlarged, 30–55 μm long, basally 2.5–5
μm wide, apically 1.5–3 μm. Conidia solitary, ellipsoidal,
apically rounded, basally narrowly truncate, aseptate, hya-
line, smooth, thick-walled, 7.5–20 × 4–7 μm.
In vitro (specimen Diederich 16919): Colonies in liquid cul-
ture pinkish, showing abundant white aerial hyphae. Hyphae
hyaline, septate, straight, branched. Conidiogenous cells and
conidia similar to those in vivo. Chlamydospores present,
hyaline, terminal, subspherical, aseptate, 8–20 µm diam.
Notes. Diederich & Braun (2009) reported a lichenicol-
ous population of Acremonium from necrotised thalli of
Cylindromonium dirinariae, Vietnam, Zhurbenko 19223. Conidiophores, conidiogenous cells, conidial mass and single conidia, in phloxine or
KOH. Scale bar: 10 μm.
Assumed asexual stage of Pronectria cf. echinulata, Spain, Mallorca,
Diederich 16919. Colonies over the thallus and apothecia of Dirina
ceratoniae. Scale bar: 500 μm.

53
Acremonium s. lat.
Physcia stellaris in Germany with particularly large co-
nidia, 11.5–20 × 5.5–7 μm, as Acremonium hypholomatis.
As perithecia of Pronectria echinulata were also present
and abundant in the collecting locality on the same host,
and as the neotype of A. hypholomatis was isolated from a
water culture of decaying leaves in Papua New Guinea, it
is likely that the lichen-inhabiting Acremonium represents
the asexual stage of P. echinulata and is not conspecic
with the genuine A. hypholomatis. Brackel (2008) col-
lected a specimen of P. echinulata in Italy on Physconia
venusta with a similar Acremonium-like anamorph hav-
ing slightly smaller conidia, (7–)8–10(–11) × (4–)4.5–6(–
6.5) μm. Conidia in a Russian specimen (Himelbrant et
al. 2018) were ‘concurrent with those given by Dieder-
ich & Braun (2009)’. Brackel & Berger (2019) collected
P. echinulata with an ‘Acremonium anamorph’ twice in
Italy, Sardinia. A sequenced specimen from Mallorca on
an unusual host (Dirina ceratoniae) is here treated as P.
cf. echinulata.
Ecology and hosts. The assumed asexual stage is known
from the thallus of Dirina ceratoniae, Physcia stellaris, Phy-
scia sp., Physconia distorta, P. enteroxantha and P. venusta.
Distribution. The assumed asexual stage is known from
Europe (Germany; Italy; Portugal; Russia; Spain). No map
is given.
Assumed asexual stage of Pronectria cf. echinulata, Spain, Mallorca, Diederich 16919. Conidiogenous cells and conidia, in water. Scale bar: 10 μm.
Assumed asexual stage of Pronectria echinulata, Italy, Brackel 4375.
Conidia, in water. Photo: W. von Brackel. Scale bar: 10 μm.

54
Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
Selected specimens of the assumed asexual stage examined. Italy:
Marche: Prov. Macerata, Cingoli, S Zentrum, 43°22’N, 13°13’E,
590 m, on Physconia venusta, 2011, Brackel 6408 (herb. Brack-
el). Sicily: Prov. di Palermo, Bosco della Ficuzza, mixed oak
forest, on Physconia venusta, 37°51’51”N, 13°24’57”E, 945 m,
2007, Brackel 4375 (herb. Brackel). Spain: Mallorca: 2 km S of
Valldemossa, 39.6930°N, 2.6342°E, 260 m, on bark of Quercus
ilex subsp. rotundifolia, on Dirina ceratoniae, 2010, Diederich
16919 (BR).
References. Diederich & Braun 2009 [Brackel & Berger 2019,
Etayo & López de Silanes 2020, Himelbrant et al. 2018].
Cylindromonium everniae Crous & Boers
in Crous et al., Persoonia 47: 243 (2021). Type: Netherlands,
Prov. Drenthe, Dwingeloo, graveyard, 52.8316°N, 6.3644°E,
10 m, isolated from Evernia prunastri, March 2021, J. Boers
(CBS H-24801 – holotype; ex-type culture CBS 148255).
Sexual stage unknown. Colonies supercial, euse, whit-
ish. Conidiophores solitary, erect, unbranched, 1(–2)-sep-
tate, hyaline, smooth, 28–35 × 3–4 μm (in vitro 40–100 ×
2.5–3 μm). Conidiogenous cells discrete, terminal, hyaline,
wall c. 0.5 μm, smooth, narrowly subulate to subcylindri-
cal, phialidic, collarette slightly enlarged, 20–26 μm long,
basally 3–3.5 μm wide, apically 1.5–2.5 μm (in vitro 35–55
μm long, basally 2.5–3 μm wide, apically 1.5–2 μm). Co-
nidia solitary, ellipsoidal to subcylindrical, apically round-
ed, basally narrowly truncate, aseptate, hyaline, smooth,
(7.5–)8–12(–13.5) × (3.5–)4–5.5 μm (in vitro (8–)12–14(–
19) × 3–4 μm).
Notes. This species was described based on a culture ob-
tained from a specimen on Evernia prunastri. Unfortunate-
ly, no in vivo description or illustrations of the specimen
were provided. J. Boers recollected an Acremonium-like
fungus on E. prunastri in the type locality with conidi-
ophores and conidiogenous cells much shorter than in the
original in vitro description. We obtained a culture and se-
quences identical to those of the type. This demonstrates
Assumed asexual stage of Pronectria echinulata, Germany, Heklau
(modied from Diederich & Braun 2009). Conidiophores and co-
nidia. Scale bar: 10 μm.
Assumed asexual stage of Pronectria echinulata, Italy, Brackel 6408. Colonies over the thallus of Physconia venusta. Photo: W. von Brackel.
Scale bar: 100 μm.

55
Acremonium s. lat.
the importance of giving in vivo descriptions of new spe-
cies, whenever possible, allowing the identication of
specimens from nature.
Ecology and host. On the necrotised and discoloured thal-
lus of Evernia prunastri.
Distribution. Europe (Netherlands), known only from the
type locality.
Specimen examined. Netherlands: Same locality as type, on Ever-
nia prunastri, 11 Dec. 2022, Boers (BR – ‘topotype’).
Reference. Crous et al. 2021.
Trichonectria hirta (A. Bloxam) Petch
Naturalist (Hull) 1937: 282 (1937); Nectria hirta Bloxam in
Currey, Trans. Linn. Soc. London 24: 158 (1863); Calonec-
tria hirta (A. Bloxam) Sacc., Michelia 1: 307 (1878). Type:
England, Leicestershire, Twycross, on decaying rails, on
Placynthiella uliginosa, leg. A. Bloxam (K ex Broome –
holotype).
Cylindromonium everniae, Netherlands, Boers (‘topotype’). A, Conidia. B, Conidiophores, conidiogenous cells and developing conidia. C,
Mycelium with lower cell of conidiophore. B–C, in phloxine. Scale bar: 10 μm.
Cylindromonium everniae
Cylindromonium everniae, Netherlands, Boers (‘topotype’). Colonies
over the thallus of Evernia prunastri. Scale bar: 200 μm.

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Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
= Calonectria vermispora Massee & Crossland, Naturalist (Hull)
1904: 4 (1904). Type: UK, England, Yorkshire, Hebden
Bridge, Hardcastle Hill, on decorticate trunk, 6 Nov. 1897, J.
Needham (K – holotype).
Ascomata supercial, reddish, hairy perithecia. Colonies
of the asexual stage supercial, pinkish, euse or in sporo-
dochium-like aggregations of conidiophores, 10–20 mm
diam. Conidiophores erect, aseptate, unbranched, hyaline,
smooth. Conidiogenous cells discrete, terminal, hyaline,
straight, phialidic with a distinct collarette when young,
40–95 μm long, basally 4–5 μm wide, apically 2.5–3 μm.
Conidia solitary, elongate ellipsoidal, fusiform to subcy-
lindrical, straight or slightly bent, apically rounded, basally
rounded or narrowly truncate, (0–)7(–11)-septate, hyaline,
smooth, 7-septate conidia (35–)40–55(–60) × (4–)5.5–8 μm.
Notes. This species was rather common in Western Europe
in the 1980s, especially on strongly eutrophic bark in pol-
luted environments, but has declined drastically after 2000.
No fresh material for molecular work could be obtained.
Ecology and hosts. Over corticolous lichens, including Le-
canora conizaeoides, Lepraria incana, Micarea prasina,
Asexual stage of Trichonectria hirta, Luxembourg, Diederich 8540. Conidiophores, conidiogenous cells and conidia. Scale bar: 10 μm.

57
Acremonium s. lat.
Physcia tenella, Placynthiella uliginosa, P. dasaea, Scoli-
ciosporum chlorococcum and S. umbrinum, usually killing
the host thalli over a surface reaching 20 mm in diam.
Distribution. Europe (Austria; Belgium; Czech Republic;
Denmark; France; Estonia; Germany; Luxembourg; Neth-
erlands; Poland; Spain; UK; Ukraine). No map is given.
Selected specimens examined. Belgium: 10 km WSW of Arlon,
large quarry on S side of road from Meix-le-Tige to Chantemelle,
at c. 2.5 km ESE of Chantemelle, 49.64°N, 5.69°E, on Placyn-
thiella dasaea, 2002, Diederich 15566 (BR). Luxembourg:
NWW of Pétange, bois d’Athus, 49.55°N, 5.84°E, on Carpinus,
on Scoliciosporum chlorococcum, 1987, Diederich 8540 (BR).
Selected references. Diederich 1989, Rossman et al. 1999.
Cylindromonium lichenicola (W. Gams) Crous
in Crous et al., Persoonia 43: 313 (2019); Acremonium licheni-
cola W. Gams, Cephalosporium-art. Schimmelpilze: 134
(1971). Type: Germany, Plon District, Schüttbrehm, isol. ex
Betula litter, May 1965, W. Gams (CBS 425.66 – holotype; K
IMI 224426 – isotype).
Syn. nov.: Cephalosporiopsis epiparasitaster Alstrup, Graphis
Scripta 8: 25 (1993). Type: Norway, Trøndelag, Grong, c. 1 km
E of Grong centre, small brook ravine NE of hill, UTM 33W
UM 7252, 100–160 m, old spruce forest, on Chaenothecopsis
parasitaster, 26 July 1993, V. Alstrup (C – holotype).
Sexual stage unknown. Colonies supercial, euse. Con-
idiophores erect, unbranched, 0–1(–3)-septate, with a short
basal cell, hyaline, thin-walled, smooth, subcylindrical to
Asexual stage of Trichonectria hirta, Luxembourg, Diederich 8540.
Colonies over the thallus of Scoliciosporum chlorococcum. Arrow
points at perithecia. Scale bar: 200 μm.
Asexual stage of Trichonectria hirta, Luxembourg, Diederich 8540.
Colonies over the thallus of Scoliciosporum chlorococcum, showing
conidiophores with conidia. Scale bar: 50 μm.
Cylindromonium lichenicola, Luxembourg, Marson. A, Colonies over the thallus of Physcia tenella. B, Conidia. C, Conidiophores, conidio-
genous cells and conidia. In water. Photos: G. Marson. Scale bars: A = 200 μm, B–C = 10 μm.

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Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
liform. Conidiogenous cells discrete, terminal, hyaline,
thin-walled, smooth, phialidic with a broad collarette, 30–60
μm long, basally 2–4 μm wide, apically 1–1.5 μm. Conidia
solitary (in vitro in a slimy mass), subcylindrical, rounded at
both apices, hyaline, smooth, thin-walled, 0–1(–3)-septate,
aseptate conidia 5–8(–20) × 1.5–2.8 μm, 1-septate 6–13 ×
2–3 μm, 2- and 3-septate 14–18(–21) × 3(–4) μm.
Notes. This species is distinguished from all other lichenicol-
ous Acremonium s. lat. species by the narrowly cylindrical co-
nidia that are rst aseptate, often become 1-septate, rarely up
to 3-septate. When Hawksworth (1979) revised the licheni-
colous hyphomycetes, the species was known only from cul-
ture, with relatively small, 0–1-septate conidia, 5–8 × 1.5–2.8
μm. Tsurkyau et al. (2016) observed conidia in nature up to 18
μm long and up to 3-septate, suggesting a much more variable
species. In a French specimen examined by us, and a Lux-
embourg specimen photographed by G. Marson (see photo-
graphic plate), conidia reached 21 × 4 μm, but otherwise are
in agreement with the description of the species.
Cephalosporiopsis epiparasitaster was described from the
apothecia of the lichenicolous Chaenothecopsis parasitaster
for an Acremonium-like fungus with erect, mostly straight,
hyaline, smooth conidiophores, 15–55 μm tall, basally c. 3
μm wide, apically 0.7–1 μm, and aseptate, hyaline, smooth,
subcylindrical conidia with rounded ends, 8–9 × 2 μm. We
have re-examined the holotype and found just a single squa-
mule of Cladonia with a few ascomata of C. parasitaster
that did not appear parasitized. We have re-opened one of
the sealed and dried original slides and transferred the con-
tent to a new slide. Many conidiophores with broad phialides
were visible, some with a minuscule, young conidium, but
no mature conidia were seen. The observed conidiophores,
together with the conidial dimensions given in the original
description, leave no doubt that this name is a younger syno-
nym of Cylindromonium lichenicola.
Ecology and hosts. Over fungi, corticolous algae, bark and
Betula litter. Facultatively lichenicolous on the thallus of
Cladonia mbriata, C. grayi, C. squamosa, Hypocenom-
yce scalaris, Hypogymnia physodes, Parmelia saxatilis,
Physcia tenella and Punctelia sp., and on apothecia of the
lichenicolous fungus Chaenothecopsis parasitaster over
squamules of Cladonia digitata.
Distribution. Lichenicolous specimens reported from Eu-
rope (Belarus; Belgium; Germany; France; Luxembourg;
Netherlands; Norway; UK: England; Ukraine) and North
America (Mexico). No map is provided.
Selected specimen examined. France: Puy-de-Dôme: Clermont-
Ferrand, Chanturgue, on Lecidella, 2023, Pinault (herb. Pinault).
References. Gams 1971, Hawksworth 1979, Tsurkyau et al. 2016
[Alstrup 1997 as Cephalosporiopsis epiparasitaster, Brackel
2010, 2015, 2023, Brand et al. 2013, Diederich 1989, Etayo &
Sancho 2023, Hitch 1995, Khodosovtsev et al. 2018, Roux 2020].
Paranectria oropensis (Ces.) D. Hawksw. & Piroz.
Canad. J. Bot. 55 (19): 2555 (1977); Sphaeria oropensis Ces.
and Nectria oropensis Ces. (alternative names), in Rabenh.,
Klotzschii Herb. Viv. Mycol., Ed. Nova, Cent. 6: 524 (1857);
Dialonectria oropensis (Ces.) Cooke, Grevillea 12 (64): 111
(1884); Cucurbitaria oropensis (Ces.) Kuntze, Revis. Gen.
Pl. 3 (3): 461 (1898); Ciliomyces oropensis (Ces.) Höhn.,
Sitzungsber. Akad. Wiss. Wien, Math.-Naturwiss., Kl., Abt. 1,
Cylindromonium lichenicola, Germany, Gams (CBS 777.69) (modi-
ed from Hawksworth 1979). In vitro conidiophores, conidiogenous
cells and conidia. Scale bars: 10 μm.
Asexual stage of Paranectria oropensis, Luxembourg, Diederich 17500. Colonies over the thallus of Lepraria nkii. Scale bar: 50 μm.

59
Acremonium s. lat.
115: 673 (1906). Type: Italy, Piedmont, Oropa, near Sanctu-
ary of Oropa, on Cladonia squamules, Sept. 1856, Cesati, Ra-
benh., Herb. Viv. Mycol. 524 (BG, C, HAL, M, S – syntypes).
Ascomata supercial, reddish, hairy perithecia. Colonies of
the asexual stage supercial, euse or sporodochium-like,
pinkish. Conidiophores erect, septate, hyaline, smooth, un-
branched or branched near the base. Conidiogenous cells
discrete, terminal, hyaline, straight, phialidic with a distinct
collarette, 50–110 μm long, basally 4–6 μm wide, apically
2.5–3 μm. Conidia solitary, elongate ellipsoidal to subcylin-
drical, straight, apically rounded, basally narrowly truncate,
(1–)3-septate, hyaline, smooth, 25–30(–40) × 5–8(–12) μm.
Notes. This is a common and widespread species with
supercial, reddish, hairy perithecia and large, muriform
ascospores attenuated at both ends. The asexual stage has
been reported and illustrated by Brackel (2015). In the ma-
terial examined by us, it is present in more than 50% of the
specimens. As most specimens have been reported from
the sexual stage, no detailed information on hosts, distribu-
tion and references is given here.
Ecology and hosts. Over corticolous lichens, including
Cladonia sp.
Distribution. Europe (incl. Italy), North America and Asia.
No map is given.
Selected specimen of the asexual stage examined. Luxembourg:
Between Alzingen and Syren, 200 m NEE of road bridge over
railways, 49.5645°N, 6.1963°E, 290 m, on Fagus, on Lepraria
nkii, 2013, Diederich 17500 (BR).
Reference concerning the asexual stage. Brackel 2015.
Acremonium pertusariae Brackel & Etayo
in Brackel et al., Mycosphere 3: 855 (2012). Type: Italy, Basilicata,
Prov. di Potenza, Bosco Teduri near Bagni, old beech forest, on
Fagus sylvatica, on Pertusaria pertusa, 40°06’26”N, 15°58’25”E,
1260 m, 16 Aug. 2010, W. & G. von Brackel 5498 (M – holotype;
herb. von Brackel 5498 – isotype; CIRM – ex-type culture).
Sexual stage unknown. Colonies supercial, euse to con-
uent, tufted, pale pinkish, with sporodochium-like orange
droplets of released conidia. Conidiophores erect, richly
branched, septate, hyaline, smooth (in vitro surface rough).
Conidiogenous cells discrete, terminal, hyaline, thin-
walled, smooth, subulate, phialidic, collarette not enlarged,
Asexual stage of Paranectria oropensis, Luxembourg, Diederich 17500.
A, Conidiophores and phialide with young conidium. B, Conidia. In
KOH. Scale bar: 10 μm.
Acremonium pertusariae, Italy, holotype (modied from Brackel et
al. 2012). Conidiophores, conidiogenous cells and conidia. Scale
bars: 10 μm.

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Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
(15–)23–45 μm long, basally 2–4 μm wide, apically 1.5–2
μm, guttulate. Conidia solitary, broadly ellipsoidal, apical-
ly rounded, basally slightly truncate or rounded, aseptate,
hyaline, smooth, guttulate, (4.5–)5–7(–8.5) × 3–4.5 μm.
Notes. This species is distinguished by the relatively short
and broad conidia, and the richly branched conidiophores.
Pronectria pertusariicola Lowen, also growing on Lepra
and Pertusaria species, is frequently accompanied by an
Acremonium-like asexual stage with similarly sized conid-
iogenous cells and conidia. Brackel et al. (2012) and Brackel
(2021) reported the co-occurrence of A. pertusariae and P.
pertusariicola and wondered whether both might be syno-
nyms. They concluded that they are probably distinct spe-
cies, as conidiophores are typically simple and verrucose in
P. pertusariicola, but branched and smooth in A. pertusari-
ae. Further, both species have been obtained in pure culture
(Brackel et al. 2012), where P. pertusariicola had longer
conidiogenous cells, 15–50 μm vs 15–30 μm, and thinner
conidia than A. pertusariae, l/w = 1.6–2.1 vs 2–2.9.
Ecology and hosts. On the thallus of Lepra albescens and
Pertusaria pertusa.
Distribution. Europe (France; Germany; Italy; Russia; Spain;
Switzerland) and Macaronesia (Canary Islands: La Palma).
Selected specimen examined. France: Pyrénées-Atlantiques: 22
km ESE of Saint-Jean-Pied-de-Port, W of Larrau, chalets d’Iraty,
43.0403°N, 1.0266°W, 1350 m, on Lepra albescens, 2015, Died-
erich 18101 (BR).
References. Brackel et al. 2012, Zhurbenko 2017 [Brackel 2015,
2023, Brackel & Puntillo 2016; swissfungi.wsl.ch].
Nectriopsis physciicola D. Hawksw. & Earl.-Benn.
in Earland-Bennett et al., Bol. Soc. Micol. Madrid 30: 245 (2006).
Type: Spain, Comunidad de Madrid, Municipio de El Boalo,
Mataelpino, 500 m S of village centre, on Fraxinus angustifola,
on apothecia and thallus of Physcia stellaris, 8 Nov. 2004, P. M.
Earland-Bennett & C. J. B. Hitch (MAF-Lich 14098 – holotype).
Ascomata supercial perithecia entirely covered by pink-
ish brown hairs. Colonies of the asexual stage supercial,
euse, white. Conidiophores erect, simple, aseptate or with
a septum close to the base. Conidiogenous cells discrete,
terminal, hyaline, thin-walled, smooth, narrowly subulate,
phialidic, collarette not enlarged, 15–55 μm long, basally
4.5–6 μm wide, apically (2–)3–4 μm. Conidia solitary,
subspherical, aseptate, hyaline, smooth, thin-walled, 10–
13.5(–18.5) μm diam.
Notes. This asexual stage is distinguished from all other
Acremonium s. lat. species by the large subspherical conidia.
Acremonium pertusariae, France, Diederich 18101. Colonies over the thallus of Lepra albescens. Scale bar: 200 μm.
Acremonium pertusariae

61
Acremonium s. lat.
Gams (1971) treated just one species with subspherical co-
nidia, A. pteridii W. Gams & Frankland, but these were much
Nectriopsis physciicola, Luxembourg, Diederich 19664. Colonies of the asexual stage over the thallus of Physcia tenella. Arrows point at
ascomata. Scale bar: 200 μm.
smaller, 2.4–2.8 × 2.2–2.4 μm. In the four known specimens,
the asexual stage developed close to ascomata.
Asexual stage of Nectriopsis physciicola, Luxembourg, Diederich 19664. A, Conidiophores, conidiogenous cells and young conidia. B. Mature
conidia. In phloxine. Scale bar: 10 μm

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Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
Ecology and hosts. The asexual stage has been collected
on the thallus and apothecia of Physcia stellaris, P. adscen-
dens, P. aipolia and P. tenella, and also on adjacent Aman-
dinea punctata, Melanohalea exasperatula, Polycauliona
candelaria and P. polycarpa. The sexual stage has further
been reported from Heterodermia, Melanohalea exasper-
ata, Physcia biziana, P. leptalea, Physconia servitii and
Xanthoria parietina. A specimen on Phaeophyscia from
Ecuador may belong to this species (Etayo 2017).
Distribution. The asexual stage is known from Europe
(Austria; France; Luxembourg), the sexual stage from Eu-
rope (Austria; France; Germany; Ireland; Luxembourg;
Netherlands; Spain; Switzerland; UK) and Asia (Russia).
Specimens with the asexual stage examined. Austria: Tyrol, Hin-
tertux, 47°06’25”N, 11°40’25”E, 1515 m, on Physcia stellaris,
Melanohalea exasperatula and Polycauliona polycarpa, 2014,
Diederich 17765 (BR; also distributed in Hafellner Lichenic.
Biota 257). France: Puy-de-Dôme: E of Saint-Ours, Beauregard,
930 m, on Physcia aipolia, 2021, Pinault (herb. Pinault). Luxem-
bourg: E of Kehlen, cemetery of Schéimerech, on Amandinea
punctata and Polycauliona candelaria, 1997, Diederich 12929
(BR); Strassen, Tossebierg, on Physcia adscendens and P. tenella,
2023, Diederich 19664 (BR).
References. Earland-Bennett et al. 2006 [Brackel 2011, 2015, 2023,
Cezanne et al. 2023, Hafellner 2017, Hitch 2010, Roux 2020, van der
Kolk et al. 2024, Zhurbenko 2014, Zimmermann & Feusi 2018; brit-
ishlichensociety.org.uk; www.wsl.ch/map_fungi; lichenportal.org].
Trichothecium roseum (Pers.) Link
Mag. Neuesten Entdeck. Gesammten Naturf. Ges. Naturf. Fre-
unde Berlin 3: 18 (1809); Trichoderma roseum Pers., Neues
Mag. Bot. 1: 92 (1794); Hyphelia rosea (Pers.) Fr., Syst. Orb.
Veg. 1: 149 (1825). Type: not located.
Nectriopsis physciicola
Trichothecium roseum, France, Pinault. A, Colonies on the thallus of Xanthoria parietina. B, Conidiophores with attached conidia. C, Same,
lateral view. D, Conidia, E Conidiophores with a few lateral branches. F, Conidiogenous cells with attached young conidia. G, Conidia. D–G,
in water. Photos: P. Pinault. Scale bars: A = 2 mm, B–C = 200 μm, E = 20 μm, D, F–G = 10 μm.

63
Acremonium s. lat.
Sexual stage unknown. Colonies supercial, euse, white
to pale pinkish. Conidiophores erect, simple to sparsely
branched, single or in loose groups, septate, hyaline, thin-
walled, smooth, 140–340 μm tall, 2.5–4 μm diam. Con-
idiogenous cells discrete, terminal, hyaline, thin-walled,
smooth or verruculose, retrogressive. Conidia hyaline,
ellipsoidal-clavate to pyriform, 1-septate, the upper cell
wider, the lower cell slightly curved, smooth- and thick-
walled, 13–25 × 6–11 μm, with an obliquely prominent
truncate base; the rst conidium is born as a blowout from
the side of the conidiogenous cell apex; the next conidium
is produced just below the rst one, but on the opposite
side; successive conidia are thus produced in a characteris-
tic zigzag patterned chain.
Notes. A well-characterized species with large, 1-septate
conidia with a curved base, produced on conidiophores in
zigzag-like patterns.
Ecology and hosts. A common and widespread saprophyte,
occasionally growing on lichens, e. g., on Athallia pyracea
and Xanthoria parietina.
Distribution. A widespread, cosmopolitan fungus, reported
as facultative lichenicolous in France and Ukraine. No map
is given.
Lichenicolous specimen examined. France: Charente-Maritime:
Île d’Oléron, on Xanthoria parietina, 2023, Pinault (herb. Pinault).
References. Hawksworth 1979 [Darmostuk 2020, Darmostuk et
al. 2018, Roux 2020].
Trichonectria rubefaciens (Ellis & Everh.) Diederich &
Schroers
in Sérusiaux et al., Lejeunia n. S. 162: 57 (1999); Nectria rube-
faciens Ellis & Everh., J. Mycol. 3: 116 (1887); Nectriopsis
rubefaciens (Ellis & Everh.) M. S. Cole & D. Hawksw., My-
cotaxon 77: 322 (2001) [as ‘rubifaciens’]. Type: USA, New
Jersey, Neweld, on dead fallen Acer limb, on the thallus of
Parmotrema cetratum, 25 Sept. 1887, J. B. Ellis (NY – lecto-
type, designated by Lowen 1995).
= Acremonium rhabdosporum W. Gams, Cephalosporium-art.
Schimmelpilze: 136 (1971); Cylindromonium rhabdosporum
(W. Gams) Crous, in Crous et al., Persoonia 43: 313 (2019).
Type: Austria, Innsbruck, near Aldranser Aim, isol. ex Clado-
nia sp., Oct. 1965, M. Gams (CBS 438.66 – holotype; K IMI
223813 – isotype).
Syn. nov.: Cylindromonium lichenophilum Crous, in Crous et al.,
Persoonia 51: 393 (2023). Type: South Africa, Mpumalanga
Prov., Buelskloof Nature Reserve, on Parmelina tiliacea, 5
Aug. 2022, P. W. Crous 3985 (CBS H-25276 – holotype; ex-
type culture CBS 150063).
incl. Trichonectria rubefaciens (Ellis & Everh.) Diederich &
Schroers subsp. cryptoramalinae Etayo & López de Silanes,
J. Fungi 2024, 10, 60. Type: Spain, Lugo, Courel, Mercurín,
42°37’58”N, 7°09’48.6”W, 670 m, grove with old chestnut
trees, on Castanea, on Ramalina fastigiata, 6 July 2021, J.
Etayo 33545 & M. E. López de Silanes (SANT-Lich. 12716 –
holotype; herb. Etayo – isotype).
Trichonectria rubefaciens, Austria, Berger 37400. Conidiophores with slimy conidial masses on the thallus of Parmelia saxatilis. Scale bars:
A = 200 μm, B = 100 μm.
Trichonectria rubefaciens, Luxembourg, Diederich 19650. Reddish
brown perithecia and white conidiophores on the thallus of Parmelia
sulcata. Scale bar: 500 μm.

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Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
Ascomata reddish brown, supercial, with a ‘crown’ of
short setae. Colonies of asexual stage supercial, euse.
Conidiophores erect, unbranched, 0–2-septate, hyaline,
thin-walled, smooth, 70–90 μm long. Conidiogenous cells
discrete, terminal, hyaline, thin-walled, smooth, narrowly
subulate to liform, phialidic with a narrow collarette,
40–80 μm long, basally 2–3.5 μm wide, apically 1–2 μm.
Conidia solitary or in a slimy mass, elongate-cylindrical,
rounded at both apices, hyaline, smooth and thin-walled,
often with annular thickenings at both ends, aseptate, 11.5–
27 × 2–3.5 μm.
Notes. This species is easily recognized by the brown,
supercial perithecia with a ‘crown’ of short, sti setae
around the ostiole and the elongate-cylindrical conidia.
It is a common parasite of macrolichens, mainly in very
humid and moist conditions, and both sexual and asexual
stages frequently co-occur. The anamorph-teleomorph
relationship has been proved by Lowen (1995) who ob-
tained the asexual stage from single ascospore isolations,
and Glenn et al. (1997) who obtained perithecia from the
asexual stage in laboratory conditions. Etayo & López de
Silanes (2024) described the new subspecies cryptorama-
linae for populations on Ramalina hosts, distinguished by
smaller perithecia, narrower asci and conidia lacking annu-
lar apical thickenings. We have observed a great variabil-
ity of conidia on Parmeliaceae hosts, including specimens
in which conidial annular thickenings were not observed.
E. g., our specimen Diederich 19650 has perithecia 120–
160 μm diam. and is thus typical T. rubefaciens s. str., al-
though conidia have no apical annular thickenings (see
photos). More studies will be needed to identify whether
specimens with smaller perithecia deserve recognition as
Asexual stage of Trichonectria rubefaciens, Luxembourg, Diederich
19650. Conidiophores, conidiogenous cells and conidia without api-
cal annular thickenings. Scale bar: 10 μm.
Trichonectria rubefaciens, Austria, Berger 37400. Long and narrow,
subcylindrical conidia, showing the annular thickenings at both ends.
Scale bar: 5 μm.
Asexual stage of Trichonectria rubefaciens, Austria, isotype (modied
from Hawksworth 1979). Conidiophore, conidiogenous cell and conidia,
showing the annular thickenings at both ends. Scale bar: 10 μm.

65
Acremonium s. lat.
Asexual stage of Pronectria santessonii, Italy, Brackel 4398 (modi-
ed from Brackel 2008). Conidiophores, conidiogenous cells and co-
nidia. Scale bars: 10 μm.
a distinct taxon. The recently described Cylindromonium
lichenophilum is morphologically and phylogenetically
indistinguishable from T. rubefaciens and is therefore con-
sidered as a younger synonym.
Ecology and hosts (specimens of subsp. cryptoramalinae
not considered here). On the thallus of Cladonia sp., Ever-
nia prunastri, Flavoparmelia caperata, Myelochroa sub-
aurulenta, Parmelia protosulcata, P. saxatilis, P. squarro-
sa, P. sulcata, Parmelina tiliacea, Parmotrema cetratum,
P. reticulatum, Platismatia glauca, Pleurosticta acetabu-
lum, Pseudevernia furfuracea, Punctelia jeckeri, P. sub-
rudecta, Ramalina farinacea, Usnea sp., Xanthoparmelia
conspersa and X. stenophylla.
Distribution. Europe (Austria; Belarus; Belgium; Czech Re-
public; Estonia; France; Germany; Italy; Latvia; Lithuania;
Luxembourg; Netherlands; Norway; Russia; Spain; Sweden;
Switzerland; UK: England, Scotland; Ukraine), Macaron-
esia (Canary Islands: La Gomera; Madeira), North America
(USA: Connecticut, Georgia, Maryland, New Jersey, New
York, North Carolina, Rhode Island, Virginia), South America
(Chile; Ecuador), Africa (South Africa) and Asia (Russia).
Selected specimens examined. Austria: Niederösterreich: Bez.
Scheibbs, Wildnisgebiet Dürrenstein, 47°46’42”N, 15°05’34”E,
on Parmelia saxatilis, 2023, Palice (herb. Berger 37400). Lux-
embourg: Strassen, 49.6231°N, 6.0425°E, 320 m, branch lying
on the ground, on Parmelia sulcata, 2022, Diederich 19650 (BR).
Madeira: Rabaçal, 32.8067°N, 17.1408°W, 1225 m, on Usnea,
2023, Diederich 19719 (BR).
Selected references. Gams 1971, Hawksworth 1979 [Brackel 2008,
2010, 2014, Brackel & Wirth 2023, Darmostuk et al. 2020, Died-
erich 1989, Diederich et al. 1991, Etayo 1996, 2010, 2017, Etayo
& López de Silanes 2008, Etayo & Sancho 2008, Hafellner 2009,
John et al. 2011, Kocourková 2009, Kuznetsova et al. 2012, Lowen
1995, Motiejūnaitė et al. 2012, 2016, Roux 2020, Tsurykau et al.
2013, van den Boom & van den Boom 2006, Zhurbenko 2014,
Zimmermann & Berger 2018, Zimmermann & Feusi 2021; GBIF;
www.verspreidingsatlas.nl; britishlichensociety.org.uk].
Pronectria santessonii (Lowen & D. Hawksw.) Lowen
Mycotaxon 39: 462 (1990); Nectriella santessonii Lowen & D.
Hawksw., Lichenologist 18: 322 (1986). Type: England, Dev-
on, Dartmouth, Warren Cove below Combe Point, on rocks
facing the sea, in the thallus of Anaptychia runcinata, 5 Aug.
1986, D. L. Hawksworth & R. Lowen 141 (K IMI 305040 –
holotype; NY, UPS – isotypes).
Ascomata reddish brown perithecia immersed in the host thal-
lus. Conidiophores erect, 0(–1)-septate (in vitro up to 3-sep-
tate), simple or branched in the middle, hyaline; wall nely
verrucose and wavy in outline. Conidiogenous cells discrete,
terminal, hyaline, phialidic, collarette not enlarged, 26–50 μm
long, basally 3–4 μm wide, apically c. 2 μm. Conidia solitary,
ellipsoidal, apically rounded, basally truncate, aseptate, hya-
line, biguttulate, (8–)8.5–10.5(–11) × 4–5(–5.5) μm (in vitro
in slimy droplets, 0(–1)-septate, 4–8 × 2.5–4 μm).
Notes. This species was typied on a sexual specimen on An-
aptychia runcinata (Lowen & Hawksworth 1986), but has
later also repeatedly been found on A. ciliaris (Berger et al.
2020). Lowen & Hawksworth (1986) obtained a single-spore
isolate from the holotype producing an Acremonium-like an-
amorph. Brackel (2008) collected the asexual stage also in na-
ture on Anaptychia ciliaris, in the presence of the sexual stage.
Trichonectria rubefaciens
Pronectria santessonii

66
Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
Ecology and hosts. On the thallus of Anaptychia runcinata
and A. ciliaris.
Distribution. Europe (Finland; France; Ireland; Italy; Lith-
uania; Russia; Spain; Sweden; UK: England).
References. Brackel 2008, 2015, Lowen & Hawksworth 1986
[Berger et al. 2020, Etayo 1998, Motiejūnaitė 2011, Puolasmaa et
al. 2008, Roux 2020, Zhurbenko & Kobzeva 2014].
Trichonectria setadpressa Etayo
Biblioth. Lichenol. 84: 134 (2002). Type: Colombia, Dept. Nar-
iño, Pasto, El Encano, vereda Sta. Isabel, S lago La Cocha,
páramo azonal, 2700 m, on Lobariella pallida, 30 July 1998,
J. Etayo 16582, S. Churchill, J. Muñoz & B. Ramírez (COL –
holotype; herb. Etayo – isotype).
Ascomata orange brown, supercial perithecia with ad-
pressed hairs. Colonies of the asexual stage pinkish
sporodochium-like aggregations of conidiophores on an
arachnoid subiculum, often with orange ‘droplets’ made of
conidia. Conidiophores reduced to the conidiogenous cell.
Conidiogenous cells discrete, terminal, hyaline, straight or
bent, 10–30 μm long, basally 1–2.5 μm wide, tapering at
the apex. Conidia solitary, subspherical to broadly ellipsoi-
dal, not truncate, aseptate, hyaline, smooth, (2–)2.5–3.8(–
4.5) × 2–2.5(–3) μm.
Notes. The asexual stage of this species is very similar to
that of Globonectria cochensis, known from the same host
genus. It is distinguished by the pinkish orange vs white
mass of conidia, and the slightly broader conidia, 2–2.5
μm vs 1.5–2.2 μm
Ecology and hosts. On the thallus of Lobariella pallida, L.
crenulata and L. subexornata.
Distribution. South America (Bolivia; Colombia; Ecuador).
References. Etayo 2002, 2017, Flakus et al. 2019.
Acremonium spegazzinii D. Hawksw.
Bull. Brit. Mus. (Nat. Hist.), Bot. 6: 195 (1979); Verticillium li-
chenicola Speg., Bol. Acad. Nac. Ci. 11: 612 [p. 234 of reprint]
(1889, as ‘lichenicolum’). Type: Brazil, on decayed thallus of
Leptogium andinum, 1880, J. Puiggari 127 (LPS 11.339 – hol-
otype). Not Acremonium lichenicola W. Gams (1971).
Sexual stage unknown. Colonies supercial, tufted, white.
Conidiophores erect, dispersed or aggregated into lax tufts
200–400 μm diam., irregularly branched, hyaline, verrucu-
lose, mainly 3–5 μm wide. Conidiogenous cells discrete,
terminal, hyaline, thin-walled, smooth to verruculose,
subulate, phialidic with a broad collarette, 15–60 μm tall,
basally 2.5–3.5 μm wide, apically 1.5–2.5 μm. Conidia
solitary, adhering in a slimy mass, elongate ellipsoidal,
rounded at both apices, aseptate, hyaline, smooth (minute-
ly verruculose?), 4–6 × 2–3 μm.
Notes. This species is distinguished from most other li-
chenicolous taxa by the branched verruculose conidi-
Asexual stage of Trichonectria setadpressa, Bolivia, Flakus 29612
(modied from Flakus et al. 2019). Conidia in LPCB. Scale bar: 10 μm.
Asexual stage of Trichonectria setadpressa, Bolivia, Flakus 29612
(modied from Flakus et al. 2019). Colonies on the thallus of Lobari-
ella pallida. Arrows point at conidial droplets. Dark reddish brown
perithecia are also present. Scale bar: 200 μm.
Trichonectria setadpressa

67
Acremonium s. lat.
ophores forming lax sporodochial tufts, and by the dimen-
sions of the conidia.
Ecology and hosts. On the thallus of Leptogium andinum, L.
cyanescens, L. cf. vesiculosum and Scytinium lichenoides.
Distribution. Europe (Italy), Macaronesia (Azores: Ter-
ceira) and South America (Brazil; Ecuador).
References. Brackel 2015, Hawksworth 1979 [Etayo 2017, 2018].
Acremonium a. spegazzinii
Notes. The two specimens examined strongly resemble
Acremonium spegazzinii, but dier by the smooth-walled
conidiophores and conidiogenous cells. The conidiophores
are irregularly branched, the conidiogenous cells 17–35
μm tall, basally 2.5–3 μm wide, and conidia are ellipsoidal,
5–6 × 2.3–2.8 μm. Molecular data will be needed to iden-
tify whether they represent a distinct species.
Ecology and hosts. On the thallus of Cladonia sp. and Lep-
raria incana.
Distribution. Europe (France; Luxembourg). No map is given.
Acremonium spegazzinii (Brazilian locality not accurate)
Acremonium a. spegazzinii, Luxembourg, Diederich 8643. A, Colonies on the thallus of Lepraria incana. B, Conidia. C–D, Conidiophores
and conidiogenous cells. B–D, in KOH. Scale bars: A = 200 μm, B–D = 10 μm.
Acremonium spegazzinii, Brazil, holotype (modied from Hawks-
worth 1979). Conidiophores, conidiogenous cells and conidia. Scale
bar: 10 μm.

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Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
Specimens examined: France: Puy-de-Dôme: Anzat-le-Luguet,
on Cladonia, 2022, Pinault (herb. Pinault). Luxembourg: NW of
Dalheim, Kinneksbierg, 49.56°N, 6.23°E, on Fagus, on Lepraria
incana, 1987, Diederich 8643 (BR).
Sarocladium strictum (W. Gams) Summerb.
in Summerbell et al., Stud. Mycol. 68: 158 (2011); Acremonium
strictum W. Gams, Cephalosporium-art. Schimmelpilze: 42
(1971). Type: Germany, Schleswig-Holstein, Kiel, Kitze-
berg, isolated from leaves of Triticum (CBS 346.70 – holo-
type).
In vitro: Sexual stage unknown. Colonies supercial, ef-
fuse. Conidiophores in slimy heads, erect, unbranched,
rarely basally branched, 0(–1)-septate, hyaline, smooth.
Conidiogenous cells discrete, terminal, hyaline, thin-
walled, smooth, narrowly subulate to subcylindrical, phia-
lidic, collarette not enlarged, 20–40(–65) μm long, basally
1.5–2(–2.5) μm wide, apically 0.5–1 μm. Conidia solitary,
ellipsoidal to subcylindrical, rounded at both apices, asep-
tate, hyaline, smooth, 4–5.5(–7) × 1–2(–3.5) μm.
Notes. Following Gams (1971), this is one of the most
common and most variable species of Acremonium s. lat.
As no in vivo description is available, and as morpho-
logical characters in culture may considerably dier from
those in nature, we provisionally accept here lichenicolous
specimens with slightly deviating measurements as con-
specic. Specimen Diederich 19649 has slightly larger co-
nidia, 4.5–6.5 × 2–2.5 μm; similarly, specimen Diederich
17487 has conidia 4.8–7 × 2–2.5 μm; Etayo (1996) reports
conidia of c. 5 × 1.5 μm; Berger & Zimmermann (2016)
report conidia of 4–5(–6) × 2.5–3.5 μm and conidiogenous
cells 15–25 × 3.5–4.5 μm.
Acremonium a. spegazzinii, France, Pinault. A, Colonies on the thallus of Cladonia sp. B, Conidia. C–D, Conidiophores and conidiogenous
cells. B, in ammonia; C–D, in Congo red. Photos: P. Pinault. Scale bar (B–D): 10 μm.
Sarocladium strictum, Luxembourg, Diederich 19649. Colonies over
dying thallus of Parmelia sulcata. Scale bar: 200 μm.
Sarocladium strictum, Luxembourg, Diederich 19649. Conidiophores
and conidia. Scale bar: 10 μm.

69
Acremonium s. lat.
Ecology and hosts. On diverse substrata, including leaves of
Triticum, but most commonly isolated from soil. Facultatively
lichenicolous on Cladonia pyxidata, Flavoparmelia caperata,
Lobaria pulmonaria, Parmelia sulcata, Physcia stellaris and
Ricasolia virens.
Distribution. Widespread, incl. Germany, lichenicolous
populations reported from Europe (France; Luxembourg;
Ukraine), Macaronesia (Madeira) and North America
(USA: Alaska). No map is given.
Specimens examined. Luxembourg: W of Steinfort, old quarry,
49.6689°N, 5.9052°E, 320 m, on Cladonia pyxidata, 2012, Died-
erich 17487 (BR); Strassen, 49.6231°N, 6.0425°E, 320 m, branch
lying on the ground, on Parmelia sulcata, 2022, Diederich 19649
(BR).
References. Gams 1971 [Berger & Zimmermann 2016, Brackel
2024, Darmostuk & Khodosovtsev 2017, Etayo 1996, Spribille
et al. 2010].
 (Ellis & Galw.) Etayo & Brackel
in Brackel et al., Mycosphere 3: 857 (2012); Dendrodochium su-
beusum Ellis & Galw., J. Mycol. 6: 33 (1890). Type: USA,
New York, Farmington, on trunk of a pear tree, on Physcia
millegrana (and adjacent Candelaria concolor), Aug. 1889,
E. Brown, Ellis & Everh., N. Am. Fungi 2394 (B, BPI, ILLS,
K, MICH, M, MU, NEB, NY, PH, WIS – syntypes).
= Dendrodochium eusum Vouaux, Bull. Trimestr. Soc. Mycol.
Fr. 30: 315 (1914), nom. inval. (Art. 36.1).
Sexual stage unknown. Colonies supercial, euse, white to
very pale orange. Conidiophores erect, forming translucent,
applanate, irregular, gelatinized sporodochia mainly 300–500
μm diam., and compacted, hyaline, branched, irregularly sub-
verticillate with 2–3 phialides at each node. Conidiogenous
cells discrete, terminal, hyaline, thin-walled, generally dis-
tinctly roughened below, terminating at about the same level,
subcylindrical to subulate, phialidic with a broad collarette,
20–30(–35) μm long, basally 3–4.5 μm wide, apically 2–3
Acremonium subeusum, France, Pinault. A, Colonies on the thallus of Physcia sp. B–D, Mycelium, conidiophores, conidiogenous cells and
conidia. E–F, Conidia. B–E, in ammoniacal Congo red; F, in Carbomer. Photos: P. Pinault. Scale bars: B = 20 μm, C–F = 10 μm.

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Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
μm. Conidia solitary, slimy in mass, broadly ellipsoidal, api-
cally rounded, basally truncate, aseptate, hyaline, smooth,
rarely walled uneven, (6–)7–10.5 × 4–6 μm.
Notes. Etayo & Pérez-Ortega (2016) suggested that this spe-
cies may represent the asexual stage of Pronectria leptaleae.
Ecology and hosts. On the thallus of Candelaria concolor,
Physcia millegrana, Physcia sp. and Physconia venusta.
Distribution. Europe (France; Spain) and North America
(USA: New York).
Additional specimen examined. France: Puy-de-Dôme:
Creste, on Physcia, 2022, Pinault (herb. Pinault).
References. Brackel et al. 2012, Hawksworth 1979 [Berger et al.
2020, Etayo & López de Silanes 2020, Etayo & Pérez-Ortega 2016].
Patriciomyces valentinianus D. Hawksw.
in Cole & Hawksworth, Mycotaxon 77: 327 (2001). Type: USA,
Missouri, Carter Co., Peck Ranch Conservation Area, vicin-
ity of Stegall, Mountain re tower, 37°45’30”N, 91°12’W,
1200-1340 ft, on dead Carya, on Physcia stellaris, 12 Oct.
1997, M. S. Cole 7394 (MIN – holotype; NY – isotype).
Sexual stage unknown. Colonies supercial sporodochia,
loose to compact, pale to bright orange, 0.4–1 mm diam.
Conidiophores erect, hyaline, smooth-walled, repeatedly
sympodially branched, septate, (3.5–)4–5.5(–6) μm thick.
Conidiogenous cells arising terminally and laterally on the
conidiophores, hyaline, smooth-walled, phialidic, collarette
not enlarged, 30–55 μm long, basally slightly constricted,
3–4(–4.5) μm diam., gradually tapering to 1–1.5 μm, prob-
ably phialidic without visible collarette. Conidia in easily
disarticulating chains, narrowly biconical, one or both apices
attenuated and truncate by narrow hila, (0–)1-septate, not con-
stricted at the septum, hyaline, smooth, 17–20 × 5.5–6.5 μm.
Notes. This species is characterized by the 1-septate conidia
that are attenuated at one or both ends. Cole & Hawksworth
(2001) were unsure whether it is an opportunist saprotroph or a
pathogen killing the host thallus. The repeated discovery of ad-
ditional specimens on Physcia suggests that it might an obligate
lichenicolous fungus, possibly conned to Physcia hosts.
Ecology and hosts. On the thallus of Physcia stellaris, P.
cf. aipolia, P. americana and P. caesia.
Distribution. North America (USA: Arkansas, Indiana,
Missouri, North Carolina, Oklahoma).
Acremonium subeusum, USA, New York, isotype (K) (modied
from Hawksworth 1979). Conidiophores, conidiogenous cells and
conidia. Scale bar: 10 μm.
Acremonium subeusum
Patriciomyces valentinianus, USA, Indiana, Scott 7990. Conidiomata
on the thallus of Physcia caesia. Scale bar: 500 μm.
Acremonium subeusum, USA, New York, isotype (M). Colonies over the
thallus of Candelaria concolor. Photo: W. von Brackel. Scale bar: 200 μm.

71
Acremonium s. lat.
Additional specimens examined. USA: Arkansas: Crawford Co.,
Ozark National Forest, Natural Dam, along Mountain Fork c. 0.2
mi N of conuence with Lee Creek, on Physcia sp., 2004, Har-
ris, 49113 (NY). Indiana: Marion Co., Sargent Road Nature Park,
39.9026°N, 86.0107°W, on corticolous Physcia caesia, 2022,
Scott 7990 (herb. Scott). North Carolina: Jackson Co., Cedar
Cli Mountain, along NC 281, 3.6 mi E of NC 107 at Tuckase-
gee, 35°15’N, 83°05’W, on P. americana, 1996, Harris 38674
(BR, NY). Oklahoma: Muskogee Co., Gruber Wildlife Manage-
ment Area, along OK 10, 3.3 mi N of Camp Gruber O.R.V. Area,
on Physcia cf. aipolia, 2004, Harris 49064 (NY).
References. Cole & Hawksworth 2001 [Diederich 2003; GBIF].
Acremonium vanderkolkii Boers & Diederich, sp. nov.
Diagnosis: Diers from all other known lichenicolous Hypocre-
ales species by an asexual sporodochial stage with elongate,
5‒11-septate conidia, 45‒75 × 2.5‒4 μm.
Etymology: Named after Henk-Jan van der Kolk, Dutch ecologist
and expert of lichenicolous fungi.
Type: Netherlands, Groningen, Rottumerplaat, 53.54°N, 6.49°E,
on a wooden post on the beach, on Lecanora sp., 10 Febr.
2023, J. Boers 16 (BR – holotype).
MycoBank: MB855945
Assumed sexual stage uncommon, orange-red, hairy, su-
percial perithecia, 135 μm diam. (n = 1). Colonies of the
asexual stage yellowish orange to pinkish sporodochium-
like aggregations of conidiophores and conidia develop-
ing over an often dense, white, supercial mycelium. My-
Patriciomyces valentinianus, USA, Indiana, Scott 7990. A, Conidia.
B, Conidiogenous cells. In water. Scale bar: 5 μm.
Patriciomyces valentinianus
Patriciomyces valentinianus, USA, Missouri, holotype (modied from
Cole & Hawksworth 2001). Conidiophores, conidiogenous cells and
conidia. Scale bars: 10 μm.
Patriciomyces valentinianus, USA, Oklahoma, Harris 49064. Conid-
ia, in KOH. Scale bar: 10 μm.

72
Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
celium immersed to more commonly supercial, hyphae
hyaline, 1.5‒2.5 μm diam. Conidiophores subcylindrical,
septate, branched, relatively short, or reduced to the con-
idiogenous cell. Conidiogenous cells monophialidic, elon-
Acremonium vanderkolkii, Netherlands. A, holotype; B, ‘topotype’. Conidiophores, conidiogenous cells and conidia, in phloxine. Scale bar: 10 μm.

73
Acremonium s. lat.
gate ellipsoidal to subcylindrical, 12‒36 × 3‒5 μm, with
a distinct, slightly aring collarette. Conidia dry, hyaline,
subcylindrical, straight to more frequently slightly curved,
irregular, attenuated and obtuse towards both apices,
5‒11-septate, not or slightly constricted at the septa, 45‒75
× 2.5‒4 μm, thin- and smooth-walled, basally indistinctly
truncate, hilum 0.8‒1.5 μm.
Notes. This species is distinguished from all other known
asexual lichenicolous Hypocreales species by the particu-
larly long and multiseptate conidia. Although repeated
attempts to culture and sequence the species failed, we
describe it nevertheless as new, based on the remarkable
morphological features. It may be related to Campylo-
carpon, but we prefer to describe it in Acremonium s. lat.,
awaiting molecular data.
Ecology and hosts. On corticolous or lignicolous lichens,
including a sterile green lichenized crust of goniocysts
(Micarea sp.?), Lecanora sp., Lecidella elaeochroma and
Lepraria sp.
Distribution. Europe (Luxembourg; Netherlands), in the
Netherlands known only from the isles in the Wadden Sea.
Additional specimens examined. Luxembourg: Dudelange, Ge-
haansbierg, 49.4886°N, 6.0606°E, 380 m, on Fagus, over Lepraria
sp. and a sterile green lichenized crust of goniocysts, 2023, Diederich
19654 (BR). Netherlands: Friesland: Ameland, 53.44°N, 5.85°E,
on wooden posts near the Wadden Sea, on Lecanora sp., 2020,
Boers 20 (herb. van der Kolk). Terschelling, dunes near Hoorn,
53.40°N, 5.35°E, on branches of Quercus robur, on Lecidella
elaeochroma, 2022, Boers 21 (herb. van der Kolk); ibid., dunes
near West-Terschelling, 53.37°N, 5.24°E, on branches of Q.
robur, on L. elaeochroma, 2022, Boers 22 (herb. van der Kolk).
Groningen: Same locality as type, on Lecidella elaeochroma, Bo-
ers 17 (BR – ‘topotype’).
Species known only from culture
Acremonium psychrophilum C. Möller & W. Gams
Mycotaxon 48: 445 (1993). Type: Antarctica, King George Isle,
near Argentinian station Jubany (now Carlini Base), isolated
from Mastodia tessellata, Dec. 1991, C. Möller (CBS 732.92
– holotype; ex-type culture).
In vitro: Sexual stage unknown. Mycelium of hyaline,
smooth, thick-walled (chondroid) hyphae, 1–2 μm diam.
Conidiophores often reduced to the conidiogenous cell, or
Acremonium vanderkolkii, Netherlands, holotype. Colonies over the
thallus of Lecanora sp. Scale bar: 500 μm.
Acremonium vanderkolkii, Netherlands, ‘topotype’. Colonies over
the thallus of Lecidella elaeochroma. Scale bar: 500 μm.
Acremonium vanderkolkii, Luxembourg, Diederich 19654. Colonies
over sterile corticolous lichen thalli. Arrow points at perithecium of
the assumed sexual stage. Scale bar: 500 μm. Acremonium vanderkolkii

74
Diederich, Ertz & Braun – Flora of Lichenicolous Fungi – Vol. 2. Hyphomycetes – 2024
a short cell, bearing several conidiogenous cells. Conidio-
genous cells straight or slightly curved, tapering towards the
apex, smooth, thick-walled, phialidic, rarely with a distinct
collarette, 20–70 μm long, basally 1.8–2.8 μm diam., api-
cally 0.8–2 μm. Conidia in slimy heads, hyaline, smooth,
ellipsoidal to subcylindrical, aseptate, 5–13.5 × 2–2.5 μm.
Notes. No in vivo description of this species is available.
The authors reported two isolates, CBS 732.92 and 139.93,
obtained from the same locality and from the same host.
The rst one was chosen as the type, whilst the second one
has been successfully sequenced by Hou et al. (2023). In
their phylogeny, the species was sister to Acremonium al-
ternatum, the type species of Acremonium.
Etayo et al. (2023) collected an Acremonium on Mastodia
tessellata on Livingston Island in Antarctica and identied
it as A. psychrophilum. It formed ‘white dots’ on the host
thallus and had simple, occasionally septate conidiophores,
conidiogenous cells 30–58 μm long, basally 2–3 μm diam.,
apically 1–2 μm, and ellipsoidal, straight, 0–1-septate co-
nidia, 5–9(–12) × 1.5–2.5 μm. They did not mention the
presence of chondroid hyphae, nor of thick-walled phial-
ides. In their phylogeny, it was sister to Acremonium ruti-
lum W. Gams, not closely related to A. alternatum. These
authors almost surely collected a distinct species on the
same host, with similar morphological characters.
Ecology and host. Isolated from the thallus of Mastodia
tessellata.
Distribution. Antarctica (South Shetland Islands), known
only from the type locality.
References. Hou et al. 2023, Möller & Gams 1993.
Excluded species
Paracosmospora physciae Crous & Boers
in Crous et al., Persoonia 48: 273 (2022). Type: Netherlands,
Prov. Drenthe, Dwingeloo, 52.8359°N, 6.3644°E, isolat-
ed from ‘Physcia tenella’, 10 March 2021, J. Boers (CBS
H-24952 – holotype; ex-type culture CBS 148942; original
specimen CBS HPC 3602).
Notes. No in vivo description of the original specimen has
been provided. A re-examination has shown that it repre-
sents hardly recognizable remnants of cf. Physcia thalli,
entirely overgrown by a blackish microbial lm. No fungus
was visible in the dried herbarium specimen, but it is obvi-
ous that the fungus was isolated from this microbial lm,
and not directly from a lichen thallus. The species has thus
to be excluded from the fungi isolated from lichens.
Acknowledgements
Curators of C (loan of type of Galloea cladoniicola), CBS (Para-
cosmospora physciae) and NY (Patriciomyces valentinianus), F.
Berger (Trichonectria rubefaciens), R. Common (C. dirinariae),
P. Scott (P. valentinianus), M. P. Zhurbenko (C. dirinariae), A.
Flakus (permission to reproduce photographic plates of Glo-
bonectria cochensis and Trichonectria setadpressa), G. Marson
(photos of C. lichenicola).
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Supplementary material
https://www.mnhn.lu/pub/FloraLichenicolousFungi