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Mycosphere Doi 10.5943/mycosphere/3/5/10
854
Notes on three lichenicolous species of
Acremonium including two new species
Brackel Wv1*, Etayo J2 and Lechat C3
1Wolfgang von Brackel, Institut für Vegetationskunde und Landschaftsökologie, Georg-Eger-Str. 1b, D-91334
Hemhofen, Germany. – e-mail: wolfgang.von.brackel@ivl-web.de
2Javier Etayo, Navarro Villoslada, 16-3° dcha., E-31003 Pamplona, Spain. – e-mail: jetayosa@pnte.cfnavarra.es
3Christian Lechat, Ascofrance, 64 route de Chizé, 79360 Villiers en Bois, France. – e-mail: lechat@ascofrance.fr
Brackel Wv, Etayo J, Lechat C 2012 – Notes on three lichenicolous species of Acremonium.
Mycosphere 3(5), 854–862, Doi 10.5943 /mycosphere/3/5/10
Two new species of lichenicolous fungi: Acremonium pertusariae on Pertusaria spp. and A.
bavaricum on Melanelixia glabratula are described. A new combination is proposed for
Dendrodochium subeffusum.
Key words – Anamorph fungi – Bionectriaceae – Hyphomycetes
Article Information
Received 4 September 2012
Accepted 14 September 2012
Published online 30 September 2012
Corresponding author: Wolfgang von Brackel – e-mail – wolfgang.von.brackel@ivl-web.de
Introduction
The genus Acremonium currently
comprises more than 100 species (Diederich
& Braun 2009); most of them are
saprophytes, but some are parasites of other
organisms (Gams 1971, Hawksworth 1972).
Hawksworth (1979) recorded four Acremo-
nium species known only from lichens and
one more fortuitously lichenicolous. Now
nine species are known as living on lichens,
of which seven are described in the genus
Acremonium and two as anamorphs of
Pronectria. Within the genus Acremonium
Gams (1971) and Morgan-Jones & Gams
(1982) described or combined the sections
Albo-lanosa, Chaetomioides, Gliomastix,
Nectroidea, and Simplex (Acremonium).
Lowen (1995) added the section
Lichenoidea, in which most of the
lichenicolous species are included.
During our studies in Spain, southern
Italy, and southern Germany, we found two
undescribed taxa of Acremonium as
probable parasites on lichens. The two new
species are described and illustrated in this
paper and a species of Dendrodochium is
recombined into Acremonium.
Materials and methods
Morphological and anatomical
observations were made using standard
microscopic techniques. Microscopic
measurements were made on hand-cut
sections mounted in water with an accuracy
up to 0.5 µm. Measurements of conidia are
recorded as (minimum–)
X
X-σ
–
X
X+σ
(–
maximum) followed by the number of
measurements. The specimens are deposited
in the private herbaria of the authors, the
holotypes of Acremonium pertusariae and
A. bavaricum in M, cultures of the
anamorphs at CIRM.
Mycosphere Doi 10.5943/mycosphere/3/5/10
855
Results
Acremonium pertusariae Brackel & Etayo,
sp. nov. Figs 1–2
MycoBank 519867
Etymology – pertaining to the host
genus Pertusaria.
Diagnosis – Coloniae superficiales,
pulviniae, pallide roseae, 200–300 µm diam.
Hyphae hyalinae, septatae, leves, 3–4 µm
latae. Conidiophora ramosa, septata,
hyalina. Cellulae conidiogenae hyalinae,
leves, subulatae, phialidicae, (15–)25–45 ×
1.5–4 µm. Conidia solitaria, levia, guttulata,
late ellipsoidea, apici rotundo, basi truncata
vel rotunda, hyalina, (4.5–)5–7(–8.5) × (3–
)3.4–4.3(–4.5) µm. Clamydosporae absunt;
teleomorphosis ignota.
Holotype – Italy, Basilicata, Prov. di
Potenza, Bosco Teduri near Bagni, old
beech forest, on Pertusaria pertusa on
Fagus sylvatica, 1260 m, 40°06′26′′N,
15°58′25′′E, W. & G. v. Brackel, 16.8.2010
(M – holotypus, hb ivl 5498 – isotypus,
culture deposited at CIRM).
Description – Colonies discrete to
confluent, superficial, tufted, pale pinkish,
building sporodochial-like cushions 200–
300 µm; mycelium partly immersed, hyphae
flexuose, thin-walled, smooth, hyaline, 3–4
µm wide. Conidiophores semi-macrone-
matous, richly branched, septate, hyaline,
smooth. Conidiogenous cells discrete,
terminal, hyaline, thin-walled, smooth,
subulate, phialidic, (15–)25–45 µm long, 2–
4 µm wide at the base and 1.5–2 µm at the
apex. Both conidiophores and conidio-
genous cells contain many small (or later
some big) oil guttules. Conidia solitary,
broadly ellipsoid, rounded at the apex,
slightly truncate at the base or rounded,
simple, hyaline, smooth, guttulate, (4.5–)5–
7(–8.5) × (3–)3.4–4.3(–4.5) µm, l/b = (1.1–
)1.3–2.4(–2.8), (n = 72). Clamydospores
absent, teleomorph unknown.
Features in culture at 25°C at day
light on Difco PDA containing 5mg/L
streptomycin– Colonies fast growing, grey-
brown whitish from above, brownish from
below, very fluffy, abundantly producing
conidia. Vegetative hyphae thin-walled,
smooth, hyaline, 2–4 µm wide.
Conidiophores semi-macronematous, bran-
ched, septate, hyaline, smooth. Conidioge-
nous cells discrete, terminal, hyaline, thin-
walled, smooth, subulate, phialidic, c. 15–30
µm long, 1.5–2 µm wide at the base and 1–
1.5 µm at the apex. Both conidiophores and
conidiogenous cells contain several
conspicious guttules. Conidia solitary,
ellipsoid, rounded at the apex, slightly
truncate at the base or rounded, simple,
hyaline, smooth, guttulate, (4–)4.5–7.4(–
9.5) × (2–)2.1–2.8(–3) µm, l/b= (1.6–)2–
2.9(–3.3) (n=20).
Distribution and hosts – Spain
(Aragón, Navarra, País Vasco), Canary
Islands (La Palma) and Italy (Basilicata), on
Pertusaria albescens, P. albescens var.
corallina, and P. pertusa, thallus and
apothecial warts. The fungus is apparently
parasitic, as the infected parts of the host
thallus become blackened and destroyed.
Uninfected parts of the thallus were healthy
and no other infecting fungus could be
found in the type and some of the Spanish
samples, but sometimes it lives together
with other fungi, especially Pronectria
pertusariicola. The latter species has
already been recorded, as A. aff. spegazzinii
D. Hawksw. in Etayo & López de Silanes
(2008) and Etayo (2010), on Pertusaria
albescens.
Discussion – The new species
belongs to the sect. Nectroidea, as the
hyphae are thin-walled and not chondroid,
and the conidiophores are richly branched. It
clearly does not belong to the sect.
Lichenoidea because of the lack of wall
thickenings in the phialides and the broadly
ellipsoid conidia (l/b ratio < 2). According
to Lowen (1995) there were only two
species of Acremonium found on lichens
that do not belong to the Lichenoidea: A.
psychrophilum (sect. Gliomastrix), and A.
strictum (sect. Acremonium); both are
distinguished from the new species by the
features of their section (chondroid hyphae
respectively unbranched conidiophores).
Also A. hypholomatis, which was recently
found growing on Physcia stellaris
(Diederich & Braun 2009), is a member of
the sect. Lichenoidea. Within the sect.
Nectrioidea only A. zeae has some similarity
Mycosphere Doi 10.5943/mycosphere/3/5/10
856
Fig. 1 – Acremonium pertusariae (holotypus) A conidiophores with conidiogenous cells and
conidia. B conidia.
with the new species (according to the key
in Gams 1971), but this species has
distinctly narrower conidia (3.5–5.8 × 1.2–
1.9 µm).
Acremonium is one of the genera
considered as anamorphs on Pronectria
(Bionectriaceae) species (Rossman et al.
1999). Five taxa of Bionectriaceae are
known as parasites on the genus Pertusaria:
Nectriopsis frangospora (on P. albescens,
P. rubefacta), N. hirta (on P. albescens, P.
pertusa), Pronectria pertusariicola (on P.
albescens, P. pertusa, Pertusaria spp.),
Pronectria sp. (sensu Etayo 2006, on
Pertusaria hymenea) and Trichonectria
pertusariae on P. amara var. slesvicensis
and P. ophtalmiza. Among these species P.
pertusariicola seems to be the more
widespread and common. It is very common
in humid beech forests in oceanic northern
Spain and not rare in the higher mountains
of southern Italy. Although not in the type
material, Acremonium pertusariae has been
collected on some occasions as well in
Spain, as in Italy, growing together or over
thallus infected with P. pertusariicola
(which might be the teleomorph of the new
species); in Spain sometimes also with a
Fusarium species which we will study in the
future. The presence of two different
conidial genera on Pertusaria thalli infected
by P. pertusariicola and the fact that several
thalli infected by Acremonium do not show
the presence of Pronectria, we cannot
ensure the teleomorph-anamorph connec-
tion. Moreover, the anamorph of P.
pertusariicola in culture is clearly distin-
guished from A. pertusariae (see below)
even in culture by larger conidiogenous
cells and longer and thiner conidia in A.
pertusariae.
Additional specimens – Spain:
Álava, entre Santa Cruz y Oteo, pequeño
cañón que atraviesa la carretera, on P.
albescens on Acer, 650 m, 42º42'13''N,
2º21'21''W, 4.9.2010, J. Etayo 26227 (hb.
Etayo). Álava, Muniain, Sª de Entzía,
robledal centenario de Muniain, on P.
albescens on Q. robur, 750 m, 42º50’N,
2º20’W, 6.2007, J. Etayo 24235 (hb. Etayo).
Ibidem, 9.2007, J. Etayo 24323 (hb. Etayo).
Ibidem, 26401 (hb. Etayo, culture deposited
Mycosphere Doi 10.5943/mycosphere/3/5/10
857
at CIRM). Álava, Kontrasta, bosque de
Quercus x pyrenaica, por sendero desde el
pueblo, 42º45'33"N, 2º16'56''W, 850 m,
26.8.2010, sobre P. albescens en grueso
Quercus sp., J.A. Azpilicueta & J. Etayo
26212 (hb. Etayo). Álava, Sª de Urbasa,
camino de Larraona a Puerto de Opacua,
cerca de la Fuente de la Sierra, en
Pertusaria pertusa sobre Acer
pseudoplatanus cercanos carretera, 990 m,
42º48'30''N, 2º19'34''W, 29.8.2010, J.A.
Azpilicueta & J. Etayo 26400, 26401 (hb.
Etayo, VIT). Huesca, Peña Ezcaurre, on P.
albescens var. corallina on Fagus, 1600-
1750 m, 28.9.2002, J. Etayo 19768 (JACA).
Navarra, Urdiain, on P. albescens on Q.
robur, 20.7.1991, J. Etayo 5962 (hb. Etayo).
Canary Isles, La Palma, Los Tilos, subida al
mirador de la Baranda, barranco or. NW., on
Pertusaria albescens, 12.3.1993, J. Etayo
651 (hb. Etayo). Italy: (together with
Pronectria pertusariicola) Basilicata, Prov.
di Potenza, between Laurenzana and Monte
Caldarosa, forest of beeches and firs, on P.
pertusa on Fagus, 1120 m, 40°24′24′′N,
15°57′26′′E, 18.8.2010, W. & G. v. Brackel
(hb ivl 5632). Type locality, (hb ivl 5600).
Material with thinner conidia –
France: Pyrenées-Atlantiques, 1 km del Col
d’Ibardin, La Redoute des Emigrés, on P.
albescens on Q. robur, 150 m, 14.3.1995, J.
Etayo 12849 & B. Marbach (hb. Etayo).
Pronectria pertusariicola – (hb.
Etayo 26401, locality already recorded
where both species live together). – Features
of the anamorph in culture – Colonies
slowly growing, pinkish orange under a
whitish cover from above, orange from
below, compact to folded in the centre,
zoned at the margins. Vegetative hyphae
thin-walled, smooth, hyaline, 3.5–5.5 µm
wide, often with swollen, short cells, 6–8 ×
7–9 µm. Conidiophores mostly branched,
septate, hyaline, rough, guttulate.
Conidiogenous cells discrete, terminal,
hyaline, thin-walled, rough, subulate,
phialidic, 15–50 µm long , c. 4 µm wide at
the base and c. 2 µm wide at the apex.
Conidia solitary, ellipsoid or slightly
irregular, rounded at both ends, simple,
hyaline, smooth, guttulate, (4.5–)4.8–6(–
6.5) × (2.5–)2.8–3.2(–3.5) µm, l/b = (1.3–
)1.6–2.1(–2.4) (n=20). Dendrodochium
subeffusum Ellis & Galw. was described on
Physcia millegrana and Candelaria
concolor from USA, and a concise
description of it appears in Hawksworth
(1979). It has all features of an
Acremonium.We have studied an isotype
(M) and there are no differences with other
members of the genus; conidiophores grow
together but do not form a distinguishable or
compact structure we can name as a
sporodochium. Another lichenicolous
species of Acremonium like A. spegazzinii
D. Hawksw., growing on tropical
Leptogium, has much more compact
conidiophores. For this reason we propose
the new combination of D. subeffusum in
the genus Acremonium:
Acremonium subeffusum (Ellis & Galw.)
Etayo & Brackel, comb. nov. Fig. 3
MycoBank 519871
Bas. Dendrodochium subeffussum
Ellis & Galw., J. Mycol. 6: 33 (1890).
USA., New York, Farmington, on
thallus of some foliaceous lichen, on
Physcia millegrana and Candelaria
concolor on trunks of pear trees, August
1889, E. Brown. (M! – isotype).
A. subeffusum has conidiogenous
cells distinctly roughened below and conidia
of (6–)7–9(–9.5) × 4.5–6 µm (Hawksworth
1972) with conidiogenous cells 20–30(–35)
µm tall, and 3–4.5 µm wide at the base and
2–3 µm at the tip.
In M another specimen named
Dendrodochium subeffusum from the USA
is listed in the catalogue (Triebel 2006–
2010: M-0041264). The host is Lobaria
quercizans (in the original label as Sticta
glomulifera, in the catalogue as
Xanthoparmelia verruculifera). We saw this
specimen and the fungus is quite similar in
several features to A. subeffusum: the
conidiophores are irregularly subverti-
cillately branched, the conidiogenous cells
are rough and the conidia are broadly
elliptic with a peg-like narrowly truncated
base. On the other hand it differs from A.
subeffusum in the much more compact
colonies, the only slightly rough and much
shorter (about 15 µm long)
Mycosphere Doi 10.5943/mycosphere/3/5/10
858
Fig. 2 – Acremonium pertusariae (holotypus): infected part of the thallus of Pertusaria
pertusa with pink cushions of conidiophores.
conidiogenous cells, and the smaller
conidia, 4.5–5 × 2.5–3 µm. More material
would be needed for further investigations
on this taxon surely different from A.
subeffusum.
Some of the specimens in Hb. Etayo
growing on Pertusaria have similar
conidiophores but the conidia are smaller
and thinner, 3.5–5 × 2–2.5 µm. For the
moment, we cannot assume that they belong
to the same species.
Acremonium bavaricum Brackel sp. nov.
Figs 5–6
MycoBank 519869
Diagnosis – Coloniae superficiales,
effusae, pellucidae. Hyphae hyalinae,
septatae, leves, 2–5 µm latae. Conidiophora
plerumque simplicia, raro septata et ramosa,
hyalina, levia. Cellulae conidiogenae
hyalinae, leves, anguste subulatae,
phialidicae, 35–50 × 1–3.5 µm. Conidia
solitaria, anguste ellipsoidea, apici rotundo,
basi truncata, hyalina, levia, (4–)4.5–5.6(–6)
× (1.5–)1.8–2.3(–2.5) µm. Clamydosporae
absunt; teleomorphosis ignota.
Holotype – Germany, Bavaria,
Oberpfalz, Kreis Regensburg, Karlswies-
bachtal, on Melanelixia glabratula on
Fraxinus excelsior, 435 m, 49°04'32,3''N,
12°18'51,6''E, W. v. Brackel, 28.10.2009 (M
– holotypus).
Description – Colonies superficial,
effuse, translucent. Mycelium partly
superficial, partly immersed, composed of
hyaline, cylindrical to irregular cells, 2–5
µm wide. Conidiophores semi-macronema-
tous, mostly simple and unbranched, rarely
septate and branched, hyaline, smooth, 3–15
× 3–5 µm. Conidiogenous cells discrete,
terminal, hyaline, thin-walled, smooth,
narrowly subulate, phialidic, 35–50 µm
long, 3–3.5 µm wide at the base and 1–1.5
µm apically. Conidia solitary, narrowly
ellipsoid, rounded at the apex, truncate at
the base, simple, hyaline, smooth-walled,
(4–)4.5–5.6(–6) × (1.5–)1.8–2.3(–2.5) µm,
l/b = (1.8–)2.1–2.3(–2.5), (n = 20), often
collecting in heads in liquid droplets at the
apices of the conidiogenous cells.
Clamydospores absent, teleomorph
unknown.
Features in culture – Colonies fast
growing, grey whitish from above, dark
grey from below, compact to folded in the
centre, slightly zoned at the margins,
Mycosphere Doi 10.5943/mycosphere/3/5/10
859
Fig. 3 – Acremonium subeffusum (isotypus) growing on corticolous lichens as Candelaria and
Physcia.
abundantly producing conidia. Vegetative
hyphae thin-walled, smooth, hyaline to very
pale brownish, 1.5–2 µm wide, septate,
single cells 10–20 µm long. Conidiophores
semi-macronematous, septate, hyaline,
smooth, or missing. Conidiogenous cells
discrete, terminal, hyaline, thin-walled,
smooth, phialidic, c. 40–45 µm long, 2 µm
wide at the base and 1 µm wide at the apex.
Conidia solitary, ellipsoid, rounded at the
apex, slightly truncate at the base, simple,
hyline, smooth, (2.5–)2.6–3.3(–3.5) × (1.5–
)1.7–2 µm, l/b = 1.3–1.9(–2.3) (n = 20).
Distribution and hosts – A.
bavaricum is known from two closely
situated localities in Germany, Bavaria. In
both specimens it is growing on Melanelixia
glabratula, mainly on the isidia. As the
isidia are slightly discoloured, the fungus is
presumed to be a weak parasite.
Discussion – With its soft colonies,
usually unbranched conidiophores and the
narrowly ellipsoid, hyaline conidia A.
bavaricum belongs to the sect. Lichenoidea.
In this section, only A. antarcticum and A.
spegazzinii have conidia of similar size.
Both are distingished from the new species
by the conidia rounded at both ends and the
different hosts (Caloplaca respectively
Leptogium). Furthermore, A. antarcticum is
distinguished by the much shorter and
narrower conidiogenous cells (15–20 × 1–2
µm), and A. spegazzinii by the verruculose
conidiophores. Following the key of Gams
(1971), no similar species of Acremonium
could be found.
On hosts of the former genus
Melanelia only two members of the
Bionectriaceae (and Nectriaceae) are
known, Paranectria oropensis and
Pronectria septemseptata. Anamorphs of
both species were not known until now.
Paranectria oropensis is widespread and
able to grow on several genera of hosts. It is
quite common in Bavaria and was found
also in the neighbourhood of the type
locality. The very rare Pronectria
septemseptata could be found only 3 km
Mycosphere Doi 10.5943/mycosphere/3/5/10
860
Fig. 4 – Acremonium on Lobaria quercizans (M-0041264), growing on the apothecial
margins.
distance from the type locality of A.
bavaricum, but on another host. So both
species seemed to be possible teleomorphs
of the new species but we have never found
them together. Recently we have found a
rich population of P. septemseptata in
northern Spain, but also there no
Acremonium found in the samples. From a
culture of the anamorph of P. septemseptata
we could see that it is clearly distinguished
from A. bavaricum in its plurilocular
conidiogenous cells and the brownish
conidia (see below). As Paranectria
oropensis also produces a different
anamorph in culture (Fusarium-like, with
fusiform and septate conidia), it is unlikely
that one of these species represents the
teleomorph of A. bavaricum.
Additional specimen – Germany,
Bavaria, Oberpfalz, Kreis Regensburg,
Otterbachtal W Bruckhäusl, on Melanelixia
glabratula on Alnus glutinosa, 400 m,
49°04'15,6''N, 12°17'26,3''E, W. v. Brackel,
28.10.2009 (hb ivl 5671).
Specimens of possible teleomorphs
found in the neighbourhood – Paranectria
oropensis: Germany, Bavaria, Oberpfalz,
Kreis Regensburg, Otterbachtal N
Unterlichtenwald, on Physcia tenella and
Lecania cyrtella on Salix fragilis, 355 m,
49°03'23''N, 12°15'59''E, W. v. Brackel,
14.11.2008 (hb ivl 5451). – Pronectria
septemseptata: Germany, Bavaria,
Oberpfalz, Kreis Regensburg, Otterbachtal
SE Heuweg, on Melanohalea elegantula on
Carpinus betulus, 390 m, 49°03'36''N,
12°16'48''E, W. v. Brackel, 14.11.2010 (hb
ivl 5449, culture deposited at CIRM).
Features of the anamorph of Pronectria
septemseptata in culture – Colonies fast
growing, whitish, becoming grey in the
centre, brownish from below, fluffy, zoned
at the margins, abundantly producing
conidia. Vegetative hyphae thin-walled,
hyaline to slightly brownish, 1.5–3.5 µm
wide, septate, single cells 10–20 µm long,
smooth, sometimes slightly rough.
Condiophores arising from the mycelium,
Mycosphere Doi 10.5943/mycosphere/3/5/10
861
Fig. 5 – Acremonium bavaricum (holotypus): A: conidiophores with mycelium,
conidiogenous cells, and conidia. B: outlines of conidia.
Fig. 6 – Acremonium bavaricum (holotypus): infected part of the thallus of Melanelixia
glabratula with the effuse, translucent colonies.
Mycosphere Doi 10.5943/mycosphere/3/5/10
862
mostly single, septate, c. 40 x 2 µm.
Conidiogenous cells integrated, terminal and
sometimes intercalary, with several (up to
20) conidiogenous loculi, dispersed over the
whole cell. Conidia ellipsoid, smooth,
slightly brownish to pale brown, both ends
rounded, sometimes with a minute scar,
with (0–)2 small guttules, (3–)3.2–3.9(–4) ×
2–2.4(–2.5) µm, l/b = (1.4–)1.5–1.9(–2)
(n=20).
Acknowledgements
We thank Dagmar Triebel, curator at
the herbarium of the Botanische
Staatssammlung München, for the loan of
specimens, and Roz Lowen (New York) for
sending separata.
References
Diederich P, Braun, U. 2009 – First
lichenicolous record of Acremonium
hypholomatis (anamorphic Ascomy-
cota). Bulletin de la Société des
Naturalistes Luxembourgeois 110,
97–100.
Etayo J. 2006 – Proyecto de estudio de los
líquenes y hongos liquenícolas de
Oieleku. URL:
http://ec.europa.eu/environment/life/
project/Projects/index.cfm?fuseactio
n=home.showFile&rep=book&fil=L
%C3%ADquenes_y_hongosliquen%
C3%ADcolas.pdf
Etayo J. 2010b – Líquenes y hongos
liquenícolas de Aragón. Guineana
16, 1–501.
Etayo J, López de Silanes ME. 2008 –
Líquenes epífitos y hongos
liquenícolas del Bosque Viejo de
Munain-Okariz (Álava, País Vasco,
España). Nova Acta Científica
Compostelana. Bioloxía 17, 5–23.
Gams W. 1971 – Cephalosporium-artige
Schimmelpilze (Hyphomycetes).
Jena: G. Fischer.
Hawksworth DL. 1972 – A large-spored
species of Acremonium sect.
Nectrioidea. Transactions of the
British Mycological Society 58,
510–512.
Hawksworth DL. 1979 – The lichenicolous
Hyphomycetes. Bulletin of the
British Museum (Natural History).
Botany 6, 183–300.
Lowen R. 1989 – Two new species of
Nectriella and an Acremonium
anamorph. Memoirs of the New
York Botanical Garden 49, 243–252.
Lowen R. 1995 – Acremonium section
Lichenoidea section nov. and
Pronectria oligospora species nov.
Mycotaxon 53, 81–95.
Morgan-Jones G, Gams W. 1982 – Notes on
Hyphomycetes. XLI. An endophyte
of Festuca arundinacea and the
anamorph of Epichloe typhina, new
taxa in one of the new sections of
Acremonium. Mycotaxon 15, 311–
318.
Rossman A Y, Samuels G J, Rogerson C T
& Lowen R. 1999 - Genera of
Bionectriaceae, Hypocreaceae and
Nectriaceae (Hypocreales, Ascomy-
cetes). Studies in Mycology 42, 1–
248.
Triebel D. 2006–2010 – The collection of
lichenicolous fungi at the Botanische
Staatssammlung München.
http://www.botanischestaatssammlun
g.de/ DatabaseClients/
BSMlichfungicoll
