New records of lichenized and lichenicolous fungi in Scandinavia

Abstract

Fourteen species of lichenized or lichenicolous fungi are reported new to either Norway or Sweden or both countries. Several of these are rare and almost unknown. The reported species are: Acarospora insignis (new to Norway), A. pyrenopsoides (Norway, Sweden), A. versicolor (Norway), Calvitimela perlata (Sweden), Lecidea degeliana (Sweden), Nephroma helveticum (Sweden), Peltula placodizans (Norway), Phacographa
protoparmeliae (Norway), Rhizocarpon pycnocarpoides (Norway, Sweden), Sarcogyne algoviae (Sweden),
Sarcogyne hypophaeoides (Norway, Sweden), Tephromela grumosa (Norway), Tremella lobariacearum
(Norway) and Tremella wirthii (Sweden). In addition Cladonia albonigra is confirmed from Sweden.
Sarcogyne hypophaeoides is lectotypified and is also reported new to Austria.

Full text

New records of lichenized and lichenicolous fungi in Scandinavia 33
New records of lichenized and
lichenicolous fungi in Scandinavia
Martin Westberg1, Einar Timdal2, Johan Asplund3, Mika Bendiksby4,
ReidarHaugan2, Fredrik Jonsson5, Per Larsson6, Göran Odelvik1,
Mats Wedin1, Ana M. Millanes7
1 Department of Botany, Swedish Museum of Natural History, P.O. Box 50007, SE-104 05 Stockholm,
Sweden 2 Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, NO-0318 Oslo, Norway
3Department of Ecology and Natural Resource Management. Norwegian University of Life Sciences, P.O. Box
5003, NO-1432, Ås, Norway 4 NTNU University Museum, Norwegian University of Science and Technology,
7491 Trondheim, Norway 5 Alsens-Ede 227, SE-830 47 Trångsviken, Sweden 6 Nordiska museet, Julita gård,
SE-643 98 Julita, Sweden 7 Departamento de Biología y Geología, Física y Química Inorgánica, Universidad
Rey Juan Carlos, C/ Tulipán s.n. E-28933 Móstoles, Spain
Corresponding author: Martin Westberg (martin.westberg@nrm.se)
Academic editor: G. Rambold|Received24 September 2015|Accepted 29 october 2015|Published 13 November2015
Citation: Westberg M, Timdal E, Asplund J, Bendiksby M, Haugan R, Jonsson F, Larsson P, Odelvik G, Wedin M,
Millanes AM (2015) New records of lichenized and lichenicolous fungi in Scandinavia. MycoKeys 11: 33–61. doi:
10.3897/mycokeys.11.6670
Abstract
Fourteen species of lichenized or lichenicolous fungi are reported new to either Norway or Sweden or both
countries. Several of these are rare and almost unknown. e reported species are: Acarospora insignis (new
to Norway), A. pyrenopsoides (Norway, Sweden), A. versicolor (Norway), Calvitimela perlata (Sweden),
Lecidea degeliana (Sweden), Nephroma helveticum (Sweden), Peltula placodizans (Norway), Phacographa
protoparmeliae (Norway), Rhizocarpon pycnocarpoides (Norway, Sweden), Sarcogyne algoviae (Sweden),
Sarcogyne hypophaeoides (Norway, Sweden), Tephromela grumosa (Norway), Tremella lobariacearum
(Norway) and Tremella wirthii (Sweden). In addition Cladonia albonigra is conrmed from Sweden.
Sarcogyne hypophaeoides is lectotypied and is also reported new to Austria.
Key words
Acarosporaceae, barcode, oristics, ITS, lichens
Copyright Martin Westberg et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
MycoKeys 11: 33–61 (2015)
doi: 10.3897/mycokeys.11.6670
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RESEARCH ARTICLE

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
34
Introduction
Although studies of the biodiversity in Scandinavia have a long and continuous tradi-
tion, several thousands of species remain to be discovered, many of which are likely
to be common. Among many of those discovered, almost nothing is known about
where they occur, how they live and what their functional roles in the ecosystems are.
To explore this poorly known biodiversity, the Swedish Taxonomy Initiative (STI)
was established in 2002, with the aim to have all multicellular plants, fungi and ani-
mals in Sweden identied and described. Numerous taxonomic projects have been
funded by the STI to investigate poorly known groups of organisms. Since 2002,
almost 2000 species have been found new to Sweden and almost 1000 are new to sci-
ence (Sundin 2014). In 2009, a similar project was established in Norway (the Nor-
wegian Taxonomy Initiative, NTI), aimed at mapping all organisms in the country.
Although the NTI does not provide funding for taxonomic research-projects, around
260 new species to science had been discovered in Norway until 2013 and the num-
ber of species new to Norway was approaching 1200 (http://www.artsdatabanken.
no/Article/Article/133228). e two countries have signed a declaration of coopera-
tion to increase the knowledge of and level of competence on biodiversity in Scan-
dinavia, and collaborative projects have emerged. Several projects in both countries
have focused on groups of lichenized and lichenicolous fungi, such as Acarosporaceae
(funded by STI), lecideoid lichens (NTI, STI), Tremellales (STI), etc. During the
last three years, lichen research groups in Oslo and Stockholm have collaborated and
arranged workshops for lichenologists involved in these projects and beyond. Crus-
tose, saxicolous lichens are in general very poorly known. Here, we add ten saxicol-
ous lichens to the Scandinavian ora, mainly found in the still poorly investigated
montane areas of Scandinavia. Also new macrolichens are continuously discovered in
Scandinavia (Frödén 2010, Hultengren et al. 2011, Klepsland 2013, Klepsland and
Timdal 2010) and here we report Nephroma helveticum new to Sweden, a rare and
declining species in Europe that needs attention and possible conservation actions.
In addition we conrm the presences of Cladonia albonigra in Sweden. Lichenicol-
ous fungi are understudied and a lot of research is needed on this group (Ihlen and
Wedin 2008). We have found three new species to the Nordic countries, one asco-
mycete, Phacographa protoparmeliae and two heterobasidiomycetes, Tremella lobari-
acearum and T. wirthii. ese new records were collected during the last eld seasons
by some of the people involved in the dierent research projects funded by the STI
and the NTI.
Material and methods
Material. We studied material collected during eld surveys (2011–2014) funded
through STI and NTI projects as well as specimens of the same species on loan from
various herbaria (GB, GZU, LD, O, S, STU, UPS). All specimens collected during
these eld trips are deposited in the herbaria O and S.

New records of lichenized and lichenicolous fungi in Scandinavia 35
Morphology. Macromorphological traits were observed using a Zeiss Stemi 2000-C
dissecting microscope. Microscopical examinations were carried out using squash prepa-
rations, hand-cut sections and freezing microtome sections cut at 16 μm. e prepara-
tions were observed in distilled water, lactophenol cotton blue, or 10% KOH (K). Micro-
scopic structures in heterobasidiomycetes were studied using handmade sections stained
with Phloxin (1% in water) after pre-treatment with KOH (5%), following the methods
of Diederich (1996), and observed with a Zeiss Axioscope-2 microscope. Macro- and
microscopic photographs were taken with a Nikon Coolpix 995 camera tted to both
microscopes or with a Nikon D300 with a macrolens with bellows. e macrophotos
were processed using focus stacking with the programs Helicon Focus and Combine ZP.
Secondary chemistry. Selected specimens were examined by thin-layer chromatog-
raphy (TLC), performed in accordance with the methods of Culberson (1972), as
modied by Menlove (1974) and Culberson and Johnson (1982).
DNA barcoding. Some specimens (Table 1) were DNA barcoded for the nuclear
ribosomal internal transcribed spacer region (ITS) through the Norwegian Barcode of
Life project (http://www.norbol.org/) or projects funded by the STI.
The species
Acarospora insignis H.Magn, 1924
MycoBank: MB 375596
Acarospora insignis H.Magn. Svensk Bot. Tidskr. 18: 329. 1924.
Holotype. SWEDEN. Härjedalen: Viken. 1878, P. J. Hellbom (GB 0129823!). TLC:
gyrophoric acid.
Table 1. List of sequenced specimens with Accession numbers in the European Nucleotide Archive or
GenBank.
Species Origin Voucher Acc. No. (ITS)
Acarospora insignis Norway, Oppland Haugan 10022 (O L-173397) LN890273
Norway, Oppland Westberg (S F265207) LN890274
Acarospora pyrenopsoides Norway, Aust-Agder Timdal 11308 (O L-163369) LN890275
Acarospora versicolor Norway, Buskerud Westberg 08-092 (S F268460) LN890276
Norway, Oppland Westberg 08-205 (S F268463) LN890277
Nephroma helveticum
Sweden, Västerbotten Jonsson 4200 (UPS L-193714) LN890278
Norway, Oppland Klepsland JK11-L559 (O L-181601) KT800006
Norway, Buskerud Hofton 13176 (O L-196491) KT800007
Rhizocarpon pycnocarpoides
Norway, Sør-Tröndelag Bendiksby et al. 12630 (O L-179560) KR780560
Norway, Oppland Rui & Timdal 12665 (O L-179903) KT800002
Norway, Buskerud Rui & Timdal 12854 (O L-184267) KT800003
Norway, Nordland Haugan 11125 (O L-183808) KT800004
Norway, Nord-Trøndelag Haugan 11128 (O L-183810) KT800005
Tephromela grumosa Sweden, Bohuslän Haugan 11501 (O L-184061) KR303667

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
36
Distribution. New to Norway. is species has until now only been known from
the type collection from central Sweden. Magnusson (1935) also mentioned a specimen
from East Greenland. At rst glance it appears to be an odd form of the common and
well-known A. fuscata with pale brown to reddish brown areoles and a black lower sur-
face that is typically spreading onto the edges of the upper surface (Fig. 1). e C+ red
reaction of the cortex (gyrophoric acid) is also present in A. fuscata. However, the round,
mostly scattered areoles with a single round apothecium and a very low hymenium, only
between 50–65 μm tall, identies A. insignis. A preliminary analysis of ITS sequences
indicates that it is a close relative to the North American species A. thamnina (ID of
specimens in GenBank of that species was conrmed by Kerry Knudsen, pers. comm.).
e Norwegian specimens were both collected on siliceous boulders in open
spruce forests. e type specimen grows on Aspicilia cinerea and Magnusson (1924)
concluded that it is a saprophyte. However, the newly found specimens do not grow
in direct association with other species. Among other species seen in the specimens,
Aspicilia cf. cinerea, Rhizocarpon grande and Trapelia obtegens were noted.
Specimens examined. NORWAY. Oppland: Lillehammer, Døsgrenda, between
Kinnlia and Åsen, alt. 500 m, 61°05.21'N, 10°20.32'E. 1 June 2011, Haugan 10022
(O L-173397); Lom municipality, Runningsgrende, Klypa, 61,7236°N, 8,0262°E. 28
June 2013, Westberg (S F265207), TLC: gyrophoric acid.
Acarospora pyrenopsoides H.Magn, 1926
MycoBank: MB 375684
Acarospora pyrenopsoides H.Magn. Acta Horti Gothob. 2: 74. 1926.
Holotype. GREENLAND. Nennese. [undated], J. Vahl (UPS L-067474!).
Distribution. New to Norway and Sweden. is is another poorly known species
that has received very little attention since Magnusson (1926) discovered it. It has until
now been known from scattered records from Greenland, Germany, Finland and Austria
(Magnusson 1935, Wirth et al. 2013). It has also been reported from Denmark and
Canada but those nds are considered erroneous (Søchting and Alstrup 2008, Knudsen
and Kocourková 2010). is is a characteristic species that has a thallus composed of
rounded, fairly large squamules (fertile squamules 0.5–3 mm wide), with an incised
or almost lobate margin (Fig. 2). e upper surface is attened with a brownish grey
colour. e lower side as well as the steep, vertical sides of the squamules are jet black
and somewhat shiny. e apothecia are immersed, at rst deeply sunken but later almost
level with the thallus. e disc is brown, somewhat shiny and can become somewhat
uneven in larger apothecia. Magnusson (1926) thought that the apothecia resembled
those in certain species of Pyrenopsis from which the epithet is derived. e diagnostic
characters include a thin poorly delimited cortex (10–15 μm), an uninterrupted algal
layer, a tall (120–140 μm), euamyloid (I+ blue), hymenium and narrow spores (4–5 ×
1.5 μm). Acarospora pyrenopsoides is a peculiar Acarospora not similar to any other species

New records of lichenized and lichenicolous fungi in Scandinavia 37
Figure 1. Acarospora insignis (O L-173397). Scale: 1 mm.
Figure 2. Acarospora pyrenopsoides (O L-163369). Scale: 1 mm.

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
38
in the genus. Magnusson (1926) considered its position in the genus to be isolated
because of its unusual characters but speculated that it may be closest to A. nitrophila.
e specimen from Norway was sequenced and a preliminary analysis (not shown)
indicates that it belongs to a group with A. peliscypha, A. molybdina and A. wahlenbergii
within Acarospora in a restricted sense (see Westberg et al. 2015). ere is however, as far
as we can see, no particular character of its morphology that indicates this relationship.
Acarospora pyrenopsoides appear to prefer humid localities. e locality in Norway
reported here, lies near a waterfall where the species was found on sloping rocks. It is
the only European site of the North American Rhizocarpon bolanderi (Tuck.) Herre,
and was found during an inventory of that species. e two Swedish localities are lo-
cated on or near lakeshores in the province of Värmland in central Sweden.
Specimens examined. NORWAY. Aust-Agder: Valle, Hallandsfossen. 28
July 2010, Timdal 11308 (O L-163369); SWEDEN. Värmland: Stora Kil par., W
steep towards the lake Fryken, S of Prästhamna. 15 May 1960, Sundell 1915a (UPS
L-126715); Trankil par., Guppviksön. 22 Aug. 1976, Sundell 11217 (UPS L-515137).
Acarospora versicolor Bagl. & Carestia, 1863
MycoBank: MB 375773
Acarospora versicolor Bagl. & Carestia. Comm. Soc. Critt. Ital. 1: 440. 1863.
Type. ITALY. Piemonte. F. Baglietto s.n. (the location of the type is unknown accord-
ing to Knudsen et al. in press).
Distribution. New to Norway. Acarospora versicolor is widespread on both sili-
ceous and calcareous rocks in Europe and western Asia and is in the Nordic countries
known from one locality in Finland and one unconrmed report from Denmark (Al-
strup et al. 1990, Knudsen et al. in press).
Acarospora versicolor belongs to the morphological group of brown Acarospora species
lacking secondary metabolites. ere are many names and many taxonomical problems
in this group but A. versicolor was recently discussed and described in detail by Knudsen
et al. (in press). Acarospora versicolor is identied by its white pruinose thallus and the
negative C-reaction of the cortex. e thallus in A. versicolor is areolate and can become
somewhat squamulose when well developed. It is mostly recognized by the presence of
a white pruina on at least parts of the thallus (Fig. 3), but the pruina is sometimes lack-
ing completely. e apothecia usually have a distinct margin and both the disc and the
margin are typically darker than the thallus, sometimes almost black. It may then remind
of a small form of A. badiofusca but the apothecia are not sessile as the mostly are in that
species and there are several other character separating these species.
e Norwegian specimens reported here all grow calcareous rocks in sun-exposed
habitats in the southern parts of the country. It is as far as we know the only one in
this group of species in Scandinavia growing on calcareous rocks. However, elsewhere
in Europe it also grows on non-calcareous rocks (Knudsen et al. in press).

New records of lichenized and lichenicolous fungi in Scandinavia 39
Specimens examined. NORWAY. Buskerud: Hole, west side of the island
Storøya, 60,04685°N, 10,2376°E. 8 June 2008, Westberg 08-092 (S F268460);
Oppland: Dovre, Hjelle, 62°03.21'N, 9°08.40'E, alt. 650 m. 10 Aug. 2010, Timdal
11757 (O L-163814, led under Lecidea degeliana); Vågå, c. 300 m NE of the farm
Viste, 61,86671°N, 9,02391°E. 13 June 2008, Westberg 08-198, 08-203, 08-205 (S
F268461, F268462, F268463).
Calvitimela perlata (Haugan & Timdal) R. Sant, 2004
MycoBank: MB 478956
Calvitimela perlata (Haugan & Timdal) R. Sant. Lichen-forming and lichenicolous fungi
of Fennoscandia: 73. 2004.
Basionym. Tephromela perlata Haugan & Timdal, Graphis Scripta 6(1): 22 (1994).
Holotype. NORWAY. Sør-Trøndelag: Oppdal municipality, Drivdalen, by the
rapids in the lower part of the river Kaldvella, 62°17'N, 9°35'E, alt. 940–980 m, ex-
posed rock face in the subalpine region. 23 July 1993, E. Timdal 7535 (O L-125!),
TLC: rangiformic acid, norrangiformic acid, unknown substances.
Distribution. New to Sweden. e species was previously known only from Norway
and Greenland (Haugan and Timdal 1994). It resembles the more common C. aglaea
(Sommerf.) Hafellner, but diers in forming white areolae (Fig. 4), slightly longer ascospores
Figure 3. Acarospora versicolor (S F268463). Scale: 1 mm.

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
40
(11.5–17 × 5–7.5 μm; Haugan and Timdal 1994), and in the presence of rangiformic and
norrangiformic acids in the thallus. In C. aglaea, the areolae are usually pale yellow, the as-
cospores shorter (9–13.5 × 5–7.5 μm), and the thallus contains atranorin, bourgeanic acid,
usnic acid (rarely lacking), and sometimes stictic acid (Haugan and Timdal 1994).
In Norway, the species grows on sun-exposed, more or less sloping rock surfaces,
often where water is trickling or near rivers or waterfalls in the subalpine and alpine
regions. e Swedish locality is a boulder in the alpine region.
Specimen examined. SWEDEN. Torne Lappmark: Låktatjåkka, 68°24.87'N,
18°19.07'E, alt. 640 m, steep face of boulder in the low alpine region. 8 July 2014,
Timdal 13464-1 (O L-195661), TLC: rangiformic acid (major). norrangiformic acid
(minor), unknown substances (minor).
Cladonia albonigra Brodo & Ahti, 1996
MycoBank: MB 415621
Cladonia albonigra Brodo & Ahti. Canad. J. Bot. 74: 1152. 1996
Holotype. CANADA. British Columbia: Queen Charlotte Islands, Graham Island, 2
mi. SE of Port Clements. 1971, Brodo 18104 & Wong (CANL).
Distribution. Conrmed for Sweden. e species is reported from the province
Torne Lappmark in northernmost Sweden by Ahti and Stenroos (2013), but no speci-
Figure 4. Calvitimela perlata (O L-195661). Scale: 5 mm.

New records of lichenized and lichenicolous fungi in Scandinavia 41
men was cited and there are no specimens from Sweden in S or UPS. We here conrm
its presence in Sweden and that province. e specimen (Fig. 5) was found on the
same rock as Calvitimela perlata referred to above.
Specimen examined. SWEDEN, Torne Lappmark: Låktatjåkka, 68°24.87'N,
18°19.07'E, alt. 640 m, steep face of boulder in the low alpine region. 8 July 2014,
Timdal 13464-2 (O L-195662), TLC: fumarprotocetraric acid (major).
Lecidea degeliana Hertel, 1970
MycoBank: MB 342410
Lecidea degeliana Hertel. Herzogia 2: 41. 1970.
Holotype. NORWAY, Troms: [Harstad, Hinnøy], Sandtorg Nordvik, ad saxum mica-
cei-schistosum aeneum fuscinigrum tinctum. 14 July 1953, G. Degelius (UPS L-108141!).
Distribution. New to Sweden. Lecidea degeliana was described by Hertel (1970)
from material collected by Degelius in northern Norway. Degelius himself (1982) re-
ported a second nd from the island of Vega in northern Norway but no further locali-
ties was mentioned by Hertel (1995) and we have not found any other literature reports
of this species although collections have slowly been accumulating in the herbarium in
Oslo. During an excursion to Sør-Trøndelag in Norway 2012, Swedish lichenologists
had the opportunity to study the species in the eld, and it was thereafter discovered in
Figure 5. Cladonia albonigra (O L-195662). Scale: 5 mm.

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
42
the Abisko area in northern Sweden in 2013 and on a second locality a few kilometers
from the rst in 2014. e species is so far only known from Scandinavia.
is small species may easily be overlooked or mistaken for a poorly developed L.
fuscoatra. It is, however, a quite distinct species once discovered. e species is initially
developing as a parasite on Acarospora spp. (Fig. 6). e thallus of L. degeliana is areolate,
up to 3 mm wide, and frequently fertile with black, rounded or angular apothecia that are
immersed in, or somewhat raised from the areoles; up to 0.2–0.6(–0.7) mm diam. In the
microscope the apothecia have a greenish or bluish green hymenium, a black or greenish
black epihymenium, and a dark brown hypothecium. e excipulum has a yellowish in-
ner part, and the outer part is dark grey. e ascospores are ellipsoid–subglobose, 5.5–7 ×
4–5 μm in the studied samples. e thallus contains the gyrophoric acid syndrome (Her-
tel 1995); thallus cortex in section K–, C+ red, excipulum in section K–, C– or C+ red.
Figure 6. Lecidea degeliana. a With the host, Acarospora sp., still visible (S F265204) b Independent
thallus (S F265201). Scale: 1 mm.

New records of lichenized and lichenicolous fungi in Scandinavia 43
Hertel (1968) described Lecidea grummannii from Kärnten, Austria, another spe-
cies growing parasitically on small brown Acarospora sp. at species is as far as we
know only known from the type. According to Hertel (1995), L. grummannii diers
from L. degeliana in having broader spores (5.2–5.5 vs 3.5–5.5 μm in L. degeliana), a
paler hypothecium, (pale brown vs dark to blackish brown); a blue-green epithecium
(vs olive- or greyish green in L. degeliana) and a paler thallus (cream coloured–whitish
to beige vs beige to pale brown). In our experience of L. degeliana these characters are
not so clear-cut and only the dark brown hypothecium seem to be a consistently dif-
ferent character compared to the description of L. grummannii.
Lecidea degeliana has mostly been found on calcareous rocks in exposed, subal-
pine habitats but also on siliceous and iron-containing rocks. ere seem to be several
dierent species of Acarospora involved as hosts but they are often sterile and mostly
unidentied by us. ey all belong to the small brown species of Acarospora s. str., and
are in one case identied as A. versicolor through sequencing of the ITS and in another
case tentatively identied morphologically as A. rugulosa.
Specimens examined. NORWAY. Hedmark: Ringsaker, the islet Holmen S of Hel-
gøya in lake Mjøsa, 60°43'N, 11°01'E, alt. 125–130 m. 11 Apr. 1982, Timdal 3230 (O
L-37925), TLC: gyrophoric acid; Ringsaker, Helgøya, the islet Holmen in lake Mjøsa,
60°43'N, 11°01'E, alt. 125 m. 27 March 1994, Haugan 3699 (O L-37924); Nordland:
Vega, Vega Is., Holand, Exposed blasted rock (clay schist) at the road near Kirköy. 4
July 1979, Degelius V-2406 (UPS L-143256); Oppland: Dovre, Verkensætri, rock out-
crop by the houses, 62°03.8 ‘N, 9°32.7 ‘E, alt. 1010-1020 m. Calciferous schist. 3 July
1995, Haugan & Timdal 8035 (O L-15096); Dovre, Hjelle, 62°03.21'N, 9°08.40'E,
alt. 650 m. 10 Aug. 2010, Timdal 11757 (O L-163814); Lom, Runningsgrende, Klypa.
Alt. 720-850 m. 61.7158°N, 8.2342°E. 28 June 2013, Westberg (S F265203), Lun-
ner, Råsted, 60°18'N, 10°37'E, alt. 330 m. S-facing, steep rock face. 19 June 1983,
Timdal 3844 (O L-37926), TLC: gyrophoric acid; Vågå, hill E of Svarthåmårbekken,
61°52.10'N, 8°59.21'E, alt. 690 m. 30 June 2013, Bendiksby, Haugan & Timdal 12978
(O L-184391); Westberg (S F265204). SWEDEN. Torne Lappmark: Jukkasjärvi par.,
Björkliden, N side of the river Rakkasjohka, just N of the trail from Björkliden Fjällby
to Nuolja, alt. 495 m. 68.4053°N, 18.6698°E. 6 Aug. 2013, Westberg (S F265201);
Björkliden, just W of the road E10 at its crossing over the river Rakkasjohka. 68.4029°N,
18.6944°E. 6 July 2014, Westberg VAR152 (S F265198).
Nephroma helveticum Ach, 1810
MycoBank: MB 357157
Nephroma helveticum Ach. Lich. Univ.: 523. 1810.
Lectotype. ‘In montibus Helvetiae, Schleicher’ (H-ACH 1470B, James & White 1987).
Distribution. New to Sweden. Nephroma helveticum is a cosmopolitan species
complex with a wide ecological amplitude and a large morphological and chemical

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
44
Figure 7. Nephroma helveticum (O L-196491) Scale: 5 mm.
variation. In Europe, however, the species is very rare and appears to have declined
considerably (Klepsland 2103, James and White 1987). ere are very few recent nds,
among them a locality in Oppland in Norway, which is the rst nd in that country
(Klepsland 2013). In the Nordic countries it is elsewhere known from a few old locali-
ties in Finland (Vitikainen 2007). Here we report it from one locality found in 2009
in Västerbotten and in addition we have found an old specimen in the herbarium in
Stockholm from Västerbotten collected by Sten Ahlner in 1945 but never reported
in the literature. e two localities are located 2.5 kilometers apart. We also report a
second nd of this species in Norway. e species is recognized by its tomentose lower
surface without papillae (Fig. 7) and its chemistry of secondary metabolites, including
the terpenoid T4 (James and White 1987). e most similar species N. resupinatum also
has a tomentose lower surface but has white papillae between the tomentum and lacks
secondary metabolites. One of the recently collected specimen from Västerbotten has
been barcoded and the ITS sequence conrms the identity as N. helveticum.
On Ahlner´s locality, Borstaberget, the bedrock consists of greenstone (porphyrite)
and the mountain has long southwest facing slopes with steep clis. Ahlner collected,
together with N. helveticum, also the rare lichen Heterodermia speciosa. On the moun-
tain Mitti-Skansberget N. helveticum was found in 2009 in two places 200 meters
apart. It was found growing on conglomerate clis in the southwest facing precipices.
Other species that were found on the clis were Peltigera rufescens, Lobaria scrobiculata,
L. pulmonaria, Fuscopannaria leucophaea and Biatora vernalis.

New records of lichenized and lichenicolous fungi in Scandinavia 45
Specimens examined. NORWAY. Buskerud: Nes municipality, Gardnosberget,
MGRS: 32V NN 0230, 2309, alt. 300 m, east-facing, steep slope below high moun-
tain wall, open spruce forest over rock eld, on boulder. 10 September 2013, Hofton
13176 (O L-196491), TLC: series of terpenoids, including T4 (major). SWEDEN.
Västerbotten: Jörn par., Borstaberget. 24 June 1945, Ahlner (S L-54838); Norsjö par.,
Mitti-Skansberget SE precipice, SE of Klövertjärnen, 9.7 km NW of Petiknäs church.
28 Sept. 2009, Jonsson 4200 (UPS L-193714).
Peltula placodizans (Zahlbr.) Wetmore, 1970
MycoBank: MB 343163
Peltula placodizans (Zahlbr.) Wetmore, Ann. Missouri Bot. Gard. 57: 179. 1970.
Basionym. Heppia placodizans Zahlbr., Bull. Torrey Bot. Club 35: 299 (1908).
Holotype. U.S.A. Arizona, Tucson, Tumamoc Hill. 1908, Blumer (W, holotype,
not seen).
Distribution. New to the Nordic countries. e species is widely distributed in
arid areas of both the Northern and Southern Hemispheres (Egea 1989). e nearest
locality is found in Northern Italy (Vinschgau in South Tyrol; Buschardt 1979), and
the species ts in the element of continental lichens with a widely disjunct distribution
from the Alps to the upper valleys of southeast Norway. Other species with a similar
disjunction include Buellia elegans Poelt, Gyalolechia desertorum (Tomin) Søchting et
al., Psora vallesiaca (Schaerer) Timdal, Toninia sculpturata (H. Magn.) Timdal, T. tau-
rica (Szatala) Oxner, T. tristis (. Fr.) . Fr., and Xanthocarpia tominii (Savicz) Frö-
dén et al. e species is recognized by the crustose to subsquamulose, dark olivaceous
brown thallus composed of areolae, which are up to 1 mm diam. and with marginal,
black, granular soralia. e Norwegian material is sterile. e other Peltula species
in the Nordic countries, P. euploca (Ach.) Poelt, diers in forming much larger, pel-
tate squamules, up to 5 mm diam. e Norwegian specimen of Peltula placodizans is
identied with some uncertainty, as the material for comparison (17 specimens from
Europe, Africa, Australia, and North and South America, borrowed from GZU), was
often more olivaceous brown than the dark brown Norwegian material (Fig. 8). Wet-
more (1970) indicates that the species is probably a complex of several taxa.
In Norway, the species was found on a vertical wall of calcareous rock in a steep,
west-facing hillside. e site has apparently previously been an open or sparsely wood-
ed pasture, but is now in the process of being transformed into spruce forest. Other
remarkable lichens collected at the site include Metamelanea caesiella (.Fr.) Henssen,
Physcia dimidiata (Arnold) Nyl., allinocarpon nigritellum (Lettau) P.M.Jørg., and
Toninia alutacea (Anzi) Jatta.
Specimen examined. NORWAY. Oppland: Sør-Fron municipality, Harpefoss,
along the trail W of farm Tåkåstad towards Mt. Lundin, 61°34.95'N, 9°52.55'E, alt.
490 m. 1 Oct. 2007, Timdal 11054 (O L-158470), TLC: no lichen substances.

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
46
Figure 8. Peltula placodizans (O L-158470). Scale: 1 mm.
Phacographa protoparmeliae Hafellner, 2009
MycoBank: MB 513175
Phacographa protoparmeliae Hafellner. Bibl. Lich. 100: 106. 2009.
Holotype. AUSTRIA, Kärnten: Hohe Tauern, Kreuzeck-Gruppe, Kalkschieferwände
in den SE-Hängen der Sensenspitze N der Turgger Alm, c. 200 m. 17 July 1978,
Hafellner 603 (GZU).
Distribution. New to the Nordic countries. Phacographa was described by Hafell-
ner 2009 who included three species in the genus. It belongs in the newly described
family Lecanographaceae within the Arthoniales (Frisch et al. 2014). Phacographa pro-
toparmeliae (Fig. 9) was originally reported from Austria and Spain (Hafellner 2009),
and a specimen was later discovered in the Murmansk region in arctic Russia (Fryday
2011). Here we report it for the rst time from the Nordic countries from two locali-
ties in Norway. Both are at old copper mines in subalpine habitats in the mining dis-
trict of Røros in central Norway.
Phacographa protoparmeliae (Fig. 9) is a lichenicolous fungus growing on the
common and widespread saxicolous lichen Protoparmelia badia. e apothecia of
Phacographa have a black margin around a dark brown disc; they are up to 1.2 mm
wide in the Norwegian material and form clustered groups of 2–6 apothecia. Micro-
scopically it is characterized by its 3-septate, spores, 22–25 × 6–7(–8) μm that are

New records of lichenized and lichenicolous fungi in Scandinavia 47
colourless but have a perispore sheath that becomes brown with age. More detailed
descriptions can be found in Hafellner (2009) and Fryday (2011).
Specimens examined. NORWAY. Sør-Trøndelag: Røros, Storwartz, at the site
of the old copper-mine, 62°37.63'N, 11°31.19'E. 15 June 2012, Westberg 12-030 (S
F265199); Røros, Klasberget, old copper mine NE of farm Svensvollen, 62°39.25'N,
11°34.06'E. 16 June 2012, Westberg 12-052 (S F265200).
Rhizocarpon pycnocarpoides Eitner, Jahresbericht der Schles, 1911
MycoBank: MB 404075
Rhizocarpon pycnocarpoides Eitner, Jahresbericht der Schles. Gesellschaft für vaterl.
Cultur 88, 2: 46. 1911.
Holotype. CZECH REPUBLIC, Krkonoše Mts, “an den alten Bergwerken im Riesen-
grunde“ (not seen).
Distribution. New to the Nordic countries. e species is apparently previously
known only from the type locality in the Krkonoše Mts, where it was collected by Eit-
ner about 1910 and by Kuťák in 1927. We have not seen the type material, but rather
one duplicate of the material distributed by Kuťák in his exsiccate (Flechtensamml.
Böhmen No. 520, O-L-184255).
Figure 9. Phacographa protoparmeliae (S F265200). Scale: 1 mm.

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
48
e species grows on rocks with a high content of iron and the thallus is rust
coloured (Fig. 10). It resembles R. oederi (Weber) Körb., which may be found in the
same localities, but diers primarily by forming larger ascospores (mainly muriform,
24–30 × 10–15 μm in R. pycnocarpoides vs. mainly 3-septate, 12–18 × 3–7 μm in R.
oederi). Furthermore, the apothecia are more sessile, more regularly rounded, and with
a thicker margin in R. pycnocarpoides, not in level with the areolae and angular to exu-
ouse as in R. oederi. An unpublished phylogenetic analysis of the genus Rhizocarpon
(Bendiksby et al. in prep.), which includes four Norwegian specimens of R. pycnocar-
poides, places the two species in sister position and well separated in the ITS marker.
Rhizocarpon pycnocarpoides has been found at ve localities in Norway and one
in Sweden. All localities are rich in rust stained rock and most sites are in or near old
copper or zinc mines.
Specimens examined. CZECH REPUBLIC. Krkonose. 1927, V. Kuťák,
Kuťák, Flechtensamml. Böhmen No. 521 (O L-184255). NORWAY. Buskerud:
Ringerike municipality, the old mine Ertlien, 60°04.14'N, 10°02.89'E, alt. 160 m.
18 May 2013, Rui & Timdal 12854 (O L-184267), TLC: No lichen substances;
Nord-Trøndelag: Namsskogan municipality, S bank of river Namsen just W of
Storholmen, 64°55.60'N, 13°08.72'E, alt. 200 m. 8 Aug. 2012, Haugan 11128 (O
L-183810), TLC: no lichen substances; Nordland: Fauske municipality, Sulitjel-
ma, Furuhaugen, site of old copper mine (in operation 1896-1921), 67°09.02'N,
15°57.92'E, alt. 260 m. 11 Aug. 2012, Haugan 11125 (O L-183808), TLC: no
lichen substances; Oppland: Lunner municipality, the old zinc mine Nysetergru-
Figure 10. Rhizocarpon pycnocarpoides (O L-184267). Scale: 1 mm.

New records of lichenized and lichenicolous fungi in Scandinavia 49
vene (in use 1889-1931), 60°15.69'N, 10°41.62'E, alt. 520 m. 16 July 2012, Rui
& Timdal 12665 (O L-179903); Sør-Trøndelag: Røros municipality, by river Orva
upstreams from the bridge at Litlstuvollen, 62°38.78'N, 11°21.22'E, alt. 700 m. 16
June 2012, Bendiksby et al. 12630 (O L-179554, L-179560), TLC: no lichen sub-
stances; Røros, Klasberget, old copper min NE of the farm Svensvollen. 62°39.25'N,
11°34.06'E. 16 June 2012, Westberg 12-061 (S F265205), TLC: no lichen sub-
stances. SWEDEN. Torne lappmark: Karesuando par., Pältsan (Bealccan), N-facing
slope of the middle peak (1444 m). Alt. 1240 m. 69,0106°N, 20,2366°E. 3 Aug.
2011, Westberg P122 (S F265206).
Sarcogyne algoviae H.Magn, 1935
MycoBank: MB 411790
Sarcogyne algoviae H.Magn. Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 9(5.1): 78. 1935.
Holotype. [GERMANY, Bayern] Obere Seealpe in der Allgäuer Alpen bei Oberstdorf,
c. 5000’. 1860, H. Rehm (S L2741!).
Distribution. New to Sweden. is is a little known species rarely reported in the
literature. Sarcogyne algoviae belongs to the morphological group in Sarcogyne with a
strongly carbonized margin (Westberg et al. 2015). e apothecia vary considerably
in size (0.3–1.8 mm) and has a raised, often glossy margin that is incised at short and
fairly regular intervals (Fig. 11a, b) forming 5–15 segments. e disc is brown to dark
reddish brown and often has one or two small carbonized accretions on the surface.
Further characters include a euamyloid hymenium, 65–105 μm tall and narrow, 1–1.5
μm wide paraphyses, ellipsoid to narrowly ellipsoid spores, 2.5–4 × 1.5 μm and a
colorless hypothecium (Fig. 11c). It can primarily be confused with S. clavus and S.
hypophaea (syn. S. privigna, see Knudsen et al. 2013) but is apparently not closely re-
lated to either of these species (see phylogeny in Westberg et al. 2015). Sarcogyne clavus
in its current sense (e.g. Magnusson 1935) always grows on siliceous rocks, has larger
apothecia (up to 6 mm wide) with a margin that is irregularily cracked and crenulate
and has a yellowish, to pale brownish hypothecium. Sarcogyne hypophaea grows on
both siliceous and calcareous rocks and has generally smaller apothecia 0.3–0.7(1.3)
mm, with a dierent appearance. e margin can be without or with rather indistinct
incisions or with fewer (3–8) incisions at irregular intervals.
We have found Sarcogyne algoviae on several localities in Scandinavia, two in the
continental parts of southern Norway and two in the Abisko area in northernmost Swe-
den. In addition we have found a few specimens from the Swedish mountains under
the name S. clavus in the herbarium UPS. In all localities it grows on soft, calcareous
schist in alpine–subalpine habitats. Sarcogyne algoviae is already included in the check-
list for Fennoscandia, originally based on a specimen from Finnmark in northernmost
Norway (Santesson et al. 2004). at specimen, however, has broadly ellipsoid spores
and stout paraphyses, c. 2.5 μm wide, and belongs to Polysporina urceolata (specimen in

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
50
Figure 11. Sarcogyne algoviae. a Apothecia (S F122537) b Apothecia (S L2741, holotype) c Section of
an apothecium showing a strongly carbonized margin and a colourless hypothecium (S F122537). Scale:
1 mm (a–b); 100 μm (c).
O, L-38325). Other literature or database records of S. algoviae from Norway must be
considered unreliable but here we also conrm its presence in Norway.
Specimens examined. NORWAY. Oppland: Dovre, Jønndalen, Mt Nonshaugen,
S precipice of the mountain, NE of farm Jønndalen, alt. 700–800 m. 12 June 2008,
Westberg 08-276 (S F122564) Lom, Bøvertun, just W of the lake Bøvertunvatnet.
Alt. 954 m. 12 June 2008, Westberg 08-165, 08-168, 08-169 (S F122535, F122537);
Lom, Runningsgrende, Klypa. Alt. 700–760 m. 61.7237°N, 8.0262°E. 28 June 2013,
Westberg (S F265202); Nordland: Vega Island, farm Dalen. 22 June 1974, Degelius
V-683 (UPS L-516318); Troms: Insula Rollöen. [undated], Norman (UPS L-680378).
SWEDEN. Härjedalen: Tännäs par., Mt Stora Mittåkläppen, the southern slope. 15
Aug. 1962, Santesson 14987b (UPS L-516445); Jämtland: Åre par., Storlien. 24 July
1950, Magnusson 22128a (UPS L-515993); Lule lappmark: Gällivare par., Vastenjau-
re. 4 Aug. 1965, Gilenstam 1186a (UPS L-103743); Torne lappmark: Jukkasjärvi par.,
Björkliden, N side of the river Rakkasjohka, just N of the trail from Björkliden Fjällby
to Nuolja, alt. 495 m. 68.4053°N, 18.6698°E. 6 Aug. 2013, Westberg (S F265208);

New records of lichenized and lichenicolous fungi in Scandinavia 51
Latnjajaure eld station, by the lake Latnjajaure c. 15 km W of Abisko. 68°20'N,
18°30'E, alt. 980–1000 m. 3 Aug. 1998, Westberg 2572 (LD).
Sarcogyne hypophaeoides Vain. ex H.Magn, 1935
MycoBank: MB 411805
Sarcogyne hypophaeoides Vain. ex H.Magn. Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 9(5.1):
84. 1935.
Lectotype. FINLAND. Tavastia australis, Luhanka, Keihäsniemi. 1873 (TUR-Vainio
25683, designated here, TUR-Vainio 25682 isotype!).
Distribution. New to Norway, Sweden and Austria. Sarcogyne hypophaeoides has
until now been known from the type material in central Finland and from one collec-
tion in Germany (Magnusson 1935). It is a characteristic but overlooked lichen that
appears to be widespread in Scandinavia. e apothecia are 0.5–1.2 mm wide, with
a reddish brown to almost black disc sometimes with a central carbonized accretion.
e margin is typically nely striated (Fig. 12a, b) and is softer and more leather-like
in texture and not as strongly carbonized and brittle as the margin in e.g., S. clavus.
e degree of carbonization varies however and in sections the central parts of the ex-
ciple may be rather weakly coloured (Fig. 12c). e hymenium is 70–90 μm tall and
has a euamyloid reaction (I+ blue) and the spores are narrow and almost bacilliform,
3.5–5.5 × 1.0 μm. Sarcogyne hypopaheoides can mainly be confused with S. clavus or S.
hypophaea but has a dark, brownish black to black hypothecium (Fig. 12c), which is a
characteristic and diagnostic feature of this species. Sarcogyne clavus has a very uneven,
cracked, crenulate margin and a yellowish to pale brown hypothecium. Sarcogyne hy-
pophaea on the other hand, has a smooth margin, which usually appears segmented
due to 2-3 deep incisions in the margin and a colorless hypothecium.
Sarcogyne hypophaeoides grows exclusively on siliceous rocks. We have found it on
exposed, horizontal or sloping seashore rocks on the west coasts of Sweden and Nor-
way, on lakeshores and on semi-exposed, vertical rocks or boulders in open coniferous
forests. It is often growing with scattered apothecia on smooth rock surfaces or along
cracks in the rock in a similar way to S. clavus and the two species have been found
growing together on at least two localities. Diculties to collect it and a supercial
similarity to S. clavus are possibly reasons why this species has been overlooked. It ap-
pears to be fairly common in humid habitats in the boreal region of Fennoscandia but
its distribution is incompletely known.
Specimens examined. AUSTRIA. Steiermark: Steirisches Randgebirge, Fishcba-
cher Alpen, im Feistriztal, ca 2 km E von Rettenegg. 14 Nov. 1998, Kocourkova &
Hafellner 46366 (GZU); NORWAY. Rogaland: Rennesøy, Fjøløy, ca 200 m. NE of
the lighthouse. 11 June 2008, Westberg 08-139 (S F123697). SWEDEN. Bohuslän:
Ljung par., Skarsjön. 31 Aug. 1916, Magnusson 17466 (UPS L-175686); Naverstad
par., Tyfteäll. 22 July 1917, Magnusson (UPS L-516234); Brastad par., Stora Bornö,

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
52
Figure 12. Sarcogyne hypophaeoides. a Apothecia (S F123697) b Apothecia (Kocourkova & Hafellner
46366 (GZU)) c Section of an apothecium showing a carbonized margin and a brownish black hypothecium
(SF265197). Scale: 1 mm (a–b); 100 μm (c).
just S of Källviken on the E side of the island, c. 750 m SE of the research station.
58.3750°N, 11.5902°E. 4 Oct. 2013, Westberg (S F265197); Dalarna: Leksand par.,
Draggberget. Hermansson 14488 (UPS L-564677); Dalsland: Skållerud par., c. 5.5
km N of Skållerud church, SW slope of Skalåsen. 58.8268°N, 12.4435°E. 5 June
2008, Westberg 08-078 (S F120302); Gästrikland: Hille par., Iggön. 13 Aug. 1946,
Ahlner (S F90566); Lycksele lappmark: Tärna par., Ume älv, Över-Umans sydvästli-
gaste vik. 18 Aug. 1960, Du Rietz 927 (UPS L-115656); Pite lappmark: Arvidsjaur
par., Pite älv, Trollforsen, nedre delen, N-sidan uppströms bron. 24 Aug. 1962, Du
Rietz 486b (UPS L-113499); Uppland: Djurö par., Runmarö, S of Kasviken, along a
forest-trail. 59,2612°N, 18,7666°E. 11 May 2009, Westberg, Millanes & Wedin 09-
308 (S F265196), Värmland: Gustav Adolf par., NW of Uddeholmshyttan. 2 Sept.
1981, Sundell 14905a (UPS L-516265); Västmanland: Grythyttan par., c. 1400 m S
of Loka Brunn, at the south tip of the lake S. Loken. 59.5926°N, 14.4844°E. 1 May
2008, Westberg 08-002 (S F119718).

New records of lichenized and lichenicolous fungi in Scandinavia 53
Tephromela grumosa (Pers.) Hafellner & Cl. Roux,1985
MycoBank: MB 103854
Tephromela grumosa (Pers.) Hafellner & Cl. Roux. Bulletin de la Société Botanique du
Centre-Ouest 7: 829. 1985.
Basionym. Lichen grumosus Pers., Ann. Bot. Usteri 14: 36. 1795. Nom. nov. Lichen
caerulescens Pers., Ann. Bot. Usteri 11: 15. 1794. Nom. illeg. (non Lichen caerulescens
Hagen 1782).
Type. Sine loc., „ad saxa arenaria (rubicunda), a Dom. Heyder primo observatus“
(Not seen).
Distribution. New to Norway. Tephromela grumosa has been expected to occur in
Norway, as it is known from a number of provinces in Sweden and Finland. It occurs in
West, Central, and North Europe and in Asia (Lambley and Purvis 2009). e species
is rather closely related to the type species of the genus, T. atra (Huds.) Hafellner &
Kalb, but it is always sorediate and rarely fertile (Fig. 13). In contrast to the genetically
very heterogenous T. atra s. lat., T. grumosa seem to be genetically rather consistent
compared to its morphology (Muggia et al. 2008, 2014). e species is furthermore
separated by their chemistries, i.e., by the presence of lichesterinic acid in T. grumosa.
e chemistry of the Norwegian specimen was conrmed by TLC. According to Mug-
gia et al (2008) T. grumosa is a cooltemperate species found on steeply inclined, acidic
siliceous rocks. In the locality in Norway, luxuriant, partly fertile specimens grew on
Figure 13. Tephromela grumosa (O L-190787). Scale: 5 mm.

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
54
steep faces of a large boulder in a stabilized S-facing talus. e rock is schistose, and
probably somewhat calciferous. e locality is situated in one of the driest areas in
Scandinavia, with an annual precipitation of c. 350 mm. In other habitats, e.g. on hard,
granitic rocks in coastal heath on the Swedish west coast, specimens of this species are
much thinner and less prominent.
Specimens examined. NORWAY. Oppland: Lom municipality, Runningsgrende,
Klypa. 61°43.41'N, 8°15.67'E, alt. 730 m. 28 June 2013, Bendiksby et al. 12357 (O
L-190787); TLC: atranorin, lichesterinic acid. SWEDEN. Bohuslän: Sotenäs munici-
pality, Ramsviklandet nature reserve, W of Haby. 58°24.50'N, 11°14.55'E, alt. 20 m.
25 Mar. 2012, Haugan 11501 (O L-184061).
Tremella lobariacearum Diederich & M. S. Christ, 1996
MycoBank: MB 415289
Tremella lobariacearum Diederich & M. S. Christ. Bibl. Lichenol. 61: 103. 1996.
Type. PORTUGAL. Madeira: Rabaçal, on Lobaria pulmonaria. 8 Apr. 1992, P. Diederich
4935 (LG – holotype; herb. Diederich – isotype; S F102418 - isotype!).
Distribution. New to the Nordic countries. Tremella lobariacearum was described
by Diederich (1996) and is currently known from Africa (Ile de la Réunion), Asia (Ja-
pan, Philippines, and Russia), Europe (British Isles, France, and Portugal), Macaron-
esia (Canary Islands and Madeira), Oceania (Papua New Guinea) and South America
(Bolivia, Colombia, Ecuador, and Peru), growing on Lobaria and Pseudocyphellaria
species (Diederich 1996, 2003, van den Boom and Etayo 2000, Etayo 2002, 2010,
Coppins et al. 2012, Flakus and Kukwa 2012). e species induces the formation of
pale-brown to dark-brown galls on the isidia or soredia of the host (Fig. 14), or oc-
casionally also directly on the thallus. e basidia are 2-celled. Two asexual morphs,
consisting of lunate conidia and asteroconidia, are often found within the galls. Tre-
mella lobariacearum is here reported from Norway and from the Nordic Countries
for the rst time, growing on Lobaria pulmonaria. e species is associated to habitats
with high humidity and oceanic inuence. e only specimen reported was collected,
together with other non-infected L. pulmonaria thalli in an area with mixed and rich
populations of Lobaria, in a Fraxinus-dominated open broad-leaved deciduous stand.
Tremella includes mainly mycoparasitic taxa that grow on a wide range of fungal
hosts, including lichenized hosts. However, mycologists and lichenologists in general
did not look much at the lichenicolous species until the rst comprehensive study
by Diederich (1996). Since then, several new species have been described (Diederich
2003, Sérusiaux et al. 2003, Diederich 2007, Zamora et al. 2011, Diederich et al.
2014, Millanes et al. 2012, 2014), and an increased interest has resulted in numerous
new records, especially in the Nordic countries (Holien and Tønsberg 1994, Alstrup
et al. 2004, 2008, Pippola and Kotiranta 2008, Westberg et al. 2008, Svensson and
Westberg 2010, Knutsson and Johansson 2011, Millanes et al. 2014, ell et al. 2014,

New records of lichenized and lichenicolous fungi in Scandinavia 55
Westberg and or 2014). Nevertheless, both the actual diversity and the distribution
area of most species are still largely unknown.
Specimen examined. NORWAY. Hordaland: Tysnes municipality, Støle,
59°59.14'N, 05°29.84'E, alt. 60 m. 6 Apr. 2008, Asplund & Larsson (S F263902).
Tremella wirthii Diederich, 1996
MycoBank: MB 415310
Tremella wirthii Diederich. Bibl. Lichenol. 61: 164. 1996.
Holotype. GERMANY. Bayern: Neu-Ulm, Holzheim, Obstgarten WSW Steinheim,
MTB 7626/2. 6 Feb 1991, V. Wirth 21713 (STU)
Distribution. New to the Nordic countries. Tremella wirthii was described by Died-
erich (1996) based on material from four localities in Germany, growing on an unidenti-
ed sterile corticolous lichen. e host was later described as Protoparmelia hypotremella
(Aptroot et al. 1997), a species similar to P. oleagina, from which it diers by its paler
colour and its isidia-like granules. Both species are among the few corticolous Protoparme-
lia species, and grow on old wooden fences and buildings, and on deciduous and conifer-
ous trees (Aptroot et al. 1997, 2001, 2004, Clerc 2004, Diederich and Sérusiaux 2000,
Figure 14. Tremella lobariacearum. Galls induced by Tremella lobariacearum on Lobaria pulmonaria
(SF263902). Scale: 1 mm.

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
56
Figure 15. Tremella wirthii. a Basidiomata on the thallus of Protoparmelia oleagina (S F262967) bBasidium
with one longitudinal septum (S F262963) c Basidium with one oblique septum (S F262963) dBasidium with
one transeverse septum (S F262963) e Catenulate conidia (S F262963). Scale: 0.5 mm (a); 10 mm (b–e).

New records of lichenized and lichenicolous fungi in Scandinavia 57
Scholz2000, Hafellner and Türk 2001, Palice et al. 2006, van den Boom et al. 2007,
Kubiak et al. 2010, Kukwa et al. 2012, Himelbrant et al. 2014). Aptroot et al. (1997)
reported the presence of Tremella wirthii on specimens of P. hypotremella from Austria and
the Netherlands. In the type locality of Protoparmelia hypotremella, in the Netherlands, T.
wirthii was found both on P. hypotremella and P. oleagina. e three Scandinavian speci-
mens grow on Protoparmelia oleagina. ey were collected on wood of an old hay drying
rack in the agricultural landscape of the Siljan Ring, an area with a calcareous bedrock, and
the only locality in Sweden were P. hypotremella has been reported. Here, however, Tre-
mella wirthii has not yet been found on that host. In the Nordic countries Protoparmelia
oleagina occurs in Norway and Sweden (Nordin et al 2015). Tremella wirthii forms dark
basidiomata, which are rarely bigger than 0.5 mm in diam., on the host thalli (Fig.15a).
Some micromorphological dierences have been found between Swedish and Central Eu-
ropean material. e specimens studied in the original description had 2–4 celled, longi-
tudinally septate basidia, and no asexual morph was reported (Diederich 1996). e three
specimens collected in Dalarna show typical basidia with 2 or 4 cells and longitudinal septa
(Fig. 15b), but also basidia with oblique (Fig. 15c) or transverse (Fig. 15d) septa are present
within the same basidioma. Moreover, in some cases there is an anamorph consisting of
catenulate conidia (individual conidia 3–6 × 3–6 μm; Fig. 15e). Despite these dierences,
we consider the material from Sweden and Central Europe to be conspecic. e basidium
morphology is extremely variable in many species of Tremella (Zamora et al. 2011, Mil-
lanes et al. 2012), and, when present, the occurrence of conidia is not constant in all speci-
mens of the same species. Since the original description was based on four specimens only,
the morphological variation within the taxon may well be greater than originally observed.
Tremella wirthii is probably overlooked in Sweden, and additional eld and herbarium
surveys could reveal its presence in places were Protoparmelia oleagina is abundant.
Specimens examined. SWEDEN. Dalarna: Orsa municipality 61°11.25'N,
14°52.43'E, alt. 255 m. 1 Oct. 2014, Jonsson FU5955 (S F262967); Dalarna: Orsa
municipality 61°11.32'N, 14°52.45'E, alt. 253 m. 1 Oct. 2014, Jonsson FU5956 (S
F262963); Mora municipality 60°55.67'N, 14°37.08'E, alt. 204 m. 2 Oct. 2014, Jon-
sson FU5957 (S F262952).
Acknowledgements
M. Westberg, A. Millanes and M. Wedin are supported by grants from e Swed-
ish Taxonomy Initiative (Svenska Artprojektet) administered by the Swedish Species
Information Centre (ArtDatabanken). M. Bendiksby, R. Haugan and E. Timdal were
supported by e Norwegian Taxonomy Initiative (Norske Artsprosjektet; proj no:
70184216) administered by the Norwegian Biodiversity Information Centre (Arts-
Databanken). DNA barcoding of R. pycnocarpoides and T. grumosa was funded by the
Norwegian Barcode of Life project (http://www.norbol.org/). We are grateful to the
curators of the herbaria mentioned for loan of material. We thank our reviewers, in
particular Paul Diederich, for valuable comments that improved the manuscript.

Martin Westberg et al. / MycoKeys 11: 33–61 (2015)
58
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