Zhurbenko, M. P. & Alstrup, V. 2004. Lichenicolous

Abstract

Four taxa are described as new –Polycoccum laursenii Zhurb., Pronectria tibellae Zhurb., Taeniolella strictae Alstrup and Sphaerellothecium araneosum var. cladoniae Alstrup & Zhurb. The combination Arthonia epicladonia (Nyl.) Alstrup &Zhurb. is proposed. Lecanora leptacinella, Niesslia cladoniicola and Roselliniella cladoniae are reported as new to Russia, Cercidospora cladoniicola, Epicladonia stenospora, Protothelenella santessonii and Taeniolella beschiana as new to Svalbard, Lichenosticta alcicornaria as new to Iceland, Epicladonia sandstedei as new to Ireland, Cercidospora punctillata as new to Jan Mayen, and Bachmanniomyces uncialicola, Lichenoconium pyxidatae and Taeniolella beschiana as new to USA. Cercidospora punctillata, Dactylospora deminuta, Protothelenella sphinctrinoidella and Rinodina egediana are reported on the genus Cladonia for the first time. A worldwide key to 77 fungi on Cladonia species is provided.

Full text

Lichenicolous fungi on Arctic Cladonia 477
Symb. Bot. Ups. 34:1
Lichenicolous fungi on Cladonia mainly from the Arctic
Mikhail P. Zhurbenko and Vagn Alstrup
Zhurbenko, M. P. & Alstrup, V. 2004. Lichenicolous fungi on Cladonia mainly from the
Arctic. – Acta Univ. Ups. Symb. Bot. Ups. 34:1, 477–499. Uppsala. ISBN 91-554-6025-9.
Four taxa are described as new – Polycoccum laursenii Zhurb., Pronectria tibellae Zhurb.,
Taeniolella strictae Alstrup and Sphaerellothecium araneosum var. cladoniae Alstrup &
Zhurb. The combination Arthonia epicladonia (Nyl.) Alstrup & Zhurb. is proposed. Lecanora
leptacinella, Niesslia cladoniicola and Roselliniella cladoniae are reported as new to Russia,
Cercidospora cladoniicola,Epicladonia stenospora,Protothelenella santessonii and Taenio-
lella beschiana as new to Svalbard, Lichenosticta alcicornaria as new to Iceland, Epicladonia
sandstedei as new to Ireland, Cercidospora punctillata as new to Jan Mayen, and Bachman-
niomyces uncialicola,Lichenoconium pyxidatae and Taeniolella beschiana as new to USA.
Cercidospora punctillata,Dactylospora deminuta,Protothelenella sphinctrinoidella and Ri-
nodina egediana are reported on the genus Cladonia for the first time. A worldwide key to 77
fungi on Cladonia species is provided.
Key words: cladoniicolous fungi, Polycoccum,Pronectria,Taeniolella,Sphaerellothecium,
key.
Mikhail P. Zhurbenko, Komarov Botanical Institute Russian Academy of Sciences, Professor
Popov 2, St. Petersburg, 197376, Russia. E-mail: zhurb@MZ3838.spb.edu
Vagn Alstrup, Botanical Museum, University of Copenhagen, Gothersgade 130, 1123 Copen-
hagen K, Denmark. E-mail: vagna@bot.ku.dk
Introduction
Cladonia is a large lichen genus containing ap-
proximately 450 species (Litterski & Ahti 2004).
So far about 85 species of lichenicolous fungi (in-
cluding several slightly lichenized ones) are
known to grow on this genus worldwide. Only
Peltigera supports more lichenicolous fungi – no
less than 130 species according to literature. How-
ever, cladoniicolous fungi have been specially
treated only in Hansen & Alstrup (1995), dealing
with fungi growing on Cladonia subgen. Cladina
in Greenland. Recent treatments of Arctic licheni-
colous fungi include Alstrup & Hawksworth 1990,
Diederich et al. 2002, Hansen & Alstrup 1995,
Ihlen 1998, Triebel 1989, Zhurbenko 2000,
Zhurbenko & Hafellner 1999 and Zhurbenko &
Santesson 1996.
During field trips to different places throughout
the Arctic, the authors have collected many fungi
growing on Cladonia. Additional findings were
made during investigation of herbarium collec-
tions. The aim of this study is to describe the new
species found and to report on lichenicolous fungi
which are insufficiently known, new to different
regions, or found on new hosts. Special emphasis
has been given to lichenicolous fungi growing on
Cladonia pocillum and other Cladonia species
with a well-developed primary thallus. The study
is mainly based on the Arctic material. However, a
few additional, more southern collections are also
included. We also found it appropriate to include a
worldwide key to almost all lichenicolous fungi
growing on Cladonia.
Material and methods
The study is based on specimens deposited in the
following herbaria: LE (194 collections, mostly
collected by MZ), C (40 collections, partly col-
lected by VA) and KPABG (18 collections of vari-

478 Mikhail P. Zhurbenko and Vagn Alstrup
Symb. Bot. Ups. 34:1
ous collectors). The collections are deposited in
LE unless otherwise indicated. The Arctic collec-
tions are from Greenland (20 collections), Sval-
bard (38), Taymyr Peninsula (19), Severnaya
Zemlya Archipelago (11), Lena River delta (14),
Wrangel Island (22), Chukotka (11), Alaska (46),
and the Canadian Arctic Archipelago (8). Extra-
Arctic regions of the Holarctic are mainly from the
Murmansk Region, Russia (18 collections). Li-
chen nomenclature mainly follows Santesson et al.
(2004).
External morphology was studied with dissect-
ing microscope (MBS-1 or Wild). For microscopic
characters hand-made sections or squash prepara-
tions were studied using LOMO MBR-3 or Rei-
chert light microscopes. Photographs were taken
with a Nikon Coolpix 5000. Macroscopic meas-
urements refer to dry herbarium specimens. Mi-
croscopic measurements were performed in water,
0.5% Lugol’s solution with and without pre-treat-
ment with K (K/I and I respectively), 10% KOH
(K), 1% Brilliant Cresyl blue (BCr), and/or in am-
moniacal erythrosin. Unless otherwise indicated,
they refer to water mounts. Microscopic measure-
ments either represent the extreme range or are
given as: (min.–){X´–SD}–X´–{X´+SD}(–max.)
rounded to the nearest 0.5 µm, where min. and
max. are the extreme values, X´the arithmetic
mean, and SD the corresponding standard devia-
tion.
Hosts
New host lichens were found for Cercidospora
cladoniicola (Cladonia subgen. Cladonia), C.
punctillata (Cladonia), Dactylospora deminuta
(Cladonia), Epicladonia simplex (C. pocillum,C.
symphycarpia), E. stenospora (C. pocillum), Li-
chenosticta alcicornaria (C. pocillum), Niesslia
cladoniicola (C. pyxidata), Phaeosporobolus alpi-
nus (C. pocillum), Phoma (Cladonia), Protothe-
lenella santessonii (C. pyxidata), P. sphinctri-
noidella (Cladonia), Rinodina egedeana (Clado-
nia) and Taeniolella beschiana (C. pyxidata,C.
symphycarpia).
The most frequently colonized Cladonia spe-
cies are C. pocillum, C. pyxidata and C. symphy-
carpia, the first one being much more hospitable
than the others. Surprisingly very few fungi were
found on Cladonia coccifera,C. gracilis,C. mac-
roceras, and C. stricta, which are common and
sometimes dominant in the Arctic tundra.
Species
The species are presented in alphabetical order. Occa-
sional notes on species morphology, geography, host
ranges, and taxonomy are provided. Lichenized fungi
are marked with asterisks. Abbreviations: MZ=Mikhail
Zhurbenko, VA=Vagn Alstrup.
Arthonia epicladonia (Nyl.) Alstrup & Zhurb.
comb. nov.
Basionym: Lecidea epicladonia Nyl., Flora 70: 132
(1887). – Scutula epicladonia (Nyl.) Sacc., Sylloge Fun-
gorum 18: 175 (1906).
Apothecia black, subglobose and often appressed,
constricted at the base, 75–250 µm in diam., im-
mersed at first, becoming sessile, discrete but ag-
gregated into groups; hymenium olive-brownish,
I–, K/I+ blue; paraphyses c. 1.5 µm wide, apically
to 4 µm and olivaceous; asci broadly clavate,
short-stalked, 28–50 ×10–17 µm, wall and tholus
K/I+ blue, 8-spored. Spores colourless, lecythi-
form (skittle-shaped), guttulate, 1-septate, (10–)
14–15.5–17.5(–20) × 5–5.5(–6) µm (n=15, in wa-
ter, I or K/I).
DISTRIBUTION AND HABITAT. This fungus has sporad-
ically been reported on Cladonia from Greenland,
Russian Arctic, Sweden, British Isles and the
mountains of Papua New Guinea (Alstrup &
Hawksworth 1990, Aptroot et al. 1997, Hawks-
worth 2003, Santesson et al. 2004, Zhurbenko
2001, Zhurbenko & Pospelova 2001, Zhurbenko
& Santesson 1996). Material examined by us from
Alaska grows on podetial and basal squamules
(moribund or not) of Cladonia pocillum and C.
pyxidata.
REMARKS. Alstrup & Hawksworth (1990) first sug-
gested that this species might belong to Arthonia
and provided a drawing. Arthonia digitatae Hafell-

Lichenicolous fungi on Arctic Cladonia 479
Symb. Bot. Ups. 34:1
ner (Hafellner 1999), A. pelvetii (Hepp) Almq. and
A. molendoi (Frauenf.) R.Sant. (Alstrup & Hawks-
worth 1990), as well as A. colombiana Etayo
(Etayo 2002), readily differ from A. epicladonia by
the I+ red hymenium. Arthonia lepidophila (Anzi)
Clauzade et al. (ined.) known on C. pyxidata in
Italy (Hafellner 1999 and further reference therein)
differs from A. epicladonia by its 3-septate spores.
It is possible that A. epicladonia has a greater range
of spore size, as we found a very similar specimen,
which differs in smaller spores (9–)10–10.5–
11.5(–12) × (3–)3.5–4–(–4.5) µm (n=62, in water,
I, or K/I) [Russia, Sakha (Yakutiya) Republic,
lower portions of Lena River, its right tribute Dal-
dyn River, c. 68°30'N, 124°00'E, on podetia of Cla-
donia pyxidata in sparse Larix forest, 1957, Lu-
kicheva. No pathogenic effect observed].
SPECIMENS EXAMINED.USA.Alaska: Northern Alaska, c.
140 km E of Kotzebue, Kobuk Valley Wilderness,
67°06'N, 159°01'W, 50 m, lichen heath with sparse Pi-
cea glauca, 2000, MZ 00446b; 67°07'N, 159°03'W, 40
m, shrubs by abrupt river bank, 2000, MZ 0032.
Bachmanniomyces uncialicola (Zopf)
D.Hawksw.
Conidia (7–)8–8.5–9.5(–11) ×(3–)3.5–4.5–5(–6)
µm (n=31, in water or erythrosin).
DISTRIBUTION AND HABITAT. Hawksworth (1981) re-
ported Bachmanniomyces uncialicola as associ-
ated with gall-like deformations on podetia of the
Cladonia uncialis-group. Later it has also been re-
ported on Cladonia furcata (Etayo & Diederich
1996), C. pyxidata (Zhurbenko 1998), C. arbus-
cula, C. portentosa and C. stellaris (Diederich
2003, Zhurbenko & Otnyukova 2001). It seems to
colonize both podetia and basal squamules, if the
latter are developed. Material examined by us from
Alaska grows on basal squamules (sometimes
swollen and flexuose) of Cladonia pocillum and C.
pyxidata. New to USA.
SPECIMENS EXAMINED.USA.Alaska: Northern Alaska,
Franklin Bluffs, 69º40'N, 148º43'W, dwarf shrub-moss
tundra, 24 Aug. 2001, Walker; c. 140 km E of Kotzebue,
Kobuk Valley Wilderness, 67°07'N, 159°03'W, 50 m, Pi-
cea glauca forest, 2000, MZ 00126.
Cercidospora cladoniicola Alstrup
Peridium brownish-olive; asci cylindrical, with
rounded apex and short stalk, (50–)52.5–57.5–
63(–65) ×9–10–10.5(–12) µm (n=13, in water or
K/I). Spores at first colourless but later pale oliva-
ceous (with shade of straw), fusiform, symmetri-
cal, straight, guttulate, with smooth wall c. 0.5 µm
thick, (1–)3-septate, constricted at the septa, (13–)
16–18–20(–21) × (3–)4–4.5–5 µm (n=33).
DISTRIBUTION AND HABITAT.Cercidospora cladonii-
cola has previously been reported on C. arbuscula,
C. arbuscula ssp. mitis and C. portentosa in Nor-
way, Denmark, British Isles and Russia (Alstrup
1997, Alstrup & Svane 1998, Coppins 1998,
Zhurbenko 2002). Material examined by us from
Svalbard grows on extensively bleached portions
of the upper side of basal squamules of C. pocillum
and C. symphycarpia. New to Svalbard.
REMARKS. In the species protologue (Alstrup 1997)
spores are given as colourless, with the lower cells
narrower than the upper ones, 16–20 × 5–6 µm.
SPECIMENS EXAMINED.Svalbard. Spitsbergen, Bünsow
Land, 2 km SW of Nordenskiöldbreen glacier, Nord-
dammen Lake, 78º38'N, 16º44'E, 10 m, moist mossy
tundra, 2003, MZ 03195 & 03243.
Cercidospora punctillata (Nyl.) R.Sant.
C. lichenicola (Zopf) Hafellner
Spores colourless, narrowly lecythiform (skittle-
shaped), (1–)3–5-septate, 12–27 ×4–7 µm (n=46,
in water or I).
DISTRIBUTION AND HABITAT.Cercidospora punctil-
lata s. lat. is common in the Arctic. It is a rather
ubiquitous species, colonizing a wide range of un-
related hosts, such as Mycobilimbia, Peltigera,
Phaeorrhiza, Protopannaria, Psoroma, Solorina
and Sphaerophorus (e.g. Hafellner 1987, Santes-
son et al. 2004, Zhurbenko & Santesson 1996),
particularly when they are in a moribund stage.
Material examined by us from the Arctic grows on
basal squamules (sometimes moribund) of Clado-
nia pocillum and C. cf. pyxidata. New to Jan
Mayen and Cladonia is a new host.

480 Mikhail P. Zhurbenko and Vagn Alstrup
Symb. Bot. Ups. 34:1
Specimens examined. Jan Mayen. Blytts Bjerg, 4 Aug.
1919, Gandrup (C). Svalbard. Spitsbergen, Norden-
skiöld Land, W coast of Grønfjorden near Aldegondab-
reen glacier, 78º00'N, 14º12'E, 50 m, moist tundra, 2003,
MZ 03189. Russia.Franz Josef Land: Hooker Island,
Cape Sedov, 80°20'N, 53°00'E, 100 m, 1930, Savicz 432
(C); Scott Keltie Island, 80°20'N, 52°20'E, 30 m, 1930,
Savicz 993a (C). Wrangel Island: Sovetskaya River ba-
sin, 71°13'N, 178°59'E, 200 m, scree tundra, 22 Aug.
1991, Schtrik.
Dactylospora deminuta (Th.Fr.) Triebel
Spores pale to medium brownish olive, fusiform
to subcylindrical, with rounded apices, (3–)5–7-
septate, rarely with additional oblique septum,
(15–)17.5–19.5–21.5(–25) ×(4.5–)5–5.5–6(–7)
µm (n=50).
DISTRIBUTION AND HABITAT.Dactylospora deminuta
is a common species in the Arctic, growing on
many terricolous lichen genera. Material exam-
ined by us from the Russian Arctic grows on mor-
ibund or healthy-looking basal squamules and po-
detia of Cladonia pocillum.
REMARKS.Cladonia is a new host genus for this
species. Other reported Dactylospora species on
Cladonia include D. cladoniicola Alstrup &
Olech, known only from the type locality at Sval-
bard on C. macrophyllodes (Alstrup & Olech
1993), and Dactylospora sp. with brown, 1-septate
spores, 10.5–15 ×5–7 µm, growing on C. pyxidata
at Taymyr Peninsula (Zhurbenko 1998).
SPECIMENS EXAMINED.Russia.Siberia: Severnaya
Zemlya, Bol’shevik Island, Akhmatova Bay, 79°04'N,
102°41'E, 40 m, coastal plain, 1996, MZ 96836; Nor-
denshel’d Archipelago in the Kara Sea, Russkii Island,
16 July 1993, Mel’nikov. Sakha (Yakutiya)Republic:
Lena River delta, 3 km E of Cape Krest-Tumsa,
72°22'N, 126°42'E, 50 m, patchy Dryas tundra, 1998,
MZ 98228.
Epicladonia sandstedei (Zopf) D.Hawksw.
Conidia (7–)9–10–11(–12) ×(2.5–)3–3.5(–4) µm
(n=52).
DISTRIBUTION AND HABITAT. Material examined by
us grows on podetia and basal squamules (usually
in galls) of Cladonia pocillum and C. pyxidata.
New to Ireland.
SPECIMENS EXAMINED.Ireland. Clare Co., N of Ennis, 19
Aug. 1952, Böcher, det. M. S. Christiansen (C). Russia.
Murmansk: Khibiny Mts., Severnyi Lyavochorr Mt., c.
67°55'N, 33°50'E, 600 m, mountain tundra, 1988, Du-
doreva 402 (KPABG, dupl. in LE); Kandalaksha Bay of
White Sea, Kolvitskaya Guba, Klochikhinskaya Mt., c.
67°05'N, 32°55'E, 500 m, boulder field in Betula nana
tundra, 1988, Dudoreva 67 (KPABG). Siberia: Taymyr
Peninsula, NW coast of Pyasino Lake, N’yapan hills,
70°05'N, 87°43'E, 100 m, herb-shrub-moss tundra, 11
July 1999, Zanokha. USA.Alaska: Northern Alaska, c.
140 km E of Kotzebue, Kobuk Valley Wilderness,
67°07'N, 159°03'W, 40 m, shrubs on abrupt river bank,
2000, MZ 00128b; Sagwon, 69°26'N, 148°40'W, 280 m,
Dryas tundra, 7 Aug. 2003, Walker; ibid., 100 m, hill
slope, moist Eriophorum-Betula nana tundra, 22 Aug.
2000, Walker.
Epicladonia simplex D.Hawksw.
Conidia (8–)8.5–9.5–10.5(–12) ×3–3.5–4 µm
(n=30, in erythrosin).
DISTRIBUTION AND HABITAT. This rather rarely re-
ported species is known from Finland, Russia,
Canada and USA (Alstrup & Cole 1998, Cole &
Hawksworth 2001, Hawksworth 1981, Zhurbenko
2004). Material examined by us grows on basal
squamules and podetia (sometimes in galls) of
Cladonia pocillum and C. symphycarpia.
SPECIMENS EXAMINED.Russia.Murmansk: Kutsa Re-
serve, Pyukhyakuru, c. 66°40'N, 29°40'E, mossy rock,
1998, Dudoreva 98132 (KPABG). USA.Alaska: North-
ern Alaska, c. 140 km E of Kotzebue, Kobuk Valley Wil-
derness, 67°06'N, 159°01'W, 50 m, lichen heath with
sparse Picea glauca, 2000, MZ 00447.
Epicladonia stenospora (Harm.) D.Hawksw.
Pycnidia dark brown near the ostiole, which is
50–75 µm in diam., colourless below, 80–125 µm
in diam., immersed. Conidia colourless, broadly
bacilliform, narrowly ellipsoidal, or irregular in
shape, with rounded apex and usually attenuated
and truncated base, simple, 8–9.5–10.5(–12) ×
(3–)3.5–4–4.5(–5) µm (n=25).

Lichenicolous fungi on Arctic Cladonia 481
Symb. Bot. Ups. 34:1
DISTRIBUTION AND HABITAT. Material examined by
us from Svalbard grows on the upper side of basal
squamules (mostly in globose gall-like swellings)
of Cladonia pocillum.E. stenospora has not previ-
ously been reported from Svalbard or from Clado-
nia pocillum.
REMARKS. Hawksworth (1981) reported that Epi-
cladonia stenospora does not produce galls and
has conidia 3–3.5 µm wide.
SPECIMEN EXAMINED.Svalbard. Spitsbergen, Bünsow
Land, 2 km SW of Nordenskiöldbreen glacier, calcifer-
ous foot of Teltfjellet Mt., 78º38'N, 16º44'E, 30 m,
Dryas-Cassiope tundra, 2003, MZ 03226.
*Lecanora leptacinella Nyl.
DISTRIBUTION AND HABITAT. Material examined by
us from the Russian Arctic grows on moribund po-
detia of Cladonia coccifera, and sometimes on
neighbouring mosses (MZ 96835). This is a rather
rare and inconspicuous arctic-alpine lichen grow-
ing on bryophytes, detritus and moribund lichens
such as Cladonia and Sphaerophorus. New to Rus-
sia.
SPECIMENS EXAMINED.Russia.Siberia: Severnaya
Zemlya, Bol’shevik Island, Mushketov glacier, 79°11'N,
102°09'E, 200 m, mountain terrace, MZ 96835; Cape
Antsev, 78°12'N, 103°17'E, 30 m, rolling coastal plain,
boulder field, 16 July 2000, Matveeva.
Lichenoconium pyxidatae (Oudem.) Petr. &
Syd.
Pycnidia 70–150 µm in diam., immersed to al-
most sessile (Hawksworth 1977 reported pycnidia
up to 100 µm in diam.). Conidia pale to medium
olive, usually broadly ellipsoidal, attenuated and
truncated at the base, occasionally globose, wall
smooth to finely verruculose, 3–3.5–4(–5) ×(2–)
2.5–3(–3.5) µm (n=54).
DISTRIBUTION AND HABITAT.Lichenoconium pyxida-
tae has been reported in the Arctic only from Franz
Josef Land (Zhurbenko & Santesson 1996). Mate-
rial examined by us from the Arctic grows on
bleached portions of basal squamules and podetia
of Cladonia pocillum and C. macroceras. New to
USA.
SPECIMENS EXAMINED.Russia.Siberia: Severnaya
Zemlya, Bol’shevik Island, E coast of Shokal’skogo
Strait, 79°16'N, 101°40'E, 20 m, coastal terrace, 1996,
MZ 96831. Sakha (Yakutiya)Republic: Lena River delta,
Bol’shaya Tumatskaya protoka, Amerika-Khaya hill,
72°35'N, 126°18'E, 50 m, 1998, MZ 98231. Wrangel Is-
land: Krasnyi Flag River basin, 71°18'N, 178°55'W, 120
m, 18 July 1997, Kholod. USA.Alaska: Northern
Alaska, Prudhoe Bay, 70°38'N, 148°56'W, 1 m, moist
polygonal graminoid-dwarf shrub tundra, 17 Aug. 2000,
Walker.
Lichenosticta alcicornaria (Linds.) D.Hawksw.
Conidia (7–)8–8.5–9.5(–11.5) ×3–3.5–4(–5) µm
(n=31).
DISTRIBUTION AND HABITAT. This common species
has been reported from many Cladonia species,
including C. pyxidata (e.g. Hafellner 1993). Mate-
rial examined by us grows on both sides of basal
squamules and podetia of Cladonia pocillum and
C. pyxidata. No pathogenic effect observed. New
to the Russian Arctic and Iceland.
SPECIMENS EXAMINED.Iceland.Nordvestur Island: V.-
Bard., Siglunesá, 5 July 1994, Kristinsson (C). Russia.
Murmansk: 1.5 km E of Kandalaksha, Krestovaya Mt.,
c. 67°10'N, 32°15'E, 15 m, mossy boulder in Pinus for-
est, 1965, Dombrovskaya 243 (KPABG); ibid., Kan-
dalaksha Bay of the White Sea, Velikii Island, c.
66°35'N, 33°20'E, mossy rocks, 1964, Piin 92
(KPABG); ibid., Cape Turii, c. 66°35'N, 34°30'E, mossy
coastal cliffs, 15 July 1989, Dudoreva (KPABG). Nenetz
Autonomous Okrug: NW of Bol’shezemel’skaya Tundra,
Bolvanskii Nos Peninsula, Cape Bolvanskii Nos,
68°14'N, 54°28'E, polygonal tundra, 18 July 1999, Lav-
rinenko.
Niesslia cladoniicola D.Hawksw. & W.Gams
Spores colourless, fusiform, with rounded apices,
1-septate, (8–)8.5–9.5–10.5(–11) × 2(–2.5) µm
(n=11).
DISTRIBUTION AND HABITAT. Material examined by
us from the Arctic grows on the moribund base of
podetia of Cladonia pyxidata. It is its second Arc-

482 Mikhail P. Zhurbenko and Vagn Alstrup
Symb. Bot. Ups. 34:1
tic report after Greenland (Hansen & Alstrup
1995). New to Russia.
REMARKS. In the species protologue (Hawksworth
1975) spore size is given as 4.5–8 × 1.5–2 µm.
SPECIMEN EXAMINED.Russia.Siberia: Taymyr Peninsula,
NW coast of Pyasino Lake, N’yapan hills, 70°05'N,
87°43'E, 100 m, Alnus fruticosa tundra, 24 July 1999,
Zanokha.
Phaeosporobolus alpinus R.Sant., Alstrup &
D.Hawksw.
Conidiomata superficial, irregularly convex to
subglobose, 20–70 µm in diam., spread as blackish
speckles over host surface. Conidia pale to me-
dium brownish olive, subglobose, 8–11 µm in
diam., composed of 2–10 (sub)globose finely ver-
ruculose cells 3.5–7 µm in diam. Alstrup &
Hawksworth (1990) reported conidia composed of
10–15(–20) cells 3–4 µm in diam.
DISTRIBUTION AND HABITAT.Phaeosporobolus alpi-
nus has mostly been reported on Ochrolechia and
Pertusaria species, but also on a number of unre-
lated lichen genera such as Baeomyces, Calo-
placa, Flavocetraria, Flavoparmelia, Lecanora,
Megaspora, Parmelia, Umbilicaria, Varicellaria
as well as Cladonia (C. borealis in Alstrup et al.
2000), but never on C. pocillum. Material exam-
ined by us grows on podetia and basal squamules
(exposed medulla on the underside of the squam-
ules and in places with destroyed cortex) of Clado-
nia pocillum. No pathogenic effect observed.
SPECIMEN EXAMINED.Svalbard. Spitsbergen, Bünsow
Land, NE of Billefjorden, calciferous foot of Teltfjellet
Mt., 78º38'N, 16º44'E, 10–50 m, Dryas-Cassiope tun-
dra, 2003, MZ 03230.
Phoma sp.
Pycnidia black, globose, 50–70 µm in diam., im-
mersed to sessile. Conidia oblong to occasionally
slightly reniform, apices usually rounded or occa-
sionally attenuated, simple, 4–5–5.5(–6) ×(1.5–)2
µm (n=50, in erythrosin).
DISTRIBUTION AND HABITAT. Material examined by
us from Svalbard grows in extensively bleached
portions of upper sides and margins of basal squa-
mules of Cladonia pocillum and C. symphycarpia.
It is not clear whether the host’s bleaching is
caused by the fungus.
REMARKS. The material represents an undescribed
species growing on various Cladonia species. It
has been mentioned in Diederich & Serusiaux
(2003, p. 105) and will be described in a forthcom-
ing paper by Diederich and Zhurbenko. There is no
other Phoma species known on Cladonia.
SPECIMENS EXAMINED.Svalbard. Spitsbergen, Bünsow
Land, 2 km SW of Nordenskiöldbreen glacier, shore of
Norddammen Lake, 78º38'N, 16º44'E, 10 m, dwarf
shrub-moss-lichen tundra, 2003, MZ 03222, 03232 &
03234.
Plectocarpon cladoniae R.Sant.
DISTRIBUTION AND HABITAT. This rather rarely re-
ported species has previously been reported in
Russia only from one locality in the Karelian Re-
public (Zhurbenko & Himelbrant 2002). Material
examined by us from the Murmansk Region grows
on the podetial squamules of Cladonia pyxidata.
SPECIMENS EXAMINED.Russia.Murmansk: Khibiny Mts.,
Nephelinovye Peski, c. 67°45'N, 33°20'E, boggy Pinus
forest, 1960, Dombrovskaya 94 (KPABG, dupl. in LE).
Polycoccum laursenii Zhurb. sp. nov.
Similis Polycocco cladoniae sed ab eo imprimis differt
sporis minoribus (7–)8.5–10.5(–12) ×(3–)3.5–4.5(–5)
µm.
Type: USA, N. Alaska, c. 140 km E of Kotzebue, mid-
dle Kobuk River on it’s left bank, Kobuk Valley Wilder-
ness, NW part of the Great Kobuk Sand Dunes, 67°06'N,
159°01'W, 50 m, depression with lichen heath with
sparse Picea glauca, on upper side of basal squamules of
Cladonia pocillum, 4 Aug. 2000, M. Zhurbenko 00448,
associated with Sphaerellothecium araneosum var. cla-
doniae (LE holotype).
ETYMOLOGY. Dedicated to Prof. Gary Laursen, the Ber-
ingian cryptogamist, who made it possible for MZ to
undertake fieldwork in Alaska.
ILLUSTRATION. Fig. 1.

Lichenicolous fungi on Arctic Cladonia 483
Symb. Bot. Ups. 34:1
Perithecia blackish, (sub)globose, without con-
spicuous ostiole, 0.1–0.15 mm in diam., immersed
to ½emergent, arising singly; peridium brown, of
textura angularis, cells 5–8 ×5–4 µm; interascal
filaments sparse, branched and anastomosing, 0.5–
1 µm in diam., tips not swollen; asci cylindrical,
bitunicate, with short apical beak, (50–)55–60–
65(–70) ×6–6.5–8(–9) µm (n=21, in water or
erythrosin), 8-spored, I–, K/I–. Spores at first col-
ourless, then pale to medium brownish-olive, solei-
form with narrower lower cell, guttulate, with ver-
ruculose walls c. 0.5 µm thick, 1-septate with sep-
tum 1 µm thick, constricted at the septum, (7–)8.5–
9.5–10.5(–12) ×(3–)3.5–4–4.5(–5) µm (n=73),
overlapping uniseriate to biseriate. Anamorph not
seen.
DISTRIBUTION AND HABITAT. Material examined by
us from Subarctic Alaska grows on the upper side
of basal squamules of Cladonia pocillum. Infected
portions of the host thalli are more or less
bleached, but this can be due to the associated
Sphaerellothecium araneosum var. cladoniae.
REMARKS. The new species belongs to the group of
Polycoccum species with small spores, and is dis-
tinguished within this group by having the narrow-
est spores. Polycoccum bryonthae (Arnold) Vûzda
differs from P. laursenii in its uniseriate and some-
what larger spores, (10–)11–13(–15) ×(4–)4.5–6
µm, occasionally developing an additional second
septum, and in being restricted to apothecia of Ca-
loplaca and Pertusaria (Hawksworth & Diederich
1988). The two other Polycoccum species known
on Cladonia are P. cladoniae Diederich &
D.Hawksw. (Hawksworth & Diederich 1988) and
P. microcarpum Diederich & Etayo (Etayo & Die-
derich 1998). Polycoccum cladoniae differs from
P. laursenii by its larger spores, (12.5–)13.5–
16.5(–22) ×6.5–8(–9) µm, which are oval with
nearly equal cells and brown, wider asci 9–12 µm
wide, and induction of swellings of the host thal-
lus. Polycoccum microcarpum differs from P.
laurseni in its larger spores (12–14.5 ×4.5–7 µm),
shorter but wider asci (30–35 ×15 µm) and
smaller perithecia (30–60(–100) µm), which are
aggregated in groups of 20–80 and immersed in
galls.
ADDITIONAL SPECIMENS EXAMINED.USA.Alaska: Northern
Alaska, c. 140 km E of Kotzebue, Kobuk Valley Wilder-
ness, 67°07'N, 159°03'W, 40 m, shrubs on abrupt river
bank, 2000, MZ 00128a.
Figure 1. Polycoccum laurse-
nii (holotype). Asci and
spores. Bar 25 µm.

484 Mikhail P. Zhurbenko and Vagn Alstrup
Symb. Bot. Ups. 34:1
Pronectria tibellae Zhurb. sp. nov.
Similis Pronectriae tenacis, sed ab ea imprimis differt
sporis angustioribus (8–)9.5–13.5(–16) × 3–4.5(–5) µm.
Type: USA, N. Alaska, coast of Norton Sound of Ber-
ing Sea, Seward Peninsula, 7 km NE of Nome, Newton
Peak, 64°33'N, 165°22'W, 240 m, open scree nival de-
pression among Salix shrubs, on Cladonia pocillum
(mostly upper side of basal squamules, but also on pode-
tia; often on bleached portions of the host thallus), 5
Sept. 2001, M. Zhurbenko 0166 (LE holotype).
ETYMOLOGY. Dedicated to Prof. Leif Tibell.
ILLUSTRATIONS. Figs 2 & 3.
Perithecia subglobose to obpyriform, without se-
tae or papillae, glabrous, 0.1–0.15 mm in diam.,
immersed and protruding only in the ostiolar area
with elevated yellow margin and intensively scarlet
center, pale yellow in immersed parts, all parts
K–, scattered to occasionally contiguous; interascal
filaments absent; asci cylindrical, with truncated
apex, 70 ×7–8 µm (n=6, in erythrosin), K–, K/I–,
with 8 overlapping uniseriate spores. Spores col-
ourless, fusiform, guttulate (often with 4 large gut-
tules), with thin verruculose walls, (0–)1-septate,
usually not constricted at the septum, but occasion-
ally (old spores?) markedly constricted, (8–)9.5–
11.5–13.5(–16) ×3–3.5–4.5(–5) (n=90, in water or
erythrosin). Anamorph not seen.
DISTRIBUTION AND HABITAT. Material examined by
us from Arctic and Subarctic Alaska grows mostly
on the upper side of basal squamules, but also on
podetia of Cladonia acuminata, C. pocillum and
C. symphycarpia. Infections are often associated
with bleached portions of the host thallus, but it is
not clear whether the fungus is the bleaching
agent.
REMARKS. Following Rossman et al. (1999), the
new fungus belongs to Pronectria. Its spores are
amongst the smallest, and particularly the narrow-
est ones in the genus. The new species differs
from, for instance, Pronectria tenacis in its nar-
rower, persistently colourless, overlapping uniseri-
ate spores, narrower asci, absence of papillae of
perithecia, and different host genus (Cladonia vs.
Collema). Four other nectrioid species are known
on Cladonia, viz. Trichonectria cladoniicola (Al-
strup & Svane) Alstrup, Nectriopsis cladoniicola
M.Cole & D.Hawksw., Nectriopsis parmeliae
(Berk. & M.A.Curtis) M.Cole & D.Hawksw., and
Paranectria oropensis (Cesati) D.Hawksw. &
Piroz. (Alstrup & Svane 1998, Cole & Hawks-
worth 2001, Hawksworth, 1982). Trichonectria
cladoniicola differs by its setose perithecia, P. oro-
pogensis by its muriform spores and Nectriopsis
cladoniicola and N. parmeliae by their superficial
perithecia.
Figure 2. Pronectria tibellae (holotype). Perithecia. Ba
r
0.25 mm.
Figure 3. Pronectria tibellae (holotype). Spores. Bar 5
µm.

Lichenicolous fungi on Arctic Cladonia 485
Symb. Bot. Ups. 34:1
ADDITIONAL SPECIMENS EXAMINED.USA.Alaska: Northern
Alaska, c. 140 km E of Kotzebue, Kobuk Valley Wilder-
ness, 67°07'N, 159°03'W, 50 m, open margin of abrupt
river bank in Picea glauca forest, 2000, MZ 00449 (as-
sociated with Sphaerellothecium araneosum var. clado-
niae); coast of Norton Sound of Bering Sea, Seward Pe-
ninsula, 7 km NE of Nome, Newton Peak, 64°33'N,
165°21'W, 240 m, open scree nival depression among
Salix shrubs, 2001, MZ 01101; 250 m, open roadside
with Salix shrubs, 2001, MZ 0167 & 01617; 220 m, Sa-
lix-dwarf-shrub-lichen-moss tundra, 2001, MZ 0165.
Protothelenella santessonii H.Mayrhofer
Asci subcylindrical, with rounded apices, 80–130
×16–18 µm, (2–)8-spored. Spores colourless,
usually oval to narrowly ellipsoidal, rarely broadly
ellipsoidal to almost globose, usually with acute
apices, symmetrical or more often attenuated at
one apex and there often with an apiculus about 2
µm long and 0.75–1 µm wide, usually (sub)muri-
form with 3–7 transverse or occasionally oblique
septa and 1 longiseptum in central segments or oc-
casionally only transversely septate, (12–)18–
21.5–25(–28) ×(6–)7.5–9–10(–12) µm (n=84),
overlapping uniseriate in ascus, the upper 4 with
the attenuated apex oriented downwards and the
lower 4 upwards.
DISTRIBUTION AND HABITAT. This rather rarely re-
ported fungus is previously known in the Arctic
only from Greenland (Alstrup 1995). Protothe-
lenella santessonii usually grows on Cladonia spe-
cies, but was once reported on Solorina crocea
(Alstrup & Cole 1998). Material examined by us
grows on basal squamules of Cladonia pocillum,
C. pyxidata and Cladonia sp. It is often associated
with bleached portions of the host thallus. New to
Svalbard and reported for the second time for
USA.
REMARKS. Mayrhofer (1987) did not mention the
characteristic spore apiculus, and the spores were
also given as somewhat wider (18–24 ×10–12
µm). The apiculus is similar to those known at one
or both apices of Protothelenella leucothelia
spores (Mayrhofer & Poelt 1985, Fig. 7). Accord-
ing to H. Mayrhofer (pers. comm.) such an apicu-
lus is a good character at species level in the genus.
SPECIMENS EXAMINED.Svalbard. Spitsbergen, Bünsow
Land, NE of Billefjorden, calciferous foot of Teltfjellet
Mt., 78º38'N, 16º44'E, 30 m, spotty Dryas tundra, 2003,
MZ 03229; Dickson Land, W of Billefjorden, Gar-
maksla Mt., 200 m, below Ca-rich rocks with bird nests,
2003, MZ 03184. Russia.Novaya Zemlya: Northern Is-
land, Sporyi Navolok Peninsula, Ledyanaya Gavan’
Bay, Cape Medvezhii, 76°15'N, 68°10'E, tundra on
limestones, 1 Sept. 1995, Kuliev. Bashkortostan Repub-
lic: Southern Urals, 10 km S of Tyulyuk, Bol’shoi
Iremel’ Mt., 54°30'N, 58°50'E, 1100 m, 3 July 1982,
Bredkina. Siberia: Severnaya Zemlya, Bol’shevik Is-
land, Akhmatova Bay, 79°04'N, 102°45'E, 20 m, coastal
terrace, 1996, MZ 96834; Taymyr Peninsula, Byrranga
Mts., Levinson-Lessing Lake, 74°24'N, 98°38'E, 150 m,
1995, MZ 95214. Siberia: Western Sayan, Mt. “2735”,
cold mountain desert, 1991, Kuvaev 2099b. Sakha
(Yakutiya)Republic: Lena River delta, 3 km E of Cape
Krest-Tumsa, 72°22'N, 126°42'E, 50 m, spotty Dryas
tundra, 1998, MZ 98227; 3 km W of Cape Krest-Tumsa,
Ebe-Khaia Island, 72°24'N, 126°40'E, 50 m, tundra with
rocky outcrops, 1998, MZ 98236; Olenek protoka, Chai-
Tumus cabin, 72°22'N, 125°40'E, 30 m, spotty tundra,
1998, MZ 98234. Wrangel Island: Pravaya Gusinaya
River valley, 71°09'N, 179°15'E, 115 m, Salix polaris
tundra, 1991, Kholod; headwaters of Gusinaya River,
71°09'N, 179°37'E, 390 m, wet Eriophorum-bryophyte
tundra, 1996, Kholod. Chukotka: Center of Chukchi Pe-
ninsula, 20 km SE of Ioni Lake, Gil’mymlinei hot
springs, 65°48'N, 173°15'W, 19 July 1977, Makarova.
USA.Alaska: Northern Alaska, Franklin Bluffs,
69°40'N, 148°43'W, 130 m, moist Carex-Dryas tundra,
17 Aug. 2000, Walker.
*Protothelenella sphinctrinoidella (Nyl.)
H.Mayrhofer & Poelt
Thallus not found. Peridium citrine. Spores col-
ourless, fusiform with attenuated apices, occasion-
ally oblong with more rounded apices, straight,
symmetrical or slightly asymmetrical, submuri-
form with 6–10 transverse or oblique septa and
usually 1 longiseptum in central segments, (22–)
24.5–27–29 ×6.5–7–7.5(–8.5) µm (n=20), over-
lapping uniseriate.
DISTRIBUTION AND HABITAT. This infrequently re-
ported species grows on soil, moribund bryophytes
and plant detritus including lichen thalli. It has
been reported on moribund foliose lichens such as
Peltigera (Mayrhofer 1987). Material examined
by us from the Subarctic grows on decaying pode-

486 Mikhail P. Zhurbenko and Vagn Alstrup
Symb. Bot. Ups. 34:1
tia and basal squamules of Cladonia sp. and neigh-
bouring bryophytes.
REMARKS.Protothelenella sphinctrinoidella usu-
ally has a visible, although thin, membrane-like
lichenized thallus.
SPECIMEN EXAMINED.Russia.Siberia: Putorana Plateau,
Yt-Kyuel’ Lake, 700 m, mountain tundra, July 1996,
Matveeva.
Rinodina egedeana (Linds.) Alstrup &
D.Hawksw.
Thallus absent. Apothecia sessile, constricted at
base, to 0.25 mm in diam.; exciple persistent,
fawn. Spores with strongly thickened apical walls
and thick median septum, 25–30 × 11–13 µm.
DISTRIBUTION AND HABITAT. Material examined by
us from Greenland grows on moribund thalli of
Cladonia pocillum associated with Sphaerel-
lothecium araneosum var. cladoniae. The species
has previously only been reported from moribund
Parmelia saxatilis and Peltigera sp. in W. Green-
land (Alstrup & Hawksworth 1990).
SPECIMEN EXAMINED.Greenland. Kronprins Christian
Land, Centrum Sø, 2 July 1917, Wulff (C).
Roselliniella cladoniae (Anzi) Matzer &
Hafellner
Spores colourless when immature but becoming
finally dark rusty tawny, ellipsoidal, occasionally
narrowly citriform or subglobose, wall smooth,
but in overmature spores disintegrated into scale-
like pieces, simple, (15–)19–21–23.5–(26) ×(9–)
9.5–11.5–13.5(–17) µm (n=42), biseriate or over-
lapping uniseriate.
DISTRIBUTION AND HABITAT. Previously known in the
Arctic only from Greenland (Alstrup 1993a). The
species is rather common in more southern regions
and grows on various Cladonia species. Material
examined by us grows on basal squamules of Cla-
donia pocillum, C. pyxidata and Cladonia sp., ei-
ther attached to the underside of the squamules or
growing through them. New to Russia.
SPECIMENS EXAMINED.Russia.Murmansk: Khibiny Mts.,
Vud’yavrchorr Mt., 67°40'N, 33° 32'E, 500 m, belt of
bent Betula forest, 1961, Dombrovskaya 4 (KPABG).
Chukotka: Center of Chukchi Peninsula, Ioni Lake, foot
of Gil’mymlinei Mt., 65 48'N, 173 15'W, sedge-moss-
lichen tundra, 3 July 1977, Makarova.
Sphaerellothecium araneosum (Arnold) Zopf
var. cladoniae Alstrup & Zhurb. var. nov.
Differt ab varietate typica in Ochrolechia peritheciis et
mycelio partim immersis, parietibus cellularibus mycelii
granulosis, ascis majoribus (20–)21–30(–36) ×(11–
)11.5–15(–16) µm, sporis angustioribus (7–)9–12.5(–
14) ×(2.5–)3–4.5(–5) µm, et hospite diverso (Cladonia).
Type: Svalbard, Spitsbergen, Nordenskiöld Land, W
coast of Grønfjorden near Aldegondabreen glacier,
78º00'N, 14º12'E, 50 m, depression with small lake,
moist graminoid-dwarf shrub-lichen-moss tundra, on
Cladonia pocillum (bleached basal squamules), 16 July
2003, M. Zhurbenko 03216 (LE holotype).
ETYMOLOGY. Refers to the host genus.
ILLUSTRATION. Fig. 4.
Mycelium dark brown (sienna coloured in water),
sparsely branched with side-branches deviating at
angles of 45–90º, superficial or semi-immersed;
young hyphae mostly straight, smooth- and thin-
walled, of cells 5–9 µm long and 3–5 µm thick, not
constricted at septa, older hyphae becoming
strongly torulose and constricted between cells,
thick-walled and covered by granules c. 1 µm high
and 2 µm in diam., to 8.5 µm thick. Perithecia
Figure 4. Sphaerellothecium araneosum var. cladoniae
(holotype). Perithecia and mycelial network. Bar 0.5
mm.

Lichenicolous fungi on Arctic Cladonia 487
Symb. Bot. Ups. 34:1
brownish-black, shiny, (sub)globose, often with
conical apices, without appendages, (25–)45–60
(–70) µm in diam., ostiolate with pore 10–15 µm
in diam., ¾immersed to superficial, arising singly
or exceptionally contiguous, usually associated
with superficial mycelial network, but in some old
infections the latter is scanty or even not observed;
exciple of c. 2 layers of polyhedral cells (4–)6–10
×(3–)5–11 µm, smooth-walled at first, the outer
layer becoming granulose like the mycelium; inte-
rascal filaments not seen; asci bitunicate, saccate,
obpyriform or ellipsoidal, with short stalk, tholus
c. 4 µm tall, sometimes with protruding narrow
central appendage, (20–)21–25.5–30(–36) ×(11–)
11.5–13.5–15(–16) µm (n=19, in water, eryth-
rosin, I, or K/I), 8-spored, ascoplast K/I+ yellow-
orange, otherwise all parts K/I–, I–, BCr–. Spores
usually colourless, rarely becoming pale olive
greyish or yellowish brown with age, sometimes
with 4 guttules, with epispore c. 1 µm thick, solei-
form or lecythiform (skittle-shaped), 1-septate,
slightly constricted at septum, (7–)9–10.5–12.5
(–14) ×(2.5–)3–4–4.5(–5) µm (n=83, in water,
erythrosin, K, or K/I), overlapping in 2–3 rows in
an ascus; sporoplast BCr+ blue.
DISTRIBUTION AND HABITAT.Sphaerellothecium ara-
neosum var. cladoniae is extremely common and
abundant at least throughout the Arctic (Fig. 5) on
Cladonia pocillum and C. symphycarpia, while
less common on C. coccifera, C. gracilis, C. mac-
roceras, C. pyxidata and C. subcervicornis. It fol-
lows the hosts in the boreal zone and southwards
to alpine habitats in the central Asian mountains.
Surprisingly it has not yet been found on collec-
tions of Cladonia species from the Alps. It mostly
grows on the upper side of basal squamules, occa-
sionally on their underside and on podetia. The
fungus is clearly pathogenic, often inducing strong
bleaching of the host thallus, not encircled by any
ring of blackened lichen tissue. In infections on C.
symphycarpia bleaching was not seen.
Figure 5. Distribution of Sphaerellothecium araneosum var. cladoniae

488 Mikhail P. Zhurbenko and Vagn Alstrup
Symb. Bot. Ups. 34:1
REMARKS. The type host lichen species of
Sphaerellothecium araneosum is Ochrolechia up-
saliensis. Alstrup & Hawksworth (1990) provided
a detailed description of this species (as Echi-
nothecium glabrum M.S.Christ., Alstrup &
D.Hawksw.) on the type host. They found the myc-
elium and ascomata to be superficial, the myc-
elium torulose, of thick-walled hyphae 5–7.5 µm
thick, walls verruculose through splitting of the
outer wall layers; ascomata 30–50(–60) µm in
diam.; ascoma wall pseudoparenchymatous of
cells 8–10 µm in diam.; asci broadly saccate, 20–
25 ×10–15 µm; spores colourless, not or slightly
constricted at septum, (7.5–)9–12(–14.5) ×(4–)
4.5–5(–5.5) µm.
The new variety differs from this description in
its partly immersed ascomata and mycelium, the
cell walls becoming coarsely granulose, bigger
asci, narrower spores, and a different host (Clado-
nia). It also seems to be more pathogenic, though
heavy infections of S. araneosum on Ochrolechia
species can also be distinctly pathogenic, often de-
stroying the hymenium of the host. Measurements
of Alstrup & Hawksworth 1990) resemble the
original description of the species. However, the
description by Roux & Triebel (1994) also based
on material from Ochrolechia and Pertusaria devi-
ates in several respects from this, as well as the one
given by Zopf (1897) based on specimens on Le-
canora badia. Possibly more taxa exist within the
Sphaerellothecium araneosum complex. Awaiting
further studies of the group we have chosen to de-
scribe the fungus as a variety, although it might
deserve another rank. There are two more species
with conspicuous dark superficial mycelial reticu-
lum growing on Cladonia species, viz. Sphaerel-
lothecium cladoniicola E.S. Hansen & Alstrup and
Echinothecium cladoniae Keissl. The former dif-
fers from S. araneosum var. cladoniae by a myc-
elium composed not only of single but often also
of a few parallel, agglutinated hyphae. It also has
entirely superficial perithecia, smaller asci c. 20 ×
10 µm, permanently colourless spores, and differ-
ent host subgenus (Cladina vs. Cladonia). The lat-
ter differs in its setose perithecia. Sphaerel-
lothecium araneosum has been reported (as Echi-
nothecium glabrum) on Cladonia pocillum, Arc-
toparmelia separata and Sphaerophorus fragilis in
Greenland (Alstrup & Hawksworth 1990). The
species on Sphaerophorus has later been described
as S. minutum Hafellner (Hafellner 1993), a com-
mon fungus in the Arctic. Following the broader
species concept, Zhurbenko (1998), Zhurbenko &
Daniëls (2003) and Zhurbenko & Pospelova
(2001) have reported Sphaerellothecium araneo-
sum on Cladonia pocillum from Taymyr and the
Canadian Arctic.
SELECTED ADDITIONAL SPECIMENS EXAMINED.Greenland.
Disko Island, Alakariaq, 18 July 1975, VA (C); Disko,
Qutdligssat, 21 July 1950, Gelting (C); N. Greenland,
Kronprins Christian Land, mountain S of Centrum Sø,
VA & Daniels (C); N. Greenland, Kap Benet, 5 June 1917,
Wulff (C); E. Greenland, Eskimonæs, 8 June 1932, Gelt-
ing (C); Nordre Strømfjord, Nuerssorfiarqap, 25 July
1958, Hansen (C); Hudson Land, Loch Fynes Bund, 3
Aug. 1929, Seidenfaden (C); Julianehåb District, Juliane-
håb, 11 Aug. 1937, Dahl (C); Mestersvig District, Skel-
dal, 27 July 1963, Spearing (C). Svalbard. Wahlenberg
Bay, c. 3 km E of Round Hill, 2 July 1931, Scholander
(C); Forsbladhamna i Bell Sund, 29 July 1926, Lynge (C);
Wijdebay, Østfjorden, 8 Aug. 1899, Wulff (C); Spitsber-
gen, Nordenskiöld Land, W of Grønfjorden near Alde-
gondabreen glacier, 78º00'N, 14º12'E, 20–100 m, grami-
noid-dwarf shrub-lichen-moss tundra, 2003, MZ 03181
& 03208; Bünsow Land, NE of Billefjorden, near Nord-
dammen Lake, 78º38'N, 16º44'E, 5–30 m, Dryas-lichen-
moss tundra, 2003, MZ 03170, 03194, 03221, 03237 &
03244. Great Britain.Wales: Dyfed, Pembrokeshire, SE
coast of Angle Peninsula, Stackpole Warren, 31 Aug.
1995, Christensen (C). Norway.Finnmark: Porsanger
herad, 1973, Aava 12175 (C). Sweden.Torne Lappmark:
N of Nissunjåkka, 28 June 1982, VA (C). Finland.Koil-
lismaa: Juuma Village, 6 July 1981, VA (C). Lapponia
Enontekiensis: Enontekiö, Toskalpahta, 3 Aug. 1955, Hu-
uskonen (C). Kuusamo: Salla, between Riekinlampi and
Pääjärvi, 6 Aug. 1937, Lehtonen & Pankakoski
(KPABG). Russia.Murmansk: near Nikel’, c. 69°25'N,
30°10'E, 1973, Dombrovskaya 73 (KPABG); N coast of
Kandalaksha Bay of the White Sea, Cape Turii, c.
66°35'N, 34°30'E, mossy coastal cliffs, 15 July 1989, Du-
doreva (KPABG). Karelia Republic: Karelia Ladogensis,
Kurkijoki, Lapinlahti, 26 July 1934, Räsänen (C). Franz
Josef Land: Hooker Island, c. 80°20'N, 53°00'E, 100 m,
1930, Savicz 1461 (C). Novaya Zemlya: Northern Island,
Sporyi Navolok Peninsula, Ledyanaya Gavan’ Bay, Cape
Medvezhii, 76°15'N, 68°10'E, tundra on limestones, 1
Sept. 1995, Kuliev. Nenetz Autonomous Okrug: NW of
Bol’shezemel’skaya Tundra, Bolvanskii Nos Peninsula,
Cape Bolvanskii Nos, 68°14'N, 54°28'E, polygonal tun-
dra, 18 July 1999, Lavrinenko. Komi Republic: Northern
Ural, headwaters of Pechora River, top of Mt. Med-

Lichenicolous fungi on Arctic Cladonia 489
Symb. Bot. Ups. 34:1
vezh’ya, 62°03'N, 59°03'E, 700 m, diabase rocks in al-
pine belt, 1997, MZ 97299. Siberia: Severnaya Zemlya,
Bol’shevik Island, E coast of Shokal’skogo Strait,
79°16'N, 101°40'E, 20 m, coastal terrace, 1996, MZ
96421; Taymyr Peninsula, Byrranga Mts., Levinson-
Lessing Lake, 74°33'N, 98°34'E, 200 m, dwarf shrub-
moss-lichen tundra, 1995, MZ 95220; Putorana Plateau,
Yt-Kyuel’ Lake, 700 m, mountain tundra, July 1996,
Matveeva; Western Sayan, Mt. “2735”, cold mountain
desert, 1991, Kuvaev 2099a. Sakha (Yakutiya) Republic:
Lena River delta, D’ieleekh Aryyta Island, 72°22'N,
126°29'E, 10 m, dwarf-shrub-lichen-moss tundra, 1998,
MZ 98224; Lena River between Sinsk and Tit-Ary,
61°04'N, 127°29'E, 150 m, Ca-rich cliffs by the river-side
with steppe, 1992, MZ 92524; Indigirka River at 8 km
NNW of Ust’-Nera, 64°38'N, 143°20'E, 600 m, open
slope with steppe, 1992, MZ 92523. Wrangel Island:
Neizvestnaya River basin, 71°13'N, 179°15'W, 190 m,
spotty Dryas tundra, 23 Aug. 1987, Kholod. Chukotka:
Chukchi Peninsula, Anadyr Bay coast near Sireniki, c.
64°30'N, 174°00'W, 200 m, tundra, 23 July 1986,
Katenin; Lavrentiya Bay, near Lavrentiya, 65°35'N,
171°00'W, Salix-forb tundra, 9 Aug. 1972, Makarova.
Kirgizstan. Alai Mts., Dugaba Valley, 3000 m, 16 July
1996, VA (C). Mongolia.Khangai: Arakhangai Aimak,
Tevshrulekh Somon, Ikh-Ar-Ul Mt., 2450 m, mountain
meadow, 1971, Biazrov 2771; Bayan-Khongar Aimak,
Gurvan-Bulak Somon, Khukh-Nur Lake bank, 2560 m,
1972, Biazrov 6713a; Uverkhangai Aimak, Burd Somon,
Us-Ula Mt., 2180 m, mountain steppe, 1978, Biazrov
5992. USA.Alaska: Northern Alaska, c. 140 km E of
Kotzebue, Kobuk Valley Wilderness, 67°02'N, 158°50'W,
50 m, open lichen heath with sparse Picea glauca and
Populus balsamifera, 2000, MZ 0052; coast of Norton
Sound of Bering Sea, Seward Peninsula, 7 km NE of
Nome, Newton Peak, 64°33'N, 165°22'W, 240 m, Salix
shrubs, 2001, MZ 0172; Toolik Lake, 68°37'N, 149°36'W,
800 m, moss-lichen-forb-dwarf-shrub spotty, 2001, MZ
01447; Deadhorse, 70°10'N, 148°28'W, moist sedge-Dr-
yas-moss tundra, 17 July 2001, Walker. Canada.Cana-
dian Arctic Archipelago: Cornwallis Island, Resolute
Bay, 74°42'N, 94°55'W, Salix arctica-moss tundra, 6 Aug.
1999, Matveeva; Ellesmere Island, Sverdrup Pass,
79°08'N, 81°27'W, 320 m, Carex-Dryas tundra,31 July
1992, Daniëls. Nunavut: Arctic Bay Point, Society Cliff
Formation, c. 73°00'N, 85°00'W, 31 July 1994, Ferris (C).
Taeniolella beschiana Diederich
Conidia (7–)7.5–9–10.5(–13) ×3.5–4–4.5(–5)
µm (n=22).
DISTRIBUTION AND HABITAT. Material examined by
us grows on both sides of basal squamules of Cla-
donia pocillum, C. pyxidata, C. symphycarpia and
C. trassi.Taeniolella beschiana has been reported
from various Cladonia species, including C. pocil-
lum (Coppins 2000). New to Svalbard and USA.
REMARKS. In the species protologue (Diederich
1992) the size of the conidia is given as 6–10 ×
2.5–4 µm.
SPECIMENS EXAMINED.Svalbard. Spitsbergen, Bünsow
Land, 2 km SW of Nordenskiöldbreen glacier, near
Norddammen Lake, 78º38'N, 16º44'E, 5 m, Dryas-
moss-lichen tundra, 2003, MZ 03219 & 03238. Russia.
Murmansk: Khibiny Mts., c. 67°45'N, 33°40'E, 1962,
Prokhorov (KPABG). Siberia: Taymyr Peninsula, Byr-
ranga Mts., Levinson-Lessing Lake, 74°24'N, 98°38'E,
150 m, tundra on limestones, 1995, MZ 95213. Wrangel
Island: Mamontovaya River basin, 71°10'N, 179°45'W,
180 m, 4 Aug. 1995, Kholod. USA.Alaska: Northern
Alaska, c. 140 km E of Kotzebue, Kobuk Valley Wilder-
ness, 67°07'N, 159°03'W, 40 m, shrubs on abrupt river
bank, 2000, MZ 00110; coast of Norton Sound of Bering
Sea, Seward Peninsula, 7 km NE of Nome, Newton
Peak, 64°33'N, 165°22'W, 240 m, nival Salix shrubs,
2001, MZ 01102; Toolik Lake, 68°37'N, 149°36'W, 770
m, rolling plain, moss-lichen-dwarf-shrub moist tundra,
2001, MZ O1454.
Taeniolella strictae Alstrup sp. nov.
Similis Taeniolellae beschianae sed ab ea imprimis dif-
fert conidiis crassioribus 8.5–12 × 5.5–6 µm.
Type: Greenland, Qaanaaq District, Siorapaluk, 50 m,
on Cladonia stricta in dwarf shrub heath, 13 Aug. 1993,
V. Alstrup 6048 (C holotype).
ETYMOLOGY. Named after the host Cladonia species.
ILLUSTRATION. Fig. 6.
Figure 6. Taeniolella strictae (holotype). Conidiophores
and conidia. Bar 10 µm.

490 Mikhail P. Zhurbenko and Vagn Alstrup
Symb. Bot. Ups. 34:1
Mycelium partly immersed and colourless, partly
superficial and dark brown, branched, rough-
walled, to 10 µm in diam. Conidiophores upright,
dense, branched, septate, 25–50 µm high and 5–7
µm thick, smooth-walled; conidiogenous cells in-
tegrated, terminal, phialidic; conidia formed in
chains, 1–2-septate, smooth-walled, 8.5–12 ×5.5–
6 µm.
DISTRIBUTION AND HABITAT. The new species is
known only from the type collection on Cladonia
stricta.
REMARKS. Two other species of Taeniolella are
known from Cladonia spp. T. cladiniicola Alstrup
has a thin mycelium penetrating the host hyphae of
C. arbuscula and C. portentosa. It is known from
Denmark and the Czech Republic (Alstrup 1993a,
1993b; Kocourková 2000). T. beschiana Diederich
has immersed but not intrahyphal mycelium, co-
nidiophores with a thick and roughened wall, and
solitary or catenate, 0–1-septate, thick-walled and
often rough-walled conidia. It is rather common, at
least in Europe, and grows on various Cladonia
species. Other species with smooth cell walls are
T. delicata M.S.Christ. & D.Hawksw., in which
the conidiophores are branched only at the base
and both lower and thinner. T. punctata
M.S.Christ. on Graphis scripta has bigger and
thicker mycelium and conidia. T. phaeophysciae
D.Hawksw. on Phaeophyscia and Physconia spp.
has much larger conidia.
Key to cladoniicolous fungi
The key is based on original data and literature
(e.g. Alstrup 1993b, 1997; Alstrup & Cole 1998;
Alstrup & Hansen 2001; Alstrup & Hawksworth
1990; Alstrup & Olech 1993; Alstrup & Svane
1998; Alstrup et al. 2004; Christiansen 1993; Die-
derich 1992, 1996, 2003; Diederich et al. 2001;
Dodge 1948; Etayo & Diederich 1996, 1998;
Hafellner 1999; Hansen & Alstrup 1995; Hawks-
worth 1975, 1977, 1978, 1981, 1982, 1990;
Hawksworth & Diederich 1988; Hawksworth &
Santesson 1990; Matzer & Hafellner 1990;
Mayrhofer 1987; Mayrhofer & Poelt 1985;
Santesson 1994; Santesson & Tønsberg 1994;
Triebel 1989). It does not include most of the dis-
tinctly lichenized species ocurring on Cladonia
such as Anzina carneonivea (Anzi) Scheid. var.
carneonivea,Cladonia luteoalba A.Wilson &
Wheldon, Diploschistes muscorum (Scop.)
R.Sant., Japewia tornoënsis (Nyl.) Tønsberg, and
Lecanora leptacinella Nyl., as they are easy to
identify with other keys, while the weakly lichen-
ized species of Merismatium and Protothelenella
are included. Excluded are also the anamorphic
species of Acremonium (A. lichenicola W.Gams,
A. rhabdosporum W.Gams) and Monocillium,
known only from culture, and some poorly known
species – Pezizella ucrainica S.Y.Kondr., Scutula
cladoniarum (Müll.Arg.) Rambold & Triebel
(ined.), Spilomium epicladoniae H.Olivier, and
Verrucaster lichenicola Tobler. Hypoxylon lichen-
icolum Höhn. was also reported from Cladonia
sp. (Höhnel 1927). However, the supposed Hy-
poxylon ascomata are found on the stromata of a
Xylaria sp. resembling a Cladonia, and they are
in fact the ascomata of that Xylaria (Læssøe, pers.
comm.). The included insufficiently known spe-
cies are given in square brackets. Outlines of the
known species distribution is provided. Author ci-
tations are given for species not treated above.
1. Fungus a basidiomycete .....................................................................................................................2
– Fungus an ascomycete or deuteromycete...........................................................................................5
2. Basidia non-septate ............................................................................................................................3
– Basidia septate....................................................................................................................................4
3. Basidiomata originating as a cottony web of pale yellow fibres, membranaceous,
merulioid; basidiospores buff or pale yellow, ellipsoid or oval, smooth, c. 4.5–5.6 ×
3.0–3.7 µm; produces conspicuous sclerotia; boreal Canada and Fennoscandia.................................
....................................................................... Leucogyrophana lichenicola Thorn, Malloch & Ginns

Lichenicolous fungi on Arctic Cladonia 491
Symb. Bot. Ups. 34:1
– Basidiomata adnate to gall inducing, often elongate and bent, or tuberculate to
irregular, waxy to gelatinous reddish brown; basidiospores 7–9 × 4.5–6 µm;
widespread........................................................... Syzygospora bachmannii Diederich & M.S.Christ.
4. Basidiomata inducing pinkish brown, convex galls, becoming subspherical, to
2 mm in diam.; basidia with one transseptum; basidiospores 7–10 × 6–8 µm;
widespread.................................................................... Tremella cladoniae Diederich & M.S.Christ.
– Galls absent; basidiomata subglobose to pulvinate, to 0.8 mm in diam.;
basidia with one longiseptum, rarely with oblique or transseptum; basidio-
spores 5.5–7.5 × 4–5.5 µm; Norway ...........................Tremella macroceratis Diederich & Hafellner
5. Fungus an ascomycete........................................................................................................................6
– Fungus a deuteromycete...................................................................................................................56
6. Ascomata perithecioid or catathecioid ...............................................................................................7
– Ascomata apothecioid .....................................................................................................................34
7. Ascomata yellow, orange or red; spores colourless............................................................................8
– Ascomata dark brown to black, without red tinge; spores colourless to dark coloured...................13
8. Spores filiform, simple, 165–175 × 5–6 µ; perithecia with subiculum; Denmark
...................................................................................................... Barya lichenophila Ferd. & Winge
– Spores much shorter, not filiform; subiculum absent.........................................................................9
9. Perithecia in upper part with stiff setae of few to many non-septate, white hairs,
sessile, pale yellow to bright orange, 0.1–0.2 mm in diam.; spores narrowly fusi-
form, 1-septate, 26–30 × 3–5 µm; Denmark ........................................................................................
........................................................................Trichonectria cladoniicola (Alstrup & Svane) Alstrup
– Perithecia without setae....................................................................................................................10
10. Spores muriform with 5–8 transverse and 1–2 longitudinal septa, (22–)25–32(–36)
× (9–)11–14(–15) µm, with cauda to 12 µm long at both apices; widespread .....................................
............................................................................. Paranectria oropensis (Ces.) D.Hawksw. & Piroz.
– Spores only transversely septate, without appendages.....................................................................11
11. Perithecia immersed, exposed only in ostiolar area which is scarlet with yellow
margin; spores fusiform, (0–)1-septate, (8–)9.5–13.5(–16) × 3–4.5(–5) µm; USA
(N. Alaska) ........................................................................................................... Pronectria tibellae
– Perithecia superficial ........................................................................................................................12
12. Spores 1-septate, broadly ellipsoid, 6.5–8.5(–9.5) × 3.5–4.5(–5) µm; peri-
thecia pale pink but more intensively pigmented around the ostiole, covered
with a loose open weft of hyphae; USA (Massachusetts)....................................................................
..................................................................................Nectriopsis cladoniicola M.Cole & D.Hawksw.
– Spores 1–3-septate, of different size within the same ascus, 8–17 × 3–7 µm
and 34–60 × 12–20 µm; widespread ....................................................................................................
..................................................Nectriopsis parmeliae (Berk. & M.A.Curtis) M.Cole & D.Hawksw.
13. Ascomata catathecia, without setae, ostiolate, black, 50–70 µm in diam.,
sessile; asci c. 36 × 14 µ; spores ellipsoid, 1-septate, colourless, 18–21 ×
5–6 µm; Greenland, Iceland, and N Europe.....Lichenopeltella cladoniarum E.S. Hansen & Alstrup
– Ascomata perithecia .........................................................................................................................14
14. Ascomata with stiff setae, superficial; spores 1-septate, colourless.................................................15
– Ascomata without setae; spores various...........................................................................................16
15. Ascomata sitting on conspicuous superficial dark reticulate mycelium; spores
8–11 × 3.5–4.5 µm; Colombia and Europe .....................................Echinothecium cladoniae Keissl.

492 Mikhail P. Zhurbenko and Vagn Alstrup
Symb. Bot. Ups. 34:1
– Conspicuous superficial dark reticulate mycelium absent; spores narrower,
4.5–11 × 1–2.5 µm; widespread .........................................................................Niesslia cladoniicola
16. Spores coloured at maturity..............................................................................................................17
– Spores mostly colourless, rarely coloured when old........................................................................24
17. Spores usually simple, occasionally 1–4-septate, ellipsoidal, fusiform, narrowly
citriform or occasionally subglobose, at first colourless, dark brown at maturity,
with verruculose and finally scaling wall, 15–52 × 6–17 µm; asci 2–8-spored, uni-
tunicate; perithecia to 0.7 mm in diam. and to 0.7 mm tall, ovoid, sometimes papillate,
black, with rugose surface, often sessile; widespread.....................................Roselliniella cladoniae
– Spores always septate, shorter than 20 µm, often paler and with olive tone;
asci bitunicate; perithecia not wider than 0.25 mm..........................................................................18
18. Spores 1(–3)-septate.........................................................................................................................19
– Spores 3-septate to muriform, often pale .........................................................................................22
19. Interascal filaments absent; spores 1(–3)-septate, 9–11.5 × 3–4 µm, soleiform,
brown; perithecia 40–70 µm in diam., at first semi-immersed, later superficial;
vegetative hyphae not forming a superficial net on the host thallus, irregularly
immersed, visible at high magnification; infected host thallus becoming character-
istically grey-tinged; in oceanic woods; Pyrénées ...............................................................................
.............................................................................Sphaerellothecium cinerascens Etayo & Diederich
– Interascal filaments present; spores 1-septate, brown to olive-brown..............................................20
20. Perithecia 30–60(–100) µm in diam., aggregated in groups of 20–80 and immersed
in gall-like warts on the host squamules; asci 30–35 × 15 µm; spores without a distinct
ornamentation, 12–14.5 × 4.5–7 µm; Great Britain and Pyrénées .......................................................
.....................................................................................Polycoccum microcarpum Diederich & Etayo
– Perithecia larger, 0.1–0.25 mm in diam., not aggregated; asci longer than 50 µm;
spores verruculose............................................................................................................................21
21. Spores (7–)8.5–10.5(–12) × (3–)3.5–4.5(–5) µm, soleiform, colourless, then pale
to medium brownish–olive, verruculose; infected thallus not bullate; USA (N. Alaska) ....................
........................................................................................................................... Polycoccum laursenii
– Spores bigger, (12.5–)13.5–16.5(–22) × 6.5–8(–9) µm, oval with nearly equal
cells, brown, verruculose; infected thallus becomes bullate; Germany and Austria
................................................................................. Polycoccum cladoniae Diederich & D.Hawksw.
22. Associated with goniocysts up to 60 µm in diam.; spores fusiform to oval,
apically attenuated............................................................................................................................23
– Not associated with goniocysts; perithecia to 175 µm in diam.; spores only trans-
versely septate to submuriform, with 3–4 trans- or oblique septa and often 1(–2)
longisepta in central segments, pale to medium greyish-olive or brown, (8–)10–13
(–16) × (4–)5–6.5(–7.5) µm, smooth, oval to narrowly ellipsoidal, sometimes more
attenuated at one apex; Eurasia and Greenland............ Merismatium heterophractum (Nyl.) Vouaux
23. Perithecia to 250 µm in diam.; spores smooth, (1–)3-septate, rarely with additional
longiseptum in central segment, pale brown or grey-brown, (10.5–)13.5–16.5(–17.5)
× (3.5–)4–5.5(–6) µm; widespread................................... Merismatium decolorans (Arnold) Triebel
– Perithecia c. 100 µm in diam.; spores minutely rugose, 3-septate with the two median
cells brown and end-cells mostly colourless or rarely brown, 13–16.5 × 5.5–7 µm;
Norway ......................................................................................... Merismatium cladoniicola Alstrup
24. Perithecia associated with superficial dark brown mycelial network; spores
1-septate; interascal filaments absent ...............................................................................................25
– Superficial dark mycelial network absent; interascal filaments present...........................................26

Lichenicolous fungi on Arctic Cladonia 493
Symb. Bot. Ups. 34:1
25. Mycelial strands of single hyphae; perithecia ¾ immersed to superficial;
asci (20–)21–30(–36) × (11–)11.5–15(–16) µm; spores (7–)9–12.5(–14)
× (2.5–)3–4.5(–5) µm, colourless, rarely becoming coloured with age; wide-
spread .......................................................................... Sphaerellothecium araneosum var. cladoniae
– Mycelial strands of both single and a few parallel, agglutinated hyphae; perithecia
superficial; asci c. 20 × 10 µm; spores 8–12 × 5 µm, colourless; widespread .....................................
...................................................................... Sphaerellothecium cladoniicola E.S.Hansen & Alstrup
26. Spores simple, 9–12 × 4–7 µm; ascomata to 0.1 mm in diam.; hymenium I+ blue;
Central Europe................................................................... [Physalospora cladoniae (Stein) Vouaux]
– Spores septate...................................................................................................................................27
27. Spores 1-septate................................................................................................................................28
– Spores 3-septate to muriform ...........................................................................................................29
28. Spores (17.5–)18–22 × (6–)7–7.5(–8) µm; widespread .......................................................................
.................................................................. Zwackhiomyces dispersus (Körb.) Triebel & Grube s. lat.
– Spores 10–14 × 3–5 µm; widespread ................. Zwackhiomyces cladoniae (C.W.Dodge) Diederich
29. Spores with only transsepta..............................................................................................................30
– Spores muriform...............................................................................................................................32
30. Perithecia immersed, breaking through the host cortex in a stellate way, to 0.15
mm in diam.; interascal filaments little branched, 1.5 µm thick, apically to 2.5 µm,
sticking to the asci; spores 3-septate, 13–14 × 5–5.5 µm; Svalbard ....................................................
............................................................................................. Stellifraga cladonicola Alstrup & Olech
– Perithecia protruding in uppermost part, not breaking cortex in a stellate way;
interascal filaments not sticking to asci and not apically enlarged...................................................31
31. Spores (1–)3–4(–8)-septate, (12–)14–23(–27) × 4.5–6(–9) µm; peridium
aeruginose; Arctic and Europe .................................................................... Cercidospora punctillata
– Spores (1–)3-septate, (13–)16–20(–21) × (3–)4–6 µm; peridium brown;
Europe and Svalbard .................................................................................Cercidospora cladoniicola
32. Distinctly lichenized; perithecia immersed in conspicuous whitish thalline
warts (red in calcium hypochloride); spores with 8–10 trans- or oblique septa
and 1–2 longisepta in central segments, sometimes with apiculus at one or both
ends, 24–36 × 9–14 µm; occasionally on decaying lichens; widespread in the N.
Hemisphere................................................. Protothelenella leucothelia (Nyl.) H.Mayrhofer & Poelt
– Not lichenized or indistinctly lichenized..........................................................................................33
33. Not lichenized; spores with 3–7 transsepta and usually 1 longiseptum in central
segments, (12–)18–25(–28) × (6–)7.5–10(–12) µm, often with apiculus at one end;
arctic-alpine in the N. Hemisphere........................................................... Protothelenella santessonii
– Lichenized thallus membrane-like or absent; spores with 6–10 trans- or oblique
septa and 1 longisepta in central segments, (22–)24.5–29(–33) × 6.5–7.5(–10) µm,
withouth apiculus; occasionally on decaying lichens; arcto-boreal in the N. Hemisphere..................
........................................................................................................ Protothelenella sphinctrinoidella
34. Apothecial disc pale, whitish, yellow, orange or brownish; spores colourless ................................35
– Apothecia blackish; spores colourless or coloured ..........................................................................37
35. Apothecia with stipe up to 1.5 mm high terminating with flat to cupulate disc up to
1.5 mm in diam., pale orange; spores 1-septate, fusiform, 11–13 × 2–3 µm; Canada
(British Columbia)......................................................... Bryoscyphus lichenicola Alstrup & M. Cole
– Apothecia not stipitate; spores narowly cylindrical to vermiform...................................................36

494 Mikhail P. Zhurbenko and Vagn Alstrup
Symb. Bot. Ups. 34:1
36. Spores (1–)3-septate, 19–25(–31) × 2.5–3 µm; apothecia sessile, plane with
slightly raised paler margin, pale orange to orange-brown; widespread..............................................
.......................................................................................Lettauia cladoniicola D.Hawksw. & R.Sant.
– Spores simple, 30–36 × 2 µm; ascomata almost perithecioid, urceolate;
disc white; exciple thick and black; Austria and Sweden .................. Stictis cladoniae (Rehm) Sacc.
37. Ascomata on blackish stalk to 1 mm tall with lenticular capitulum to
0.3 mm in diam., closed at first, later opening; spores 1-septate, brown,
(6–)7–9(–10) × 2–3(–3.5) µm; Europe.....................Chaenothecopsis pusilla (Ach.) A.F.W.Schmidt
– Ascomata of different habitus ..........................................................................................................38
38. Ascomata without exciple, sometimes irregular ..............................................................................39
– Ascomata with exciple .....................................................................................................................48
39. Spores dark.......................................................................................................................................40
– Spores colourless; apothecia arthonioid...........................................................................................42
40. Hymenium divided into loculi by brown sterile tissue; apothecia up to
2.5 mm in diam., at first immersed then somewhat prominent, plane to
slightly convex; asci 4-spored; spores (4–)5-septate, 20–27 × 7–8 µm,
halonate; N. Europe....................................................................................... Plectocarpon cladoniae
– Hymenium not divided into loculi by brown sterile tissue; apothecia smaller,
to 0.3 mm in diam., sessile, convex, constricted at the base to sometimes shortly
stipitate; asci 8-spored; spores 1-septate, without halo....................................................................41
41. Spores (7–)6.5–8.5(–11) × 3–3.5(–4.5) µm, upper cell slightly thicker and longer
or not than the lower cell, medium to dark brown, distinctly verrucose, readily
breaking into two semi-spores along the septum; hymenium colourless, K+ bluish
green; apothecia 150–270 µm in diam.; USA and Europe...................................................................
......................................................................................Abrothallus cladoniae R.Sant. & D.Hawksw.
– Spores narrower, 6.5–9 × 2–3 µm, upper cell much thicker and longer than the
lower cell, pale brown, verruculose, hardly breaking into two semi-spores; hyme-
nium yellowish brown, K–; apothecia 100–220 µm in diam.; USA (Florida, Missouri).....................
...................................................................................Abrothallus pezizicola Diederich & R.C.Harris
42. Spores 3-septate, 10–15 × 3–5 µm; Italy..........[Arthonia lepidophila (Anzi) Clauzade et al. (ined.)]
– Spores 1-septate................................................................................................................................43
43. Hymenium K+ purple, I+ red; spores 12–15 × 4.5–5.5 µm; Colombia ...............................................
................................................................................................................. Arthonia colombiana Etayo
– Hymenium not purple in K...............................................................................................................44
44. Hymenium I–, K/I+ blue, olive-brown.............................................................................................45
– Hymenium I+ red or blue then red...................................................................................................46
45. Spores (10–)14–17.5(–20) × 5–5.5(–6) µm; widespread .................................. Arthonia epicladonia
– Spores (9–)10–11.5(–12) × (3–)3.5–4(–4.5) µm; Siberia ............................ Arthonia cf. epicladonia
46. Epihymenium blackish; spores 9–15 × 4–6 µm; widespread...............................................................
..................................................................................................Arthonia molendoi (Frauenf.) R.Sant.
– Epihymenium brown........................................................................................................................47
47. Asci 35–45 × 15–20 µm, 4–8-spored; spores 7–13 × 3–5 µm; widespread .........................................
........................................................................................................... Arthonia pelvetii (Hepp) Almq.
– Asci smaller, 20–25 × 12–15 µm, 8-spored; spores 9–11 × 3–4.5 µm;
Austria, Luxembourg..............................................................................Arthonia digitatae Hafellner
48. Spores dark brown............................................................................................................................49
– Spores colourless..............................................................................................................................52

Lichenicolous fungi on Arctic Cladonia 495
Symb. Bot. Ups. 34:1
49. Ascomata lirellate, with split-like opening; spores colourless, becoming brown
at maturity, 3-septate, 14–17 × 5–6 µm; Hawaii .............. Opegrapha cladoniicola Ertz & Diederich
– Ascomata rounded, constricted at base ............................................................................................50
50. Spores with thick walls and thick septum penetrated by narrow canal, 1-septate,
25–30 × 11–13 µm; Greenland..............................................................................Rinodina egedeana
– Spores of different type....................................................................................................................51
51. Spores 1-septate, 33–37 × 12–14 µm; Svalbard............ Dactylospora cladoniicola Alstrup & Olech
– Spores (3–)5–7-septate, (15–)17.5–21.5(–25) × (4.5–)5–6(–7) µm; widespread ................................
........................................................................................................................Dactylospora deminuta
52. Spores septate...................................................................................................................................53
– Spores simple ...................................................................................................................................54
53. Spores acicular, 12–?28-septate, 80–110(–120) × (2.5–)3–4(–5) µm, parallel
in ascus; apothecia sessile to substipitate, urceolate, to 4 mm in diam., with
prominent margin; exciple and epihymenium aeruginose; infested host parts con-
spicuously aeruginose; Europe, N. America ............. Arthrorhaphis aeruginosa R.Sant. & Tønsberg
– Spores 1–2-septate, 13–15(–16) × 5–6.5 µm; apothecia concave, becoming plane
to convex with age, strongly constricted at base, to 0.7 mm in diam., sessile;
Greenland, Canada (Northwest Territories) ................... Scutula cladoniicola Alstrup & D.Hawksw.
54. Ascomata with marginal hairs 35–45 µm long, urceolate, up to 0.25 mm in diam.,
sessile; spores ellipsoid, 8–10 × 2–3 µm; usually on sand covering lichen thalli;
Greenland ........................................................................... Skyttea arenicola Alstrup & E.S. Hansen
– Ascomata without marginal hairs, flat to convex .............................................................................55
55. Epithecium blue-black to dark green; hymenium pale bluish green; hypothecium
purplish brown; spores c. 10 × 4 µm; ascomata immersed at first; causes dis-
figuration; W. Europe ................................................................................[Lecidea cladoniaria Nyl.]
– Epithecium and hypothecium dark brown; hymenium castaneous; spores
narrowly ellipsoid to fusiform, 6–12 × 2–4 µm; ascomata sessile, becoming
convex, to 0.3 mm in diam.; widespread..............................................................................................
......................................................... Phaeopyxis punctum (A.Massal.) Rambold, Triebel & Coppins
56. Conidiomata stromatic, subglobose, to 75 µm in diam., black, superficial,
forming (dark) brown, smooth, irregularly globose, multicellular conidia
c. 10–15 µm in diam., composed of 10–15(–20) subglobose individual cells
3–4 µm in diam.; widespread ......................................................................Phaeosporobolus alpinus
– Conidiomata not distinctly stromatic; conidia not as above.............................................................57
57. Conidia formed in pycnidia..............................................................................................................58
– Conidia not enclosed in pycnidia .....................................................................................................72
58. Conidia branched, colourless ...........................................................................................................59
– Conidia unbranched..........................................................................................................................61
59. Conidia with four X-shaped, cylindrical, 1–3-septate arms, each 18–32 µm
long and 1–2.5 µm in diam.; pycnidia dark brown, subglobose, ostiolate,
immersed; Central Europe............... Cladoniicola staurospora Diederich, Van den Boom & Aptroot
– Conidia unicellular, non-septate, Y-shaped with three arms; pycnidia white to brownish...............60
60. Arms of conidia c. 2–3 µm broad, without filiform appendages at the apices;
widespread.............................................................Cornutispora lichenicola D.Hawksw. & B.Sutton
– Arms of conidia 3–4 µm broad, with filiform appendages at the apices;
widespread.................................................................................................. Cornutispora ciliata Kalb

496 Mikhail P. Zhurbenko and Vagn Alstrup
Symb. Bot. Ups. 34:1
61. Conidia colourless............................................................................................................................62
– Conidia brown, widespread species .................................................................................................69
62. Pycnidia usually associated with galls .............................................................................................63
– Pycnidia usually not associated with galls .......................................................................................66
63. Two types of conidia produced within the same pycnidium, macroconidia irregularly
ellipsoidal to subglobose, of (1–)2–4(–5)cells, 8–15 × 4.5–8 µm, and microconidia
subcylindrical, simple, 3.5–4.5 × 1–1.4 µm; pycnidia subsphaerical, without ostiole,
dark brown to black, 50–100 µm in diam., up to 30 immersed in a gall to 1.2 mm in
diam.; Belgium............................................................Stromatopogon cladoniae Diederich & Sérus.
– Pycnidia with only one type of conidia; wall (sub)colourless becoming orange to
brown only in exposed parts; sometimes not associated with galls ................................................64
64. Conidia lens-shaped to pyriform, apically attenuated, often more markedly so and
also truncated at the base, simple, (7–)8–10(–11) × (3–)4–5.5(–6) µm; pycnidia
completely immersed in galls, to 200 µm in diam.; widespread.........Bachmanniomyces uncialicola
– Conidia subcylindrical to narrowly ellipsoid, truncated at the base by a scar;
pycnidia immersed or superficial .....................................................................................................65
65. Conidia simple, (8–)8.5–10.5(–12) × 3–4 µm, mature pycnidia sessile on galls,
to 175 µm in diam.; widespread ..........................................................................Epicladonia simplex
– Conidia (0–)1-septate, (7–)9–12(–14) × (2.5–)3–4 µm; pycnidia immersed
in galls, to 125 µm in diam.; widespread ....................................................... Epicladonia sandstedei
66. Conidia 3-septate, 18–22 × 3.5–5 µm, broadly fusiform, with the base often
more attenuated and weakly curved; Czech Republic..........................................................................
......................................................................... Keissleriomyces sandstedeanus (Keissl.) D.Hawksw.
– Conidia simple or 1-septate..............................................................................................................67
67. Conidia with apices usually rounded or occasionally slightly attenuated, oblong to
occasionally slightly reniform, simple, 4–5.5(–6) × (1.5–)2 µm; pycnidia black,
globose, 50–70 µm in diam., immersed to sessile; Svalbard ..............................................Phoma sp.
– Conidia often apically attenuated or truncated.................................................................................68
68. Conidia born only at the apices of conidiogenous cells, simple or exceptionally
1-septate, subcylindrical to cymbiform, not curved, narrowed and abruptly trun-
cated at the base by a scar, 7.5–11 × 3–3.5 µm; pycnidia immersed, possibly
sometimes induce galls; Eurasia ................................................................... Epicladonia stenospora
– Conidia born both apically and laterally, simple, lacriform, often slightly
curved, distinctly attenuated and pointed at the base, (6–)6.5–10(–11.5) ×
(2–)3–4.5(–6) µm; pycnidia often semi-immersed; widespread ...............Lichenosticta alcicornaria
69. Pycnidia (80–)100–175(–200) µm in diam., semi-immersed; conidiogenous
cells (5–)6–8(–11) × (2–)2.5–4 µm; conidia (sub)globose, truncated at the
base, almost smooth-walled, (2.5–)3–4.5(–6) µm in diam.; widespread .............................................
............................................................................................... Lichenoconium xanthoriae M.S.Christ.
– Pycnidia smaller, up to 110 µm in diam...........................................................................................70
70. Pycnidia (20–)30–50(–60) µm in diam., immersed; conidiogenous cells
(3.5–)4–5(–6) × (2–)3–3.5(–4) µm, conidia subglobose, not distinctly truncated
at the base, distinctly verruculose, 2–3.5(–4) µm in diam.; strongly
pathogenic, induces necrotic patches on the thalli; widespread...........................................................
............................................................................ Lichenoconium erodens M.S.Christ. & D.Hawksw.
– Pycnidia usually wider than 50 µm; conidiogenous cells longer; less pathogenic species .............71

Lichenicolous fungi on Arctic Cladonia 497
Symb. Bot. Ups. 34:1
71. Conidia subglobose to ovoid, truncated at the base, almost smooth-walled,
(2–)2.5–3.5(–4) × 2–3 µm; pycnidia mainly superficial, 50–80(–100) µm in diam.,
conidiogenous cells (5–)6–9(–11) × 1.5–2.5(–3) µm; N. Hemisphere ...... Lichenoconium pyxidatae
– Conidia subglobose, not distinctly truncated at the base, distinctly verruculose,
3–4(–5) µm in diam.; pycnidia immersed to superficial, (40–)50–80(–110)
µm in diam.; conidiogenous cells (5–)7–9(–11) × (2–)2.5–3.5 µm; widespread .................................
..........................................................................................Lichenoconium usneae (Anzi) D.Hawksw.
72. Conidia formed in sporodochia........................................................................................................73
– Conidia formed directly from mycelium..........................................................................................74
73. Sporodochia blackish, convex, to 120 µm in diam.; conidia unevenly brown,
verruculose, variable in form, more or less lobed, simple, 7–14(–16) × 6–10(–12)
µm; in very shaded Fagus woods in the Pyrénées..........Milospium lacoizquetae Etayo & Diederich
– Sporodochia rose, red or orange, to 200 µm in diam.; conidia 11–15 × 7–10 µm;
widespread................................... Marchandiomyces corallinus (Roberge) Diederich & D.Hawksw.
74. Colonies inconspicuous, without any superficial mycelium, of pale brown,
up to 50 µm long, 1–3-septate, occasionally branched phialophores ending in
a lageniform phialide; conidia colourless, simple, cylindrical with truncate ends,
2.5–4 × 0.5–0.8 µm; Svalbard ..........................................................Chalara lichenicola M.S.Christ.
– Colonies conspicuous; conidiophores and conidia dark brown ........................ .............................75
75. Mycelium c. 2 µm thick, internal in host hyphae; conidiophores short; conidia
usually simple and 7–8 × 3.5–4 µm, or rarely 1-septate and 11–14 × 3.5–4.5 µm,
smooth-walled; mature colonies give the host surface a grey-brown velvety ap-
pearance; on Cladonia subgen. Cladina; Denmark and Czech Republic ............................................
............................................................................................................Taeniolella cladniicola Alstrup
– Mycelium thicker, superficial or immersed, but not internal in host hyphae ...................................76
76. Mycelial hyphae 2–3.5 µm thick; conidiophores 40–50 × 3–5.5 µm, with rough wall;
conidia solitary or occasionally catenate by 2, 0–1-septate, weakly verruculose,
(7–)7.5–9–10.5(–13) × 3.5–4–4.5(–5) µm; colonies exhibit conspicuous blackish
bristles; widespread...........................................................................................Taeniolella beschiana
– Mycelium of immersed colourless hyphae 4–5 µm thick and superficial, dark
brown, rough-walled, branched hyphae to 10 µm thick; conidiophores 25–50 ×
5–7 µm, smooth-walled; conidia formed in chains, 1–2-septate, smooth-walled,
8.5–12 × 5.5–6 µm; Greenland..............................................................................Taeniolella strictae
Acknowledgements
MZ is grateful to Prof. Rolf Santesson, who first
suggested him that Sphaerellothecium araneosum
on Cladonia pocillum might represent an unde-
scribed taxon. Dr. Helmut Mayrhofer is thanked
for comments on Protothelenella taxonomy. MZ is
greatly indebted to Dr. Lev Biazrov, Dr. Sergei
Kholod, Dr. Irina Makarova, Dr. Adrian Katenin,
Dr. Nadya Matveeva, Prof. Fred Daniels, Dr. Don-
ald A. Walker, Dr. Lida Zanokha, Prof. Vladimir
Kuvaev, and the others who put their lichen collec-
tions at his disposal. The field studies of MZ dur-
ing the past 12 years, which contributed to this
study, have been supported by many organisations
and funds, the recent ones being grants from the
National Park Service (Nos. PX9830-93-062,
PX9830-92-385, PX9830-0-0451, PX9830-0-
0472, and PX9830-0-0512) made to Gary Laursen
of the University of Alaska Fairbanks (UAF) and
the “Spitsbergen-2003” expedition of the Arctic
and Antarctic Research Institute (St.-Petersburg,
Russia). The study of MZ has been also financially
supported by the Russian Fund for Basic Re-
searches (RFBR grant no 03-04-49400) and basic
research programme of the Russian Academy of

498 Mikhail P. Zhurbenko and Vagn Alstrup
Symb. Bot. Ups. 34:1
Sciences “Biodiversity of Russia”. We are also in-
debted to Dr. Rod Seppelt for improving the Eng-
lish.
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