Lichenicolous fungi growing on Thamnolia, mainly from the Holarctic, with a worldwide key to the known species

Abstract

Twenty species of lichenicolous fungi are reported on Thamnolia species and discussed. Epithamnolia karatyginii gen. et sp. nov., Capronia thamnoliae sp. nov., Cercidospora epithamnolia sp. nov., C. thamnogalloides sp. nov., C. thamnoliae sp. nov., and Sphaerellothecium thamnoliae sp. nov. (var. thamnoliae, var. taimyricum) are described from Thamnolia. Dacampia thamnoliicola and Phoma thamnoliae are introduced ad interim. Cercidospora lecidomae is reduced to synonymy with C. punctillata. Polycoccum vermicularium is new to Asia, Odontotrema santessonii and O. thamnoliae are new to North America, Cladosporium licheniphilum is new to the Arctic, Thamnogalla crombiei is new to Greenland and Svalbard, Stigmidium frigidum is new to Mongolia and confirmed in the USA, Lichenopeltella thamnoliae is new to Bolivia. Cladosporium licheniphilum and Phaeospora arctica are newly documented on Thamnolia and Lichenopeltella thamnoliae on Thamnolia papelillo var. subsolida. Thamnolia vermicularis, supporting 23 species of lichenicolous fungi, is shown to be the 15th most hospitable lichen species in the world. A worldwide key to 23 species of fungi known to occur on Thamnolia is provided.

Full text

Lichenicolous fungi growing on Thamnolia, mainly from the
Holarctic, with a worldwide key to the known species
Mikhail P. ZHURBENKO
Abstract: Twenty species of lichenicolous fungi are reported on Thamnolia species and discussed.
Epithamnolia karatyginii gen. et sp. nov., Capronia thamnoliae sp. nov., Cercidospora epithamnolia sp.
nov., C. thamnogalloides sp. nov., C. thamnoliae sp. nov., and Sphaerellothecium thamnoliae sp. nov. (var.
thamnoliae, var. taimyricum) are described from Thamnolia.Dacampia thamnoliicola and Phoma
thamnoliae are introduced ad interim.Cercidospora lecidomae is reduced to synonymy with C. punctillata.
Polycoccum vermicularium is new to Asia, Odontotrema santessonii and O. thamnoliae are new to North
America, Cladosporium licheniphilum is new to the Arctic, Thamnogalla crombiei is new to Greenland and
Svalbard, Stigmidium frigidum is new to Mongolia and confirmed in the USA, Lichenopeltella thamnoliae
is new to Bolivia. Cladosporium licheniphilum and Phaeospora arctica are newly documented on
Thamnolia and Lichenopeltella thamnoliae on Thamnolia papelillo var. subsolida.Thamnolia vermicularis,
supporting 23 species of lichenicolous fungi, is shown to be the 15th most hospitable lichen species in
the world. A worldwide key to 23 species of fungi known to occur on Thamnolia is provided.
Key words: lichen, new records, new taxa, thamnoliicolous fungi
Accepted for publication 27 August 2011
Introduction
Within the last eight years, the number of
known lichenicolous fungal species has risen
by about one quarter, roughly from 1300 to
1700 [Lawrey & Diederich 2003; author’s
database, derived from Culberson et al.
(2011)], which corresponds to a long-
established opinion that the diversity of this
hostal group of fungi is yet to be revealed.
However, specific estimates of their species
richness vary from 2000 (Hawksworth 1991)
to 5000–7000 (Lawrey & Diederich 2003).
Taking into consideration that 1) the value of
the Lichenicolous Index (ratio of species of
lichenicolous fungi per species of lichens
within an area) for the best known myco-
biotas of the world is about 0·2 (Zhurbenko
2007), 2) the known and estimated numbers
of lichen species in the world vary from
17 500 to 28 000 (Feuerer & Hawksworth
2007; Lücking 2008), 3500 to 5500 species
of lichenicolous fungi could be expected
worldwide, which means that approximately
30–50% of them are presently described.
This corresponds with the results of studies
of fungi inhabiting particular lichen genera,
such as Pilophorus,Stereocaulon,Tephromela
or Thamnolia, where about 15–35% of
the species occurring are new to science
(Zhurbenko & Triebel 2005; Hafellner 2007;
Zhurbenko 2010a; present study). Such
studies are also optimal to show various
aspects of ‘host-parasite’ relationships, for
instance, how many fungi can invade one
lichen species. The world record in that
respect seems to belong to Peltigera rufescens
(Weiss) Humb., supporting 54 species of
lichenicolous fungi (author’s database).
The oroarctic lichen genus Thamnolia con-
sists of three species: a widely distributed,
cosmopolitan T. vermicularis (Sw.) Schaer.,
T. papelillo R. Sant., known from South
America, and T. juncea R. Sant., known from
Papua New Guinea (Santesson 2004). Until
M. P. Zhurbenko: Laboratory of the Systematics
and Geography of Fungi, Komarov Botanical
Institute, Russian Academy of Sciences, Professor
Popov 2, St.-Petersburg, 197376, Russia. Email:
zhurb58@gmail.com
The Lichenologist 44(2): 147–177 (2012) © British Lichen Society, 2012
doi:10.1017/S0024282911000739

now, 13 species of lichenicolous fungi were
known to occur on Thamnolia, four of which
were specifically treated in Norway (Ihlen
1995). The results of the revision of fungi
found on Thamnolia vermicularis within the
Holarctic (with one find on T. papelillo from
the Neotropics) are presented here. They
nearly double the number of thamnoliicolous
fungi species now reaching 23, including
eight (35%) described herewith. A new an-
amorphic lichenicolous fungal genus and a
synonym within Cercidospora are introduced.
Since no key to fungi growing on Thamnolia
has been published previously, one is pro-
vided here, along with notes on their
taxonomy, hosts, and geography.
Materials and Methods
The study is based on 190 samples of lichenicolous fungi
on Thamnolia vermicularis, obtained from no less than
1500–2000 host specimens examined from the Holarc-
tic, and one sample on T. papelillo from South America.
Furthermore, 37 LE herbarium specimens were revised.
The material was examined and photographed using
Zeiss microscopes Stemi 2000–CS and Axio Imager A1
equipped with Nomarski differential interference con-
trast optics. Microscopical examination was carried out
on material mounted in water, 10% KOH (K), Lugol’s
iodine, directly (I) or after a KOH pre-treatment (K/I),
or Brilliant Cresyl blue (BCr). The length, breadth, and
length/breadth ratio (l/b) of asci and ascospores are
given (where possible) as: (min−){x−SD}−{x+SD}
(−max), where min and max are the extreme values,x
the arithmetic mean, and SD the corresponding stan-
dard deviation. Measurements of the ascus sizes were
rounded to the nearest 1 m, those of the ascospores and
conidia to the nearest 0·5 m. The Ascomycota classifi-
cation follows Lumbsch & Huhndorf (2007). Terms for
simple plane shapes, used for describing asci and asco-
spores, mostly follow Stearn (1992: 539). The estima-
tion of total diversity of thamnoliicolous fungi in the
Holarctic was based on a simple estimator proposed by
Turing (Chao & Shen 2003), based on the proportion
of single occurrences (singletons) in the sample
&I
6
=− × , where fis the number of singletons and
Sis the number of known species. All examined speci-
mens are housed in the mycological herbarium of the V.
L. Komarov Botanical Institute in St. Petersburg (LE).
The following specimens have been examined and
compared with the species studied: Cercidospora lecido-
mae Zhurb. & Triebel: LE 207636 (paratype), LE
260308, LE 260555. Cercidospora parva Hafellner &
Ihlen: LE 260155, LE 260054, LE 233214. Cercidospora
punctillata (Nyl.) R. Sant.: LE 206999, LE 207003,
LE 207007, LE 207012, LE 207779, LE 233253, LE
260095, LE 260166, LE 260177, LE 260245, LE
260276, LE 260296.
The Taxa
Capronia thamnoliae Zhurb. sp. nov.
MycoBank no.: MB 563049
Fungus lichenicola in thallis lichenum generis Thamno-
lia crescens. Similis speciei Capronia normandinae, sed
differt praecipue ascosporis minoribus, (11–)13–18
(–29) × (5–)6–8(−8·5) m, et hospite diverso.
Typus: USA, Alaska, Great Kobuk Sand Dunes,
67°05#N, 159°00#W, 45 m elev., Dryas-lichen-moss
vegetation among sparse Picea glauca, on moribund
bases of Thamnolia vermicularis var. subuliformis thalli,
13 August 2000, M. Zhurbenko 003 (LE 260400—
holotypus).
(Fig. 1)
Vegetative hyphae pale brown, cylindrical
to torulose, 2–3 m wide, septate, with cells
4–15 m long, scarcely branched, origin-
ating from the base of ascomata, mostly
superficial.
Ascomata perithecioid, black, subglobose
to occasionally pyriform, (70–)90–160 m
diam., with an ostiole c.15m diam., usually
densely setose throughout, semi-immersed
to mostly sessile, arising singly, dispersed.
Setae dark brown, subulate, the tips rounded,
15–75 m tall, 2–3 m wide, base swollen
6–9 m wide, thick-walled, the edges smooth
or occasionally crenate, non-septate or rarely
with 1(–2 ?) septa, unbranched. Exciple
medium brown, in surface view composed of
angular pseudoparenchymatous cells (textura
angularis) 4–13 m across, K+ acquiring a
vague grey shade. Hymenial gel I+ coral to
rose, K/I+ blue. Hamathecium not observed.
Asci clavate to subcylindrical, with strongly
thickened apical wall, often penetrated by an
internal apical beak (best developed in im-
mature asci), (56–)60–72(–76) × (13–)14–
18(–21) m(n= 29), 8-spored, I−, K/I−.
Ascospores initially hyaline, then pale to me-
dium grey (septa sometimes with a brown
shade), mostly elliptic to narrowly elliptic,
occasionally very narrowly elliptic, broadly
elliptic, rotund, narrowly ovate, or narrowly
obovate, the apices attenuated or occasion-
ally rounded, (11–)13·5–18·0(–29) × (5–)6–
148 THE LICHENOLOGIST Vol. 44

8(–12) m, l/b = (1·3–)1·8–2·8(–5·8) (n=
154), mostly submuriform, with (0–)3–5(–7)
transverse septa and as a rule with longitudi-
nal or oblique septa in 1–3(–4) segments
(very rare, probably when immature, non-
septate), not or rarely constricted at the septa
(when overmature), smooth-walled, non-
halonate, often guttulate, arranged biseriate
to partly overlapping and uniseriate in the
ascus.
Anamorph not observed.
Notes. Thirteen of c. 60 previously known
species of Capronia Sacc. (Herpotrichiellaceae,
Chaetothyriales) grow on lichens (Untereiner
2000; Lawrey & Diederich 2011) [Capronia
triseptata (Diederich) Etayo does not belong
to Capronia (P. Diederich, pers. comm.,
2011)], all being exclusively lichenicolous.
Nearly all of them are restricted to a particu-
lar lichen host genus [with the exception of
Capronia hypotrachynae Etayo & Diederich
known on Hypotrachyna and Menegazzia
(Etayo & Diederich 1998; von Brackel
2010b) and perhaps C. epilobarina S. Y.
Kondr. & D. J. Galloway, which was de-
scribed from Lobaria and was reported (with
‘cf.’) also on Erioderma (Etayo 2002)]; none
has been found on Thamnolia yet. Ascospore
colour, size, and septation are among the
most reliable morphological characters for
the species delimitation within the genus.
Seven lichenicolous Capronia species also
have submuriform (or muriform) as-
cospores, characteristic of Capronia thamno-
liae. Of these, Capronia epilobarina differs
from the latter in its greyish brown, narrower
ascospores up to 7 m wide (Kondratyuk &
Galloway 1995; Etayo & Diederich 1996b);
C. guatemalense Etayo & van den Boom has
larger ascospores (22–27 × 9·0–11·5 m),
which are brownish from the beginning and
have more septa (Etayo & van den Boom
2006); C. hypotrachynae has slightly narrower
ascospores (12–19 × 5·5–7·5 m) and asep-
tate and much shorter setae, measuring
5–35 × 3–4 m, often less than 10 m long
(Etayo & Diederich 1998); C. leopoldiana
Etayo has longer and narrower ascospores
[(15–)17–22(–24) × 4–5·5 m], shorter
setae (15–40 × 3–4 m), and smaller
ascomata (70–100 m diam.) (Etayo &
Sancho 2008); C. magellanica Etayo differs in
its shorter and narrower ascospores [(11–)
13–16(–17·5) × 4–6 m], which only occa-
sionally have 1 longitudinal septum in some
cells, shorter setae (20–45 × 4–5 m), and
the I−, K/I− reaction of the hymenial gel
F. 1.Capronia thamnoliae. A, ascoma (holotype); B, asci with spores in K/I (LE 260410); C, setae in K/I (holotype).
Scales:A=100m;B&C=10m.
2012 Holarctic thamnoliicolous fungi—Zhurbenko 149

(Etayo & Sancho 2008); C. normandinae
R. Sant. & D. Hawksw. has pale olivaceous
brown, slightly larger ascospores [(13–)15–
21(–27) × 7·5–9·0 m], tending to appear
densely dictyosporous due to guttules and
putative distosepta, and non-septate setae,
mostly developing around the ostiole (Hawk-
sworth 1990); C. pseudonormandinae Dieder-
ich has pale brown, smaller and less septate
ascospores (12·5–16 × 6–7·5 m), and non-
septate, shorter setae, measuring 10–
30 × 2·5–4 m (Aptroot et al. 1997).
Distribution and host. Circumpolarly
known in the Holarctic from arctic and al-
pine tundras and northern boreal forests.
Saprotrophic or commensalistic on old or
moribund portions of Thamnolia vermicularis
thalli.
Additional specimens examined [all on Thamnolia ver-
micularis var. subuliformis (Ehrh.) Schaer.]. Canada:
Canadian Arctic Archipelago: Axel Heiberg Island, Bunde
Fjord, 80°30#N, 94°36#W, 30 m, 1999, N. Matveeva
(LE 260410).—Norway: Troms County: Skibotndalen
Valley, 69°15#N, 20°23#E, 950 m, 2003, M. Zhurbenko
(LE 260458).—Russia: Taimyr Peninsula: Byrranga
Mts., Levinson-Lessinga Lake, 74°30–33#N, 98°26–
33#E, 120–300 m, 1994, M. Zhurbenko (LE 207477c);
ibid., 1995, M. Zhurbenko (LE 260390, LE 260598).
Cercidospora epithamnolia Zhurb. sp.
nov.
MycoBank no.: MB 563050
Fungus lichenicola in thallis lichenum generis Thamno-
lia crescens. Similis speciei Cercidospora solearispora, sed
differt praecipue ascosporis minoribus, (12–)13–16
(–19) × (3–)3·5–4·5(–5) m, et hospite diverso.
Typus: Russia, Severnaya Zemlya, Bolshevik Island,
W coast of Mikoyana Bay, 79°18#N, 101°55#E, 10 m
elev., polar desert, on thalli of Thamnolia vermicularis
var. subuliformis, 21 July 1996, M. Zhurbenko 96908 (LE
232954—holotypus).
(Fig. 2A, B,D&F)
Vegetative hyphae indistinct.
Ascomata perithecioid, black, often with a
greenish or bluish grey shade when slightly
shielded by host tissues, smooth, subglobose,
90–120 m diam., with an ostiole of 10–
25 m diam., immersed to slightly protrud-
ing in the ostiolar area, arising singly or
occasionally contiguous, aggregated into
groups of up to 100 or more. Exciple dark
bluish green with a grey shade above, hyaline
below, in surface view resembling textura an-
gularis (cells mostly 3–5 m across), K−.
Hymenial gel I−, K/I−. Hamathecium of
sparse or sometimes indistinct paraphysoids,
non-capitate, c. 1·5 m wide, septate,
scarcely branched. Asci subcylindrical or
moderately thickened in lower part, wall 1–2
m thick apically and 0·5–1·5 m laterally,
sometimes with a small ocular chamber, with
an indistinct or short stalk, (40–)44–60(–
63) × 10–13(–14) m(n= 12; in K elon-
gated up to 76 m), 8-spored, I−, K/I−.
Ascospores hyaline, strongly heteropolar, nar-
rowly soleiform or skittle-shaped with nar-
rower to attenuated lower part and rounded
ends, (12–)13–16(–19) × (3–)3·5–4·5(–5)
m, l/b = (2·9–)3·1–4·1(–5·0) (n= 40), with
(0–)1 central septum, constricted at the sep-
tum only in K, smooth-walled, halo not ob-
served, sometimes with a few conspicuous
guttules per cell, arranged irregularly biseri-
ate in the ascus.
Anamorph not observed.
Notes. Sixteen of 27 Cercidospora Körb.
species previously described have exclusively
or mainly 1-septate ascospores. Of these, the
following five species are most similar to
C. epithamnolia by colour of exciple and
mainly 8-spored asci: C. caudata Kernst.
s. lat. (grows on Caloplaca and Xanthopeltis),
C. epipolytropa (Mudd) Arnold (on Lecanora),
C. parva (on Baeomyces), C. solearispora
Calat., Nav.-Ros. & Hafellner (on Aspicilia
and Circinaria) and C. verrucosaria (Linds.)
Arnold (on Megaspora) (Ihlen 1998; Navarro-
Rosinés et al. 2004, 2009; specimens of
Cercidospora parva were examined for com-
parison). Cercidospora caudata is dis-
tinguished from C. epithamnolia by its larger
ascospores
[(14–)16–24(–30) × (3–)4–6(–7)
m], which
are usually strongly heteropolar,
with the lower cells curved and attenuated as
a tail; C. epipolytropa has larger ascospores
[(14–)15–19(–22) × (4·5–)5–6(–7) m]; C.
parva has a more green exciple, 4–8-spored
asci and is a weak parasite; C. solearispora
differs in its larger ascospores [(15–)17–21(–
22) × (4·5–)5–6(–7) m] with much shorter
150 THE LICHENOLOGIST Vol. 44

and narrower lower cells; C. verrucosaria has
a more brown exciple and larger ascospores,
measuring (14·5–)15·5–19·5(–23) × (4·5–)
5–6(–7) m. Moreover, all of them grow on
different host genera (most Cercidospora
species are restricted to a particular host
genus) and have better developed paraphys-
oids. Comparison with C. thamnogalloides
Zhurb., which also has 1-septate ascospores,
is provided in Table 1. Though scarce inter-
ascal filaments are not typical for Cercido-
spora, they have been reported for C.
cladoniicola Alstrup, C. macrospora (Uloth)
Hafellner & Nav.-Ros. (type species of the
genus) and C. thamnoliicola Ihlen (Ihlen
1995; Alstrup 1997; Navarro-Rosinés et al.
2004).
Distribution and host. Known from polar
desert and arctic tundra of Asia. Commen-
salistically growing throughout healthy-
looking thalli of Thamnolia vermicularis.
Additional specimens examined (all on Thamnolia ver-
micularis var. subuliformis). Russia: Severnaya Zemlya:
Bolshevik Island, Akhmatova Bay, 79°04#N, 102°45#E,
10 m, 1996, M. Zhurbenko (LE 260574); ibid.,
Shokalskogo Strait, 79°18#N, 101°40#E, 20 m, 1996,
M. Zhurbenko (LE 233044). Taimyr Peninsula: Byrranga
Mts., Levinson-Lessinga Lake, 74°33#N, 98°26#E,
100 m, 1995, M. Zhurbenko (LE 260490). Wrangel
Island: Naskhok River, 71°25#N, 178°20#W, 10 m,
1998, S. Kholod (LE 260590, LE 260530).
F. 2.Cercidospora epithamnolia A, B, D & F; A, infection habitus (LE 260574); B, ascomata (LE 233044) D, ascus
in water (LE 260590); F, ascospores in K (LE 260590). Cercidospora thamnogalloides (holotype)C,E&G;C,
infection habitus, note gall; E, asci in I; G, ascospore in water, note halo. Scales:A=1mm;B=200m; C =
500 m; D−G = 10 m.
2012 Holarctic thamnoliicolous fungi—Zhurbenko 151

Cercidospora thamnogalloides Zhurb.
sp. nov.
MycoBank no.: MB 563051
Fungus lichenicola in thallis lichenum generis
Thamnolia crescens. Similis speciei Cercidospora
galligena, sed differt praecipue ascomatibus minoribus,
80–120 m diam., ascosporis angustioribus, (12–)14–
17(–19) × (3·5–)4–5(–5·5) m, et hospite diverso.
Typus: Russia, Komi Republic, Northern Ural,
Pechora-Ilych State Reserve, headwaters of Pechora
River, Yanypupuner Range, Mt. “981”, 62°05#N,
59°06#E, 800 m elev., dwarf shrub alpine tundra, on
thalli of Thamnolia vermicularis var. subuliformis,30June
1997, M. Zhurbenko 97394 (LE 260510—holotypus).
(Fig. 2C,E&G)
Vegetative hyphae indistinct.
Ascomata perithecioid, black, smooth, sub-
globose, 80–120 m diam., with an ostiole
mostly of 10–20 m diam., immersed to
slightly protruding in the ostiolar area, aris-
ing singly, aggregated in groups of up to 100
or more. Exciple dark bluish grey and 15–
20 m thick above, hyaline and 10–15 m
thick below, in surface view resembling
textura angularis (cells 5–12 m across), K−.
Hymenial gel I−, K/I−. Hamathecium of
sparse or sometimes indistinct paraphysoids,
non-capitate, 1·5–2·0 m wide, septate,
scarcely branched. Asci obclavate to narrowly
elliptic, occasionally subcylindrical, apical
wall strongly thickened only in immature
asci, rarely with an indistinct ocular cham-
ber, with an indistinct or short stalk, (41–)
46–60(–72) × (10–)12–16(–18) m(n= 34;
in K elongated up to 85 m), 8-spored, I−,
K/I−. Ascospores hyaline, distinctly hetero-
polar, narrowly soleiform, skittle-shaped, or
narrowly oblanceolate, lower cell narrower
than the upper one, usually with rounded ends,
(12–)14–17(–19) × (3·5–)4–5(–5·5) m, l/b
= (2·7–)3·0–3·8(–4·1) (n= 57), with 1 cen-
tral septum, not constricted at the septum,
smooth-walled, with distinct halo 1·5–2·0
m thick, usually with a few conspicuous
guttules per cell best seen in K, arranged
biseriate to partly diagonally uniseriate in the
ascus.
Anamorph not observed.
Etymology. The specific epithet reflects
external similarity of the new species to
Thamnogalla crombiei (Mudd) D. Hawksw.
Notes. Just two previously known species of
Cercidospora regularly induce galls: C. cecidi-
iformans Grube & Hafellner (on Rhizocarpon)
and C. galligena Hafellner & Nav.-Ros. (on
Aspicilia and Circinaria). However, occasional
production of cecidia was also observed in C.
macrospora [on the Lecanora muralis (Schreb.)
Rabenh. group], C. solearispora, and C. stereo-
caulorum (Arnold) Hafellner (on Stereocaulon)
(Hafellner 1993; Navarro-Rosinés et al.
2004, 2009; Zhurbenko 2010a). Cercidospora
cecidiiformans differs from C. thamnogalloides
in its larger ascomata (130–150 m diam.),
T 1.Distinguishing characteristics of Cercidospora species growing on Thamnolia (based on the author’s own data).
Character C. epithamnolia C. thamnogalloides C. thamnoliae C. thamnoliicola
Ascomata aggregated aggregated scattered aggregated
Paraphysoids sparse to indistinct sparse to indistinct abundant sparse to indistinct
Exciple bluish green bluish grey bluish green olive-brown
Spores per ascus 8 8 (4–)8 (2–)4(–6)
Ascospores
length (m) (12–)13–16(–19) (12–)14–17(–19) (12·5–)16·5–21·0 (–27·5) (11–)14·0–17·5(–21)
breadth (m) (3–)3·5–4·5(–5) (3·5–)4–5(–5·5) (4–)4·5–5·5(–8) (4·0–)4·5–5·5(–6·5)
l/b ratio (2·9–)3·1–4·1(–5·0) (2·7–)3·0–3·8(–4·1) (2·5–)3·2–4·2(–5·4) (2·2–)2·6–3·4(–4·2)
septa (0–)1 1 (1–)3(–4) (1–)2–3
halo absent always present occasionally present absent or indistinct
Pathogenicity commensal
gall-inducer, mild
pathogen
mild pathogen or
saprotroph mild pathogen
152 THE LICHENOLOGIST Vol. 44

olive-brown exciple, wider ascospores (13–
19 × 5–8 m), and the I+ red, K/I+ blue
reaction of the hymenial gel; C. galligena has
larger ascomata (mainly 120–190 m diam.),
a brown to violet-black exciple, (4–)6–8-
spored asci, and occasionally simple, some-
what wider ascospores [(13·5–)14–19 × 5–
6(–7) m], being only slightly heteropolar; C.
macrospora has larger ascomata (150–250 m
diam.), 4(–8)-spored asci, and longer, rarely
slightly heteropolar, ascospores [(19–)20–
25(–30) × 4–6(–7) m]; C. solearispora dif-
fers in its larger ascomata (160–230 m
diam.) and larger ascospores [(15–)17–21
(–22) × (4·5–)5·0–6·0(–7·0) m] with a very
different shape and size of lower and upper
cells; C. stereocaulorum has larger, sometimes
superficial ascomata [100–200(–300) m
diam.], (2–)4(–8)-spored asci, and (1–)3
(–6)-septate, larger ascospores, measur-
ing (13–)18·5–25·5(–30) × (4–)5–7(–8) m.
Moreover, all these species grow on different
hosts and all, except C. macrospora, have
abundant paraphysoids. Galls induced by the
new species strongly resemble those caused
by Thamnogalla crombiei. Curiously, the latter
species was also found in the type collection.
Distribution and host. Known from alpine
tundra in the type locality in the Northern
Ural at the boundary between Europe and
Asia. The species induces gall-like swellings
on healthy-looking thalli of Thamnolia ver-
micularis. The swellings are concolorous with
the host thalli or pale brownish vin-
aceous. Host thalli are sometimes contorted
under infection.
Additional specimen examined.Russia: Northern Ural:
headwaters of Pechora River, Yanypupuner Range,
62°05#N, 59°06#E, 800 m, on Thamnolia vermicularis
var. subuliformis, 1997, M. Zhurbenko (LE 260560).
Cercidospora thamnoliae Zhurb. sp.
nov.
MycoBank no.: MB 563052
Fungus lichenicola in thallis lichenum generis Thamno-
lia crescens. Similis speciei Cercidospora ochrolechiae, sed
differt praecipue ascosporis (1–)3(–4)-septatis, et hos-
pite diverso.
Typus: Russia, Taimyr Peninsula, Byrranga Mts., N
of Levinson-Lessinga Lake, 74°33#N, 98°26#E, 180 m
elev., arctic tundra, on thalli of Thamnolia vermicularis
var. subuliformis, 26 August 1995, M. Zhurbenko 95571
(LE 260533—holotypus).
(Fig. 3A–D)
Vegetative hyphae indistinct.
Ascomata perithecioid, black, smooth,
subglobose, (60–)90–130(–200) m diam.
(ascomata of more than 150 m diam. were
observed only in LE 260528), with an ostiole
of 15–30 m diam., almost always immersed,
rarely slightly protruding in the ostiolar area
or exceptionally 1/3 exposed (only in LE
260528), arising singly, dispersed. Exciple
dark bluish green often with a grey shade and
30–40 m thick above, hyaline below, in
surface view resembling textura epidermoidea
or textura angularis (cells 3–10 m across),
K−. Hymenial gel I−, K/I−. Hamathecium of
abundant paraphysoids, non-capitate, 1·5–
2·0 m wide, septate with individual cells
6–15 m long, rather frequently branched
and anastomosed. Asci subcylindrical or usu-
ally moderately thickened in lower or some-
times central part, wall 1·5–5·0 m thick
apically and 0·5–1·5 m laterally, often with
a small ocular chamber, with an indistinct
or short stalk, (41–)53–69(–80) × (10–)11–
15(–18) m(n= 100; in K elongated up to
112 m), (4–)8-spored, I−, K/I−. Ascospores
hyaline, usually distinctly heteropolar, nar-
rowly soleiform with a narrower to attenu-
ated lower part, rarely almost narrowly
elliptic, upper or both ends often rather
acute, (12·5–)16·5–21·0(–27·5) × (4–)4·5–
5·5(–8) m, l/b = (2·5–)3·2–4·2(–5·4) (n=
308), with (1–)3(–4) septa (4-septate spores
occurred only in LE 260528), usually not or
only slightly constricted at the septa (mark-
edly constricted only when overmature or in
K), smooth walled, with many small guttules
best seen in K, arranged irregularly (often
diagonally) uni- or more commonly biseriate
in the ascus; a halo, 1·0–1·5 m thick, was
sometimes observed around spores within
the asci.
Anamorph not observed.
Notes. The following eight previously de-
scribed Cercidospora species regularly have
2012 Holarctic thamnoliicolous fungi—Zhurbenko 153

ascospores with three or more septa: C.
alpina Ihlen & Wedin (on Stereocaulon), C.
cladoniicola (on Cladonia), C. decolorella
(Nyl.) O. E. Erikss. & J. Z. Yue (on terricol-
ous algal films, Mycobilimbia,Peltigera,Proto-
pannaria, and Solorina), C. ochrolechiae
Zhurb. (on Ochrolechia and Pertusaria), C.
pluriseptata (Nyl.) Zopf (on Lecanora), C.
punctillata (on Biatora,Buellia,Cladonia,
Lecanora,Micarea,Mycobilimbia,Peltigera,
Phaeorrhiza,Pilophorus,Protopannaria,Pso-
roma,Solorina, and Sphaerophorus), C. soror
Obermayer & Triebel (on Arthrorhaphis), and
C. stereocaulorum (on Stereocaulon) (Nylander
1866; Alstrup et al. 1994; Hafellner &
Obermayer 1995; Zhurbenko et al. 1995;
Zhurbenko & Santesson 1996; Alstrup 1997;
Zhurbenko 2002, 2008, 2009a,b, 2010b;
Zhurbenko & Alstrup 2004; Zhurbenko &
Triebel 2005; Ihlen & Wedin 2007; von
Brackel 2010a; specimens of Cercidospora
punctillata examined for comparison). Cerci-
dospora alpina differs from C. thamnoliae in its
much larger (200–350 m diam.), semi-
immersed ascomata and longer ascospores
[(18–)19·5–33(–43) × (4–)4·5–6·5(–7) m]
with up to 7 septa; C. decolorella has some-
what larger ascomata (100–250 m diam.)
and ascospores often with 4–5 septa; C. cla-
doniicola readily differs in its olive-brown ex-
ciple; C. ochrolechiae is most similar to the
new species, differing from the latter mainly
in the occasional presence of 5-septate as-
cospores; C. pluriseptata has a brown exciple
and ascospores with up to 7 septa; C. punctil-
lata differs in its somewhat larger and mark-
edly erumpent ascomata, larger ascospores
[(14–)18·5–25·0(–33) × (4–)4·5–6·5(–9) m]
F. 3.Cercidospora thamnoliae A–D; A, ascomata (LE 260570); B, exciple in surface view in K (LE 260600); C,
interascal filaments in K/I (LE 260600); D, asci with spores in I (LE 232882). Cercidospora thamnoliicola E, asci with
spores in water (LE 260420). Scales:A=100m; B–E = 10 m.
154 THE LICHENOLOGIST Vol. 44

with (1–)3–5(–6) septa, and distinct patho-
genicity; C. soror is clearly distinguished by its
(2–)4-spored asci; C. stereocaulorum has
larger, sometimes almost superficial asco-
mata [100–200(–300) m diam.], (2–)4(–8)-
spored asci, and larger ascospores [(13–)
18·5–25·5(–30) × (4–)5–7(–8) m] with up
to 6 septa. Additionally, all these species grow
on different host genera. The comparison
with C. thamnoliicola, which also has many
3-septate ascospores, is provided in Table 1.
Features such as markedly exposed asco-
mata of more than 150 m diam., the occur-
rence of 4-septate ascospores, and the clear
bleaching of host tissues were observed only
in LE 260528, the inclusion of which signifi-
cantly broadened the new species concept.
That deviating sample resembles some speci-
mens of Cercidospora punctillata agg., which
thus might be the fifth Cercidospora species
growing on Thamnolia. However, the latter
species is the only one in the genus with an
extremely wide host range, which suggests
that it might be heterogenous. It is note-
worthy in this connection that 20 of 27
Cercidospora species are confined to one
host genus, the exceptions being, apart from
C. punctillata [syn. C. decolorella var. licheni-
cola (Zopf) O. E. Erikss. & J. Z. Yue], C.
decolorella (actually indistinguishable from
the latter), C. caudata,C. galligena,C.
ochrolechiae,C. solearispora, and C. werneri
Nav.-Ros., Calat. & Hafellner. Another con-
sideration is that in LE 260528 Cercidospora
thamnoliae occurred on an abnormally robust
host thallus, which might be the reason for
developing comparatively large and exposed
ascomata. Further material is needed to con-
firm the suggested broad species concept or
make it narrower.
Specimens examined of Cercidospora tham-
noliae also fit the protologue of C. lecidomae
Zhurb. & Triebel, the species distinguished
from the earlier described C. punctillata
(Nyl.) R. Sant. mainly by its smaller and less
septate ascospores (Zhurbenko & Triebel
2003). However, careful comparison of ad-
ditional material of Cercidospora lecidomae
and C. punctillata revealed that ascospores
of both species are (1–)3–5(–6)-septate and
almost indistinguishable in size: (15·0–)
18·5–27·0(–35·5) × (4·5–)5·0–6·5(–8·0) m
(n= 92) vs. (14–)18·5–25·0(–33) × (4–)4·5–
6·5(–9) m(n= 479). No additional dis-
criminating characters between the two
species were found. As a consequence, Cerci-
dospora lecidomae Zhurb. & Triebel [Biblio-
theca Lichenologica 86: 206 (2003)] is
reduced here to synonymy (syn. nov.) with
Cercidospora punctillata (Nyl.) Sant. [Lichen
forming and Lichenicolous fungi of Fennoscan-
dia 83 (2004); basionym: Verrucaria punctil-
lata Nyl. Flora 67: 223 (1884)]. Cercidospora
lecidomae has been reported on Thamnolia
vermicularis by Kukwa & Flakus (2009). Al-
though I did not examine this material, it is
likely to belong to Cercidospora thamnoliae.
Distribution and host. Known from polar
deserts and arctic tundra of the American
and Russian Arctic. Mostly found on old or
decaying thalli of Thamnolia vermicularis,
sometimes possibly causing local bleaching
of host tissues (their distinct discoloration
was observed only in LE 260528).
Additional specimens examined [on Thamnolia vermicu-
laris var. subuliformis (15), var. vermicularis (Sw.) Schaer.
(2)]. Canada: Canadian Arctic Archipelago: Ellef
Ringnes Island, Isachsen Bay, 78°47#N, 103°33#W,
40 m, 2005, N. Matveeva (LE 260316, LE 260526).—
Russia: Nenets Region: Malozemel’skaya Tundra,
Seduiyakha River, 68°23#N, 53°15#E, 1998, O.
Lavrinenko (LE 232884). Sverdrupa Island in Kara Sea:
74°33#N, 79°25#E, 10 m, 1992, Yu. Kozhevnikov (LE
260483). Severnaya Zemlya: Bolshevik Island, Mikoyana
Bay, 79°18#N, 101°55#E, 10 m, 1996, M. Zhurbenko
[LE 210306, erroneously reported as Cercidospora tham-
noliicola in Zhurbenko (2008); LE 260573]; ibid.,
Shokalskogo Strait, 79°16#N, 101°40#E, 20 m, 1996,
M. Zhurbenko (LE 233034, LE 232844, LE 232813, LE
260583, LE 260528); Sedova Archipelago, 79°25#N,
91°40#E, 1930, V. Savicz [LE 207056, erroneously
reported as C. thamnoliicola in Zhurbenko & Santesson
(1996)]. Novosibirskie Islands: Zhokhova Island, 76°08#
N, 152°45#E, 1989, M. Samarskii [LE 207223, errone-
ously reported as C. thamnoliicola in Karatygin et al.
(1999)]. Wrangel Island: Klark River, 71°07#N, 178°14#
W, 100 m, 1998, S. Kholod (LE 232882); Naskhok
River, 71°24#N, 178°08#W, 5 m, 1998, S. Kholod (LE
260570, LE 260600); Neozhidannaya River, 71°02#N,
179°10#E, 110 m, 1992, S. Kholod (LE 260374a).
Cercidospora thamnoliicola Ihlen
Graphis Scripta 7(1):18 (1995); type: Norway, Nord-
land, on Thamnolia vermicularis var. subuliformis,Norman
(O—holotype, n. v.).
2012 Holarctic thamnoliicolous fungi—Zhurbenko 155

Notes. Considering that the newly exam-
ined specimens reveal some discrepancies
with the protologue of this insufficiently
known species (Ihlen 1995), its full descrip-
tion is provided here.
Vegetative hyphae indistinct.
Ascomata perithecioid, black, smooth, sub-
globose to broadly ovate, 50–125 m diam.,
with an ostiole of 15–20(–40) m diam.,
immersed to slightly protruding in the osti-
olar area, arising singly or sometimes con-
tiguous, aggregated in groups of up to 100 or
more at the apices or central parts of the host
thalli. Exciple medium to dark olive-brown to
olivaceous buff and 17–25 m thick above,
hyaline and 8–12 m thick below, in surface
view resembling textura epidermoidea or
textura angularis (cells 3–8 m across), K+
yellowish brown. Hymenial gel I−, K/I−.
Hamathecium of sparse or sometimes indis-
tinct, non-capitate, c. 1·5 m wide, septate,
scarcely branched paraphysoids. Asci cylin-
drical or slightly thickened in the central or
lower part, wall 1–3 m thick apically and
0·5–1·5 m laterally, ocular chamber not
observed (with shallow ocular chamber
from the protologue), (35–)46–62(–65) ×
(7–)8–10 m(n= 46; in K elongated up to
76 m), (2–)4(–6)-spored (almost always
with 4 well-developed spores, very rare with
2–3 normal and 1–2 abortive spores or with
4 normal and 2 abortive spores) [37–41 ×
6–8(–12) m, 4(–6)-spored taken from the
protologue], I−, K/I− (Fig. 3E). Ascospores
hyaline, usually distinctly heteropolar,
narrowly elliptic, mostly with narrower to
attenuated lower or occasionally upper part
and often with rather acute ends, (11–)14·0–
17·5(–21) × (4·0–)4·5–5·5(–6·5) m, l/b =
(2·2–)2·6–3·4(–4·2) (n= 158), with (1–)2–3
septa (3-septate according to the proto-
logue), not or sometimes slightly constricted
at the septa, smooth-walled, distinct halo not
observed (halonate in the protologue), often
with a few conspicuous guttules per cell best
seen in K, uniseriate or diagonally arranged
in the ascus.
Anamorph not observed. Infected host
parts usually are slightly greyish and ‘scurfy’,
sometimes also somewhat curved or
swollen.
Poorly developed paraphysoids (also
stressed in the protologue), asci with an in-
distinct ocular chamber, and K+ reaction of
the exciple are not typical for Cercidospora.
This evidently rare species has been confused
with the more frequent C. thamnoliae (see
above under the latter species) and thus was
in fact formerly known only from two finds in
the Norwegian mountains (Ihlen 1995).
Here it is confirmed for this type region and
also reported new to the Asian Arctic.
The occurrence of four Cercidospora
species on one host species (Table 1) is re-
markable for lichenicolous fungi in general,
but not for this particular lichenicolous
fungal genus, where almost half of the host
lichen genera support more than one Cerci-
dospora species. For instance, three Cercido-
spora species are known on Aspicilia and
Circinaria (C. galligena,C. solearispora and
C. werneri) and another three on Caloplaca
[C. caudata,C. epicallopisma Arnold and C.
epicarphinea (Nyl.) Grube & Hafellner]
(Navarro-Rosinés et al. 2004, 2009). It may
also reflect the high heterogeneity of Tham-
nolia vermicularis populations (Nelsen &
Gargas 2009a,b). It is also possible that three
Cercidospora species with indistinct paraphys-
oids, treated above (Table 1), might be better
disposed in another genus.
Specimens examined [all in arctic or alpine tundras on
Thamnolia vermicularis var. subuliformis (2), var. vermicu-
laris (1)]. Norway: Troms County: Skibotndalen Valley,
69°16#N, 20°23#E, 950 m, 2003, M. Zhurbenko (LE
260460).—Russia: Taimyr Peninsula: Byrranga Mts.,
Levinson-Lessinga Lake, 74°24#N, 98°38#E, 150 m,
1995, M. Zhurbenko (LE 260420). Chukchi Peninsula:
Provideniya, 64°27#N, 173°11#W, 100 m, 2001, M.
Zhurbenko (LE 260440).
Cladosporium licheniphilum Heuchert
& U. Braun
Herzogia 19: 12 (2006); type: Russia, Altai, on Pertusaria
alpina Hepp ex H. E. Ahles, 1999, Davydov (LE—
holotype!).
Notes. New to the Arctic. Thamnolia is a
new host genus.
Specimen examined.Russia: Taimyr Peninsula: Byr-
ranga Mts., Levinson-Lessinga Lake, 74°32#N, 98°33#
E, 250 m, arctic tundra, on damaged portions of
156 THE LICHENOLOGIST Vol. 44

Thamnolia vermicularis var. subuliformis thalli, 1995, M.
Zhurbenko (LE 260309, HAL 2444 F; det. U. Braun).
Dacampia thamnoliicola Zhurb. ad int.
(Fig. 4)
Ascomata perithecioid, black, subglobose,
100–200 m diam., ostiolate, erumpent to
sessile. Exciple brown, in surface view similar
to textura angularis, composed of thick-
walled, angular, pseudoparenchymatous
cells. Hamathecium formed by well-
developed interascal filaments, which are
1·5–3·5 m wide, septate, and branched.
Asci fissitunicate, clavate, with a distinct foot,
70–100 × 22–24 m(n= 5), 8-spored.
Ascospores initially pale, then dark brown,
elliptic, with rounded or rather acute ends,
(23–)25–29(–31·5) × (10·5–)11–13(–14) m,
l/b = (1·8–)2·1–2·5(–2·7) (n= 45), muriform
when mature with 5 − 7 transverse septa and
a longitudinal or oblique septum in central
cells, slightly constricted at the septa, wall
ornamented (×1000), with halo 1·5–2·5 m
thick, guttules not observed, arranged diago-
nally uniseriate to biseriate in the ascus.
On ± decaying podetia of the host.
Notes. As noted by Halici & Hawksworth
(2008), the lichenicolous species in the
Dacampiaceae, with the exception of Pyr-
enidium actinellum Nyl., are almost always
host-specific. None of the 13 previously
known Dacampia A. Massal. species has been
found on Thamnolia. Among them, asco-
spores with up to 7 trans-septa were known
in D. cyrtellae Brackel (on Lecania), D. leca-
niae Kocourk. & K. Knudsen (on Lecania),
D. leptogiicola (D. Hawksw.) D. Hawksw. (on
Collema,Leptogium, and Pannaria), and D.
muralicola Halici & D. Hawksw. (on Proto-
parmeliopsis) (Hawksworth 1975; Etayo &
Breuss 1996; Halici & Hawksworth 2008;
Lopez de Silanes et al. 2009; von Brackel
2010b; Kocourková & Knudsen 2010).
Dacampia cyrtellae differs from D. thamnolii-
cola in its smaller ascomata (110–160 m
diam.) and smaller ascospores [(19·5–)21·2–
24·9(–26·0) × (6·5–)6·8–8·3(–9) m]; D. le-
caniae seems to be the closest to this
presumably new species, including the pres-
ence of a conspicuous gelatinous sheath, but
has somewhat smaller ascospores, measuring
(21–)22·8–26·5(–28) × (8–)9·5–12·0 m; D.
leptogiicola has smaller ascospores (21–25 ×
5·0–6·5 m); and D. muralicola has some-
what smaller ascospores [21–26(–31·5) ×
F. 4.Dacampia thamnoliicola (LE 207531b). A, asco-
mata; B, ascospore in water, note halo; C, immature
ascus in water; D, interascal filaments in water; E, asci
with spores in water. Scales:A=100m; B–E = 10 m.
2012 Holarctic thamnoliicolous fungi—Zhurbenko 157

(7·0–)9·0–12·5(–14·5) m] and smaller as-
comata (95–120 m diam.). Ascospores of
Dacampia thamnoliicola are not that typical
for the genus in being halonate and orna-
mented; the first feature also shared by D.
lecaniae and D. rubra Halici, Candan & Calat.
and the second one by D. rhizocarpicola D.
Hawksw., D. rubra, and D. xanthomendozae
Etayo & Halici (Halici & Hawksworth 2008;
Halici et al. 2009a,b).
Specimen examined.Russia: Severnaya Zemlya:
Sedova Archipelago, 79°25#N, 91°40#E, on Thamnolia
vermicularis var. vermicularis, 1930, V. Savicz 2017b (LE
207531b).
Epithamnolia Zhurb. gen. nov.
MycoBank no.: MB 563053
Genus lichenicola ad Coelomycetes pertinens. Conidi-
omata plus minusve nigra, subglobosa ad cupuliformia,
cum muris textura globulosa vel porrecta, subimmersa
ad superficialia, singularia, dispersa. Conidiophora des-
unt. Cellulae conidiogenae enteroblasticae, phialidicae,
non prolifericae, hyalinae, lagenulatae vel anguste ellip-
ticae. Conidia acrogena, hyalina, cylindrica, (0–)1(–2?)-
septata, laevia.
Typus: Epithamnolia karatyginii Zhurb.
Genus lichenicolous, belonging to the
Coelomycetes.
Conidiomata pycnidial, blackish, rugose,
irregularly subglobose, but becoming cupuli-
form with age, initially almost closed, then
with an irregular opening, subimmersed in
thalli of Thamnolia species to finally superfi-
cial, arising singly, dispersed. Exciple in cross-
section dark to medium brown throughout,
in surface view resembling textura angularis/
globulosa or textura porrecta, K−. Conidi-
ophores absent. Conidiogenous cells lining the
pycnidial cavity, enteroblastic, phialidic,
with indistinct collarette and periclinal thick-
ening, not proliferating, hyaline, lageniform
to narrowly elliptic, smooth-walled. Conidia
acrogenous, arising singly, not catenate, hya-
line, cylindrical, with obtuse to sometimes
slightly truncated ends, ± straight, (0–)1
(–2?)-septate, not constricted at the septum,
smooth-walled, sometimes with sparce in-
conspicuous guttules.
Notes. Amongst the non-lichenicolous pyc-
nidial fungal genera, the new genus mainly
recalls Ascochyta Lib. agg., Clypeopycnis
Petr., Didymochaeta Sacc. & Ellis, Gelatino-
pycnis Dyko & B. Sutton, Pocillopycnis Dyko
& B. Sutton emend. DiCosmo, Pseudocenan-
gium P. Karst., and Septopatella Petr. (Dyko
& Sutton 1979; DiCosmo 1980; Sutton
1980; Mel’nik 2000). However, there are
significant differences. In Ascochyta agg. the
conidiomata are non-cupulate; in Clypeopyc-
nis the conidiomata are non-cupulate, and
conidiophores are present; in Didymochaeta
Sacc. & Ellis the conidiomata are non-
cupulate, and the conidiogenous cells have a
distinct collarette and a periclinal thickening;
in Gelatinopycnis the conidiomata are closed
and have a complex wall, and conidiophores
are present; in Pocillopycnis conidiophores are
present, the conidiogenous cells are sympo-
dially proliferating, and the conidia are holo-
blastic, lunate to sigmoid, 3–10-septate; in
Pseudocenangium conidiophores are present
and the conidiogenous cells are annelidic; in
Septopatella conidiophores are present, and
the conidiogenous cells are sympodially pro-
liferating. Considering the coelomycetes
known to occur on lichens, Epithamnolia
karatyginii, the type species of this monotypic
genus, resembles species of Hainesia Ellis &
Sacc. in having finally cupulate conidiomata
and hyaline, cylindrical, septate conidia
(Etayo & Diederich 1996a; von Brackel
2009). However, the latter are easily dis-
tinguished from the new species by their
branched conidiophores (Punithalingam &
Spooner 1997). Epitham
nolia karatyginii may
also resemble the lichen
ized Woessia fusari-
oides D. Hawksw., Poelt & Tsch.-Woess.,
which differs from the former in having
white, shallow cupuliform to disc-like conidi-
omata, falcate, non-septate conidia and in
the lignicolous life habit (Hawksworth &
Poelt 1986).
Epithamnolia karatyginii Zhurb. sp. nov.
MycoBank no.: MB 563054
Fungus lichenicola in thallis lichenum generis Thamno-
lia crescens. Conidiomata 110–220 m diam. Cellulae
conidiogenae (5–)6·5–9(–10) × (1·5–)2–3 m. Conidia
(14–)18·5–27(–32) × (1–)1·5–2(–2·5) m.
Typus: Canada, British Columbia, Wells Gray Pro-
vincial Park, Mt. Raft, 51°44#N, 119°50#W, 2100 m
158 THE LICHENOLOGIST Vol. 44

elev., alpine tundra with sparse Picea engelmannii,on
moribund thalli of Thamnolia vermicularis var. subuli-
formis, 3 August 2002, M. Zhurbenko 02343 (LE
260498—holotypus).
(Fig. 5)
Vegetative hyphae pale brown, somewhat
flexuose, 2·5–5·0 m wide, scarcely septate,
constricted at the septa, immersed.
Conidiomata pycnidial, blackish, glossy,
rugose, irregularly subglobose at first, be-
coming cupuliform with age, 110–220 m
diam., initially almost closed, then with an
irregular opening (20–)40–60(–100) m
across, exposing the pale brown interior, sub-
immersed to finally superficial, arising singly,
dispersed. Exciple in cross-section dark to
medium brown throughout, basally of 3–5
cell layers, in surface view resembling textura
angularis/globulosa (cells 3·5–10·0 m across)
or textura porrecta, K−. Conidiophores absent.
Conidiogenous cells lining the pycnidial cavity,
enteroblastic, phyalidic, with indistinct col-
larette and periclinal thickening, not prolifer-
ating, hyaline, lageniform to narrowly
elliptic, (5–)6·5–9·0(–10) × (1·5–)2–3 m
(n= 39), smooth-walled. Conidia abundant,
acrogenous, arising singly, not catenate, hya-
line, cylindrical, with obtuse to sometimes
slightly truncated ends, ± straight, (14–)
18·5–27·0(–32) × (1·0–)1·5–2·0(–2·5) m,
l/b = (6·0–)9·8–17·4(–21·3) (n= 122), (0–)
1(–2?)-septate, not constricted at the sep-
tum, smooth-walled, sometimes with sparce
inconspicuous guttules.
Etymology. The species is named after the
eminent Russian mycologist Dr Igor V.
Karatygin, the main author of the first
F. 5.Epithamnolia karatyginii (holotype). A, conidiomata; B, exciple in water; C, conidiogenous cells and conidia
in water; D, conidia in water. Scales:A=100m; B–D = 10 m.
2012 Holarctic thamnoliicolous fungi—Zhurbenko 159

catalogue of the Russian Arctic fungi
(Karatygin et al. 1999).
Distribution and host. Known from polar
deserts, arctic and alpine tundras of North
America and Eurasia. On ± damaged basal
or apical parts of Thamnolia vermicularis
thalli, sometimes intermixed with Sphaerello-
thecium thamnoliae Zhurb. or Stigmidium
frigidum (Th. Fr. ex Sacc.) Alstrup & D.
Hawksw.
Additional specimens examined [on Thamnolia vermicu-
laris var. subuliformis (5), var. vermicularis (1)]. Russia:
Severnaya Zemlya: Bolshevik Island, Shokalskogo Strait,
79°16#N, 101°40#E, 20 m, 1996, M. Zhurbenko (LE
232933). Northern Ural: headwaters of Pechora River,
Yanypupuner Range, 62°05#N, 59°06#E, 800 m, 1997,
M. Zhurbenko (LE 260538a, intermixed with Sphaerello-
thecium thamnoliae var. thamnoliae). Kola Peninsula:
Barents Sea coast, Voron’ya River mouth, 69°09#N,
35°50#E, 20 m, 1997, M. Zhurbenko (LE 260448).
Taimyr Peninsula: Byrranga Mts., Bolshaya Bootankaga
River, 74°30#N, 97°40#E, 350 m, 1995, M. Zhurbenko
(LE 232994b, intermixed with Stigmidium frigidum).
Yakutiya: Laptev Sea coast, Tiksi, 71°40#N, 128°40#E,
50 m, 1998, M. Zhurbenko (LE 260444b, intermixed
with Stigmidium frigidum).
Geltingia associata (Th. Fr.) Alstrup &
D. Hawksw.
Meddel. Grønl., Biosc.31: 33 (1990); type: Spitsbergen,
Danskøn, 1861, Malmgren (UPS—lectotype, n. v.).
Note. This fungus mostly grows on
Ochrolechia spp., but has also been recorded
from Thamnolia (Walker 1970; Zhurbenko &
Santesson 1996; Diederich et al. 2010).
Specimen examined.Russia: Taimyr Peninsula:
Byrranga Mts., Levinson-Lessinga Lake, 74°31#N,
98°36#E, 250 m, arctic tundra, on moribund thalli
of Thamnolia vermicularis var. subuliformis, 1995, M.
Zhurbenko (LE 260468).
Lichenoconium usneae (Anzi) D.
Hawksw.
Persoonia 9(2):185 (1977); type: Italy, on Usnea filipen-
dula Stirt. agg. (K—isotype, n. v.).
Note. This common coelomycete has been
recorded from no less than 29 lichen genera,
including one previous record on Thamnolia
(Hafellner & Türk 1995).
Specimens examined.Russia: Taimyr Peninsula:
Byrranga Mts., Levinson-Lessinga Lake, 74°32–33#N,
98°26–33#E, 180–300 m, arctic tundra, on partly mori-
bund thalli of Thamnolia vermicularis var. subuliformis,
1994, M. Zhurbenko (LE 207532c); ibid., 1995, M.
Zhurbenko (LE 260544).
Lichenopeltella thamnoliae R. Sant.
Thunbergia 28: 7 (1998); type: Colombia, Boyacá, on
Thamnolia vermicularis var. vermicularis, 1972, Cleef
(UPS—holotype, n. v.).
Notes. Previously known from Colombia,
Venezuela, Ecuador, and Peru (Santesson
1998; Etayo 2010). New to Bolivia. Tham-
nolia papelillo var. subsolida (M. Satô) R. Sant.
is a new host variety.
Specimen examined.Bolivia: 25 km S of Sorata, 1 km
NE of Laguna Ajuyani, 15°53#S, 68°39#W, 4250 m, on
Thamnolia papelillo var. subsolida, 2000, A. Elvebakk
(LE).
Merismatium thamnoliicola Alstrup &
E. S. Hansen
Graphis Scripta 12: 43 (2001); type: Greenland, Thule
District, on Thamnolia vermicularis, 1992, Alstrup (C—
holotype, n. v.).
Vegetative hyphae pale to medium brown,
1·5–2·5 m wide.
Ascomata perithecioid, black, subglobose,
often with a small papilla, 100–150 m
diam., 1/4–3/4 protruding (superficial ac-
cording to the protologue). Hymenial gel I+
brownish orange, K/I+ blue. Ascospores ini-
tially hyaline, then pale to medium oliva-
ceous buff, snuff brown, or greyish brown, at
maturity with contrasting darker septa, ellip-
tic to oblanceolate (Fig. 6), constricted lower
ends noted in the protologue were observed
only in some overmature spores and are
evidently the result of spore destruction,
(14–)16·5–24·0(–34) × (6–)6·5–8·5(–9) m,
l/b = (1·6–)2·1–3·3(–4·3) (n= 89) (22–
30 × 8·5–9·5 m according to the proto-
logue), with (3–)5–7(–8) transverse septa
and a longitudinal or oblique septum in cen-
tral or all cells, smooth-walled.
Notes. Mostly found on moribund bases of
host thalli, pathogenicity not observed. The
species was described and known previously
from two collections in Greenland (Alstrup
& Hansen 2001) and thus is new to Eurasia.
Examination of new material revealed some
160 THE LICHENOLOGIST Vol. 44

discrepancies with its protologue, described
above.
Specimens examined (all in polar desert, arctic or al-
pine tundras on Thamnolia vermicularis var. subuliformis).
Russia: Severnaya Zemlya: Bolshevik Island, Akhma-
tova Bay, 79°04#N, 102°45#E, 10 m, 1996, M. Zhur-
benko (LE 260554); Sedova Archipelago, 79°25#N,
91°40#E, 1930, V. Savicz (LE 260430, LE 260450).
Taimyr Peninsula: Byrranga Mts., Bolshaya Bootankaga
River, 74°20#N, 98°05#E, 400 m, 1991, V. Kuvaev
(LE 260473). Eastern Sayan Mts.: Kryzhina Range,
headwaters of Belyi Kitat River, 53°59#N, 95°32#E,
1500 m, 2009, M. Zhurbenko (LE 260373).
Odontotrema santessonii Zhurb.,
Etayo & Diederich
Lichenologist 34: 495; type: Russia, Chukchi Peninsula,
on Thamnolia vermicularis var. subuliformis, 1971, Yurtsev
(LE 207696—holotype!; hb. Diederich—isotype!).
Notes. Ascomata up to 450 m diam., fi-
nally sometimes almost sessile. Hymenium I+
brownish red to coral throughout. Ascospores
hyaline, narrowly oblanceolate, (15–)18–
23(–25) × (3–)3·5–4·5(–5) m, l/b = (3·8–)
4·7–5·9(–6·6) (n= 39), with (3–)7–8 trans-
verse septa and often a few longitudinal or
oblique septa in central cells, arranged ± in
two 4-spored fascicles in the ascus. Patho-
genicity not observed. New to North
America.
Specimens examined [all in polar desert or arctic tun-
dra on Thamnolia vermicularis var. subuliformis (5), var.
vermicularis (4)]. Canada: Canadian Arctic Archipelago:
Axel Heiberg Island, Bunde Fjord, 80°30#N, 94°36#W,
30 m, 1999, N. Matveeva (LE 260543).—Russia:
Severnaya Zemlya: Bolshevik Island, Akhmatova Bay,
79°04#N, 102°45#E, 10 m, 1996, M. Zhurbenko (LE
232853). Taimyr Peninsula: Byrranga Mts., Bolshaya
Bootankaga River, 74°30#N, 97°40#E, 200 m, 1995,
M. Zhurbenko 95309 (LE 232814). Yakutiya: Laptev
Sea coast, Tiksi, 71°39–40#N, 128°40–45#E, 50 m,
1998, M. Zhurbenko (LE 232854, LE 260524, LE
260324); Lena River delta, Stolb Island, 72°24#N,
126°40#E, 50 m, 1998, M. Zhurbenko (LE 260364).
Wrangel Island: Neizvestnaya River, 71°11#N, 179°15#
W, 170 m, 1987, S. Kholod (LE 232962). Chukotka:
Gil’mymlineiveem River, 65°48#N, 173°15#W, 1977,
I. Makarova (LE 233094).
Odontotrema thamnoliae Zhurb.,
Diederich & Etayo
Lichenologist 34: 497; type: Russia, Chukchi Peninsula,
on Thamnolia vermicularis var. vermicularis, 1971,
Makarova (LE 207695—holotype!).
Notes. Hymenium I+ orange except the
uppermost part, which is I+ blue. Ascal wall
I+ blue at the apex, I− below. Sometimes
slightly bleaching the host tissues. New to
North America.
Specimens examined [all in arctic or alpine tundras on
Thamnolia vermicularis var. subuliformis (3), var. vermicu-
laris (3)]. USA: Alaska: Seward Peninsula, 7 km NE of
Nome, 64°33#N, 165°21#W, 220 m, 2001, M. Zhur-
benko (LE 260376).—Russia: Kola Peninsula: Barents
F. 6.Merismatium thamnoliicola, old ascospores in I
(LE 260554). Scale = 10 m.
2012 Holarctic thamnoliicolous fungi—Zhurbenko 161

Sea coast, Voron’ya River mouth, 69°09#N, 35°50#E,
20 m, 1997, M. Zhurbenko (LE 260464). Western Sayan
Mts.: Kulumys Range, 52°51#N, 93°17#E, 1700 m,
2010, M. Zhurbenko (LE 260588). Taimyr Peninsula:
Byrranga Mts., Levinson-Lessinga Lake, 74°24#N,
98°38#E, 150 m, 1995, M. Zhurbenko (LE 260597).
Yakutiya: Laptev Sea coast, Tiksi, 71°37#N, 128°54#E,
70 m, 1998, M. Zhurbenko (LE 233074). Chukotka:
Kolyuchinskaya Bay, Yuniveem River mouth, 66°40#N,
173°54#W, 20 m, 1980, A. Katenin &M. Zhurbenko
(LE 232864).
Phaeospora arctica Horáková &
Alstrup
Graphis Scripta 6: 61 (1994); type: Greenland, Disco
Island, on Arctocetraria andreievii (Oxner) Kärnefelt & A.
Thell, 1983, Poelt (PRM 842918—holotype, n. v.).
Ascomata black, subglobose, c. 120 m
diam., semi-immersed. Hymenial gel appears
I+ pale red. Hamathecium not observed.
Asci subcylindrical to obclavate, with a long
internal apical beak, c. 55–60 × 11–14 m,
6–8-spored, I−. Ascospores hyaline to pale
cinnamon-brown, narrowly elliptic (mostly)
to elliptic and often slightly narrower below,
(10–)15·5–20·5(–22) × (4·5–)5–6(–8·0) m,
l/b = (1·7–)2·7–3·9(–4·7) (n= 51), (1–)3-
septate, not constricted at the septa, smooth-
walled, without halo, guttulate, biseriate in
the ascus. Growing on the bleached, possibly
moribund base of the host’s podetia.
Notes. The specimen fits well the species
concept of Phaeospora arctica, except staining
of the hymenial gel with I, not reported be-
fore (Horáková & Alstrup 1994; Zhurbenko
& Santesson 1996). The species was formerly
known on Allocetraria,Arctocetraria, and
Arctoparmelia, thus Thamnolia being a new
host genus.
Specimen examined.USA: Alaska: Great Kobuk Sand
Dunes, 67°05#N, 158°58#W, 50 m, on Thamnolia
vermicularis var. subuliformis, 2000, M. Zhurbenko (LE
260359).
Phoma thamnoliae Zhurb. ad int.
(Fig. 7)
Vegetative hyphae pale brown, 3–5(–7) m
diam., septate, constricted at the septa.
Conidiomata pycnidial, black, subglobose,
indistinctly papillate, 75–110 m diam., with
an ostiole of c.10m diam., subimmersed to
almost sessile. Pycnidial wall brown, darker
around the ostiole, in surface view com-
posed of angular pseudoparenchymatous
cells (textura angularis) 4–10 m across, K−.
Pycnidial gel I−, K/I−. Conidiogenous cells
not observed clearly, probably ampulliform,
c.5×2m. Conidia hyaline, oblong to occa-
sionally elliptic, with rounded ends, (3·5–)4–
5(–5·5) × (1·5–)2·0(–2·5) m, l/b = (1·6–)
2·0–2·6(–2·8) (n= 135), non-septate,
smooth-walled, with 2 − 3 conspicuous,
usually apical, guttules, abundant, arising
singly. Infected parts of the host thalli are
bleached or pale red.
Notes. The rather scant material does not
permit a formal description of this evidently
new Phoma Sacc. species. Most (c. 70%) of
F. 7.Phoma thamnoliae (LE 260349). A, squashed
peridium in water; B, conidia in water. Scales: A&B=
10 m.
162 THE LICHENOLOGIST Vol. 44

the 22 currently recognized lichenicolous
species of Phoma are restricted to a single
host genus, and none of them has been
reported on Thamnolia (Icmadophilaceae,
Pertusariales). The only other Phoma species
hosted by a Pertusariales genus (Dibaeis)is
P. maculiformans Ihlen (Ihlen 1998), which
readily differs from P. thamnoliae in its larger
pycnidia (120–200 m diam.) and non-
guttulate, longer conidia, measuring 5–6
(–7) × 2·0–2·5 m. The shape and size of the
conidia are among the most important diag-
nostic morphological characters in Phoma
taxonomy. Of the other lichenicolous species
referred to the genus, Phoma thamnoliae is
most similar in these characters to P. aggre-
gata Calat. & Etayo, P. cladoniicola Dieder-
ich, Kocourk. & Etayo, P. grumantiana
Zhurb. & Diederich, P. lecanorina Diederich,
P. melanohaleicola D. Hawksw. & Earl.-
Benn., P. peltigerae (P. Karst.) D. Hawksw.,
and P. rozziana Etayo (Hawksworth 1981;
Diederich 1986; Etayo 1996; Calatayud &
Etayo 2001; Hawksworth & Cole 2004;
Earland-Bennett et al. 2006; Diederich et al.
2007; Etayo & Sancho 2008). However,
Phoma aggregata (on Diploschistes) is clearly
separated from the new species by the minute
pycnidia (30–50 m diam.), aggregated in
dense groups of up to 200; P.cladoniicola
(on Cladonia) has larger conidia [(4–)4·5–6·0
(–7·5) × (2–)2·5–3·0(–3·5) m]; P. gruman-
tiana is very similar to the studied material in
the sizes of pycnidia (50–100 m diam.) and
conidia [(3–)4–5(–6) × 1·5–2·0(–3) m], but
could be distinguished by its host selection
(Cladonia, Lecanorales); P. lecanorina (on
Lecanora and Flavoparmelia) differs in its
smaller pycnidia (15–60 m diam.), the
peridium, being dark green above and sub-
hyaline below, and narrower conidia (3·2–
5·0 × 1·2–1·6 m); P. melanohaleicola (on
Melanohalea) has slightly longer conidia (5–
5·5 × 2–2·5 m); P. peltigerae (on Peltigera)
has larger pycnidia (up to 200 m diam.)
and conidia [(4–)4·5–6·0(–7) × 2·0–2·5(–3)
m]; and P. rozziana (on Pseudocyphellaria)
has smaller pycnidia, measuring 50–80 m
diam. Phoma thamnoliae might also be
compared with Briancoppinsia cytospora
(Vouaux) Diederich, Ertz, Lawrey & van den
Boom [syn. Phoma cytospora (Vouaux) D.
Hawksw.], growing on a wide range of differ-
ent host genera (Diederich et al. 2012). How-
ever, that species differs in its smaller
pycnidia (40–80 m diam.), the peridium
reacting K+ dark olivaceous, pycnidial gel
reacting I+ and K/I+ red, and basally trun-
cate, slightly curved, longer conidia, measur-
ing 5–7 × 1·5–2 m.
Specimens examined (both on upper parts of podetia of
Thamnolia vermicularis var. subuliformis). USA: Alaska:
Great Kobuk Sand Dunes, 67°05#N, 158°58#–159°00#
W, 50 m, Dryas-lichen-moss vegetation among sparse
Picea glauca, 2000, M. Zhurbenko 00226, 00468 (LE
260369, LE 260349).
Polycoccum vermicularium (Linds.) D.
Hawksw.
Bull. Br. Mus. nat. Hist., Bot.14(2):172 (1985); type:
Falkland Isles, on Thamnolia vermicularis, 1842, Hooker
(E—holotype, n. v.).
Rich material now available of this species
permits us to improve or complete its pre-
vious descriptions (Hawksworth 1985;
Hawksworth & Diederich 1988; Ihlen 1995).
Ascomata perithecioid, black, subglobose,
50–120 m diam., ostiolate, initially com-
pletely immersed (sometimes in few layers),
then mostly protruding in the ostiolar area,
arising singly, sometimes adjacent or a few
confluent, aggregated in dense groups.
Hamathecium of well-developed interascal
filaments, which are 1·5–3·5 m wide, sep-
tate, branched, composed of individual cells
4–12 m long. Asci elongate-clavate, often
with a distinct ocular chamber, with a short
foot, (55–)61–75(–80) × 20–26(–30) m
(n= 30), 8-spored, I− (except plasma be-
coming red-brown). Ascospores long hyaline,
then olive and finally dark brown, walls and
septum of young pale olive spores markedly
darker than lumina, narrowly obovate to
elliptic, upper cell usually wider than lower
one, sometimes equally-celled, (13–)15·5–
20·0(–28) × (6–)7–9(–12) m, l/b = (1·5–)
1·9–2·5(–3·3) (n= 165), (0–)1-septate, not
to markedly constricted at the septum,
wall smooth, not verruculose as stated in
Hawksworth (1985), occasionally with halo
1–2 m thick (mostly around young hyaline
2012 Holarctic thamnoliicolous fungi—Zhurbenko 163

spores), often with one big guttule in each
cell, arranged irregularly biseriate in the
ascus.
Conidiogenous cells ± pyriform, 6–8 × 2–
3m. Conidia hyaline, ± oblong, straight to
slightly bent, (3–)4–6(–7·5) × (1–)1·5 m
(n= 63), non-septate.
Notes. Infected host tissues become grey,
then black, sometimes exhibit brown gall-like
swellings up to 2 mm diam., or disintegrate.
Macroscopically the infections are similar to
those of Stigmidium frigidum. New to Asia
(Russia and Mongolia).
Specimens examined [all in polar desert, arctic or al-
pine tundras, or northern boreal forest on Thamnolia
vermicularis var. subuliformis (22), var. vermicularis (7)].
USA: Alaska: Great Kobuk Sand Dunes, 67°05#N,
158°58#W, 50 m, 2000, M. Zhurbenko (LE 260339).—
Norway: Troms County: Skibotndalen Valley, 69°16#N,
20°23#E, 950 m, 2003, M. Zhurbenko (LE 260306).—
Russia: Kola Peninsula: Barents Sea coast, Voron’ya
River mouth, 69°09#N, 35°50#E, 20 m, 1997, M.
Zhurbenko (LE 260414). Nenetz Region: Bolshoi
Tsinkovyi Island, 70°27#N, 58°40#E, 1997, V.
Shevchenko (LE 232924). Altai Mts.: Ukok Tableland,
49°19#N, 87°36#E, 2500 m, 1996, T. Lunke (LE
260453). Western Sayan Mts.: Chernoe Lake, 52°49#N,
94°07#E, 1570 m, 2010, M. Zhurbenko (LE 260320);
Kulumys Range, 52°51–52#N, 93°15–17#E, 1700 m,
2010, M. Zhurbenko (LE 260326, LE 260360); Oiskii
Range, headwaters of Olen’ya River, 52°48#N,
93°15#E, 1650 m, 2010, M. Zhurbenko (LE 260380a).
Eastern Sayan Mts.: Kryzhina Range, headwaters of
Belyi Kitat River, 53°59#–54°01#N, 95°27#–95°33#E,
1800 m, 2009, M. Zhurbenko (LE 260403b, LE
260393). Taimyr Peninsula: Byrranga Mts., Levinson-
Lessinga Lake, 74°34–37#N, 98°33–47#E, 150–250 m,
1995, M. Zhurbenko (LE 233054, LE 260424); ibid.,
Bolshaya Bootankaga River, 74°30#N, 97°40#E,
350 m, 1995, M. Zhurbenko (LE 260593); ibid.,
Krasnaya River, 74°35#N, 98°08#E, 200 m, 1994, M.
Zhurbenko (LE 232803, LE 260563 − anamorph); Mt.
“217” near junction of Pravaya Uboinaya and Uboinaya
Rivers, 73°25#N, 82°51#E, 150 m, M. Zhurbenko
(LE 260363). Yakutiya: Laptev Sea coast, Tiksi, 71°37–
39#N, 128°45–54#E, 70 m, 1998, M. Zhurbenko (LE
232833, LE 260344a, LE 260348); Lena River delta,
3 km E of Cape Krest-Tumsa, 72°22#N, 126°42#E,
50 m, 1998, M. Zhurbenko (LE 232914). Wrangel Island:
Naskhok River, 71°24#N, 178°08#W, 5 m, 1998, S.
Kholod (LE 260580); Neizvestnaya River, 71°20#N,
179°29#W, 1986, A. Dobrysh (LE 260397 − ana-
morph). Komandorskie Islands: Bering Island, Nikolskoe,
55°12#N, 165°57#E, 1980, A. Dombrovskaya
(KPABG, LE 232804). Chukotka: Baranikha, 68°30#N,
168°16#E, 1971, A. Galanin (LE 260553); ibid., 1971,
I. Makarova (LE 260319 − anamorph); Chaplinskie hot
springs, 64°25#N, 172°30#W, 1957, V. Gavrilyuk & P.
Gagarin (LE 232934); Inchoun, 66°15#N, 170°20#W,
1975, I. Makarova (LE 232874).—Mongolia: Bayan-
Khongor Region: N of Khukh-Nur Lake, 47°31#N,
98°33#E, 3000 m, 1972, L. Biazrov 5804 (LE 260474).
Dzabkhan Region: Tarbagatai Range, Solongotuin-Daba
Pass, 48°15#N, 98°54#E, 2650 m, 1973, L. Biazrov
2134 (LE 260494).
Sphaerellothecium thamnoliae Zhurb.
sp. nov.
MycoBank no.: MB 563055
Fungus lichenicola in thallis lichenum generis Thamno-
lia crescens. Similis speciei Stigmidium frigidum, sed
differt hyphis semper copiosis, bene visibilibus, ascoma-
tibus minoribus, (30–)50(–80) m diam., sessilibus vel
semi-immersis, dispersis, ascosporis interdum 2–3-
septatis et halonatis.
Typus: USA, Alaska, Kobuk Valley Wilderness, junc-
tion of Kobuk River and Kavet Creek, 67°07#N,
159°03#W, 50 m elev., Dryas-lichen-moss vegetation
among sparse Picea glauca, on thalli of Thamnolia
vermicularis var. subuliformis, 13 August 2000, M.
Zhurbenko 00212 (LE 260567—holotypus).
(Fig. 8)
Vegetative hyphae medium brown, with
sculptured surface, composed of elongate
cells 3·5–9·0 m wide, septate, markedly
constricted at the septa, branched (often
more or less rectangularly) and anastomo-
sed, BCr+ blue-green, I−, mostly superficial,
abundant, forming a conspicuous dense
reticulum.
Ascomata perithecioid, black, subglobose
to somewhat conical above, (30–)50(–80)
m diam., with an ostiole of 5–15 m diam.,
without appendices or projections, sessile to
semi-immersed, dispersed. Exciple in surface
view brown throughout, sculptured, similar
to textura angularis, composed of a few layers
of angular, pseudoparenchymatous cells (3–)
5–8(–12) m across, BCr+ blue-green, K−.
Hymenial gel I−, K/I−. Hamathecium not ob-
served. Asci elliptic to pyriform, occasionally
with a short foot, endoascus thickened
above, distinct ocular chamber not observed,
size significantly differing in the two varieties
described below, 8-spored, BCr−, I−, K/I−
(except plasma reacting BCr+ blue, I+, and
K/I+ yellow to orange). Ascospores colourless
and smooth-walled or rarely (when over-
mature ?) pale to medium brown and with a
164 THE LICHENOLOGIST Vol. 44

rough wall, obskittle-shaped to narrowly
obovate, usually with a wider and sometimes
shorter upper cell, occasionally located up-
side down in the ascus, size significantly dif-
fering in the two varieties described below,
almost always 1-septate, but non-septate
hyaline and 2–3-septate brown spores were
also observed, mostly constricted at the
septa, sometimes with halo 0·5–2·0 m thick,
usually with a few big guttules in each cell,
irregularly arranged in 2–3 rows in the ascus,
wall and septum BCr−, plasma BCr+ blue.
Conidiomata pycnidial. Conidia hyaline,
bacilliform, 2·0–4·5 × 0·5 m.
Notes. The material examined (42
specimens) exhibits significant variability in
the development of vegetative hyphae, size of
the ascomata, degree of their immersion in the
host thalli, and size of asci and ascospores.
The species is typified on the specimen repre-
senting the group with abundantly developed
superficial hyphae, comparatively small,
mostly sessile ascomata, and small asco-
spores. It can be opposed to the group dis-
tinguished by the larger asci and ascospores.
However, there is no sharp delimitation be-
tween the two groups (Table 2), which are
therefore recognized here as varieties.
F. 8.Sphaerellothecium thamnoliae. A, C & D, var. thamnoliae; A, infection habitus (holotype); C, asci in water (LE
260558); D, ascospores in I, note halo (holotype);B&E,var. taimyricum; B, infection habitus (LE 232894); E,
ascospores in water, note halo (holotype). Scales:A&B=500m; C–E = 10 m.
2012 Holarctic thamnoliicolous fungi—Zhurbenko 165

In spite of the absence of visible interascal
filaments, the species is disposed within
Sphaerellothecium Zopf vs. Stigmidium Trevis.
due to its brown, thick-walled, sculptured,
comparatively thick and short-celled vegeta-
tive hyphae with swollen individual cells,
forming a conspicuous superficial reticu-
lum, and non-pseudotetrablastic ascospores
(generic delimitation according to: Triebel
1989; Roux & Triebel 1994; Calatayud &
Triebel 1999, 2001). Up to now, no Sphaerel-
lothecium species have been reported from
Thamnolia. Together with Sphaerellothecium
araneosum (Arnold) Zopf, S. icmadophilae (R.
Sant.) Zhurb. and S. stereocaulorum Zhurb.
& Triebel, the new species belongs to the
group with 1(–3)-septate, hyaline asco-
spores, occasionally becoming brown with
age (Santesson 1984; Roux & Triebel 1994;
Zhurbenko & Triebel 2008). Sphaerellothe-
cium araneosum differs from both varieties of
S. thamnoliae in its sometimes distinct inter-
ascal and ostiolar filaments and different
sizes of asci (34–45 × 14–16 m) and asco-
spores [(12·5–) 13·5–17·0 (–22·0) × (3·5–)
5–7 (–7·5) m], S. icmadophilae has non-
halonate ascospores of intermediate size (14–
19 × 4·5–5·5 m), and S. stereocaulorum has
slightly smaller ascomata (20–50 m diam.),
smaller asci [(23–)25–34(–38) × (10–)11–
16(–19) m], non-halonate, smaller asco-
spores [(9–)10–13(–16) × 3·0–4·5(–6·0) m]
and grows in the epinecral layer of the host
thallus. Characters distinguishing Sphaerello-
thecium thamnoliae from Stigmidium frigidum
are summarized in Table 2.
Distribution and host. Circumpolarly
known in the Holarctic from polar deserts,
arctic and alpine tundras. Mainly growing on
bases of Thamnolia vermicularis thalli; occa-
sionally throughout whole thallus, particu-
larly when they are prostrate. Also found on
galls induced in the host by Thamnogalla
crombiei. Evidently pathogenic as infected
host parts are bleached.
Sphaerellothecium thamnoliae var.
thamnoliae Zhurb.
(Fig. 8A,C&D)
T 2.Distinguishing characteristics of Sphaerellothecium and Stigmidium taxa growing on Thamnolia (based on the author’s own data).
Characters Stigmidium frigidum Sphaerellothecium thamnoliae
var. thamnoliae var. taimyricum
Vegetative hyphae usually immersed and inconspicuous superficial, conspicuous
Ascomata
size (m diam.) (30–)50–80(–120) (30–)50(–80)
immersion in the host thalli mostly protruding only in the ostiolar area sessile to semi-immersed
dispersion on the host thalli aggregated in groups dispersed
location on the host thalli mostly on tops mostly on bases
Size of asci (m) (33–)40–52(–60) × (13–)15–21(–33) (26–)31–41(–55) × (10–)14–20(–21) 47–64(–85) × (14–)17–27(–33)
Ascospores
size (m) (11–)13·5–16·5(–20) × (4·0–)5·0–6·5(–7·5) (8·5–)11·0–13·5(–16·0) × (3·5–)4·5–5·5
(–6·5)
(12·5–)14·0–19·5(–24·0) ×
(4·5–)5–8(–9·0)
septa 1 (0–)1(–3)
halo absent sometimes present
Malformations of host thalli sometimes present absent
166 THE LICHENOLOGIST Vol. 44

Notes. The type variety of the species is
mainly characterized by comparatively
small asci, measuring (26–)31–41(–55)
× (10–)14–20(–21) m(n= 129) and asco-
spores, measuring (8·5–)11·0–13·5(–16·0)
× (3·5–)4·5–5·5(–6·5) m, l/b = (1·8–)2·3–
2·9(–3·9) (n= 502; including free over-
mature spores).
Specimens of Sphaerellothecium thamnoliae var.
thamnoliae examined [all in arctic or alpine tundras or
northern boreal forest on Thamnolia vermicularis var.
subuliformis (16), var. vermicularis (5)]. USA: Alaska:
Great Kobuk Sand Dunes, 67°02–06#N, 158°50#–
159°01#W, 50 m, 2000, M. Zhurbenko (LE 260518, LE
260318, LE 260338, LE 260387).—Canada: Canadian
Arctic Archipelago: Prince Patrick Island, Mould Bay,
76°14#N, 119°18#W, 2004, D. Walker (LE 260388).—
Greenland: Mellem Land: Skovfjorden, Qinngua River
mouth, 61°15#N, 45°30#W, 150 m, 2005, M. Zhur-
benko (LE 260368).—Svalbard: Dickson Land: W coast
of Billefjorden, 78°37#N, 16°20#E, 200 m, 2003,
M. Zhurbenko (LE 260577).—Norway: Troms County:
Skibotndalen Valley, 69°15#N, 20°23#E, 950 m, 2003,
M. Zhurbenko (LE 260478).—Russia: Kola Peninsula:
Barents Sea coast, Voron’ya River mouth, 69°09#N,
35°50#E, 20 m, 1997, M. Zhurbenko (LE 260398).
Northern Ural: headwaters of Pechora River, Yanypu-
puner Range, 62°05#N, 59°06#E, 800 m, 1997, M.
Zhurbenko (LE 260538b). Taimyr Peninsula: Byrranga
Mts., Levinson-Lessinga Lake, 74°30–32#N, 98°30–
33#E, 120–300 m, 1994, M. Zhurbenko (LE 207477a);
ibid., 1995, M. Zhurbenko (LE 260428); Enisei Gulf,
Sibiryakova Island, 72°55#N, 79°00#E, 10 m, 1989, V.
Kuvaev (LE 260558). Yakutiya: Laptev Sea coast, Tiksi,
71°37#N, 128°54#E, 70 m, 1998, M. Zhurbenko (LE
232964, LE 260344b); Lena River delta, Stolb Island,
72°24#N, 126°40#E, 50 m,1998, M. Zhurbenko (LE
260378). Wrangel Island: Klark River, 71°05#N,
178°16#E, 110 m, 1998, S. Kholod (LE 260508);
Neizvestnaya River, 71°14#N, 179°23#W, 130 m,
1987, S. Kholod (LE 260548). Chukotka: Baranikha,
68°30#N, 168°16#E, 1971, I. Makarova (232944b, LE
260599); Yanrakynnot, 64°53#N, 172°30#E, 1976, A.
Sytin (LE 260408).
Additional specimens examined of Sphaerellothecium
thamnoliae var. thamnoliae, formerly published as
Stigmidium frigidum (Zhurbenko & Santesson 1996;
Zhurbenko & Pospelova 2001; Zhurbenko 2008): LE
210405, LE 207538, LE 207540, LE 207541, LE
207543, LE 207544.
Sphaerellothecium thamnoliae var.
taimyricum Zhurb. var. nov.
MycoBank no.: MB 563057
Differt ab varietate Sphaerellothecium thamnoliae var.
thamnoliae ascis et ascosporis majoribus.
Typus: Russia, Taimyr Peninsula, Byrranga Mts., N
of Levinson-Lessinga Lake, 74°34#N, 98°47#E, 250 m
elev., arctic tundra, on thalli of Thamnolia vermicularis
var. subuliformis, 20 August 1995, M. Zhurbenko 95586
(LE 260438—holotypus).
(Fig. 8B & E)
Etymology. The specific epithet refers to
Taimyr Peninsula in Siberia, where the holo-
type was found.
Notes. This variety is mainly distinguished
from the type variety by its larger asci, measur-
ing 47–64(–85) × (14–)17–27(–33) m(n=
15), and ascospores, measuring (12·5–)14·0–
19·5(–24·0) × (4·5–)5–8(–9·0) m, l/b =
(1·7–)2·2–3·0(–3·8) (n= 159). Furthermore,
the variety taimyricum seems to differ from
the variety thamnolia in having a more im-
mersed mycelium, mostly semi-immersed
and slightly larger ascomata, and a better
developed halo around the ascospores. It is
so far also known only from the Arctic.
Specimens of Sphaerellothecium thamnoliae var. tai-
myricum examined (all in polar desert or arctic tundra on
Thamnolia vermicularis var. subuliformis). Russia: Sever-
naya Zemlya: Bolshevik Island, Akhmatova Bay, 79°01#
N, 102°43#E, 60 m, 1996, M. Zhurbenko (LE 232894);
Oktyabr’skoi Revolutsii Island, Cape Nekrasova, 80°02#
N, 99°19#E, 2007, M. Gavrilo (LE 260418). Taimyr
Peninsula: Byrranga Mts., Levinson-Lessinga Lake,
74°24–31#N, 98°36–46#E, 150–300 m, 1995, M.
Zhurbenko (LE 260328, LE 260568); Uboinaya River
mouth, 73°39#N, 82°22#E, 20 m, 1990, M. Zhurbenko
(LE 260503). Novosibirskie Islands: Zhokhova Island,
76°08#N, 152°45#E, 50 m, 1989, M. Samarskii (LE
232974). Wrangel Island: Neozhidannaya River, 71°01#
N, 179°09#E, 100 m, 1992, S. Kholod (LE 260488).
Chukotka: Iskaten’Pass near km 32 Egvekinot−Iul’tin
Highway, 66°35#N, 179°10#W, 1971, I. Makarova (LE
260578).
Additional specimens examined of Sphaerellothecium
thamnoliae var. taimyricum, formerly published as
Stigmidium frigidum (Zhurbenko & Santesson 1996;
Karatygin et al. 1999): LE 207531a, LE 207534, LE
207535, LE 207536, LE 207542, LE 207545, LE
207546.
Stigmidium frigidum (Th. Fr. ex Sacc.)
Alstrup & D. Hawksw.
Meddel. Grønl., Biosc.31: 67 (1990); type: Canada,
British Columbia, Rocky Mountains, Hector, c. 51°25#
N, 116°21#W (I. Brodo, pers. comm.), on Thamnolia
2012 Holarctic thamnoliicolous fungi—Zhurbenko 167

vermicularis var. subuliformis, 27 July 1885, J. Macoun
(E—neotype, photographs examined).
(Fig. 9A & B)
Vegetative hyphae pale to medium brown,
composed of elongate cells 2·5–7·0 m wide,
septate, constricted at the septa, scarcely
branched, BCr+ blue-green, I−, immersed to
occasionally superficial, rather sparse, usu-
ally inconspicuous.
Ascomata perithecioid, black, subglobose,
(30–)50–80(–120) m diam., sometimes
with an indistinct papilla, with an ostiole
of 10–20 m diam., initially completely
immersed (sometimes in a few layers), then
mostly protruding in the ostiolar area or
F. 9.Stigmidium frigidum. A, infection habitus (LE 260444); B, ascomata (LE 232994); C, asci with spores in
water, note dark over-mature ascospores (LE 260354). Scales:A=1mm;B=200m;C=10m.
168 THE LICHENOLOGIST Vol. 44

occasionally up to half erumpent, arising sin-
gly or sometimes adjacent or a few confluent,
aggregated in groups of up to 500. Exciple in
surface view brown, smooth, similar to
textura angularis, composed of a few layers of
thick-walled, angular, pseudoparenchyma-
tous cells (3–)5–8(–12) m across, BCr+
blue-green, K+ olive. Hymenial gel I−, K/I−.
Hamathecium not observed. Asci elliptic to
pyriform, endoascus thickened above, some-
times with a small ocular chamber, foot not
observed, (33–)40–52(–60) × (13–)15–21
(–33) m(n= 62), 8-spored, BCr−, I−, K/I−
(except plasma reacting BCr+ blue, I+ and
K/I+ yellow to orange). Ascospores colourless
or occasionally pale to medium brown (when
over-mature ?) (Fig. 9C), obskittle-shaped
to narrowly obovate, usually with a wider
and shorter upper cell, occasionally located
upside down in the ascus, (11–)13·5–16·5
(–20) × (4·0–)5·0–6·5(–7·5) m, l/b = (1·7–)
2·3–2·9(–4·1) (n= 214), 1-septate, usually
constricted at the septum, smooth-walled,
halo not observed, usually with a few big
guttules in each cell, irregularly diagonally
arranged in 2–3 rows in the ascus, wall and
septum BCr−, plasma BCr+ blue.
Anamorph not observed.
Notes. The species was originally described
from Greenland as “Sphaeria n. 10” by Fries
(1879: 370) and later formally introduced as
“Epicymatia frigida Sacc. ex Fr. Hedw. 1881,
p. 60 (Sphaeria n. 10)” (Saccardo 1882:
572). According to this description, the type
specimen had dark torulose superficial hy-
phae and ascospores of 15–18 × 5–7 m, and
thus may well represent Sphaerellothecium
thamnoliae var. taimyricum. According to
Alstrup & Hawksworth (1990: 67), this
original material was lost and the authors
neotypified the species on John Macoun’s
“Canadian Lichens” no. 180 (E—neotype)
and combined it into Stigmidium. The local-
ity on the neotype label (“Hector, B.C.”) is
indistinctly handwritten, which was evi-
dently the reason for erroneously referring
it to Alaska (Hawksworth 1980: 180). The
examination of the neotype by R. Yahr
(pers. comm.), together with the notes of
Hawksworth (op. cit.), leave little doubt that
it fits the species concept presented here (see
below). This concept also corresponds to the
description of Stigmidium frigidum in Ihlen
(1995). The only deviating character of the
species reported by the latter author is the
4(–6)-spored asci, not typical for Stigmidium
in general and not observed in the examined
material.
Usually growing on the upper parts or even
tips of Thamnolia vermicularis thalli, but
occasionally observed on their middle or
basal parts. Evidently pathogenic, as causing
bleaching or even destruction of the host
tissues; heavy infections also induce various
malformations of the host thalli which, for
example, become bent, twisted and/or some-
what swollen (Fig. 9A). Superficially the in-
fections are similar to those of Polycoccum
vermicularium.
Typical Stigmidium frigidum can be readily
distinguished from Sphaerellothecium thamno-
liae, even with a dissecting microscope, by its
numerous, more or less immersed ascomata,
crowded at the tips of the host thalli, and the
absence of a visible mycelium; malforma-
tions of the host thalli are also diagnostic
(Table 2). Confusion can be caused by the
occasional development of superficial veg-
etative hyphae. However, in well-developed
specimens it is clearly seen that such hyphae
are associated only with some groups of
ascomata and not forming a reticulum.
New to Mongolia; a first verified report
from the USA.
Specimens examined [all in polar deserts, arctic or
alpine tundras on Thamnolia vermicularis var. subuliformis
(13), var. vermicularis (8)]. USA: Alaska: Brooks Range,
headwaters of Atigun River near mile 248 Dalton High-
way, 68°07#N, 149°28#W, 1400 m, 2001, M. Zhur-
benko (LE 260356); Kotzebue, 66°53#N, 162°31#W,
30 m, 2000, M. Zhurbenko (LE 260406).—Russia:
Severnaya Zemlya: Bolshevik Island, Mikoyana Bay,
79°18#N, 101°55#E, 10 m, 1996, M. Zhurbenko (LE
232943). Taimyr Peninsula: Byrranga Mts., Bolshaya
Bootankaga River, 74°30#N, 97°40#E, 350 m, 1995,
M. Zhurbenko (LE 232994a); ibid., Levinson-Lessinga
Lake, 74°24–34#N, 98°26–47#E, 60–300 m, 1994, M.
Zhurbenko (LE 207612); ibid., 1995, M. Zhurbenko (LE
232984, LE 232923, LE 232823, LE 232863, LE
260523, LE 260334, LE 260564, LE 260354); Dikson
Island, 73°30#N, 80°20#E, 30 m, 1990, M. Zhurbenko
(LE 260353); Ragozinka River mouth, 72°48#N,
80°53#E, 30 m, 1990, M. Zhurbenko (LE 260413).
Yakutiya: 60 km E of Cherskii, Belaya Strelka Mts.,
2012 Holarctic thamnoliicolous fungi—Zhurbenko 169

68°40#N, 162°40#E, 400 m, 1975, A. Egorova (LE
260454); Laptev Sea coast, Tiksi, 71°39–40#N,
128°40–45#E, 50–70 m, 1998, M. Zhurbenko (LE
260484, LE 260444a). Chukotka: Baranikha, 68°30#N,
168°16#E, 1971, I. Makarova (LE 233084);
Gil’mymlineiveem River, 65°48#N, 173°15#W, 1977,
I. Makarova (LE 232953).—Mongolia: Dzabkhan
Region: Mt. Otgon-Tenger, 47°35#N, 97°32#E,
3250 m, 1976, L. Biazrov 7513 (LE 260384).
Additional examined specimens of Stigmidium frigidum
published in Zhurbenko & Santesson (1996): LE 207533,
LE 207537, LE 207539.
Thamnogalla crombiei (Mudd) D.
Hawksw.
Notes Roy. Bot. Gard. Edinburgh 38(1):178 (1980); type:
Scotland, Ben Lawers, on Thamnolia vermicularis, 1864,
Crombie (BM—lectotype, n. v.).
Notes. Ascomata perithecioid, 75–125 m
diam., immersed or rarely erumpent. Exciple
in cross-section orange-brown throughout,
K+ becoming more intensively red-orange.
Hamathecium of filiform, non-capitate, c.1m
wide, septate, scarcely branched interascal
filaments. Asci cylindrical, with a long foot,
(50–)70–90 × 5–6 m(n= 17), 8-spored.
Ascospores hyaline, mostly narrowly elliptic to
narrowly oblong, with rounded ends, (7–)8–
11·5(–15) × (2–)2·5–3·0(–4) m, l/b = (2·1–
)2·8–4·0(–5·2) (n= 113) [(7–)9·6(–12) ×
(2·5–)3·7(–5·0) m, l/b = 2·6 according to
Hoffmann & Hafellner (2000)], non-septate,
smooth-walled, non-halonate, often with a
few guttules, arranged uniseriate and over-
lapping to biseriate in the ascus. Inducing
concolorous or pale coloured (cinnamon,
pinkish buff, peach, flesh pink, or grey),
finally bullate and basally constricted galls
throughout the host thalli. Otherwise patho-
genicity not observed.
This is a frequent and cosmopolitan tham-
noliicolous fungus yet is reported here for the
first time from Greenland and Svalbard.
Specimens examined [all in polar deserts, arctic or
alpine tundras, or northern boreal forest on Thamnolia
vermicularis var. subuliformis (26), var. vermicularis
(14)]. USA: Alaska: Kotzebue, 66°53#N, 162°31#W,
30 m, 2000, M. Zhurbenko (LE 260386, LE 260493);
ibid., 1961, B. Neiland (LE 260466); Great Kobuk
Sand Dunes, 67°06#N, 159°01#W, 50 m, 2000, M.
Zhurbenko (LE 260336); Seward Peninsula, Bering
Land Bridge National Preserve, 65°50#N, 164°32#W,
612 m, 2002, J. Roth (LE 260423); ibid., 66°17#N,
165°19#W, 22 m, 2002, J. Roth (LE 260383, LE
260433).—Canada: British Columbia: Wells Gray
Provincial Park, Mt. Raft, 51°44#N, 119°50#W,
2100 m, 2002, M. Zhurbenko (LE 260456).—
Greenland: Mellem Land, Skovfjorden, Qinngua
River mouth, 61°15#N, 45°30#W, 150 m, 2005, M.
Zhurbenko (LE 260370); Tugtugtooq Island SW of
Narssaq, 2 km SW of Blue Moon Lake, 60°51#N,
46°25#W, 15 m, 2005, M. Zhurbenko (LE 260340, LE
260330).—Svalbard: Bünsow Land: Norddammen
Lake, 78°38#N, 16°44#E, 10 m, 2003, M. Zhurbenko
(LE 260350).—Norway: Troms County: Skibotndalen
Valley, 69°13–19#N, 20°21–29#E, 70–650 m, 2003,
M. Zhurbenko (LE 260416, LE 260366).—Russia:
Kola Peninsula: Barents Sea coast, Voron’ya River
mouth, 69°09#N, 35°50#E, 20 m, 1997, M. Zhur-
benko (LE 260534); ibid., 4 km SSE of Dal’nie Ze-
lentsy, 69°05#N, 36°07#E, 100 m, 1997, M.
Zhurbenko (LE 260594); ibid., 17 km N of Tumannyi,
69°01#N, 35°48#E, 100 m, 1997, M. Zhurbenko (LE
260584); Khibiny Mts., headwaters of Kaskasnyunjok
Creek, 67°46#N, 33°49#E, 2007, M. Zhurbenko (LE
233042). Nenets Region: Dolgii Island, 69°18#N,
58°52#E, 1998, V. Shevchenko (LE 232824). Northern
Ural: headwaters of Pechora River, Yanypupuner
Range, Mt. “981”, 62°05#N, 59°06#E, 800 m, 1997,
M. Zhurbenko (LE 260404, LE 260504). Western
Sayan Mts.: Kulumys Range, 52°51#N, 93°17#E,
1700 m, 2010, M. Zhurbenko (LE 260310); Oiskii
Range, headwaters of Olen’ya River, 52°48#N, 93°15#
E, 1650 m, 2010, M. Zhurbenko (LE 260380b). East-
ern Sayan Mts.: Kryzhina Range, headwaters of Belyi
Kitat River, 54°01#N, 95°27#E, 1800 m, 2009, M.
Zhurbenko (LE 260403a). Severnaya Zemlya: Bolshevik
Island, Akhmatova Bay, 79°04#N, 102°45#E, 10–
20 m, 1996, M. Zhurbenko (LE 233064, LE 260314,
LE 260514); ibid., Mikoyana Bay, 79°18#N, 101°55#
E, 10 m, 1996, M. Zhurbenko (LE 232904); ibid.,
Shokalskogo Strait, 79°18#N, 101°40#E, 20 m, 1996,
M. Zhurbenko (LE 232834). Taimyr Peninsula: Byr-
ranga Mts., Levinson-Lessinga Lake, 74°33#N,
98°43#E, 100 m, 1995, M. Zhurbenko (LE 260434);
Ragozinka River mouth, 72°48#N, 80°53#E, 25 m,
1990, M. Zhurbenko (LE 260513); Uboinaya River
mouth, 73°36#N, 82°22#E, 10 m, 1990, M. Zhur-
benko (LE 260463). Yakutiya: Lena River delta, 3 km
E of Cape Krest-Tumsa, 72°22#N, 126°42#E, 50 m,
1998, M. Zhurbenko (LE 233014). Magadan Region:
Magadan, Mt. Nagaevskaya, 59°34#N, 150°45#E,
400 m, 2003, N. Sazanova (LE 260443). Wrangel Is-
land: Gusinaya River, 71°08#N, 179°10#E, 1991, V.
Shtrik (LE 232982); Naskhok River, 71°24#N,
178°08#W, 5 m, 1998, S. Kholod (LE 232932); Ne-
ozhidannaya River, 71°02#N, 179°10#E, 110 m,
1992, S. Kholod (LE 260374b); Tundrovaya River,
71°30#N, 179°45#W, 1996, S. Kholod (LE 233004).
Chukotka: Baranikha, 68°30#N, 168°16#E, 1971, I.
Makarova (LE 232944a); Chaplinskie hot springs,
64°25#N, 172°30#W, 1957, V. Gavrilyuk & P.
Gagarin (LE 232843).
170 THE LICHENOLOGIST Vol. 44

Discussion
To date, 23 species representing 18 genera of
lichenicolous fungi are known on Thamnolia
worldwide, of which 18 species (see key be-
low; c. 80%) and two genera (Epithamnolia
and Thamnogalla D. Hawksw.; c. 10%) have
been reported only from this host genus. All
these species inhabit Thamnolia vermicularis,
six of them (Endococcus thamnoliae,Lichen-
opeltella thamnoliae,Odontotrema santessonii,
Polycoccum vermicularium,Stigmidium frigi-
dum, and Thamnogalla crombiei) are also
known on T. papelillo (Etayo 2010), and none
has yet been found on T. juncea. Of the 190
newly studied fungal specimens growing on
Thamnolia vermicularis, 144 (76%) were
found on its var. subuliformis and 46 (24%) on
var. vermicularis. This disproportion might
merely reflect the fact that the former variety
is more abundant in the Northern Hemi-
sphere than the latter (Thomson 1984).
The genus Thamnolia has no species of
lichenicolous fungi in common with the
other genera of the lichen family Icma-
dophilaceae and is much more ‘hospitable’
compared to other genera in the family:
Dibaeis (hosts 5 species of lichenicolous
fungi), Icmadophila (5), Pseudobaeomyces (0),
Siphula (1 ?), Siphulella (0) (author’s data-
base).
In the richness of lichenicolous mycobiota
which it supports, Thamnolia vermicularis
ranks 15th worldwide after Peltigera rufescens
(hosts 54 species of lichenicolous fungi), Cla-
donia pyxidata (L.) Hoffm. (35), Xanthoria
parietina (L.) Th. Fr. (34), Peltigera didactyla
(With.) J.R. Laundon (32), P. canina (L.)
Willd. (30), P. praetextata (Flörke ex
Sommerf.) Zopf (29), Lobaria pulmonaria
(L.) Hoffm. (28), Parmelia saxatilis (L.) Ach.
(28), P. sulcata Taylor (27), Cladonia pocillum
(Ach.) Grognot (26), Peltigera polydactylon
(Necker) Hoffm. (26), P. aphthosa (L.)
Willd. (25), Physcia stellaris (L.) Nyl. (25),
and Circinaria calcarea (L.) A. Nordin, S.
Savic´ & Tibell (24) (author’s database). It is
noteworthy that 14 of these 15 most hospita-
ble lichen species of the world occur in the
Arctic, and at least 11 of them thrive there
(Kristinsson et al. 2010).
The following taxa of Thamnolia-
inhabiting fungi were the most common in
the Holarctic: Thamnogalla crombiei (21%
of 190 specimens examined), Polycoccum ver-
micularium (15%), Sphaerellothecium tham-
noliae var. thamnoliae (12%), Stigmidium
frigidum (11%), Cercidospora thamnoliae
(9%), and Odontotrema santessonii (5%).
The study ends in formally describing six,
and introducing another two, new species of
thamnoliicolous fungi, which is about 35%
of their known total number. However, some
of their specimens still remain unnamed.
Among these are a Lasiosphaeriopsis-like
pyrenomycete with brown, 4 − 6-septate
ascospores, c. 33–38 × 10–13 m (1996, M.
Zhurbenko 96910); a fungus with a closed
fruit body containing filiform, hyaline, 4–5-
septate diaspores, c. 58–69 × 1·5–2 m
(1998, S. Kholod); a gall-inducing hyphomy-
cete (1989, A. Katenin); and a rather com-
mon fungus with thick dark superficial
hyphal strands and conspicuous black dis-
coid sterile fruit-like structures, resembling
species of Lichenostigma Hafellner subgenus
Lichenogramma Nav.-Ros. & Hafellner. One
more unidentified coelomycete on Tham-
nolia with pleurogenous conidiogenous cells
was mentioned by Hawksworth (1980: 180).
Four of 21 named Thamnolia-dwelling fungal
species found in the Holarctic were repre-
sented by single finds. According to a Turing
estimator, this suggests that about 80% of
these fungi were detected during the survey,
or that their real diversity in the Holarctic is
no less than 26 species.
Three of the 23 fungal species that have
been reported from Thamnolia were not
found in the Holarctic lichen collections
examined: Cornutispora intermedia Punith. &
D. Hawksw., Endococcus thamnoliae Etayo
& R. Sant., and Odontotrema intermedium
Diederich, Zhurb. & Etayo (Diederich et al.
2002; Etayo 2010); all are known only
from a single or a few finds, which makes it
futile to speculate about their geography.
On the other hand a fourth, Lichenopeltella
thamnoliae, seems to be rather common in
South America (Etayo 2010) and thus its
absence in the Holarctic might not be
accidental. More observations from the
2012 Holarctic thamnoliicolous fungi—Zhurbenko 171

extra-holarctic regions are needed to answer
questions about the distribution patterns of
thamnoliicolous fungi and their infrageneric
preferences within Thamnolia.
Key to the lichenicolous fungi growing on Thamnolia
Based on data in Hawksworth (1977), Santesson (1998), Diederich et al. (2002, 2010),
Punithalingam (2003), Heuchert & Braun (2006), Etayo (2010), and the present publica-
tion. Species of lichenicolous fungi not specific to Thamnolia are given in parentheses.
1 Anamorphic fungus .................................2
Teleomorphic fungus ................................8
2(1) Hyphomycete, conidiophores usually with divergent terminal branches, conidio-
genous cells integrated, mostly terminal, with a single or few distant enteroblastic-
percurrent proliferations, conidia pale brown or yellowish brown, usually in
branched acropetal chains, subglobose or limoniform to ellipsoid-subcylindrical,
aseptate (3·5–8·0 × 3–5 m) or 1-septate (7–13 × 5–7 m)...........
.........................(Cladosporium licheniphilum)
Coelomycete .....................................3
3(2) Conidia dark brown, subglobose, aseptate, (2·5–)3·0–4·0(–5·0) m diam.,
ornamented (× 1000), pycnidia (40–)50–80(–110) m diam. .........
............................ (Lichenoconium usneae)
Conidia hyaline ...................................4
4(3) Conidia with (1–)3(–4) divergent arms ....... (Cornutispora intermedia)
Conidia without arms ................................5
5(4) Conidia (0–)1(–2?)-septate, cylindrical, (14–)18·5–27·0(–32) × (1·0–)1·5–2·0
(–2·5) m...................... Epithamnolia karatyginii
Conidia aseptate ...................................6
6(5) Conidia 2·0–4·5 × 0·5 m, bacilliform .......................
................. anamorph of Sphaerellothecium thamnoliae
Conidia over 0·5 m wide ..............................7
7(6) Conidia (3·0–)4–6(–7·5) × (1–)1·5 m, oblong ..................
................... anamorph of Polycoccum vermicularium
Conidia (3·5–)4–5(–5·5) × (1·5–)2·0(–2·5) m, oblong to occasionally elliptic . .
............................Phoma thamnoliae ad int.
8(1) Hymenium at least partly exposed at maturity, ascospores hyaline ........9
Hymenium not exposed at maturity........................ 12
9(8) Ascospores aseptate, subglobose to shortly ellipsoid, 6–10 × 5·0–7·5m.....
...............................(Geltingia associata)
Ascospores septate, ascomata urceolate ......................10
10(9) Ascospores with (3–)6–8(–9) transverse septa and often a few longitudinal or oblique
septa in central cells, (12·5–)18–23(–25·0) × (3–)3·5–4·5(–5) m .......
............................ Odontotrema santessonii
Ascospores with less septa, never submuriform..................11
172 THE LICHENOLOGIST Vol. 44

11(10)Ascospores 1(–2)-septate, (9–)11–15(–18) × 2·5–3·0(–3·5)m..........
............................Odontotrema thamnoliae
Ascospores 3-septate, (10·0–)10·5–12·5(–13·5) × (3–)3·5–4·0(–4·5)m .....
...........................Odontotrema intermedium
12(8) Ascomata catathecia, non-setose, asci 4-spored, ascospores hyaline, 1-septate, 11·5–
16 × 2·5–5 m, without setulae..........Lichenopeltella thamnoliae
Ascomata perithecioid, ascospores hyaline or pigmented.............13
13(12)Ascomata setose, ascospores initially hyaline, then pale to medium grey, with (0–)3–
5(–7) transverse septa and usually longitudinal or oblique septa in central cells,
(11 –)13·5–18·0(–29) × (5–)6–8(–12) m....... Capronia thamnoliae
Ascomata without setae ..............................14
14(13)Ascospores pigmented, at least when mature ...................15
Ascospores not or only occasionally pigmented.................. 19
15(14)Ascospores muriform ............................... 16
Ascospores only transversely septate .......................17
16(15)Interascal filaments present, ascospores ornamented (×1000), (23–)25–29
(–31·5) × (10·5–)11–13(–14) m...... Dacampia thamnoliicola ad int.
Interascal filaments absent, ascospores smooth-walled, (14–)16·5–24·0(–34) × (6–)
6·5–8·5(–9) m................. Merismatium thamnoliicola
17(15)Ascospores (1–)3-septate, hyaline to pale cinnamon-brown, (10–)15·5–20·5(–22)
× (4·5–)5–6(–8·0) m...................(Phaeospora arctica)
Ascospores (0–)1-septate ............................. 18
18(17)Interascal filaments absent, ascospores 9·0–12·5 × 3·5–5·0 m..........
.............................Endococcus thamnoliae
Interascal filaments well developed, ascospores (13–)15·5–20·5(–28) × (6–)7–9
(–12) m..................... Polycoccum vermicularium
19(14)Ascomata immersed in distinct bullate galls ................... 20
Ascomata not associated with bullate galls ....................21
20(19)Exciple orange-brown throughout, interascal filaments well developed, ascospores
aseptate, (7–)8·0–11·5(–15) × (2–)2·5–3·0(–4) m, without halo........
............................. Thamnogalla crombiei
Exciple bluish grey, interascal filaments sparse to indistinct, ascospores 1-septate,
(12–)14–17(–19) × (3·5–)4–5(–5·5) m, halonate ................
.........................Cercidospora thamnogalloides
21(19)
Exciple bluish green or olive-brown above, hyaline below, vegetative hyphae incon-
spicuous, ascomata subimmersed, interascal filaments indistinct or abundant . 22
Exciple brown throughout, vegetative hyphae conspicuous or not, ascomata sub-
immersed to sessile, interascal filaments absent ................24
22(21)Exciple olive-brown above, asci mainly 4-spored, interascal filaments sparse to
indistinct, ascospores (11–)14·0–17·5(–21) × (4·0–)4·5–5·5(–6·5) m, (1–)2–3-
septate, without halo .............. Cercidospora thamnoliicola
Exciple bluish green above, asci only/mainly 8-spored .............. 23
2012 Holarctic thamnoliicolous fungi—Zhurbenko 173

23(22)Interascal filaments abundant, ascospores (1–)3(–4)-septate, (12·5–)16·5–21·0
(–27·5) × (4–)4·5–5·5(–8) m, occasionally halonate ..............
............................Cercidospora thamnoliae
Interascal filaments sparse to indistinct, ascospores (0–)1-septate, (12–)13–16
(–19) × (3–)3·5–4·5(–5) m, without halo ....Cercidospora epithamnolia
24(21)Ascospores (1–)3-septate, hyaline to pale cinnamon-brown, (10–)15·5–20·5
(–22) × (4·5–)5·0–6·0(–8·0) m .............(Phaeospora arctica)
Ascospores only or mainly 1-septate .......................25
25(24)Vegetative hyphae usually inconspicuous, ascomata (30–)50–80(–120) m diam.,
mostly subimmersed and aggregated on tips of host thalli, ascospores 1-septate,
(11–)13·5–16·5(–20) × (4·0–)5·0–6·5(–7·5) m, without halo .........
.............................. Stigmidium frigidum
Vegetative hyphae forming a conspicuous dark reticulum, ascomata (30–)50(–80)
m diam., semi-immersed to sessile, dispersed, usually on bases of host thalli,
ascospores (0–)1(–3)-septate, sometimes halonate ...............26
26(25)Asci (26–)31–41(–55) × (10–)14–20(–21) m, ascospores (8·5–)11·0–13·5(–16·0)
× (3·5–)4·5–5·5(–6·5) m.............................
...............Sphaerellothecium thamnoliae var. thamnoliae
Asci 47–64(–85) × (14–)17–27(–33) m, ascospores (12·5–)14·0–19·5(–24·0)
× (4·5–)5– 8(–9) m.. Sphaerellothecium thamnoliae var. taimyricum
I am indebted to Irina Makarova, Sergei Kholod, Nadya
Matveeva, Adrian Katenin, J. Roth, Mikhail Samarskii,
Vladimir Shevchenko, Olga Lavrinenko, Nina
Sazanova, Thomas Lunke, Arve Elvebakk, Donald
Walker and Lev Biazrov for putting their undetermined
lichen collections at my disposal. Wolfgang von Brackel,
Paul Diederich, Dagmar Triebel, Karen Dillman,
Vadim Mel’nik, Ernie Brodo, and Brian Coppins are
thanked for their various help, as well as Uwe Braun for
the revision of Cladosporium licheniphilum and Rebecca
Yahr and Louise Olley for the examination of the Stig-
midium frigidum neotype.
R
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