Lichenicolous Fungi from Northwest Caucasus, Russia

Abstract

Zhurbenko, M. P. & Kobzeva, A. A. 2014. Lichenicolous fungi from Northwest Caucasus, Russia. — Herzogia 27: 377–396. One hundred species of lichenicolous fungi are reported from Northwest Caucasus; 64 of them are new to the Caucasus. Hainesia aeruginascens, H. brevicladoniae, Minutoexcipula calatayudii, Neobarya peltigerae, Opegrapha anomea, O. rotunda, Polycoccum arnoldii, P. kaernefeltii, Pronectria echinulata and P. santessonii are newly reported for Asia and Russia. Abrothallus cetrariae, Pyrenochaeta xanthoriae, Taeniolella delicata, Trichonectria anisospora and Zwackhiomyces echinulatus are new to Asia, but not Russia. Cercidospora melanophthalmae, Lichenostigma rupicolae, Marchandiobasidium aurantiacum and Stigmidium xanthoparmeliarum are new to Russia, but not Asia. Endococcus incrassatus and Xanthoriicola physciae are new to Asian Russia. Lichenochora rinodinae is first reported from outside the Arctic. Finds of specimens presumably belonging to the insufficiently known species Merismatium cladoniicola, Phacopsis usneae and Sphaerellothecium gowardii are discussed. Dactylospora deminuta is newly reported on Gyalecta foveolaris and Fuscopannaria praetermissa, Neobarya peltigerae on Peltigera polydactylon and Taeniolella delicata on Lecanora intumescens. Nephroma is a new host genus for Paranectria oropensis and Caloplaca for Polycoccum kaernefeltii.

Full text

Herzogia 27 (2), 2014: 377–396 377
Lichenicolous fungi from Northwest Caucasus, Russia
Mikhail P. Zhurbenko* & Anastasiya A. Kobzeva
Abstract: Zhurbenko, M. P. & Kobzeva, A. A. 2014. Lichenicolous fungi from Northwest Caucasus, Russia. –
Herzogia 27: 377−396.
One hundred species of lichenicolous fungi are reported from Northwest Caucasus; 64 of them are new to the
Caucasus. Hainesia aeruginascens, H. brevicladoniae, Minutoexcipula calatayudii, Neobarya peltigerae, Opegrapha
anomea, O. rotunda, Polycoccum arnoldii, P. kaernefeltii, Pronectria echinulata and P. santessonii are newly reported
for Asia and Russia. Abrothallus cetrariae, Pyrenochaeta xanthoriae, Taeniolella delicata, Trichonectria anisospora
and Zwackhiomyces echinulatus are new to Asia, but not Russia. Cercidospora melanophthalmae, Lichenostigma
rupicolae, Marchandiobasidium aurantiacum and Stigmidium xanthoparmeliarum are new to Russia, but not Asia.
Endococcus incrassatus and Xanthoriicola physciae are new to Asian Russia. Lichenochora rinodinae is first reported
from outside the Arctic. Finds of specimens presumably belonging to the insufficiently known species Merismatium
cladoniicola, Phacopsis usneae and Sphaerellothecium gowardii are discussed. Dactylospora deminuta is newly
reported on Gyalecta foveolaris and Fuscopannaria praetermissa, Neobarya peltigerae on Peltigera polydactylon
and Taeniolella delicata on Lecanora intumescens. Nephroma is a new host genus for Paranectria oropensis and
Caloplaca for Polycoccum kaernefeltii.
Zusammenfassung: Zhurbenko, M. P. & Kobzeva, A. A. 2014. Lichenicole Pilze aus dem Nordwestkaukasus,
Russland. – Herzogia 27: 377−396.
Einhundert Arten lichenicoler Pilze werden aus dem Nordwestkaukasus gemeldet, von denen 64 neu für den
Kaukasus sind. Hainesia aeruginascens, H. brevicladoniae, Minutoexcipula calatayudii, Neobarya peltigerae,
Opegrapha anomea, O. rotunda, Polycoccum arnoldii, P. kaernefeltii, Pronectria echinulata und P. santessonii sind
neu für Asien und Russland. Abrothallus cetrariae, Pyrenochaeta xanthoriae, Taeniolella delicata, Trichonectria
anisospora und Zwackhiomyces echinulatus sind neu für Asien, aber nicht für Russland. Cercidospora melanoph-
thalmae, Lichenostigma rupicolae, Marchandiobasidium aurantiacum und Stigmidium xanthoparmeliarum sind
neu für Russland, aber nicht für Asien. Endococcus incrassatus und Xanthoriicola physciae sind neu für den asia-
tischen Teil Russlands. Lichenochora rinodinae wurde erstmals außerhalb der Arktis gefunden. Diskutiert werden
Funde, die wahrscheinlich zu den ungenügend bekannten Arten Merismatium cladoniicola, Phacopsis usneae und
Sphaerellothecium gowardii gehören. Dactylospora deminuta wird neu auf Gyalecta foveolaris und Fuscopannaria
praetermissa dokumentiert, Neobarya peltigerae auf Peltigera polydactylon und Taeniolella delicata auf Lecanora
intumescens. Nephroma ist eine neue Wirtsgattung für Paranectria oropensis und Caloplaca für Polycoccum kaer-
nefeltii.
Key words: Biodiversity, biogeography, taxonomy, ecology, Asia.
Introduction
So far data on lichenicolous fungi of the Caucasus have been obtained mainly in the course
of floristic lichen studies. They have recently been summarized in a first synopsis comprising
72 species (Zhurbenko & Otte 2012). Later on Urbanavichus & Ismailov (2013) and
* Corresponding author

378 Herzogia 27 (2), 2014
Calatayud et al. (2013) reported another 23 species new to the region. The present paper is
based on the first special field survey of lichenicolous fungi, resulting in many biogeographic
novelties, including 64 species newly reported for the Caucasus. However, the currently re-
vealed richness of regional lichenicolous mycota (160 species) still seems to remain far from
the expected real one, as compared to those of the better studied European mountainous sys-
tems. Pertinent notes on taxonomy, geographical distribution and hosts of insufficiently known
species are provided.
Material and methods
This study is based on 199 specimens of lichenicolous fungi collected by M. P. Zhurbenko,
mainly in or nearby the Teberda State Biosphere Reserve in August 2012. All specimens were
identified or revised by the first author using Zeiss microscopes Stemi 2000−CS and Axio Imager
A1 equipped with Nomarski differential interference contrast optics (DIC). The examination
was done in water, 10 % KOH (K), Lugol’s iodine, directly (I) or after a KOH pre-treatment
(K/I), 10 % HNO3 (N), or brilliant cresyl blue (BCr). The length, breadth and length/breadth
ratio (l/b) of asci, ascospores and conidia are given as: (min−){X−SD}−{X+SD}(−max),
where min and max are the extreme values, X the arithmetic mean and SD the corresponding
standard deviation. Measurements were taken from water mounts, unless otherwise indicated.
The nomenclature of the host lichens mostly follows Blinkova et al. (2004). Considering the
geographic distribution of the treated species, we follow view disposing the Caucasus in Asia.
Examined specimens are housed in the mycological herbarium of the V.L. Komarov Botanical
Institute in St.-Petersburg (LE-Fungi).
Collecting localities in Northwest Caucasus, Russia
Karachaevo-Cherkesiya Republic
1: 3 km NE of Dombai settlement, 43°18'N/41°38'E, alt. 1700 m, Picea-Abies-Fagus-Betula forest.
2: 10 km E of Dombai settlement, right bank of Gonachkhir River, 43°18' N/41°42'E, alt. 1600−1700 m, Picea-
Abies-Fagus-Betula forest.
3: Teberda State Biosphere Reserve, 5 km W of Teberda town, NE slope of Mt. Khatipara, 43°27'N/41°41'E, alt.
2800 m, mountain tundra and rocks.
4: Teberda State Biosphere Reserve, near confluence of Baduk and Teberda Rivers, 43°22'33''N/41°41'35''E, alt.
1450 m, Picea-Abies-Fagus forest.
5: 1 km W of Dombai settlement, along road to Alibek mountaineering camp, 43°17'52''N/41°35'37''E, alt. 1800 m,
Picea-Abies-Fagus forest.
6: 6 km E of Teberda town, Dzhemagatskoe Canyon, right bank of Goralykol River, 43°27'13''N/41°49'11''E, alt.
2130 m, mixed forest and subalpine meadow with boulders.
7: Teberda town, vicinities of Teberda State Biosphere Reserve office, 43°27'N/41°44'E, alt. 1400 m, Picea-Abies-
Fagus forest.
8: vicinities of Dombai settlement, top of Mt. Musat-Cheri, 43°17'22''N/41°40'48''E, alt. 3000 m, mountain tundra
and rocks.
9: vicinities of Teberda town, 43°27'N/41°44'E, alt. 1400 m, Pinus forest and pasture.
10: Teberda town, left bank of Teberda River at 1 km upstream of Teberda State Biosphere Reserve office,
43°26'41''N/41°44'06''E, alt. 1350 m, on freestanding deciduous trees in a pasture.
11: Teberda town, left bank of Teberda River at 1.5 km upstream of Teberda State Biosphere Reserve office,
43°25'28''N/41°43'50''E, alt. 1350 m, on boulders in sparse mixed forest.
12: slope of Mt. Malaya Khatipara above Teberda State Biosphere Reserve office, 43°26'08''N/41°43'15''E, alt. 2000
m, Pinus forest and shaded rocks.

Zhurbenko & Kobzeva: Lichenicolous fungi from Northwest Caucasus, Russia 379
13: Teberda town, vicinities of Teberda State Biosphere Reserve office, 43°27'N/41°44'E, alt. 1400 m, Picea-Abies
forest.
14: vicinities of Teberda town, output of Dzhemagatskoe Canyon, 43°28'02''N/41°46'54''E, alt. 1450 m, pasture,
mixed forest and boulders.
15: 5 km S of Teberda town, right bank of Ulu-Murudzhu River, 43°23'14''N/41°42'20''E, alt. 1450 m, Picea-Abies forest.
16: 2 km W of Teberda town, road to Mukhinskii pass along Mukha River, 43°28'10''N/41°42'46''E, alt. 1500 m,
mixed forest.
Stavropol’skii Territory
17: Shadzhatmaz Plateau, 18 km S of Kislovodsk town, 43°44'N/42°39'E, alt. 2100 m, alpine meadows and limestone
cliffs.
18: Pyatigorsk town, foot of Mt. Mashuk, 44°03'48''N/43°05'44''E, alt. 590 m, on deciduous trees in mixed forest.
The species
Fungus species new to the Caucasus are denoted by an asterisk “*”. The collector M. P.
Zhurbenko is abbreviated in the list as MZ.
Findings of some host lichens are also noteworthy. Placidiopsis cinerascens has not been
reported from Russia according to Urbanavichus (2010). Bacidia rubella, Biatora subdu-
plex, Bryoria capillaris, Flavopunctelia flaventior, Gyalecta foveolaris, Melanelia tominii and
Melanelixia fuliginosa, as well as indistinctly lichenized Thelenella muscorum (6: on mosses
above boulder by a river, 22.08.2012, MZ 1218b, LE 261372b), are new to Teberda State
Biosphere Reserve according to Blinkova et al. (2004).
Abrothallus bertianus De Not.
4: on Melanelixia fuliginosa (thallus), 20.08.2012, MZ 12168, LE 261436.
Apothecia superficial, usually applanate, strongly constricted at the base, 150 –250 µm diam.,
occasionally slightly greenish pruinose. Epithecium violet, with orange brown crystals of pru-
ina, K+ blue green. Hymenium violet above, colourless below, K+ blue green. Hypothecium me-
dium olive brown. Ascospores pale to medium olive brown, granulose, (10.4−)10.8−12.4(−13.8) ×
(4.2−)4.4−5.6(−6.2) µm, l/b = (1.9−)2.1−2.5(−2.8) (n = 36). Vouauxiomyces anamorph with more or
less obpyriform smooth conidia with rounded apex and truncated base, measuring (5.2−)5.6−7.0(−7.9)
× (3.5−)3.9−4.5(−5.0) µm, l/b = (1.2−)1.3−1.7(−1.9) (n = 36).
*Abrothallus caerulescens I.Kotte
6: on Xanthoparmelia stenophylla (discs of apothecia, thallus), 22.08.2012, MZ 1249, LE 261451; 11: on X. steno-
phylla (thallus), 27.08.2012, MZ 1254, LE 261481.
Apothecia 0.2– 0.4(– 0.5) mm diam., often strongly convex, surface rough, rarely with slight whitish
pruina, scattered, discrete or rarely confluent. Epihymenium dark brown, sometimes with olive or
purple hue, here and there with orange speckles, K+ green; hymenium sometimes purplish above.
Ascospores (12.5−)13.0 −15.0(−16.6) × (4.2−)5.2−6.4(−7.4) µm, l/b = (1.9−)2.1−2.7(−3.2) (n = 83).
*Abrothallus cetrariae I.Kotte
1: on Platismatia glauca (thallus, mainly on galls induced by Nesolechia oxyspora), 14.08.2012, MZ 12167, LE
261465.
New to Asia.
Abrothallus cf. chrysanthus Stein
2: on Usnea dasypoga (dark branch base), 15.08.2012, MZ 12209, LE 261241.
Apothecia up to 0.2 mm diam. Epithecium medium to dark brown with purple tinge, containing yellow-
orange speckles, K+ fleetingly green. Hymenium c. 30 – 40 µm tall, colourless to pale purple above,

380 Herzogia 27 (2), 2014
which does not fit observations of Stein (1879) and Etay o & van den Boom (2006), who reported
the hymenium being almost hyaline, only narrowly brown above, or yellowish and without violet gra-
nules. Hypothecium medium to dark brown, K+ olive. Ascospores pale to medium brown, verruculose,
(7.0−)8.1−10.3(−12.2) × (2.9−)3.3−4.3(−4.9) µm, l/b = (2.0−)2.2−2.8(−3.1) (n = 28), 8 per ascus.
The species was formerly reported in Russia from Dagestan Republic in the Caucasus (Urbanavichus
& Ismailov 2013).
Abrothallus cf. microspermus Tul.
10: on Flavopunctelia flaventior (thallus), 29.08.2012, MZ 1223, LE 261291; 15: on F. flaventior (thallus),
31.08.2012, MZ 1227, LE 261361.
Ascomata often somewhat flattened from above, 0.2– 0.5 mm diam., occasionally slightly pruinose.
Epihymenium dark brown with orange speckles, K+ olive green; hymenium more or less colourless;
hypothecium medium brown. Ascospores medium brown, verruculose, (0 –)1-septate, upper cell broa-
der than lower one, (11.3−)12.3−15.5(−18.0) × (4.7−)5.2−6.4(−7.6) µm, l/b = (1.9−)2.1−2.5(−3.2) (n =
83). According to Tulasne (1852) ascospores of the species are much smaller, 9.5−11 × 3.2−4.8 µm.
Based on material from the type host Flavoparmelia caperata, Diederich (2011) reported for the
species epruinose apothecia and also somewhat smaller ascospores, (11.4 –)11.7–13.9(–15.9) ×
(4.6 –)4.8 –5.5(–5.8) µm, l/b = (2.0 –)2.2–2.8(–3.1) (n = 20).
So far the species was known in Russia from Southern Ural (Urbanavichene et al. 2013) and
Dagestan Republic in the Caucasus (Urbanavichus & Ismailov 2013).
Abrothallus parmeliarum (Sommerf.) Arnold
1: on Parmelia sulcata (thallus), 14.08.2012, MZ 1225, LE 261274; 7: on P. sulcata (on galls, associated with
Nesolechia oxyspora), 22.08.2012, MZ 1240b, LE 261251b; 14: on P. sulcata (thallus), 31.08.2012, MZ 1232, LE
261484.
Abrothallus prodiens (Harm.) Diederich & Hafellner
4: on Hypogymnia physodes (thallus), 20.08.2012, MZ 12104, LE 261387.
Ascomata convex, constricted at the base, occasionally slightly pruinose, 150 –500 µm diam.
Epithecium covered by yellowish or reddish brown crystals. Hymenium more or less violet, parti-
cularly above. Ascospores medium olive brown to brown, granulate, (10.0−)11.2−14.2(−16.6) ×
(4.3−)5.0−6.2(−6.8) µm, l/b = (1.9−)2.1−2.5(−2.7) (n = 26).
Arthonia epiphyscia Nyl.
6: on Physcia aipolia (thallus), 22.08.2012, MZ 12207, LE 261454.
*Arthonia excentrica Th.Fr.
12: on Lepraria sp. (thallus) growing on mossy rock, 28.08.2012, MZ 1298, LE 261207.
*Arthonia molendoi (Frauenf.) R.Sant.
8: on Caloplaca tiroliensis (apothecia), 24.08.2012, MZ 1282, LE 261011.
*Arthonia stereocaulina (Ohlert) R.Sant.
3: on Stereocaulon alpinum (phyllocladia), 17.08.2012, MZ 1267, LE 261121; 18.08.2012, MZ 1268, LE 261171.
Arthophacopsis parmeliarum Hafellner
1: on Parmelia saxatilis (thallus), 14.08.2012, MZ 1237, LE 261214; 4: on P. saxatilis (thallus), 20.08.2012, MZ
1221, LE 261234; 5: on P. omphalodes (thallus), 21.08.2012, MZ 1236, LE 261244; 6: on P. sulcata (thallus),
22.08.2012, MZ 1238b, LE 261224b; 12: on P. sulcata (thallus), 28.08.2012, MZ 1222a, LE 261491a.
Formerly known in Russia from Adygeya Republic in the Caucasus (Zhurbenko & Otte 2012),
Chukotka (Zhurbenko 2009a) and the Kamchatka Peninsula (Zhurbenko et al. 2012b).
*Athelia arachnoidea (Berk.) Jülich
10: on Physconia distorta (thallus), 27.08.2012, MZ 12202b, LE 261414b.

Zhurbenko & Kobzeva: Lichenicolous fungi from Northwest Caucasus, Russia 381
*Bachmanniomyces uncialicola (Zopf) D.Hawksw.
3: on Cladonia arbuscula (thallus), 19.08.2012, MZ 12211a, LE 261391a.
*Biatoropsis usnearum Räsänen
1: on Usnea dasypoga (thallus), 14.08.2012, MZ 12141, LE 261385; MZ 12142a, LE 261375a; 2: on U. dasypoga
(thallus), 15.08.2012, MZ 12208, LE 261341; 4: on U. dasypoga (thallus), 20.08.2012, MZ 12143a, LE 261255a;
14: on U. intermedia (thallus), 31.08.2012, MZ 12147a, LE 261425a; 16: on U. intermedia (thallus), 2.09.2012,
MZ 12139a, LE 261495a.
Formerly known in Asian Russia from the Sverdlovsk Region (Shiryaev et al. 2010) and the
Kamchatka Peninsula (Zhurbenko et al. 2012b).
Briancoppinsia cytospora (Vouaux) Diederich, Ertz, Lawrey & van den Boom
4: on Hypogymnia physodes (thallus), 20.08.2012, MZ 12105, LE 261447.
Carbonea vitellinaria (Nyl.) Hertel
3: on Candelariella vitellina (thallus), 18.08.2012, MZ 1270, LE 261081.
Cercidospora melanophthalmae Nav.-Ros., Calat. & Hafellner
3: on Rhizoplaca melanophthalma (hymenium of apothecia), 17.08.2012, MZ 12173, LE 261285; 8: on R. melano-
phthalma (hymenium of apothecia), 24.08.2012, MZ 12174, LE 261395.
Ascomata 150 –300 µm diam. Asci 4 – 8-spored. The examined material differs from the species proto-
logue (Calatayud et al. 2013) in having larger and less elongated ascospores, (18.0−)21.7−26.1(−28.0)
× (5.3−)6.4−8.2(−8.8) µm, l/b = (2.7−)3.0−3.6(−4.0) (n = 29) vs. (16 –)18 –22(–24) × (4 –)5 – 6.5(–
7) µm, l/b = (2.4 –)2.9 – 4.2(– 4.8), and in blackening of infested hymenium. However, similarly larger
ascospores have been reported for the specimen of this species from the Himalayas (Zhurbenko
2013a). The material also resembles C. macrospora (Uloth) Hafellner & Nav.-Ros., which is typi-
cally associated with lichens of the Lecanora saxicola group, has 4-spored or only exceptionally
8-spored asci and more elongated ascospores, measuring (19 –)20 –25(–30) × 4 – 6(–7) µm, l/b =
(3.0 –)3.8 –5.4(– 6.7) (Calatayud et al. 2013).
The species is known from many regions of the Holarctic, including Caucasus (Armenia), but is new
to Russia (Vondrák & Etayo 2007, Calatayud et al. 2013, Zhurbenko 2013a).
*Clypeococcum cetrariae Hafellner
3: on Flavocetraria cucullata (lobes), 18.08.2012, MZ 1272, LE 261071; on F. cucullata (lobes), 19.08.2012, MZ
12214, LE 261282; on Cetraria islandica (lobes), 19.08.2012, MZ 12215, LE 261263.
Asci narrowly subcylindrical or slightly swollen in lower half, (41−)46−56(−62) × 10−12(−13) µm (n
= 15, in water or K/I), 4-spored. Ascospores hyaline then pale to dark greyish olive, narrowly obovoid,
(12.0−)14.5−17.3(−19.6) × (4.5−)5.1−6.3(−7.7) µm, l/b = (2.1−)2.5−3.1(−3.6) (n = 84), (0−)1-septate,
not or rarely slightly constricted at the septum, often with 1−2 large guttules in each cell, wall c. 1 µm
thick, verruculose, mostly obliquely biseriate in the ascus. Infected host parts become bleached and
then turn to black.
Formerly known in Russia from the arctic Yakutiya (Zhurbenko 2002), Evenkiya (Zhurbenko &
Zhdanov 2013) and the Sokhondo Reserve in the Transbaikal Region (Zhurbenko & Yakovchenko
2014).
Corticifraga fuckelii (Rehm) D.Hawksw. & R.Sant.
16: on Peltigera praetextata (on galls induced by Refractohilum peltigerae), 2.09.2012, MZ 12223b, LE 261483b.
*Corticifraga peltigerae (Fuckel) D.Hawksw. & R.Sant.
2: on Peltigera polydactylon agg. (bleached portions of thallus), 15.08.2012, MZ 12224, LE 261413.
*Dactylospora deminuta (Th.Fr.) Triebel
3: on Gyalecta foveolaris (damaged apothecia), 18.08.2012, MZ 1289, LE 261031; on Fuscopannaria praetermissa
(thallus), 18.08.2012, MZ 1292, LE 260981; 8: on Biatora subduplex (thallus), 24.08.2012, MZ 1286, LE 260951.
Gyalecta foveolaris and Fuscopannaria praetermissa are new host species.

382 Herzogia 27 (2), 2014
Dactylospora lobariella (Nyl.) Hafellner
1: on Lobaria pulmonaria (thallus), 14.08.2012, MZ 12118, LE 261467; 2: on L. amplissima (thallus), 15.08.2012,
MZ 1231, LE 261331; on L. pulmonaria (thallus), 15.08.2012, MZ 12117, LE 261337; 4: on L. pulmonaria (thal-
lus), 20.08.2012, MZ 12116, LE 261307; 5: on L. pulmonaria (thallus), 21.08.2012, MZ 12115, LE 261327.
Often associated with bleached/damaged portions of the host lobes.
Dactylospora parasitica (Flörke) Zopf
4: on Pertusaria albescens (thallus), 20.08.2012, MZ 12135, LE 261336.
*Endococcus incrassatus Etayo & Breuss
17: on Placidiopsis cf. cinerascens (thallus), 5.09.2012, MZ 1294, LE 261217.
Ascomata ovoid to pyriform, 150−250 µm diam. Ascospores ellipsoid, broadly ellipsoid, ovoid or ob-
long, occasionally slightly curved, ends rounded or sometimes more or less acute, initially hyaline,
then olive brown and finally medium brown, (11.1−)12.0−14.6(−17.0) × (6.3−)7.0−9.0(−10.5) µm,
l/b = (1.2−)1.4−2.0(−2.4) (n = 61), (0−)1-septate, not or slightly constricted at the septum, cells equal
or the lower one narrower, verruculose, usually with one large guttule in each cell, non-halonate.
Pathogenicity not observed.
Formerly known from Mexico (Etayo & Breuss 2001), the USA (Knudsen & Kocourková 2008),
Russia (Northern Ural) (Zhurbenko et al. 2012a) and India (Zhurbenko 2013a). New to Asian
Russia.
*Epicladonia sandstedei (Zopf) D.Hawksw.
1: on Cladonia ochrochlora (podetia), 14.08.2012, MZ 125b, LE 261452b; 4: on Cladonia sp. (basal squamules)
growing on lignum, 20.08.2012, MZ 1220, LE 261412.
Formerly known in Asian Russia from the Taimyr Peninsula (Zhurbenko & Alstrup 2004) and the
Sokhondo Reserve in the Transbaikal Region (Zhurbenko & Yakovchenko 2014).
*Hainesia aeruginascens Brackel
1: on Hypogymnia physodes (thallus), 14.08.2012, MZ 12101, LE 261277.
Formerly known only from Germany (Brackel 2014). New to Asia and Russia.
*Hainesia brevicladoniae Diederich & van den Boom
3: on Cladonia gracilis (podetia), 17.08.2012, MZ 127, LE 261382; on C. pocillum (basal squamules), 18.08.2012,
MZ 1216, LE 261462.
Conidia bacilliform, aseptate or sometimes with a hardly visible septum, (9.6−)13.3−16.9(−18.6) ×
(1.1−)1.3−1.5(−1.6) µm, l/b = (6.4−)8.9−12.3(−16.3) (n = 73), with small guttules. In the species pro-
tologue conidia were reported being always aseptate and slightly narrower, viz. (13.5 –)15.3 –17.5(–
18.0) × (1.0 –)1.0 –1.1(–1.2) µm, l/b = (11.5 –)14.0 –16.9(–17.5) (n = 20). (Diederich & van den
Boom 2013). Causes bleaching of host tissues.
This recently described species was so far known from Belgium, France, Germany and the Netherlands
(Diederich & van den Boom op. cit., Brackel 2014). New to Asia and Russia.
Illosporiopsis christiansenii (B.L.Brady & D.Hawksw.) D.Hawksw.
1: on Physcia aipolia (damaged thallus) and adjacent bark, 14.08.2012, MZ 12205, LE 261315; 18: on Phaeophyscia
orbicularis (damaged thallus), 9.09.2012, MZ 12153, LE 261335.
Formerly known in Asian Russia from Adygeya Republic in the Caucasus (Zhurbenko & Otte 2012).
*Illosporium carneum Fr.
2: on Peltigera polydactylon agg. (upper and occasionally lower sides of lobes) and adjacent mosses, 15.08.2012,
MZ 12229b, LE 261442b.
Intralichen christiansenii (D.Hawksw.) D.Hawksw. & M.S.Cole
17: on Candelariella vitellina (hymenium of apothecia), 5.09.2012, MZ 12136, LE 261396.

Zhurbenko & Kobzeva: Lichenicolous fungi from Northwest Caucasus, Russia 383
Lichenochora cf. lepidiotae (Anzi) Etayo & Nav.-Ros.
3: on Fuscopannaria praetermissa (thallus), 18.08.2012, MZ 1290, LE 260911.
Ascospores (0 –)1-septate, (18.0−)22.8−32.2(−38.0) × (5.3−)5.7−6.9(−7.5) µm, l/b = (2.4−)3.4−5.4(−6.4)
(n = 15), 8 per ascus. Etayo & Navarro-Rosinés (2008) reported ascospores of the species some-
what narrower, viz. (20.5 –)23 –32(–34.5) × (4 –)4.5 – 6 µm, l/b = (3.5 –)4.0−6.4(– 6.8). The examined
material also resembles L. inconspicua Hafellner, which has somewhat wider ascospores, measuring
(19−)20.5−35(−39) × 6−7.5(−9) µm, l/b = (2.9−)3.3−5.0(−5.7) (Navarro-Rosinés et al. 1998).
The species was formerly known in Russia from the Caucasus (Krasnodar Territory and Kabardino-
Balkaria Republic) (Etayo & Navarro-Rosinés 2008, Zhurbenko & Otte 2012).
*Lichenochora rinodinae Zhurb.
3: on Rinodina turfacea (thallus), 18.08.2012, MZ 1266, LE 261101.
This recently described species was so far known only from the Arctic (Zhurbenko 2013b).
*Lichenoconium erodens M.S.Christ. & D.Hawksw.
1: on Hypogymnia physodes (thallus), 14.08.2012, MZ 12112, LE 261497; on Platismatia glauca (thallus),
14.08.2012, MZ 12154, LE 261225; 3: on Cetraria islandica (dark decaying lobe bases), 18.08.2012, MZ 1281, LE
260971; 4: on Hypogymnia physodes (thallus), 20.08.2012, MZ 12108, LE 261427; 7: on Lecanora sp. (hymenium
of apothecia) growing on bark, 22.08.2012, MZ 12156, LE 261455; on Melanelia stygia (hymenium of apothecia),
22.08.2012, MZ 1239, LE 261411; on Bryoria chalybeiformis (blackened branch base), 22.08.2012, MZ 1244, LE
261371; 10: on Physconia distorta (thallus), 29.08.2012, MZ 12210a, LE 261211a; 13: on Pseudevernia furfuracea
(thallus), 28.08.2012, MZ 12157, LE 261445.
Lichenoconium lecanorae (Jaap) D.Hawksw.
3: on Rhizoplaca chrysoleuca (apothecia, occasionally thallus), 17.08.2012, MZ 1277, LE 260961; 18.08.2012,
MZ 1284, LE 261051; on Brodoa intestiniformis (darkened lobe bases), 18.08.2012, MZ 1279, LE 260991; 6: on
Rhizoplaca chrysoleuca (apothecia), 22.08.2012, MZ 1278, LE 261161.
*Lichenoconium pyxidatae (Oudem.) Petr. & Syd.
4: on Cladonia coniocraea (damaged podetia), 20.08.2012, MZ 12217, LE 261333; 6: on C. pyxidata (bleached
portions of podetia), 22.08.2012, MZ 129, LE 261222; 15: on C. chlorophaea s. l. (bleached portions of podetia),
1.09.2012, MZ 1215, LE 261243.
Conidiomata 50−125 µm diam. Conidia subglobose to obovoid, sometimes truncated at the base,
pale to medium brown or yellowish brown, smooth or occasionally indistinctly verruculose,
(2.3−)2.5−3.1(−3.5) µm, l/b = (1.0−)1.1−1.3(−1.7) (n = 66). Conidiogenous cells subcylindrical or
slightly swollen in lower half, (6.7−)7.7−9.9(−11.0) × (2.1−)2.3−2.7(−3.0) (n = 33).
Lichenoconium usneae (Anzi) D.Hawksw.
2: on Anaptychia ciliaris (bleached lobe portions), 15.08.2012, MZ 12129, LE 261406.
*Lichenopeltella peltigericola (D.Hawksw.) R.Sant.
2: on Peltigera polydactylon agg. (underside of lobes including rhizines), 15.08.2012, MZ 12229a, LE 261442a;
16: on P. polydactylon agg. (lower and occasionally upper side of lobes), 2.09.2012, MZ 12230, LE 261232.
Asci (26−)29−37(−42) × 11−13 µm (n = 20), 4-spored. Ascospores (14.3−)15.8−19.0(−21.7) ×
(3.8−)4.1−4.9(−5.4) µm, l/b = (2.9−)3.6−4.2(−4.5) (n = 42), setulae very rarely observed.
*Lichenopuccinia poeltii D.Hawksw. & Hafellner
4: on Parmelia sulcata (thallus), 20.08.2012, MZ 1241, LE 261304; 5: on P. cf. sulcata (thallus), 21.08.2012, MZ
1243, LE 261444.
Formerly known in Russia from the Taimyr (Zhurbenko 2009a) and Kamchatka (Zhurbenko et al.
2012b) Peninsulas.
*Lichenosticta alcicornaria (Linds.) D.Hawksw.
1: on Cladonia ochrochlora (lower side of basal squamules, rarely podetia), 14.08.2012, MZ 125a, LE 261452a;
4: on C. coniocraea (lower side of basal squamules, rarely podetia), 20.08.2012, MZ 12218, LE 261312; 9: on C.
pyxidata (lower and occasionally upper side of basal squamules, rarely podetia), 25.08.2012, MZ 126, LE 261492.

384 Herzogia 27 (2), 2014
*Lichenostigma alpinum (R.Sant., Alstrup & D.Hawksw.) Ertz & Diederich
3: on Ochrolechia sp. (thallus) growing on mosses, 18.08.2012, MZ 1274, LE 2610211; 7: on Lecanora intume-
scens (thallus), 23.08.2012, MZ 12149, LE 261306; 10: on Flavoparmelia caperata (somewhat damaged thallus),
29.08.2012, MZ 1226c, LE 261311c.
The species has only rarely been recorded on Lecanora (Brackel 2009, Etayo 2010). Specimen LE
261306 growing on this host genus is characterized by conidiomata 30 –50(–70) µm diam. and pale to
dark olive brown, smooth conidia, measuring (10.0−)11.4−14.8(−15.5) × (8.0−)9.1−11.3(−12.4) µm,
l/b = (1.0−)1.2−1.4(−1.6) (n = 16), composed of up to 30 cells, 3.0 – 4.5 × 2.7– 4.1 µm each.
Lichenostigma cosmopolites Hafellner & Calat.
6: on Xanthoparmelia stenophylla (thallus), 22.08.2012, MZ 1248, LE 261301; 11: on X. stenophylla (thallus),
27.08.2012, MZ 1250, LE 261421; 14: on X. stenophylla (thallus), 31.08.2012, MZ 1251, LE 261221.
The host thalli are slightly bleached under severe infection.
This cosmopolitan species was so far reported in Russia from Republic of Karelia (Alstrup et al.
2005), the Transbaikal Region (Zhurbenko & Yakovchenko 2014) and the Jewish Autonomous
Region (Zhurbenko 2014).
Lichenostigma maureri Hafellner
1: on Usnea intermedia (thallus, underside of apothecia), 14.08.2012, MZ 12137a, LE 261405a; on U. dasypoga
(thallus), 14.08.2012, MZ 12137b, LE 261405b; MZ 12142b, LE 261375b; 4: on U. dasypoga (thallus), 20.08.2012,
MZ 12143b, LE 261255b; on U. intermedia (thallus, underside of apothecia), 20.08.2012, MZ 12145, LE 261435
(anamorph and teleomorph); 12: on U. intermedia (thallus, underside of apothecia), 28.08.2012, MZ 12144, LE
261266 (anamorph and teleomorph); 13: on Pseudevernia furfuracea (thallus), 28.08.2012, MZ 12148, LE 261265
(anamorph and teleomorph); 14: on U. intermedia (thallus, underside of apothecia), 31.08.2012, MZ 12147b, LE
261425b; 16: on U. intermedia (thallus, underside of apothecia), 2.09.2012, MZ 12139b, LE 261495b; MZ 12140a,
LE 261475a (anamorph and teleomorph).
The host thalli are slightly bleached under severe infection.
It is noteworthy that in the specimens LE 261475a, LE 261435 and LE 261266 we observed within
ascomata of Lichenostigma maureri additional medium olive to brown (K+ more olive), subglobose
fructifications 15 –30 µm diam. of unclear origin. They contain pale olive to brown (K+ more olive),
oblong, ellipsoid or broadly ellipsoid, often more or less truncated at one end, (3.4−)4.1−5.3(−6.0) ×
(2.0−)2.2−3.0(−3.7) µm, l/b = (1.3−)1.5−2.1(−2.5) (n = 58, in water or K), (0 –)1-septate diaspores
(Fig. 1).
*Lichenostigma rupicolae Fern.-Brime & Nav.-Ros.
6: on saxicolous Pertusaria sp. (apothecia, thallus) and occasionally directly on rock, 22.08.2012, MZ 12124, LE
261486.
This recently described species was so far known only from France (Fernandez-Brime et al. 2010),
Spain (Etayo 2010, Fernandez-Brime et al. 2010) and Turkey (Fernandez-Brime et al. 2010,
Yazici et al. 2011). New to Russia.
*Lichenothelia rugosa (G.Thor) Ertz & Diederich
9: on Diploschistes muscorum (thallus, occasionally apothecia), 25.08.2012, MZ 12120, LE 261487; 11: on D.
scruposus (thallus, occasionally apothecia), 27.08.2012, MZ 12126, LE 261456.
Some ascomata are associated with a thin green crustose lichenized thallus, which fits the observations
of Diederich (1986). The lichenized thallus was mainly observed on the hymenium, but occasionally
also on the thalline margins of the host apothecia.
Formerly known in Asian Russia from Altai (Zhurbenko & Davydov 2000) and Chukotka
(Zhurbenko 2009a).
*Marchandiobasidium aurantiacum Diederich & Schultheis
1: on Physcia stellaris (apothecia, thallus), 14.08.2012, MZ 12206, LE 261415.
The species is common and widespread in Europe (Diederich et al. 2003), but is new to Russia.

Zhurbenko & Kobzeva: Lichenicolous fungi from Northwest Caucasus, Russia 385
Marchandiomyces corallinus (Roberge) Diederich & D.Hawksw.
10: on Flavoparmelia caperata (thallus), 29.08.2012, MZ 1226e, LE 261311e; on epiphytic Ramalina sp.
(thallus), 29.08.2012, MZ 1297, LE 261287; mainly on Usnea subfloridana, also on adjacent Parmelia sulca-
ta and Pseudevernia furfuracea (thalli), 29.08.2012, MZ 12146, LE 261365; 11: on Melanelia tominii (thallus),
30.08.2012, MZ 12134, LE 261256; on Xanthoparmelia stenophylla (thallus), 30.08.2012, MZ 1255, LE 261381;
14: on X. stenophylla (thallus), 31.08.2012, MZ 1256, LE 261231; 18: on Physcia stellaris and adjacent Xanthoria
parietina (thalli), 9.09.2012, MZ 12151, LE 261345.
*Merismatium cf. cladoniicola Alstrup
17: on Cladonia pocillum (basal squamules), 5.09.2012, MZ 1217b, LE 261362b.
Ascomata 150−200 µm diam. Ascospores ellipsoid, hyaline to evenly pale brown, (9.5−)10.6−14.0(−15.4)
× (4.5−)5.1−6.1(−6.4) µm, l/b = (1.6−)1.8−2.6(−3.2) (n = 14), 0−3-septate. In the species protologue
(Alstrup 1997), ascospores were reported as being 3-septate, with brown median cells and mostly
hyaline end-cells.
So far the species was known only from the type collection from Norway growing on Cladonia ciliata.
Fig. 1: Squashed fruit with diaspores of unclear origin located inside regular fruit of Lichenostigma maureri (in K,
LE 261266). Bar: 10 µm.

386 Herzogia 27 (2), 2014
*Merismatium heterophractum (Nyl.) Vouaux
16: on Cladonia pyxidata (basal squamules) and adjacent plant remnants and film of cyanobacteria, 2.09.2012, MZ
1213b, LE 261242b.
*Minutoexcipula calatayudii V.Atienza
4: on Hypogymnia tubulosa (thallus), 20.08.2012, MZ 12113, LE 261257.
Conidia (6.3−)7.0−8.6(−9.2) × (3.3−)3.8−4.4(−4.6) µm, l/b = (1.4−)1.6−2.1(−2.7) (n = 28).
The species was formerly known from Spain (Atienza 2002) and Germany (Brackel 2014). New to
Asia and Russia.
Muellerella erratica (A.Massal.) Hafellner & V.John
7: on Lecanora sp. (apothecia) gowing on bark, 23.08.2012, MZ 12165, LE 261485; 17: on saxicolous Acarospora
sp. (thallus), 5.09.2012, MZ 12166, LE 261355; on Caloplaca sp. (thallus) growing on limestone, 5.09.2012, MZ
12161, LE 261476.
Muellerella hospitans Stizenb.
18: on Bacidia rubella (hymenium of apothecia), 8.09.2012, MZ 12159, LE 261226; 9.09.2012, MZ 12163, LE
261316.
Muellerella lichenicola (Sommerf.) D.Hawksw.
7: on Lecanora chlarotera (apothecia, thallus), 23.08.2012, MZ 12160, LE 261416; 18: on Caloplaca cerina (hy-
menium of apothecia), 8.09.2012, MZ 12164, LE 261325; MZ 12158, LE 261276; 9.09.2012, MZ 12162, LE
261246.
Muellerella pygmaea (Körb.) D.Hawksw.
3: on Lecanora intricata (apothecia, thallus), 18.08.2012, MZ 1283, LE 260941; 17: on Circinaria contorta (thal-
lus), 5.09.2012, MZ 12131a, LE 261446a.
Muellerella ventosicola (Mudd) D.Hawksw.
3: on Ophioparma ventosa (thallus), 17.08.2012, MZ 1280, LE 261191.
Nectriopsis lecanodes (Ces.) Diederich & Schroers
2: on Peltigera membranacea (decaying lobe bases), 15.08.2012, MZ 12227, LE 261473; 9: on P. horizontalis (da-
maged portions of thallus, occasionally apothecia), 25.08.2012, MZ 12225, LE 261403; 15: on P. canina (decaying
lobe bases), 1.09.2012, MZ 12226, LE 261392; 16: on P. canina (decaying lobe bases), 2.09.2012, MZ 12228, LE
261482.
*Neobarya peltigerae Lowen, Boqueras & Gomez-Bolea
16: on Peltigera polydactylon (upper and occasionally lower lobe sides), 2.09.2012, MZ 12234, LE 261332.
Formerly known from Spain, Scotland (Candoussau et al. 2007) and the USA (Zhurbenko 2009c).
Peltigera polydactylon is a new host species. New to Asia and Russia.
Nesolechia oxyspora (Tul.) A.Massal. s. l.
1: on Platismatia glauca (thallus), 14.08.2012, MZ 12152, LE 261295; 4: on Parmelia saxatilis (thallus),
20.08.2012, MZ 1228, LE 261201; 6: on Parmelia sulcata (thallus), 22.08.2012, MZ 1240a, LE 261251a; 11: on
Xanthoparmelia stenophylla (thallus), 27.08.2012, MZ 1253, LE 261441; 12: on P. sulcata (thallus), 28.08.2012,
MZ 1222b, LE 261491b.
*Odontotrema thamnoliae Zhurb., Diederich & Etayo
3: on Thamnolia vermicularis var. subuliformis (thallus), 17.08.2012, MZ 1262, LE 260901; MZ 1260, LE 261102.
*Opegrapha anomea Nyl.
1: on Variolaria amara (thallus, including soralia), 14.08.2012, MZ 12175, LE 261466.
New to Asia and Russia.
*Opegrapha rotunda Hafellner
10: on Physconia distorta (thallus), 27.08.2012, MZ 12201a, LE 261404a.

Zhurbenko & Kobzeva: Lichenicolous fungi from Northwest Caucasus, Russia 387
Ascomata up to 0.3 mm diam. Ascospores (13.8−)14.8−19.4(−22.5) × 4.5−5.5(−6.0) µm, l/b =
(2.7−)2.8−4.0(−5.0) (n = 17), (2–)3-septate, with halo 1–2 µm thick. Hymenium I+ blue and red.
Formerly known from Austria, Italy, Portugal and Scotland (Hafellner 1994, van den Boom &
Etay o 2000, Brackel 2008, Smith et al. 2009). New to Asia and Russia.
*Paranectria oropensis (Ces.) D.Hawksw. & Piroz.
10: on Cetrelia olivetorum (soralia), 27.08.2012, MZ 12169, LE 261215; 11: on Nephroma parile (thallus, mainly
soralia), 27.08.2012, MZ 12133, LE 261346.
Formerly known in Russia from the vicinities of Krasnoyarsk in Siberia (Zhurbenko & Otnyukova
2001). The species is known on various host genera, mainly of corticolous macrolichens, Nephroma
being a new one.
*Phacopsis cephalodioides (Nyl.) Triebel & Rambold
1: on Hypogymnia physodes (thallus), 14.08.2012, MZ 12109, LE 261347; 4: on H. physodes (thallus), 20.08.2012,
MZ 12111, LE 261397.
Phacopsis huuskonenii Räsänen
2: on Bryoria capillaris and Bryoria sp. (thalli), 15.08.2012, MZ 1246, LE 261261; 15: on B. capillaris (thallus),
1.09.2012, MZ 1245, LE 261321.
*Phacopsis cf. usneae C.W.Dodge
2: on Usnea dasypoga (thallus), 15.08.2012, MZ 12212a, LE 261401a.
Ascomata subglobose, sessile, c. 0.1 mm diam., black, glossy, dispersed. Epihymenium and hyme-
nium dark bluish green, hypothecium dark greyish reddish purple; section of apothecium K−, N+
purple. Paraphyses 1.8 –3.3 µm diam., not or slightly swollen at the apex, often somewhat swollen
between the septa, sometimes scarcely branched. Asci clavate, sometimes with a strongly thickened
apex, c. 35−43 × 11−19 µm, 8-spored. Ascospores hyaline, narrowly ellipsoid, (7.6−)8.4−10.8(−12.9)
× (2.8−)3.0−3.8(−5.0) µm, l/b = (1.8−)2.3−3.3(−4.2) (n = 25), aseptate, smooth, non-halonate, gut-
tulate, irregularly biseriate in the ascus. It is noteworthy that the examined ascomata are intimately
intermixed with conidiomata of Pseudoseptoria usneae, which might represent their anamorph.
There are some essential inconsistencies between the examined material and various descriptions of
Phacopsis usneae. According to Etayo & Sancho (2006) the species has ascomata up to 3.5 mm
diam., a colourless to olivaceous or brownish hymenium, a brown (N−) to blackish blue (N+ violet)
epihymenium, covered by a layer of colourless gel, apically swollen paraphyses 6 –9 µm thick, and lar-
ger ascospores 11–15 × 5−6(–7) µm. However, these authors note that the species is variable which fits
observations of Hawksworth & Iturriaga (2006), who reported its apothecia being up to 0.75 mm
diam., usually aggregated in mounded clumps, hymenium colourless, epihymenium dark blue-grey,
paraphyses mainly 3 – 4 µm thick, but sometimes up to 10 µm wide at the tips, ascospores 9.5 –10.5
× 5.5 – 6.5 µm. In the species protologue its paraphyses were reported being 4 µm in diam., with not
thickened tips, and the ascospores fusiform, 10 –12 × 3.5 – 4 µm (Dodge 1948). The examined material
also resembles Lecanora usneicola Etayo, but that species readily differs in having a colourless hyme-
nium and hypothecium (Etayo & Sancho 2006).
So far Phacopsis usneae was known only from the Antarctic, subantarctic islands and Tierra del
Fuego, growing on Usnea antarctica and U. trachycarpa (Etayo & Sancho 2006, Hawksworth &
Iturriaga 2006).
*Phoma cf. epiphyscia Vouaux
2: on Anaptychia ciliaris (lobe tips), 15.08.2012, MZ 12128, LE 261496.
Conidiomata 125 –150 µm diam., protruding in the upper part, inducing strong bleaching of the
host tissues. Conidia mostly narrowly oblong, (5.4−)5.9−6.7(−7.1) × (2.4−)2.5−2.9(−3.0) µm, l/b =
(2.0−)2.2−2.6 (n = 27), usually with a small guttule near each end.
Phoma epiphyscia has been neotypified on a specimen on Phaeophyscia sciastra by Alstrup &
Hawksworth (1990), who reported its conidia being up to 7 × 3 µm, l/b = 2−2.3, but did not provide a

388 Herzogia 27 (2), 2014
detailed description of the neotype. Phoma physciicola Keissl., the other species of the genus known to
grow on the members of Physciaceae, differs from our material in having broadly ellipsoid, somewhat
shorter and wider conidia, 4−5.5(−6) × 2.5−3.5(−4) µm (Diederich 2004) with a length/breadth ratio
of 1.5−1.7 (Alstrup & Hawksworth 1990).
*Phoma peltigerae (P.Karst.) D.Hawksw.
2: on Peltigera sp. (upper lobe side), 15.08.2012, MZ 12219, LE 261203.
The host thallus is strongly bleached under the infection.
Formerly known in Asian Russia from the Taimyr Peninsula (Zhurbenko 2009b) and Olekminskii
Reserve in Yakutiya (Zhurbenko & Vershinina 2014).
Plectocarpon lichenum (Sommerf.) D.Hawksw.
1: on Lobaria pulmonaria (thallus), 14.08.2012, MZ 12122, LE 261367; 2: on L. pulmonaria (upper and occasi-
onally lower sides of lobes), 15.08.2012, MZ 12121, LE 261267; 4: on L. pulmonaria (thallus), 20.08.2012, MZ
12123, LE 261437.
*Polycoccum arnoldii (Hepp) D.Hawksw.
17: on Diploschistes muscorum (apothecia, thallus), 5.09.2012, MZ 12119, LE 261377.
The ascospores are somewhat narrower than reported in Hawksworth & Diederich (1988), viz.
(9.7−)9.9−11.9(−13.7) × (3.9−)4.1−4.9(−5.2) µm, l/b = (2.0−)2.1−2.7(−3.0) (n = 24) vs. 9−11.5(–13)
× 4.5 – 6.5(–7) µm.
New to Asia and Russia.
*Polycoccum kaernefeltii S.Y.Kondr.
18: on Caloplaca cerina (discs of apothecia), 8.09.2012, MZ 12130, LE 261386.
Ascomata perithecioid, subglobose, 100−150 µm diam., protruding in the upper part, aggrega-
ted. Exciple evenly medium to dark brown, 20 –30 µm thick above, c. 15 µm thick below, K+ oli-
ve. Interascal filaments 1.5 –2.5 µm diam., septate, not or scarcely branched. Ascospores narrowly
obovoid (wider above), pale brown, K+ olive, (1–)2(–3)-septate, mostly constricted at the septa,
(16.7−)18.8−21.8(−23.7) × (8.0−)8.6−10.0(−11.2) µm, l/b = (1.7−)2.0−2.4(−2.7) (n = 59, in water,
K or K/I), with several conspicuous guttules, smooth-walled, non-halonate, 8 per ascus (Fig. 2).
Clearly pathogenic, damaging and blackening host hymenia. In the species protologue its ascomata
were reported larger, (200 –)250 –350(– 450) µm diam., and immersed, with only the ostiole visible
(Kondratyuk 2008).
So far the species was known only from Australia growing on Teloschistes chrysophthalmus. Caloplaca
is a new host genus. New to Asia and Russia.
*Polycoccum pulvinatum (Eitner) R.Sant.
16: on Physcia aipolia (thallus), 2.09.2012, MZ 12191a, LE 261314a; 18: on P. caesia (thallus), 5.09.2012, MZ
12186, LE 261464.
Ascospores ellipsoid to narrowly obovoid, medium to dark olive, (0 –)1-septate, sometimes slightly con-
stricted at the septum, (13.5−)14.3−17.1(−19.5) × (6.4−)6.9−8.3(−9.0) µm, l/b = (1.7−)1.9−2.3(−2.5)
(n = 23), verrucose, often with one large and many small guttules in each cell, usually with halo
1–3 µm thick, more or less uniseriate in the ascus. Conidiomata immersed, intermixed with ascomata.
Conidia bacilliform, hyaline, aseptate, c. 4.5 × 1.5 µm.
*Pronectria echinulata Lowen
16: on Physcia aipolia (thallus, margin and disc of apothecia), 2.09.2012, MZ 12195b, LE 261324b.
Ascomata 150 –200 µm diam. Exciple orange or brownish orange, medium coloured above, pale to
colourless below, not changing colour in K and lactic acid. Asci c. 75−90 × 10−13.5 µm. Ascospores
hyaline, (11.6−)12.9−14.9(−15.8) × (5.7−)6.9−8.5(−9.3) µm, l/b = (1.5−)1.6−2.0(−2.3) (n = 36), 1-sep-
tate, smooth or finely echinate to verruculose, uniseriate in the ascus.
New to Asia and Russia.

Zhurbenko & Kobzeva: Lichenicolous fungi from Northwest Caucasus, Russia 389
*Pronectria santessonii (Lowen & D.Hawksw.) Lowen
2: on Anaptychia ciliaris (thalline margin and base of apothecia), 15.08.2012, MZ 12127, LE 261426.
Ascomata subglobose, 75 –175 µm diam., usually protruding only in the ostiolar area, pale to medi-
um orange-yellow-brown, with rather indistinct, sometimes darker papilla, aggregated. Ascospores
hyaline, narrowly ellipsoid or occasionally narrowly obovate, verruculose, (9.5−)12.9−16.7(−18.7)
× (3.7−)4.4−5.4(−6.0) µm, l/b = (1.9−)2.5−3.7(−4.7) (n = 60), (0 –)1-septate, not constricted at the
septum, usually with 1–2 large guttules in each cell, often with a halo 1–2 µm thick. Causes heavy
bleaching of the host tissues.
In the species protologue its ascomata were reported dark red, turning almost black with age,
190 –240 µm wide, ascospores slightly pale brown when mature, slightly constricted at the septum,
with sometimes finely echinulate wall, (10−)12−18 × 4.5−8 µm (Lowen & Hawksworth 1986).
Pronectria sp. and P. tincta (Fuckel) Lowen growing on the same host species differ from the exami-
ned material in having longer ascospores, 15−23 × 6−7 µm and 17−20 × 4−5 µm respectively (Lowen
& Hawksworth 1986).
New to Asia and Russia.
*Pseudoseptoria usneae (Vouaux) D.Hawksw.
2: on Usnea dasypoga (thallus), 15.08.2012, MZ 12212b, LE 261401b (intimately intermixed with ascomata of
Phacopsis cf. usneae); 8: on Usnea sp. (somewhat damaged branches), 22.08.2012, MZ 12138, LE 261235.
Fig. 2: Ascomata section of Polycoccum kaernefeltii with the ascospores (in K, LE 261386). Bar: 10 µm.

390 Herzogia 27 (2), 2014
Conidiomata 75 –150 µm diam. Wall dark bluish green outside and greyish reddish purple inside.
Conidia (10.6−)11.9−14.3(−17.0) × (2.5−)3.0−3.6(−4.3) µm, l/b = (2.7−)3.4−4.6(−5.7) (n = 93, in wa-
ter or K).
Formerly known in Russia from the Irkutsk Region (Zhurbenko & Vershinina 2014).
Pyrenidium actinellum Nyl.
3: on Peltigera rufescens (thallus), 18.08.2012, MZ 12221, LE 261302; 9: on P. elisabethae (thallus), 25.08.2012,
MZ 12220, LE 261463; 11: on Diploschistes scruposus (thallus), 27.08.2012, MZ 12125, LE 261206.
Ascomata 100 – 400 µm diam., aggregated in clusters and partly confluent. Exciple and hamatheci-
um in the ostiolar region bluish green. Ascospores pale to dark brown, difference in colour intensity
of the central and end cells not observed, (15.8−)18.6−22.2(−24.7) × (6.7−)7.4−9.0(−9.7) µm, l/b =
(2.0−)2.2−2.8(−3.3) (n = 65), (0 –)1–3-septate.
*Pyrenochaeta xanthoriae Diederich
18: on Xanthoria parietina (more or less damaged apothecia), 9.09.2012, MZ 1293, LE 261357.
Formerly known in Russia from Republic of Karelia (Alstrup et al. 2005). New to Asia.
*Reconditella physconiarum Hafellner & Matzer
10: on Physconia distorta (margins of damaged lobes), 29.08.2012, MZ 12193, LE 261344.
Formerly known in Asian Russia from Chukotka (Zhurbenko 2009b).
*Refractohilum peltigerae (Keissl.) D.Hawksw.
15: on Peltigera cf. praetextata (thallus), 1.09.2012, MZ 12222, LE 261383; 16: on P. praetextata (thallus),
2.09.2012, MZ 12223a, LE 261483a.
Conidia (14.2−)16.0−20.4(−23.8) × (5.5−)6.8−8.8(−10.5) µm, l/b = (1.7−)1.9−2.7(−3.4) (n = 49, in water
or K). The examined material fits the species protologue (Keissler 1920), but differs from its description
in Hawksworth (1977) in the following: 1) distinct percurrent proliferations of the conidiogenous cells
with annellations were not observed; 2) the conidia are not always aseptate, but very rarely also 1-septa-
te; 3) the base of the conidia is not distinctly thickened and refractive. Similar material without distinct
annellations of the conidiogenous cells has been previously erroneously reported from the Krasnoyarsk
Territory of Russia as Hawksworthiana peltigericola (D.Hawksw.) U.Braun (Zhurbenko 2012).
*Rhagadostoma lichenicola (De Not) Keissl.
3: on Solorina crocea (thallus), 18.08.2012, MZ 1265, LE 261091.
*Rhymbocarpus neglectus (Vain.) Diederich & Etayo
8: on Lepraria sp. (thallus) growing on mosses, 24.08.2012, MZ 1285, LE 261061.
*Skyttella mulleri (Willey) D.Hawksw. & R.Sant.
2: on Peltigera sp. (decaying thallus), 15.08.2012, MZ 12232, LE 261423.
Ascomata 150−175 µm diam. Paraphyses 1.5−2.5 µm diam. Ascospores (11.0−)11.7−14.1(−16.9) ×
(3.1−)3.5−4.1(−4.7) µm, l/b = (2.6−)2.9−3.9(−4.8) (n = 22). There are some discrepancies with the
species description in Hawksworth & Santesson (1988), where its ascomata were reported being
(0.2−)0.3−0.7(−1) mm diam., paraphyses (3−)3.5−4 µm diam., and ascospores (8.5−)10−12(−14) ×
2.5−3.5(−5) µm. However, these characters of the species seem to be rather variable as was shown in
the material from the Kamchatka Peninsula of Russia (Zhurbenko et al. 2012b).
Formerly known in Russia from Republic of Karelia (Schiefelbein & Rätzel 2005) and the
Kamchatka Peninsula (Zhurbenko et al. 2012b).
Sphaerellothecium cladoniae (Alstrup & Zhurb.) Hafellner
17: on Cladonia pocillum (basal squamules), 5.09.2012, MZ 1217a, LE 261362a.
*Sphaerellothecium cf. gowardii Alstrup & M.S.Cole
16: on Placidium squamulosum (thallus), 5.09.2012, MZ 12100, LE 261227.

Zhurbenko & Kobzeva: Lichenicolous fungi from Northwest Caucasus, Russia 391
So far the species was known only from the type collection in British Columbia of Canada growing on
Acarospora schleicheri (Alstrup & Cole 1998). Except for the host genus, the examined specimen
completely fits the species protologue. However, the latter is rather short and evidently based on scant
material as it does not show variation of the ascospore size. Additional collections are necessary to
clarify the species concept and thereafter the specimen identification.
Sphaerellothecium reticulatum (Zopf) Etayo
6: on Parmelia sulcata (thallus), 22.08.2012, MZ 1240c, LE 261251c; on P. sulcata (thallus), 22.08.2012, MZ
1238a, LE 261224a; 13: on P. sulcata (thallus), 28.08.2012, MZ 1229, LE 261294; 14: on P. sulcata (thallus),
31.08.2012, MZ 12188, LE 261284.
*Sphaerellothecium thamnoliae Zhurb. var. thamnoliae
3: on Thamnolia vermicularis var. subuliformis (thallus), 19.08.2012, MZ 12216, LE 261432.
*Stigmidium frigidum (Th.Fr. ex Sacc.) Alstrup & D.Hawksw.
3: on Thamnolia vermicularis var. subuliformis (thallus), 17.08.2012, MZ 1259, LE 261041; 8: on T. vermicularis
var. subuliformis (thallus), 24.08.2012, MZ 1264, LE 261001.
*Stigmidium pumilum (Lettau) Matzer & Hafellner
6: on Physcia caesia (thallus), 22.08.2012, MZ 12190, LE 261384; 11: on P. phaea (thallus), 30.08.2012, MZ
12189, LE 261374.
Heavily infected parts of the host thalli become grey.
*Stigmidium xanthoparmeliarum Hafellner
11: on Xanthoparmelia stenophylla (thallus), 27.08.2012, MZ 1252, LE 261431.
Walls of asci and ascospores BCr−. Causes distinct bleached spots on the host thallus.
New to Russia.
*Syzygospora bachmannii Diederich & M.S.Christ.
6: on Cladonia pyxidata (podetia), 22.08.2012, MZ 1218a, LE 261372a; 16: on C. pyxidata (podetia), 2.09.2012,
MZ 1213a, LE 261242a.
Formerly known in Asian Russia from the Sverdlovsk Region (Shiryaev et al. 2010) and the Sokhondo
Reserve in the Transbaikal Region (Zhurbenko & Yakovchenko 2014).
*Taeniolella beschiana Diederich
3: on Cladonia arbuscula (thallus), 19.08.2012, MZ 12211b, LE 261391b.
*Taeniolella delicata M.S.Christ. & D.Hawksw.
10: on Lecanora intumescens (apothecia, thallus), 27.08.2012, MZ 12177a, LE 261216a.
Formerly known in Russia from Republic of Karelia (Alstrup et al. 2005). Lecanora intumescens is
a new host species. New to Asia.
*Thamnogalla crombiei (Mudd) D.Hawksw.
3: on Thamnolia vermicularis var. subuliformis (thallus), 17.08.2012, MZ 1258, LE 261072; MZ 1261, LE 261082;
8: on T. vermicularis var. subuliformis (thallus), 24.08.2012, MZ 1257, LE 261062.
*Tremella cetrariicola Diederich & Coppins
4: on Tuckermannopsis chlorophylla (thallus), 20.08.2012, MZ 12171b, LE 261356b; 13: on T. chlorophylla (thal-
lus), 28.08.2012, MZ 12132b, LE 261296b.
Tremella hypogymniae Diederich & M.S.Christ.
1: on Hypogymnia physodes (thallus), 14.08.2012, MZ 12114, LE 261477; 5: on H. physodes (thallus), 20.08.2012,
MZ 12107, LE 261407.
*Trichonectria anisospora (Lowen) van den Boom & Diederich
1: on Hypogymnia physodes (damaged thallus), 14.08.2012, MZ 12102, LE 261297.
Formerly known in Russia from Republic of Karelia (Alstrup et al. 2005). New to Asia.

392 Herzogia 27 (2), 2014
Vouauxiella lichenicola (Linds.) Petr. & Syd.
10: on Lecanora intumescens (hymenium of apothecia), 27.08.2012, MZ 12150, LE 261376.
Formerly known in Russia from the Kaliningrad Region (Dedkov et al. 2007) and Adygeya Republic
(Zhurbenko & Otte 2012).
*Xanthoriicola physciae (Kalchbr.) D.Hawksw.
18: on Xanthoria parietina (apothecia, thallus), 9.09.2012, MZ 1295, LE 261247.
Formerly known in Russia from the Leningrad Region (Kuznetsova et al. 2012). New to Asian
Russia.
Xenonectriella cf. leptaleae (J.Steiner) Rossman & Lowen
10: on Physcia aipolia (more or less damaged lobes), 27.08.2012, MZ 12199, LE 261334; on Physconia distorta
(bleached parts of thallus and discs of apothecia), 29.08.2012, MZ 12194, LE 261424.
Ascomata protruding only in the ostiolar area to occasionally sessile, dark red in the exposed parts,
subglobose, sometimes with distinct papilla, 200−250 µm diam. Exciple orange to orange yellow, not
changing colour in K and lactic acid. Ascospores ellipsoid to broadly ellipsoid, usually with rounded
ends, initially hyaline and smooth, then pale to medium reddish orange and prominently tubercu-
late, (12.3−)12.9−18.5(−23.5) × (6.7−)7.8−9.0(−9.7) µm, l/b = 1.4−2.4(−3.4) (n = 39; LE 261424)
or (11.3−)11.5−15.1(−17.4) × (7.2−)7.4−8.8(−9.3) µm, l/b = 1.4−2.0(−2.4) (n = 14; LE 261334),
(0 –)1-septate, usually slightly constricted at the septum, mostly with one large guttule in each cell,
uniseriate in the ascus (Fig. 3).
Fig. 3: Asci and ascospores of Xenonectriella cf. leptalea at different stages of maturation (in water, LE 261334).
Bar: 10 µm.

Zhurbenko & Kobzeva: Lichenicolous fungi from Northwest Caucasus, Russia 393
Rossman et al. (1999) reported for the species smaller ascospores, 8−12 × 6.5−8 µm, and an exciple
becoming pale brown to black in K and yellow in lactic acid. The examined material is conspecific with
the specimen on Physconia distorta from the Caucasus (LE 260846) reported by Zhurbenko & Otte
(2012) as Xenonectriella cf. leptaleae, which also has an exciple not changing colour in K and lactic
acid and bigger ascospores, (8 –)9.5 –13.5(–18) × (4.5 –)5.5 –7.5(– 8.5) µm. The examined collections
are reminiscent of Pronectria echinulata Lowen also growing on members of Physciaceae. However,
the latter clearly differs in having hyaline to slightly brownish, spinulose ascospores (Rossman et al.
1999, Etay o 1998).
So far the species was known in Asian Russia from Bastak Reserve in the Jewish Autonomous Region
(Zhurbenko 2014).
*Xenonectriella ornamentata (D.Hawksw.) Rossman
3: on Peltigera lepidophora (decaying lobes), 18.08.2012, MZ 1288, LE 260921.
Formerly known in Asian Russia from the Taimyr Peninsula and Chukotka (Zhurbenko 2009b).
Fig. 4: Conidiomata of Hainesia sp. on Physconia distorta intimately intermixed with ascomata of Zwackhiomyces
echinulatus (in K, LE 261434a). Bar: 10 µm.

394 Herzogia 27 (2), 2014
*Zwackhiomyces echinulatus Brackel
10: on Physconia distorta (thallus, margins and discs of apothecia), 27.08.2012, MZ 12187, LE 261204; MZ
12198a, LE 261434a.
Ascomata superficial, 75−175 µm diam., black, more or less glossy and rough, usually crow-
ded to concrescent. Ascospores hyaline to occasionally pale brown, (17.7−)20.3−25.3(−27.2) ×
(6.2−)7.9−10.3(−11.4) µm, l/b = (1.9−)2.2−2.8(−3.2) (n = 59), 4−8 per ascus. The infected thallus so-
metimes is slightly bleached, the infected hymenium distinctly damaged. In the specimen LE 261434a
the ascomata are intimately intermixed with the conidiomata of Hainesia sp. containing conidia c.
20−45 × 2−3.5 µm (Fig. 4.).
Formerly known from Italy (Brackel 2008) and the Tula Region of Russia (Zhurbenko &
Gudovicheva 2013). New to Asia.
Acknowledgements
Mikhail P. Zhurbenko is indebted to Alexandr E. Kovalenko, Vladimir G. Onipchenko, Vladimir N. Khramtsov, Viktor
A. Senik and Irina A. Senik for various kinds of help and support during his field studies. Wolfgang von Brackel,
Paul Diederich, Javier Etayo, Ave Suija, Uwe Braun and Kenan Yazici are thanked for the help in identification of
some critical specimens and/or valuable discussion. The study was financially supported by the grant of the Russian
Foundation for Basic Research No. 14-04-01031 «Lichenicolous fungi of Northwest Caucasus».
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Mycologie 32: 429−437.

396 Herzogia 27 (2), 2014
Zhurbenko, M. P. 2002. Arthonia triebeliae (Arthoniales), a new lichenicolous fungus from the Arctic. − Mycol.
Progress 1: 137–141.
Zhurbenko, M. P. 2009a. Lichenicolous fungi and some lichens from the Holarctic. – Opusc. Philolichenum 6:
87–120.
Zhurbenko, M. P. 2009b. Lichenicolous fungi and lichens from the Holarctic. Part II. – Opusc. Philolichenum 7:
121−186.
Zhurbenko, M. P. 2009c. New and interesting lichenicolous hypocrealean fungi from the Northern Hemisphere. −
Sydowia 61: 177–188.
Zhurbenko, M. P. 2012. New records of lichenicolous fungi from «Stolby» Reserve (Krasnoyarsk Territory). −
Novitates Systematicae Plantarum non Vascularium (St.-Petersburg) 46: 92–95. (in Russ., Engl. summ.)
Zhurbenko, M. P. 2013a. A first list of lichenicolous fungi from India. – Mycobiota 3: 19 –34.
Zhurbenko, M. P. 2013b. Lichenicolous fungi and some allied lichens from the Canadian Arctic. – Opusc.
Philolichenum 12: 180−197.
Zhurbenko, M. P. 2014. Lichenicolous fungi from Far East of Russia. − Folia Cryptog. Estonica 51: 113 –119.
Zhurbenko, M. P. & Alstrup, V. 2004. Lichenicolous fungi on Cladonia mainly from the Arctic. − Acta Univ. Ups.
Symb. Bot. Ups. 34: 477– 499.
Zhurbenko, M. P. & Davydov, E. A. 2000. Lichenicolous fungi and some lichens from the Russian Altai, southern
Siberia. − Folia Cryptog. Estonica 37: 109 –118.
Zhurbenko, M. P. & Gudovicheva, A. V. 2013. Zwackhiomyces echinulatus and other lichenicolous fungi from Tula
Region of Russia. − Mikologiya i fitopatologiya [Mycology and phytopathology, St.-Petersburg] 47: 19 –20. (in
Russ., Engl. summ.).
Zhurbenko, M. P. & Otnyukova, T. N. 2001. Lichenicolous fungi from the Sayan-Tuva Mountains, Southern
Siberia, Russia. − Folia Cryptog. Estonica 38: 79 – 84.
Zhurbenko, M. P. & Otte, V. 2012. Lichenicolous fungi from the Caucasus: new records and a first synopsis. –
Herzogia 25: 235 –244.
Zhurbenko, M. P. & Vershinina, S. E. 2014. Opegrapha bryoriae sp. nov. and other lichenicolous fungi from Asian
Russia. – Herzogia 27: 93−109.
Zhurbenko, M. P. & Yakovchenko, L. S. 2014. A new species, Sagediopsis vasilyevae, and other lichenicolous
fungi from Zabaikal’skii Territory of Russia, southern Siberia. – Folia Cryptog. Estonica 51: 121–130.
Zhurbenko, M. P. & Zhdanov, I. S. 2013. Melaspilea galligena sp. nov. and some other lichenicolous fungi from
Russia. – Folia Cryptog. Estonica 50: 89 –99.
Zhurbenko, M. P., Hermansson, J. & Pystina, T. N. 2012. Endococcus incrassatus new to Eurasia and some other
lichenicolous fungi from the Komi Republic of Russia. – Graphis Scripta 24: 36 –39.
Zhurbenko, M. P., Himelbrant, D. E., Kuznetsova, E. S. & Stepanchikova, I. S. 2012. Lichenicolous fungi from
the Kamchatka Peninsula, Russia. – Bryologist 115: 295−312.
Manuscript accepted: 11 October 2014.
Addresses of the authors
Mikhail P. Zhurbenko, Laboratory of the Systematics and Geography of Fungi, Komarov
Botanical Institute, Russian Academy of Sciences, Professor Popov 2, St.-Petersburg, 197376
Russia. E-mail: zhurb58@gmail.com
Anastasiya A. Kobzeva, same address. E-mail: anastasiakobzeva9023@gmail.com