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90 Herzogia 33 (1), 2020: 90 –138
Lichens and allied non-lichenized fungi of virgin forests in the
Caucasus State Nature Biosphere Reserve (Western Caucasus,
Russia)
Gennadii *, Jan , Irina &
Abstract: , G.,, J.,, I.,, Z. &, J. 2020. Lichens and allied
non-lichenized fungi of virgin forests in the Caucasus State Nature Biosphere Reserve (Western Caucasus, Russia). –
Herzogia 33: 90 –138.
We report on 659 epiphytic and epixylic species recorded from seven one-hectare plots established along an altitudinal
gradient in a virgin forest of the Caucasus State Nature Biosphere Reserve. A total of 564 species are lichens, 61 are li-
chenicolous fungi and 34 are allied non- or facultatively lichenized fungi. one hundred forty – nine species (116 lichens,
17 lichenicolous and 16 saprophytic fungi) are new to the Northern Caucasus, including 133 species (104 lichens, 15
lichenicolous and 14 saprophytic fungi) that are new to the Caucasus Mountains. Fifty species are reported from Russia for
the first time: 37 lichens (Andreiomyces obtusaticus, Bacidina mendax, Biatora aegrefaciens, B. bacidioides, B. chrysan-
thoides, Biatorella dryophila, Buellia iberica, Cliostomum haematommatis, Endohyalina ericina, Fellhanera christian-
senii, Gyalidea minuta, Japewia aliphatica, Lecanora barkmaniana, L. subravida, Lecidea strasseri, Leptogium hibern-
icum, Lithothelium hyalosporum, L. phaeosporum, L. septemseptatum, Loxospora cristinae, Melanelixia epilosa, Micarea
nowakii, M. perparvula, Opegrapha trochodes, Orcularia insperata, Parvoplaca servitiana, Phylloblastia inexpectata,
Psoroglaena stigonemoides, Ptychographa xylographoides, Ramonia dictyospora, R. luteola, Rinodina polysporoides,
Thelopsis flaveola, Topelia jasonhurii, Verrucaria hegetschweileri, Wadeana minuta, Waynea giraltiae), nine lichenicol-
ous fungi (Arthonia vorsoeensis, Didymocyrtis melanelixiae, Epigloea urosperma, Muellerella polyspora, Phacographa
zwackhii, Pronectria pilosa, Rhymbocarpus pubescens, Taeniolella friesii, Unguiculariopsis acrocordiae) and four non-
lichenized saprophytic fungi (Cyrtidula major, Karschia cezannei, Kirschsteiniothelia recessa, Pseudotryblidium neesii).
The ratio of macrolichens ranges between 26.5 – 40 % and rises with elevation. Lichens with a trentepohlioid photobi-
ont are represented by 15 –51 species per plot and their species richness decreases with elevation. The species richness
of cyanolichens is substantial in all plots (15 –28 species) reflecting a negligible effect of acidification/air pollution.
Beech and fir are the most preferred phorophytes, but the vast majority of lichen species have low substrate specificity.
Species richness per plots ranged between 236 and 379. The highest richness was found in a plot outside the Caucasian
Reserve and we recommend its inclusion into the protected area.
Zusammenfassung: , G.,, J.,, I.,, Z. &, J. 2020. Flechten
Rußland). – Herzogia 33: 90 –138.
wurden 659 epiphytische und epixyle Arten nachgewiesen. Davon sind 564 Flechten, 61 lichenicole Pilze und 34
-
Andreiomyces ob-
tusaticus, Bacidina mendax, Biatora aegrefaciens, B. bacidioides, B. chrysanthoides, Biatorella dryophila, Buellia
* corresponding author
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 91
iberica, Cliostomum haematommatis, Endohyalina ericina, Fellhanera christiansenii, Gyalidea minuta, Lecanora
barkmaniana, L. subravida, Lecidea strasseri, Leptogium hibernicum, Lithothelium hyalosporum, L. phaeosporum, L.
septemseptatum, Loxospora cristinae, Melanelixia epilosa, Micarea nowakii, M. perparvula, Opegrapha trochodes,
Orcularia insperata, Parvoplaca servitiana, Phylloblastia inexpectata, Psoroglaena stigonemoides, Ptychographa
xylographoides, Ramonia dictyospora, R. luteola, Rinodina polysporoides, Thelopsis flaveola, Topelia jason-
hurii, Verrucaria hegetschweileri, Wadeana minuta, Waynea giraltiae), 9 lichenicole Pilze (Arthonia vorsoeensis,
Didymocyrtis melanelixiae, Epigloea urosperma, Muellerella polyspora, Phacographa zwackhii, Pronectria pilosa,
Rhymbocarpus pubescens, Taeniolella friesii, Unguiculariopsis acrocordiae) und 4 nicht lichenisierte saprophytische
Pilze (Cyrtidula major, Karschia cezannei, Kirschsteiniothelia recessa, Pseudotryblidium neesii).
-
-
ne Rolle spielen. Geringe Artenvielfalt und geringe Abundanz nitrophiler Arten indizieren unbedeutende Immissionen
-
ße Mehrheit der Flechten zeigt eine geringe Substratspezifität. Die Artenvielfalt je Plot bewegt sich zwischen 236
und 379. Die höchste Vielfalt wurde in einem Plot außerhalb des Reservates festgestellt und wir empfehlen dessen
Aufnahme in das Schutzgebiet.
Key words: Biodiversity, epiphytes, hot-spots, lichen inventory, lichenized and lichenicolous fungi.
Introduction
The highest biodiversity among terrestrial biomes is associated with forests ( 2009).
However, most woodlands in Europe today are managed plantations, often with non-native
trees and with very low biodiversity. Primeval forests have become rare and are one of the
most endangered habitats ( et al. 2018a). The Russian North-western Caucasus is
outstanding with its 1.3 million hectares of undisturbed primeval forests ( 2017).
105 thousand hectares of undisturbed primeval fir and fir-beech forests are protected in the
Caucasus State Nature Biosphere Reserve (Caucasus Reserve in following text, et al.
1990). The Western Caucasus is a place where endangered, rare, endemic and relict plant and
animal species are concentrated ( et al. 2006). It is one of the world’s biodiversity
hot-spots ( & 2002) and one of the most diverse regions in Russia (
et al. 2001). It forms a border between two phytogeographical regions: the Euxinian province
of the Euro-Siberian region and the Hyrcanian province of the Irano-Turanian region. Its loca-
tion within the mid-latitudinal temperate zone, high topographical variation and proximity to
-
tions to alpine and nival conditions. It also harbors a high diversity of tree species and forest
types ( et al. 2011, et al. 2018).
Although lists of lichens collected in NW Caucasus have been already published by
(1997), & (2004, 2014, 2015),
& (2005), and (2005, 2007), fir-beech virgin forests in the Caucasus
(1997) published a lichen
-
ported from the territory of the Caucasus Reserve. Subsequent floristic research resulted in 518
& 2004). More
recently, &
Plateau (North Division of the Caucasus Reserve), mainly from alpine habitats, but also from
2) in the northern part of the Kamennoe
More (Stone Sea) Range where 330 epiphytic and epixylic species were recorded (including
30 species of lichenicolous fungi). In the Western and Southern parts of the Caucasus Reserve,
92 Herzogia 33 (1), 2020
and epixylic species were recorded ( & 2014, 2016). Another
90 species of lichenicolous fungi were reported for the Caucasus Reserve by &
(2016) and (2017). Altogether, 1050 species (among them about 550
lichens, 160 lichenicolous fungi and 21 lichen-allied saprobic fungi.
In 2016, we pursued lichen inventories along an altitudinal gradient in untouched fir-beech
forests of the northern part of the Caucasus Reserve, focused on local lichen diversity hot-
spots (following et al. 2018). Ecological and conservational implications of the
results have been published separately ( et al. 2019b), but the impressive floristic
results merit a publication of its own and are published here.
Study area
The Caucasus State Nature Reserve was established in 1924 and declared as an UNESCO
Biosphere Reserve in 1979 ( et al. 1990). In 1999, the entire territory of the Caucasus
Reserve (280 000 hectares) was included in the UNESCO World Natural Heritage Sites as
“Western Caucasus”. Approximately two thirds of the territory of the Caucasus Reserve is
covered with virgin forests; the rest of the area is occupied by subalpine and alpine meadows,
nival communities, and glaciers. Coniferous forests in the NW Caucasus are mainly formed by
Nordmann fir (Abies nordmanniana). Fir and fir-beech forests are the predominant habitat at
600 –1800 m above the sea level. Fir trees may have stunning sizes; more than 60 m in height
and about two meters in diameter at breast height ( et al. 1990). The eastern beech
(Fagus orientalis) reaches smaller sizes; slightly above 40 m in height and 100 cm in diam-
eter. Fir trees often reach 400 –500 years of age, and sometimes even 700 years, the eastern
beech reaches 280 –300 years but sometimes even 380 years ( 2011). Trautvetter
maple (Acer trautvetteri) is another forest dominant and other trees involved are: maple tree
species (Acer campestre, A. laetum, A. platanoides), lime tree (Tilia begoniifolia), hornbeam
(Carpinus betulus), ash (Fraxinus excelsior), birch (Betula litwinowii, B. pubescens) and pine
(Pinus kochiana, = P. sylvestris) ( et al. 1990). Other tree and shrub species are listed
at the beginning of the lichen list with abbreviations (see below).
According to the classification of terrestrial ecosystems (biomes) by et al. (2018), the
study area belongs to the North-Western Caucasus regional orobiome of the North Caucasian
group with nemoral coniferous-deciduous and deciduous forests. Altitude belts in this orobi-
Quercus robur) – 200 –350 m
Quercus petraea) – 350 –500 m a.s.l.; 3) middle
mountain forests dominated by eastern beech (Fagus orientalis) – 500 – 650 m a.s.l.; 4) middle
mountain beech-fir forests with occassional occurrences of Acer pseudoplatanus, Carpinus
betulus, Tilia begoniifolia, Ulmus glabra etc. – 650 –1500 m a.s.l.; 5) mountain beech-fir for-
ests with Acer trautvetteri, Betula litwinowii, Sorbus aucuparia, Salix sp. etc. – 1500 –1800 m
Rhododendron caucasicum and sedge-herb meadows – 1800 –2300 m a.s.l. and 8) alpine belt –
2300 –3000 m a.s.l. ( 2016).
In the surveyed area, the Belaya River basin at the village Guzeripl (Fig. 1) at 670 m a.s.l., the
average annual temperature is about 8.2°C (-2°C in January, 18°C in July), the annual aver-
age precipitation 1130 mm (maximum 1540 mm). At 1400 –1700 m a.s.l., the average annual
temperature is about 5°C, (-3°C in January, 15°C in July) and the annual average precipitation
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 93
1200 mm (maximum 2000 mm). At 1810 –2020 m a.s.l., the temperature is about 3°C, (-7°C
in January, 13°C in July), the precipitation 2000 mm (maximum 2800 mm; et al.
1982, 2009).
Materials and methods
Our field research was carried out from 6th to 18th June 2016, and organized with the aim to detect
a maximum number of lichen species in the study area. We concentrated efforts on epiphytic and
epixylic lichen species, allied non- or facultatively lichenized fungi and lichenicolous fungi. The
research was performed in seven square 1-hectare plots positioned in structurally diverse for-
est habitats along an altitudinal gradient from 700 to 1920 m (Fig. 1). Criteria for plot selection
followed the hot-spot approach described by et al. (2018). The survey was done as a
multi-expert inventory by five of the authors (GU, IU, JM, JV, ZP), following methods described
by et al. (2016). Approximately 40 person-days were spent in the field. GPS coor-
dinates are defined in the WGS-84 system. Specimens were identified using primarily routine
methods (light microscopy and spot tests). The researchers identified their specimens individu-
-
mens, especially sterile crusts, were identified using thin layer chromatography (in solvents A,
B’ and C) according to the methods summarized by et al. (2001). Some specimens were
Fig. 1. Maps showing the surveyed region and locations of plots 1–7 (basis of maps: http://opengeodata.ru).
94 Herzogia 33 (1), 2020
Fig. 2. Plot 1 – forest with Abies nordmanniana, Acer campestre, Alnus glutinosa, Carpinus betulus, Corylus avellana,
Fagus orientalis and Quercus
sequenced for nuclear ITS (ITS1, 5.8S, ITS2) and / or mitochondrial SSU DNA barcodes, fol-
lowing methods by et al. (2018b). A standard nucleotide BLASTN search (
et al. 1990, Appendix) was used for a confirmation of species identity.
The list of taxa is in alphabetical order. The data presented for each species include: presence
on plots, substrata, collectors and a count of voucher specimens. Voucher specimens were col-
lected for most of the recorded species and are deposited in PRA (JV, ZP), LE (IU) and in the
List of sampled 1-hectare plots
1.
or dense forest with frequent Abies nordmanniana, Acer campestre, Alnus glutinosa, Carpinus betulus, Corylus
avellana, Fagus orientalis and Quercus
2.
sparse forest with frequent Abies nordmanniana, Carpinus betulus, Fagus orientalis, Quercus spec. and Tilia
begoniifolia, and with forest gaps covered by Rhododendron ponticum
3.
Abies nordmanniana, Acer trautvetteri and Fagus orientalis
& 14 VI 2016.
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 95
4. Acer trautvetteri with spots of dense
forest dominated by Abies nordmanniana and Fagus orientalis
12 & 13 VI 2016.
5. Betula litwinowii,
with single trees of Abies nordmanniana and Fagus orientalis and with spots of Rhododendron caucasicum
6. Abies
nordmanniana and Fagus orientalis, with a forest gap covered by Rhododendron ponticum scrubs, alt.710 –740 m
7.
-
dant trees: Abies nordmanniana, Acer trautvetteri, Betula litwinowii, Fagus orientalis, Sorbus aucuparia, Salix
caprea, Taxus baccata, Ulmus glabra, and also Pinus kochiana
16, 17 & 18 VI 2016 (Fig. 3). This site is not protected. It is situated 150 m beyond the northern boundary of the
Caucasus Reserve.
Results
New records for Russia, Caucasus or North Caucasus are indicated as R, C and NC, respec-
tively, before the name of the respective species. The nomenclature mainly follows
et al. (2018), (2010) [absent taxa in the previous reference] and &
(2018), except the taxa from the families Ramalinaceae and Teloschistaceae (
Fig. 3. Plot 7 – mixed forest with predominant Abies nordmanniana, Acer trautvetteri and Betula litwinowii on lime-
96 Herzogia 33 (1), 2020
et al. 2013, et al. 2018) along with few relevant recent novelties ( et al.
2018, & 2018, et al. 2019). Several recently described taxa not in-
taxon and amended with a reference. Substrate and abbreviations:
AC – Acer campestre, AG – Alnus glutinosa, AN – Abies nordmanniana, AP – Acer platanoides, AT – Acer trautvet-
teri, BL – Betula litwinowii, CA – Corylus avellana, CB – Carpinus betulus, FE – Fraxinus excelsior, FO – Fagus
orientalis, IC – Ilex colchica, JC – Juniperus communis, LO – Lonicera sp., PK – Pinus kochiana, QU – Quercus
sp., RP – Rhododendron ponticum, SA – Sorbus aucuparia, SC – Salix caprea, SN – Sambucus nigra, ST – Sorbus
torminalis, TB – Taxus baccata, Ti – Tilia begoniifolia, UG – Ulmus glabra, VA – Vaccinium arctostaphylos, VO –
Viburnum orientale, bry – bryophytes, le – leaves, ne – needles, tw – twigs.
Lichens
Absconditella lignicola
Acolium inquinans (Sm.) A.Massal. – 4: AN/snag (JM1, ZP1).
Acrocordia cavata (Ach.) R.C.Harris – 1, 7: AN, AT, CB, FE, SN (GU2, IU1, JM1, JV2, ZP2).
Acrocordia gemmata (Ach.) A.Massal. – 1, 2, 3, 4, 5, 6, 7: AN, AP, AT, CA, CB, FO, QU, Ti, UG (GU11,
IU4, JM4, JV7, ZP11).
Agonimia allobata (Stizenb.) P.James – 1, 2, 3, 5, 6: AN, FO, QU, Ti (IU1, JM2, JV6, ZP5).
Agonimia borysthenica Dymytrova, Breuss & S.Y.Kondr. – 1, 2, 6: AG, FO, stump (GU2, JM3, JV3,
ZP6). The species was recorded only in ‘lowland’ forest plots below 1000 m. Quite recently it has been
reported as new to Russia and Caucasus from Caspian forests in Dagestan ( et al. 2017).
Agonimia flabelliformis Halda, Czarnota & Guz.-Krzem. – 1, 2, 6: AG, AN, log, stump (GU3, IU1, JM2,
JV5, ZP3).
Agonimia repleta Czarnota & Coppins – 1, 2, 3, 4, 5, 6: AG, AN, AT, CB, FO, UG (GU2, IU3, JM2,
JV9, ZP6).
Agonimia tristicula (Nyl.) Zahlbr. – 1, 2, 3, 4, 7: AT, FO, QU, Ti, UG (IU1, JM4, JV4, ZP6).
Agonimia vouauxii (B. de Lesd.) M.Brand & Diederich – 7: UG/bry (JV1).
Alectoria sarmentosa (Ach.) Ach. – 3: AN (IU1, JV1).
Alyxoria ochrocheila (Nyl.) Ertz & Tehler – 4: FO (JM1, JV1).
Alyxoria varia (Pers.) Ertz & Tehler – 1, 2, 3, 4, 6, 7: AN, AP, AT, CB, FE, FO, QU, UG, log, snag
(GU13, IU12, JM2, JV15, ZP5).
Amandinea punctata (Hoffm.) Coppins & Scheid. – 1, 2, 3, 4, 5, 6, 7: AN, AT, BL, ST, TB, snag, stump
(GU13, IU5, JV10, ZP4).
Amandinea aff. punctata – 4, 7: AN (JV4, ZP3). Apothecia resemble Buellia schaereri or Amandinea
punctata, but yellow, K+ red pigment present in hymenium. Specimens were seen by H. Mayrhofer,
but left unidentified. The mtSSU sequence of the sample JV16429 is very distinct from sequences of
B. schaereri and A. punctata; sequences of Diplotomma species are the most similar (97 % similarity)
according to the BLASTN-search.
Anaptychia ciliaris (L.) Körb. – 2, 3, 4, 5, 7: AN, AT, BL, FO, SA, Ti, UG (GU2, IU3, JM1, JV4, ZP1).
Anaptychia crinalis
RC Andreiomyces obtusaticus
atic acid.
Anisomeridium biforme (Borrer) R.C.Harris – 1, 4, 6: AG, AT, CB, FO (GU3, JV1, ZP2).
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 97
Anisomeridium polypori (Ellis & Everh.) M.E.Barr – 1, 2, 3, 4, 5, 6, 7: AN, AT, CB, FO, SA (GU6, JM4,
JV9, ZP7).
C Anzina carneonivea (Anzi) Scheid. – 5: BL/wood (JM2).
C Aquacidia trachona (Ach.) Aptroot – 1: QU (ZP1).
Arthonia cf. bueriana (J. Lahm ex Arnold) Zahlbr. – 4: AN (IU1, JV1, ZP1). In Russia and the Caucasus,
this species was collected before from Krasnodar Territory (specimen in GLM; V. Otte in litt.). Our
-
et al. (2018) A. bueriana is a non-lichenized species. Our specimens are lichenized with trentep-
ohlioid algae and fit best the description of this species provided in the monograph of Arthoniaceae
by (1937), who regarded this taxon for a lichen containing algae of Trentepohlia type.
Optionally the Caucasian material could represent epiphytic populations of usually saxicolous taxon
Arthonia granitophila Th.Fr. Caucasian material deserves further study. Specimen ZP23213 is filed
under the latter name in the herbarium PRA.
Arthonia didyma Körb. – 1, 2, 3, 6, 7: AN, CB, FO, SA, Ti (GU5, IU4, JM3, JV7, ZP5).
Arthonia dispersa (Schrad.) Nyl. – 1, 2, 5: AN, CA, FO (GU1, IU1, JV1).
C Arthonia faginea Müll.Arg. – 3, 4, 5, 7: AN, AT, CB, FO, SC, UG (GU1, JM4, JV8, ZP6).
Arthonia fuliginosa (Turner & Borrer) Flot. – 2: RP (ZP1). Small specimen ZP22922 with purplish-
(e.g. 1937) and it was also confirmed by A. Frisch. This species has been reported for the
first time for the Caucasus and Russia from the neighbouring territory of the Republic of Adygea
(-
Arthonia helvola (Nyl.) Nyl. – 4, 7: AN, wood of snag (JM1, ZP2)
C Arthonia incarnata Th.Fr. ex Almq. – 4, 7: AN, snag (IU1, JV2).
Arthonia mediella Nyl. – 3, 4, 5, 6, 7: AN, AT, FO, SA, snag (GU3, JM2, JV3, ZP8).
C Arthonia patellulata Nyl. – 7: AT, SA (GU1, ZP1).
Arthonia punctiformis Ach. – 1, 2, 3, 4, 5, 6, 7: BL, CA, FO, SA, UG (GU1, IU1, JV9).
Arthonia radiata (Pers.) Ach. – 1, 2, 3, 4, 5, 6, 7: AN, AT, BL, CB, FO, SA, Ti, UG (GU17, IU11, JM2,
JV13, ZP6).
Arthonia ruana A.Massal. – 1, 2: IC/tw, Q, UG/tw (JV1, ZP2).
Arthonia spadicea Leight. – 1, 2, 3, 4, 5, 6, 7: AN, AG, AT, CB, FO, QU, SC, ST, log, snag (GU6, IU2,
JM1, JV6, ZP1).
Arthonia vinosa Leight. – 1, 3, 7: AN, AT, PK, snag (GU1, IU2, JV4, ZP3). Some specimens produce
Trimmatostroma; sporodochia con-
together with sporodochia.
Arthopyrenia analepta (Ach.) A.Massal. – 1, 4, 5, 6, 7: AN/tw, FO/tw, BL, SA (GU3, JM2, JV3, ZP2).
Arthopyrenia cerasi (Schrad.) A.Massal. – 2, 7: SA/tw, Ti/tw (GU1, JV2).
Arthopyrenia salicis A.Massal. – 2, 3, 5: AN/tw, FO/tw, UG/tw (GU3, JV1).
C Arthopyrenia subcerasi (Vain.) Zahlbr. – 1, 4, 6: AT, FO/tw (GU1, JV2).
C Arthothelium scandinavicum Th.Fr. – 3, 4, 6, 7: AN, AT (GU2, IU1, JM3, JV7, ZP6).
98 Herzogia 33 (1), 2020
Arthothelium spectabile Flot. ex A.Massal. – 1, 2: CB, QU, Ti (JM1, JV1, ZP1).
Aspicilia cinerea auct. brit. (cf.) – 4: FO (ZP1). TLC: norstictic acid. The specimen ZP21467 was coll-
ected at complete basis of leaning Fagus. The identification is tentative because no apothecia were
glossy prothallus and pycnidia containing relatively long conidia, c. 10 –15 × 1 µm, suggest this broad-
ly circumscribed, primarily saxicolous taxon following et al. (2009).
Athallia cerinelloides (Erichsen) Arup, Frödén & Søchting – 4, 7: AT, FO, LO, VO (IU1, JM1, JV4,
ZP3).
Athallia pyracea (Ach.) Arup, Frödén & Søchting – 3, 4, 5: AT, SN (JM1, JV2).
Bacidia absistens (Nyl.) Arnold – 1: AG (IU1).
Bacidia albogranulosa
Central and Eastern Europe (see et al. 2018a).
Bacidia arceutina (Ach.) Arnold – 1, 2, 6: AC, AG, AN, CA, CB, FE, FO, QU, Ti, UG, log (GU8, IU1,
JM3, JV7, ZP7).
Bacidia biatorina (Körb.) Vain. – 1, 2: AN, FO, QU, RP (GU1, IU2, ZP1).
Bacidia fraxinea Lönnr. – 1, 3, 6: AN, AT, SN (GU1, IU2, JM2).
Bacidia friesiana (Hepp) Körb. – 1, 3, 6: AN, FO, SN (GU3, IU1, JM2, JV2, ZP1).
Bacidia laurocerasi (Delise ex Duby) Zahlbr. – 1, 2, 3, 6: AG, AN, CA, CB, FO, RP, UG (GU5, JM2,
JV5, ZP7).
Bacidia polychroa (Th.Fr.) Körb. – 1, 2: AC/tw, AP, CA, CB, FE, SN (GU2, IU4, JV4, ZP2).
C ‘Bacidia’ propinqua
of Bilimbia sabuletorum but it is a distinct species of the genus Bilimbia
Bacidia rosella (Pers.) De Not. – 1, 2, 3, 6: AN, CB, FO, Ti (GU5, IU3, JM2, JV1, ZP3).
Bacidia rubella (Hoffm.) A.Massal. – 1, 2, 3, 4, 6, 7: AN, AT, CA, FO, QU, SA, UG (GU6, IU7, JV4,
ZP3).
Bacidia suffusa (Fr.) A.Schneid. – 1: AN (IU1).
Bacidina apiahica
Bacidina assulata
Bacidina chloroticula
wood (GU3, JM2, JV4, ZP5).
Bacidina delicata
RC Bacidina mendax Czarnota & Guz.-Krzem. – 1, 6: AC/tw, AN/tw, CA, UG/tw (JM1, JV3, ZP3). B.
mendax is new to Russia, but the record of B. neosquamulosa from the Leningrad region may belong
to that species as its image and short description provided by et al. (2017) fit better to B.
mendax ( & 2018).
C Bacidina modesta
Bacidina phacodes
Bacidina pycnidiata (Czarnota & Coppins) Czarnota & Guz.-Krzem. – 1, 3, 4, 6, 7: AT/bry, FO/liver-
worts, LO, log, snag (GU1, JM1, JV6, ZP3).
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 99
Bactrospora dryina (Ach.) A.Massal. – 1, 6: AN, AG, QU (GU2, IU2, JM3, JV2, ZP1).
Bellicidia incompta
(GU1, IU1, JM3, JV2, ZP2).
RC Biatora aegrefaciens Printzen – 3: AN/wood of branch (ZP1). It is a well recognizable species within
the genus due to its three-septate ascospores, amyloid (I + bluish) exciple and presence of gyrophoric
acid in apothecia (
( 1995, et al. 2002, et al. 2018). The taxon is a promising bioindicator
species. Due to its rarity and ecological requirements it was suggested to be a relictual old-growth
forest species ( et al. 2002).
Biatora albohyalina (Nyl.) Bagl. & Carestia – 1, 2, 3: AN, snag (GU1, IU2, ZP2).
RC Biatora bacidioides Printzen & Tønsberg – 1, 2: CB (JM1, ZP1). TLC: gyrophoric acid, argopsin.
This sorediate taxon with affinities to Biatora beckhausii was described from Picea orientalis forests
& 2003) hence the presence
in the Caucasus was expected. Recently it was recorded also from humid Fagus sylvatica forests in
the Eastern Carpathians ( et al. 2018b, et al. 2018). The collected Caucasian ma-
terial was sterile and identified by its extensively sorediate thalli containing both gyrophoric acid and
argopsin.
Biatora beckhausii
JV6, ZP8).
C Biatora chrysantha (Zahlbr.) Printzen – 1, 2, 4, 5, 7: AN, AT, BL, FO, RP, log (GU1, IU7, JM9, JV10,
ZP8). TLC: gyrophoric acid. Identity of one sterile specimen (JV16141) confirmed by ITS sequence.
Since the majority of vouchers is sterile, we cannot exclude that a part of the material from higher
altitudes belongs to the following species.
RC Biatora chrysanthoides Printzen & Tønsberg – 5: BL, snag (JM1, ZP4). TLC: gyrophoric acid in the
only in the subalpine forest plot during our survey. The species has been reported only once (Sweden,
et al. 2019) since the original description from northern Norway and Pacific North America
( & 2003). It is morphologically hardly distinguishable from Biatora chrysantha,
if not fertile. Fortunately, the Caucasian specimens are fertile and with apothecia containing gyro-
phoric acid, the main discriminating feature from Biatora chrysantha ( & 2003).
Biatora efflorescens (Hedl.) Räsänen – 3, 4, 5, 7: AG, AN, AT, BL, CB, FO, PK, SA, TB, log, snag
(GU15, IU19, JM5, JV11, ZP15). The specimen on wood of snag (JV16449) with unusual phenotype
resembling Violella fucata. Identification confirmed by ITS sequence data.
Biatora fallax Hepp – 4, 5, 7: AN, BL, snag (IU1, JM1, JV7, ZP1).
Biatora globulosa
JV8, ZP12).
Biatora helvola Körb. ex Hellb. – 7: FO (ZP1).
Biatora meiocarpa (Nyl.) Arnold – 7: AN/tw (ZP1). The specimen ZP23254 is macroscopically similar
to B. helvola or extreme morphs of Lecania cyrtella s. lat. with a receding thalline margin, but with
distinct, almost rectangular lumina of excipular hyphae as large as 7 × 3.5 µm in water, enlarged
paraphyse-tips and presence of filiform conidia (30 × 1.5 µm). The species was identified based on the
description in (1994).
Biatora aff. meiocarpa – 5: FO (ZP1). The specimen ZP23529 is a non-sorediate taxon with an immersed
thallus and small pallid apothecia containing small ellipsoid ascospores, not exceeding 10 × 3.5 µm.
100 Herzogia 33 (1), 2020
Apparently a member of the B. meiocarpa-group (sensu 2014) based on the exciple charac-
ter and enlarged paraphyses ends.
Biatora mendax Anzi – 1, 2, 3, 4, 5, 6, 7: AN, BL, CB, FO, QU, RP, UG (GU10, IU1, JM4, JV8, ZP22).
Biatora ocelliformis (Nyl.) Arnold – 1, 2, 3, 4, 6, 7: AN, AP, AT, CB, FO, RP, SA, TB, log, snag, stump
(GU12, IU5, JV5, ZP11).
Biatora pontica Printzen & Tønsberg – 1, 2, 3, 6: AG, AN, CB, FO, RP, UG (GU5, IU2, JM8, JV5,
-
mens were collected in lowland sampling plots. The identification of two sterile specimens (JM10212,
JV17846) was supported by mtSSU and ITS sequences.
Biatora radicicola Printzen, Palice & J.P.Halda – 4: FO (ZP5). One of the Caucasian specimens is para-
type of the species (ZP21105). The characteristic ecology, e.g. exposed roots and bases of trees subjec-
ted to water spray and/or sites with a prolonged snow cover (see et al. 2016) was emended
et al. 2019).
C Biatora vacciniicola (Tønsberg) Printzen – 5: FO (JM1, ZP1). TLC: gyrophoric acid. Sterile speci-
mens were compared to typical vouchers from Scandinavia. It is a mostly northern species preferring
-
tains ( 1995, & 1999, et al. 2018) reaching to Northern Ural Mts
on the northeast ( et al.
European range of the species to the southeast.
Biatora vernalis (L.) Fr. – 1, 6: AG/bry, CB, FO/bry (IU2, JM2, JV2, ZP2).
Biatora veteranorum Coppins & Sérus. – 1, 2, 3, 7: AN, snag, stump (GU3, IU2, JM2, JV4, ZP3).
RC Biatorella dryophila
without margin, 0.15 – 0.2 mm diam.; epithecium granular, K+ purple; spores globose 1.2–2 µm diam.
& (1977) and Piccolia ochrophora, but we
consider it a separate species.
Biatoridium delitescens (Arnold) Hafellner – 1: AC/branch (ZP2). Only recently recorded for the first
time from the Caucasus ( & 2016)
Biatoridium monasteriense J.Lahm ex Körb. – 1, 2, 3, 4, 5, 6, 7: AT, FO, UG (GU3, IU1, JM3, JV6,
ZP5). Only recently recorded for the first time from the Caucasus ( &
2018).
C Bibbya vermifera
Bilimbia microcarpa (Th.Fr.) Th.Fr. – 1, 5, 7: FO, UG (JM1, JV1, ZP2).
Bilimbia sabuletorum (Schreb.) Arnold – 1, 2, 3, 4, 5, 7: AC, AN, AP, AT, CA, FO/bry, QU, UG/bry
(GU2, IU2, JM2, JV9, ZP7).
Blastenia anatolica
is a paratype of the species (
mountains.
Blastenia herbidella
IU18, JM1, JV11, ZP4). Identification of four specimens confirmed with ITS sequences (
et al. 2019a).
Blastenia hungarica (H.Magn.) Arup, Søchting & Frödén – 2, 3, 5, 7: AN/tw, CB/tw, FO/tw, TB/tw
(GU1, IU1, JM1, JV2, ZP1).
Bryobilimbia hypnorum
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 101
Bryobilimbia sanguineoatra
CB, FO/bry, log (IU3, JM4, JV9, ZP6). Distinguished from B. hypnorum by mainly simple, narrowly
ellipsoid ascospores, < 5 µm wide, with a smooth instead of a warted perispore ( et al. 2014).
Bryoria americana
Bryoria bicolor
Bryoria capillaris
JM4, JV2, ZP3).
Bryoria fuscescens
JM3, JV3, ZP2).
Bryoria implexa
IU4, ZP3).
Bryoria kuemmerleana
Bryoria nadvornikiana
SA, log (GU17, IU12, JM3).
Bryoria smithii
negative Pd spot reaction. It resembles B. bicolor
Bryoria vrangiana
Bryostigma muscigenum (Th.Fr.) Frisch & G.Thor (incl. Arthonia microsticta Vain.) – 1, 2: AN/ne
(GU2, IU1, JV1, ZP3); 4, 7: AN, FO/bry, log (JM2, JV4, ZP2). Specimens on Abies needles may be
referrable to Arthonia microsticta, but B. muscigenum and A. microsticta are possibly conspecific.
& (2009) describe an wide range of substrata for B. muscigenum (sub Arthonia
muscigena) including Abies needles for B. muscigenum. Identity of one specimen on wood (ZP21403)
confirmed by A. Frisch.
C Buellia arborea Coppins & Tønsberg – 7: BL, PK/wood, snag (ZP2). The identification of the sterile
sorediate specimen ZP22088 was confirmed by TLC (atranorin, placodiolic acid). By its delimited
(semi-)urceolate soralia the species is quite similar to Buellia griseovirens and Xylographa vitiligo
when sterile, but well recognizable by its chemistry. It preferably grows on hard, slowly decaying
wood of conifers ( 1992a).
Buellia disciformis (Fr.) Mudd – 2, 3, 4, 5, 7: AN, AT, BL, FO, SC (GU4, IU5, JM2, JV9, ZP7).
Buellia erubescens Arnold – 1, 2, 6, 7: CB, FO, Ti, UG, stump (GU4, IU2, JM2, JV5, ZP1).
Buellia griseovirens (Turner & Borrer ex Sm.) Almb. – 1, 2, 3, 4, 5, 7: AN, AT, BL, CB, FO, PK, RP, TB,
log, snag, stump (GU14, IU8, JM5, JV8, ZP9). TLC: atranorin, norstictic acid.
RC Buellia iberica Giralt – 4, 7: AT (GU1, IU1, JM2). Thallus whitish, K+ yellow (atranorin); large
ascospores of Callispora-type, 17–23 × 8.5 µm, with microrugulate surface; hymenium without oil
droplets.
Buellia schaereri De Not. – 3, 4, 7: AN, AT, snag (GU3, JM3, JV6, ZP6).
Byssoloma leucoblepharum (Nyl.) Vain. – 1, 6: AN/ne+tw, FO/wood (GU4, IU2, JM1, JV4, ZP4).
Calicium glaucellum Ach. – 2, 3, 4, 7: PK/wood, SA/wood, snag, stump (GU4, IU3, JM2, JV6, ZP4).
Calicium lenticulare Ach. – 3, 4, 6, 7: FO, PK/wood, snag, stump (GU2, IU3, JM2, JV3, ZP1).
C Calicium parvum Tibell – 7: AN/wood (IU1).
Calicium salicinum Pers. – 1, 2, 3, 4, 6, 7: AN, AT/wood, BL, CB, log, snag, stump (GU12, IU4, JM5,
JV3, ZP4).
102 Herzogia 33 (1), 2020
Calicium trabinellum (Ach.) Ach. – 7: AN/wood, PK/wood, snag, stump (GU1, IU2, JM1, JV2, ZP2).
Calicium viride Pers. – 2, 3, 6: AN, snag (GU2, JM1, JV1, ZP1).
Caloplaca cerina (Hedw.) Th.Fr. – 1, 4, 5, 7: AN, AP, AT, BL, CA, FO, L, SA, SC, UG (GU8, IU9, JM3,
JV8, ZP6).
Caloplaca haematites
Caloplaca lucifuga G.Thor – 1, 2, 3, 6: AG, AN, AT, Ti (IU4, JM1, JV2).
Caloplaca monacensis (Leder.) Lettau – 4, 7: AT, UG (GU1, JV2, ZP2).
Caloplaca obscurella (J.Lahm ex Körb.) Th.Fr. – 7: AT (GU1).
C Caloplaca sorocarpa (Vain.) Zahlbr. – 5, 7: BL, log (JM1, JV1, ZP1).
Caloplaca stillicidiorum (Vahl) Lynge – 1, 2, 4, 7: AN/bry, AT, FO/bry, SA, SN, QU, UG/bry (GU2, IU5,
JM2, JV5, ZP1).
C Caloplaca turkuensis (Vain.) Zahlbr. – 4, 7: AT, UG (JM1, JV2, ZP4). Identifications of two sterile
specimens confirmed with mtSSU sequences.
Candelaria concolor
Candelariella efflorescens
(GU8, IU3, JM2, JV2, ZP1).
Candelariella faginea Nimis, Poelt & Puntillo – 2, 4, 5, 6, 7: AN, AT, BL, FO, UG (GU10, IU11, JM3,
JV7).
Candelariella lutella (Vain.) Räsänen – 2, 5: AN, FO, SA (GU2, JM1, ZP1).
Candelariella xanthostigma (Ach.) Lettau – 2, 3, 4, 5, 7: AN, AT, BL, FO, SA, UG (GU5, IU1, JM1,
JV6, ZP7).
Catillaria nigroclavata (Nyl.) Schuler – 1, 2, 3, 4, 6, 7: AG, AN/+ne, CA, CB, FO, QU, RP, SN, Ti, UG –
mostly tw, dead wood (GU10, IU2, JM3, JV7, ZP8).
Catinaria atropurpurea
(GU5, JM2, JV8, ZP9). The specimen JV15993 from plot 4 was identified as Catinaria aff. atropur-
purea
(maybe identical with the taxon in the note below C. atropurpurea in (2009).
Cetraria islandica (L.) Ach. – 5: BL (IU1).
Cetraria sepincola (Ehrh.) Ach. – 5: BL (GU1, IU2, JM1, JV1, ZP2).
Cetrelia cetrarioides (Delise ex Duby) W.L.Culb. & C.F.Culb. – 1, 2, 3, 5, 6, 7: AN, BL, CB, FO, RP, SC
(GU6, IU5, JM2, JV4, ZP2). TLC: atranorin, perlatolic acid, anziaic acid, 4-O-methylolivetoric acid.
Cetrelia chicitae (W.L.Culb.) W.L.Culb. & C.F.Culb. – 1, 2: CB, FO, QU, log (GU1, IU3, JV1, ZP1).
Cetrelia monachorum (Zahlbr.) W.L.Culb. & C.F.Culb. – 1, 2, 3, 4, 5, 6, 7: AN, AT, BL, CA, CO, FO,
RP, SC, Ti, UG (GU8, IU2, JM7, JV6, ZP15). TLC (A, B’, C): atranorin, imbricaric acid, perlatolic
acid, anziaic acid, 4-O-dimethylimbricaric acid.
Cetrelia olivetorum (Nyl.) W.L.Culb. & C.F.Culb. – 1, 2, 4, 5, 6, 7: AG, AN, AT, BL, CB, FO,
RP, SC, Ti, UG (GU13, IU9, JM3, JV3, ZP6). TLC: atranorin, olivetoric acid, anziaic acid,
4-O-demethylmicrophyllinic acid.
Chaenotheca brachypoda (Ach.) Tibell – 2, 3, 4: AN, snag (JM1, JV1, ZP1).
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 103
Chaenotheca brunneola (Ach.) Müll.Arg. – 1, 2, 3, 4, 7: BL/wood root, snag, stump (GU2, IU3, JM1,
JV1, ZP2).
C Chaenotheca chlorella (Ach.) Müll.Arg. – 4: wood (JM1).
Chaenotheca chrysocephala (Turner ex Ach.) Th.Fr. – 2, 3, 4, 6, 7: AN, PK/wood, snag (GU4, IU2,
JV2).
Chaenotheca ferruginea (Turner ex Sm.) Mig. – 2, 7: AN, PK/wood (GU1, JV1).
Chaenotheca furfuracea (L.) Tibell – 2, 4, 6, 7: AN, AT, BL/wood root, FO, snag (GU2, IU1, JV1).
Chaenotheca gracilenta (Ach.) Mattsson & Middelb. – 1, 4, 7: AC, AN, AT (ZP1).
Chaenotheca hispidula (Ach.) Zahlbr. – 1, 2, 4, 7: AN, AT, QU, snag (GU2, IU1, JM2, JV3, ZP2).
Chaenotheca laevigata
Chaenotheca phaeocephala (Turner) Th.Fr. – 4, 6, 7: AN, log, snag (GU1, IU2, JM2, JV3, ZP4).
Chaenotheca stemonea (Ach.) Müll.Arg. – 2, 3, 4, 5, 6, 7: AN, BL/wood root, FO, PK, QU (GU2, IU2,
JM3, JV6).
Chaenotheca trichialis (Ach.) Th.Fr. – 2, 3, 4, 6, 7: AG, AN, AT, CB, PK, snag, stump (GU5, IU4, JV3).
Chaenotheca xyloxena
ZP1).
Cheiromycina flabelliformis
JM3, JV4, ZP4). Two specimens from the collections for this project (ZP21103, 21313) were listed in
the phylogenetic study by et al. (2017).
Cheiromycina petri
Two specimens from the collections for this project (ZP 21311, 21312) were listed in the phylogenetic
study by et al. (2017).
C Cheiromycina reimeri Printzen – 2: snag (ZP1). This taxon was expected to occur in the Great Caucasus
2007). The
identification is tentative because the conidia are getting brown in part; otherwise the conidia fit well
to the description by (2007), i.e. short terminal branches, extralarge conidiogenous cell etc.
Chrysothrix caesia (Flot.) Ertz & Tehler – 1: AN/tw (ZP1).
Chrysothrix candelaris (L.) J.R.Laundon – 1, 2, 3, 4, 6, 7: AG, AN, AP, AT, CB, FO, PK, QU, Ti, snag,
stump (GU19, IU16, JV4, ZP4).
Cladonia caespiticia
Cladonia carneola (Fr.) Fr. – 5: BL (ZP1).
Cladonia cenotea (Ach.) Schaer. – 2, 3, 5, 7: BL, PK, SA, log, stump (IU2).
Cladonia chlorophaea
IU1, JM3, ZP1).
Cladonia coccifera (L.) Willd. – 7: AN (ZP1).
Cladonia coniocraea
stump (GU11, IU19).
Cladonia digitata (L.) Hoffm. – 1, 2, 3, 4, 5, 7: BL, log, snag, stump (IU1, JV1).
Cladonia fimbriata (L.) Fr. – 2, 3, 4, 5, 6, 7: AN, AT, BL, FO, RP, SC, log, snag (GU1, IU1, JM2, ZP1).
Cladonia macilenta Hoffm. – 1, 3, 4, 7: snag, stump (JV1, ZP1).
104 Herzogia 33 (1), 2020
Cladonia merochlorophaea Asahina – 3: FO (JV2).
Cladonia norvegica Tønsberg & Holien – 1: log, stump (IU1).
C Cladonia novochlorophaea
acids. Differs from C. homosekikiaca, which has the same chemistry, by esorediate podetia with a
well-developed, verrucose cortex.
Cladonia ochrochlora
Cladonia parasitica (Hoffm.) Hoffm. – 1: QU, log, stump (IU2, JV1, ZP1).
Cladonia pleurota
Cladonia pyxidata (L.) Hoffm. – 2, 4, 5, 7: AT, BL, FO, QU, SC, snag (GU2, IU3, JV2, ZP1).
Cladonia squamosa Hoffm. – 2, 5, 7: BL, PK, RP, SA (IU1).
Cliostomum corrugatum (Ach.: Fr.) Fr. – 2, 3, 4, 7: AN, AP, AT, BL, PK, QU, snag, stump (GU8, IU7,
JM3, JV5, ZP5).
Cliostomum griffithii (Sm.) Coppins – 1, 2, 3, 4, 6, 7: AN, snag (GU2, IU2, JM3, JV2, ZP3).
RC Cliostomum haematommatis
ZP5). TLC & LC-MS: atranorin and 2'-O-methylperlatolic acid (or its close derivative), sometimes with
two related compounds visible on TLC plates. The BLASTN-search did not find a very close match to
our sequences. The most similar sequences belonged to Ramalina species and Lecania baeomma; their
and chemistry correspond well to the type material of C. haematommatis except the missing pycnidia
et
al. 2006) and Switzerland ( & 2019).
Coenogonium luteum
(GU1, JM1, JV2, ZP3).
Coenogonium pineti
(GU13, IU7, JM2, JV5, ZP2).
Collema flaccidum (Ach.) Ach. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, CA, CB, FO, QU, SA, UG
(GU7, IU8, JV3, ZP8).
Collema furfuraceum (Arnold) Du Rietz – 7: AN, AT (GU1, IU2, ZP1).
Collema nigrescens (Huds.) DC. – 2, 3, 4, 5, 7: AT, BL, FO, QU, SA, SC, UG (GU4, IU5, JM3, JV5,
ZP2).
Collema subflaccidum Degel. – 1, 2, 3, 4, 6, 7: AN, AT, CA, CB, FE, FO, QU AT, FO, UG (GU7, IU5,
JM3, JV3, ZP3).
Collema subnigrescens Degel. – 3, 5, 7: AT, BL, FO, SA, SC, UG (GU4, IU5, JV1).
C Dictyocatenulata alba Finley & E.F.Morris – 3, 4, 5, 6: AN, FO (GU1, IU1, JM1, JV1, ZP3).
Elixia flexella (Ach.) Lumbsch – 4, 7: AN/snag, PK/snag (GU1, ZP2).
Enchylium tenax (Sw.) Gray – 7: UG/bry (JV1).
RC Endohyalina ericina (Nyl.) Giralt, van den Boom & Elix – 7: AN (ZP1). This is an oceanic species,
occurring locally along the Atlantic coast of Europe as far north as Ireland and of America as far south
and Italy ( et al. 2010, et al. 2014, 2016). This record is quite unexpected as the
unique hosting also other oceanic lichen species with mostly Mediterranean distribution which are
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 105
otherwise rare in the Caucasus ( &2014). The present material fits
well the description given by et al. (2010) in having a distinct grey verrucose thallus and bu-
-
ned both apically as well as at the septum and tend to be paler towards the ends. The septum is formed
lately, and the ontogeny is referable to the type B. The material was too sparse for TLC analyses for
confirmation of characteristic compounds (diploicin, fulgidin) and filiform conidia (characteristic for
this genus) were not found, however the identification was confirmed by H. Mayrhofer.
Eopyrenula leucoplaca (Wallr.) R.C.Harris – 1: CA (GU1).
Evernia divaricata (L.) Ach. – 1, 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, CB, FO, PK, QU, RP, SA, SC, TB, UG,
snag, stump (GU14, IU18, JV1, ZP1).
Evernia prunastri (L.) Ach. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, CA, CB, FO, QU, RP, log, snag
(GU14, IU5).
Fellhanera bouteillei
JM2, JV4, ZP3).
RC Fellhanera christiansenii
C Fellhanera gyrophorica Sérus., Coppins, Diederich & Scheid. – 1: FO (JV1, ZP1).
Fellhanera subtilis
Fellhanera viridisorediata Aptroot, M.Brand & Spier – 1, 6: AN/ne+tw (GU3, ZP1).
Fellhaneropsis myrtillicola (Erichsen) Sérus. & Coppins – 1, 6: AN/ne+tw (GU3, IU1, ZP1).
Flavoparmelia caperata (L.) Hale – 1, 2, 3, 4, 5, 6, 7: AN, BL, FO, CB, QU, RP, SA, Ti, UG, snag (GU5,
IU5, JV1, ZP2).
Flavoplaca flavocitrina (Nyl.) Arup, Frödén & Søchting – 2: QU (JV1).
Frutidella furfuracea (Anzi) M.Westb. & M.Svenss. – 3, 4, 5, 7: AN, BL, FO (GU1, JM3, JV2, ZP3).
Fuscidea arboricola Coppins & Tønsberg – 1, 4, 7: AN, BL, FO, PK (JM2, JV2, ZP7).
Fuscidea cyathoides
C Fuscidea recensa
1992a). This was also the case for the sterile
Betula.
Fuscopannaria mediterranea (Tav.) P.M.Jørg. – 1, 2, 7: AN, AT, FO, QU, Ti (GU2, IU3, JM1).
Gabura fasciculare (L.) P.M.Jørg. – 4, 5, 7: AT, BL, FO, SA, SC, UG (GU4, IU4, JM4, ZP3).
Graphis scripta (L.) Ach. s. lat. – 1, 2 (+ “betulina” type), 3 (+ “pulverulenta” type), 4, 6: AG, AN, AP,
AT, CA, CB, FO, QU, Ti, UG, VA (GU20, IU12, JM2, JV9, ZP7). The recent study based on both
molecular and morphological characters showed that some putative species are nested within the G.
scripta-complex, but they do not fully correspond to the taxa that were distinguished on the basis of
apothecium morphology ( et al. 2015).
Gyalecta carneola (Ach.) Hellb. – 1, 2, 3: AN, AP, AT, CB, FO, QU, RP, Ti (GU8, IU2, JM2, JV4, ZP3).
Gyalecta derivata Schuler & Zahlbr. – 3: AT, FO (IU1, JM1, JV3, ZP2). Our specimens may be identi-
fied as Gyalecta croatica (sensu 1958) distinguished from G. derivata by shorter and broader
ascospores (17–28 × 4.5 – 6 µm) containing 4 –9 cells. Our observations of some specimens from a
British herbarium (E) revealed substantial variation in spore shape and septation of G. derivata, which
included the range given for croatica.
106 Herzogia 33 (1), 2020
Gyalecta fagicola (Hepp ex Arnold) Kremp. – 2, 3, 4, 7: AN, AT, CB, FO (IU3, JM1, ZP4).
Gyalecta flotowii Körb. – 1, 2, 3: AN, AP, CB, FO, QU (GU2, IU1, JM2, JV2, ZP2).
Gyalecta herculina (Rehm) Baloch, Lumbsch & Wedin – 2, 3, 4, 5, 7: AN, AT, FO, SA, UG (GU9, IU4,
JM6, JV10, ZP12). Specimens JV15841 and JV16395 are with pycnidia. Conidia of two types: (a)
bacilliform, 5 × 1 µm and (b) long and sigmoid, c. 50 × 2 µm.
Gyalecta ophiospora
JV6, ZP6).
Gyalecta truncigena (Ach.) Hepp – 2, 3, 6: AP, CB, FO, QU (GU3, IU1, JM1, JV2, ZP6). Material from
plot 2 apparently includes two taxa. Except typical G. truncigena with ellipsoid-fusiform ascospores,
some specimens (e.g. ZP23488) produce spores approaching G. flotowii by the shape but are slightly
pointed at the apices, more richly muriform (more than 10 cells in the optical view), and the septa are
not oblique as in the latter species.
Gyalecta ulmi (Sw.) Zahlbr. – 2, 3, 6: AP, FO, QU (GU1, JM1, JV2, ZP3).
Gyalectidium setiferum
RC Gyalidea minuta-
has recently been summarized by & (2012). Our epixylic occurrence of the species
Central Europe.
Gyalideopsis helvetica
mainly on wood, log, snag (GU5, IU3, JM4, JV5, ZP8).
NC Gyalideopsis piceicola
& 2012).
Gyalolechia flavorubescens (Huds.) Søchting, Frödén & Arup – 1: CB (JV1).
Halecania viridescens Coppins & P.James – 1, 2, 3, 4, 6: AN, CA, CB, FO, RP, UG (GU2, JM1, JV5,
ZP6). In Russia and the Caucasus, this species was collected for the first time from the Republic
growing on branches and young stems of deciduous trees, often accompanied with Catillaria nig-
roclavata and other slightly nitrophytic lichens. Apothecia are produced rarely and were not obser-
Scoliciosporum sarothamni or Trapeliopsis flexuosa
red). Probably a widespread lichen in Russia.
Hazslinszkya gibberulosa (Ach.) Körb. – 1, 2, 3, 6, 7: AC, AN, AP, CB, FE, FO, QU, UG (GU1, JM3,
JV5, ZP12).
Heterodermia japonica (M.Satô) Swinscow & Krog – 1, 2, 3, 4, 6, 7: AN, AT, FO, QU (GU5, IU2, JM1,
JV4, ZP1).
Heterodermia speciosa (Wulfen) Trevis. – 1, 2, 4, 5, 7: AN, AP, AT, BL, CA, CB, FO, SA (GU3, IU7,
JM2, ZP2).
Hyperphyscia adglutinata
ZP2).
Hypocenomyce scalaris (Ach.) M.Choisy – 7: PK, snag (GU2).
Hypogymnia austerodes (Nyl.) Räsänen – 4, 5, 7: AN, AT, BL, FO, snag (GU1, IU3, JM2, JV1, ZP3).
Hypogymnia bitteri (Lynge) Ahti – 3, 4, 5: AN, AT, BL (GU1, IU1, ZP2).
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 107
Hypogymnia physodes (L.) Nyl. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, CB, FO, PK, QU, RP, SA, SC,
TB, Ti, log, snag, stump (GU17, IU13, JV1, ZP3).
Hypogymnia tubulosa (Schaer.) Hav. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, FO, PK, QU, RP, SA, SC,
TB, snag, stump (GU13, IU12, ZP1).
Hypogymnia vittata (Ach.) Parrique – 3, 4, 5, 7: AN, BL, FO, SC (GU1, IU3, JM2, JV4, ZP1).
Hypotrachyna laevigata (Sm.) Hale – 1: AN/ne+tw (GU1).
Hypotrachyna revoluta
Icmadophila ericetorum (L.) Zahlbr. – 2, 4, 7: log (GU, IU, JM, JV, ZP – only field records).
Imshaugia aleurites (Ach.) S.L.F.Mey. – 7: AN, BL, PK, sang (IU1, JM1, JV2).
Inoderma byssaceum (Weigel) Gray – 1, 2, 3, 6: AN, AT, CB, FO, QU, snag (GU4, IU5, JM2, JV6, ZP4).
RC Japewia aliphatica-
racterized by its brown, often areolate thallus composed of tiny blastidia that frequently develop into
et al.,
unpublished).
Japewia subaurifera Muhr & Tønsberg – 5, 7: PK, SC, snag (GU1, IU1, JV1, ZP1). TLC: Japewia
(1992).
Japewia tornoënsis (Nyl.) Tønsberg – 7: AN, BL, PK, snag (GU2, JM1, JV3, ZP4).
Lathagrium auriforme
Lecania croatica (Zahlbr.) Kotlov – 1, 2, 3, 4, 5, 6, 7: AG, AN, AT, CA, CB, FO, QU, RP, SN, Ti,
UG (GU7, IU12, JM5, JV10, ZP11). Lichen with a distinct green thallus around soralia, resembling
Mycobilimbia epixanthoides
ITS sequence data. The specimen ZP23048 is fertile.
Lecania cyrtella (Ach.) Th.Fr. – 1, 2, 3, 4, 5, 6, 7: AN, AT, BL, CA, FO, JC, SA, UG, log, snag (GU10,
IU5, JM4, JV8, ZP14).
Lecania cyrtellina (Nyl.) Sandst. – 1, 3, 4, 5, 7: AN, AT, FO, log, snag (GU7, IU2, JV1, ZP2).
Lecania naegelii (Hepp) Diederich & van den Boom – 1, 2, 3, 4, 6, 7: AG, AN, CA, CB, FO, LO, QU,
Ti, UG, VO, snag (GU16, IU1, JV8, ZP6).
Lecanora albella (Pers.) Ach. – 2: CB (GU1, JM1, JV1).
Lecanora albellula (Nyl.) Th.Fr. – 2, 3, 4, 5, 7: AN, PK/wood, snag, stump (GU4, IU3, JM2, ZP3).
Lecanora allophana Nyl. – 2, 4, 5, 7: AN, AP, AT, FO, SA, UG (GU3, IU5, JM2, JV4, ZP6).
C Lecanora aff. anopta
Ascospores ellipsoid, 7–10 × 3 – 4.5 µm. Conidia cylindrical, slightly curved, c. 5 – 8 × 1 µm. The spe-
cimen ZP21226 recalls pale morps (green pigment deficient) of L. anopta by its finely pruinose, see-
mingly biatorine apothecia (early receding thalline margin is however visible on section), by unevenly
inspersed guttulae (golden sclerotized ascospores) within the hymenium and by frequent pycnidia with
slightly curved conidia (see
separate species, distinct from Lecanora anopta due to its different chemistry and narrower ascospores.
Lecanora argentata (Ach.) Malme – 1, 2, 3, 4, 6, 7: AG, AN, AT, CB, FO, ST, Ti, UG (GU6, IU6, JM1,
JV3, ZP7).
RC Lecanora barkmaniana
specimen JM10602 was identified based on the match of ITS and mtSSU data following a BLASTN
108 Herzogia 33 (1), 2020
search (100 % identity and 76 % coverage for ITS; >99 % and 100 % coverage for mtSSU). Both
Caucasian specimens are sterile. This sorediate taxon was recently dealt with in more detail by
et al. (2017).
Lecanora cadubriae (A.Massal.) Hedl. – 7: AN, PK (JV1, ZP1).
Lecanora carpinea (L.) Vain. – 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, CB, FO, QU, RP, SA, TB (GU22, IU11,
JM6, JV1).
Lecanora chlarotera Nyl. – 2, 3, 4, 5, 6, 7: AN, AT, CA, CB, FO, SA (GU7, IU5, JM4, JV1, ZP2).
Lecanora cinereofusca H.Magn. – 1, 2, 6: CB, FO (JM2, JV2, ZP2).
NC Lecanora compallens
zeorin. Sterile specimen ZP22341 is tentatively identified by its chemistry and delimited soralia (so-
mewhat urceolate at the beginning). In Transcaucasia, this species has been recently reported from
Armenia ( et al. 2014).
Lecanora expallens Ach. – 1, 2, 3, 4, 5, 6, 7: AN, AG, CB, snag, stump (GU2, JM3, JV7, ZP2). Only
sterile specimens recorded, often with a low concentration of xanthones.
Lecanora exspersa Nyl. – 3, 4, 5, 7: AN, AT, BL, FO, log, snag, stump (GU8, IU3, JM9, JV11, ZP17).
TLC: atranorin, nephrosteranic acid or sometimes one additional fatty acid. Sequences from the
et al. (2017).
Lecanora glabrata (Ach.) Malme – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, CA, CB, FO, QU, SA, ST, Ti
(GU22, IU8, JM10, JV7, ZP10). Both chemotypes reported by (2014) were present in the
study area.
Lecanora hypoptoides (Nyl.) Nyl. – 7: AN, snag (GU1, JV1).
Lecanora intumescens (Rebent.) Rabenh. – 1, 2, 3, 4, 5, 7: AT, BL, CA, FO, SA (GU8, IU7, JM2, JV7,
ZP5).
Lecanora leptyrodes (Nyl.) Degel. – 2, 3, 4, 5, 7: AN, AP, AT, BL, FO, SA, SC (GU8, IU9, JM4, JV4).
Lecanora mughicola Nyl. – 7: AN/wood, PK/wood (ZP2). TLC: isousnic acid. Traces of cinereorufa-
green pigment observed, broader ascospores than in L. saligna (4 –5 µm).
C Lecanora phaeostigma (Körb.) Almb. – 4, 5, 7: AN/wood, PK/wood, log, snag (GU2, IU1, JM1, JV2,
ZP2).
Lecanora praesistens Nyl. – 4, 5: AT, FO, SC (GU4, IU1, JM2). A member of the Lecanora subfusca
group with 16-spored asci.
Lecanora pulicaris (Pers.) Ach. – 1, 2, 3, 4, 5, 6, 7: AN, AT, BL, FO, PK, SA, SC, ST, snag (GU3, IU7,
JM4, JV3, ZP5). Very unusual morphotypes with a distinctly crenulate margin, which is not typical
for this species, occured in the area. Additionally, the chemotype with fumarprotocetraric acid was
missing; only atranorin and roccellic acid were recorded by TLC. Identity of several specimens was
confirmed by the results of BLASTN searches based on ITS (>98 % identity) and mtSSU (>99 %)
sequences.
Lecanora saligna (Schrad.) Zahlbr. – 3, 4, 5, 7: AT, FO, SC, snag (GU1, IU1, JM2, JV2, ZP1).
NC Lecanora sarcopidoides (A.Massal.) A.L.Sm. – 4: AN/snag (ZP1). TLC: pseudoplacodiolic acid.
Microconidia (often gently bent) 4 –5 × 0.8 µm. In Transcaucasia, this species has been reported from
1983).
Lecanora stanislai -
rin, usnic acid. Recently described species similar to L. expallens, but without xanthones (
et al. 2017). Two Caucasian specimens (JM10367, JV14920) are paratypes. It differs
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 109
from L. compallensL. compallens). Identifications
Lecanora subcarpinea Szatala – 3, 5: AT, BL, FO (GU3, IU1).
Lecanora subintricata (Nyl.) Th.Fr. – 7: snag (IU1, JM1, JV1, ZP2).
RC Lecanora subravida Nyl. – 1, 2: snag (JM1, ZP1). TLC: usnic, isousnic, pseudoplacodiolic and
squamatic acids. Macroconidia broadly falcate, 9 –10 × 3 –3.3 µm. The taxon of the Lecanora saligna
historical localities ( & 2008).
Lecanora symmicta (Ach.) Ach. – 1, 2, 5, 6, 7: AT, BL, FO, QU, Ti, UG, snag (GU4, IU2, JM3, JV1,
ZP2).
Lecanora thysanophora R.C.Harris – 1, 2, 3, 4, 5, 6: AG, AN, AP, AT, BL, CA, CB, FO, QU, SC, Ti
Lecidea albofuscescens Nyl. – 7: AN, SA (JV1, ZP1).
C Lecidea apochroeella Nyl. – 1: wood (JM1). Hypothecium brown (K-); epihymenium granular, brow-
Lecidea coriacea Holien & Palice – 3, 7: AN/wood, log (JM1, JV1, ZP5). This old-growth forest species
related to Lecidea betulicola is easily identifiable due to the content of secalonic acid A in hypothe-
cium (KOH + golden-yellow). It was described quite recently ( et al. 2016) and some of our
specimens are paratypes.
Lecidea nylanderi (Anzi) Th.Fr. – 1, 7: FO, PK, snag (GU1, IU1, JM2, JV2, ZP2).
NC Lecidea sphaerella Hedl. – 2, 3, 4, 7: AN, AT, BL, FO, UG (JM2, JV5, ZP2). In Transcaucasia, this
species has been reported from Georgia ( 1983).
RC Lecidea strasseri Zahlbr. – 2: QU/bry (JV1). Similar to Bryobilimbia sanguineoatra, but with a
Lecidea ber-
engeriana
Lecidella achristotera
Lecidella elaeochroma (Ach.) M.Choisy s. lat. – 1, 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, CB, FO, QU, RP, SA,
SC, Ti, UG, snag (GU9, IU5, JM2, JV21, ZP22).
Lecidella euphorea
Lecidella flavosorediata
(GU2, IU2, JM6, JV5, ZP7). TLC: granulosin, arthothelin, +1 additional xanthone with lower Rf va-
lue. Identification of JM11114 confirmed by ITS and mtSSU.
Lecidella laureri (Hepp) Körb. – 2, 3, 4, 6, 7: AN, AP, FO, TB (GU7, IU2, ZP1).
Lepra albescens (Huds.) Hafellner – 1, 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, CB, FO, QU, RP, SA, Ti, UG
(GU12, IU8, JV6).
Lepra amara (Ach.) Hafellner – 1, 2, 3, 4, 5, 6, 7: AN, AT, BL, CB, FO, QU, RP, SA, SC, TB, Ti, log
(GU11, IU20, JV4, ZP2).
C Lepra borealis
fumarprotocetraric acid (in solvents A & C). Both samples correspond also morphologically with L.
borealis.
Lepra ophthalmiza (Nyl.) Hafellner – 2, 3: AN, FO, ST (JM2, JV2, ZP1). TLC: unidentified aliphatic
compounds.
110 Herzogia 33 (1), 2020
Lepra trachythallina (Erichsen) Lendemer & R.C.Harris – 1, 2, 6: CB, FO, QU, Ti (GU5, IU3, JM3,
JV6, ZP3). TLC: thamnolic acid.
Lepra waghornei (Hult.) Lendemer & R.C.Harris – 1, 2: CB, FO (GU1, JM1, ZP3). TLC: norstictic acid.
Lepraria eburnea J.R.Laundon – 2: CB/bry, QU/bry (GU2, IU1, ZP2). TLC: chemotype I – alectorialic
and protocetraric acids; chemotype II – alectorialic, barbatolic, psoromic and 2'-O-demethylpsoromic
acids.
Lepraria elobata Tønsberg – 1, 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, FO, UG, log (GU2, IU6, JM3, JV4, ZP1).
TLC: stictic, constictic acids, zeorin, atranorin.
Lepraria finkii (B.de Lesd.) R.C. Harris – 1, 2, 3, 4, 5, 6, 7: AG, AN, AT, BL, CB, FO, QU, RP, Ti, UG,
snag (GU28, IU4, JV19, ZP7). TLC: stictic, constictic acids, zeorin, atranorin.
Lepraria incana (L.) Ach. – 1, 2, 3, 4, 6, 7: AG, AN, AT, BL, FO, QU, snag (GU2, IU1, JM4, JV10,
UV+ white reaction.
Lepraria jackii
and roccellic acids.
Lepraria rigidula (B.de Lesd.) Tønsberg – 1, 2, 3, 4, 5, 6, 7: AG, AN, AT, CB, FO, PK, QU, RP, SA, snag/
bry (GU3, IU2, JM2, JV5, ZP4). TLC: atranorin, nephrosteranic acid.
Lepraria vouauxii (Hue) R.C.Harris – 1, 2, 3, 4, 6: AP, AT, CB, FO, Ti (JM2, JV3, ZP4). TLC: pannaric
acid 6-methylester.
Leptogium burnetiae C.W.Dodge – 1, 2, 3, 4, 6, 7: AN, AP, AT, CA, FE, FO, QU, SA, Ti, UG (GU6,
IU8, JM4, JV4, ZP5). MtSSU sequence of JV15904 has 99 % identity with L. burnetiae from Spain
and with L. pedicellatum from Korea. Our samples match the description of L. burnetiae: tomentum
on the lower thallus surface of cylindrical cells, isidia coralloid forming patchy clusters on the upper
surface; thallus colour blue-grey. A red-listed species in the Russian Federation ( et al. 2008).
Leptogium cyanescens (Rabenh.) Körb. – 1, 2, 6: AN, AT, CA, CB, FE, FO, QU, RP, SN, Ti, UG, log
(GU7, IU9, JM4, JV4, ZP5).
RC Leptogium hibernicum
striate with nodular isidia; lower surface uniformly finely pubescent-tomentose; tomental hairs c.
Leptogium saturninum
bry (GU1, IU9, JM5, JV8, ZP5). Specimen JV15903 was confirmed by a mtSSU sequence (similarity
99 –100 % with L. saturninum according to Blast).
Letharia vulpina (L.) Hue – 7: PK/wood, snag (GU & IU (photodocumentation), JM, JV, ZP – field
records). A red-listed species in the Russian Federation ( et al. 2008).
Lichenomphalia umbellifera (L.: Fr.) Redhead, Lutzoni, Moncalvo & Vilgalyis – 2, 3, 7: log, stump
(IU3, JV1).
RC Lithothelium hyalosporum (Nyl.) Aptroot – 1, 2, 6: AN, CB, FO (GU1, IU4, JM1, JV1, ZP3). Our
specimens have ascospores usually 20 –28 µm long. (2006) reported ascospores 14 –20(–24)
µm long. This is the first report of the genus for Russia and Caucasus.
RC Lithothelium phaeosporum (R.C.Harris) Aptroot – 1: FO (IU1). This species is closely related to L.
septemseptatum, but has only 3 instead of 5 –7 septa ( 2006).
RC Lithothelium septemseptatum (R.C.Harris) Aptroot – 1: QU (ZP1). Perithecia with reddish-brown
Mature ascospores usually 6-celled, getting red-brown, c. 38 × 13.5 µm.
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 111
Lobaria pulmonaria (L.) Hoffm. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, CA, CB, FO, QU, RP, SA,
Ti, UG, log (GU12, IU14, JV1, ZP1). A red-listed species in the Russian Federation ( et al.
2008).
Lobarina scrobiculata (Scop.) Nyl. ex Cromb. – 1, 2, 3, 4, 5, 6, 7: AN, BL, FO, QU, RP, SA, SC (GU2,
IU5, JM3, ZP3).
C Lopadium disciforme (Flot.) Kullh. – 1, 3, 4, 6, 7: AN, AT, BL, FO, QU, snag (GU2, IU1, JM3, JV1,
ZP2).
RC Loxospora cristinae
Identification of this recently described species has been confirmed by TLC – the samples contained
2'-O-methylperlatolic acid with traces of (up to four) related compounds. Several samples were also
Switzerland ( et al. 2018, et al. 2018, & 2019).
NC Loxospora elatina (Ach.) A.Massal. – 1, 2, 3, 6, 7: AG, AN, AT, FO, PK, QU (GU3, JM5, JV7, ZP4).
1983).
Marchantiana asserigena (J.Lahm) Søchting & Arup – 1, 2: UG/tw, FO/tw (IU1, JV1). Specimen
JV16394 confirmed with ITS sequence data.
Maronea constans (Nyl.) Hepp – 1, 2, 6: AN, FO, RP (GU2, JM2, JV2, ZP5).
Megalaria grossa (Pers. ex Nyl.) Hafellner – 1, 2: AN, QU (JM2, ZP1).
Megalaria laureri (Th. Fr.) Hafellner – 1, 2, 3, 6: AN, AP, CB, FO, QU, RP, ST, Ti (GU12, IU1, JM4,
JV6, ZP12).
C Megalospora porphyritis -
men JV15146 (TLC: pannarin, zeorin), matches 99 % the Australian sample Kantvilas 370/09 of
Megalospora tuberculosa). It should be noted, the authors of the phylogenetic study, where this sam-
ple was included ( & 2012), followed a broad concept of the species by
(1983, 1986) who included pannarin-containing North American M. porphyritis ( 1983) and
later, also non-sorediate Australian M. inflexa ( 1986) under M. tuberculosa, as ‘chemical strain
B’. The same authors ( & 2012) have shown that M. tuberculosa (sensu Sipman)
is non-monophyletic taxon, but left the question of the species circumscription open. M. porphyritis
has recently been reported as new to Russia as an example of “American/Asian disjunction” (
M. tuberculosa sensu (1983, 1986)
shows much larger distribution including S Brazil and Australia. More detailed studies with larger
sampling are necessary to resolve species boundaries in this complex.
RNC Melanelixia epilosa (J.Steiner) A.Crespo et al. – 3, 4, 5, 7: AN, AT, BL, FO, SA, SC, UG (GU8,
IU10, JM3, JV3, ZP2). We sequenced one specimen (ITS and mtSSU) and confirmed M. epilosa
(>99 % for ITS). The presence of M. glabra among non-sequenced specimens remains possible. In
Transcaucasia, M. epilosa has been recently reported from Armenia and Georgia ( et al. 2016).
Melanelixia glabratula (Lamy) Sandler & Arup – 1, 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, CB, FO, RP, SA,
Ti, log, snag (GU10, IU7, ZP1).
Melanelixia subargentifera (Nyl.) O.Blanco et al. – 3, 4, 6: AT, CB, FO (GU2, JM1).
Melanelixia subaurifera (Nyl.) O.Blanco et al. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AT, BL, CB, FE, FO, QU,
RP, TB, Ti, log, snag (GU30, IU10, ZP2).
Melanohalea exasperata (De Not.) O.Blanco et al. – 2, 3, 4, 5, 7: AN, AT, BL, CB, FO, SA, SC, UG
(GU12, IU6, JV4, ZP4).
Melanohalea exasperatula (Nyl.) O.Blanco et al. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, CB, FO, RP,
SA, SC, TB, snag (GU22, IU12, JV2, ZP4).
112 Herzogia 33 (1), 2020
Menegazzia subsimilis (H. Magn.) R.Sant. – 1, 2, 3, 6: AG, CB, FO (GU1, JM1, JV3, ZP1).
Menegazzia terebrata (Hoffm.) A.Massal. – 1, 2, 4, 5, 6: AG, AN, CB, BL, FO, QU (GU3, IU1, JM2).
C Micarea anterior (Nyl.) Hedl. – 7: AN/snag (ZP1).
C Micarea botryoides (Nyl.) Coppins – 1, 4, 6, 7: log, snag (JV3, ZP2).
Micarea denigrata (Fr.) Hedl. – 3, 4, 5, 7: log, snag, stump (GU3, JM5, JV5, ZP7). TLC: gyrophoric and
5-O-methylhiascic acids.
C Micarea elachista (Körb.) Coppins & R.Sant. – 7: PK/snag (JV1, ZP1).
Micarea globulosella (Nyl.) Coppins – 1, 3, 4, 5: AN, BL, FO (GU2, IU2, JM2, JV4, ZP2). TLC and spot
tests: gyrophoric acid not detected in most specimens. Possibly M. synotheoides is involved here too;
the specimen ZP21248 shows faintly C+ reddish reaction.
C Micarea hedlundii Coppins – 2, 7: log (JM1, JV1).
Micarea melaena (Nyl.) Hedl. – 1, 7: AN, PK/wood, QU/snag, stump (IU2, JM2, JV2, ZP1).
C Micarea melaeniza Hedl. – 2, 3: AN/wood, snag (IU3, ZP2).
Micarea micrococca (Körb.) Gams ex Coppins – 1, 2, 3, 4, 5, 6, 7: AG, AN/wood, CB/wood, QU/wood,
log, snag, stump (GU5, IU10, JV10, ZP6).
Micarea misella (Nyl.) Hedl. – 1, 2, 3, 4, 5, 7: log, snag, stump (GU1, IU1, JM4, JV10, ZP8).
Micarea nigella Coppins – 1, 3, 4, 7: BL/wood root, log, snag, stump (GU3, JM1, JV5, ZP2). Pycnidia
often do not contain any traces of purple (K+ green) pigment (character of M. melaeniza), but the pig-
ment is always present in the hypothecium.
RC Micarea nowakii Czarnota & Coppins – 4, 5, 7: wood (JM3). TLC: micareic acid. Identity confirmed
by mtSSU barcoding (97–99 % in BLASTN search).
Micarea peliocarpa (Anzi) Coppins & R.Sant. – 1, 2, 3, 4, 5, 6, 7: AG, AN, CB/wood, FO, log, snag,
stump (GU5, IU3, JM2, JV8, ZP4).
RC Micarea perparvula (Nyl.) Coppins & Printzen – 3: log (JM1). The voucher corresponds well to a
sample from Italy (JM6933) identified by B. Coppins.
Micarea prasina Fr. – 1, 2, 3, 4, 5, 7: AN, BL, QU, log, snag, stump (GU6, IU2, JM8, JV6, ZP7).
Micarea pusilla
from the M. micrococca group, characterized by very small (up to 0.2 mm in diam.) whitish apothecia,
usually very thin and membranaceous thallus and small ascospores (7–9 × 2–3 µm). It occurs espe-
cially on wood of coniferous trees (mainly stumps of Picea abies) in old-growth as well as managed
forests ( et al. 2019).
Micarea soralifera
snag, stump (GU7, IU6, JV12). TLC: micareic acid. This newly described species is characterized by
the thallus developing distinct, mostly delimited green soralia, the presence of micareic acid and the
Sedifolia et al. 2016). M. sorali-
fera can be confused in the field with several other sorediate Micarea species and Trapelia corticola,
which is very common and often grows in the same localities as M. soralifera. T. corticola, however,
contains gyrophoric acid (soralia and thallus C+ red). Most specimens are sterile with at least initially
C Micarea tomentosa Czarnota & Coppins – 2, 7: AN/snag (JV1, ZP1).
Multiclavula mucida (Pers.) R.H.Petersen – 6, 7: log (GU1).
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 113
Mycobilimbia carneoalbida
ZP1).
Mycobilimbia epixanthoides
1, 2, 3, 4, 5, 6, 7: AN, AT/bry, BL, CB, FO/bry, QU/bry, RP, Ti/bry, UG/bry (IU1, JM7, JV4, ZP8).
Thallus similar to well developed Lecania croatica
-
med as M. epixanthoides with ITS and mtSSU data (99 % similarities with available M. epixanthoides
Mycobilimbia tetramera
5, 7: AT/bry, UG/bry, log/bry (IU1, JM3, JV4).
C Mycoblastus alpinus (Fr.) Th.Fr. ex Hellb. – 7: AN/log, snag (JV1, ZP1). TLC: atranorin, usnic acid,
cf. planaic acid. This sorediate taxon is dealt as synonym under its fertile counterpart Mycoblastus
affinis (Schaer.) T.Schauer by et al. (2018). Here we follow the traditional concept to separate
these two distinctive morpho-ecotypes as distinct species.
Myelochroa aurulenta
and leucotylic acid, cf. secalonic acid A. Spot reaction of medulla Pd-.
Myelochroa metarevoluta (Asahina) Elix & Hale – 1: AN/ne+tw, AT, CA, FO (GU3, ZP1). TLC: atra-
norin, galbinic acid, 3 – 4 major terpenoids (zeorin, leucocytin, leucocytic acid and derivatives), cf.
salazinic acid. Spot reaction of medulla Pd+ orange.
Myriolecis persimilis
TB, UG (GU10, IU5, JM2, JV6, ZP8).
Myriolecis sambuci (Pers.) Clem. – 4, 7: AT, FO (GU1, IU1, JM2, JV2, ZP1).
Nephroma bellum
ZP3).
Nephroma helveticum Ach. – 2: FO, RP (GU1, JV1).
Nephroma parile (Ach.) Ach. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, CB, FO, QU, RP, SA, SC, ST,
UG, log (GU11, IU17, JM2, JV3, ZP5).
Nephroma resupinatum (L.) Ach. – 2, 3, 4, 5, 7: AP, AT, BL, FO, SA, SC, UG (GU2, IU6, JV3, ZP2).
C Normandina acroglypta (Norman) Aptroot – 1, 2, 3, 4, 5, 6, 7: AN, BL, CB, FO, QU, SA – often as-
crustose lichens when without perithecia. Collected Caucasian material is sterile with greenish areola-
of deciduous trees. TLC of the specimen ZP22076 revealed presence of zeorin, which was previously
discovered by Tønsberg (see Norvegian Lichen Database) and also reported by et al. (2014)
for this species. The presence of zeorin clearly separates strongly sorediate morphs of N. acroglypta
from Mycobilimbia epixanthoides, a species with a similar appearance and ecology.
Normandina pulchella (Borrer) Nyl. – 1, 2, 4, 6: AC, AG, AN, AT, BL, CA, CB, FO, QU, RP, UG – often
associated with liverworts (GU3, IU3, JM2, JV1, ZP2).
Ochrolechia alboflavescens (Wulfen) Zahlbr. – 4, 5, 7: AN, BL, PK (GU2, IU1, JM3, JV3, ZP3). TLC:
variolaric acid, lichesterinic, protolichesterinic acids. A richly fertile specimen (ZP22813) contained
atranorin and gyrophoric acid in addition.
Ochrolechia androgyna (Hoffm.) Arnold – 1, 3, 6, 7: AN, AT, BL, FO, TB (GU1, JM1, JV3, ZP3). TLC:
Ochrolechia arborea
snag (GU1, IU1, JM4, JV4, ZP4). TLC: gyrophoric and lecanoric acids, lichexanthone. UV+ orange.
114 Herzogia 33 (1), 2020
Ochrolechia bahusiensis H.Magn. – 1, 2, 3, 4, 3, 7: AN, AT, BL, CB, FO, RP, SA, snag (IU1, JM4, JV1,
ZP11). TLC: lecanoric and gyrophoric acids, murolic acid complex. UV+ white thallus.
C Ochrolechia mahluensis Räsänen – 3, 4: AT (JM2, JV1). TLC: gyrophoric and lecanoric acids.
Ochrolechia microstictoides Räsänen – 2, 4, 5, 7: AN, BL, CB/wood, PK, log, snag (GU1, IU1, JM4,
JV1, ZP3). TLC: only variolaric acid. K-, P+ yellow, C+ yellow, UV+ white.
Ochrolechia pallescens (L.) A.Massal. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AT, BL, CB, FO, RP, SA, SC, UG,
stump (GU12, IU8, JM4, JV4, ZP13). TLC: variolaric acid, murolic acid, gyrophoric acid. One speci-
men collected by JM confirmed by ITS and mtSSU.
Ochrolechia szatalaensis Verseghy – 2, 3, 4, 5, 7: AN, AT, BL, CB, FO, RP, SA, snag (GU5, IU3, JM1,
JV3, ZP3). TLC: variolaric acid.
Ochrolechia trochophora (Vain.) Oshio – 1, 6: CB, FO (JM1, JV2, ZP3). TLC: gyrophoric and lecanoric
acids. Identification of the specimen JV15442 was confirmed by mtSSU and ITS barcode.
Ochrolechia turneri (Sm.) Hasselrot – 2, 4, 7: AN, QU, snag (IU1, ZP2).
Opegrapha niveoatra (Borrer) J.R.Laundon – 1, 2, 3, 4, 6, 7: AG, AN, AT, FO, PK/wood, Ti, snag (GU8,
IU7, JM3, JV8, ZP8).
RC Opegrapha trochodes Coppins, F.Berger & Ertz – 1, 2, 3, 4, 6, 7: AN, AT, CB, FO, QU, UG (GU1,
IU3, JM3, JV3, ZP7).
Opegrapha vermicellifera (Kunze) J.R.Laundon – 1, 2, 3, 4, 6: AN, AT, CB, FO, QU, Ti (GU3, IU2,
JM3, JV5, ZP3).
RC Orcularia insperata (Nyl.) Kalb & Giralt – 2: AN/tw (ZP1). The genus Orcularia has recently been
segregated from the artificial genera Buellia and Rinodina ( & 2011) based on polarilo-
cular ascospores (OrculariaAmandinea). Orcularia in-
sperata
hemispheres. In Europe the epithet biloculata (either as Buellia or Rinodina) was used for this species,
which shows an oceanic-Mediterranean distribution ( et al. 2009). The Caucasian specimen
is rather small, with apothecia superficially resembling Catillaria nigroclavata, but containing charac-
teristic, only slightly pigmented brownish polarilocular ascospores fitting the description in &
(2011). This is the first report of the genus for Russia and Caucasus.
Pannaria conoplea (Ach.) Bory – 1, 2, 3, 4, 5, 7: AN, AP, AT, BL, CB, FO, QU, RP, SA, SC, UG (GU3,
IU3, JM4, JV4, ZP2).
Parmelia barrenoae
Parmelia ernstiae Feuerer & A.Thell – 1, 2, 3, 4, 5, 6, 7: AN, AT, FO, QU, RP, SA (GU9, IU9, JM3, ZP6).
Parmelia saxatilis (L.) Ach. – 2, 3, 4, 5, 7: AN, AT, BL, FO, PK, QU, SA, TB, log, snag (GU4, IU8).
Some specimens resembling P. ernstiae by a strongly pruinose thallus upper surface, but confirmed as
P. saxatilis by ITS sequences.
Parmelia serrana
and mtSSU.
Parmelia submontana
IU3, JM2, ZP1).
Parmelia sulcata Taylor – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, CB, FO, PK, QU, RP, SA, SC, TB, Ti,
log, snag, stump (GU41, IU26, JM1, ZP2).
Parmeliella triptophylla (Ach.) Müll.Arg. – 1, 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, CB, CA, FE, FO, QU, RP,
SA, Ti, UG (GU13, IU12, JM1, JV8, ZP7).
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 115
Parmelina carporrhizans
IU5, JV5, ZP5).
Parmelina pastillifera (Harm.) Hale – 2, 3, 4, 5, 7: AN, AT, BL, CB, FO, RP, UG (GU4, IU3, JV1, ZP1).
Parmelina quercina (Willd.) Hale – 3, 4, 5, 6, 7: AN, AT, BL, Ti (GU1, JV1, JM2).
Parmeliopsis ambigua (Wulfen) Nyl. – 3, 4, 5, 6, 7: AN, AP, AT, BL, FO, PK, log, snag, stump (GU9,
IU11, JV1).
Parmeliopsis hyperopta (Ach.) Arnold – 3, 4, 5, 7: AN, BL, FO, PK, log, snag (GU1, IU4, ZP1).
Parmotrema perlatum (Huds.) M.Choisy – 1, 2, 6: AG, AN, CB, FO, RP, SN, UG (GU5, IU4, JM4, JV1,
ZP3). TLC: stictic acid complex, atranorin.
RC Parvoplaca servitiana (Szatala) Arup, Søchting & Frödén – 3, 4: AN/tw (IU1, JV1). Specimen
JV16132 confirmed with ITS sequence data.
Parvoplaca tiroliensis (Zahlbr.) Arup, Søchting & Frödén – 7: UG/bry (GU1, JV1). Tiny yellow apothe-
Ulmus glabra (not its typical ecology). Specimen
JV15751 confirmed with ITS and mtSSU sequence data.
Peltigera collina (Ach.) Schrad. – 1, 2, 3, 4, 6, 7: AG, AN, AP, AT, CB, FO, QU, RP, SA, Ti, UG, log
(GU4, IU7, JM2, JV1, ZP1).
Peltigera degenii Gyeln. – 2, 3, 4, 5, 7: AN, FO, log, snag (GU1, IU5, JM1, JV2).
Peltigera horizontalis (Huds.) Baumg. – 1, 2, 3, 6, 7: AN, AT, FO, QU, SA, log, snag (GU1, IU2).
Peltigera neopolydactyla (Gyeln.) Gyeln. – 2, 6: log, snag (IU1, ZP1).
Peltigera polydactylon
JV3).
Peltigera praetextata
SA, UG, log, snag (GU5, IU10, JV2, ZP2).
Pertusaria coccodes (Ach.) Nyl. – 1, 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, CB, FO, ST, log, snag (GU5, IU2,
JM2, JV4, ZP7).
Pertusaria constricta Erichsen – 1, 2, 3, 4, 6, 7: AN, AP, AT, CA, CB, FO, SA (GU6, IU2, JM4, JV4,
ZP3).
Pertusaria coronata (Ach.) Th.Fr. – 2, 3, 4, 5, 6, 7: AN, AT, BL, CB, FO, PK/wood, QU, SA, Ti, snag
(GU13, IU11, JM4, JV3, ZP6).
Pertusaria flavida (DC.) J.R.Laundon – 2, 3, 4, 6, 7: AN, AT, BL, FO, Ti (GU1, IU1, JM1, JV1, ZP2).
TLC: thiophaninic and an additional xanthone.
Pertusaria leioplaca DC. – 1, 2, 3, 4, 6, 7: AN, AT, CA, CB, FO, RP, SA, UG (GU9, IU2, JM3, JV8,
ZP6). TLC: stictic acid, coronatone. A variable material that may comprise more taxa; while material
from the montane plot 7 displays distinctly convex ascocarps with 4 – 6-spored asci, the specimen
from the ‘lowland’ plot 1 (ZP23406) has shallow ascocarps and 6 – 8 spored asci recalling ‘Pertusaria
alpina’.
Pertusaria pertusa
C Pertusaria pupillaris (Nyl.) Th.Fr. – 3, 4, 5, 7: AN, BL, snag (JM1, JV5, ZP3). TLC: protocetraric acid
or fumarprotocetraric acid (ZP22074).
Pertusaria sommerfeltii
Phaeophyscia ciliata (Hoffm.) Moberg – 1: SN (GU1).
116 Herzogia 33 (1), 2020
Phaeophyscia endophoenicea (Harm.) Moberg – 1, 2, 3, 4, 5, 6, 7: AN, AP, AT, FO, UG (GU6, IU9,
JM3, JV3, ZP3).
Phaeophyscia orbicularis
Phaeophyscia pusilloides (Zahlbr.) Essl. – 1, 6: AC, AG, AN, CA, CB, FO, SN, Ti, UG (GU12, IU9,
JM2, JV4, ZP5).
Phaeophyscia rubropulchra (Degel.) Essl. – 1, 2, 3, 6: AG, AN, AT, CB, FO (GU2, IU1, JM2, JV2,
ZP3).
Phlyctis agelaea (Ach.) Flot. – 1, 2, 3, 6: AN, AT, CA, CB, FO, QU, RP, Ti (GU3, IU4, JM2, JV4, ZP2).
Phlyctis argena (Spreng.) Flot. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, CB, FO, QU, TB, Ti, UG, snag
(GU17, IU8, JM1, JV2, ZP1).
RC Phylloblastia inexpectata
correspond to the description of this species by et al. (2007). This is the first report of the
genus for Russia and Caucasus.
Phyllogyalidea phyllophila
Physcia adscendens (Fr.) H.Olivier – 1, 3, 4, 6, 7: AG, AN, AP, AT, FO (GU9, IU9, JM1, JV2, ZP1).
Physcia aipolia (Ehrh. ex Humb.) Fürnr. – 1, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, CA, CB, FO, SA, SC,
Ti, UG (GU6, IU9, JM2, JV2, ZP6).
Physcia cf. biziana (A.Massal.) Zahlbr. – 5: BL (JM1). A single small thallus fits well with a description
of this species (e.g. & 2012), but according to BLASTN, our sample of P.
biziana
GU247178, GU247212).
Physcia stellaris (L.) Nyl. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, FO, SA, SC, UG (GU13, IU5, JV4,
ZP4).
Physcia tenella (Scop.) DC. – 1, 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, CB, FO, RP, SA, TB, Ti, UG (GU14,
IU8, JV1, ZP1).
Physcia tribacia (Ach.) Nyl. – 4: AT (IU1, JM1).
Physciella chloantha (Ach.) Essl. – 1, 6: AC, AG, AN, CA, CB, FO, SN, UG (GU3, IU2, ZP2).
Physconia detersa (Nyl.) Poelt – 3, 4, 6: AT, FO (JV2, ZP3).
Physconia distorta (With.) J.R.Laundon – 1, 2, 3, 4, 5, 6, 7: AC, AN, AP, AT, BL, CA, CB, FE, FO, SA,
SC, Ti, UG (GU18, IU25, JM2, JV3, ZP3).
Physconia perisidiosa (Erichsen) Moberg – 3, 4, 5, 7: AN, AP, AT, FO, UG (GU2, IU6, JM3, JV1, ZP1).
NC Piccolia ochrophora (Nyl.) Hafellner – 1, 2, 4: AT, QU, SN (JM1, JV2). Specimen JV14926 repre-
sents an anamorph with distinct white pycnidia and sphaerical conidia. In Transcaucasia, this species
has been recently reported from Armenia ( et al. 2015).
Placynthiella dasaea (Stirt.) Tønsberg – 2, 3, 4, 5, 7: AN/wood, FO/wood, log, snag, stump (GU5, IU3,
JM2, JV2, ZP5).
Placynthiella icmalea (Ach.) Coppins & P.James – 1, 2, 3, 4, 5, 7: AN, BL, CB/wood, FO, PK/wood, log,
snag, stump (GU9, IU10, JV3, ZP4).
Platismatia glauca (L.) W.L.Culb. & C.F.Culb. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AT, BL, FO, PK, SA, TB,
snag (GU5, IU5, JV1).
Polycauliona candelaria (L.) Frödén, Arup & Søchting – 7: AN (ZP1).
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 117
Polycauliona polycarpa (Hoffm.) Frödén, Arup & Søchting – 7: UG (JV1).
Porina aenea (Wallr.) Zahlbr. – 1, 2, 3, 4, 6, 7: AN, AP, AT, CB, FO, QU, RP, SA, log (GU16, IU3, JM3,
JV8, ZP6).
Porina borreri
Porina byssophila (Körb. ex Hepp) Zahlbr. – 2, 3: CB, FO (ZP2). Similar to P. aenea but with a diffe-
rent pigmentation, giving a KOH+ greyish blue reaction. Identity of the specimen ZP24020 confir-
med by A. Orange. Previously it was recorded only once from the Caucasus as a saxicolous species
( & 2004). Recently it has been shown it is more frequent species than
previously believed and often growing as an epiphyte as well (2013).
Porina leptalea (Durieu & Mont.) A.L.Sm. – 1, 2, 3, 6: AN, AT, FO, QU, SN (IU1, JM2, JV2, ZP4).
Porina oxneri R.Sant. – 1, 6: AN/ne+tw (GU4, IU2, JM1, JV3, ZP1).
Porina pseudohibernica Tretiach – 2, 3, 4, 6, 7: AN, AT, FO, QU, Ti (JM3, JV3, ZP5). Only recently
recorded for the first time from the Caucasus and Russia ( et al. 2017).
Porina rosei Sérus. – 1: CB (JV1). Sterile, isidiate specimen; thalllus green-grey; isidia long, ±branched,
fragile, with distinct papillae of outer mycobiont cells, but isidia not monilliform; cortex of isidia
rather thin, indistinct.
Protoparmelia oleagina (Harm.) Coppins – 7: wood (JM1).
Pseudevernia furfuracea (L.) Zopf – 2, 3, 4, 5, 7: AN, AT, BL, FE, FO, PK, RP, SA, SC, TB, snag (GU4,
IU9, ZP2).
Pseudoschismatomma rufescens (Pers.) Ertz & Tehler – 3: AP, AT (IU1, JM1, JV2, ZP1).
C Psoroglaena abscondita
JV2, ZP2).
Psoroglaena dictyospora
log, snag (GU3, IU2, JM1, JV9, ZP10).
RC Psoroglaena stigonemoides (Orange) Henssen – 2: Ti (ZP1). A sterile specimen with distinctly papil-
late filaments under the microscope; branchlets 8 –10 µm broad with tiny photobiont cells, seemingly
arranged in two rows.
RC Ptychographa xylographoides Nyl. – 4: snag (IU1). Apothecia persistently narrow and elongate; ex-
ascospores simple, 8.5 –13 × 4.5 – 6.5 µm. This is the first report of the genus for Russia and Caucasus.
Punctelia borreri (Sm.) Krog – 5, 6: BL, Ti (GU1, ZP1).
Punctelia jeckeri (Roum.) Kalb – 1: UG (IU1).
C Puttea exsequens (Nyl.) Printzen & Davydov – 1, 2, 6: FO, QU – wood, log, snag (GU2, IU1, JM2,
JV4, ZP4).
Pycnora praestabilis (Nyl.) Hafellner – 7: PK/wood, snag (GU2, JV1, ZP2).
C Pycnora xanthococca (Sommerf.) Hafellner – 7: PK/wood (JV1).
Pyrenula chlorospila (Nyl.) Arnold – 1, 2: AP, CA (GU2).
Pyrenula coryli A.Massal. – 1: FO (JV1).
Pyrenula laevigata (Pers.) Arnold – 1, 2, 6: AN, AP, CA, CB, FO (GU2, IU3, JV3, ZP2).
Pyrenula nitida (Weigel) Ach. – 1, 2, 3, 4, 5, 6: AG, AN, AP, AT, CA, CB, FO, QU, ST, Ti, UG (GU15,
IU6, JM2, JV9, ZP5).
118 Herzogia 33 (1), 2020
Pyrenula nitidella
IU1, JM2, JV11).
Ramalina calicaris (L.) Fr. – 2, 3, 4, 5, 7: AN, AP, AT, BL, CB, FO, RP, SA, SC (GU4, IU14, JM3, JV3,
secondary metabolites (ZP22953).
Ramalina europaea
Ramalina farinacea (L.) Ach. – 1, 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, CA, CB, FO, QU, RP, SA, Ti, snag,
stump (GU19, IU15, ZP7). TLC: usnic acid (present or absent), protocetraric acid (always present),
norstictic acid with a satellite (connorstictic acid) (present or absent); two chemotypes revealed: I –
with norstictic acid (ZP21641, 22696) and II – without norstictic acid (ZP22932, 23201, 23412).
Ramalina fraxinea (L.) Ach. – 3, 4, 5, 7: AN, AT, BL, FO, SA, SC (GU1, IU1, JM1, ZP1).
Ramalina obtusata (Arnold) Bitter – 1, 2, 3, 4, 5, 6, 7: AG, AN, AP, AT, BL, CB, FO, PK, QU, RP, TB,
Ti, log, snag (GU16, IU15, JM3, JV4, ZP8).
Ramalina panizzei De Not. – 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, CB, FO, SA, SC (GU1, IU2, JM2,
ZP22766 was confirmed by ITS barcode.
Ramalina pollinaria (Westr.) Ach. s. lat. – 1, 2, 3, 4, 6, 7: AG, AN, AT, CA, CB, FO, QU, Ti (GU4,
IU3, JM3, JV1). Three specimens collected by JM represent R. pollinaria s. str., other specimens may
represent R. europaea.
Ramalina roesleri (Hochst. ex Schaer.) Hue – 1: dead tw of Prunus
Ramalina sinensis Jatta – 2, 3, 4, 5, 7: AN, AT, BL, CB, FO, SC (GU3, IU7, JV1, ZP1).
Ramalina thrausta (Ach.) Nyl. – 1, 2, 3, 6: AN, FO, RP, Ti (GU1, IU1, JV3, ZP2).
Ramboldia elabens (Fr.) Kantvilas & Elix – 7: PK/wood, snag (JM1, ZP1).
Ramonia chrysophaea
RC Ramonia dictyospora Coppins – 1: SN (JV1).
RC Ramonia luteola
Reichlingia leopoldii Diederich & Scheid. – 3: AT (JM1). TLC: 2'-O-methylperlatolic acid (major), con-
fluentic acid (minor). Based on molecular data it is recognized as a member of Arthoniomycetes, and
was emended to include also Arthonia zwackhii, a chemically concordant fertile species ( et al.
2014). In Russia and the Caucasus, this species was collected for the first time from the Republic of
Adygea and from Krasnodar Territory (specimen in GLM; V. Otte in litt.).
Ricasolia amplissima (Scop.) De Not. – 1, 2, 3, 4, 6, 7: AN, AP, AT, BL, CB, FE, FO, QU, SA, UG (GU2,
IU2, JV1, ZP4). A red-listed species in the Russian Federation ( et al. 2008). The fruticose
growth form of Ricasolia amplissima (“Dendriscocaulon
as photobionts) has a rather different ecology and distribution compared to its bi- or tripartite foliose
photomorph (always with chlorophyte photobiont, sometimes with additional cyanobacteria in cepha-
lodia). In the Caucasus, it often grows independent of the foliose R. amplissima
it can inhabit other lichen species (e.g., Cetrelia sp., Parmelina sp. and other Parmeliaceae). We are
listing the records of the “Dendriscocaulon
form: 3, 4, 7: AN, BL, FO, SA (GU2, IU4, JM2).
Rinodina albana (A.Massal.) A.Massal. – 1, 2, 3, 4, 5, 6, 7: AC, AN, AT, BL, CA, CB, FO, SA, UG, snag
(GU3, JM6, JV7, ZP8).
C Rinodina buckii Sheard – 1: FO (ZP2). TLC: zeorin, pannarin. Fertile material (ZP22823) with
teichophila-type ascospores; verified by H. Mayrhofer. The species was recently described as E North
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 119
American / E Asian taxon by
region. Presence of the species in the Caucasus represents a large range expansion.
C Rinodina capensis Hampe – 1, 2, 3, 4, 5, 7: AN, AT, BL, FO, RP, log, snag (GU5, IU4, JM5, JV4, ZP7).
C Rinodina efflorescens Malme – 1, 2, 3, 4, 5, 7: AN, AT, BL, CB, FO, Ti, snag (GU3, IU1, JM3, JV6,
ZP3). TLC: pannarin, secalonic acid A.
C Rinodina excrescens Vain. – 1, 7: AN, FO (GU1, IU1, JM1, JV1). TLC: pannarin (traces).
Rinodina exigua (Ach.) Gray – 1, 2, 4: AG, AN, FO (GU1, IU1, JM1, ZP3).
C Rinodina freyi H.Magn. – 1, 2, 6: AN, CA, FO (GU2, JV2, ZP2).
Rinodina griseosoralifera Coppins – 4, 7: AT, BL, UG (GU1, IU1, JV2).
C Rinodina malangica (Norman) Arnold – 5: AT, SA (JV1, ZP2). The most similar mtSSU sequences
(94 –95 % in BLASTN) of the sterile specimen collected by JV belong to Rinodina species. Specimen
ZP21628 bears few apothecia.
Rinodina orculata Poelt & M.Steiner – 1, 2, 3, 4, 5, 7: AN, AT, BL, CB, FO, JC, PK/wood, SA, SC, TB,
snag (GU7, IU6, JM5, JV14, ZP21).
RC Rinodina polysporoides Giralt & H.Mayrhofer – 1, 2, 6: AP, FO, Ti (GU4, JM2). The sample
JM10654 identified by H. Mayrhofer.
Rinodina pyrina (Ach.) Arnold – 3, 4, 5: AN, CB (GU2, ZP1).
C Rinodina sheardii Tønsberg – 3, 4, 5, 7: AN, AT, BL, FO (JM1, JV2, ZP4). TLC: Secalonic acid A, ze-
orin, unidentified UV+ blue substance (both prior and after charring) at level of atranorin, not forming
a visible spot. Caucasian material is sterile, forming delimited, brightly yellowish-greenish soralia on a
see 1992b).
Rinodina sophodes (Ach.) A.Massal. – 2, 3, 4, 5, 7: AP, AT, BL, CB, FO, SA, UG (GU4, IU2, JM4, JV9,
ZP7).
Rinodina subpariata (Nyl.) Zahlbr. – 1, 2, 3, 4, 5, 6, 7: AG, AN, AT, BL, CB, FO, RP, SA, Ti, stump
(GU9, IU3, JM6, JV10, ZP7).
C Rinodina tenuis Müll. Arg. – 1: FO (JM1). TLC: pannarin, zeorin. Identified by H. Mayrhofer.
NC Rinodina trevisanii (Hepp) Körb. – 1, 4, 5, 7: AN, AT, BL, CA, FO, SC, log, snag (GU2, IU2, JM4,
JV3, ZP4). In Transcaucasia, this species has been reported from Georgia ( &
2007).
C Rinodina willeyi Sheard & Giralt – 1, 6: AG, FO (JV2, JM3, ZP1). TLC: pannarin, zeorin, traces of
terpenoids. JM specimens identified by H. Mayrhofer. Caucasian material is sterile but corresponds
well morphologically and chemically to description of the species ( et al. 2012). It was recently
et al. 2017).
C Ropalospora viridis (Tønsberg) Tønsberg – 1, 2, 6: CB, FO (JM2, JV1, ZP2).
Rostania occultata
ZP1).
C Sagedia aff. mastrucata
out as Aspicilia grisea sensu British authors ( et al. 2010). Sagedia mastrucata is the closest
-
blished data).
120 Herzogia 33 (1), 2020
C Sarcosagium campestre (Fr.) Poetsch & Schied. – 4, 6: AN/wood, snag (JM1, ZP2).
C Schaereria corticola
(ZP21398, 23342): gyrophoric and 5-O-methylhiascic acid (trace). Persistently delimited brown sora-
lia. MtSSU sequence (JV16547) has >97 % identity with S. corticola (sample Tønsberg 28432).
Schismatomma pericleum (Ach.) Branth & Rostr. – 1, 2, 3, 4, 6, 7: AN, FO, TB, Ti (GU3, IU5, JM3,
JV4, ZP1).
NC Sclerophora amabilis (Tibell) Tibell – 2, 3: FO (JV1, ZP1). In Transcaucasia, this species has been
1998).
Sclerophora farinacea Chevall. – 2, 3, 6: AP, FO (JM1, JV1, ZP1).
Sclerophora pallida (Pers.) Y.J.Yao & Spooner – 1, 2, 3, 4: FO, QU (GU2, JM2, ZP1).
NC Sclerophora peronella (Ach.) Tibell – 4: AT (GU1). In Transcaucasia, this species has been reported
1998).
Scoliciosporum chlorococcum
(IU1, JV1, ZP3).
C Scoliciosporum sarothamni-
mens may represent the very similar S. gallurae.
Scoliciosporum schadeanum
Scoliciosporum umbrinum (Ach.) Arnold – 1, 2, 3, 4, 5, 6, 7: AN, AT, BL, CB, FO, RP, SA, SC, TB,
UG – mainly tw, snag (GU11, IU5, JM3, JV12, ZP11).
Scutula circumspecta
CB, FO, QU, UG (GU1, IU1, JM5, JV5, ZP11).
Scytinium gelatinosum
Scytinium lichenoides
Ti, UG/bry, log (GU7, IU9, JV2, ZP2).
Scytinium pulvinatum
RP, ST, Ti/bry, UG (GU2, IU3, JM1, JV1, ZP2).
Scytinium subtile
Scytinium teretiusculum
UG, snag (GU4, IU5, JM4, JV7, ZP7).
C Steinia geophana (Nyl.) Stein – 2, 3, 4, 5: log, snag (JV3, ZP1).
C Strangospora microhaema (Norman) R.A.Anderson – 1: AC/tw (ZP1).
C Strangospora moriformis (Ach.) Stein – 7: PK (ZP1).
Strigula jamesii (Swinscow) R.C.Harris – 2: CB (ZP1).
Strigula stigmatella (Ach.) R.C.Harris – 2, 3, 4, 6, 7: AT, CB, FO, QU, SA, Ti – incl. bryophytes (GU5,
IU2, JM4, JV12, ZP12).
Tephromela atra (Huds.) Hafellner – 2, 3, 4, 5, 7: AN, AP, AT, CB, FO, SA (GU4, IU8, JV5, ZP7).
Tetramelas chloroleucus (Körb.) A.Nordin – 5, 7: AT, BL, SA (ZP5).
Tetramelas triphragmioides (Anzi) A.Nordin & Tibell – 3, 5: AN, AT, BL (GU1, IU2, JV2, ZP1).
Thelenella muscorum (Fr.) Vain. – 2, 3, 4: AT/bry, CB/bry, FO/bry (IU1, JM1, JV2, ZP1).
C Thelenella pertusariella (Nyl.) Vain. – 3, 4, 5, 6, 7: AG, FO, SC (GU1, JV3, ZP6).
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 121
Thelocarpon epibolum Nyl. – 3: AN, FO (also on Peltigera sp.), log (JV2).
C Thelocarpon intermediellum Nyl. – 4: snag (IU1, JM1).
C Thelocarpon lichenicola
RNC Thelopsis flaveola Arnold – 2, 4, 5: AP, AT, FO, Ti (GU2, IU2, JM2, JV4, ZP5). In Transcaucasia,
& 2012).
Thelopsis rubella Nyl. – 2, 3, 4, 6: AN, CB, FO, QU, Ti (IU1, JV2, ZP7).
Thelotrema lepadinum (Ach.) Ach. – 1, 2, 6: AN, CB, FO, RP, Ti (GU1, IU2, JM3, JV2, ZP4).
C Toensbergia leucococca
Toniniopsis subincompta
BL, CB, FO, SA, SN, UG, log, snag (GU11, IU6, JM3, JV18, ZP12).
RC Topelia jasonhurii
to the description of this species in et al. (2013). In the field, Topelia jasonhurii can
resemble some Thelopsis species, but it differs for example by muriform ascospores. This taxon has
Trapelia corticola Coppins & P.James – 1, 2, 3, 4, 5, 6, 7: AG, AN/wood, QU, log, snag, stump (GU5,
IU3, JM4, JV14, ZP3).
Trapeliopsis flexuosa (Fr.) Coppins & P.James – 1, 2, 3, 4, 5, 7: BL, PK, log, snag, stump (GU1, IU2,
JM1, ZP2).
C Trapeliopsis gelatinosa
C Trapeliopsis glaucolepidea (Nyl.) Gotth. Schneid. – 3, 7: snag (JM1, JV2).
Trapeliopsis granulosa (Hoffm.) Lumbsch – 3, 7: AN, PK, snag (JV2).
Trapeliopsis pseudogranulosa Coppins & P.James – 1, 2, 3, 5: BL, snag (JM2, JV1, ZP2).
Trapeliopsis viridescens (Schrad.) Coppins & P.James – 1, 2, 3, 4, 7: FO, log, snag, stump (GU1, JV).
Tuckermannopsis chlorophylla (Willd.) Hale – 2, 3, 4, 5, 7: AN, AT, BL, CB, FO, PK, QU, TB, snag
(GU3, IU10, JV1, ZP1).
Usnea articulata (L.) Hoffm. – 1, 2, 3, 4, 5, 7: AN, BL, CB, FO, PK, QU, RP, UG (GU1, IU2, JM1, JV4,
ZP2).
Usnea barbata (L.) F.H.Wigg. – 2, 3, 4, 5, 6, 7: AN, AT, BL, FO, RP (GU4, IU4, JM6). Selected speci-
mens verified by P. Clerc.
Usnea cavernosa
and 5 formed a compacted medulla.
Usnea dasopoga (Ach.) Nyl. – 1, 2, 3, 4, 5, 6, 7: AN, BL, CB, FO, PK, RP, snag (GU5, IU4, JM3, JV6,
ZP1).
Usnea flavocardia Räsänen – 1: AN (JM1).
Usnea florida (L.) F.H.Wigg. – 1, 2, 3, 4, 5, 6, 7: AN, AT, BL, CB, FO, QU, RP, SA, SC, UG, log, snag
(GU13, IU7, JV2, ZP3). A red-listed species in the Russian Federation ( et al. 2008).
Usnea glabrescens (Nyl. ex Vain.) Vain. var. glabrescens – 1, 2, 3, 4, 6, 7: AN, CB, FO, RP, snag (GU9,
IU3).
var. fulvoreagens Räsänen – 1, 2, 4, 6: AN, CB, FO, QU, Ti (GU6, IU2, JM2, ZP3).
Usnea hirta (L.) F.H.Wigg. – 5, 7: AN, BL, PK, snag (GU1, JV1, ZP1).
122 Herzogia 33 (1), 2020
Usnea intermedia (A.Massal.) Jatta – 2, 3, 4, 5, 6, 7: AN, AP, AT, BL, CB, FO, SA, SC, snag (GU6, IU12,
JM4, JV3, ZP6).
Usnea longissima Ach. – 1, 2, 3, 6: AN, CA, CB, FO, QU, RP (GU10, IU5, JM, JV, ZP2).
Usnea perplexans Stirt. – 1, 3, 4, 5, 7: AN, AT, BL, FO, SA, SC, snag (GU6, IU7, JM5, JV2, ZP3).
Specimens collected by JM verified by P. Clerc.
Usnea subfloridana Stirt. – 1, 2, 3, 4, 5, 6, 7: AN, AT, BL, CB, FO, QU, PK, RP, QU (GU3, IU3, JM3,
JV5, ZP4).
Usnea substerilis
Usnea wasmuthii Räsänen – 2, 6: FO (JM3). Specimens collected by JM analyzed by TLC and verified
by P. Clerc.
Usnocetraria oakesiana
Vahliella saubinetii (Mont.) P.M.Jørg. – 1, 2, 7: AN, AT, FO, QU, Ti (GU2, IU3, JM2).
Varicellaria hemisphaerica
QU, RP, SA, Ti, snag (GU6, IU10, JM4, JV5, ZP3). TLC: gyrophoric and lecanoric acids (+ an un-
Verrucaria breussii Diederich & van den Boom – 1, 2, 6: AG, FO (JV4, ZP1). All specimens have no
involucrellum and the exciple is colourless in the lower part. As for the spore size the material is not
homogenous and more taxa may be included. Specimen JV15453 contains usually smaller ascospores
(c. 16 –20 µm long), but at least some larger ascospores (20 –25 µm long) were also present. Specimen
ZP22986 fits well this taxon with ascospores c. 23 × 10.5 µm.
RC Verrucaria hegetschweileri Körb. ex Nyl. (nom. illeg.) non (Naegeli ex Hepp) Garov. – 2: QU (ZP1).
wall reaching to the base; ascospores ellipsoid to ovoid (subpyriform) 12–16.5 × 6 –7 µm.
Verrucaria cf. lignicola
the base. The voucher ZP23166 was identified by O. Breuss. It is similar to Verrucaria bryoctona with
-
V. bryoctona the involucrellum is developed, but hardly discernible from the perithecial
wall, and the thallus is more or less areolate-squamulose but not granular.
Vezdaea aestivalis (Ohlert) Tscherm.-Woess & Poelt – 1, 2: Ti, log (ZP2).
C Vezdaea retigera Poelt & Döbbeler – 1, 6: AG, FO, log (JV1, ZP2). Sterile specimens identified on
basis of characteristic goniocysts with short spines.
C Vezdaea rheocarpa Poelt & Döbbeler – 2, 7: AN, QU (JM1, ZP1). Sterile specimen ZP22936 with
C Violella fucata (Stirt.) T.Sprib. – 2, 5, 6, 7: AN, BL, CB, FO, PK/wood (JM3, ZP3). TLC: atranorin,
fumarprotocetraric acid.
Vulpicida pinastri (Scop.) J.-E. Mattsson & M.J. Lai – 2, 5, 7: AN, BL, FO, RP, SC, stump (GU2, IU4,
JM, JV1, ZP1).
RC Wadeana minuta Coppins & P.James – 4: FO (JV1). This is the first report of the genus for Russia
and Caucasus.
RC Waynea giraltiae van den Boom – 7: UG (GU1, ZP1). Characteristic species when fertile. Hairy
squamules are somewhat similar to those of unrelated Agonimia opuntiella when young, but distinctly
larger with age. In the specimen ZP21598 only one apothecium is present, habitually very similar to
2010, et al. 2018, et al. 2018).
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 123
Xanthomendoza ulophyllodes (Räsänen) Søchting, Kärnefelt & S.Y.Kondr. – 4, 7: AT (JM, JV2).
Specimen JV16088 was confirmed by ITS barcode (98 –100 % identity by BLASTN). Other speci-
mens with similar morphology are suggested to belong here too.
Xanthoria parietina (L.) Th.Fr. – 2, 3, 6: AN, UG (GU1, JV1, ZP1).
Xylographa parallela (Ach.) Fr. – 2, 3, 4, 5, 7: CB/wood, PK/wood, log, snag, stump (GU3, IU2, JV3,
ZP1). TLC (ZP22099): stictic acid (major), norstictic acid (minor). Specimens tentatively identified as
Xylographa pallens, following the monograph by et al. (2014), were revised as X. parallela
on the basis of mtSSU and ITS barcode.
C Xylographa soralifera Holien & Tønsberg – 3, 4, 7: log, snag, stump (GU3, JV2, ZP1).
C Xylographa trunciseda
ZP4).
Xylographa vitiligo (Ach.) J.R.Laundon – 2, 3, 4, 7: AN/wood, PK/wood, log, snag, stump (GU2, IU4,
JM4, JV4, ZP1).
Zwackhia viridis (Ach.) Poetsch & Schied. – 1, 2, 4, 6: AN, CB, FO, Ti (GU5, IU2, JM2, JV7, ZP3).
Lichenicolous fungi
Abrothallus bertianus De Not. – 2, 3, 4, 5: on Melanelixia glabratula, M. subaurifera, Melanohalea
exasperata, AT, BL, FO (GU2, IU2).
C Abrothallus nephromatis Suija & Pérez-Ortega – 5: on Nephroma parile, SC (GU1, IU1).
Abrothallus parmeliarum (Sommerf.) Arnold – 5: on Parmelia ernstiae, P. sulcata, BL, FO, SC (GU2,
IU2, JM1, ZP1).
Abrothallus peyritschii (Stein) Kotte – 5: on Vulpicida pinastri, BL (IU1, JM1, ZP1).
C Abrothallus suecicus (Kirschst.) Nordin – 5: on Ramalina panizzei, AT (GU1).
Acolium sessile (Pers.) Arnold – 3: on Pertusaria coccodes, AN (JV1).
C Arthonia biatoricola Ihlen & Owe-Larss. – 3, 4, 5, 7: on Biatora efflorescens, AN, AT, PK/wood, snag,
stump (GU3, IU3, JV3, ZP1).
Arthonia epiphyscia Nyl. – 1, 5: on Physcia stellaris, Physciella chloantha, BL, SN (GU1, IU1).
Arthonia phaeophysciae Grube & Matzer – 1: on Phaeophyscia pusilloides, AN (GU1).
Arthonia subfuscicola (Linds.) Triebel – 4, 7: on Lecanora carpinea, AN, FO, SA (GU2, IU1).
RC Arthonia vorsoeensis Alstrup – 3: on Amandinea punctata, AT (GU1). Hymenium 40 –50 µm high,
gel I+ blue; asci arthonioid, clavate, apically I-, 8-spored; ascospores l-septate, 9 –11 × 3.5 – 4 µm, long
remaining colourless ( et al. 2004).
Arthophacopsis parmeliarum Hafellner – 5, 7: on Parmelia sulcata, SA, SC, TB (GU2, IU2).
C Arthrorhaphis aeruginosa R.Sant. & Tønsberg – 7: on Cladonia sp. (JM1).
Bachmanniomyces punctum (A.Massal.) Diederich & Pino-Bodas – 7: on Cladonia coniocraea, PK
(IU1).
Biatoropsis usnearum Räsänen s. str. – 2: on Usnea sp. (JM1).
Chaenothecopsis brevipes Tibell – 2, 6: on Inoderma byssaceum, AN, QU (JV2).
NC Chaenothecopsis vainioanaArthonia sp., PK/wood, snag (GU1, IU1). In
2006).
124 Herzogia 33 (1), 2020
Dactylospora deminuta (Th.Fr.) Triebel – 2: on Mycobilimbia epixanthoides, QU (ZP1).
Dactylospora lobariella (Nyl.) Hafellner – 2, 4: on Lobaria pulmonaria, AT, CB (GU2, JM1).
Dactylospora parasiticaLepra albescens (JM1).
C Didymocyrtis cf. cladoniicola Lecanora
carpinea, SA (IU1).
RC Didymocyrtis melanelixiaeParmelia sulcata, BL (IU1).
RC Epigloea urosperma Döbbeler – 2: on Placynthiella icmalea, log (IU1, JV1). Asci 34 µm long, with
32 ascospores; ascospores l-septate, 8.4 × 2 µm with tail 4 µm.
Heterocephalacria bachmannii (Diederich & M.S.Christ.) Millanes & Wedin – 1: on Cladonia conio-
craea, log, stump (GU2).
Homostegia piggotiiParmelia sulcata, BL, Ti (GU1, ZP1).
Intralichen christiansenii Lecanora intumescens, FO
(GU1).
Kalchbrenneriella cyanescens (Kalchbr.) Diederich & M.S.Christ. – 7: on Usnea subfloridana, AN
(IU1). In Russia and Caucasus, this species was collected for the first time from the Republic of
Adygea (specimen in GLM; V. Otte in litt.).
Lichenoconium lecanorae Hypogymnia sp., Lecanora carpinea,
Parmeliopsis ambigua, AT, SA, TB (GU1, IU2, JM1).
Lichenoconium usneaeMelanohalea exasperata, Physconia perisidiosa,
Ramalina panizzei, AN, AT (GU3).
Lichenopeltella peltigericolaPeltigera praetextata, BL (IU1).
Lichenostigma alpinumCladonia coniocraea,
stump (IU1).
C Monodictys cellulosa S.Hughes – 7: on Arthonia vinosa, snag (GU1).
C Monodictys epileprariaLepraria rigidula, RP (GU1).
Muellerella lichenicolaCaloplaca cerina, FO (IU1).
RC Muellerella polyspora Hepp ex Müll.Arg. – 2, 4: on Arthonia radiata, FO (GU2). Asci multispored;
ascospores 5 –7 × 2.5 –3 µm; perithecia becoming ± superficial.
Myxophora leptogiophilaCollema flaccidum, QU
(ZP1).
Opegrapha rotunda Hafellner – 4: on Physconia distorta, AT (IU1).
Perigrapha superveniens (Nyl.) Hafellner – 4, 5: on Parmelia sulcata, AN, BL (JV1, ZP1).
RNC Phacographa zwackhii Phlyctis argena, AN, AT
(IU1, JM2, ZP2). In Transcaucasia, this species has been recently reported from Armenia (
et al. 2015).
Plectocarpon lichenum Lobaria pulmonaria, CB, FO, QU (GU1,
IU2, JM1).
Plectocarpon scrobiculatae Diederich & Etayo – 5: on Lobarina scrobiculata, BL (IU1).
RC Pronectria pilosa Etayo & López de Silanes – 5: on Collema subnigrescens, SC (IU1). Originally
decribed from Collema furfuraceum. Ascomata yellowish, 50 – 80 µm in diam., with normally setose
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 125
papilla around ostiole; ascospores ellipsoid to almost fusiform, hyaline, 1-septate, slightly constricted
in the septum, not ornamented, 10 –13.5(–15) × 3.3 –3.5 µm (et al. 2009).
Pseudoseptoria usneae Usnea perplexans, Usnea sp., AT, BL, FO
(GU1, IU2).
Raesaenenia huuskoneniiBryoria nadvor-
nikiana, Bryoria sp., AN, AT, FO, PK (GU6, IU4).
RC Rhymbocarpus pubescens (Etayo & Diederich) Diederich & Etayo – 4: on Lepraria finkii, AT
(GU1). The species is well characterized within the genus by its hairy exciple and association with
Lepraria. Another species occurring on the same host genus, R. neglectus (Vain.) Diederich & Etayo,
& 1998, &2000).
Sclerococcum serusiauxii Boqueras & Diederich – 4: on Parmelina carporrhizans, AT (GU1).
Sclerococcum simplexLepra trachythallina, CB (IU1).
Sphaerellothecium propinquellum (Nyl.) Cl.Roux & Triebel – 7: on Lecanora carpinea, AT (GU1).
Sphaerellothecium pumilum (Lettau) Nav.-Ros., Cl.Roux & Hafellner – 7: on Physcia aipolia, SA (IU1).
Sphinctrina anglica Nyl. – 7: on Protoparmelia oleagina, wood (JM1).
Sphinctrina turbinata (Pers.) De Not. – 2, 4: on Pertusaria coronata, P. pertusa, CB, FO (GU3, ZP1).
Stigmidium buelliae Zhurb. & Himelbrant – 7: on Buellia disciformis, FO (GU1).
Stigmidium congestum (Körber) Triebel – 4, 7: on Lecanora carpinea, L. chlarotera, L. leptyrodes, L.
pulicaris, AN, AT, SA (GU5, IU2, JM2, ZP1).
Stigmidium euclineVaricellaria hemisphaerica, AN, Ti, snag (GU2, IU2).
Stigmidium lecidellae Triebel, Cl.Roux & Le Coeur – 4: on Lecidella elaeochroma, FO (ZP1).
Taeniolella delicataLecidella flavosorediata, snag (GU1).
RC Taeniolella friesii (Hepp) Hafellner – 2, 3, 6: on Strigula stigmatella, AT, CO, FO, QU (GU2, IU1,
JV2, ZP3). On thallus of Strigula stigmatella
Tremella cetrariicola Diederich & Coppins – 3, 7: on Tuckermannopsis chlorophylla, FO (GU1, JV1).
Tremella hypogymniae Diederich & M.S.Christ. – 4: on Hypogymnia sp. (JM1).
RC Unguiculariopsis acrocordiae (Diederich) Diederich & Etayo – 2: on Acrocordia gemmata, CO
(ZP1). U. acrocordiae is well recognizable due to its tiny reddish/brownish ascomata with hairy
margin and its strict association with Acrocordia gemmata ( & 2000 &
2018).
Vouauxiella lichenicola (Linds.) Petr. & Syd. – 5: on Lecanora praesistens, AT (IU1).
Not or doubtfully lichenized fungi
C Actidium hysterioides Fr. – 2: wood (ZP, no voucher).
NC Agyrium rufum (Pers.) Fr. – 3, 7: AN/wood (ZP2). In Transcaucasia, this species has been reported
1986).
C Anisomeridium macrocarpum (Körb.) V.Wirth – 1, 2, 4, 6: AT, CA, FO, QU (GU2, JV2).
Atichia glomerulosa (Ach.) Stein – 2, 4: AN/ne, RP/le (GU1, ZP2).
Chaenothecopsis debilis (Sm.) Tibell – 3, 4, 7: FO/snag, wood (IU2, JM1, JV1).
Chaenothecopsis pusilla (Ach.) A.F.W.Schmidt – 2, 7: snag (GU2, JV1).
126 Herzogia 33 (1), 2020
Chaenothecopsis pusiola (Ach.) Vain. – 3, 4, 7: AN/wood, snag, stump (GU1, IU1, JV1, ZP1).
Chaenothecopsis rubescens Vain. – 7: AN (IU1).
Chaenothecopsis savonica (Räsänen) Tibell – 7: AN, snag (GU1, IU2).
Chaenothecopsis viridireagens
Cryptodiscus pallidus (Pers.) Corda – 1, 2, 5, 7: log, snag, stump (GU1, IU2, JM2, JV1, ZP3).
RC Cyrtidula major (Nyl.) Vain. – 5: BL (GU1, JM1).
Dennisiella babingtonii
C Exarmidium hemisphaericum (Fr.) Aptroot – 2, 4: log, snag (GU3, IU1). Ascospores 28 –30 × 8 –10
µm.
C Exarmidium inclusum (Pers.) Aptroot – 2, 3, 4, 7: log, snag (JV4, ZP3).
C Hysterium pulicare (Lightf.: Fr.) Pers. – 1: QU (ZP1).
RC Karschia cezannei Ertz & Diederich – 2: QU, Ti (ZP3). This recently described corticolous taxon
has been formerly apparently confused with members of Melaspilea s. lat., well distinguishable from
them on account of the periphyses in inner part of the apothecial margin and amyloid asci (see
&
taxon.
NC Kirschsteiniothelia aethiops
this species has been recently reported from Georgia ( 1986).
RC Kirschsteiniothelia recessa
Ascospores large, 18 –20 × 7 µm.
C Melaspileella proximella (Nyl.) Ertz & Diederich – 1, 2, 3, 4, 7: AC, AN, FO (JV2, ZP5).
C Microcalicium ahlneri Tibell – 1, 7: QU/wood, snag (IU1, ZP1).
Microcalicium disseminatum (Ach.) Vain. – 2, 4: AN, snag (GU2).
Mycocalicium subtile (Pers.) Szatala – 2, 3, 4, 7: AN/wood, AT/wood, PK/wood, QU/wood, SA/wood,
snag, stump (GU9, IU6, JM4, JV6, ZP2).
Mycomicrothelia wallrothii
C Mycoporum cf. antecellens (Nyl.) R.C.Harris – 4, 7: AN, AT, snag (GU1, JV2). Not lichenized, but
-
cial wall K-; asci with a distinct thollus; slender paraphysoids present; conidia c. 4 –5 × 1 µm.
Naetrocymbe fraxini (A.Massal.) R.C.Harris – 5: SA (JM1).
Naetrocymbe punctiformis (Pers.) R.C.Harris – 1, 3, 4, 5, 7: AN/tw, AT/tw, BL, CB, FO/tw, SA, Ti, UG/
tw (GU4, IU2, JM1, JV7, ZP1).
Peridiothelia fuliguncta
RC Pseudotryblidium neesii (Flot.) Rehm – 2, 6, 7: AN (ZP3). This is a distinctive non-lichenized leotia-
Abies
with lichens at bases of fir-trees. It was even regarded as a lichenicolous fungus by some earlier authors
(see 2011). The Caucasian material grows along with sterile thalli of Loxospora elatina
but evidently not directly associated with it. This species is quite easily identifiable due to its relatively
turning violet by KOH and two-celled ascospores in cylindrical asci are the most distinctive features
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 127
for the taxon (see 2011). Until now it was reported only from Europe, exclusively from
Abies albaAbies
alba and A. nordmanniana.
Rebentischia massalongii (Mont.) Sacc. – 1, 4, 7: AN, AT, UG (IU2, JV1). Only recently recorded for the
first time from the Caucasus and Russia ( & 2018).
C Rhizodiscina lignyota (Fr.) Hafellner – 7: wood (ZP1).
Sarea difformis (Fr.) Fr. – 3: AN (JM1).
Stenocybe pullatula (Ach.) Stein – 1: AG, CB (IU3).
Stictis radiata (L.) Pers. – 1, 2: dead twig of deciduous trees, Ti (IU1, ZP1).
We found a total of 659 species (ca. 16.1 % of the lichen flora of Russia, and ca. 32 % of the
lichen flora of the Northern Caucasus) in seven 1-hectare plots, including 564 lichenized, 61
lichenicolous and 34 allied non- or facultatively lichenized fungi that are often studied by
lichenologists. Plots 2, 4 and 7 had the most diverse lichen flora with more than 300 recorded
species; the lowest species richness, below 250 species, was observed in plots 5 and 6 (Table
1). Microlichens predominate in our species list – of 564 species, 407 were microlichens (ca.
72 % of lichenized taxa). The ratio of macrolichensis was higher at upper altitudes (plots 4, 5
and 7; Table 1).
Table 1. Total species richness and species richness in morpological and ecological groups observed in the investi-
gated plots. Plots are ordered according to increasing altitude.
Plot 1
700 m
Plot 6
720 m
Plot 2
940 m
Plot 3
1460 m
Plot 4
1720 m
Plot 7
1830 m
Plot 5
1900 m Total
Lichens 298 230 331 288 314 340 224 564
microlichens 219 161 235 199 215 232 134 407
macrolichens 79 69 96 89 99 108 90 157
% micro-/macrolichens 73.5/26.5 70/30 71/29 69/31 68/32 68/32 60/40 72/28
cyanolichens 17 17 28 20 22 25 15 34
the same, % 5.7 7.4 8.5 6.9 7 7.4 6.8 6
with trentepohlioid photobiont 50 39 51 43 38 27 15 79
the same, % 16.8 16.9 15.4 14.9 12.1 8 6.7 14
Lichenicolous fungi 3 4 16 10 21 20 17 61
Non-lichenized fungi 10 2 17 8 13 19 5 34
All groups 311 236 364 306 348 379 246 659
We found high diversity in the genera Lecanora (29 species), Arthonia s. lat. (21), Bacidia
s. lat. (18), Biatora (18), Caloplaca s. lat. (18), Micarea (18), Rinodina (18), Pertusaria s. lat.
(15), Usnea (14), Chaenotheca (13), Ochrolechia (10), Ramalina (10) and Gyalecta (8). On
the other hand, we found only a few species and low abundances of nitrophilous lichen genera,
e.g. Physconia (3 species), Xanthoria s. lat. (4), Phaeophyscia (5) and Physcia (6).
Abies nordmanniana (378
species) and Fagus orientalis (353), lower numbers are from Acer trautvetteri (256), Betula
litwinowii (199) and Carpinus betulus (184), and the lowest numbers are from Quercus (137),
Ulmus glabra (124), Sorbus aucuparia (120), Tilia begoniifolia (89) and others. Wood-
dwelling species (223) were recorded from snags (188) and logs (108).
128 Herzogia 33 (1), 2020
Discussion
Significant extensions of known geographical range
149 species (116 lichens, 17 lichenicolous fungi, 16 saprobic fungi) are new to the Northern
Caucasus (= Russian part of the Caucasian Mts), including 133 species (104 lichens, 15 li-
chenicolous fungi, 14 saprobic fungi) new to the Caucasus Mts.
Lichen species new to Russia are Andreiomyces obtusaticus, Bacidina mendax, Biatora
aegrefaciens, B. bacidioides, B. chrysanthoides, Biatorella dryophila, Buellia iberica,
Cliostomum haematommatis, Endohyalina ericina, Fellhanera christiansenii, Gyalidea minu-
ta, Japewia aliphatica, Lecanora barkmaniana, L. subravida, Lecidea strasseri, Leptogium
hibernicum, Lithothelium hyalosporum, L. phaeosporum, L. septemseptatum, Loxospora cris-
tinae, Melanelixia epilosa, Micarea nowakii, M. perparvula, Opegrapha trochodes, Orcularia
insperata, Parvoplaca servitiana, Phylloblastia inexpectata, Psoroglaena stigonemoides,
Ptychographa xylographoides, Ramonia dictyospora, R. luteola, Rinodina polysporoides,
Thelopsis flaveola, Topelia jasonhurii, Verrucaria hegetschweileri, Wadeana minuta and
Waynea giraltiae.
Lichenicolous fungi new to Russia are Arthonia vorsoeensis, Didymocyrtis melanelixiae,
Epigloea urosperma, Muellerella polyspora, Phacographa zwackhii, Pronectria pilosa,
Rhymbocarpus pubescens, Taeniolella friesii and Unguiculariopsis acrocordiae.
Non-lichenized fungi new to Russia are Cyrtidula major, Karschia cezannei,
Kirschsteiniothelia recessa and Pseudotryblidium neesii.
Genera new to Russia are Andreiomyces, Lithothelium, Orcularia, Phylloblastia, Topelia and
Wadeana.
Another five lichen species new to Russia have been recently described and our specimens
are already published: Bacidia albogranulosa ( et al. 2018a), Biatora radicicola
( et al. 2016), Blastenia anatolica ( et al. 2019a), Lecanora stanislai
( et al. 2017) and Lecidea coriacea ( et al. 2016).
Pronectria pilosa ( et al. 2009),
Cliostomum haematommatis, Endohyalina
ericina, Leptogium hibernicum, etc. (
Aegean Islands, Greece for Buellia iberica ( &
the Carpathians Mts for Verrucaria hegetschweileri ( et al. 2006). Moreover, the range
of Topelia jasonhurii
Ecological aspects
The general characteristics of the observed diversity correspond with data obtained from an
altitudinal gradient in Carpathian beech dominated forests ( et al. 2018a,
et al. 2015, 2018). All plots tend to harbour rare species that characteristically occur in old-
growth forests with a long lasting continuity. In contrast, species considered nitrophilous (e.g.,
frequent European species of Teloschistaceae and Physciaceae) have low species richness and
low abundances in all plots which possibly indicates a negligible effect of air pollution caused
by emissions of fixed nitrogen.
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 129
In plots at low altitudes (below 1000 m), we recorded numerous species and genera character-
istic for a warm-humid or oceanic climate (e.g., Bactrospora dryina, Byssoloma leucoblephar-
um, Fellhanera spp., Gabura fasciculare, Hypotrachyna laevigata, H. revoluta, Lithothelium
spp., Maronea constans, Myelochroa aurulenta, M. metarevoluta, Parmotrema perlatum,
Phylloblastia inexpectata, Phyllogyalidea phyllophila, Thelotrema lepadinum, Topelia jason-
hurii and Vahliella saubinetii). In plots at high altitudes, we recorded some typically subal-
pine species preferring cool-humid climate (e.g., Anzina carneonivea, Caloplaca sorocarpa,
Parvoplaca tiroliensis, Rinodina magelanica and Tetramelas chloroleucus) and boreal species
(e.g., Cetraria sepincola, Fuscidea recensa, Japewia subaurifera, Ptychographa xylograph-
oides and Pycnora xanthococca).
The number of cyanolichen species ranged 15 –28 species per plot (total 34 species; Table
1). Although these are seemingly low numbers, they are substantially higher than in ancient
forests which were affected by former or present air pollution. For example, old-growth for-
est plots in the Czech Republic are often without cyanolichens, or only one or few Peltigera
-
anolichens is demonstrated for other regions affected by air pollution, e.g. the New Forest in
England ( 2010). Lichens with a trentepohlioid photobiont ranged 15 –51 species
per plot (total 79 species; Table 1). They were distinctly more represented in plots at low al-
titudes correspondingly with results by et al. (2017) and et al. (2018).
Ratio of foliose plus fruticose lichens (macrolichens) versus crustose lichens (microlichens)
in our surveyed plots was between 26.5 – 40 % and it rose with altitude. The same trend was
et al. 2018).
Positive correlation of macrolichens ratio with altitude may be explained by high demands of
macrolichens for humidity and light accessability that both increase with altitude (et
al. 1997, 2010, et al. 2016).
Substrate specificity and selectivity of epiphytic lichens
et al.
2009, et al. 2009, 2012, et al. 2013). In our study, the greater species
richness was found on silver fir and beech which is not a surprising fact as both tree species are
of species recorded from birch is caused by the birch dominance in the subalpine plot.
Although beech and fir are the most preferred substrata, most lichen species have a low speci-
ficity to fir or beech. About 90 % of recorded species were observed on three or more types of
substrates. Species recorded on less than three types of substrata may be either substrate spe-
cific (e.g. Gyalidea minuta and Multiclavula mucida on logs and snags, and Stenocybe pullat-
ula mostly on Alnus twigs), or were rarely recorded (e.g. Cetraria sepincola, Lecanora albella,
Physcia tribacia and Usnocetraria oakesiana). In the latter case, we suggest a lower substrate
specificity, which could be tested by extended sampling. Among 223 lichenized species occur-
ring on wood (logs, snags, stumps), only a low portion (56 species) is strictly lignicolous (i.e.
Plot 7 – the most diverse lichen flora was found beyond the protected area
The present study is focused on the unique area in the NW Caucasus designated as a UNESCO
130 Herzogia 33 (1), 2020
some even new to Russia. However, the highest species richness (379 species per hectare)
was found in the seventh plot which is situated beyond the borderline of the Caucasus Reserve
(details in the list of studied sites). The lichenological value of this plot is not only in the large
number of species but also in the presence of rare and threatened species. Five species are
et al. 2008). Among them,
Letharia vulpina, Lobaria pulmonaria and Usnea florida are vulnerable (VU), and Leptogium
burnetiae and Ricasolia amplissima are near threatened (NT). The high diversity of lichens in
Plot 7 can be explained by its large forest habitat diversity, the high number of tree species and
local hot-spot that we located within our research, we strongly recommend an extension of the
Caucasus Reserve to include woodlands beyond its current northern boundary.
Acknowledgements
(Cladonia), O. Breuss (Wien) (Verrucaria cf. lignicola), P. Clerc (Genève) (Usnea), B. Coppins (Edinburgh) (Micarea
perparvulaBacidia s.lat.), A. Frisch (Trondheim) (ArthonialesLecanora
stanislai), H. Mayrhofer (Graz) (Amandinea & RinodinaBryoria), A. Orange (Cardiff) (Porina
byssophila). We received support from the long-term research development grant RVO [67985939] and the Russian
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Communicated by: Holger Thüs
Addresses of the authors
Gennadii Urbanavichus, Institute of North Industrial Ecology Problems, Kola Science Centre,
Russia. E-mail: g.urban@mail.ru
Irina Urbanavichene, Komarov Botanical Institute, Russian Academy of Sciences, Professor
Popov Str. 2, 197376 St Petersburg, Russia. E-mail: urbanavichene@gmail.com
136 Herzogia 33 (1), 2020
Czech Republic & University of South Bohemia, Faculty of Biological Sciences, Department
Appendix.
Taxon Sample ITS mtSSU
Amandinea sp. (A. aff. punctata) -MK778513
Bacidia albogranulosa MK158339 MK158334
Bacidia albogranulosa MK158340 MK158335
Biatora sp. (B. amylacea ined.) MK778585 MK778516
Biatora efflorescens MK778586 MK778517
Biatora chrysantha MK778587 -
Biatora pontica MK778588 MK778518
Biatora pontica -MK778519
Biatora vacciniicola -MK778520
Blastenia anatolica Z. Palice 21647 (PRA) MK778590 -
Blastenia herbidella MK778591 -
Buellia schaereri MK778592 MK778521
Caloplaca turkuensis - MK778522
Caloplaca turkuensis - MK778523
Candelariella faginea MK778596 MK778524
Catinaria sp. MK778597 -
Cliostomum haematommatis MK778598 MK778525
Cliostomum haematommatis MK778599 MK778526
Fellhanera sp. MK778600 -
Gyalecta herculina MK778584 MK778515
Gyalecta herculina MK778601 -
Gyalectaceae sp. - MK778527
Lecania croatica MK778602 -
Lecania cf. cyrtella Z. Palice 21641 (PRA) MK778603 MK778528
Lecania cf. cyrtella G. Urbanavichus s.n. (hb. GU) - MK778582
Lecanora allophana KY548050 KY502421
Lecanora argentata MK778604 MK778529
Lecanora barkmaniana MK778605 MK778530
et al.: Lichens of virgin forests in the Caucasus Reserve (Russia) 137
Taxon Sample ITS mtSSU
Lecanora carpinea s.str. MK778606 MK778531
Lecanora carpinea s.str. MK778607 MK778532
Lecanora carpinea s.str. MK778608 MK778533
Lecanora carpinea s.str. - MK778534
Lecanora cinereofusca - MK778535
Lecanora exspersa KY548053 KY502420
Lecanora exspersa KY548054 KY502419
Lecanora exspersa KY502415 KY548057
Lecanora exspersa MK778609 MK778536
Lecanora leptyrodes - MK778537
Lecanora leptyrodes MK778610 MK778538
Lecanora pulicaris MK778611 MK778539
Lecanora pulicaris MK778612 MK778540
Lecanora pulicaris - MK778541
Lecanora pulicaris Z. Palice 23005 (PRA) - MK778542
Lecanora sp. - MK778543
Lecanora sp. MG076967 -
Lecanora stanislai - MK778544
Lecanora stanislai - MK778545
Lecidea sp. MK778613 MK778546
Lecidella aff. flavosorediata MK778614 -
Lecidella flavosorediata MK778615 MK778547
Leptogium burnetiae - MK778548
Leptogium saturninum - MK778549
Loxospora cristinae MK778617 MK778550
Loxospora cristinae - MK778551
Loxospora cristinae - MK778552
Loxospora cristinae - MK778553
Loxospora cristinae MK778619 -
Loxospora cristinae MK778620 -
Loxospora cristinae MK778621 MK778554
Marchantiana asserigena MK778593 -
Megalospora porphyritis MK778622 MK778555
Melanelixia epilosa MK778623 MK778556
Micarea nowakii - MK778557
Micarea nowakii - MK778558
138 Herzogia 33 (1), 2020
Taxon Sample ITS mtSSU
Mycobilimbia epixanthoides MK778624 MK778559
Mycoblastus sp. MK778625 MK778560
Ochrolechia pallescens MK778626 MK778561
Ochrolechia trochophora MK778627 MK778562
Parmelia barrenoae MK778628 -
Parmelia ernstiae MK778629 MK778563
Parmelia saxatilis MK778630 MK778564
Parmelia saxatilis MK778631 MK778565
Parmelia saxatilis MK778632 -
Parmelia serrana MK778633 MK778566
Parmelia serrana MK778634 MK778567
Parmelia serrana MK778635 MK778568
Parmelia submontana - MK778569
Parvoplaca servitiana MK778594 MK778570
Parvoplaca tiroliensis MK778595 -
Pertusaria pupillaris - MK778571
Physcia cf. biziana MK778637 MK778572
Ramalina panizzei Z. Palice 22766 (PRA) MK778638 -
Rinodina griseosoralifera - MK778573
Rinodina malangica - MK778574
Rinodina sp. MK778639 MK778575
Rinodina sheardii MK778640 -
Rinodina sheardii - MK778576
Sagedia aff. mastrucata Z. Palice 23193 (PRA) MK778583 MK778514
Scoliciosporum cf. umbrinum MK778641 MK778577
Scoliciosporum sp. MK778642 MK778578
Schaereria corticola - MK778579
Tetramelas sp.Z. Palice 23089 (PRA) MK778643 -
unidentified lichen MK778644 MK778580
Xanthomendoza ulophyllodes MK778645 -
Xylographa parallela MK778636 -
Xylographa parallela MK778589 MK778581
Xylographa parallela Z. Palice 22099 (PRA) MK778618 -
