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21Revision of crustose Teloschistaceae (lichenized Ascomycota) from the Russian Far East based on herbarium materials of the Komarov Botanical Institute
I. V. Frolov1, L. V. Gagarina2
'Botanical Garden of the Ural Branch of the Russian Academy of Sciences, Ekaterinburg, Russia 2Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia Corresponding author. I. V. Frolov, ivfrolov@gmail.com
Abstract. A revision of 47 specimens of crustose Teloschistaceae (so-called "Caloplaca s. lat.") from the Russian Far East in the herbarium LE belonging to 21 species is resulted in re-identification of 18 specimens and new records. Blastenia furfuracea from the Kamchatka Peninsula is new to the Russian Far East. Caloplaca fraudans is new to the Kamchatka Peninsula. Records of Blastenia ferruginea, B. hungarica, Caloplaca haematites and Solitaria chrysophthalma are doubtful for the Far East. Placodium haematites var. ussuriense Tomin [= Caloplaca haematites var. ussuriensis (Tomin) Zahlbr.] is lectotypified and synonymized under Caloplaca stillicidiorum s. lat.
Keywords: Blastenia furfuracea, Caloplaca fraudans, Caloplaca haematites var. ussuriensis, Chu-kotka, Kamchatka, Primorye Territory.
Ревизия накипных лишайников семейства Teloschistaceae с российского
Дальнего Востока по материалам гербария Ботанического института
им. В. Л. Комарова РАН
И. В. Фролов1, Л. В. Гагарина2
'Ботанический сад УрО РАН, Екатеринбург, Россия 2Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия Автор для переписки. И. В. Фролов. ivfrolov@gmail.com
Резюме. В гербарии Ботанического института им. В. Л. Комарова РАН (LE) обнаружено 47 образцов накипных лишайников семейства Teloschistaceae (род Caloplaca s. lat.), собранных на территории российского Дальнего Востока. Они относятся к 21 виду. Восемнадцать образцов были переопределены. Blastenia furfuracea с Камчатки является новым видом для российского Дальнего Востока. Caloplaca fraudans является новым видом для Камчатки. Находки Blastenia ferruginea, B. hungarica, Caloplaca haematites и Solitaria chrysophthalma сомнительны для Дальнего Востока. Обозначен лектотип названия разновидности Placodium haematites var. ussuriense (= Caloplaca haematites var. ussuriensis), которая сведена в синонимы к Caloplaca stillicidiorum s. lat.
Ключевые слова: Blastenia furfuracea, Caloplaca fraudans, Caloplaca haematites var. ussuriensis, Камчатка, Приморье, Чукотка.
The family Teloschistaceae Zahlbr. includes more than 1000 crustose species and most of them were formerly placed in the genus Caloplaca Th. Fr. However, several years ago Arup et al. (2013) completely revised the taxonomy of Teloschistaceae and
https://doi.org/10.31111/nsnr/2020.54.1.139
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retained within Caloplaca s. str. only 12 species. Nevertheless, often lichenologists still unite crustose members of the family under the name "Caloplaca s. lat.". During the last decade a lot of new species of Caloplaca s. lat. were described and the concepts of others were changed (e. g., Sochting et al., 2008; Arup, 2009; Vondrak et al., 2009, 2013a, 2016; Kondratyuk et al., 2011; Frolov et al., 2016).
Caloplaca kamczatica (Savicz) Savicz was the first species described from the Russian Far East (Savicz, 1914, as Placodium kamczaticum Savicz). Tomin (1926) described from the vicinity of Vladivostok a new species Caloplaca gordejevii (Tomin) Oxner ex Khodos. (as Placodium gordejevii Tomin) and a new variation Caloplaca haematites var. ussuriensis (Tomin) Zahlbr. (as Placodium haematites var. ussuriense Tomin). In the list of species Lichens of Russia (Spisok..., 2010) 56 species of Caloplaca s. lat. were reported for the Russian Far East (excluding the Republic of Sakha). The list included mainly common European, Arctic-alpine and boreal species of the genus. That was surprising for such a unique territory within Russia and indicated scarcity of our knowledge on the group in the Russian Far East, which was partly demonstrated by describing of several new Caloplaca s. lat. species from this part of Russia after 2010 by Kondratyuk et al. (2011, 2013, 2014, 2015).
In the frame of the revision of Caloplaca s. lat. in the lichens herbarium of the Komarov Botanical Institute RAS (LE) we studied specimens from the Russian Far East (here we do not include the Republic of Sakha to the region). Considering our insufficient knowledge on diversity of the family Teloschistaceae in the region as well as recent changes in the concepts of many species, we decided to publish the list of the Caloplaca s. lat. taxa from the Russian Far East deposited in the herbarium LE. The majority of the revised specimens were previously published mainly in Khodosovstev et al. (2004) and Makarova, Katenin (2009). Several specimens seem to be published here for the first time.
Material and methods
Specimens of Caloplaca s. lat. deposited in the herbarium of the Komarov Botanical Institute (LE) represented the material for the study. Measurements of morphological characters follow Vondrak et al. (2013b). All microscopic observations were based on hand-cut sections mounted in water, without chemical treatments. We follow the taxonomy of the family Teloschistaceae proposed by Arup et al. (2013). We use square brackets in citation of a specimen label when the information is absent in an original label and have been added by us. A question mark is applied in case of uncertainty in correct reading of a label or administrative attachment of a locality. We use quotes for the names of lichens on the original labels if we were not able to find a name in Index Fungorum or MycoBank. If it is not specified in the text, the specimen was identified by the first author.
Results
We revised forty-seven specimens of Caloplaca s. lat. mainly from Kamchatka Territory and Chukotka Autonomous Area as well as from Khabarovsk Territory, Maga-
dan Region, and Primorye Territory. The majority of the specimens were collected by V. P. Savicz in Kamchatka and I. I. Makarova in Chukotka. Sporadic collections from the southern part of the Russian Far East belong to I. A. Galanina, T. P. Gordeev, A. N. Oxner, and others. We re-identified 18 specimens and compiled a list of 21 species.
List of species
Athallia pyracea (Ach.) Arup et al. — Kamchatka Territory: [Elizovo District, Elizovo] (Zavoyko), along bank of Avacha River, 1908, Savicz 6056, LE L s. n. [probably det. Savicz sub Placodium cerinum (Hedw.) Nägeli ex Hepp, det. Khodosovtsev, 2004, sub Caloplaca pyracea (Ach.) Zwackh and published in Khodosovtsev et al. (2004)].
A. saxifragarum (Poelt) Arup et al. — Magadan Region: in the specimen LE L-13863 (see Caloplaca stillicidiorum).
Blastenia ammiospila (Wahlenb.) Arup et al. — Chukotka Autonomous Area: NE of Chukotka Peninsula, downstream of Utaveem River, left bank, near Mt. Gynron, 1991, Makarova, LE L-631 [det. Makarova sub Caloplaca ammiospila (Wahlenb.) H. Olivier and probably published without a number in Makarova, Katenin (2009)]; Kamchatka Territory: [Petropavlovsk-Kamchatskii], Zavoyko, on bark of shrub, 1908, Savicz 6150, LE L-14080 [probably det. Savicz sub "Placodium ferrugineum var. ammiospila", det. Khodosovtsev, 2004, sub Caloplaca hungarica H. Magn. and published in Khodosovtsev et al. (2004)]; [Elizovo District, on way from village] Malki [to village] Ganaly, on bark of Populus, 1909, Savicz 5113, LE L-14076 [probably det. Savicz sub "Placodium ferrugineum var. corticolum", det. Khodosovtsev, 2004, sub Caloplaca hungarica and published in Khodosovtsev et al. (2004)].
B. furfuracea (H. Magn.) Arup et al. — Kamchatka Territory: [Elizovo District], village Nachiki, Uzdach River, on bark, 1908, Savicz 5738, LE L-14078 [probably det. Savicz sub Placodium ferrugineum (Huds.) Hepp, det. Khodosovtsev, 2004, sub Caloplaca hungarica and published in Khodosovtsev et al. (2004)]. New to the Russian Far East.
Bryoplaca jungermanniae (Vahl) Sochting et al. — Chukotka Autonomous Area: Chaun-skii District, SW coast of Chaun Bay, riverhead Pineyveen, the middle part of the slope, sedge-forb steppe, 1982, Slinchenkova plot 3, LE L-7235 [det. Makarova sub Caloplaca tetrasporella (Nyl.) H. Olivier]; 32nd km of road Egvekinot-Iul'tin, 1971, Makarova, LE L-616 [det. Makarova sub C. jungermanniae (Vahl) Th. Fr.]; NE of Chukotka Peninsula, downstream of river Che-gitun, junction of Vetgyvayam River, 1991, Makarova, LE L-628 [det. Makarova sub C. jungermanniae and probably published without a number in Makarova, Katenin (2009)]; SE of Chukotka Peninsula, NW coast of Penkichnei Bay, lower part of S slope, overgrown slide-rock, 1978, Katenin plot 90, LE L-14257 (det. Makarova, 1980, sub C. jungermanniae).
B. tetraspora (Nyl.) Sochting et al. — Kamchatka Territory: [Elizovo District ?], second half of pass from valley of Shchapina River to Kronotskaya side, alpine zone, 1909, Savicz 6350, LE L-14256 [det. Savicz sub "Placodium jungermanniae var. globosum Savicz nov. variet.", det. Khodosovtsev, 2004, sub Caloplaca tetraspora (Nyl.) H. Olivier and published in Khodosovtsev et al. (2004)]; Chukotka Autonomous Area: in the specimen LE L-616 (see Bryoplaca junger-manniae).
Note: the name Placodium jungermanniae var. globosum Savicz is a nomen nudum and is not therefore validly published [ICN (Shenzhen Code) Art. 38.1(a)].
Caloplaca borealis (Vain.) Poelt — Chukotka Autonomous Area: in the specimen LE L-631 (see Blastenia ammiospila).
C. caesiorufella (Nyl.) Zahlbr. — Chukotka Autonomous Area: Pitlekai, 1878, Almquist, LE L-14122 [det. Vainio sub Placodium ferrugineum var. ferrugineofuscum Vain., published in
Wainio (1909)]; NE of Chukotka Peninsula, junction of rivers Chegitun and Vetgyvayam, 1991, Makarova, LE L-1316 [probably det. Makarova sub Caloplaca jungermanniae and then probably revised and published without a number in Makarova, Katenin (2009) sub Caloplaca cf. caesiorufella]; in the specimen LE L-628 (see Bryoplaca jungermanniae); Kamchatka Territory: Ust'-Kamchatsk District, ca. 24 km SE of Kozyrevsk, Kamchatka River basin, SW slope of Ushkovskii volcano, tundra with dwarf shrubs and graminoids, on lignum of Salix spheno-phylla A. K. Skvortsov, 1550 m a. s. l., 2004, Himelbrant and Kuznetsova K-101-04, LE L-6937, det. Himelbrant and Stepanchikova, 2008.
C. etesiae (Nyl.) Du Rietz — Kamchatka Territory: [Elizovo District], cape in Rakovaya Bay, before colony of lepers, 1909, Savicz 5025, LE L-14356, det. Khodosovtsev, 2004 [probably det. Savicz sub "Placodium murorum var. major f. scopulorum"; published in Khodosovtsev et al. (2004)].
C. fraudans (Th. Fr.) H. Olivier — Kamchatka Territory: [Mil'kovo District, former] village Mashura, on wood of bridge, 1909, Savicz 5480, LE L-14077 [probably det. Savicz sub "Placodium ferrugineum var. genuinum", det. Khodosovtsev, 2004, sub Caloplaca hungarica and published in Khodosovtsev et al. (2004)]. New to the Kamchatka Peninsula.
C. fulvolutea (Arnold) Jatta — Kamchatka Territory: [Kronotskii Nature Reserve], Shchapinskie hot springs, 1909, Savicz 6384, det. Khodosovtsev, 2004, LE L-14255 [probably det. Savicz sub Placodium jungermanniae var. subolivacea (Th. Fr.) Vain.].
Note: published as the new species to Russia by Khodosovtsev et al. (2004). However, the specimen looks much damaged and correctness of the identification is doubtful for us.
C. gordejevii (Tomin) Oxner ex Khodos. — Kamchatka Territory: Elizovo District, middle stream of Kronotskaya River, forest with Betula sp. and Populus suaveolens Fisch., on bark of Populus, 1982, Mikulin 2-118-d, LE L-14187 [det. Mikulin sub Caloplaca aurantiaca (Lightf.) Th. Fr., det. Khodosovtsev and Himelbrant, 2004, and published in Khodosovtsev et al. (2004)]; [Elizovo District], between Koryaki and Nachiki, "Narrow plateau" by Koryatskaya [River],
1908, Savicz 6038, LE L-14188, 14189 [probably det. Savicz sub Placodium aurantiacum var. salicinum (Schrad.) Anzi, det. Khodosovtsev and Himelbrant, 2004, and published in Khodosovtsev et al. (2004)]. Primorye Territory: [Vladivostok], Reineke Island, E slope of main summit of island, on bark of trees, 1915, Gordeev, LE L-448, lectotype, selected in Khodosovtsev et al. (2004); Vladivostok, teum (?) Ljan-tschi-che (?), on bark of Quercus mongolica Fisch. ex Turcz., 1927, Oxner 10747/a, LE L-14190, det. Oxner.
C. kamczatica (Savicz) Savicz — Kamchatka Territory: along mouth of Nalachevaya River, 1909, Savicz 6564, LE L-451, lectotype, selected in Khodosovtsev et al. (2004), LE L-450, 452, isolectotypes; Kronotskii Gulf, seashore, 1909, Savicz 6584, LE L-454; along Vakhil' River,
1909, Savicz 6504, LE L-453.
C. stillicidiorum (Vahl) Lynge, 1921, Kunglige Norske Videnskabers Selskab Skrifter 15: 4.
Syn. nov. Placodium haematites (Chaub.) Anzi var. ussuriense Tomin, 1926, Bull. South. Ussuri Branch State Russ. Geograph. Soc. 12: 218. = Caloplaca haematites var. ussuriensis (Tomin) Zahlbr., 1930 [1931], Zahlbruckner's Cat. Lich. Univ. 7: 143. — Type: [Russia, Primorye Territory, Ussuriisk], village Sinelovka, slopes of Khunkhus Mt., on rocks, 1915, Gordeev, LE L-14901 (lectotype, designated here; Fig. 1).
Chukotka Autonomous Area: NE of Chukotka Peninsula, downstream of Chegitun River, near the junction of Vetgyvayam River, 1991, Makarova, LE L-664 [probably published without a number in Makarova, Katenin (2009) sub Caloplaca cerina]; Kamchatka Territory: [Elizovo District], cape near lake "Solenoe" before entrance to Rakovaya Bay, on coastal rocks, 1909, Savicz 4999, LE L-14130 [det. Savicz sub Placodium gilvum var. stillicidiorum (Vahl) Vain., det. Khodosovtsev, 2004, sub Caloplaca cerina (Hedw.) Th. Fr. and published in Khodosovtsev
Fig. 1. Lectotype of Placodium haematites (Chaub.) Anzi var. ussuriense Tomin (1915, Gordeev, LE L-14901). Scale bar — 0.5 mm.
et al. (2004)]; [Kronotskii Nature Reserve], Shchapinskie hot springs, 1909, Saviez 6174, LE L-14131 [probably det. Saviez sub Placodium gilvum var. shrhurkii (?) Th. Fr. (?), det. Kho-dosovtsev, 2004, sub Caloplaca cerina and published in Khodosovtsev et al. (2004)]; Magadan Region: E part of Wrangel Island, Krasnyi Flag River, calcareous scree loamy slope, in spotty Dryas integrifolia Vahl tundra, on dead Saxifraga sp., 100 m a. s. l., 1997, Kholod, LE L-13863 [det. Zhurbenko, 2002, sub Caloplaca celata Th. Fr. and probably published without a number in Kholod, Zhurbenko (2005)]; Primorye Territory: see above the lectotype of Placodium haematites var. ussuriense.
Note: apothecia morphology of the lectotype of Placodium haematites var. ussuriense (up to 1 mm diam., thick tortuous thalline margin, absence of well-developed proper margin) corresponds to the genus Caloplaca s. str., but not to C. haematites group. Its ecology corresponds to the epilithic forms of C. stillicidiorum s. lat.
Here we follow a wide species concept of C. stillicidiorum proposed by Soun et al. (2011) whoes molecular phylogeny demonstrated that the species is polyphyletic and consists of several lineages which may be only partly interpreted on the basis of their morphology and ecology.
C. trassii Galanina et S. Y. Kondr. — Primorye Territory: [Ussuriisk], 10 km SE of Korfo-vka, upper part of slope, oak-birch forest, 2006, Galanina, LE L-10595, det. Galanina, 2011 [published in Kondratyuk et al. (2011)]; Khasanskii District, Zarubino, in the garden on apricot, 2010, Galanina, LE L-10596, det. Galanina, 2011.
Caloplaca sp. — Kamchatka Territory: in the specimen Savicz 6056 (see Athallia pyracea), probably published in Khodosovtsev et al. (2004) sub Caloplaca haematites (Chaub.) Zwackh. The specimen likely belongs to the genus Athallia Arup et al.
Gyalolechia oxneri (S. Y. Kondr. et Sochting) Sochting et al. — Primorye Territory: Islands of Bay of Great Peter, Popova Island, young Alneta forest, on Fraxinus sp. (60 cm diam.), 1989, Kondratyuk, det. Kondratyuk, LE L-1236.
G. ussuriensis (Oxner et al.) Vondrak — Kamchatka Territory: [Ust'-Kamchatsk District ?, former] village Tolbachik, along river Tolbachik, on aspen, 1909, Savicz 5440, LE L-13973
[probably det. Savicz sub Placodium citrinum (Hoffm.) Hepp, det. Khodosovtsev, 2004, sub Ca-loplaca chrysophthalma Degel. and published in Khodosovtsev et al. (2004)]; [Mil'kovo District], between village Kirganik and [former] village Mashura, on aspen, 1909, Savicz 5477, LE L-13974 [probably det. Savicz sub Placodium citrinum; probably published as Savicz 5473 (Ca-loplaca chrysophthalma) in Khodosovtsev et al. (2004)]. Primorye Territory: Khankaiskii District, 2 km of village Rubinovka, floodplain of Malokanovka River, on Salix sp., 2007, Galanina P-1, LE L-10592, det. Galanina [published in Kondratyuk et al. (2011)]; Ussuriisk District, Borisovskoe plateau, 6 km of Mt. Strelkovaya, herb-rich Quercus mongolica forest on gentle slope, 510 m a. s. l., 2008, Yakovchenko, LE L-10593, det. Galanina, 2012.
Polycauliona verruculifera (Vain.) Arup et al. — Kamchatka Territory: Elizovo District, mouth of river Semyachik, 10 km to N, oceanic island between spring Goryachii and river Per-vaya Rechka, on coastal rocks, 1985, Mikulin 2-095-d, LE L-14172 [probably det. Mikulin sub Caloplacagranulosa (Müll. Arg.) J. Steiner, det. Khodosovtsev, 2004, sub C. verruculifera (Vain.) Zahlbr. and published in Khodosovtsev et al. (2004)]. Khabarovsk Territory: Iony Island in the Sea of Okhotsk, 1948, Pavlinovplot3, LE L-14898 (probably det. Pavlinov sub C. verruculifera).
Rufoplaca sp. — Kamchatka Territory: Elizovo District, Kronotskii Peninsula, upper stream of Bolshoy Chapsly River, summit 1100 m a. s. l., on stones, 1981, Mikulin 1-120-d, LE L-14081 [probably det. Mikulin sub Caloplaca ferruginea (Huds.) Th. Fr., the specimen mentioned in Khodosovtsev et al. (2004) as dubious in discussion under C. (Blastenia) ferruginea]; slope to S from valley of ridge "Morskoi", Tar'inskaya Bay, Avachinskii Gulf, sparse Betula forest on slope, on rocks, 1908, Savicz, LE L-14079 (probably det. Savicz sub Placodium ferrugineum); Khabarovsk Territory: [district unknown], village Malmyzh, on rock, 1909, Fedtschenko 964, LE L-14124 (probably det. Fedtschenko sub P. ferrugineum).
Xanthoria sp. — Kamchatka Territory: Petropavlovsk[-Kamchatskii], Mt. Nikol'skaya, rocks in forest, 1908, Savicz 351, LE L-14355 (probably det. Savicz sub "Placodium murorum var. major").
List of re-identified specimens
Caloplaca celata (Kholod, LE L-13863) ^ C. stillicidiorum
C. cerina (Savicz 4999, LE L-14130) ^ C. stillicidiorum
C. cerina (Savicz 6174, LE L-14131) ^ C. stillicidiorum
C. chrysophthalma (Savicz 5440, LE L-13973) ^ Gyalolechia ussuriensis
C. chrysophthalma (Savicz 5477, LE L-13974) ^ Gyalolechia ussuriensis
C. ferruginea (Mikulin 1-120-d, LE L-14081) ^ Rufoplaca sp.
C. haematites (Savicz 6056, LE L s. n.) ^ Caloplaca sp.
C. hungarica (Savicz 6150, LE L-14080) ^ Blastenia ammiospila
C. hungarica (Savicz 5113, LE L-14076) ^ Blastenia ammiospila
C. hungarica (Savicz 5738, LE L-14078) ^ Blastenia furfuracea
C. hungarica (Savicz 5480, LE L-14077) ^ C. fraudans
C. jungermanniae (Makarova, LE L-1316) ^ C. caesiorufella
C. tetrasporella (Slinchenkovaplot 3, LE L-7235) ^ Bryoplaca jungermanniae
Placodium ferrugineum (Savicz, LE L-14079) ^ Rufoplaca sp.
P. ferrugineum (Fedtschenko 964, LE L-14124) ^ Rufoplaca sp.
P. ferrugineum var. ferrugineofuscum (Almquist, LE L-14122) ^ Caloplaca caesiorufella P. haematites var. ussuriense (Gordeev, LE L-14901, lectotype) ^ Caloplaca stillicidiorum s. lat. P. murorum var. major (Savicz 351, LE L-14355) ^ Xanthoria sp.
Discussion
During the revision we found one species new to the Russian Far East (Blastenia furfuracea) and one species new to the Kamchatka Peninsula (Caloplaca fraudans). Blastenia furfuracea occurs on tree bark or weathered wood in the boreal zone. In Russia it is known from the Ural Mts, Altai Mts and from the Kodar Ridge in Trans-Baikal Territory (Vondrak et al, 2019). Caloplaca fraudans is an arctic species growing on eutrophicated wood or non-calciferous rock. In Russia it is known from the arctic and northern regions of the European part and Siberia (Spisok..., 2010).
After revision of the crustose Teloschistaceae in the herbarium LE and investigation corresponding literature sources reports of some species for the Russian Far East became doubtful for us. Below we discuss such species.
Comments on some doubtful taxa
Blastenia ferruginea (Huds.) A. Massal. [= Caloplaca ferruginea (Huds.) Th. Fr.]. According to Vondrak et al. (2019) the species occurs only in Mediterranean and Western Europe. Reports from the Russian Far East most likely belong to corticolous specimens of Blastenia ammiospila, as well as to Caloplaca caesiorufella, Rufoplaca spp. and others.
B. hungarica (H. Magn.) Arup et al. [= Caloplaca hungarica H. Magn.]. According to Vondrak et al. (2019) the species is restricted to Europe and adjacent Mediterranean regions. Reports from the Russian Far East most likely belong to corticolous specimens of Blastenia ammi-ospila, Caloplaca fraudans etc.
Caloplaca celata Th. Fr. On the base of molecular data Vondrak et al. (2012) suggested that C. celata represents a variant of C. stillicidiorum s. lat. lacking anthraquinones.
C. etesiae (Nyl.) Du Rietz and Polycauliona brattiae (W. A. Weber) Arup et al. Caloplaca (Polycauliona) brattiae W. A. Weber was synonymized with C. etesiae by Khodosovtsev et al. (2004). However, it was not accepted by Arup et al. (2013) and McCune (2017). We did not study this issue.
C. haematites (Chaub.) Zwackh. So far doubtless records of the species are only known from Europe and adjacent Mediterranean regions. Reports from the Russian Far East most likely belong to Caloplaca cerina s. lat. and C. stillicidiorum s. lat.
Solitaria chrysophthalma (Degel.) Arup et al. [= Caloplaca chrysophthalma Degel.]. So far doubtless records of the species are only known from Europe. Reports from the Russian Far East most likely belong to other yellow sorediate crusts — Calogaya bryochrysion (Poelt) Vondrak, Gyalolechia epiphyta (Lynge) Vondrak and G. ussuriensis.
Acknowledgements
We are grateful to L. A. Konoreva for the idea of the project and comprehensive assistance during its implementation. We also thank the anonymous reviewer and the editor for the revision of the manuscript and their valuable comments. The first author thanks the stuff of the Laboratory of Lichenology and Bryology of the Koma-rov Botanical Institute RAS (especially D. E. Himelbrant and S. V. Chesnokov) for hospitality and help during his work in the herbarium LE. This study was supported by the Russian Foundation for Basic Research (projects no 17-34-50115 and no 19-04-00074). I. V. Frolov worked in the frame of the national project of the Institute Botanical Garden (Russian Academy of Sciences, Ural Branch).
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Литература
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Frolov I., Vondrák J., Fernández-Mendoza F., Wilk K., Khodosovtsev A., Halici M. G. 2016. Three new, seemingly-cryptic species in the lichen genus Caloplaca (Teloschistaceae) distinguished in two-phase phenotype evaluation. Annales botanici Fennici 53(3-4): 243-262. https://doi.org/10.5735/085.053.0413
Khodosovtsev A., Kuznetzova E., Himelbrant D. 2004. Lichen genus Caloplaca on the Kamchatka Peninsula (Russian Far East). Botanica Lithuanica 10(3): 195-208.
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Kondratyuk S. Y., Elix J. A., Galanina I. A., Yakovchenko L. S., Karnefelt I., Thell A. 2011. Four new Caloplaca species (Teloschistaceae, Ascomycotina). Folia cryptogamica Estonica 48: 17-23.
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Kondratyuk S., Lakós L., Tschabanenko S., Skirina I., Galanina I., Oh S.-O., Hur J.-S. 2014. Calo-placa kedrovopadensis sp. nova and some new lichens from the Primorsky Region, Russia. Acta botanica Hungarica 56(1-2): 125-140. https://doi.org/10.1556/ABot.56.2014.1-2.11
Kondratyuk S. Y., Lakós L., Zarei-Darki B., Haji Moniri M., Tchabanenko S., Galanina I., Yakovchenko L., Hooshmand F., Ezhkin A., Hur J. 2013. Five new Caloplaca species (Teloschistaceae, Asco-mycota) from Asia. Acta botanica Hungarica 55(1-2): 41-60. https://doi.org/10.1556/ABot.55.2013.1-2.4
[Makarova, Katenin] Макарова И. И., Катенин А. Е. 2009. Лишайники природно-этнического парка «Берингия» (северо-восток Чукотского полуострова). Новости систематики низших растений 43: 172-189.
McCune B. 2017. Microlichens of the Pacific Northwest. Vol. 2. Key to the Species. Corvallis, Oregon: 755 p.
[Savicz] Савич В. П. 1914. Новые виды и формы лишайников Камчатки. Известия Император-скаго Ботанического сада Петра Великаго 14(1-6): 111-128.
Sochting U., Lorentsen L. B., Arup U. 2008. The lichen genus Caloplaca (Ascomycota, Lecanoromy-cetes) on Svalbard. Notes and additions. Nova Hedwigia 87(1-2): 69-96. https: //doi.org/10.1127/0029-5035/2008/0087-0069
Soun J., Vondrák J., Sochting U., Hrouzek P., Khodosovtsev A., Arup U. 2011. Taxonomy and phylo-geny of the Caloplaca cerina group in Europe. The Lichenologist 43(2): 113-135. https://doi.org/10.1017/S0024282910000721
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Vondrák J., Frolov I., Arup U., Khodosovtsev A. 2013b. Methods for phenotypic evaluation of crus-tose lichens with emphasis on Teloschistaceae. Chornomorskii botanichnii zhurnal 9(3): 382-405. https://doi.org/10.14255/2308-9628/13.93/6
Vondrák J., Frolov I., Davydov E., Urbanavichene I., Chesnokov S., Zhdanov I., Muchnik E., Kono-reva L., Himelbrant D., Tchabanenko S. 2016. The extensive geographical range of several species of Teloschistaceae: Evidence from Russia. The Lichenologist 48(3): 171-189. https://doi.org/10.1017/S0024282916000116
Vondrák J., Frolov I., Kosnar J., Arup U., Veselská T., Halici G., Malícek J., Sochting U. 2019. Substrate switches, phenotypic innovations and allopatric speciation formed taxonomic diversity within the lichen genus Blastenia. Journal of systematics and evolution. https: //doi.org/10.1111/j se. 12503
Vondrák J., Frolov I., Ríha P., Hrouzek P., Palice Z., Nadyenina O., Halici G., Khodosovtsev A., Roux C. 2013a. New crustose Teloschistaceae in Central Europe. The Lichenologist 45(6): 701-722. https://doi.org/10.1017/S0024282913000455
Vondrák J., Ríha P., Arup U., Sochting U. 2009. The taxonomy of the Caloplaca citrina group (Telo-schistaceae) in the Black Sea region; with contributions to the cryptic species concept in liche-nology. The Lichenologist 41(6): 571-604. https://doi.org/10.1017/S0024282909008317
Vondrák J., Soun J., Vondráková O., Fryday A., Khodosovtsev A., Davydov E. 2012. Absence of anthraquinone pigments is paraphyletic and a phylogenetically unreliable character in the Telo-schistaceae. The Lichenologist 44(3): 401-418. https://doi.org/10.1017/S0024282911000843
Wainio E. 1909. Lichenes in viciniis hibernœ expeditionis Vegœ prope pagum Pitlekai in Sibiria septentrionali a D-re E. Almquist collecti. Arkiv för Botanik 8(4): 1-175.
