Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 53(1): 107–142. 2019

Lichens — Лишайники

The first lichenological survey in Koryakia (Northern Kamchatka, ) — the last unexplored part of Beringia

D. E. Himelbrant1, 2, I. S. Stepanchikova1, 2, T. Ahti3, V. Yu. Neshataeva2

1St. Petersburg State University, St. Petersburg, Russia 2Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia 3Botanical Museum, Finnish Museum of Natural History, University of Helsinki, Finland Corresponding author: D. E. Himelbrant, [email protected]

Abstract. The first lichenological inventory in Koryakia has resulted in the list of 315 species reported from Parapolsky Dale, within and in vicinities of the Koryak State Reserve. Altogether 46 species are published from the Kamchatka Territory for the first time, includingLecanographa grumulosa new to Russia, East Asia and Beringia; Cercidospora trypetheliza, Lecania dubitans, Per- tusaria borealis, Piccolia ochrophora, Protoparmelia cupreobadia, Rimularia badioatra and Stran- gospora moriformis new to Russian Far East; Abrothallus bertianus, Cladonia strepsilis, Physciella melanchra, Rimularia badioatra, Sclerococcum parasiticum, Sphinctrina leucopoda and Strangospora moriformis new to Beringia. The lichen diversity of the study area is relatively poor due to natural reasons. Comparison with neighboring regions (, Chukotka, Magadan Region, Yakutia and Alaska) shows that the lichen flora of Parapolsky Dale contains almost no specific spe- cies. The majority of the species recorded here are also known from neighboring regions, especially Alaska and Kamchatka Peninsula. Keywords: lichen flora, new records, Beringia, Kamchatka, Koryak State Reserve, Koryakia.

Первые сведения о лихенофлоре Корякии (Северная Камчатка, Россия) — последней неизученной территории Берингии

Д. Е. Гимельбрант1, 2, И. С. Степанчикова1, 2, Т. Ахти3, В. Ю. Нешатаева2

1Санкт-Петербургский государственный университет, Санкт-Петербург, Россия 2Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия 3Ботанический музей, Финский музей естественной истории, университет Хельсинки, Финляндия Автор для переписки: Д. Е. Гимельбрант, [email protected]

Резюме. Результатом первых лихенологических исследований в Корякии стал список из 315 видов, обнаруженных на территории Парапольского Дола, в границах и в окрестностях Корякского государственного заповедника. Всего 46 видов приведены впервые для Камчатского края, в том числе Lecanographa grumulosa — вид, новый для России, Восточной Азии и Берингийского экорегиона; Cercidospora trypetheliza, Lecania dubitans, Pertusaria borealis, Piccolia ochrophora, Protoparmelia cupreobadia, Rimularia badioatra и Strangospora morifor­mis — https://doi.org/10.31111/nsnr/2019.53.1.107 107 Himelbrant et al. The first lichenological survey in Koryakia

виды, новые для Российского Дальнего Востока; Abrothallus bertianus, Cladonia, Physciella mel- anchra, Rimularia badioatra, Sclerococcum parasiticum, Sphinctrina leucopoda и Strangospora mori- formis — виды, новые для Берингии. Лихенофлора района исследований относительно бедна по естественным причинам. Сравнение с лихенофлорами ближайших регионов (полуостров Камчатка, Чукотка, Магаданская обл., Якутия и Аляска) показало, что лихенофлора Параполь- ского Дола практически не содержит каких-либо специфичных для нее видов. Подавляющее большинство выявленных видов известно с территории Аляски и полуостровной части Камчатки. Ключевые слова: лихенофлора, новые находки, Берингия, Камчатка, Корякский государ- ственный заповедник, Корякия. Introduction The geographical affinities between the lichen diversities of the Pacific regions of Northern Asia and North America are clearly predictable due to well-known palaeo- geographical processes in the Beringian Ecoregion (see Brigham-Grette, 2004; Cook et al., 2005; Waltari et al., 2007; Krasnov et al., 2015) and long-distance transporta- tion of diaspores, but still insufficiently investigated. One of the most important rea- sons is poor knowledge about lichen diversity of the Russian regions in North Pacific.

Fig. 1. Map of the study area. An investigated area is surrounded by circle. 108 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 53(1): 107–142. 2019

­Koryakia is one of the last lichenologically fully unexplored large territories in Russia, as well as in Beringia. Koryakia (formerly Koryak Autonomous Okrug, now part of Kamchatka Territo- ry) occupies the northernmost part of the Kamchatka Peninsula and adjacent mainland area. The Koryak State Reserve is the only reserve in Koryakia; it consists of three sepa- rate parts: “Mys Govena” (S part of the Goven Peninsula), “Bukhta Lavrova” (Lavrova Bay), and “Parapolsky dol” (Parapolsky Dale). The total area of its terrestrial part is about 3188 km2 (Kronotsky..., 2018). The investigated part Parapolsky Dale (1764 km2) lies in the north-western part of Koryakia in a tectonic depression between the Penzhin- sky Range and Koryakskoe Highland (Fig. 1). Parapolsky Dale is biogeographically one of the key parts of Kamchatka and represents a natural landscape barrier against the mainland in the narrowest part of the peninsula. Numerous animals have here the limits of their ranges, including different insects (see Mutin, 2014), mammals like moose and other forest species. The main type of landscape of Parapolsky Dale is swampy lowland (elevation mainly from 50 to 100 m a. s. l.) with a dense river net and thousands of lakes of various sizes. The largest of them is Lake Talovskoe (44 km2) (Gusakov, 1998). Al- though Parapolsky Dale is situated within the hypoarctic zone, shrub communities com- posed of Pinus pumila (Pallas) Regel or Betula divaricata Ledeb. (= Betula middendorffii Trautv. et Mey.), or more rarely Alnus fruticosa Rupr. [= Duschekia fruticosa (Rupr.) Pouzar; Alnus alnobetula (Ehrh.) K. Koch subsp. fruticosa (Rupr.) Raus], are distributed only on slopes of the depression and small local hills. Same positions are also occupied by lichen- and moss-dwarf shrub tundras with Betula exilis Sukacz. [= Betula nana L. subsp. exilis (Sukaczev) Hultén], Empetrum nigrum L., Ledum decumbens (Ait.) Lodd. ex Steud. (= Rhododendron decumbens D. Don ex G. Don), Vaccinium uliginosum L., and V. vitis-idaea L. The most part of the lowland is covered by swamps with Eriophorum vaginatum L. and Carex spp., shores of swamps and lakes are overgrown by Salix spp. communities. Forest communities are represented only by floodplain forests withSalix arbutifolia Pall. [= Chosenia arbutifolia (Pall.) A. Skvorts.], Populus suaveolens Fisch., Salix schwerinii E. Wolf, and S. udensis Trautv. et Mey. along the banks of main rivers (Neshataeva et al., 2017). Special lichenological studies in Parapolsky Dale, as well as in the whole Koryakia, have not been conducted ever before. Material and methods The specimens were collected by Dmitry E. Himelbrant (abbreviated as DH) and Valentina Yu. Neshataeva in August 2016 in Parapolsky Dale (within the Koryak State Reserve and its vicinities), Penzhinsky District of Kamchatka Territory. Alto- gether 42 localities were investigated: 35 standard sample plots (20 × 20 m in forest communities and 10 × 10 m in shrub communities, tundras or swamps, otherwise in natural boundaries of the community), where the lichen diversity on each substrate was described as detailed as possible, and 7 additional plots, where only individual substrates and species were recorded. All geographical coordinates are given in the coordinate system WGS 1984.

109 Himelbrant et al. The first lichenological survey in Koryakia

Chromatography was performed by Irina S. Stepanchikova according to the stand- ard techniques of high performance thin-layer chromatography using solvent systems A and B (Orange et al., 2001). The nomenclature of taxa generally follows recently published checklists for Scandinavia, Alps and North America (Nordin et al., 2011; Hafellner, Türk, 2016; Esslinger, 2018) and special taxonomic papers and monographs (Jahns, 1981; Dombrovskaya, 1996; Fryday, 2002; Halonen et al., 2009; Kukwa, 2011; Westberg, Arup, 2011; Spribille et al., 2014; Davydov, 2017; Sheard et al., 2017; ­Zhdanov, 2017). The specimens were mainly identified by DH, Irina Stepanchikova, and Teuvo Ahti (Cladonia spp.), if otherwise, the identifier’s names are indicated in the species list. Representative specimens are deposited in the herbaria of the Botanical Museum, University of Helsinki (H), Komarov Botanical Institute RAS (LE), and Department of Botany, St. Petersburg State University (LECB). Information on the presence of species in neighboring regions is mostly based on published records, otherwise sources are cited in the species list. Literature used for distribution analysis is cited below. Kamchatka Territory: 1 (Nylander, 1887), 2 (Trass, 1963), 3 (Mikulin, 1986), 4 (Mikulin, 1987), 5 (Mikulin, 1988), 6 (Dobrysh, 1993), 7 (Dombrovskaya, 1996), 8 (Neshataeva et al., 2003), 9 (Neshataeva et al., 2004), 10 (Khodosovtsev et al., 2004), 11 (Titov et al., 2004), 12 (Neshataeva et al., 2005), 13 (Kuznetsova, Himelbrant, 2006), 14 (Davydov et al., 2011), 15 (Zhurben- ko et al., 2012), 16 (Himelbrant et al., 2014), 17 (Spribille et al., 2014), 18 (Galani- na et al., 2017), 19 (Sheard et al., 2017). Chukotka Autonomous Area: 20 (Jahns, 1981), 21 (Andreev et al., 1996), 22 (Galanina, Galanin, 1999), 23 (Fryday, 2002), 24 (Urbanavichus, Urbanavichene, 2005), 25 (Makarova, Katenin, 2009), 26 (Zhur- benko, 2009b), 18 (Galanina et al., 2017), 27 (Zhurbenko, Pino-Bodas, 2017). Maga- dan Region: 28 (Andreev, 1978), 29 (Andreev, 1984), 30 (Korolev, Tolpysheva, 1980), 31 (Kotlov, 1993), 32 (Søchting, 1994), 33 (Kotlov, 1995), 34 (Makryi, Zheludeva, 2012), 35 (Zheludeva, 2014), 36 (Zheludeva, 2015), 37 (Zhurbenko, Zheludeva, 2015), 38 (Davydov, 2017), 18 (Galanina et al., 2017), 39 (Zheludeva, 2017). Repub- lic of Sakha (Yakutia): 40 (Poryadina, 2005), 41 (Zhurbenko et al., 2005b), 42 (Zhur- benko, 2009a), 26 (Zhurbenko, 2009b), 43 (Vershinina et al., 2012), 44 (Miadlikow­ ska et al., 2014), 45 (Chesnokov et al., 2015), 46 (Galanina, 2015), 47 (Vershinina et al., 2015), 48 (Chesnokov et al., 2016), 49 (Galanina, 2016), 50 (Chesnokov et al., 2017), 18 (Galanina et al., 2017), 51 (Zhdanov, 2017), 27 (Zhurbenko, Pino-Bodas, 2017). Alaska: 52 (Nylander, 1885), 1 (Nylander, 1887), 53 (Hadwen, Palmer, 1922), 54 (Magnusson, 1929), 55 (Thomson, 1979), 56 (Thomson, 1984), 57 (Gowan, 1989), 58 (Talbot et al., 1992), 32 (Søchting, 1994), 59 (Thomson, Ahti, 1994), 60 (Talbot et al., 1997), 61 (Thomson, 1997), 62 (Geiser et al., 1998), 63 (Brodo et al., 2001), 64 (Talbot et al., 2001), 65 (Mayrhofer, Sheard, 2002), 66 (Ahti, 2007), 67 (Talbot et al., 2007), 68 (McCune et al., 2009), 42 (Zhurbenko, 2009a), 69 (Esslinger, Dillman, 2010), 70 (Spribille et al., 2010), 71 (Westberg, Arup, 2011), 17 (Spribille et al., 2014), 72 (Dillman et al., 2012), 73 (Stehn et al., 2015), 74 (McCune et al., 2018).

110 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 53(1): 107–142. 2019

Sampling locations Standard sample areas. Koryak Nature Reserve, Parapolsky Dale: 1 — NE shore of Lake Talovskoe, 61°20'31.1"N, 164°47'00.8"E, 52 m a. s. l., Pinus pumila and Betula divaricata shrubs with Sphagnum sp., 09 VIII 2016, DH Tal-1; 2 — same place, 61°20'34.3"N, 164°46'59.8"E, 49 m a. s. l., Salix pulchra Cham. shrubs with Rubus arcticus Trautv., 09 VIII 2016, DH Tal-2; 3 — E to Lake Talovskoe, W slope of hill chain, 61°20'20.6"N, 164°48'38.1"E, 127 m a. s. l., Pinus pumila shrubs with Rubus cha- maemorus L., Vaccinium vitis-idaea and Ledum decumbens, 09 VIII 2016, DH Tal-3; 4 — same place, W slope of hill chain, 61°20'19.4"N, 164°48'56.9"E, 135 m a. s. l., Carex sp.-lichen-dwarf shrub tundra with Alnus fruticosa shrubs, 09 VIII 2016, DH Tal-4; 5 — same place, top of the hill in hill chain, 61°18'16.9"N, 164°45'55.0"E, 197 m a. s. l., open rocky outcrops with lichens, 10 VIII 2016, DH Tal-5; 6 — N shore of Lake Talovskoe, 61°21'46.0"N, 164°49'03.9"E, 74 m a. s. l., Salix pulchra shrubs with Calamagrostis purpurea (Trin.) Trin., 11 VIII 2016, DH Tal-6; 7 — NE to Lake Talovskoe, top of small hill in hill chain, 61°21'02.9"N, 164°50'11.2"E, 180 m a. s. l., small open rocky outcrops with lichens, 12 VIII 2016, DH Tal-7; 8 — same place, 61°21'07.6"N, 164°50'32.3"E, 211 m a. s. l., stone tundra with lichens, 12 VIII 2016, DH Tal-8; 9 — NE shore of Lake Talovskoe, 61°20'38.3"N, 164°47'57.5"E, 48 m a. s. l., Betula divaricata shrubs with Ledum decumbens and Vaccinium uliginosum, 12 VIII 2016, DH Tal-9; 10 — same place, 61°20'34.5"N, 164°47'05.6"E, 50 m a. s. l., Rubus chamaemorus-Sphagnum spp. bog with Ledum decumbens and lichens, 13 VIII 2016, DH Tal-10; 11 — same place, 61°20'32.8"N, 164°47'07.0"E, 50 m a. s. l., Eriophorum vaginatum bog, 13 VIII 2016, DH Tal-11; 12 — same place, 61°20'26.7"N, 164°47'16.6"E, 57 m a. s. l., Eriophorum vaginatum bog, 13 VIII 2016, DH Tal-12; 13 — same place, 61°20'34.5"N, 164°48'09.6"E, 55 m a. s. l., tundra with Eriophorum vaginatum, lichens, and scattered Alnus fruticosa shrubs, 13 VIII 2016, DH Tal-13; 14 — NE to Lake Talovskoe, base of the hill in hill chain, 61°20'37.4"N, 164°48'22.5"E, 57 m a. s. l., old Pinus pumila shrubs with Rho- dodendron aureum Georgi and mosses, 14 VIII 2016, DH Tal-14; 15 — same place, 61°20'37.4"N, 164°48'23.0"E, 55 m a. s. l., old Alnus fruticosa shrubs with Spiraea beauverdiana Schneid., 14 VIII 2016, DH Tal-15; 16 — NE to Lake Talovskoe, bank of small river, 61°20'40.2"N, 164°48'03.6"E, 46 m a. s. l., Salix pulchra shrubs with Rubus arcticus, 14 VIII 2016, DH Tal-16; 17 — NE to Lake Talovskoe, base of the hill in hill chain, 61°20'37.0"N, 164°48'21.0"E, 54 m a. s. l., Eriophorum vaginatum bog with Carex spp. and lichens, 14 VIII 2016, DH Tal-17; 18 — left bank of Ichiginynvayam, fluvial valley, 61°25'04.7"N, 165°01'24.8"E, 55 m a. s. l., Eriophorum vaginatum-Sphagnum spp. bog, 18 VIII 2016, DH Ichig-22; 19 — same place, 61°25'05.7"N, 165°01'24.6"E, 56 m a. s. l., lichen tundra, 18 VIII 2016, DH Ichig-23; 20 — right bank of Ichiginynvayam, fluvial valley, 61°24'40.8"N, 165°01'53.5"E, 70 m a. s. l., Salix arbutifolia floodplain forest with Populus suaveolens, Salix udensis, Alnus fruticosa and Calamagrostis purpurea, 18 VIII 2016, DH Ichig-24; 21 — left bank of Ichiginynvayam, fluvial valley, 61°25'25.0"N, 165°01'54.4"E, 69 m a. s. l., Carex spp.-Sphagnum spp. bog, 19 VIII 2016, DH Ichig-25; 22 — same place, 61°25'30.0"N, 165°01'36.8"E, 61 m a. s. l., Betula divaricata shrubs with lichens, 19 VIII 2016, DH Ichig-26; 23 — same place, 61°25'29.0"N, 165°01'43.7"E, 61 m a. s. l., Carex spp.-Sphagnum spp. bog, 19 VIII 2016, DH Ichig-27; 24 — same place, 61°25'47.3"N, 165°02'01.7"E, 64 m a. s. l., Carex spp.-dwarf shrub-lichen tundra, 20 VIII 2016, DH Ichig-28; 25 — same place, 61°25'46.5"N, 165°02'02.8"E, 64 m a. s. l., Eriophorum vaginatum-Carex spp.-Sphagnum spp. bog, 20 VIII 2016, DH Ichig-29; 26 — same place, 61°25'40.0"N, 165°02'34.3"E, 64 m a. s. l., Populus sua- veolens floodplain forest with Alnus fruticosa and Calamagrostis purpurea, 20 VIII 2016, DH Ichig-30; 27 — same place, 61°25'40.8"N, 165°02'33.0"E, 60 m a. s. l., Alnus fruticosa shrubs with Calama- grostis purpurea, 20 VIII 2016, DH Ichig-31; 28 — left bank of Katalyanajvayam, floodplain terrace, 61°24'34.3"N, 165°02'02.0"E, 66 m a. s. l., dwarf shrub-moss tundra, 21 VIII 2016, DH Ichig-32; 29 — same place, 61°24'30.0"N, 165°01'56.6"E, 71 m a. s. l., Eriophorum vaginatum-Carex spp. bog, 21 VIII 2016, DH Ichig-33; 30 — left bank of Katalyanajvayam, fluvial valley, 61°24'39.7"N, 165°02'02.9"E, 61 m a. s. l., Salix arbutifolia floodplain forest with Populus suaveolens, Salix schwerinii, Alnus fruti- cosa and Calamagrostis purpurea, 21 VIII 2016, DH Ichig-34; 31 — fluvial valley between Ichiginyn- vayam and Kujul rivers, near unnamed lake, 61°28'07.2"N, 165°02'01.7"E, 56 m a. s. l., lichen tundra

111 Himelbrant et al. The first lichenological survey in Koryakia with Arctous alpina (L.) Niedenzu and Eriophorum vaginatum, 22 VIII 2016, DH Ichig-35. Vicinity of Koryak Nature Reserve, Ametistovoe gold mines: 32 — E of Ametistovoe gold mines, base of the hill in hill chain, near the road, 61°18'06.5"N, 165°00'57.6"E, 144 m, small stone-birch forest with Spiraea beauverdiana and Calamagrostis purpurea on the hill slope, 16 VIII 2016, DH Am-18; 33 — same place, 61°18'14.0"N, 165°01'08.4"E, 301 m, stone tundra (kurum), 16 VIII 2016, DH Am-19; 34 — N of Ametistovoe gold mines, wide plain fluvial valley of Ichiginynvayam, right bank of Ichig- inynvayam, 61°19'14.1"N, 165°55'30.2"E, 109 m, graminoid lichen tundra, 16 VIII 2016, DH Am-20; 35 — same place, 61°19'15.2"N, 165°55'24.6"E, 101 m, Salix pulchra shrubs with Rubus arcticus and Calamagrostis purpurea on the river bank, 16 VIII 2016, DH Am-21.

Additional collection points. Koryak Nature Reserve, Parapolsky Dale: a1 — E to Lake Talovskoe, W slope of hill chain, 61°20'28.8"N, 164°49'28.0"E, 177 m a. s. l., stone tundra with lichens, 09 VIII 2016, DH Tal-1ad; a2 — E to Lake Talovskoe, top of the hill in hill chain, 61°18'24.6"N, 164°46'12.4"E, 213 m a. s. l., small open rocky outcrops, 10 VIII 2016, DH ­Tal-2ad; a3 — NE to Lake Talovskoe, top of the hill in hill chain, 61°21'03.7"N, 164°50'33.0"E, 223 m a. s. l., open rocky outcrops, 12 VIII 2016, DH Tal-3ad; a4 — same place, 61°21'14.8"N, 164°49'38.1"E, 75 m a. s. l., Alnus fruticosa shrubs, 12 VIII 2016, DH Tal-4ad; a5 — left bank of Ichiginynvayam, fluvial valley, 61°25'42.9"N, 165°02'23.9"E, 60 m a. s. l., Betula divaricata shrubs with lichens, 20 VIII 2016, DH Ichig-5ad; a6 — fluvial valley between Ichiginynvayam and Kujul rivers, near unnamed lake, 61°27'58.0"N, 165°01'30.4"E, 55 m a. s. l., Carex spp.-Sphagnum spp. bog with lichens, 22 VIII 2016, DH Ichig-6ad; a7 — vicinity of Lake Talovskoe, hill chain, [61°21'N, 164°51'E], ca. 200 m a. s. l., tundra with lichens, 08.2016, Neshataeva 1627.

Results and discussion An annotated list of species of Parapolsky Dale and vicinities is presented be- low. The species reported for the first time for the Kamchatka Territory are marked with !, lichenicolous fungi with #, non-lichenized fungi with +. Each species is ac- companied with the list of substrates and localities. Lichen substances are given for TLC-analyzed specimens. The frequency of occurrence within the study area was estimated and is indicated in square brackets: [R1] “rare — single record” — the spe- cies was found once, [R] “rare” — recorded in 2–7 standard sample areas or more than in one additional plot, [O] “occasionally”— 8–14, [F] “frequent”— 15–21, [C] “common”— 22–28, [VC] “very common”— 29–35 standard sample areas (the latter category is absent in the analyzed material). For the species known from the Kam- chatka Territory and/or neighboring regions, abbreviations of the regions are listed: Ala — Alaska, Chu — Chukotka Autonomous Area, Kam — Kamchatka Territory [within the limits of Kamchatka Peninsula and Commander (Komandorskie) Is- lands], Mag — Magadan Region, Yak — Republic of Sakha (Yakutia). References are given after each abbreviation (in brackets), one for each region; numbers correspond to those in the list of publications (see Material and methods). Special comments are given when needed.

!#Abrothallus bertianus De Not. — on thallus of Melanohalea olivacea on bark of Alnus fruticosa; 27 [R1]; LE. — New to Beringia and northern part of the Russian Far East. The nearest known locality is Amur River Basin within the Khabarovsk Territory (Zhurbenko, Tugi, 2013).

112 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 53(1): 107–142. 2019

#A. parmeliarum (Sommerf.) Arnold — on thallus of Parmelia omphalodes subsp. omphal- odes on rocks; 5 [R1]; LE. — Kam (15), Mag (37), Yak (45), Ala (70). Acarospora fuscata (Schrad.) Th. Fr. — on rocks; 7, a2 [R]; LE. — Kam (16), Chu (21), Yak (40), Ala (61).

!A. molybdina (Wahlenb.) A. Massal. — on rocks; 5 [R1]; LE. — Chu (21), Yak (40), Ala (54).

!Adelolecia pilati (Hepp) Hertel et Hafellner — on rocks; 5 [R1]; H. — Chu (21), Yak (40), Ala (70). Alectoria ochroleuca (Hoffm.) A. Massal. — on soil and siliceous rocks; 4, 7, 8, 17, 19, 24, 31, 33, 34, a1 [O]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). !Amandinea cacuminum (Th. Fr.) H. Mayrhofer et Sheard — on rocks; 5, 7, a2 [R]; H, LE. — Chu (21), Yak (40), Ala (65). A. punctata (Hoffm.) Coppins et Sheid. — on bark of Betula divaricata, Alnus fruticosa, Pi- nus pumila and lignum; 3, 7, 15, 22, a1, a7 [R]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (73). Amygdalaria panaeola (Ach.) Hertel et Brodo — on rocks; 7, 8 [R]; H. — Kam (1), Chu (21), Mag (33), Yak (40), Ala (70).

A. subdissentiens (Nyl.) Mas. Inoue et Brodo — on rocks; 8 [R1]; H. — Kam (6), Chu (21), Ala (59). Arctocetraria andrejevii (Oxner) Kärnefelt et A. Thell — on soil; 24, 34 [R]; LE. — Kam (16), Chu (21), Mag (28), Yak (40), Ala (64). Arctoparmelia incurva (Pers.) Hale — on rocks; 8, 33, a1, a3 [R]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). A. separata (Th. Fr.) Hale — on rocks and primary soil; 7, 8, 33, a1, a3 [R]; LE (sub Cetraria nigricans). — Kam (16), Chu (21), Mag (30), Yak (40), Ala (60).

!#Arthonia digitatae Hafellner — on thallus of Cladonia cf. macilenta on soil; 21 [R1]; LE. — Chu (27), Yak (27), Ala (27).

+A. punctiformis Ach. — on bark of Salix pulchra; 6 [R1]; LE. — Kam (16), Ala (62). A. radiata (Pers.) Ach. — on bark of Alnus fruticosa, Salix pulchra, S. udensis; 6, 20 [R]; LE. — Kam (12), Yak (47), Ala (70). +Arthopyrenia analepta (Ach.) A. Massal. — on bark of Alnus fruticosa, Pinus pumila, Salix pulchra; 1, 3, 16, 32, 35, a4, a7 [R]; LE. — Kam (16), Chu (25), Ala (70).

!Arthrorhaphis alpina (Schaer.) R. Sant. — on primary soil on rocks; 8 [R1]; LE. — Chu (21), Ala (70).

A. citrinella (Ach.) Poelt — on soil; a6 [R1]; H. — Kam (16), Chu (25), Yak (40), Ala (70). Asahinea chrysantha (Tuck.) W. L. Culb. et C. F. Culb. — on soil and siliceous rocks; 5, 7, 8, 18, 19, 24, 25, 31, 33 [O]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (60). A. scholanderi (Llano) W. L. Culb. et C. F. Culb. — on rocks; 5, 8, 33 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). Aspicilia cinerea (L.) Körb. — on rocks; 5, 7 [R]; H. — Kam (13), Chu (21), Mag (33), Yak (40), Ala (70).

!Athallia holocarpa (Hoffm.) Arup, Frödén et Søchting — on rocks; a2 [R1]; LE. — Chu (21), Yak (40), Ala (70).

A. pyracea (Ach.) Arup, Frödén et Søchting — on bark of Populus suaveolens; 20 [R1]; LE (det. Ivan Frolov). — Kam (10), Yak (50), Ala (1). Baeomyces carneus Flörke — on soil; 8, 12, 19, 34 [R]; LE (sub Cetraria nigricans). Thalli contain norstictic and stictic acids. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (60).

113 Himelbrant et al. The first lichenological survey in Koryakia

B. placophyllus Ach. — on soil; 5, 8, 34 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

B. rufus (Huds.) Rebent. — on soil; 8 [R1]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). !Biatora albidula Willey — on bark of Alnus fruticosa, Populus suaveolens, Salix arbutifolia, S. udensis, Sorbus sibirica; 20, 26, 27, 30 [R]; LE. — Kam [Kamchatka Territory, Milkovo Dis- trict, , Levaya Schapina River basin, right bank of the Ipuin River, 55°05'30"N, 160°00'31"E, 340 m a. s. l., Picea ajanensis Fisch. ex Trautv. et C. A. Mey forest with mosses, on bark of Sorbus sambucifolia (Cham. et Schltdl.) M. Roem., 07 VIII 2009, Himelbrant, Stepanchikova K-07-09, H], Ala (70). B. efflorescens (Hedl.) Räsänen — on bark of Salix pulchra, S. schwerinii; 16, 30 [R]; LE. — Kam (16), Ala (70). B. pallens (Kullh.) Printzen — on bark of Betula divaricata, Alnus fruticosa, Pinus pumila, Salix pulchra and lignum; 1–3, 9, 14, 16, 20, a4, a7 [R]. — Kam (16), Yak (47), Ala (70).

B. sphaeroidiza (Vain.) Printzen et Holien — on bark of Betula divaricata; 3 [R1]. — Kam (9), Chu (21), Ala (72). !Blastenia ammiospila (Wahlenb.) Arup, Søchting et Frödén — on bark of Salix pulchra; 2, 6, 16 [R]; LE (det. Ivan Frolov). — Chu (21), Yak (40), Ala (70).

B. hungarica (H. Magn.) Arup, Søchting et Frödén — on bark of Salix arbutifolia; 20 [R1]; LE (det. Ivan Frolov). — Kam (16). Bryobilimbia hypnorum (Lib.) Fryday, Printzen et S. Ekman — on mosses; 2 [R1]; H. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (68). Bryocaulon divergens (Ach.) Kärnefelt — on soil and siliceous rocks; 4, 5, 8, 19, 24, 31, 34, a2 [R]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (60). Bryoria nitidula (Th. Fr.) Brodo et D. Hawksw. — on rocks, soil and plant debris; 5, 7, 8, 19, 24, 31, 33, 34 [O]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). B. simplicior (Vain.) Brodo et D. Hawksw. — on bark of Alnus fruticosa, Populus suaveolens; 26, 27 [R]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). Buellia disciformis (Fr.) Mudd — on bark of Alnus fruticosa, Populus suaveolens, Salix arbu- tifolia, S. udensis; 20, 26, 30, 32 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). B. ectolechioides (Vain.) Erichsen — on rocks; 7, 19 [R]; H (sub Ionaspis obtecta and Mela- nohalea infumata). — Kam (16), Chu (21), Mag (33). B. erubescens Arnold — on bark of Salix pulchra; 16, 35 [R]; LE (sub Rinodina septentrio­ nalis). — Kam (16), Ala (70).

Caloplaca ahtii Søchting — on bark of Populus suaveolens; 26 [R1]; LE (det. Ivan Frolov). — Kam (16), Mag (32), Yak (40), Ala (32). C. borealis (Vain.) Poelt — on bark of Betula ermanii, Alnus fruticosa, Populus suaveolens, Salix arbutifolia, S. pulchra; 2, 6, 16, 20, 30, 32 [R]; LE (det. Ivan Frolov). — Kam (16), Yak (40), Ala (70). C. cerina (Hedw.) Th. Fr. — on bark of Populus suaveolens, Salix pulchra; 20, 35 [R]; LE. — Kam (10), Chu (21), Yak (40), Ala (73).

C. stillicidiorum (Vahl) Lynge — on reindeer horn; 8 [R1]; LECB. — Kam (1), Ala (61).

Candelaria pacifica M. Westb. et Arup — on mosses; 5 [R1]; LE. — Kam (UPS), Ala (71).

!Candelariella arctica (Körb.) R. Sant. — on rocks; 5 [R1]; LE. — Chu (21), Ala (64). !C. coralliza (Nyl.) H. Magn. — on rocks; 5, 7 [R]; LE. — Kam (Kamchatka Territory, Elizovo District, Vachkazhets volcano, 53°03'41.7"N, 157°56'49.8"E, 804 m a. s. l., tundra with big stones, on siliceous stone, 21 VIII 2018, Himelbrant, Stepanchikova K-Vachk-6a, LECB), Ala (74).

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C. efflorescens R. C. Harris et W. R. Buck — on bark of Alnus fruticosa, Populus suaveolens, Salix arbutifolia, Sorbus sibirica; 20, 26, 27 [R]; LE. — Kam (16), Ala (70). C. lutella (Vain.) Räsänen — on bark of Salix arbutifolia, S. pulchra; 2, 6, 16, 20 [R]; LE. — Kam (16), Mag (33), Yak (40), Ala (61). C. vitellina (Hoffm.) Müll. Arg. — on rocks; 5, 7 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

C. xanthostigma (Ach.) Lettau — on bark of Salix arbutifolia; 20 [R1]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (64). !Catillaria erysiboides (Nyl.) Th. Fr. — on lignum of Salix udensis; 20, a1 [R]; LE. — Kam (Kamchatka Territory, Milkovo District, Kronotsky Nature Reserve, Levaya Schapina River basin, right bank of the Ipuin River, 55°05'18"N, 160°00'45"E, 350 m a. s. l., spruce forest with mosses, on wood of Picea ajanensis, 07 VIII 2009, Himelbrant, Stepanchikova K-05-09, H), Ala (70).

Catinaria atropurpurea (Schaer.) Vězda et Poelt — on bark of Salix arbutifolia; 30 [R1]; LE. — Kam (16), Yak (50), Ala (70). !#Cercidospora trypetheliza (Nyl.) Hafellner et Obermayer — on thallus of Arthrorhaphis alpina on primary soil; 8 [R1]; LE. — Yak (42). New to Russian Far East. Cetraria islandica (L.) Ach. subsp. islandica — on soil; 22, 31 [R]; H, LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. kamczatica Savicz — on soil; 31 [R1]; H. — Kam (16), Chu (21), Mag (34), Yak (26), Ala (60). C. laevigata Rassad. — on soil; 4, 5, 7, 8, 10, 12, 13, 17–19, 21, 23–25, 34 [F]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. nigricans Nyl. — on soil and siliceous rocks; 8, a6 [R]; H, LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. sepincola (Ehrh.) Ach. — on bark of Betula ermanii, B. exilis, B. divaricata, Alnus frutico- sa, Pinus pumila, Salix pulchra, Vaccinium uliginosum; 1, 3, 4, 6, 7, 9, 10, 12, 14–16, 18, 21, 22, 27, 30, 32, 35, a4 [F]; LE. — Kam (16), Chu (21), Mag (30), Yak (47), Ala (70). Cetrariella delisei (Bory ex Schaer.) Kärnefelt et A. Thell — on soil; 24, 34 [R]. — Kam (16), Chu (25), Mag (30), Yak (40), Ala (70).

C. fastigiata (Delise ex Nyl.) Kärnefelt et A. Thell — on soil; 24 [R1]; H. — Kam (16), Chu (21), Yak (40), Ala (60). !Cetrelia alaskana (W. L. Culb. et C. F. Culb.) W. L. Culb. et C. F. Culb. — on soil; 5 [R1]; LE. — Chu (26), Yak (40), Ala (64). — Rare terricolous foliose lichen covered with white pseudocyphellae, well visible in the field (Plate I, 1).

Chaenotheca stemonea (Ach.) Müll. Arg. — on bark of Betula ermanii; 32 [R1]. — Kam (11), Yak (45), Ala (73). Chrysothrix chlorina (Ach.) J. R. Laundon — on rocks; 5, 7, 8 [R]. — Kam (3), Chu (21), Ala (70).

Cladonia acuminata (Ach.) Norrl. — on soil; 16 [R1]; LE. Thallus contains atranorin and trace of norstictic acid. — Kam (16), Chu (21), Mag (31), Yak (40), Ala (73).

!C. albonigra Brodo et Ahti — on plant debris; a7 [R1]; LE. Thallus contains fumarprotoce- traric and grayanic acids. — Ala (73). — New to the northern part of the Russian Far East. The nearest known locality is in Primorye Territory (H). C. amaurocraea (Flörke) Schaer. — on soil and bark of Pinus pumila; 1, 4, 5, 7–14, 17–19, 21–25, 28, 29, 31, 33, 34 [C]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70).

115 Himelbrant et al. The first lichenological survey in Koryakia

Plate I. 1 — Cetrelia alaskana (LE, Tal-5); 2 — Cladonia nitens (H, Ichig-28); 3a, 3b — Cladonia wainioi (H, Ichig-29); 4a, 4b — Lecanographa grumulosa (H, Tal-5). Scale bars: 1–3a, 3b — 1 cm, 4a, 4b — 1 mm.

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C. arbuscula (Wallr.) Flot. subsp. arbuscula (incl. subsp. beringiana Ahti) — on soil and bark of Salix pulchra; 1, 3–5, 7–13, 17–19, 21–25, 28, 29, 31, 33–35, a2 [C]; LE (sub Lepra panyrga). — Kam (16), Chu (21), Mag (30), Yak (40), Ala (60).

C. bacilliformis (Nyl.) Sarnth. — on lignum of Pinus pumila; a6 [R1]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (73).

C. bellidiflora (Ach.) Schaer. — on soil; a6 [R1]; H. — Kam (5), Chu (21), Mag (33), Yak (40), Ala (70). C. borealis S. Stenroos — on soil and bark of Salix pulchra; 5, 8, 10, 12, 18, 19, 21, 24, 34, 35, a7 [O]; LE. — Kam (16), Chu (H), Mag (H), Yak (40), Ala (70). C. carneola (Fr.) Fr. — on soil; 5, 19, 24 [R]. — Kam (16), Chu (21), Mag (30), Yak (H), Ala (70). C. cenotea (Ach.) Schaer. — on soil; 1, 4, 10–13, 18, 21, 24, 25, 28, 29 [O]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

C. chlorophaea (Flörke ex Sommerf.) Spreng. — on soil; 17 [R1]; LE. Thallus contains fu- marprotocetraric acid. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70).

C. coccifera (L.) Willd. — on soil; 8 [R1]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. coniocraea (Flörke) Spreng. — on bark of Betula ermanii, Salix arbutifolia, S. pulchra, Sorbus sibirica, soil; 11, 16, 20, 27, 32 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (64). C. cornuta (L.) Hoffm. — [F]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70): subsp. cornuta — on bark of Betula divaricata, Pinus pumila, Populus suaveolens, Salix arbu- tifolia, S. udensis, soil; 1, 3, 4, 8–12, 14, 20, 21, 24–26, 28–31; subsp. groenlandica (E. Dahl.) Ahti — on soil; 4, 13, 18. C. crispata (Ach.) Flot. var. crispata — on soil; 3, 4, 10, 31 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. cyanipes (Sommerf.) Nyl. — on soil; 9, 10, 12, 18, 21, 25, 28, 29, 31 [O]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). C. deformis (L.) Hoffm. — on soil; 3, 4, 11, 12, 18, 21, 24, 25, 28, 29, 31, 34 [O]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (73).

C. digitata (L.) Hoffm. — on soil; 4 [R1]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (73).

C. ecmocyna Leight. — on soil; a7 [R1]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. fimbriata (L.) Fr. — on bark of Betula divaricata and Salix arbutifolia, lignum and soil; 9, 21, 30 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (73). C. gracilis (L.) Willd. — [F]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70): subsp. turbinata (Ach.) Ahti — on soil; 29; subsp. vulnerata Ahti — on soil; 1, 4, 7–11, 13, 18, 21, 22, 24, 25, 28, 29, 31. C. kanewskii Oxner — on soil; 24, 34 [R]; H, LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

C. libifera Savicz — on soil; 18 [R1]; H. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (68). C. macilenta Hoffm. — on lignum of Pinus pumila and soil; 11, 18, 19, 21, 29, 31, a6 [R]; H. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. macroceras (Delise) Hav. — on soil; 4, 5, 7, 19 [R]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). C. macrophylla (Schaer.) Stenh. — on soil and bark of Salix pulchra; 5, 10, 16 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70).

117 Himelbrant et al. The first lichenological survey in Koryakia

C. macrophyllodes Nyl. — on soil; 5, 7 [R]. — Kam (6), Chu (21), Mag (30), Ala (70). C. maxima (Asahina) Ahti — on soil; 3, 4, 7, 9, 22, 28, 31 [R]. — Kam (16), Chu (21), Ala (70). C. nipponica Asahina — on soil; 8, 19, 24, a6 [R]; H, LE. — Kam (5), Chu (21), Mag (28), Yak (40), Ala (56). C. nitens Ahti — on soil; 24 [R1]; H. — Kam (UPS), Chu (LE), Mag (LE), Yak (LE), Ala (66). — The species has slender podetia with centrally proliferating narrow scyphi and contrast melanotic basal parts (Plate I, 2). C. phyllophora Hoffm. — on soil; 19, 28 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. pleurota (Flörke) Schaer. — on soil, bark of Betula ermanii and Pinus pumila; 5, 7, 8, 10, 12, 18, 21, 24, 25, 29, 32 [O]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. pyxidata (L.) Hoffm. — on soil, mosses and bark of Salix pulchra; 2, 5, 7–9, 16, 18, 19, 21, 24, 28, 29, 31, 34, 35 [F]; H, LE. Thalli contain fumarprotocetraric acid. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. rangiferina (L.) F. H. Wigg. — on soil and bark of Salix pulchra; 1, 3, 4, 7–11, 18, 19, 21, 22, 24, 25, 28, 29, 31, 35 [F]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. rei Schaer. — on soil; 9, 21 [R]. — Kam (16), Chu (21), Mag (33), Yak (49), Ala (59). C. scabriuscula (Delise) Nyl. — on soil and bark of Pinus pumila; 2, 9, 14 [R]; H. — Kam (16), Chu (21), Yak (40), Ala (70). C. squamosa Hoffm. — on soil; 4, 5, 7, 8, 10, 18, 19, 24, 25, 34 [O]. — Kam (16), Chu (21), Yak (40), Ala (73). C. stellaris (Opiz) Pouzar et Vězda — on soil; 4, 7, 8, 11, 12, 17, 19, 22, 24, 25, 31, 33, 34 [O]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. straminea (Sommerf.) Flörke — on soil; 7, 8, 12, 19, 21, 31, 34 [R]; H, LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (60).

!C. strepsilis (Ach.) Grognot — on soil; 34 [R1]; H. — New to Beringia and northern part of Russian Far East. The nearest known locality is Sikhote-Alin Range within Primorye Territory of Russia (Tchabanenko, 2002). C. stygia (Fr.) Ruoss — on soil; 1, 4, 7, 8, 10–13, 17–19, 21, 22, 24, 25, 28, 29, 31, 33, 34 [F]. — Kam (16), Mag (39), Yak (40), Ala (70). C. subfurcata (Nyl.) Arnold — on soil; 8, 9, 12, 18, 21, 24, 25, 29, 33 [O]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

C. subulata (L.) F. H. Wigg. — on soil; 31 [R1]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (73). C. sulphurina (Michx.) Fr. — on soil; 4, 11, 18, 25, 28, 29 [R]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

C. turgida Hoffm. — on soil; 18 [R1]. — Kam (16), Chu (21), Mag (30), Ala (56). C. uliginosa (Ahti) Ahti — on soil and bark of Salix pulchra; 18, 21, 34, 35 [R]. — Kam (16), Yak (40), Ala (68). C. uncialis (L.) F. H. Wigg. — on soil; 3–5, 7, 11, 18, 19, 21, 24, 25, 28, 29, 31, 34 [O]; H. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). C. verticillata (Hoffm.) Schaer. — on soil; 8, 19, 34 [R]. — Kam (16), Chu (25), Mag (30), Yak (40), Ala (73). C. wainioi Savicz — on soil; 19, 25, 31 [R]; H, LE. — Kam (2), Chu (22), Ala (70). — The species resembles C. arbuscula, but differs in having patchy cortex, brownish tips of podetia (Plate I, 3a and 3b) and K+ yellow medullary reaction.

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Cliostomum griffithii (Sm.) Coppins — on bark of Salix arbutifolia; 30 [R1]; LE (sub Toni­ niopsis subincompta). — Kam (9), Yak (49), Ala (73).

Collema furfuraceum (Arnold) Du Rietz — on bark of Populus suaveolens; 20 [R1]; LE. — Kam (16), Yak (40), Ala (70).

Dactylina arctica (Richardson) Nyl. — on soil; 18 [R1]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). Dibaeis baeomyces (L. f.) Rambold et Hertel — on soil; 12, a2, a6 [R]; H (sub Arthrorhaphis citrinella). — Kam (16), Chu (21), Mag (36), Yak (40), Ala (73). !Elixia flexella (Ach.) Lumbsch — on lignum; a1 [R1]; LE. — Ala (70). New to the northern part of the Russian Far East. The nearest known locality in Russia is Sikhote-Alin Range within Primorye Territory (Tchabanenko, 2002).

Euopsis granatina (Sommerf.) Nyl. — on rocks; 8 [R1]; H. — Kam (16), Chu (21), Mag (33), Ala (70).

Evernia mesomorpha Nyl. — on bark of Pinus pumila; 7 [R1]. — Kam (16), Chu (21), Mag (36), Yak (40), Ala (73). Flavocetraria cucullata (Bellardi) Kärnefelt et A. Thell — on soil; 4, 5, 7, 8, 10–13, 17–19, 21, 24, 25, 29, 31, 34 [F]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (60). F. minuscula (Elenkin et Savicz) Ahti, Poryadina et Zhurb. — on soil; 4, 8, 11, 17, 19, 21, 24, 25, 29, 31 [O]. — Kam (16), Mag (34), Yak (48), Ala (73). F. nivalis (L.) Kärnefelt et A. Thell — on soil; 4, 5–11, 17–19, 24, 25, 28, 29, 31, 33, 34 [F]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). Frutidella furfuracea (Anzi) M. Westb. et M. Svensson — on bark of Betula divaricata, Al- nus fruticosa, Pinus pumila; 3, 14, 15, 21, 22, 32 [R]; LE. Thalli contain sphaerophorin. — Kam (16), Ala (70). !Fuscopannaria praetermissa (Nyl.) P. M. Jørg. — on rocks, soil and bark of Salix pulchra; 2, 5 [R]; H, LE. — Kam (Kamchatka Territory, Milkovo District, SE of Kozyrevsk, Tolbachik volcano, slope of Kruglenkaya side cone, 55°43'23"N, 160°22'32"E, 1153 m a. s. l., tundra, on plant debris, 05 VIII 2008, Himelbrant, Stepanchikova K-09-08, LECB), Chu (21), Yak (40), Ala (70). Gowardia nigricans (Ach.) Halonen, Myllys, Velmala et Hyvärinen — on rocks, soil and bark of Pinus pumila; 5, 7, 19, 31, a3 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). Hypogymnia bitteri (Lynge) Ahti — on bark of Betula ermanii, Pinus pumila, Salix pulchra; 1, 16, 32 [R]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (73). H. physodes (L.) Nyl. — on bark of Betula divaricata, Pinus pumila, siliceous rocks; 1, 5, 7, 22 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). H. pseudophysodes (Asahina) Rassad. — on bark of Betula divaricata, Pinus pumila; 1, 3, 7, 14, 22 [R]; LE. — Kam (16). H. subobscura (Vain.) Poelt — on soil, plant debris, mosses; 5, 8, 19 [R]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (60).

H. tubulosa (Schaer.) Hav. — on bark of Pinus pumila; 3 [R1]. — Kam (9), Mag (39), Yak (40), Ala (70). Icmadophila ericetorum (L.) Zahlbr. — on mosses; 8, 10, 25, a2 [R]. — Kam (S), Chu (21), Mag (30), Yak (40), Ala (70). !Immersaria athroocarpa (Ach.) Rambold et Pietschm. — on rocks; 5, 7 [R]; H. — Chu (21), Mag (33), Yak (40), Ala (70).

Imshaugia aleurites (Ach.) S. L. F. Meyer — on bark of Pinus pumila; 7 [R1]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70).

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Ionaspis obtecta (Vain.) R. Sant. — on rocks; 7, 8, 19 [R]; H. — Kam (16). Japewia tornoёnsis (Nyl.) Tønsberg — on bark of Betula ermanii, B. exilis, B. divaricata, Alnus fruticosa, Pinus pumila and lignum; 1, 3, 7, 9, 10, 14, 15, 18, 20–22, 27, 32, a1 [O]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). !Lambiella insularis (Nyl.) T. Sprib. — on thallus of Lecanora bicincta on siliceous rock; a2

[R1]; LE. — Chu (26), Yak (26), Ala (72). — Lichenicolous lichen, forming dark brown islets on host thallus. Lasallia caroliniana (Tuck.) E. A. Davydov, Peršoh et Rambold — on rocks; 5, 7, 8, 33, a1 [R]; LE. — Kam (16), Mag (38), Yak (47), Ala (73). Lecania cyrtella (Ach.) Th. Fr. — on bark of Salix arbutifolia, Populus suaveolens; 20, 30 [R]; LE. — Kam (4), Yak (40). L. cyrtellina (Nyl.) Sandst. — on bark of Populus suaveolens, Salix arbutifolia, S. pulchra; 20, 30, 35 [R]; LE. — Kam (16), Yak (46).

!L. dubitans (Nyl.) A. L. Sm. — on bark of Salix arbutifolia; 20 [R1]; LE. — Ala (72). New to Russian Far East. The nearest published locality in Russia is Republic of Buryatia (Budaeva, 2012). !Lecanographa grumulosa (Dufour) Egea et Torrente — on thallus of Lecanora rupicola on siliceous rock; 5 [R1]; H. — New to Russia, Beringia and East Asia. Known from Europe, Africa (Hafellner, 2009), West Asia (Kinalioğlu, 2010) and North America (Esslinger, 2018). Facul- tatively lichenicolous lichen. Anatomical characteristics of the specimen are in accordance to the description of the species (see Smith et al., 2009); our specimen develops own thallus with Trentepohlia photobiont over the thallus of Lecanora rupicola (Plate I, 4a and 4b).

Lecanora albellula (Nyl.) Th. Fr. — on bark of Betula ermanii; 32 [R1]; LE. — Kam (16), Yak (47), Ala (58). !L. allophana Nyl. (incl. f. sorediata Nyl. ex Vain.) — on bark of Salix arbutifolia, Populus suaveolens; 20, 26 [R]. Thalli contain atranorin and terpenoids. — Chu (21), Yak (40), Ala (67). L. bicincta Ramond — on rocks; 5, 7, a2 [R]; LE. — Kam (16), Chu (21), Mag (39), Yak (40), Ala (70). L. boligera (Norman ex Th. Fr.) Hedl. — on bark of Betula divaricata, Alnus fruticosa, Pinus pumila, Salix pulchra and lignum; 1, 3, 9, 14–16, 20, 22, 35 [O]; LE. — Kam (16), Ala (70). L. cadubriae (A. Massal.) Hedl. — on bark of Alnus fruticosa; 15, 32 [R]. — Kam (16). L. chlarotera Nyl. — on bark of Betula ermanii, B. divaricata, Alnus fruticosa, Pinus pumila, Salix pulchra; 1, 3, 7, 14–16, 20, 22, 30, 32, a4, a7 [O]; LE. Thalli contain atranorin, gangaleoidin and roccellic acid. — Kam (16), Mag (31), Yak (45), Ala (73). L. circumborealis Brodo et Vitik. — on bark of Alnus fruticosa, Betula exilis, B. divaricata, Pinus pumila, Salix pulchra; 1, 3, 7, 9, 10, 15–18, 22, 27, 32, 35 [O]; LE. — Kam (16), Chu (21), Ala (70). L. fuscescens (Sommerf.) Nyl. — on bark of Betula divaricata, Alnus fruticosa, Pinus pumila, Populus suaveolens, Salix pulchra, Vaccinium uliginosum and lignum of Pinus pumila; 1, 3, 4, 7, 9, 10, 12, 14–17, 21, 22, 26, 32, 34, 35, a7 [F]; LE. — Kam (16), Chu (21), Yak (40), Ala (70). L. intricata (Ach.) Ach. — on rocks; 5, 7, 8, 33 [R]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). L. polytropa (Ehrh. ex Hoffm.) Rabenh. — on rocks; 5, 7, 8, 33, a2 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (60).

!L. rupicola (L.) Zahlbr. — on rocks; 5 [R1]; H (sub Lecanographa grumulosa). –– Chu (21), Mag (29), Yak (40), Ala (58). L. subintricata (Nyl.) Th. Fr. — on bark of Betula ermanii, B. divaricata, Pinus pumila, Po­ pulus suaveolens; 1, 20, 32 [R]. — Kam (16), Yak (47), Ala (61).

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L. symmicta (Ach.) Ach. — on bark of Betula ermanii, B. exilis, B. divaricata, Alnus fruticosa, Pinus pumila, Populus suaveolens, Salix arbutifolia, S. pulchra, S. schwerinii, S. udensis, Sorbus sibirica, Vaccinium uliginosum, lignum of Sorbus sibirica and plant debris; 1–6, 9, 10, 12, 13, 16, 18–21, 26, 27, 30, 32, 35, a4 [F]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

!Lecidea auriculata Th. Fr. subsp. auriculata — on rocks; 33 [R1]; H (sub Protoparmelia cupreobadia). — Chu (21), Mag (33), Yak (40), Ala (60). L. erythrophaea Flörke ex Sommerf. — on bark of Salix arbutifolia, Populus suaveolens, Sorbus sibirica; 20, 26, 27, 30 [R]; LE. — Kam (9), Mag (33), Yak (47), Ala (70). Lecidella elaeochroma (Ach.) M. Choisy — on bark of Alnus fruticosa, Populus suaveolens, Salix arbutifolia, S. schwerinii, S. udensis; 20, 26, 30 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (55). L. euphorea (Flörke) Hertel — on bark of Populus suaveolens, Salix pulchra, Sorbus sibirica and lignum; 16, 20, 26, 27 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (64). Lepra dactylina (Ach.) Hafellner — on rocks, soil, plant debris, mosses and bark of Betula divaricata; 7, 8, 18, 19, 21, 24, 25, 29, 33 [O]; H (sub Sclerococcum parasiticum). — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). !L. excludens (Nyl.) Hafellner — on rocks; 5, 8 [R]; H, LE. Thalli contain norstictic acid and unknown fatty acid. — Chu (21), Mag (33), Yak (40), Ala (59). L. ophthalmiza (Nyl.) Hafellner — on bark of Betula divaricata, Pinus pumila; 17, 22 [R]; LE. Thalli contain 4 fatty acids and unknown substances. — Kam (8), Ala (70). L. panyrga (Ach.) Hafellner — on soil, plant debris, bark of Betula exilis and B. divaricata; 5, 7, 8, 10, 17–19, 22, 24, 29, 34, a1, a2 [O]; LE. Thalli contain 4 fatty acids and unknown sub- stances. — Kam (16), Chu (21), Mag (33), Yak (48), Ala (60).

Lepraria neglecta (Nyl.) Erichsen — on soil; 5 [R1]; LE. Thallus contains alectorialic acid. — Kam (16), Chu (21), Yak (40), Ala (70).

Leptogium cyanescens (Rabenh.) Körb. — on bark of Salix pulchra; 2 [R1]; LE. — Kam (8), Yak (47), Ala (70). L. saturninum (Dicks.) Nyl. — on bark of Populus suaveolens; 20, 26 [R]; H, LE. — Kam (8), Chu (21), Mag (36), Yak (40), Ala (70).

+Leptorhaphis epidermidis (Ach.) Th. Fr. — on bark of Betula divaricata; 9 [R1]; LE. — Kam (9), Yak (40), Ala (74). Lichenomphalia umbellifera (L.: Fr.) Redhead, Lutzoni, Moncalvo et Vilgalys — on soil, plant debris and mosses; 18, 19, 21, 24, 32 [R]. — Kam (16), Yak (40), Ala (70).

Lobaria linita (Ach.) Rabenh. — on soil; 9 [R1]. — Kam (16), Chu (21), Mag (36), Yak (40), Ala (70). L. scrobiculata (Scop.) DC. — on bark of Populus suaveolens, siliceous rocks, mosses; 5, 26 [R]; LE (sub Rinodina turfacea). — Kam (8), Chu (21), Ala (70).

Lopadium coralloideum (Nyl.) Lynge — on soil, plant debris; 19 [R1]; H. — Kam (16), Chu (21), Yak (40), Ala (61). L. pezizoideum (Ach.) Körb. — on soil; 5, 19 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). Melanelia hepatizon (Ach.) A. Thell — on rocks; 5, 7, 8, 33, a1, a3 [R]; H, LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). M. stygia (L.) Essl. — on rocks; 5, 7, 8, 33 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). Melanohalea infumata (Nyl.) O. Blanco, A. Crespo, Divakar, Essl., D. Hawksw. et ­Lumbsch — on rocks; 7 [R1]; H, LE. — Kam (13), Chu (21), Mag (33), Yak (40), Ala (70).

121 Himelbrant et al. The first lichenological survey in Koryakia

M. olivacea (L.) O. Blanco, A. Crespo, Divakar, Essl., D. Hawksw. et Lumbsch — on bark of Betula ermanii, B. divaricata, Alnus fruticosa, Pinus pumila, Populus suaveolens, Salix arbutifolia, S. pulchra, S. schwerinii, S. udensis and lignum of Sorbus sibirica; 1–4, 6, 7, 10, 13–16, 20, 22, 26, 27, 30, 32, 35, a4, a7 [F]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). M. septentrionalis (Lynge) O. Blanco, A. Crespo, Divakar, Essl., D. Hawksw. et Lumbs- ch — on bark of Betula ermanii, B. divaricata, Alnus fruticosa, Populus suaveolens, Salix arbuti- folia, S. pulchra; 1, 6, 9, 16, 20, 21, 30, 32 [O]; LE (sub Lecanora chlarotera). — Kam (16), Chu (21), Mag (33), Yak (40), Ala (73).

!Micarea melaena (Nyl.) Hedl. — on mosses; 24 [R1]; LE. — Kam (Kamchatka Territo- ry, Milkovo District, Kronotsky Nature Reserve, Levaya Schapina River basin, 55°08'38"N, 159°59'24"E, 370 m a. s. l., spruce forest with mosses and horsetails, on wood of Picea ajanensis, 13 VIII 2009, Himelbrant, Stepanchikova K-14-09, H), Chu (21), Mag (33), Yak (40), Ala (61). M. prasina Fr. — on lignum of Salix udensis; 20, 26 [R]; LE. Thalli contain micareic acid. — Kam (16), Yak (40), Ala (70). !Miriquidica leucophaea (Flörke ex Rabenh.) Hertel et Rambold — on rocks; 5 [R1]; H. — Chu (21), Mag (33), Yak (40), Ala (55). Montanelia disjuncta (Erichsen) Divakar, A. Crespo, Wedin et Essl. — on rocks; 5, 7, a2 [R]; LE (sub Aspicilia cinerea). — Kam (16), Chu (21), Mag (36), Yak (40), Ala (73). M. panniformis (Nyl.) Divakar, A. Crespo, Wedin et Essl. — on rocks; 5, 7, 8, 33, a1 [R]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

M. sorediata (Ach.) Divakar, A. Crespo, Wedin et Essl. — on rocks; 7 [R1]. — Kam (16), Chu (21), Yak (40), Ala (73). Mycobilimbia carneoalbida (Müll. Arg.) S. Ekman et Printzen — on bark of Salix arbutifo- lia, Populus suaveolens, mosses; 20, 26, 30 [R]; LE. — Kam (16), Chu (21), Yak (40), Ala (73). M. epixanthoides (Nyl.) Vitik., Ahti, Kuusinen, Lommi et T. Ulvinen ex Hafellner et Türk — on mosses; 20, 26 [R]. — Kam (16), Ala (72). M. tetramera (De Not.) Vitik., Ahti, Kuusinen, Lommi et T. Ulvinen ex Hafellner et Türk — on bark of Populus suaveolens; 26 [R1]; LE. — Kam (9), Chu (21), Yak (45), Ala (70). Mycoblastus sanguinarius (L.) Norman — on bark of Betula divaricata, Pinus pumila, sili- ceous rocks; 3, 5, 7, 14, 17, 22, 33 [R]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (60). +Mycocalicium subtile (Pers.) Szatala — on bark of Salix arbutifolia and lignum of Sorbus sibirica; 27, 30 [R]; LE. — Kam (16), Chu (21), Yak (43), Ala (61).

Myriolecis dispersa (Pers.) (Pers.) Śliwa, X. Zhao et Lumbsch — on reindeer horn; 8 [R1]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

M. straminea Ach. — on rocks; 8 [R1]; H. — Kam (5), Chu (21), Ala (60). Nephroma arcticum (L.) Torss. — on soil; 5, a6 [R]. — Kam (8), Chu (21), Mag (30), Yak (40), Ala (70). N. bellum (Spreng.) Tuck. — on bark of Salix pulchra, S. udensis; 2, 16, 20, 35 [R]; H (sub Nephroma parile). — Kam (8), Chu (22), Mag (39), Yak (40), Ala (70).

N. expallidum (Nyl.) Nyl. — on soil; a5 [R1]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (64). N. parile (Ach.) Ach. — on bark of Betula divaricata, Populus suaveolens, Salix arbutifolia, S. pulchra; 1, 2, 20, 26 [R]; H; LE. — Kam (16), Chu (21), Mag (36), Yak (40), Ala (70). Ochrolechia frigida (Sw.) Lynge s. lat. [incl. O. frigida, O. gonatodes (Ach.) Räsänen, O. ­inaequatula (Nyl.) Zahlbr.] — on soil, plant debris, bark of Betula divaricata, Pinus pumila and Salix pulchra; 3–5, 7, 8, 10, 16, 18, 19, 21, 22, 24, 25, 28, 29, 31, 34 [F]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

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O. mahluensis Räsänen — on bark of Betula ermanii, Pinus pumila, Salix pulchra; 2, 14, 32, a1, a7 [R]; H (sub Lopadium coralloideum). Thalli contain gyrophoric and lecanoric acids. — Kam (16), Ala (74). O. subplicans (Nyl.) Brodo subsp. subplicans — on rocks; 5, 8 [R]; H, LE. — Kam (5), Chu (21), Ala (60). — Crustose lichen with very distinctive shape of thallus and apothecia (Plate II, 2). Ophioparma lapponica (Räsänen) Hafellner et R. W. Rogers — on rocks; 5, 33 [R]; H. — Kam (6), Mag (LE), Yak (40), Ala (70). Parmelia omphalodes (L.) Ach. subsp. omphalodes — on rocks, soil, plant debris and mos­ ses; 5, 7, 8, 19, 33, a1–a3 [R]; H, LE. Thalli contain salazinic acid and atranorin. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70).

P. pinnatifida Kurok. — on rocks; 5 [R1]; LE. — Kam (16), Ala (70). P. saxatilis (L.) Ach. — on bark of Pinus pumila, siliceous rocks and primary soil; 5, 7 [R]; LE. Thalli contain atranorin, lobaric and salazinic acids. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

P. skultii Hale — on rocks; 5 [R1]; LE. — Kam (16), Yak (41), Ala (73).

P. squarrosa Hale — on bark of Betula ermanii; 32 [R1]; LE. — Kam (16), Mag (36), Ala (70). P. sulcata Taylor — on bark of Betula ermanii, Alnus fruticosa, Pinus pumila, Populus sua­ veolens, Salix arbutifolia, S. schwerinii, S. udensis, siliceous rocks; 5, 7, 20, 26, 27, 30, 32 [R]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). Parmeliopsis ambigua (Wulfen) Nyl. — on bark of Betula ermanii, B. divaricata, Alnus fru- ticosa, Pinus pumila, Salix pulchra, Vaccinium uliginosum; 1–3, 7, 9, 10, 12, 14–17, 21, 22, 27, 30, 32, 35, a4, a7 [F]; H (sub Arthopyrenia analepta). — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). P. hyperopta (Ach.) Arnold — on bark of Alnus fruticosa, Betula ermanii, B. divaricata, Pinus pumila, Salix pulchra and lignum; 1, 3, 7, 9, 10, 14–17, 20, 22, 27, 32, 35, a4 [O]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70).

Peltigera canina (L.) Willd. — on soil; 9 [R1]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (60). P. collina (Ach.) Schrad. — on bark of Salix arbutifolia, Populus suaveolens, Salix pulchra; 16, 20, 26, 30 [R]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (60).

P. didactyla (With.) J. R. Laundon — on soil; 2 [R1]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70).

P. extenuata (Norrl. ex Vain.) Lojka — on bark of Salix pulchra; 2 [R1]. — Kam (16), Yak (48), Ala (70).

!P. latiloba Holt.-Hartw. — on mosses; 2 [R1]; H. — Yak (44), Ala (70).

P. lepidophora (Vain.) Bitter — on bark of Salix pulchra; 2 [R1]; LE. — Kam (16), Chu (21), Yak (40), Ala (70). P. leucophlebia (Nyl.) Gyeln. — on soil, mosses and bark of Salix pulchra; 2, 9, 19, 21, 28, 35 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). P. malacea (Ach.) Funck — on soil; 19, 34 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). P. membranacea (Ach.) Nyl. — on soil; 9 [R1]; LE. — Kam (16), Chu (21), Yak (40), Ala (70). P. neopolydactyla (Gyeln.) Gyeln. — on soil; 2 [R1]. — Kam (16), Yak (40), Ala (70). P. polydactylon (Neck.) Hoffm. — on soil; 9, 10 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70).

123 Himelbrant et al. The first lichenological survey in Koryakia

Plate II. 1a, 1b — Pilophorus vegae (H, Tal-7); 2 — Ochrolechia subplicans (LE, Tal-5); 3 — Protoparmelia cupreobadia (H, Am-19); 4 — Rimularia badioatra (H, Tal-5); 5 — Stereocaulon spinosum (LE, Tal-1ad). Scale bars: 1a — 1 mm, 1b — 1 cm, 2–5 — 1 mm.

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P. praetextata (Flörke ex Sommerf.) Zopf — on soil; 2 [R1]. — Kam (12), Chu (21), Mag (39), Yak (40), Ala (70). P. rufescens (Weiss) Humb. — on soil and bark of Salix pulchra; 6, 7, 19 [R]; LE. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70). P. scabrosa Th. Fr. — on soil, mosses; 9, 10, 21, 25, 29 [R]. — Kam (16), Chu (21), Mag (30), Yak (40), Ala (70).

!Pertusaria borealis Erichsen — on bark of Pinus pumila; 7 [R1]; LE. Thallus contains fumar­ protocetraric and protocetraric acids. — Ala (70). New to Russian Far East. The nearest known localities in Russia are in Ural Mountains (e. g., Paukov, Teptina, 2013). P. carneopallida (Nyl.) Anzi — on bark of Alnus fruticosa and lignum; 15, 20, 27, 30, 32 [R]; LE. — Kam (16), Chu (21), Yak (40), Ala (70).

P. coriacea (Th. Fr.) Th. Fr. — on plant debris; 5 [R1]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

P. leioplaca DC. — on bark of Salix udensis; 20 [R1]; LE. — Kam (8), Ala (58). P. oculata (Dicks.) Th. Fr. — on soil; 7, 8, 18 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). P. sommerfeltii (Flörke ex Sommerf.) Fr. — on bark of Populus suaveolens, Salix arbutifolia, S. pulchra, S. udensis; 2, 20, 26, 30 [R]; LE. Thalli contain stictic acid and 2,7-dichlorolichexan- thone. — Kam (12), Chu (21), Yak (47), Ala (61). +Phaeocalicium compressulum (Nyl. ex Vain.) A. F. W. Schmidt — on bark of Alnus fruti- cosa; 15, 20, 26, 27, 30, 32, a4 [R]; LE. — Kam (11), Chu (21), Yak (48), Ala (70).

+P. tremulicola (Norrl. ex Nyl.) Tibell — on bark of Populus suaveolens; 30 [R1]; LE. — Kam (11). Phaeophyscia denigrata (Hue) Moberg — on bark of Alnus fruticosa, Populus suaveolens; 20, 26 [R]; LE. — Kam (12), Mag (34), Yak (40). P. kairamoi (Vain.) Moberg — on bark of Alnus fruticosa, Populus suaveolens, Salix arbuti- folia, S. udensis, Sorbus sibirica; 20, 26, 27, 30 [R]; H, LE. — Kam (16), Mag (36), Yak (40), Ala (70).

P. rubropulchra (Degel.) Essl. — on bark of Populus suaveolens; 20 [R1]; LE. — Kam (12).

!Phylliscum demangeonii (Moug. et Mont.) Nyl. — on rocks; 8 [R1]; H. — Chu (21), Ala (70). Physcia aipolia (Ehrh. ex Humb.) Fürnr. — on bark of Alnus fruticosa, Populus suaveolens, Salix arbutifolia, S. pulchra, S. schwerinii, S. udensis, Sorbus sibirica; 2, 20, 26, 27, 30 [R]; LE. — Kam (16), Chu (25), Mag (33), Yak (40), Ala (70).

P. alnophila (Vain.) Loht., Moberg, Myllys et Tehler — on bark of Alnus fruticosa; 26 [R1]; LE. — Kam (18), Chu (18), Mag (18), Yak (18), Ala (73). P. caesia (Hoffm.) Fürnr. — on rocks, reindeer horn; 5, 7, 8 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). P. dubia (Hoffm.) Lettau — on bark ofBetula ermanii, Alnus fruticosa, siliceous rocks; 5, 7, 26, 32, a2 [R]; H (sub Phaeophyscia denigrata). — Kam (16), Chu (21), Mag (36), Yak (40), Ala (70).

!Physciella melanchra (Hue) Essl. — on rocks; 5 [R1]; LE. — New to Beringia and northern part of Russian Far East. The nearest known locality is Sakhalin Island (Tchabanenko, 2006). Physconia detersa (Nyl.) Poelt — on bark of Populus suaveolens and primary soil on rocks; 5, 26 [R]; LE. — Kam (8), Chu (21), Yak (40), Ala (64).

P. grumosa Kashiw. et Poelt — on rocks and primary soil; 5 [R1]; LE. — Kam (16), Ala (69).

125 Himelbrant et al. The first lichenological survey in Koryakia

P. leucoleiptes (Tuck.) Essl. — on rocks; 5 [R1]; H. — Kam (13), Chu (24).

!Piccolia ochrophora (Nyl.) Hafellner — on bark of Salix arbutifolia; 20 [R1]; LE. — Ala (74). New to Russian Far East. The nearest published locality in Russia is Sayan Mountains (Sedelnikova, 2001). !Pilophorus vegae Krog — on rocks and primary soil; 7, 8 [R]; H, LE. Thalli contain at- ranorin, stictic acid and zeorin. — Chu (20), Ala (70). — Characterized by having esorediate corticate pseudopodetia with partly blackish medulla, usually sterile. The specimen kept in H has several developed apothecia, those being recorded for the first time for this species (Plate II, 1a and 1b).

Placynthiella dasaea (Stirt.) Tønsberg — on mosses; 18 [R1]; H. — Kam (16), Ala (70). P. icmalea (Ach.) Coppins et P. James — on soil, plant debris, mosses, bark of Betula ermanii and Salix arbutifolia; 5, 10, 18–21, 32, 34 [O]; LE. — Kam (16), Yak (40), Ala (70). P. uliginosa (Schrad.) Coppins et P. James — on soil, plant debris and bark of Betula ermanii; 10, 24, 25, 32 [R]. — Kam (16), Chu (21), Yak (40), Ala (70). Polycauliona candelaria (L.) Frödén, Arup et Søchting — on rocks, manured by birds; 5, 7 [R]; LE. — Kam (8), Chu (21), Mag (39), Yak (40), Ala (70).

!Polysporina simplex (Taylor) Vězda — on rocks; 5 [R1]; LE. — Chu (21), Yak (40), Ala (61). Porpidia crustulata (Ach.) Hertel et Knoph — on rocks; 8, 19 [R]; H. — Kam (1), Chu (21), Mag (33), Yak (40), Ala (55).

P. flavicunda (Ach.) Gowan — on rocks; a6 [R1]; H. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). P. macrocarpa (DC.) Hertel et A. J. Schwab — on rocks; 7, 8, 33 [R]. — Kam (16), Mag (33), Yak (40), Ala (70).

!P. melinodes (Körb.) Gowan et Ahti — on rocks; 8 [R1]; H. — Chu (21), Yak (40), Ala (57). Protopannaria pezizoides (Weber) P. M. Jørg. et S. Ekman — on soil; 5, 19 [R]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). Protoparmelia badia (Hoffm.) Hafellner — on rocks; 5, 7, 8 [R]. — Kam (16), Chu (25), Mag (33), Yak (40), Ala (70).

!P. cupreobadia (Nyl.) Poelt — on rocks; 33 [R1]; H. — Yak (51), Ala (72). — Rare saxico- lous species with distinctive dark brown glossy crustose thallus (Plate II, 3). Psoroma hypnorum (Vahl.) Gray — on soil and bark of Betula divaricata; 19, 21 [R]. — Kam (16), Yak (40), Ala (70). Ramalina almquistii Vain. — on rocks; 5, 7, a3 [R]; LE. Thalli contain usnic and sekikaic acids. — Kam (2), Chu (21), Mag (34), Yak (40), Ala (60). R. roesleri (Hochst. ex Schaer.) Hue — on rocks; 5, 7 [R]; LE. Thalli contain usnic and sekikaic acids. — Kam (16), Chu (21), Mag (36), Ala (70). Ramboldia cinnabarina (Sommerf.) Kalb, Lumbsch et Elix — on bark of Pinus pumila, Rho- dodendron aureum; 1, 7, 14 [R]. — Kam (16), Ala (70).

Rhizocarpon geographicum (L.) DC. — on rocks; 5 [R1]; H. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). R. grande (Flörke ex Flot.) Arnold — on rocks; 5, 19 [R]; H. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (64). R. infernulum (Nyl.) Lynge f. infernulum — on rocks; 8, 19 [R]; H. — Kam (1), Chu (23), Ala (52).

Rhizoplaca chrysoleuca (Sm.) Zopf — on rocks; 5 [R1]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (64).

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!Rimularia badioatra (Kremp.) Hertel et Rambold — on rocks; 5 [R1]; H. — New to Beringia and Russian Far East. The nearest known locality is Kodar Range, the Trans-Baikal Territory (Chesnokov et al., 2018). — Rarely recorded saxicolous lichen with crustose brownish thallus containing gyrophoric acid, and contorted apothecia (Plate II, 4). Rinodina cinereovirens (Vain.) Vain. — on bark of Betula ermanii, Salix arbutifolia, Alnus fruticosa, Populus suaveolens; 20, 26, 32 [R]; LE. — Kam (19), Ala (74). R. septentrionalis Malme — on bark of Betula divaricata, Alnus fruticosa, Populus suaveo- lens, Salix arbutifolia, S. pulchra; 2, 6, 16, 20, 22, 26, 30, 35 [O]; LE. — Kam (19), Chu (21), Yak (40), Ala (70). R. turfacea (Wahlenb.) Körb. — on plant debris, thallus of Lobaria scrobiculata on siliceous rock and dead thallus of Peltigera sp. on soil; 5, 19 [R]; LE. — Kam (19), Chu (21), Mag (33), Yak (40), Ala (64).

R. willeyi Sheard et Giralt — on bark of Betula ermanii; 32 [R1]; LE. Thalli contain pan- narin. — Kam (19).

Ropalospora lugubris (Sommerf.) Poelt — on rocks; 7 [R1]; H. — Kam (16), Chu (21), Yak (40), Ala (70). Rostania occultata (Bagl.) Otálora, P. M. Jørg. et Wedin — on bark of Populus suaveolens;

20 [R1]; LE. — Kam (9), Ala (73). Rusavskia elegans (Link) S. Y. Kondr. et Kärnefelt — on rocks, manured by birds; 5, 7 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). R. sorediata (Vain.) S. Y. Kondr. et Kärnefelt — on rocks and primary soil; 5, a3 [R]; LE. — Kam (16), Chu (21), Yak (40), Ala (73). !Sagedia mastrucata (Wahlenb.) A. Nordin, Savić et Tibell — on rocks; 5, 7, 8, a2 [R]; H, LE. Thalli contain norstictic acid. — Chu (21), Ala (74).

!+Sarea difformis (Fr.) Fr. — on resin of Pinus pumila; 1 [R1]. — Kam (Kamchatka Terri- tory, Milkovo District, Kronotsky Nature Reserve, Levaya Schapina River basin, 55°08'29"N, 159°58'17"E, 340 m a. s. l., spruce forest with mosses, on resin of Picea ajanensis, 12 VIII 2009, Himelbrant, Stepanchikova K-12-09, H). +S. resinae (Fr.: Fr.) Kuntze — on resin of Pinus pumila; 1, 3, 7 [R]. — Kam (16), Mag (36), Yak (40). !#Sclerococcum montagnei Hafellner — on thallus of Lecanora bicincta on siliceous rock; a2 [R1]; LE. — Chu (26). !#Sclerococcum parasiticum (Flörke) Ertz et Diederich — on thalli of Lepra dactylina and L. panyrga on soil and plant debris; 8, 33, a1 [R]; H, LE. — New to Beringia and northern part of the Russian Far East. The nearest known locality is Amur River basin, Khabarovsk Territory of Russia (Zhurbenko, Tugi, 2013). Scoliciosporum chlorococcum (Graewe ex Stenh.) Vězda — on bark of Betula ermanii;

32 [R1]. — Kam (16), Yak (45), Ala (58).

S. umbrinum (Ach.) Arnold — on rocks; 19 [R1]. — Kam (16), Yak (40), Ala (70). Scutula circumspecta (Vain.) Kistenich, Timdal, Bendiksby et S. Ekman — on bark of Salix arbutifolia; 20, 30 [R]; LE. — Kam (9), Yak (50), Ala (74). Scytinium teretiusculum (Wallr.) Otálora, M. Jørg. et Wedin — on bark of Populus suaveo- lens; 26 [R1]; LE (sub Candelariella efflorescens). — Kam (16), Yak (50), Ala (70). Sphaerophorus fragilis (L.) Pers. — on rocks; 5, 7, 8, 33, a1 [R]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). S. globosus (Huds.) Vain. — on soil; 5, 7, 8, 18, 19, 21, 24, 29, 31, 33 [O]; H (sub Cladonia wainioi). — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

127 Himelbrant et al. The first lichenological survey in Koryakia

!#Sphinctrina leucopoda Nyl. — on thallus of Lecanora rupicola on siliceous rock; 5 [R1]; H (sub Lecanographa grumulosa). — New to Beringia and northern part of Russian Far East. The nearest known locality in Russia is Sakhalin Island (Titov, 2006).

Steinia geophana (Nyl.) Stein — on mosses; 25 [R1]; H. — Kam (16), Ala (42). Stereocaulon alpinum Laurer — on soil; 5, 7 [R]. — Kam (7), Chu (21), Mag (30), Yak (40), Ala (64).

S. apocalypticum Nyl. — on soil; 33 [R1]; H. — Kam (7), Chu (21), Mag (33), Yak (40), Ala (56).

S. depreaultii Delise ex Nyl. — on rocks; 8 [R1]; LE. — Kam (7), Chu (21). S. glareosum (Savicz) H. Magn. — on soil; 5, 35 [R]; LE. — Kam (7), Chu (21), Mag (33), Yak (40), Ala (70). S. paschale (L.) Hoffm. — on soil and siliceous rocks; 5, 7, 9, 18, 19, 21, 31, 33, 34 [O]; H. — Kam (7), Chu (21), Mag (30), Yak (40), Ala (70). S. spinosum Dombr. — on rocks and primary soil; 8, a1 [R]; LE. Thalli contain atranorin and porphyrilic acid. — Kam (7). — The species is morphologically similar (and probably related) to S. depreaultii, from which it differs by developing digitate phyllocladia additionally to rounded ones (Plate II, 5), and grayish cephalodia.

S. subcoralloides (Nyl.) Nyl. — on rocks; a3 [R1]; LE. — Kam (7), Chu (21), Mag (33), Yak (40), Ala (60). S. symphycheilum I. M. Lamb — on soil and siliceous rocks; 7, 8 [R]. — Kam (7), Chu (21), Ala (70). S. wrightii Tuck. — on soil and siliceous rocks; 7, 8, 33, a1 [R]; LE. — Kam (7), Chu (21), Mag (30), Yak (40), Ala (53).

!Strangospora moriformis (Ach.) Stein — on bark of Betula ermanii; 32 [R1]; LE. — New to Beringia and Russian Far East. The nearest known locality in Russia is Khamar-Daban Range within the Republic of Buryatia (Urbanavichene, 2015). Tephromela atra (Huds.) Hafellner — on rocks; 5, 7 [R]; H. — Kam (5), Chu (21), Mag (33), Yak (40), Ala (60). Tetramelas triphragmioides (Anzi) A. Nordin et Tibell — on bark of Populus suaveolens, Salix arbutifolia, S. udensis; 20, 26 [R]; LE. — Kam (13), Ala (74). Thamnolia vermicularis (Sw.) Schaer. — [F]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (60): var. vermicularis — on soil and siliceous rocks; 4, 5, 7, 8, 12, 13, 17–19, 21, 22, 24, 25, 28, 29, 31, 34, a1; var. subuliformis (Ehrh.) Schaer. — on rocks; 5, 33.

!Thelenella modesta (Nyl.) Nyl. — on bark of Populus suaveolens; 20 [R1]; LE. — Ala (74). New to the northern part of Russian Far East. The nearest known locality in Russia is Sikhote- Alin within the Primorye Territory (Tchabanenko, 2002). Toensbergia leucococca (R. Sant.) Bendiksby et Timdal — on bark of Alnus fruticosa, Bet- ula ermanii, B. divaricata, Pinus pumila, Rhododendron aureum; 1, 3, 7, 14, 15, 21, 22, 27, 32 [O]; LE. — Kam (16), Yak (H), Ala (70). Toninia populorum (A. Massal.) Kistenich, Timdal, Bendiksby et S. Ekman — on bark of Salix arbutifolia, Populus suaveolens; 20, 30 [R]; LE. — Kam (16), Yak (49). Toniniopsis subincompta (Nyl.) Kistenich, Timdal, Bendiksby et S. Ekman — on bark of Populus suaveolens, Salix arbutifolia, S. pulchra, S. schwerinii, mosses; 16, 20, 26, 27, 30, 35 [R]; LE. — Kam (9), Chu (21), Yak (47), Ala (70).

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Trapeliopsis flexuosa (Fr.) Coppins et P. James — on lignum and plant debris; 19, 31, a1 [R]; LE (sub Elixia flexella). — Kam (16), Yak (40), Ala (70). T. granulosa (Hoffm.) Lumbsch — on soil; 5, 7, 24, 28, 34 [R]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

Tremolecia atrata (Ach.) Hertel — on rocks; 8 [R1]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). Tuckermannopsis ciliaris (Ach.) Gyeln. s. lat. — on bark of Betula exilis, B. divaricata; 1, 4, 9 [R]. — Kam (16), Chu (21), Mag (36), Yak (H), Ala (63).

Umbilicaria cinereorufescens (Schaer.) Frey — on rocks; 5 [R1]; LE. — Kam (14), Chu (21), Mag (38), Yak (40), Ala (70).

U. deusta (L.) Baumg. — on rocks; 5 [R1]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). U. hyperborea (Ach.) Hoffm. var.hyperborea — on rocks; 5, 7, 8, a1 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). U. proboscidea (L.) Schrad. — on rocks; 5, 7, 8, 33, a1 [R]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70). U. torrefacta (Lightf.) Schrad. — on rocks; 7, 8, 33 [R]. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

U. vellea (L.) Hoffm. — on rocks; 5 [R1]; LE. — Kam (14), Chu (21), Mag (35), Yak (40), Ala (70). !Varicellaria rhodocarpa (Körb.) Th. Fr. — on bark of Pinus pumila; 3, 14 [R]; LE. Thalli contain lecanoric acid and lichexanthone. — Chu (21), Yak (40), Ala (61). Vulpicida juniperinus (L.) J.-E. Mattson et M. J. Lai — on bark of Betula divaricata, Pinus pumila; 1, 3, 7, 9, 10, 14, 17, 22 [O]; LE. — Kam (16), Chu (21), Mag (33), Yak (40), Ala (55). V. pinastri (Scop.) J.-E. Mattson et M. J. Lai — on bark of Alnus fruticosa, Betula ermanii, B. exilis, B. divaricata, Pinus pumila, Rhododendron aureum, Salix arbutifolia, S. pulchra, S. schwer- inii, S. udensis, Vaccinium uliginosum; 1, 3, 4, 7, 9, 10, 12, 14–17, 20–22, 26, 27, 30, 32, 35, a4, a7 [F]; LE (sub Lecanora albellula). — Kam (16), Chu (21), Mag (33), Yak (40), Ala (70).

Xylographa parallela (Ach.: Fr.) Fr. — on lignum of Pinus pumila; 34 [R1]; LE. — Kam (17), Ala (17).

X. vermicularis T. Sprib. — on lignum; a1 [R1]; LE. Thallus contains stictic, confriesiic (ma- jor) and friesiic (minor) acids. — Kam (17), Ala (17).

The revealed diversity of lichens and allied fungi of Parapolsky Dale counts 315 spe- cies: 300 lichens, 7 lichenicolous fungi and 8 non-lichenized saprobic fungi related to lichens. Altogether 46 species are published from the Kamchatka Territory for the first time, including Lecanographa grumulosa new to Russia, East Asia and Beringia; Cercidospora trypetheliza, Lecania dubitans, Pertusaria borealis, Piccolia ochrophora, Protoparmelia cupreobadia, Rimularia badioatra and Strangospora moriformis new to Russian Far East; Abrothallus bertianus, Cladonia strepsilis, Physciella melanchra, Rim- ularia badioatra, Sclerococcum parasiticum, Sphinctrina leucopoda and Strangospora moriformis new to Beringia. Although the list of lichenicolous fungi for the study area is obviously incomplete and needs special research, the lichen diversity has been stud- ied rather comprehensively and is just predictably poor. The most interesting habitats, including rocks, are concentrated on sloping edges of the Parapolsky Dale, the bottom is relatively monotonous and boring for lichenol-

129 Himelbrant et al. The first lichenological survey in Koryakia ogist. The richest biotopes are rocky outcrops (147 lichen species, 46.7 % of the stud- ied biota) and tundra of different types (139 species, 44.1 %). Relatively high is the lichen diversity of shrub communities (altogether 108 species, 34.3 %), dominated by Salix pulchra (54 species), Betula divaricata (51 species), Pinus pumila (45 species) and Alnus fruticosa (31 species). Forest communities are not common in Parapolsky Dale. The forest lichens count 89 species (28.3 % of the studied biota), of them 73 spe- cies were recorded in floodplain forests, and 36 species only — in stone-birch forest. Bogs occupy significant part of the study area, but they have relatively poor lichen biota (78 species, 24.8 %). An average number of species per standard sample plot is 33.8±3.5: 11–34 species per sample plot in shrub communities, 3–42 in bogs, 33–57 in forests, 11–77 in tundras, and 93–98 in rocky outcrops. Most diverse are epiphytic lichens (124 species, 39.4 % of the studied biota), the richest phorophytes are Salix pulchra (46 species), Populus suaveolens (43 species), Sa- lix arbutifolia and Pinus pumila (38 species each), Alnus fruticosa (35 species), Betula divaricata (33 species), B. ermanii (27 species), Salix udensis (15 species); few lichens were also recorded on bark of Salix schwerinii and Sorbus sibirica (8 species each), Bet- ula exilis (7 species), Vaccinium uliginosum (5 species), Rhododendron aureum (3 spe- cies). Soil is the second richest substrate, it has 114 species (36.2 % of the studied biota). Rocks are rare in Parapolsky Dale and occur only on its boards. Altogether 95 saxicolous species (30.2 %) were recorded; the lichens found here are typical for si- liceous (non-calcareous) stones. Deadwood is poor represented and has relatively low lichen diversity (21 species, 6.7 % of the studied biota); 19 species (6.0 %) were found on mosses, 14 species (4.4 %) — on plant debris, 10 species (3.2 %) — on lichen thalli (including 7 fungi and 3 lichens), also 3 species were recorded on reindeer horn, and 2 saprobic fungi — on resin of Pinus pumila. Within the study area, the majority of species have rare [R] frequency of occur- rence (altogether 270 species or 85.7 % of the studied biota; of them, 117 were record- ed once). Only two species (0.6 %) — namely, Cladonia amaurocraea and Cladonia arbuscula — are “common” [C], 16 species (5.1 %) were found “frequently” [F] (Ce- traria laevigata, C. sepincola, Cladonia cornuta, C. gracilis, C. pyxidata, C. rangiferina, C. stygia, Flavocetraria cucullata, F. nivalis, Lecanora fuscescens, L. symmicta, Melano- halea olivacea, Ochrolechia frigida s. l., Parmeliopsis ambigua, Thamnolia vermicularis, Vulpicida pinastri), and 27 species (8.6 %) — “occasionally” [O]. Comparison with neighboring regions (Kamchatka Peninsula, Chukotka, Magadan Region, Yakutia and Alaska, for most of them at least 600–1000 lichen species have been reported) shows that the lichen flora of Parapolsky Dale contains almost no spe- cific species. Half of the studied biota (150 species) are known from all adjacent re- gions (Kamchatka, Chukotka, Magadan, Alaska) and Yakutia. Most species recorded in Parapolsky Dale are known also from Alaska (288 species, 91.4 % of the studied li- chen biota), Kamchatka outside Koryakia (278 species, 88.3 %), Yakutia (241 species, 76.5 %), Chukotka (219 species, 69.5 %) and Magadan (178 species, 56.5 %). Further statistical similarity analysis of species lists would show obviously different degree

130 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 53(1): 107–142. 2019 of knowledge of regional lichen floras: e. g., Alaska is well-studied, unlike Magadan. Anyway, the obtained numbers are enough to show high similarity and low specificity of the studied biota relative to neighboring regions, including both Beringian (Kam- chatka, Chukotka, Alaska) and more remote (Magadan, Yakutia) territories. Most species discovered in Koryakia have rather wide geographical ranges. Of special interest are species supposed to be Amphiberingian (Thomson, 1984) in wide sense, or endemic for North Pacific and adjacent territories (including Eastern Sibe- ria and Yukon): Hypogymnia pseudophysodes, Stereocaulon spinosum (both known from Asia only), Amygdalaria subdissentiens, Cetraria kamczatica, Cladonia kanewskii, C. nipponica, C. wainioi, Ochrolechia subplicans, Pilophorus vegae, Stereocaulon apoca- lypticum, and S. wrightii. Most of these species, however, are known not from Beringia only, but also from neighboring inland territories (e. g., Yakutia). Some other species were considered before as Amphiberingian as well (e. g., Asahinea spp., Cetrelia alaska- na), but nowadays their ranges are known to be wider, up to circumpolar in some cases. However, the distribution of lichen species in Northern Asia requires further studies. Acknowledgements We would like to thank the head and the staff of Kronotsky Nature Reserve for organization and support of the expedition, as well as inspectors of Koryaksky Nature Reserve Anatoly Sorokin and Alexandr Zyryanov for friendly help during field stud- ies. We are grateful to Ivan Frolov (Botanical Garden of the Urals Branch of Russian Academy of Sciences, Yekaterinburg) for revision of Caloplaca specimens, to Toby Spribille (Edmonton, Canada) for help in finding information on species distribution in Alaska, and also to Leena Myllys and Saara Velmala (Helsinki, Finland) for assis- tance in organizing laboratory studies. The study was carried out within the frame- work of the institutional research project “Flora of lichens and bryophytes of Russia and phytogeographically important regions” (no. АААА-А19-119020690077-4) of the Komarov Botanical Institute RAS and supported by Russian Foundation for Ba- sic Research (expedition: grant 16-05-00736; laboratory research: grants 18-05-60093 and 19-05-00805). References Ahti T. 2007. Further studies on the Cladonia verticillata group (Lecanorales) in East Asia and wes­ tern North America. Bibliotheca Lichenologica 96: 5–19. Andreev M. P. 1978. The lichens of research station “Aborigen” (Tenkin District of the Magadan Region). Botanicheskii zhurnal 63(11): 1626–1632. (In Russ. with Engl. abstract). Andreev M. P. 1984. Lichenes epilithici insulae Ionae (Mare Ochotense). Novosti sistematiki nizshikh rastenii 21: 143. (In Russ. with Latin abstract). Andreev M., Kotlov Y., Makarova I. 1996. Checklist of lichens and lichenicolous fungi of the Russian Arctic. The Bryologist 99(2): 137–169. https://doi.org/10.2307/3244545 Brigham-Grette J. 2004. Paleoenvironmental Conditions in Western Beringia before and during the Last Glacial Maximum. Entering America. Northeast Asia and the Beringia Before the Last Glacial Maximum. Salt Lake City: 1–57. Brodo I. M., Duran Sharnoff S., Sharnoff S. 2001. Lichens of North America. New Haven; London: 795 p.

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