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59New data on diversity of lichens and lichenicolous fungi of the Amur Region and the Russian Far East
E. S. Kuznetsova1, 2, S. V. Dudov3, 4, O. I. Ryabenko3
'St. Petersburg State University, St. Petersburg, Russia 2Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia 3Lomonosov Moscow State University, Moscow, Russia 4Zeya State Nature Reserve, Amur Region, Zeya, Russia Corresponding author. E. S. Kuznetsova, [email protected]
Abstract. The lichen biota of the Amur Region (southern Russian Far East) was studied in the course of several geobotanical expeditions. In total 38 species of lichens and three lichenicolous fungi are reported for the first time for the Amur Region. Among them four species (Fulgidea oligospora, Peltigera frippii, Psora globifera, and Rhizoplaca subdiscrepans) are new to the Russian Far East and nine more — to the southern Russian Far East. Four and 38 species are new to the Zeya Reserve and Tokinsko-Stanovoy National Park, respectively.
Keywords: distribution of lichens, lichenicolous fungi, new records, Zeya Nature Reserve, Stanovoy Range, Tokinsko-Stanovoy National Park, Tukuringra Range.
Новые данные о разнообразии лишайников Амурской области и Российского Дальнего Востока
Е. С. Кузнецова1, 2, С. В. Дудов3, 4, О. И. Рябенко3
'Санкт-Петербургский государственный университет, Санкт-Петербург, Россия 2Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия 3Московский государственный университет им. М. В. Ломоносова, Москва, Россия
4Зейский государственный природный заповедник, Амурская обл., Зея, Россия Автор для переписки. Е. С. Кузнецова, [email protected]
Резюме. Лихенобиота Амурской обл. (юг Дальнего Востока России) была изучена в ходе нескольких геоботанических экспедиций. Всего для Амурской обл. впервые отмечено 38 видов лишайников и три вида лихенофильных грибов. Из них четыре вида являются новыми для Дальнего Востока России (Fulgidea oligospora, Peltigera frippii, Psora globifera и Rhizoplaca subdiscrepans) и девять — для юга Дальнего Востока России. Четыре вида являются новыми для Зейского заповедника и 38 — для Токинского-Станового национального парка.
Ключевые слова: распространение лишайников, лихенофильные грибы, новые находки, Зейский заповедник, Становой хребет, Токинско-Становой национальный парк, хребет Туку-рингра.
There is a great number of publications devoted to the lichens and allied fungi of the southern Russian Far East, that includes the Amur Region, Jewish Autonomous Region, Primorye Territory, the Sakhalin Region, and Khabarovsk Territory in part.
https://doi.org/10.31111/nsnr/2022.562357
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All available data were consolidated by S. I. Tchabanenko in the lichen checklist of the southern Russian Far East (Tchabanenko, 2002), counting 943 species. In subsequent years several dozens of publications about lichens of this area appeared (e. g., Joneson, 2004; Pchelkin, 2008; Velikanov, Skirina, 2012; Ezhkin, Galanina, 2014; Zhurbenko, 2014; Skirina, 2015, 2016; Tchabanenko et al, 2018; Konoreva et al, 2020; Chesnokov, Konoreva, 2021; Davydov et al, 2021a, b; Skirina, 2021; Skirina et al, 2021).
The lichens of the Amur Region are still investigated insufficiently, and therefore their diversity is not properly estimated. Up to date, most researchers have investigated lichens within three nature reserves — Norsky, Khingansky, and Zeysky. The lists of Norsky and Khingansky reserves counted 308 (Pchelkin, Pchelkina, 2013) and 63 (Mikulin, 1998) lichen species, respectively. Some results of lichenological investigations of Zeysky Reserve were published in the last decades of 20th century (Petelin et al, 1981; Tolpysheva et al, 1981; Stetsura, 1986; Makryi, Stetsura, 1987; Tolpysheva, Zhiryakova, 1988, 1990; Guseva et al, 1993). The list of lichens compiled on the base of these publications counted 51 species only (Urbanavichus, Urbanavichene, 2004). In 2017, we added 38 species of lichens and lichenicolous fungi to the list of Zeysky Reserve (Kuznetsova, Dudov, 2017), so till now it had included 89 species lichens and lichenicolous fungi. The aim of this paper is to improve our knowledge about lichen diversity of the Amur Region.
The Stanovoy Range extends along the border between the Amur Region and the Republic of Sakha (Yakutia) and represents the watershed of the Amur and Lena rivers. The southeastern part of the range, where the material was collected, is called the Toko-Stanovik (Tokinsky Stanovik in Russian). This area includes the highest part of the range with absolute heights up to 2000 m a. s. l. and relative differences between the tops of the ridges and the valleys bottoms exceeding 1000 m. There is a large variety of rocks of different genesis, both acidic and basic composition. Calcareous rock outcrops are common, especially in the area of the Tas-Balagan pass.
Climate of the Toko-Stanovik can be considered as temperate continental, very cold and rather humid. The total annual precipitation ranges from 504 mm to 744 mm (data from CHELSA model, Karger et al, 2017), with maximum in summer season. Winters are cold with average temperatures of -42 to -36 °C, summer temperatures rise to +15-17 °C.
The altitudinal zonation includes three vegetation belts (Borisova et al, 2020). The lower taiga belt is represented by boreal forests with larch [Larix gmelinii (Rupr.) Kuzen.] and spruce [Picea ajanensis (Lindl. et Gord.) Fisch. ex Carr.] in the tree layer. The forest borderline lays on heights of 1300-1400 m a. s. l. depending on slope aspects. Above the taiga belt till 1700 m a. s. l. the subalpine belt is situated with the narrow strip of birch krummholz [Betula lanata (Regel) V. N. Vassil.] and widespread elfin woodlands with Siberian dwarf pine [Pinuspumila (Pall.) Regel]. A distinctive feature of the Stanovoy Range from other mountain systems of the Far East is the presence of subalpine meadow communities above the upper border of the forest. Above 1600-1700 m a. s. l. there is mountain tundra, mainly with the dominance of prostrate
dwarf shrubs and forbs. Its floristic composition and community structure highly depend on the chemistry of the underlying rock.
For conservation of intact high-mountain ecosystems, a regional protected area was established here in 2010, which received the status of the Tokinsky-Stanovoy National Park in 2019.
To the south-west of the Stanovoy Range the Tukuringra Range is situated. It is a mid-mountain range with altitudes ranging from 250 to 1400 m a. s. l. (maximum 1608 m a. s. l.). The range sublatitudinally distributes for approximately 300 km. From east to west, the continentality of the climate increases (and natural complexes change in accordance with it). The climate of the Tukuringra Range is continental but characterized by several monsoon features. Winters are long and severe, with shallow snow cover. Summers are warm and rainy. The average temperature of January is -30.6 °C and one of July is +18.2 °C. The total annual precipitation ranges between 500 mm in foothills to more than 1500 mm in upper belts of mountains. The conditions change with the longitude and turn to colder and more continental in the western part of the range, or warmer and more monsoon in the eastern one (Vasiljev et al., 1985).
The lower belt of the Tukuringra Range ranges from 300 to 1000-1300 m a. s. l. and occupies the major part of the slopes. It is covered by boreal forests or taiga consisting of larch (Larix gmelinii) and spruce (Picea ajanensis) communities. In the eastern part of the range, a belt of hemiboreal forests is distinguished with the participation of manchurian plant species. Especially on the warmest southern positions hemiboreal larch and oak-black birch (Quercus mongolica Fisch. ex Ledeb., Betula davurica Pall.) complexes are represented. The subalpine belt above 1000-1200 m a. s. l. is defined by Siberian dwarf pine elfinwoods. The dwarf shrub-lichen mountain tundra develop on the flattened peaks of the mountains above 1400 m a. s. l. The western part of the Tukuringra Range is characterized by a severe and more continental climate. Floris-tically poor larch sparse forests are widely distributed there. Spruce forests occupy much smaller areas than in the eastern part of the range.
Zeya Nature Reserve is situated in the Zeya District, in the eastern part of the Tukuringra Range. This area has been protected since 1963.
Materials and Methods
Material was collected in the Amur Region during several expeditions (Fig. 1) in 2018-2021 in the frame of the vegetation surveys. On the Stanovoy Range in 2018, in order to take inventory of the biota of the national park planned for establishment, a hiking route was passed with a caravan of reindeers from the Anachan River to the Zeya with subsequent rafting to the Zeya reservoir. S. V. Dudov collected the material in the valley of the Tok River (sampling location 1), on the volcanic plateau in the upper reaches of the Uliagir and Inarogda rivers (locations 2-5), in the highlands on the watershed of the Tuksani and Sivaktylyak Pervyi rivers (locations 6-7), in the lower reaches of the Sivaktylyak Pervyi River and in the valley of the Zeya River (locations 8-10).
In August 2020, O. I. Ryabenko worked at the Bolshiye Tuksani-Zeya mountain pass (location 25). In July 2021, Dudov collected the herbarium at the Tas-Balagan mountain pass during radial routes (locations 11-20, 26, 27). In July — August 2021 Rya-benko collected lichens in the Zeya Valley during rafting down the Zeya River from the Sivaktylyak Pervyi River to the mouth of the Bolshoy Okonon River (locations 21-24, 29-34). In July 2019, Dudov and Ryabenko collected material in the hard-to-reach western part of the Tukuringra Range (locations 35-38), which had not been previously visited by botanists. The valley of the Pravaya Tynda River with tributaries was examined and an excursion was made to the highest point of the range — the Mount Tukuringra (1608 m a. s. l.). Several specimens cited in this report were collected by Ryabenko in the Zeya Nature Reserve (locations 28, 29) and in its vicinity.
The material was identified by E. S. Kuznetsova. Dissecting microscope, light microscope, and UV light cabinet were used for the identification of lichen specimens. Chroma-tography was performed by I. S. Stepanchikova according to the standard techniques of high performance thin-layer chromatography using solvent systems A, B, and C (Orange et al, 2001). Geographical coordinates are given in the coordinate system WGS 1984.
Sampling locations (Fig. 1)
Amur Region, Zeya District, Stanovoy Range, Tokinsko-Stanovoy National Park: 1 — right side of the Tok River valley, 4.5 km above the inflow of the Uliagir River, 55°39'14.6"N, 129°39'41.3"E, 1032 m a. s. l., base-rich outcrops, 10 VII 2018, Dudov; 2 — upper stream of the Inarogda River, 55°41'57.9"N, 129°58'21.9"E, 1420 m a. s. l., dwarf birch community, 13 VII 2018, Dudov; 3 — ibid, 55°41'39.2"N, 129°57'38.8"E, 1511 m a. s. l., screes of big rock fragments with lichens, 14 VII 2018, Dudov; 4 — watershed of the Inarogda and Uliagir rivers, 55°42'18.4"N, 129°58'08.2"E, 1460 m a. s. l., sparse larch-spruce woodland, 15 VII 2018, Dudov; 5 — volcanic plateau in the upper reaches of the Inarogda River, 55°42'02"N, 130°00'38"E, 1483 m a. s. l., sparse larch woodland, 16 VII 2018, Dudov; 6 — watershed of the Bolshie Tuksani and Sivaktylyak Pervyi rivers, 55°44'03.5"N, 130°31'19.3"E, 1572 m a. s. l., forb meadow on snowbed, 25 VII 2018, Dudov; 7 — ibid, 55°44'00.0"N, 130°31'14.4"E, 1585 m a. s. l., dwarf shrubs tundra, 25 VII 2018, Dudov; 8 — valley of the Zeya River, above the inflow of the Sivaktylyak Pervyi River to the Zeya River, 55°37'42.4"N, 130°42'08.4"E, 802 m a. s. l., river terrace, 28 VII 2018, Dudov; 9 — ibid., willow shrubs on the river bank, 28 VII 2018, Dudov; 10 — lower reaches of the Syvaktyliak Pervyi River, 55°38'09"N, 130°40'28"E, 833 m a. s. l., rock cliffs, 29 VII 2018, Dudov; 11 — range above the cirque to the W of the Tas-Balagan pass, 55°44'02.3"N, 130°48'07.0"E, 1720 m a. s. l., dwarf shrubs tundra, 9 VII 2021, Dudov; 12 — pass Tas-Balagan, 55°44'35.5"N, 130°45'32.7"E, 1851 m a. s. l., dwarf shrubs tundra, 14 VII 2021, Dudov; 13 — vicinity of the Tas-Balagan pass, 55°44'15.2"N, 130°46'32.6"E, 1355 m a. s. l., spruce forest with Siberian dwarf pine, 17 VII 2021, Dudov; 14 — range above the cirque to the W of the Tas-Balagan pass, 55°43'36.9"N, 130°48'31.3"E, 2014 m a. s. l., dwarf shrubs tundra, 18 VII 2021, Dudov; 15 — ibid, 55°43'35.7"N, 130°48'39.2"E, 2012 m a. s. l., dwarf shrubs tundra, 19 VII 2021, Dudov; 16 — vicinity of the Tas-Balagan pass, 55°44'31.5"N, 130°46'11.3"E, 1549 m a. s. l., grass and dwarf shrubs tundra, 23 VII 2021, Dudov; 17 — Tas-Balagan pass, 55°44'35.9"N, 130°46'33.4"E, 1469 m a. s. l., dwarf shrubs tundra, 25 VII 2021, Dudov; 18 — ibid, 55°44'31.5"N, 130°46'26.1"E, 1475 m a. s. l., tundra peat bog, 25 VII 2021, Dudov; 19 — vicinity of the Tas-Balagan pass, 55°44'22.0"N, 130°46'44.3"E, 1459 m a. s. l., tundra peat bog, 25 VII 2021, Dudov; 20 — ibid, 55°44'30.1"N, 130°47'26.1"E, 1589 m a. s. l., tundra peat bog, 26 VII 2021, Dudov; 21 — left bank of the Zeya River, 350 m above the inflow of the Sivaktylyak Pervyi River, 55°37'42.3"N, 130°42'22.5"E, 780 m a. s. l., larch forest, 29 VII 2021, Ryabenko; 22 —ibid, 55°37'39.2"N, 130°42'07.3"E, 790 m a. s. l., sparse woodland,
Fig. 1. Sampling locations (white circles). Thick gray lines indicate the boundaries
of protected areas.
30 VII 2021, Ryabenko; 23 — left side of the Zeya River valley, 4 km downstream of the Urum River inflow, 55°32'34.1"N, 130°39'30.8"E, 792 m a. s. l., larch forest, 2 VIII 2021, Ryabenko; 24 — left bank of the Zeya River, 4.2 km downstream of the Urum River inflow, 55°32'35.7"N, 130°38'23.2"E, 756 m a. s. l., sparse larch woodland, 2 VIII 2021, Ryabenko; 25 — vicinity of the Zeya River source, 55°44'46.6"N, 130°34'39.1"E, 1625 m a. s. l., dwarf-shrubs and lichen tundra, 3 VIII 2020, Ryabenko; 26 — pass Tas-Balagan, 55°44'41.4"N, 130°46'12.2"E, 1546 m a. s. l., dwarf shrubs tundra, 6 VII 2021, Dudov; 27 — vicinity of the pass Tas-Balagan, 55°44'38.2"N, 130°46'16.6"E, 1552 m a. s. l., dwarf shrubs tundra, 7 VII 2021, Dudov. Zeya District, Zeysky Nature Reserve: 28 — Tukuringra Range, hilltop above the cordon "Goltsy", 54°07'29.0"N, 126°55'53.9"E, 1293 m a. s. l., mountain tundra, 6 VIII 2019, Ryabenko; 29 — south-eastern slope of cape between the Rudenko and Gilui rivers, 54°00'58.9"N, 127°27'23.3"E, 303 m a. s. l., screes of big rock fragments, mixed forest, 8 VII 2021, Ryabenko. Zeya District, Stanovoy Range: 30 — right bank of the Karaurakkan River, 30 m above the inflow to the Zeya River, 55°26'45.2"N, 130°35'44.7"E, 686 m a. s. l., sparse larch forest, 4 VIII 2021, Ryabenko; 31 — valley of the Zeya River, 730 m to the NW of the Karaurakkan River inflow, 55°27'00.1"N, 130°35'13.5"E, 740 m a. s. l., larch forest, 4 VIII 2021, Ryabenko; 32 — right bank of the Zeya River, 150 m downstream of the Karaurakkan River inflow, 55°26'40.5"N, 130°35'45.4"E, 736 m a. s. l., larch forest, 4 VIII 2021, Ryabenko; 33 — valley of the Zeya River between boggy billabong and the Karaurakkan River, 55°26'47.1"N, 130°35'38.0"E, 691 m a. s. l., larch forest, 5 VIII 2021, Ryabenko; 34 — right bank of the Zeya River, 850 m below of the Okonon River inflow, 55°19'18.6"N, 130°23'21.7"E, 619 m a. s. l., spruce forest, 7 VIII 2021, Ryabenko. Tynda District, Tukuringra Range: 35 — left bank of the Pravaya Tynda River, 6 km above the mouth, 54°23'19.8"N, 125°37'00.8"E, 803 m a. s. l., spruce forest, 17 VII 2019, Dudov; 36 — summit surface of Mount Tukuringra (1608 m a. s. l.), 54°26'24.3"N, 125°28'14.7"E, 1499 m a. s. l., lichen dwarf-shrub tundra, 19 VII 2019, Dudov; 37 — left bank of the Pravaya Tynda River, 8 km above the mouth, 54°23'35.7"N, 125°36'08.1"E, 899 m a. s. l., sparse larch woodland, 22 VII 2019, Dudov; 38 — valley of the Shatamy River, 54°19'28.6"N, 125°36'52.7"E, 843 m a. s. l., peat bog, 26 VII 2019, Dudov.
Results and Discussion
In the annotated species list presented below the following symbols and abbreviations are used: # — lichenicolous fungi, !!! — new species to the Russian Far East, !! — new species to the southern Russian Far East, ! — new species to the Amur Region. The species in the annotated list are accompanied by the data on their sampling locations, substrates, herbaria numbers, and distribution.
Specimens are deposited in the herbarium of the Laboratory of Lichenology and Bryology, Komarov Botanical Institute RAS (LE). The nomenclature of the cited taxa mainly follows recently published checklists for Scandinavia (Westberg et al., 2021) and the continental United States and Canada (Esslinger, 2021).
!#Abrothallus peyritschii (Stein) Kotte — 23, on thallus of Vulpicida pinastri (Scop.) J.-E. Mattsson et M. J. Lai on bark of Larixgmelinii, LE L-17867; 30, on thallus of V. pinastri on bark, LE L-17868. Distribution in the southern Russian Far East: Sakhalin Region (Zhurben-ko, Ohmura, 2019).
!!#Arthrorhaphis aeruginosa R. Sant. et Tonsberg — 26, on thallus of Cladonia sp. on soil, LE L-17869; 36, on thallus of C. cervicornis (Ach.) Flot. on soil, LE L-17870. Distribution in the Russian Far East: Kamchatka Territory (Himelbrant et al, 2021).
!Bryocaulon divergens (Ach.) Karnefelt — 12, on soil, LE L-17871; 15, on soil, LE L-17872. Distribution in the southern Russian Far East: Khabarovsk and Primorye territories (Tcha-banenko, 2002; Velikanov, Skirina, 2012; Skirina, 2016), Sakhalin Region (Sakhalin I.) (Skirina et al, 2021).
!Calicium trabinellum (Ach.) Ach. — 23, on bark of Larixgmelinii, LE L-17873. Distribution in the southern Russian Far East: Khabarovsk (Vekikanov, Skirina, 2012) and Primorye (Skirina, 2016) territories, Sakhalin Region (Sakhalin and Kunashir islands) (Tchabanenko, 2002; Skirina et al., 2021).
!Catillaria nigroclavata (Nyl.) Schuler — 4, on bark of Picea ajanensis, LE L-17874. Distribution in the southern Russian Far East: Khabarovsk and Primorye territories (Tchabanenko, 2002).
!Cetraria odontella (Ach.) Ach. — 21, on primary soil, LE L-17875. Distribution in the southern Russian Far East: Khabarovsk and Primorye territories (Tchabanenko, 2002).
!Cladonia kanewskii Oxner — 6, on soil, LE L-17876. Distribution in the southern Russian Far East: Jewish Autonomous Region (Skirina, 2015), Khabarovsk Territory, Sakhalin Region (Sakhalin I.) (Tchabanenko, 2002; Skirina et al, 2021).
!C. maxima (Asahina) Ahti — 13, on soil, LE L-17878; 38, on soil, LE L-17879. Distribution in the southern Russian Far East: Khabarovsk and Primorye territories, Sakhalin Region (Sakhalin and Paramushir islands) (Tchabanenko, 2002; Velikanov, Skirina, 2012; Skirina et al., 2021).
!C. phyllophora Hoffm. — 12, on soil, LE L-17880; 17, on soil, LE L-17884; 18, on soil, LE L-17883; 21, on soil, LE L-17882. Distribution in the southern Russian Far East: Jewish Autonomous Region (Skirina, 2015), Khabarovsk and Primorye territories (Tchabanenko, 2002), Sakhalin Region (Sakhalin I.) (Tchabanenko et al, 2018).
!C. straminea (Sommerf.) Flörke — 25, on soil, LE L-17885; 37, on soil, LE L-17886. Distribution in the southern Russian Far East: Primorye Territory and Sakhalin Region (Sakhalin and Kunashir islands) (Tchabanenko, 2002).
!C. strepsilis (Ach.) Grognot — 28, on soil, LE L-17887. Distribution in the southern Russian Far East: Primorye Territory (Tchabanenko, 2002).
!C. stygia (Fr.) Ruoss -15, on soil, LE L-17889; 20, on soil, LE L-17890; 25, on soil, LE L-17888. Distribution in the southern Russian Far East: Sakhalin Region (Sakhalin I.) (Ko-noreva et al., 2020).
!C. trassii Ahti — 19, on soil, LE L-17891. Distribution in the southern Russian Far East: Jewish Autonomous Region (Skirina, 2021).
!!C. uliginosa (Ahti) Ahti — 31, on soil, LE L-17892. Distribution in the Russian Far East: Kamchatka Territory (Himelbrant et al., 2014).
Coccocarpia erythroxyli (Spreng.) Swinscow et Krog — 15, on soil, LE L-17926; 29, on stone. New to the Tokinsko-Stanovoy National Park. The species is under protection in the Amur Region (Krasnaya..., 2009).
C. palmicola (Spreng.) Arv. et D. J. Galloway — 21, on stone. New to the Tokinsko-Stano-voy National Park. The species is under protection in Russian Federation (Krasnaya., 2008) and in the Amur Region (Krasnaya., 2009).
!Collema furfuraceum (Arnold) Du Rietz — 8, on limestone, LE L-15915. Distribution in the southern Russian Far East: Jewish Autonomous Region (Skirina, 2015), Khabarovsk and Primorye territories (Tchabanenko, 2002), Sakhalin Region (Kunashir I.) (Ezhkin, 2020).
!Dactylina arctica (Richardson) Nyl. — 15, on soil, LE L-17893. Distribution in the southern Russian Far East: Khabarovsk Territory and Sakhalin Region (Sakhalin I.) (Tchabanenko, 2002).
!!D. ramulosa (Hook.) Tuck. — 12, on plant debris and soil, LE L-17894. Distribution in the Russian Far East: Chukotka Autonomous Area (Andreev et al., 1996) and Kamchatka Territory (Himelbrant et al, 2014).
! Diploschistes scruposus (Schreb.) Norm. — 21, on stone, LE L-17895. Distribution in the southern Russian Far East: Primorye Territory (Tchabanenko, 2002).
!!! Fulgidea oligospora (Timdal) Bendiksby et Timdal — 33, on burned lignum, LE L-17896. Thallus contains thamnolic and alectorialic acids. In Russia it is known from the Republic of Yakutia (Timdal, 2001). The species is morphologically close to Xylopsora friesii (Ach.) Bendiksby et Timdal, and can hardly be recognized without chemical studies. Thallus of X. friesii contains friesic acid only (Timdal, 2001; Himelbrant, Urbanavichus, 2008).
!Hypogymnia subobscura (Vainio) Poelt — 28, on soil, LE L-17897. Distribution in the southern Russian Far East: Khabarovsk and Primorye territories (Tchabanenko, 2002), Sakhalin Region (Sakhalin I.) (Konoreva et al., 2020).
!!Lecanora subintricata (Nyl.) Th. Fr. — 23, on bark of Larix gmelinii, LE L-17898; 33, on burned lignum, LE L-17899. Distribution in the Russian Far East: Kamchatka Territory (Himelbrant et al., 2014).
!!Lecidea turgidula Fr. — 23, on bark of Larix gmelinii, LE L-17900. Distribution in the Russian Far East: Kamchatka Territory (Himelbrant et al., 2014).
!#Lichenostigma maureri Hafellner — 35, on thallus of Usnea longissima Ach. on twigs of Larix gmelinii, LE L-17901. Distribution in the southern Russian Far East: Khabarovsk and Primorye territories (Zhurbenko, 2014).
!Melanelia stygia (L.) Essl. — 2, on stone, LE L-15922; 3, on stone, LE L-17927. Distribution in the southern Russian Far East: Jewish Autonomous Region (Skirina, 2015), Khabarovsk and Primorye territories, Sakhalin Region (Sakhalin I.) (Tchabanenko, 2002; Skirina, 2016).
Nephromopsis laureri (Kremp.) Kurok. — 24, on bark; 30, on bark of coniferous tree. New to the Tokinsko-Stanovoy National Park. The species is under protection in Russian Federation (Krasnaya..., 2008)
!Ochrolechia frigida (Sw.) Lynge — 14, on soil, LE L-17904; 28, on soil, LE L-17903. Distribution in the southern Russian Far East: Jewish Autonomous Region (Skirina, 2015), Primorye Territory, and Sakhalin Region (Sakhalin I.) (Tchabanenko, 2002; Skirina, 2016; Skirina et al., 2021).
!O. inaequatula (Nyl.) Zahlbr. — 12, on soil, LE L-17905. Distribution in the southern Russian Far East: Primorye Territory (Skirina, 2016).
!Parmelia cochleata Zahlbr. — 21, on stone, LE L-17906; 23, on stone, LE L-17907. Distribution in the southern Russian Far East: Primorye Territory (Skirina, 2016).
!!!Peltigera frippii Holt.-Hartw. — 10, on rock cliffs, LE L-15920. In Russia it is known from the Murmansk Region (Urbanavichus et al., 2008), Republic of Komi (Plusnin, 2007), Krasnoyarsk (Zhurbenko, 1996) and Trans-Baikal (Chesnokov et al, 2015) territories. It is easily distinguished by smooth, maculate upper surface, crisp appearance, and fan-shaped veins (Vitikainen, 2007).
!! Placidiopsis pseudocinerea Breuss — 22, on mossy rock, LE L-17909. Distribution in the Russian Far East: Kamchatka Territory (Himelbrant et al., 2021).
!Placynthiella icmalea (Ach.) Coppins et P. James — 23, on plant debris, LE L-17911; 24, on upturned roots, LE L-17912; 29, on mossy soil, LE L-17913; 30, on lignum, LE L-17914; 32, on lignum, LE L-17915. Distribution in the southern Russian Far East: Jewish Autonomous Region (Skirina, 2015) and Primorye Territory (Skirina, 2016).
!!P. uliginosa (Schrad.) Coppins et P. James — 24, on soil, LE L-17916 Distribution in the Russian Far East: Chukotka Autonomous Area (Andreev et al., 1996), Magadan Region (Zheludeva, Makryi, 2018), and Kamchatka Territory (Himelbrant et al., 2019).
!Polyblastidium japonicum (M. Sato) Kalb — 34, on bark of coniferous tree, LE L-17917. Distribution in the southern Russian Far East: Jewish Autonomous Region (Skirina, 2015), Primorye Territory, and Sakhalin Region (Kunashir I.) (Tchabanenko, 2002).
!!!Psora globifera (Ach.) A. Massal. — 5, on soil, LE L-15916; 8, on calciferous boulder, LE L-15911. In Russia it is known from Republic of Buryatia (Urbanavichene, Urbanavichus,
2009), Krasnoyarsk Territory (Zhurbenko, 2021), Republic of Yakutia (Poryadina, 2005), and Trans-Baikal Territory (Makry, 2005). The species can be confused with Romjularia lurida (Ach.) Timdal which differs in having flattened to weakly convex apothecial discs, with a more persistent exciple darker than the disc (Timdal, 2007).
!!Psorula rufonigra (Tuck.) Gotth. Schneid. — 8, on limestone, LE L-15919. Distribution in the Russian Far East: Magadan Region (Zhurbenko, 2003).
!R. thrausta (Ach.) Nyl. — 4, on twigs of Picea ajanensis, LE L-17918. Distribution in the southern Russian Far East: Khabarovsk Territory (Yakovchenko et al, 2013) and Sakhalin Region (Sakhalin I.) (Davydov et al, 2021b).
!!!Rhizoplaca subdiscrepans (Nyl.) R. Sant. — 1, on rocks, LE L-15891. In Russia it is known from Republic of Karelia (Fadeeva et al, 2007), Republic of Dagestan (Ismailov, Urba-navichus, 2018), Orenburg Region (Merkulova, Urbanavichus, 2006), Altay Territory (Davydov, 2011), and Irkutsk Region (Makryi, 2008). It can be distinguished from R. chrysoleuca (Sm.) Zopf and R. melanophthalma (DC.) Leuckert et Poelt by having numerous and crowded, distinctly convex to plicate and at most vaguely umbilicate thallus subunits, with lower surface lacking greenish or bluish black tinges (Ryan, 2002).
!Solorina saccata (L.) Ach. — 9, on limestone, LE L-15904; 22, on mossy rock, LE L-17920. Distribution in the southern Russian Far East: Primorye Territory and Sakhalin Region (Sakhalin I.) (Tchabanenko, 2002).
!Sphaerophorus fragilis (L.) Pers. — 3, on boulders, LE L-15884; 12, on soil, LE L-17921. Distribution in the southern Russian Far East: Khabarovsk and Primorye territories, Sakhalin Region (Sakhalin I.) (Tchabanenko, 2002).
!!Spilonema revertens Nyl. — 1, on rocks, LE L-15907. Distribution in the Russian Far East: Magadan Region (Zhurbenko, 2003).
!Stereocaulon subcoralloides (Nyl.) Nyl. — 1, on rocks, LE L-15881. Distribution in the Russian Far East: Khabarovsk and Primorye territories, Sakhalin Region (Sakhalin, Iturup, and Kunashir islands) (Tchabanenko, 2002).
!Vulpicida tilesii (Ach.) J.-E. Mattsson et M. J. Lai — 11, on soil, LE L-17922; 16, on soil, LE L-17923; 26, on soil, LE L-17924; 27, on soil, LE L-17925. Distribution in the southern Russian Far East: Khabarovsk Territory (Tchabanenko, 2002).
Of 44 species presented here, five are recorded for the first time for the Russian Far East, eight — for the southern Russian Far East, and 28 — for the Amur Region. Four and 38 species are new to the Zeya Reserve and Tokinsko-Stanovoy National Park, respectively. In total, three protected lichen species were found. Coccocarpia palmi-cola and Nephromopsis laureri are protected in Russian Federation (Krasnaya..., 2008), whereas Coccocarpia erythroxyli and C. palmicola are protected in the Amur Region (Krasnaya., 2009). Nephromopsis laureri is recommended for inclusion in the next edition of the Red Data Book of the Amur Region.
According to available publications, the list of lichens and allied fungi of the Amur Region hardly exceeds 450 species (Petelin et al., 1981; Tolpysheva et al., 1981; Stet-sura, 1986; Makryi, Stetsura, 1987; Tolpysheva, Zhiryakova, 1988, 1990; Guseva et al, 1993; Mikulin, 1998; Urbanavichus, Urbanavichene, 2004; Pchelkin, Pchelkina, 2013; Kuznetsova, Dudov, 2017). Taking into account the new data, about 500 species are now known for the Amur Region. For a more precise assessment of the inventory of
lichen biota of the Amur Region and definition of exact number of species, a revision of the herbarium collections is necessary.
Most of the species listed here for the first time for the Amur Region are widespread in the southern Russian Far East and are common components of tundra communities (e. g., Bryocaulon divergens and Ochrolechia frigida). This indicates a low level of knowledge of the regional lichen biota.
The group of lichenicolous fungi traditionally treated with lichens is purely studied in the Amur Region. At present, only four species are known, three of them are cited in this article. Obviously, this ecological group requires further investigation. Most of the species presented here are foliose and fruticose lichens collected mainly on soil and rocks in tundra and taiga communities. The study of all types of substrates, primarily various phorophytes, and a detailed study of crustose species is likely to significantly expand our knowledge of the lichen biota of the region. So, it can be expected that lichen list of the Amur Region will increase significantly in case of comprehensive studies.
Acknowledgments
We express our gratitude to the administration of the Zeya State Nature Reserve represented by S. Yu. and E. V. Ignatenko, and S. A. Podolsky for support of this study. I. S. Stepanchikova is thanked for help with performing of chromatography. The authors also thank the reviewers and editors for valuable comments and corrections. The study of E. S. Kuznetsova was carried out within the framework of the institutional research project "Flora and systematics of algae, lichens, and bryophytes of Russia and phytogeographically important regions of the world" (no. 121021600184-6) of the Komarov Botanical Institute RAS. The study of S. V. Dudov was supported by the Lomonosov Moscow State University research project 121032500090-7.
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