CC BY
23Four rare liverwort species: distribution, ecology, taxonomy
E. V. Sofronova1, A. D. Potemkin2
'Institute for Biological Problems of Cryolithozone, SB RAS, Yakutsk, Russia 2Komarov Botanical Institute, Russian Academy of Sciences, St. Petersburg, Russia Corresponding author: E. V. Sofronova, [email protected]
Abstract. Distribution, ecology and taxonomy of four rare liverwort species Frullania davurica, Leje-unea alaskana, Marchantia romanica, Scapania sphaerifera, which were recorded many times in collections from the Republic of Sakha (Yakutia), are compiled and analyzed. Worldwide distribution maps of Lejeunea alaskana, Marchantia romanica, Scapania sphaerifera are provided. Taxonomic status of all four species needs to be tested on the basis of molecular studies of materials through their ranges. Sporophytes of Lejeunea alaskana are described for the first time.
Keywords: Frullania davurica, Lejeunea alaskana, Marchantia romanica, Scapania sphaerifera, Marchantiophyta, habitat, range, Eastern Verkhoyanje Mountains, Republic of Sakha (Yakutia), Russia, Siberia.
Четыре редких вида печеночников: распространение, экология, таксономия
Е. В. Софронова1, А. Д. Потемкин2
'Институт биологических проблем криолитозоны СО РАН, Якутск, Россия 2Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия Автор для переписки: Е. В. Софронова, [email protected]
Резюме. Обобщены и проанализированы данные по распространению, экологии и таксономии Frullania davurica, Lejeunea alaskana, Marchantia romanica, Scapania sphaerifera, неоднократно выявленных в сборах из Республики Саха (Якутия). Приведены карты распространения в мире Le-jeunea alaskana, Marchantia romanica, Scapania sphaerifera. Таксономический статус всех четырех видов требует молекулярных исследований по материалам, собранным на протяжении всего ареала. Впервые описаны шорофиты Lejeunea alaskana.
Ключевые слова: Frullania davurica, Lejeunea alaskana, Marchantia romanica, Scapania sphaerifera, Marchantiophyta, ареал, местообитания, Восточное Верхоянье, Республика Саха (Якутия), Россия, Сибирь.
Distribution of Holarctic liverwort species is still insufficiently studied. Extensive territories of the Russian Far East, Siberia as well as southern adjacent areas still need careful exploration. The extensive liverwort studies in the Russian Far East and Siberia revealed in the last decades a number of new to science, previously unknown or poorly known liverwort species to Russia (e.g., Potemkin, 2000, 2003, 2004; Bakalin, Vilnet, 2009; Bakalin et al, 2012; Mamontov, Vilnet, 2013; Potemkin, Mamontov, 2013; Sofronova et al, 2013; Potemkin et al., 2015; Mamontov et al., 2018). Their discoveries in Russia and subsequent records have enhanced our knowledge regarding their distribution and origin. Among such species
https://doi.org/10.31111/nsnr/2018.52.2.505
505
first recorded for Russia in the last century or described from Russia are Frullania davurica, Lejeunea alaskana, Marchantía romanica and Scapania sphaerifera. For a long time they were known in Russia from only a few locations.
Ongoing field explorations and the acquisition of collections in the Eastern Verkhoyanje Mountains, Republic of Sakha (Yakutia), resulted in the discovery of new locations. The goal of this present study is to provide generalized additional information on the distribution and ecology of these four liverwort species and to discuss briefly their taxonomy.
Frullania davurica Hampe, 1845, Syn. Hepat.: 422. = F. davurica subsp. jackii (Gottsche) S. Hatt., 1976, Bull. Natl. Sci. Mus., Tokyo, B 2: 21.
Frullania davurica was described from Dahuria (Gottsche et al., 1844) without certain definition of locality. Dahuria is located within present boundaries of the Republic of Bury-atia, Trans-Baikal Territory, Amur Region of Russia. At present F. davurica is known from a number locations in Russia and listed below adjacent Asian countries.
F. davurica is closely related to European F. jackii Gottsche. Distinction of F. davurica from F. jackii at the species level was confirmed by Hentschel et al. (2009) on the basis of two specimens from the Russian Far East and Switzerland respectively. Confirmation of their distinction needs more extensive material obtained through their distributional range.
Ecology. On shaded rocks in forest belt usually in pure mats on rocks, with ?neutral to basic pH.
Distribution. Russia: from West Siberia to the Russian Far East; China (Piippo, 1990; Zhu, 2006); Korea (Choi, 2013); Japan: Hokkaido, Honshu, Shikoku, Kyushu (Yamada, Iwatsuki, 2006). In Russia it is known from southern taiga to forest steppe and from mountain taiga to mountain deciduous coniferous forests (Potemkin, Sofronova, 2009).
Until recently F. davurica was unknown from the Republic of Sakha but it was recorded from West and South Siberia, and the Russian Far East (Potemkin, Sofronova, 2009). In 2010 it was found in South Yakutia from the Aldan Upland (Sofronova, 2017). Report of F. davurica from Upper Kuranakh River, Suntar-Khayata Range (25 VIII 2017, Sofronova, SASY), is northernmost in the world.
Selected specimens examined: Russia, Republic of Sakha (Yakutia), Eastern Yakutia, Sun-tar-Khayata Range, Upper Kuranakh River, 62°48'47.3'N, 138°55'59.0"E, 1048 m a. s. l., small pure continuous cover (less than 100 cm2) on rather dry north-west faced rocky outcrop on brook bank in forest belt, 25 VIII 2017, Sofronova, SASY; South Yakutia, Aldan Upland, Timpton River Basin, Kurung-Khonku River Mouth, 57°30N, 126°33', 650 m a. s. l., large pure continuous cover (up to 1 m2) on rather dry stone under ledge of north-west facing cliff (shaded place) in forest belt, 16 VIII 2010, Sofronova, SASY, LE; Trans-Baikal Territory (former Chita Region), Sakhondinsky Biosphere Reserve, Ende River Valley, 49°27' N, 110°50'E, 1161 m a. s. l., on large boulders at base of slope, 15 VII 2010, O. M. Afonina, LE; Sakhalin Region, Kuril Islands, Shikotan Island, Aerodromnaya Bay, 43°48' N, 146°50'E, coastal slopes, on rocks and turf-covered stony deposits, 10 IX 2006, T. I. Nyushko 15.1-06, LE.
Lejeunea alaskana (R. M. Schust. et Steere) Inoue et Steere, 1978, J. Hattori Bot. Lab. 44: 330. = Hygrolejeunea alaskana R. M. Schust. et Steere, 1958, Bull. Torrey Bot. Club 85(3): 190-196, f. 1. 1958. (Fig. 1)
Fig. 1. Global distribution of Lejeunea alaskana.
Lejeunea alaskana is still insufficiently known in Russia. The actual first collection of this species for Russia was mistakenly listed as L. cavifolia (Ehrh.) Lindb. (Sofronova, 2003). In 2012 it was recorded for Russia for the first time from the Magadan Region, the Russian Far East (Bakalin et al., 2012). L. alaskana differs from L. cavifolia in mostly larger ovoid underleaves, 0.4-0.7 vs. 0.3-0.4 the dorsal lobe, with often ± closed very narrow even slit-like (vs. open with divergent sides) more shallow sinus 0.25-0.4 (0.45) vs. 0.3-0.5 the length. The shape of underleaves and sinus apparently has primary importance for differentiation of these species. Difference in the size of plants of L. alaskana and L. cavifolia, which was listed as the primary distinctive character in the key by Bakalin et al. (2012), overlaps in these species and generally cannot be used to separate them.
In collections from Yakutia (30 VIII 2017, Sofronova, SASY ), plants with mature capsules have been found for the first time. Mature perianth is on a short, often longly stipitate and fragile branch, subtended by 1-2 pairs of bracts. Recorded androecia are variable in position: 2-3 pairs of male bracts on short intercalary branches below perianth, up to 6 pairs of bracts below perianth or up to 20 pairs on main shoot. Spores are typical for the genus, large, deep green, ca. 35-50 x 40-50(75) ^m, with regularly papillose surface with papillae ca. 1 ^m in diam. Seta cross section with 12 outer cell rows and 4 inner cell rows similar in size to inner cells.
Ecology. Lejeunea alaskana is recorded at 700-1250 m a. s. l. in Yakutia to 2000 m a. s. l. in Buryatia (Tubanova et al., 2017), mostly in the tundra belt with a few records from the lower altitudes in the forest belt. Ecological requirements of L. alaskana are quite distinctive. It grows on soil over rocks and usually associates with characteristic tundra com-
munities with calciphilous Odontoschisma macounii (Austin) Underw., Scapania simmonsii Bryhn et Kaal., Frullania subarctica Vilnet, Borovich. et Bakalin as well as with neutrophilous Blepharostoma trichophyllum (L.) Dumort. and Sphenolobus minutus (Schreb.) Berggr. Such association differs from that of Lejeunea cavifolia, which usually occurs together with acidophilous and neutrophilous epiphytic or saxicolous species characteristic of forest communities.
Distribution. Russia: Republic of Sakha (Yakutia) (Sofronova, 2003, as L. cavifolia; Sofronova, Kopyrina, 2016; present report), Krasnoyarsk Territory (Fedosov et al, 2015), Republic of Buryatia (Tubanova et al, 2017), Trans-Baikal Territory (Afonina et al., 2013), Magadan Region (Bakalin et al, 2012), Khabarovsk Territory (Bakalin, 2015); USA: Alaska (Schuster, Steere, 1958; Steere, Inoue, 1978); Canada: Northwest Territories and Yukon (Hong, Vitt, 1977; Steere, Scotter, 1978). Arctic-mountain species.
Yakutian records of Lejeunea alaskana from the Arkchan Plateau (21 VIII 2005, A. P. Isaev, SASY; 30 VIII 2014, L. I. Kopyrina, SASY) are northernmost in Russia.
Conservation. Included in Red Data Book of Yakutia (Krasnaya..., 2017), category 36 — rare species with sporadic scattered small populations through the extensive range.
Selected specimens examined: Russia, Republic of Sakha (Yakutia), North-Eastern Yakutia, Arkachan Plateau, Middle Fedor-Uryage River, near the confluence of the Mangazeyka River, 65°42N, 130°09'E, ~ 1000 m a. s. l., dwarf shrubs-lichen-moss Larix forest, on soil, few plants and minute continuous cover (up to 1 cm2) together with Odontoschisma macounii, Scapania simmonsii, 21 VIII 2005, A. P. Isaev, SASY; ibidem, Middle Sirelende River, near the site Vertical'nyi, 65.69104°N, 130.05692°E, 1029 m a. s. l., tundra belt, on soil in Rhododendron-lichen tundra, few plants mixed with Frullania subarctica, Ptilidium ciliare, 30 VIII 2014, L. I. Kopyrina, SASY; Eastern Yakutia, Suntar-Khayata Range, Kurbelyakh River Valley near the mouth, 63°07N, 139°04'E, 900 m a. s. l., on rocky outcrop with dwarf shrubs-moss communities near small lake, small pure continuous cover on very wet soil, 13 VII 1999, Sofronova, SASY; ibidem, Kamenny Creek (tributary Vostochnaya Khandyga River), 63°04N, 139°05'E, 1250 m a. s. l., tundra belt, under snowfields on N-facing slope, in small pure continuous cover on moist soil covering stones of stream bank, 25 VII 1999, Sofronova, SASY; Sette-Daban Range, Middle Sakkyryr River, 62°48'13.2"N, 138°24'21.3"E, 1022 m a. s. l., subalpine shrub belt on northern slope, dwarf shrubs-moss Betula divaricata communities, on degrading Ditrichum sp., small continuous cover mixed with Sphenolobus minutus, with mature capsules, 30 VIII 2017, Sofronova, SASY; ibidem, Middle Allakh-Yun River, near the Allakh-Yun Settlement, Sellyakh Creek, 61°06N, 137°58'E, 700 m a. s. l., forest belt, on moist soil of brook bank, few plants mixed with Blepharostoma trichophyllum, Radula complanata, Scapania simmonsii, 29 VIII 2000, Sofronova, SASY; Republic of Buryatia, Yuzhnomuysky Range, Dzherginsky Reserve, 8 km south of source of Dzhirga River, rush tundra, 55°02N, 111°41'E, 2000 m a. s. l., 17 VII 2002, O. A. Anenkhonov Ky-02/36, LE; ibidem, source of Burguzin River, forest-tundra belt, on rocks covered by mosses, 55°07'24.2"N, 111°52'58.2"E, 1798 m a. s. l., 1 VIII 2013, Yu. S. Mamontov 386/2, LE; USA, Alaska, Driftwood camp, near headwaters of the Utukok River, north slope of De Long Mountains, Brooks Range, approx. 68°53N, 161°10'W, wet depression in north facing tundra hillside, 4-16 VIII 1951, W. C. Steere 16895, LE (paratype); ibidem, 68°54'N, 161°20'W, 500-700 m a. s. l., wet tundra on ridges, 11 VIII 1951, W. C. Steere 16782, LE (paratype).
Marchantia romanica (Radian) D. G. Long, Crand.-Stotl., L. L. Forrest et J. C. Villarreal, 2016, Phytotaxa 252(1): 78. = Bucegia romanica Radian, 1903, Bull. Herb. Inst. Bot. Bucarest 3-4: 4. (Fig. 2)
Despite this species was described from Romania and known for a long time from Car-patian Mts in Europe only at the end XX century and beginning of XXI century it became known from many locations in Siberia and western North America (Schljakov, 1973; Po-temkin, 1995; Konstantinova et al, 2014).
Marchantia romanica is a remarkable species of the genus because of specific morphology of the thallus cross section with empty air chambers situated in 1-3 tiers. It was distinguished as the separate genus Bucegia Raddi until recent molecular study of the family Marchantiaceae (Bischler) Lindley (Long et al, 2016). Structure of male and female carpo-cephala resembles that of M. quadrata Scop. [= Preissia quadrata (Scop.) Nees].
Ecology. In tundra and forest belts, usually in rocky niches, on calcareous soil, mostly as few plants. Occasionally associated with Plagiochila porelloides (Torr. ex Nees) Lin-denb., Marchantia quadrata, Tritomaria heterophylla R. M. Schust., Scapania gymnosto-mophila, Eocalypogeia schusteriana (R. M. Schust.) R. M. Schust., Oleolophoziaperssonii H. Buch et S.W. Amell) L. Söderstr., De Roo et Hedd.
Distribution. Norway: Svalbard (Konstantinova et al., 2014); Czech Republic, Slovakia, Poland, Romania (Schumacker, Vana, 2005), Ukraine: Carpathian Mts (Konstantinova et al., 2014); Russia: Krasnoyarsk (Bakalin et al., 2016) and Trans-Baikal (Mamontov, Afonina, 2015) territories, Republic of Sakha (Yakutia), Chukotka, Vrangel Island, Magadan Region; Canada: British Columbia, Alberta (Konstantinova et al., 2014); USA: Alaska (Potemkin, 1995; Konstantinova et al., 2014). Arctic-mountain species.
Conservation. Protected in Russia: category and status 2, species with decreasing population size (Krasnaya..., 2008).
Specimens examined and known records: Russia, Republic of Sakha (Yakutia), Northern Yakutia, Tuora-Sis Range, Sietachan Creek (tributary Lena River), 71°05N, 127°30'E, stone placer between stones on N-facing slope, 03 VII 1956, A. E. Katenin, V. V. Petrovsky (Schljakov, 1973, specimen not found); North-Eastern Yakutia, Orulgan Sis Range, Khobol River (tributary Bytantay River), 67°08N, 128°59'E, 1294 m a. s. l., tundra belt, on moist soil in cracks of rock outcrops on a mountain top, few plants, 22 VI 2006, Sofronova, SASY; ibidem, Sulda Tekeekit River (tributary Undulung River), 67°04N, 128°15'E, 1250 m a. s. l., tundra belt, on moist soil in cracks of rock outcrops on N-facing slope, few plants, 30 VI 2006, Sofronova, SASY; ibidem, Enigan Toloono River (tributary Menkere River), 68°13.707N, 128°07.610'E, 900 m a. s. l., tundra belt, on moist soil covering stones of rock outcrops between shale on E-facing slope near the top of the mountain, few plants, 07 VIII 2011, Sofronova, SASY; Arkachan Plateau, Middle Fedor-Uryage River, near the confluence of the Mangazeyka River, 65°42N -130°09'E, ~ 1000 m a. s. l., forest belt, on soil in niches between the stones of rockfields on northern slope, few plants, 21 VIII 2005, A. P. Isaev, SASY; Eastern Yakutia, Suntar-Khayata Range, Kamenny Creek (tributary Vostochnaya Khandyga River), 63°04'N, 139°05'E, 1250-1400 m a. s. l., tundra belt, on wet stream bank under snowfields among the rubble on N-facing slope, few plants, 25 VII 1999, Sofronova, SASY; ibidem, Upper Kuranakh River, 62°48'48.1"N, 138°55'54.1"E, 1077 m a. s. l.), forest belt, on very wet soil in the grooves of west faced rocky outcrops, few plants, 25 VIII 2017, Sofronova, SASY; Sun-tar-Khayata Range, Knory Creek (tributary Agayakan River), Mus-Khaya Mt. surroundings, ~ 62°30N, 140°58'E, 1880 m a. s. l., tundra belt, on soil near the creek, few plants, 15 VII 2011,M. S. Ignatov, SASY.
Scapania sphaerifera H. Buch et Tuom., 1936, Mem. Soc. Fauna Fl. Fenn. 11: 228. (Fig. 3)
Scapania sphaerifera was described and known from a single collection in the Murmansk Region (Kola Peininsula), Russia (Buch, Tuomikoski, 1936). After its first record in South Siberia (Váña, Soldán, 1985) it was found in numerous localities in Siberia and the Russian Far East. The species is easily distinguished due to characteristic ± coarsely dentate leaves and sphaerical 4-8-celled gemmae (Konstantinova, Potemkin, 1994; Potemkin, Sofronova, 2009). It occupies an isolated taxonomic position within the genus and is distinguished as a separate section Sphaeriferae Müll. Frib. ex Konstant. et Potemkin.
Ecology. Scapania sphaerifera is recorded at 700-1859 m a. s. l., in the forest and tundra belt. It grows usually on acid rocks in sheltered niches and associates with Tetralophozia setiformis (Ehrh.) Schljakov, Sphenolobus saxicola (Schrad.) Steph., S. minutus (Schreb. ex D. Crantz) Berggr., Scapania microdonta (Mitt.) Müll. Frib., S. spitsbergensis (Lindb.) Müll. Frib., S. hyperborea J0rg., Trilophozia quinquedentata (Huds.) Bakalin, Lophoziopsis excisa (Dicks.) Konstant. et Vilnet.
Distribution. Russia: Murmansk Region (Schljakov, 1981; Konstantinova, Potemkin, 1994), Republics of Sakha (Yakutia) (Sofronova et al, 2014, 2015), Altai (Váña, 1991-1992; Schuster, Konstantinova, 1996), Republic of Tuva (Váña, Soldán, 1985), Republic of Buryatia (Konstantinova, Potemkin, 1994; Kazanovsky, Potemkin, 1995; Konstantinova, Savchenko, 2008), Krasnoyarsk Territory (Vasiljev, 1992; Konstantinova, Vasil-jev, 1994; Fedosov et al., 2015), Trans-Baikal Territory (Mamontov et al., 2011), Irkutsk Province (Váña, Soldán, 1985; Krasnaya., 2008, 2010); Khabarovsk Territory (Bakalin, 2009, 2015; Choi et al., 2012), Primorsky Territory, Sakhalin Province, Kamchatka Terri-
Fig. 3. Global distribution of Scapania sphaerifera.
tory, Magadan Province, Chukotka Province (Choi et al, 2012). Known from Russia only. Mountain species.
Conservation. Protected in Russia: category and status—3a: rare species (Krasnaya..., 2008).
Specimens examined: Russia, Republic of Sakha (Yakutia), North-Eastern Yakutia, Or-ulgan Sis Range, Upper Tumara River (tributary Bytantay River), 68°13.924N, 128°40.001'E, 1340 m a. s. l., tundra belt, on moist soil covering stones of rocky outcrops on northern slope, with Scapania hyperborea and S. spitsbergensis, few plants with gemmae, 4 VIII 2011, Sofronova, SASY; Orulgan Sis Range, Khobol River Basin (tributary Bytantay River), 67°04'-67°05N, 128°48'-129°03'E, 950-1300 m a. s. l., few plants and minute continuous cover usually mixed with Tetralophozia setiformis, on moist fine earth, soil or soil covering stones in niches between the stones of rockfields or cracks in rock in forest and tundra belt, always with gemmae, June, July 2006, Sofronova, SASY; Orulgan Sis Range, Sobolokh-Mayan River (tributary Lena River), ~ 67°05N, 126°28'E, few plants on stone scree in Pinus pumila community on SW-facing slope, 23 VIII 1991, with gemmae, S. N. Kirillina, SASY; Orulgan Sis Range, Undyulyung River (tributary Lena River), ~ 66°13N, 126°59'E, few plants in the niches between the stones of rock-fields on N-facing slope in forest belt, 12 VII 1990, with gemmae, Sofronova, SASY; Western Verkhoyanie Range, Tukulan River Basin (tributary Aldan River), ~ 63°51'-64°01N, 132°40'-133°11'E, few plants in the niches between the stones of rockfields in forest belt, with gemmae, IX 1990, S. N. Kirillina, Sofronova, SASY; Eastern Yakutia, Ulakhan-Chistay Range, Upper Gyrbynya River (tributary Moma River), 64°38N, 146°39'E; ~ 1150 m a. s. l., few plants mixed with Tetralophozia setiformis, on moist soil in the niches between the stones of rockfields on E-facing slope in forest belt, with gemmae, 27 VII 2003, Sofronova, SASY; Suntar-Khayata Range, Vostochnaya Khandyga River Basin, 63°05'-63°08N, 139°02'-139°10'E, 850-1600 m a. s. l., few plants or minute continuous cover mixed with Scapania microdonta, Sphenolobus
saxicola, Tetralophozia setiformis, etc., on moist fine earth, soil covering stones or rarely decaying wood in niches between the stones of rockfields or rock outcrops in forest and tundra belt, always with gemmae, 1998, 1999, Sofronova, SASY; Suntar-Khayata Range, Upper Dyby River, 62°44'44.0"N, 139°05'06.3"E, 1000 m a. s. l., minute continuous cover in mat of Tetralophozia setiformis, on fine earth in niches between the stones of rockfields on north-western slope in forest belt, with gemmae, 27 VIII 2017, Sofronova, SASY; Suntar-Khayata Range, Upper Knory River (Agayakan River Basin), surroundings of Mus-Khaya Mt., 62°33'-62°35N, 140°56'-141°06'E; 1603-1738 m a. s. l., few plants and small continuous cover mixed with Trilophozia quinquedentata, Lophoziopsis excisa, Scapania spitsbergensis, on soil in niches between the stones of rockfields in forest and tundra belt, with gemmae, VII 2011, Sofronova, SASY; Southern Yakutia, Tokinsky Stanovik Range, Algama River Basin (tributary Gonam River), ~ 55°46N, 129°26'E, 900 m a. s. l., few plants with Sphenolobus minutus, S. saxicola, on moist fine earth in the niches between the stones of rockfields on W-facing slope in forest belt, with gemmae, 09 VII 2000, Sofronova, SASY; Udokan Range, Upper At-Bastaakh River (tributary Khani River), 57°14N, 120°28'E, 1300 m a. s. l., small continuous cover on moist fine earth in the niches between the stones of rockfields on N-facing slope in tundra belt, with gemmae, 13 VII 2002, Sofronova, SASY; Republic of Buryatia, South Muya Range, Valley of Barguzin River, 55°12'26.3"N, 111°41'41.9"E ... in niche between boulders, 1475 m a. s. l. Yu. S. Mamontov YuSM-392-1, LE; Khamar-Daban Range, Pereemnaya River Valley, mixed forest, on boulder, with per. and gemmae, 15 VIII 1989, S. G. Kazanovsky, LE; Northern macroslope of Khamar-Daban Mts., 51°23'57"N, 105°16'59"E, ., 780 m a. s. l., on shaded wet wall of big rock, with Marsupella emarginata, 11 VIII 2002, N. A. Konstantinova, LE; Trans-Baikal Territory, Krasniy Chikoy District, Malkhanskiy Range, 55°44'15"N, 110°22'52"E... in rock cracks, 1360 m a. s. l., with gemmae, ... 14 VIII 2011, I. V. Czenyadjeva Cz-28-11, LE; Sokhondinskiy Biosphere Reserve, Upper Bukukun River, 49°42N, 111°02'E, 1859 m a. s. l., on rocks, with gemmae, 20 VII 2008, O. M. Afonina #05908/3, LE; Primorsky Territory, Shkotovsky District, Pidan Mt., 700 m a. s. l., 43°05N, 132°42'E, with gemmae, 6 IX 2010, Bakalin P-39-10-10, LE.
Phytogeographic discussion
All species in question have a peculiar distribution pattern being rare worldwide and having comparatively regular occurrence through mountains of still fragmentary bryologically explored eastern Siberia and isolated locations in Europe and the western North America. These facts point at their broader and ± continuous ranges in the past and their fragmentation by ?Holocene glaciation.
The range of every species has specific features. Frullania davurica has mostly an Asian distribution pattern restricted by forest belt and isolated from closely related European F. jackii by Central European Plain. At present both species are usually sterile and their spore dissemination does not occur often. Lejeunea alaskana has amphipacific distribution pattern with many locations in eastern Siberia and western North America (Fig. 1). Marchantia romanica has European occurrence in Romania, Czech and Slovak Republics separated from other remote mountain locations (Fig. 2). Rare spore production, growth in sheltered niches and large size of spores (45-50 ^m) likely hamper distribution of M. romaica. Known range of Scapania sphaerifera (Fig. 3) is restricted by Eurasia, and has a single location in Europe, several in the Russian Far East, and numerous locations in Siberia. S. sphaerifera like M. romanica grows usually in sheltered niches. Despite constant production of gemmae only their slow dissemination via animals appears to be possible. Moreover the plants of S. sphaerifera are usually sterile that hampers dissemination. The facts above point at the relic distribution of these three species. On the contrary Lejeunea
alaskana appears to be rather young and efficiently disseminated species. Being monoicous it frequently produces gametoecia and probably spores.
Acknowledgments
Work of E. V. Sofronova was carried out within research project 0376-2018-0001 № AAAA-A17-117020110056-0 of Institute for Biological problems of Cryolitozone SB RAS. Work of A. D. Potemkin was carried out within the institutional research project «Tax-onomic diversity and ecology of bryophytes of Russia» (AAAA-A18-118032790222-1) of Komarov Botanical institute. Field work was partly supported by Russian foundation for Basic Research, projects 15-44-05134, 16-04-01156.
References
Afonina O. M., Mamontov Yu. S., Czernyadjeva I. V. 2013. New and rare liverwort and moss species from
Trans-Baikal Territory. Botanicheskii zhurnal 98(11): 1427-1440. (In Russ. with Engl. summary). Bakalin V. A. 2009. New liverwort records from Khabarovsk Territory. 1. Arctoa 18: 249-287.
https://doi.org/10.15298/arctoa.18.19 Bakalin V. A. 2015. Tardoki-Yani Range (North Sikhote-Alin, Pacific Russia) — the largest southern refugium
of arctic-alpine liverwort flora in East Asia. Arctoa 24: 314-326. https://doi.org/10.15298/arctoa.24.27 Bakalin V. A., Fedosov V. E., Borovichev E. A., Yanov A. V. 2016. Liverworts of Putorana Plateau (East Siberia): an updated checklist. Arctoa 25: 369-379. https://doi.org/10.15298/arctoa.25.29
Bakalin V., Choi S. S., Ermolenko A. 2012. Lejeunea alaskana (R. M. Schust. & Steere) Inoue & Steere, a new species for the Russian liverwort flora. Arctoa 21: 193-196. https://doi.org/10.15298/arctoa.21.18
Bakalin V. A., Vilnet A. A. 2009. Two new species of Jungermanniaceae from Asiatic Russia. Arctoa 18: 151-162. httpsVdoi.org/10.15298/arctoa.18.08
Buch H., Tuomikoski R. 1936. Scapania spaerifera spec. nova auctore Buch & Tuomikoski. Memoranda
Societatis pro Fauna et Flora Fennica 11: 228-230. Choi S. S. 2013. Taxonomy of the liverworts and hornworts in Korea. PhD thesis. Jeonju: 606 p.
Choi S. S., Bakalin V. A., Sun B.-Y. 2012. Scapania andMacrodiplophyllum in the Russian Far East. Botanica Pacifica 1: 31-95. https://doi.org/10.17581/bp.2012.01104
Fedosov V. E., Borovichev E. A., Ignatova E. A., Bakalin V. A. 2015. The bryophyte flora of Eriechka River upper course (SE Taimyr), with comments on the first record of Pseudoditrichum mirabile in Asia. Arctoa 24(1): 165-186. https:Zdoi.org/10.15298/arctoa.24.17
Gottsche C. M., Lindenberg J. B. G., Nees ab Esenbeck C. G. 1844. Synopsis Hepaticarum. Hamburgi: 834 p.
Hentschel J., Konrat M. J. von, Pocs T., Schafer-Verwimp A., Shaw A. J., Schneider H., Heinrichs J. 2009. Molecular insights into the phylogeny and subgeneric classification of Frullania (Frullaniaceae, Porel-lales). Molecular Phylogenetics and Evolution 52: 142-156. Hong W. S., Vitt D. H. 1977. The Hepaticae of the Yukon Territory, Canada. The Bryologist 80: 461-469. Kazanovsky S. G., Potemkin A. D. 1995. K flore pechenochnykh mkhov khrebta Khamar-Daban (Yuzh-noye Pribaykal'ye) [On the hepatic flora of Khamar-Daban (South Pribaikalje)]. Novosti sistematiki nizshikh rastenii 30: 98-110. (In Russ.).
Konstantinova N. A., Potemkin A. D. 1994. Studies on Scapania (Dum.) Dum., Hepaticae. Scapania sphaerifera Buch et Tuomik. Annales Botanici Fennci 31: 121-126.
Konstantinova N. A., Savchenko A. N. 2008. Diversity and phytogeography of hepatics of Siberia (Russia). Bryology in the New Millennium. Kuala Lumpur: 155-172.
Konstantinova N. A., Vasiljev А. N. 1994. On the hepatic flora of Sayan Mountains (South Siberia). Arctoa 3: 123-132. https://doi.org/10.15298/arctoa.03.07
Konstantinova N. A., Vilnet A. A., Stefanu^ S. 2014. On the distribution and variability of Bucegia romanica Radian. Arctoa 23: 137-144. https://doi.org/10.15298/arctoa.23.11
Krasnaya kniga Rossiyskoy Federatsii (rasteniya i griby) [Red Data Book of Russian Federation (plants and fungi)]. 2008. Moscow: 855 p. (In Russ.).
Krasnaya kniga Irkutskoy oblasti [Red Data Book of the Irkutsk Region]. 2010. Irkutsk: 481 p. (In Russ.).
Krasnaya kniga Respubliki Sakha (Yakutiya). T. 1: Redkiye i nakhodyashchiyesya pod ugrozoy ischezno-veniya vidy rasteniy i gribov [Red Data Book of the Republic of Sakha (Yakutia). Vol. 1: Rare and endangered species of plants and fungi]. 2017. Moscow: 412 p. (In Russ.).
Long D. G., Forrest L. L., Villarreal J. C., Crandall-Stotler B. J. 2016. Taxonomic changes in Mar-chantiaceae, Corsiniaceae and Cleveaceae (Marchantiidae, Marchantiophyta). Phytotaxa 252(1): 77-80. https://doi.org/10.11646/phytotaxa.252.1.9
Mamontov Yu. S., Afonina O. M. 2015. New and rare liverwort and moss species for Trans-Baikal Territory from the Syulban River Valley (Kodar Range, Stanovoe Upland). Botanicheskii zhurnal 100(12): 1311-1321. (In Russ. with Engl. summary).
Mamontov Yu. S., Konstantinova N. A., Afonina O. M. 2011. New records. New liverwort records from Zabaikal'sky Territory. 5. Arctoa 20: 247-268. https://doi.org/10.15298/arctoa.20.20
Mamontov Y. S., Konstantinova N. A., Vilnet A. A., Potemkin A. D., Sofronova E. V., Gamova N. S. 2018. On resurrection of Marsupella parvitexta Steph. (Gymnomitriaceae, Marchantiophyta) as a semi cryptic species of the genus Gymnomitrion. Nova Hedwigia 106(1-2): 81-101. https://doi.org/10.1127/nova_hedwigia/2017/0466
Mamontov Yu. S., Vilnet A. А. 2013. Anastrophyllum ellipticum Inoue (Jungermanniales, Marchantiophyta), a new species for Russian liverwort flora. Arctoa 22: 151-158. https://doi.org/10.15298/arctoa.22.24
Piippo S. 1990. Annotated catalogue of Chinese Hepaticae and Anthocerotae. Journal of the Hattori Botanical Laboratory 68: 1-192.
Potemkin A. D. 1995. Contribution to the knowledge of the liverworts of North America. Fragmenta Flo-ristica et Geobotanica 40: 323-338.
Potemkin A. D. 2000. An updated list of liverworts of the Severnaya Zemlya Archipelago (East Siberian High Arctic) with description of new species, Scapania matveyevae. Arctoa 9: 95-100. https://doi.org/10.15298/arctoa.09.07
Potemkin A. D. 2003. New and little known for Russia hepatics from Kamchatka and Promorskiy Territory (Russian Far East). Arctoa 12: 75-82. (In Russ. with Engl. abstract). https://doi.org/10.15298/arctoa.12.05
Potemkin A. D. 2004. Heteroscyphus (Geocalycaceae)-a new liverwort genus for Russia. Arctoa 13:
67-72. (In Russ. with Engl. summary). https://doi.org/10.15298/arctoa.13.09
Potemkin A. D., Mamontov Yu. S. 2013. On Syzygiella nipponica (Adelanthaceae, Marchantiophyta) and its first record in Russia. Arctoa 22: 167-172. https://doi.org/10.15298/arctoa.22.27
Potemkin A. D., Mamontov Y. S., Borovichev E. A., Fedosov V. E., Sofronova E. V. 2015. The genus As-cidiota C. Massal. (Porellaceae, Marchantiophyta) in North Asia. Journal of Bryology 37(1): 49-55. https://doi.org/10.1179/1743282014Y.0000000125
Potemkin A. D., Sofronova E. V. 2009. Pechenochniki i antocerotovye Rossii. T. 1 [Liverworts and hornworts of Russia. Vol. 1]. St. Petersburg; Yakutsk: 368 p. (In Russ. and Engl.).
Schljakov R. N. 1973. Pechenochnyy mokh Bucegia romanica Radian na severe Yakutii [Bucegia romanica Radian (Hepaticae) in the north of Yakutia]. Novosti sistematiki nizshikh rastenii 10: 285-286.
Schljakov R. N. 1981. Pechenochnyye mkhi Severa SSSR. Vyp. 4. Pechenochniki: Yungerman-nievye — Skapanievye [Liverworts of the North of the USSR. Vol. 4. Liverworts: Jungermanniace-ae — Scapaniaceae]. Leningrad: 221 p. (In Russ.).
Schumacker R., Váña J. 2005. Identification keys to the liverworts and hornworts of Europe andMacaron-esia (distribution and status). 2nd ed. fully revised and updated. Poznan: 211 p.
Schuster R. M., Konstantinova N. A. 1996. Studies on the distribution of critical arctic/subarctic Hepaticae with special reference to taxa found in Russia. Lindbergia 21: 26-48.
Schuster R. M., Steere W. C. 1958. Hygrolejeunea alaskana sp. n., a critical endemic of Northern Alaska. Bulletin of the Torrey Botanical Club 85: 188-196.
Sofronova E. V. 2003. Pechenochnye mkhi yakutskoy chasti Vostochnogo Verkhoyan'ya. Kand. Diss. [He-patics of the Yakutian part of East Verkhoyanie Range. Cand. Diss.] St. Petersburg: 213 p. (In Russ.).
Sofronova E. V. 2017. A contribution to the liverwort flora of Timpton River Basin (Aldan Upland, Southern Yakutia). Arctoa 26(1): 81-90. https://doi.org/10.15298/arctoa.26.08
Sofronova E. V., Mamontov Yu. S., Potemkin A. D. 2013. Frullania ignatovii (Porellales, Marchantiophy-ta), a new species from Siberia. Novosti sistematiki nizshikh rastenii 47: 334-343. (In Russ. and Engl.).
Sofronova E. V., Ivanova E. I., Mikhaleva L. G., Poryadina L. N. 2014. Rare lichens, mosses, liverworts and fungi from the Republic Sakha (Yakutia), Russia. Folia Cryptogamica Estonica 51: 89-102. https://doi.org/10.12697/fce.2014.51.10
Sofronova E. V., Kopyrina L. I. 2016. A contribution to the liverwort flora of Endybal River Basin (Arkachan Plateau, Verkhoyansky Range, Yakutia). Arctoa 25(2): 380-385. https://doi.org/10.15298/arctoa.25.30
Sofronova E. V., Potemkin A. D., Mamontov Yu. S., Sofronov R. R. 2015. Liverworts of the Mus-Khaya Mountain (Yakutia, Asiatic Russia). Arctoa 24(1): 156-164. https://doi.org/10.15298/arctoa.24.16
Steere W. C., Inoue H. 1978. The Hepaticae of Arctic Alaska. Journal of the Hattori Botanical Laboratory 44: 251-345.
Steere W. С., Scotter G. W. 1978. Additional bryophytes from Nahanni National Park and vicinity, Northwest Territories, Canada. Canadian Journal of Botany 56: 234-244.
Tubanova D. Ya., Mamontov Yu. S., Afonina O. M., Potemkin A. D. 2017. New and rare species in the moss and liverwort flora of the Republic of Buryatia. Botanicheskii zhurnal 102(10): 1442-1454. (In Russ. with Engl. summary).
Váña J. 1991-1992. Phytogeographically important hepatics from Altai Mts. (Russia). Novitates botanicae Universitatis Carolinae, Praha 7: 27-32.
Váña J., Soldán Z. 1985. Some new and phytogeographically interesting bryophytes from Central Siberia. Abstracta Botanica 9 (Suppl. 2): 123-144.
Vasiljev A. N. 1992. Konspekt flory mokhoobraznykh v zapovednikakh «Stolby» i Sayano-Shushenskom [Bryophytes of Reserves «Stolby» and Sayano-Shushenskiy]. Krasnoyarsk: 112 p. (In Russ.).
Yamada К., Iwatsuki Z. 2006. Catalog of the hepatics of Japan. Journal of the Hattori Botanical Laboratory 99: 1-106.
Zhu R.-L. 2006. New checklist of Chinese liverworts, hornworts and takakiophytes. Shanghai: 25 p.
Литература
[Afonina et al.] Афонина О. M., Мамонтов Ю. С., Чернядьева И. В. 2013. Новые и редкие виды печеночников и мхов для Забайкальского края. Ботанический журнал 98(11): 1427-1440.
Bakalin V. A. 2009. New records. New liverwort records from Khabarovsk Territory. 1. Arctoa 18: 249287. https://doi.org/10.15298/arctoa.18.19
Bakalin V. A. 2015. Tardoki-Yani Range (North Sikhote-Alin, Pacific Russia)—the largest southern refugium of arctic-alpine liverwort flora in East Asia. Arctoa 24(2): 314-326. https://doi.org/10.15298/arctoa.24.27
Bakalin V., Choi S. S., Ermolenko A. 2012. Lejeunea alaskana (R. M. Schust. & Steere) Inoue & Steere, a new species for the Russian liverwort flora. Arctoa 21: 193-196. https://doi.org/10.15298/arctoa.21.18
Bakalin V.A., Fedosov V.E., Borovichev E.A., Yanov A.V. 2016. Liverworts of Putorana Plateau (East Siberia): an updated checklist. Arctoa 25: 369-379. https://doi.org/10.15298/arctoa.25.29
Bakalin V. A., Vilnet A. A. 2009. Two new species of Jungermanniaceae from Asiatic Russia. Arctoa 18: 151-162. https:7doi.org/10.15298/arctoa.18.08
Buch H., Tuomikoski R. 1936. Scapania spaerifera spec. nova auctore Buch & Tuomikoski. Memoranda Societatis pro Fauna et Flora Fennica 11: 228-230.
Choi S. S. 2013. Taxonomy of the liverworts and hornworts in Korea. PhD thesis. Jeonju: 606 p.
Choi S. S., Bakalin V. A., Sun B.-Y. 2012. Scapania andMacrodiplophyllum in the Russian Far East. Botanica Pacifica 1: 31-95. https://doi.org/10.17581/bp.2012.01104
Fedosov V. E., Borovichev E. A., Ignatova E. A., Bakalin V. A. 2015. The bryophyte flora of Eriechka River upper course (SE Taimyr), with comments on the first record of Pseudoditrichum mirabile in Asia. Arctoa 24(1): 165-186. https:/doi.org/10.15298/arctoa.24.17
Gottsche C. M., Lindenberg J. B. G., Nees ab Esenbeck C. G. 1844. Synopsis Hepaticarum. Hamburgi: 834 p.
Hentschel J., Konrat M. J. von, Pocs T., Schafer-Verwimp A., Shaw A. J., Schneider H., Heinrichs J. 2009. Molecular insights into the phylogeny and subgeneric classifi cation of Frullania (Frullaniaceae, Porel-lales). Molecular Phylogenetics and Evolution 52: 142-156.
Hong W. S., Vitt D. H. 1977. The hepaticae of the Yukon Territory, Canada. The Bryologist 80: 461-469.
[Kazanovsky, Potemkin] Казановский С. Г., Потемкин А. Д. 1995. К флоре печеночных мхов хребта Хамар-Дабан (Южное Прибайкалье). Новости систематики низших растений 30: 98-110.
Konstantinova N. A., Potemkin A. D. 1994. Studies on Scapania (Dum.) Dum., Hepaticae. Scapania sphaerifera Buch et Tuomik. Annales Botanici Fennici 31: 121-126.
Konstantinova N. A., Savchenko A. N. 2008. Diversity and phytogeography of hepatics of Siberia (Russia). Bryology in the New Millennium. Kuala Lumpur: 155-172.
Konstantinova N. A., Vasiljev А. N. 1994. On the hepatic flora of Sayan Mountains (South Siberia). Arctoa 3: 123-132. https://doi.org/10.15298/arctoa.03.07
Konstantinova N. A., Vilnet A. A., Stefanu^ S. 2014. On the distribution and variability of Bucegia romanica Radian. Arctoa 23: 137-144. https://doi.org/10.15298/arctoa.23.! 1
[Krasnaya...] Красная книга Российской Федерации (растения и грибы). 2008. М.: 855 с.
[Krasnaya...] Красная книга Иркутской области. 2010. Иркутск: 481 с.
[Krasnaya.] Красная книга Республики Саха (Якутия). Т. 1. Редкие и находящиеся под угрозой исчезновения виды растений и грибов. 2017. М.: 412 с.
Long D. G., Forrest L. L., Villarreal J. C., Crandall-Stotler B. J. 2016. Taxonomic changes in March-antiaceae, Corsiniaceae and Cleveaceae (Marchantiidae, Marchantiophyta). Phytotaxa 252(1): 77-80. https://doi.org/10.11646/phytotaxa.252.1.9
[Mamontov, Afonina] Мамонтов Ю.С., Афонина О.М. 2015. Новые и редкие виды печеночников и мхов для Забайкальского края из долины р. Сюльбан (хребет Кодар, Становое нагорье). Ботанический журнал 100(12): 1311-1321.
Mamontov Yu. S., Konstantinova N. A., Afonina O. M. 2011. New records. New liverwort records from Zabaikal'sky Territory. 5. Arctoa 20: 247-268. https://doi.org/10.15298/arctoa.20.20
Mamontov Y. S., Konstantinova N. A., Vilnet A. A., Potemkin A. D., Sofronova E. V., Gamova N. S. 2018. On resurrection of Marsupella parvitexta Steph. (Gymnomitriaceae, Marchantiophyta) as a semi cryptic species of the genus Gymnomitrion. Nova Hedwigia 106(1-2): 81-101. https://doi.org/10.1127/nova_hedwigia/2017/0466
Mamontov Yu. S., Vilnet A. А. 2013. Anastrophyllum ellipticum Inoue (Jungermanniales, Marchantiophyta), a new species for Russian liverwort flora. Arctoa 22: 151-158. https://doi.org/10.15298/arctoa.22.24
Piippo S. 1990. Annotated catalogue of Chinese Hepaticae and Anthocerotae. Journal of the Hattori Botanical Laboratory 68: 1-192.
Potemkin A. D. 1995. Contribution to the knowledge of the liverworts of North America. Fragmenta Flo-ristica et Geobotanica 40: 323-338.
Potemkin A. D. 2000. An updated list of liverworts of the Severnaya Zemlya Archipelago (East Siberian High Arctic) with description of new species, Scapania matveyevae. Arctoa 9: 95-100. https://doi.org/10.15298/arctoa.09.07
[Potemkin] Потемкин A. Д. 2003. Новые и малоизвестные для России печеночники с Камчатки и из Приморского Края (Российский Дальний Восток). Arctoa 12: 75-82. https://doi.org/10.15298/arctoa.12.05
[Potemkin] Потемкин А. Д. 2004. Heteroscyphus (Geocalycaceae, Hepaticae)—новый род печеночников для России. Arctoa 13: 67-72. https://doi.org/10.15298/arctoa.13.09
Potemkin A. D., Mamontov Yu. S. 2013. On Syzygiella nipponica (Adelanthaceae, Marchantiophyta) and its first record in Russia. Arctoa 22: 167-172. https://doi.org/10.15298/arctoa.22.27
Potemkin A. D., Mamontov Y. S., Borovichev E. A., Fedosov V. E., Sofronova E. V. 2015. The genus As-cidiota C. Massal. (Porellaceae, Marchantiophyta) in North Asia. Journal of Bryology 37(1): 49-55. https://doi.org/10.1179/1743282014Y.0000000125
[Potemkin, Sofronova] Потемкин A. Д., Софронова E. В. 2009. Печеночники и антоцеротовыеРоссии. Т. 1. СПб.; Якутск: 368 с.
[Schljakov] Шляков Р. Н. 1973. Печеночный мох Bucegia romanica Radian на севере Якутии. Новости систематики низших растений. 10: 285-286.
[Schljakov] Шляков Р. Н. 1981. Печеночные мхи Севера СССР. Вып. 4. Печеночники: Юнгерманниевые-Скапаниевые. Л.: 221 с.
Schumacker R., Vana J. 2005. Identification keys to the liverworts and hornworts of Europe andMacaron-esia (distribution and status). 2nd ed. fully revised and updated. Poznan: 211 p.
Schuster R. M., Konstantinova N. A. 1996. Studies on the distribution of critical arctic/subarctic Hepaticae with special reference to taxa found in Russia. Lindbergia 21: 26-48.
Schuster R. M., Steere W. C. 1958. Hygrolejeunea alaskana sp. n., a critical endemic of Northern Alaska. Bulletin of the Torrey Botanical Club 85: 188-196.
[Sofronova] Софронова Е. В. 2003. Печеночные мхи якутской части Восточного Верхоянья. Дисс. ... канд. биол. наук. СПб.: 213 c.
Sofronova E. V. 2017. A contribution to the liverwort flora of Timpton River Basin (Aldan Upland, Southern Yakutia). Arctoa 26(1): 81-90. https://doi.org/10.15298/arctoa.26.08
[Sofronova et al.] Софронова Е. В., Мамонтов Ю. С., Потемкин А. Д. 2013. Frullania ignatovii (Porel-lales, Marchantiophyta) — новый вид из Сибири. Новости систематики низших растений 47: 334-343.
Sofronova E. V., Ivanova E. I., Mikhaleva L. G., Poryadina L. N. 2014. Rare lichens, mosses, liverworts and fungi from the Republic Sakha (Yakutia), Russia. Folia Cryptogamica Estonica 51: 89-102. https://doi.org/10.12697/fce.2014.51.10
Sofronova E. V., Kopyrina L. I. 2016. A contribution to the liverwort flora of Endybal River Basin (Arkachan Plateau, Verkhoyansky Range, Yakutia). Arctoa 25(2): 380-385. https://doi.org/10.15298/arctoa.25.30
Sofronova E. V., Potemkin A. D., Mamontov Yu. S., Sofronov R. R. 2015. Liverworts of the Mus-Khaya Mountain (Yakutia, Asiatic Russia). Arctoa 24(1): 156-164. https://doi.org/10.15298/arctoa.24.16
Steere W. C., Inoue H. 1978. The Hepaticae of Arctic Alaska. Journal of the Hattori Botanical Laboratory 44: 251-345.
Steere W. С., Scotter G. W. 1978. Additional bryophytes from Nahanni National Park and vicinity, Northwest Territories, Canada. Canadian Journal of Botany 56: 234-244.
[Tubanova et al.] Тубанова Д. Я., Мамонтов Ю. С., Афонина О. М., Потемкин А.Д. 2017. Новые и редкие виды мхов и печеночников во флоре республики Бурятия. Ботанический журнал 102(10): 1442-1454.
Vana J. 1991-1992. Phytogeographically important hepatics from Altai Mts. (Russia). Novitates botanicae Universitatis Carolinae, Praha 7: 27-32.
Váüa J., Soldán Z. 1985. Some new and phytogeographically interesting bryophytes from Central Siberia. Abstracta Botanica 9 (Suppl. 2): 123-144.
[Vasiljev] Васильев А. Н. 1992. Конспект флоры мохообразных в заповедниках «Столбы» и Саяно-Шушенском. Красноярск: 112 с.
Yamada К., Iwatsuki Z. 2006. Catalog of the hepatics of Japan. Journal of the Hattori Botanical Laboratory 99: 1-106.
Zhu R.-L. 2006. New checklist of Chinese liverworts, hornworts and takakiophytes. Shanghai: 25 p.
