Novosti Sistematiki Nizshikh Rastenii 52(2): 505–518
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Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 52(2): 505–518. 2018 Four rare liverwort species: distribution, ecology, taxonomy E. V. Sofronova1, A. D. Potemkin2 1Institute 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 collec- tions 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 1Институт 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 cпорофиты Lejeunea alaskana. Ключевые слова: Frullania davurica, Lejeunea alaskana, Marchantia romanica, Scapania sphaeri- fera, Marchantiophyta, ареал, местообитания, Восточное Верхоянье, Республика Саха (Якутия), Рос- сия, Сибирь. Distribution of Holarctic liverwort species is still insufficiently studied. Extensive terri- tories 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; Potem- kin 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 Sofronova, Potemkin. Four rare liverwort species: distribution, ecology, taxonomy first recorded for Russia in the last century or described from Russia are Frullania davurica, Lejeunea alaskana, Marchantia 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 moun- tain 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 con- tinuous 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°30′N, 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) 506 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 52(2): 505–518. 2018 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 differ- entiation 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 cap- sules 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 × 40–50(75) μm, with regularly papillose surface with papillae ca. 1 μm 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 dis- tinctive. It grows on soil over rocks and usually associates with characteristic tundra com- 507 Sofronova, Potemkin. Four rare liverwort species: distribution, ecology, taxonomy munities with calciphilous Odontoschisma macounii (Austin) Underw., Scapania simmonsii Bryhn et Kaal., Frullania subarctica Vilnet, Borovich. et Bakalin as well as with neutrophi- lous 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 com- munities. 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: Alas- ka (Schuster, Steere, 1958; Steere, Inoue, 1978); Canada: Northwest Territories and Yukon (Hong, Vitt, 1977;