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15On Frullania usamiensis (Frullaniaceae, Marchantiophyta) and its first record for Russia
Yu. S. Mamontov1, 2, T. I. Koroteeva3, E. V. Sofronova4, A. D. Potemkin5
'Tsitsin Main Botanical Garden of the Russian Academy of Sciences, Moscow, Russia 2Polar-Alpine Botanical Garden-Institute, Kola Science Centre of the Russian Academy of Sciences,
Kirovsk-6, Russia
3Institute of Marine Geology and Geophysics, Far Eastern Branch of the Russian Academy of Sciences, Yuzhno-Sakhalinsk, Russia
4Institute of Biological Problems of Cryolithozone, Siberian Branch of the Russian Academy
of Sciences, Yakutsk, Russia 5Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia Corresponding author. A. D. Potemkin, potemkin_alexey@binran.ru
Abstract. Frullania usamiensis is recorded for the first time for Russia from Kunashir Island, the southernmost island of the Greater Kuril Chain. It is the northernmost locality of the species, which was previously known from Japan (Honshu, Shikoku and Kyushu), China (coastal provinces Fujian and Liaoning) and the Republic of Korea. Frullania usamiensis is distinguished by (1) relatively large, emarginate or shallowly bilobed underleaves with obtuse lobes and mostly rounded sinus, (2) characteristic leaf lobules which are transversely elongated (when inflated), about x 1.5-1.9 as wide as long, with incurved rostral portion, and (3) smooth perianth with three keels. Description and photomicrographs of F. usamiensis from Kunashir Island are provided together with discussion on its ecology, variation and differentiation from having much in common F. kagoshimensis and F. taradakensis.
Keywords: Frullania usamiensis, liverworts, Kunashir Island, Russia.
O Frullania usamiensis (Frullaniaceae, Marchantiophyta) и первой находке этого вида в России
Ю. С. Мамонтов1, 2, Т. И. Коротеева3, Е. В. Софронова4, А. Д. Потемкин5
Главный ботанический сад им. Н. В. Цицина РАН, Москва, Россия 2Полярно-альпийский ботанический сад-институт им. Н. А. Аврорина КНЦ РАН, Кировск-6,
Россия
3Институт морской геологии и геофизики ДВО РАН, Южно-Сахалинск, Россия 4Институт биологических проблем криолитозоны СО РАН — обособленное подразделение
ФИЦ ЯНЦ СО РАН, Якутск, Россия 5Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия Автор для переписки. А. Д. Потемкин, potemkin_alexey@binran.ru
Резюме. Новый для России вид печеночников Frullania usamiensis выявлен на Кунаши-ре — самом южном из островов Большой Курильской гряды. Это наиболее северная находка данного вида, который ранее был известен только в Японии (Хонсю, Сикоку и Кюсю), Китае (пров. Ляонин, Фуцзянь) и в Республике Корея. Для F. usamiensis характерна следующая
https://doi.org/10.31111/nsnr/2020.54.1243
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комбинация признаков: (1) амфигастрии крупные, почти цельные на верхушке или выемчатые, или с мелкой, в основном закругленной вырезкой и тупыми до закругленных лопастями; (2) брюшные лопасти поперечно вытянутые, с соотношением длины к высоте 1.5-1.9 к 1, с почти прямой или изогнутой вниз ростральной областью; (3) периантий с тремя выступающими килями и гладкой (без выростов) поверхностью. Приводятся описание и микрофотографии F. usamiensis, а также обсуждение экологии, вариабельности и отличий этого вида от сходных F. kagoshimensis и F. taradakensis.
Ключевые слова: Frullania usamiensis, печеночники, Кунашир, Россия.
During the study of the liverwort flora of the Kuril Islands by T. I. Koroteeva, a characteristic species of the genus Frullania Raddi, F. usamiensis Steph., was discovered for the first time for Russia. It was described in 1897 by Stephani (1897) from the middle of Honshu, Japan (Usami, Shizuoka Prefecture) and later was reported as not rare in Japan (Stephani, 1910). Horikawa (1934) described F. yakushimensis Horik. from Yakushima Island, however the latter was treated as a form of F. usamiensis by Hattori (1951) and synonymized with it by Kamimura (1961). After its description, F. usamiensis became known in the mountains of middle and southern Japan (Shikoku, Kyushu, Tsuishima and Yakushima islands) and the Korean Peninsula, where it occurs mostly on trees, in the deciduous forest zone (Kamimura, 1961). Since the study by Kamimura (1961), F. usamiensis was discovered in Liaoning (Fu, Fang, 1994) and Fu-jian (Zhang et al., 2011), the coastal provinces of China. In Kunashir Island the species was collected on bark of fir [Abies sachalinensis (F. Schmidt) Mast.], along the coast of Kunashira Strait between Kunashir and Hokkaido islands. Below we provide the description and photomicrographs of F. usamiensis from Kunashir Island together with discussion on its ecology, variation and differentiation from having much in common F. kagoshimensis Steph. and F. taradakensis Steph.
Materials and methods
The description is based on the studied specimen from Kunashir Island. Photomicrographs were obtained with Leitz Wetzlar Orthoplan light microscope equipped with a Nikon D700 digital camera. In order to better illustrate the three-dimensional objects, photomicrographs were combined from several optical sections using the software package HeliconFocus.
Results
Frullania usamiensis Steph., 1897, Bull. Herb. Boissier 5: 91. (Plates I, II)
Plants in thin dense mats, green to deep dark brown, loosely adhering to substrate, irregularly pinnately branched; shoots to 1.5 cm long and 1.8-2.2 mm wide. Stems rounded, 155-245 |im in diameter, light brown. Dorsal leaf lobes slightly concave and imbricate (in view from above), sometimes spreading to weakly squarrose, widely re-niform to widely obovate, 1.0-1.3 mm long, 1.0-1.2 mm wide, with length/width ratio ca. (0.85)0.9-1.0(1.1) / 1; antical base not auriculate, or only weakly so, extending ca. stem-width across and beyond the farther edge of stem, postical base not cordate;
apex of lobes broadly rounded; margins entire. Lobules inflated, 0.1-0.2 of the size of the lobes, transversely elongated, 290-480 |im wide, 180-290 |im high, with length/ width ratio ca. 1.5-1.9 / 1, widely cucullate; the upper part distinctly saccate, inflated to the substrate, antero-upper part inflated, postero-upper part inclined or inflated; the rostral portion distinct, tubular; the beak distinct, nearly straight or incurved toward the lobule base, rather short, rounded to obtuse, hoof-shaped (not hooked), without apical hollow; the mouth truncate, rarely nearly straight, compressed, both ventral and dorsal valves wrapped up inside the mouth, the mouth opening to the subtrate, the keel ca. 0.3-0.4 of the lobule width. Stylus indistinct, filiform, 1(2) cells wide at the base, 4-6 cells long, sometimes ending by slime papilla. Cells in the lobe middle rounded hexagonal, with medium to rather large, usually convex (rarely concave) trigones, with intermediate thickenings near the base of the lobes; near the lobe apex the trigones small to medium, the intermediate thickenings usually absent or small and indistinct; the median cells ca. (18)22-27(32) x (20)27-35(40) ^m, the marginal cells smaller, nearly quadrate or slightly elongated, (9)11-18 x (12)15-22(26) |im; the cells near the base larger, (17)22-30 |im x (22)30-48(54) |im. Oil-bodies seen in inner female bracts only, 8-11 per cell, brownish, granulate, elliptical, 2.5-5 x 5.5-6.5(7) |im long. Ocelli lacking. Underleaves nearly transversely inserted, (750)950-1000 |im wide, (600)700-960 |im long, 4-5(6) times as wide as stem, widely reniform or obovate or orbicular, widest near the middle, often a little or distinctly above, the base almost rounded or obcuneate, undecurrent, with line of insertion slightly arched; the apex emarginated or bilobed ca. 0.1-0.15 of the length, sinus broad and rounded (rarely acute), margins plane, entire; lobes obtuse. Asexual reproduction unknown. Dioecious. Androecia terminal on short lateral branches, usually subglobose (button-shaped), compact or shortly spicate, of 4-5(7) pairs of densely imbricate bracts. Bracts sub-equally bilobed, lobules near the middle usually with small, filiform stylus, 1-2 cells at base and up to 4 cells long. Androecia with 3-4 bracteoles; male bracteoles free, with entire margins; upper bracteoles very small and almost quadrate, 70-160 |im wide, 70-160 |im long, bifid ca. 0.25-0.7 of the length with obtuse lobes and ± U-shaped sinus; the bracteoles near the base almost rectangular, 60-200 |im wide, 270-310 |im long, bifid ca. 0.2-0.4 of the length, with obtuse lobes and ± U-shaped sinus. Anther-idia (1)2-3 per pract, with a long stalk of 16-28 cells. Unfertilized gynoecia terminal on main shoots or on lateral branches. Female bracts unequally bilobed, with entire margins, bracts divided (0.5)0.6-0.7 of the length; lobes slightly concave, broadly oval to ovate, 690-870 |im wide, 1260-1430 |im long, apex rounded to slightly acuminate; lobules ovate to almost triangular, slightly canaliculate to nearly plane, 380-470 |im wide, 1050-1190 |im long, apex acute to acuminate; stylus at base of lobules, filiform, 1-2(3) cells at base and (2)3-8 cells long, sometimes ending by slime papilla, additional cilia at the base 1-3 cells long. Female bracteoles slightly connate at the base with one of the bracts, 420-800 |im wide, 580-1150 |im long, at the base with 1-celled cilia and rounded outgrowth, bilobed 0.25-0.45 of the length, sinus ± U-shaped, lobes narrowly triangular, with 1 large obtuse tooth or distinctly angular at one side, lobes ± direc-
Fig. 1. Frullania usamiensis (from Koroteeva 15-10, KPABG). 1 — portion of female shoot with perianth; 2-4 — sterile shoot fragments; 5, 6 — portions of male shoot showing androecia. Scale bars: 1-6 — 500 |im.
Fig. 2. Frullania usamiensis (from Koroteeva 15-10, KPABG). 1-7 — underleaves; 8, 10 — enlarged leaf lobules; 9 — leaf lobe and lobule. Scale bars: 1-10 — 250 |m.
ted forward, apex acute to apiculate. Perianth half-exserted, obovate, 1100-1530 |im wide, 1750-1930 |im long, with two lateral keels and one high and sharp ventral keel; the surface smooth; the apex subtruncate or ± retuse, with a beak 140-170 |im long; the beak apex crenulate or smooth.
Distribution. Japan — Honshu, Shikoku, Kyushu including Tsushima I., Yaku-shima I. (Kamimura, 1961; Yamada, Iwatsuki, 2006; M. Higuchi e-mail to A. D. Po-temkin of 27 XII 2019), Republic of Korea (Kamimura 1961; Hong, 1997; Choi, 2013), China (Fu, Fang, 1994; Zhang et al., 2011), Russia — Kunashir Island (present study).
Ecology. According to Kamimura (1961), F. usamiensis grows on bark (branches or trunks), rarely on rocks, in the deciduous forest zone, occurring on Abies, Tsuga, Betula spp., Fagus, Acer spp., Prunus spp., etc., mostly at altitudes 890-1800 m a. s. l. in Shikoku and Kyushu, and 600-1100 m a. s. l. in the middle and northern Japan. In Kunashir Island, F. usamiensis was collected on bark of Abies sachalinensis, in deciduous coniferous forest, 123 m a. s. l., in pure mat about 25 cm2. In China it was collected on rocks and trunks (Fu, Fang, 1994).
Specimens examined: Russia, Sakhalin Region, Kunashir Island, vicinity of Tretyakovo Settlement, deciduous coniferous forest, 43°58'36"N, 145°39'56"E, 123 m a. s. l., on bark of Abies sachalinensis about 1 m above ground, 13 VIII 2015, Koroteeva 15-10 (female plants with perianths and male plants — KPABG, LE, MHA, SASY). Japan, Honshu, Nagano Prefecture, Mt. Kisokama, ca. 1200 m a. s. l., on bark, 22 VIII 1955, Iwatsuki (Hepaticae Japonicae, Ser. 10 (1958), No. 468 — LE); Shikoku, Ehime Prefecture, Mt. Nakatsu, summit area with Sasa and Hydrangea paniculata, ca. 1500 m a. s. l., 25 XI 1999, Deguchi 34977 (Bryophytes of Asia, Fasc. 10 (2003), No. 246 — KPABG).
Discussion
Among the species of the genus Frullania known in Russia, F. usamiensis is a remarkable species having comparatively large shoots, transversely elongated lobules and shallowly bilobed underleaves. The species F. taradakensis known from the south of the Russian Far East, is similar to F. usamiensis in rather large shoots and the shape of leaf lobules and underleaves. However, F. usamiensis differs from F. taradakensis in the following characters: (1) underleaves in F. usamiensis are ± obcuneate, widest in the middle or a little above, often narrowed in basal part, not or slightly narrowed to the apex, whereas in F. taradakensis the underleaves are cordate, widest at most below the middle, gradually narrowed to the apex; (2) underleaves in F. usamiensis are slightly emarginate at apex, or sometimes nearly truncate, or about i/s-bilobed, with rounded lobes, usually with broad and rounded (rarely acute) sinus, whereas in F. taradakensis the underleaves are Vs-bilobed at apex (rarely up to with the
lobes widely triangular, subacute to obtuse, usually with rather narrow and obtuse sinus.
The species F. fauriana Steph., F. kagoshimensis and F. osumiensis (S. Hatt.) S. Hatt. known from Japan resemble F. usamiensis in leaf lobule shape and shallowly bilobed
underleaves. However, F. fauriana and F. osumiensis differ from F. usamiensis in dentate in upper half underleaves.
Frullania kagoshimensis, which is most closely related to F. usamiensis, differs from the latter in Vs-^-bilobed underleaves with acute triangular lobes and always narrow, obtuse to acute sinus. Kamimura (1961) mentioned also that F. kagoshimensis is smaller than F. usamiensis (1.5-2 mm wide with leaves vs. 2.5-2.8 mm wide). However, the plants of F. usamiensis from Kunashir Island, the northern border of the distribution of this species, are smaller and correspond in their size to F. kagoshimensis measured in Kamimura (1961). The similar size is characteristic of F. usamiensis plants from the Middle Honshu Island identified and distributed by S. Hattori with exsiccatae (He-paticae Japonicae, Ser. 10 (1958), No. 468). At this basis we consider that most diagnostic characters of this species are broad underleaves with blunt lobes and shallow and mostly rounded sunus. Noteworthy, F. kagoshimensis has similar range in Japan (Honshu, Shikoku, Kyushu, Ryukyu, Yakushima, Amammi-oshima islands, Bonin) but occurs in lower latitudes (Kamimura, 1961; Yamada, Iwatsuki, 2006). Therefore its distribution northwards is less probable.
Acknowledgments
We are grateful to Dr. Masanobu Higuchi and Dr. Rui-Liang Zhu for providing information on recent data on distribution of Frullania usamiensis in China and Japan. The work was partially supported by the Russian Foundation for Basic Research under grants 18-04-00594 and 19-04-01270. The study by A. D. Potemkin was carried out in the framework of the institutional research project of Komarov Botanical Institute of the Russian Academy of Sciences "Flora and systematics of lichens and bryophytes of Russia and phytogeographically important regions" (project AAAA-A18-118032790222-1). The study by E. V. Sofronova was carried out within the framework of the institutional research project AAAA-A17-117020110056-0 of the Institute for Biological Problems of Cryolithozone SB RAS.
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