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12Rinodina sibirica (Physciaceae, lichenized Ascomycota) in Eurasia
I. A. Galanina1, S. V. Chesnokov2, 3, D. E. Himelbrant3, 4, E. A. Davydov5, A. K. Ezhkin6, T. M. Kharpukhaeva7, L. A. Konoreva2, 8, E. S. Kuznetsova3, 4, L. N. Poryadina9, I. S. Stepanchikova3, 4, L. S. Yakovchenko1, E. V. Zheludeva10
1Federal Scientific Center of East Asian Terrestrial Biodiversity of the Far Eastern Branch
of the Russian Academy of Sciences, Vladivostok, Russia 2Botanical Garden-Institute of the Far Eastern Branch of the Russian Academy of Sciences,
Vladivostok, Russia
3Komarov Botanical Institute of the Russian Academy of Science, St. Petersburg, Russia 4St. Petersburg State University, St. Petersburg, Russia 5Altai State University, Barnaul, Russia 6Institute of Marine Geology and Geophysics of the Far Eastern Branch of the Russian Academy
of Sciences, Yuzhno-Sakhalinsk, Russia 7Institute of General and Experimental Biology of the Siberian Branch of the Russian Academy
of Sciences, Ulan-Ude, Russia 8The Polar-Alpine Botanical Garden-Institute of the Kola Science Centre of the Russian Academy
of Sciences, Kirovsk, Russia "Institute for Biological Problems of Cryolithozone of the Siberian Branch of the Russian Academy
of Sciences, Yakutsk, Russia 10Institute of Biological Problems of the North of the Far Eastern Branch of the Russian Academy
of Sciences, Magadan, Russia Corresponding author. I. A. Galanina, gairka@yandex.ru
Abstract. The paper provides data on morphology, ecology, differentiation and distribution of the crustose, epiphytic and epixylic lichen Rinodina sibirica in Eurasia. An anatomical and morphological description of R. sibirica is provided based on 138 specimens studied. Based on these records, we show that the distribution of the species from the Russian Far East is ranging from the Kamchatka Peninsula to the Mongolian border in the south and the Krasnoyarsk Territory and the Tyumen Region in the west. Current data suggest that R. sibirica is an American-Asian species with a wide Amphiberingian range, only slightly extending to Europe in the Ural Mts. Keywords: biogeography, disjunction, range, Asia.
Лишайник Rinodina sibirica (Physciaceae) в Евразии
И. А. Галанина1, С. В. Чесноков2, 3, Д. Е. Гимельбрант3, 4, Е. А. Давыдов5, А. К. Ежкин6, Т. М. Харпухаева7, Л. А. Конорева2, 8, Е. С. Кузнецова3, 4, Л. Н. Порядина9, И. С. Степанчикова2, 4, Л. С. Яковченко1, Е. В. Желудева10
Федеральный научный центр биоразнообразия наземной биоты Восточной Азии ДВО РАН,
Владивосток, Россия
https://doi.org/1031111/nsnr/2021.55.2393
393
2Ботанический сад-институт ДВО РАН, Владивосток, Россия 3Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия 4Санкт-Петербургский государственный университет, Санкт-Петербург, Россия
5Алтайский государственный университет, Барнаул, Россия 6Институт морской геологии и геофизики ДВО РАН, Южно-Сахалинск, Россия 'Институт общей и экспериментальной биологии СО РАН, Улан-Удэ, Россия 8Полярно-альпийский ботанический сад-институт КНЦ РАН, Кировск, Россия 9Институт биологических проблем криолитозоны СО РАН, Якутск, Россия 10Институт биологических проблем севера ДВО РАН, Магадан, Россия Автор для переписки: И. А. Галанина, gairka@yandex.ru
Резюме. Статья посвящена морфологии, экологии и распространению накипного эпифит-ного и эпиксильного лишайника Rinodina sibirica в Евразии. Приведены отличия от наиболее сходных видов. Анатомо-морфологическое описание Rinodina sibirica дано на основе 138 изученных образцов. На основании полученных данных мы показываем распространение вида с Дальнего Востока России от полуострова Камчатка до границы с Монголией на юге, а также до Красноярского края и Тюменской области на западе. Полученные данные позволяют считать, что R. sibirica является американо-азиатским видом с широким амфиберингийским ареалом, лишь слегка затрагивающим Европу в пределах Уральских гор.
Ключевые слова: биогеография, дизъюнкция, ареал, Азия.
The distribution of some species of the genus Rinodina (Ach.) Gray is still poorly known in Eurasia. One of the biggest gaps in the Holarctic is the Asian part of Russia and Rinodina remains poorly studied in East Asia as a whole. Although, recent taxo-nomic treatments of Rinodina (Mayrhofer, Moberg, 2002; Giralt, 2010; Sheard, 2010; Sheard et al., 2011, 2017) have provided much new information on the distribution of its species. Sheard (2010) suggested that Rinodina sibirica H. Magn. may exhibit an example of a species with a broad Beringian type of distribution (Hulten, 1937).
This lichen was first named Rinodina sibirica by K. G. W. Lang about 1910 when he identified A. K. Cajander's specimens from Yakutia [see his handwriting on the labels kept in the Herbarium of Botanical Museum, University of Helsinki (H)]. But he never published it because he died in a railway accident in Sweden in 1911, and A. H. Magnusson possibly adopted the name from his labels (Teuvo Ahti, pers. comm.). Rinodina sibirica was described from collections made by Magnus Brenner in the vicinity of Yeniseysk, Tomsk, and Tobolsk (Siberia) on the bark of Prunuspadus L. and Sorbus sp. (Magnusson, 1936). Subsequently, the species was also mentioned on the bark of Salix caprea L., S.pentandra L., S. viminalis L., Betula sp., Larix sp. and on wood in several places along such rivers as the Yenisey, Ob, Irkut, Lena, and Aldan (Magnusson, 1947). Then for a long period R. sibirica was rarely mentioned in literature until it was found in Mongolia (Golubkova, 1981), in the Sokhondinsky State Reserve of the Trans-Baikal Territory (Insarov, Pchelkin, 1989), and the Altai and Sayan Mountains (Sedelnikova, 1990, 2001, 2007).
The species has also been recorded from the Baikalsky State Reserve (Urbanavichus, Urbanavichene, 2004), as well as from the Republic of Sakha (Yakutia) and the Magadan
Region (Kotlov, 2008). Unfortunately, specimens of Rinodina sibirica from the last regions were not found in the Herbarium of Komarov Botanical Institute of the Russian Academy of Sciences in St. Petersburg (LE) or the Herbarium of Institute of Biological Problems of the North FEB RAS in Magadan (MAG). In our latest studies, R. sibirica was noted in the Trans-Baikal and Khabarovsk Territories, in the Magadan Region, on Sakhalin Island and the Kamchatka Peninsula (Sheard et al, 2017).
The species was also found in North America in the forest-tundra and tundra on bark and wood (Thomson, 1997, as Rinodina granulans Vain., see Sheard, 2010).
After starting the study, we realized that Rinodina sibirica is very common in the Republic of Buryatia, the Trans-Baikal Territory and Republic of Sakha (Yakutia), but is often incorrectly identified by lichenologists, which leads to an incorrect understanding of its distribution. Therefore, an attempt was made to study the collections from the above-mentioned regions in more detail and compile a map of the range of R. sibirica in Eurasia. In this paper, we show a more complete distribution of this species and introduce many new localities for R. sibirica in Asian part of Russia. In addition, we made an anatomical and morphological description of R. sibirica based on the 138 samples studied.
Material and Methods
During the study, herbarium material stored in the herbaria of Russia (ALTB, LE, LECB, MAG, SASY, UUH, VLA), Finland (H), and the authors' collections from different regions of the Far East of Russia, Republic of Sakha (Yakutia), Trans-Baikal Territory, Republic of Buryatia were studied. The collections of A. V. Galanin from midstream of the Vilyui River in the Republic of Sakha (Yakutia) (the region of tukulans — sand dunes) were also studied. Also in this work, we studied two specimens collected by Brenner in 1876 from Yeniseisk in the Krasnoyarsk Territory and determined by Magnusson (LE). Specimens from all regions were also sent to John W. Sheard, for comparison with material from North America. To make the distribution map of Rinodina sibirica we used specimens that were previously determined by Sheard in the Herbarium of the University of Helsinki (H) which have not been published earlier. Materials were studied and identified by the first author in the laboratory of Botany of the Federal Research Center for Biodiversity of the Far East Branch of the Russian Academy of Sciences.
The investigation of anatomical and morphological features of R. sibirica was made using the microscopes Zeiss Axioplan 2 and Stemi 2000-C. The study of spore structures and their measurement were made using the immersion oil under the magnification of 1000 x. The majority of character ranges is given from minimal to maximal states of variation. The sizes of apothecia and spores are given in the same way, with the average (bolded) value. The number of measurements (N) is indicated in parentheses after the size of spores and apothecia.
Results and Discussion
The most detailed description of the species Rinodina sibirica was made by Sheard (2010) based on samples from North America. Below we present a description of
R. sibirica based on the 138 samples from Russia, which is closely comparable with that of Sheard (2010). The first description of R. sibirica made by Magnusson (1936) was based on a few specimens from only one region of Western Siberia. Our description of the species is based on numerous specimens from a large area of Russia where the species is widespread and found on various tree species.
Rinodina sibirica H. Magn., 1936, Svensk Bot. Tidskr. 30: 261. — Type: [Russia] Siberia, Jenisejsk, Kolmogorova, 59°30'N, 3 X 1876, M. Brenner 462c [S — lectotype, designated by Sheard, 2010: 184]. (Plates I, II)
= Rinodina sophodioides H. Magn., 1947, Acta Horti Gotob. 17: 226. — Holotype: [Russia, Siberia, Tomsk Region], Narym, 2 VI 1876, M. Brenner 654f, S.
Thallus thin, indefinite or more often discrete, dark gray to often buff or brown, consisting of scattered or merging irregular areoles, the size of the areoles is 0.1-0.4 mm wide, areole surfaces flat and often smooth, matte or sometimes shiny, prothallus and vegetative propagules absent or gray prothallus is rarely present (Plate I: 1-6).
Apothecia erumpent (young apothecia in the process of development break through the cortical layer and crater-like with a very wide base), becoming broadly attached and then narrow at the base (sessile), scattered, numerous and as a rule abundant on the thallus, 0.50-0.75(1.10) mm in diam. (N = 100); disc dark brown to black, flat at first, then becoming convex, sometimes hemispherical; thalline margin entire, often thin, first lighter than the thallus, then of the same color, up to 0.05-0.10 mm wide, permanent, sometimes excluded in apothecia with a strongly convex disc. Thalline margin 50-90 jm wide in the lateral part, (50)80-120 jm in the widest part closer to the base of apothecia (Plate II: 1); cortex 10-20 jm wide in the lateral part and up to 50 jm at the base; the epinecral layer 5-10 jm wide; cortex 10-20 jm wide laterally and to 40-45 jm at the base; crystals absent from cortex and medulla; cells of the cortex to 5-6 jm wide, usually pigmented; algal cells to 8-16 jm in diam.; proper exciple in the lateral part of apothecia to 10 jm wide, pigmented, brown, expanding to the top to 10-20(40) jm; hypothecium colorless to slightly yellowish (30)50-80(100) jm high; hymenium 80-110 jm high; paraphyses 1.5-2.5 jm wide, often conglutinate, apices of paraphyses to 3.0-5.5 jm wide, with dispersed pigment forming red-brown, bright epihymenium (Plate II: 6); asci 65-80 x 15-24 jm.
Ascospores eight per ascus (sometimes with asynchronous development), type A development (apical cell wall thickening occurs after division), Physcia-Physco-nia-type (Plate II: 2-5), (17.0)20.0-21.5(25.5) x (8.5)10.0-11.5(13.0) jm (N = 560). The lumina angular at the beginning of spore development, the cell wall at the apex and septum is thickened (Physcia-type) (Plate II: 5), then as the spores mature the lumina become rounded and the cell wall thinner (Plate II: 3, 4). However, mature spores with rounded lumina often retain slight apical thickening. The torus is well developed and the cell walls are dark-colored (Plate II: 2).
Chemistry. Spot tests all negative.
Plate I. Thallus and apothecia of Rinodina sibirica. 1 — thallus with typical flat scab-shaped areoles; 2, 4 — thallus with seldom convex areoles; 3, 6 — thallus with areoles merging to become continuous; 5 — gray prothallus is rarely present.
Scale bars: 1-6 — 0.5 mm.
Ecology. The species most often grows on the bark of Betula spp., also on the bark of Alnus spp., Chosenia arbutifolia (Pall.) A. K. Skvortsov, Picea spp., Populus suaveolens Fisch., Prunus padus, Salix spp., Sorbus sambucifolia (Cham. et Schltdl.) M. Roem., branches of Pinus spp., and also on wood in different types of forest: coniferous, mixed birch forests, sparse forests, on free-standing trees in steppe communities, and in floodplain forests in the northern parts of its distribution.
Notes. Rinodina sibirica is characterized by a developed thallus of separate irregularly shaped areoles, narrowly attached mature apothecia, with the disc often becoming hemispherical, while young apothecia can be "erumpent". Ascospores occupy an intermediate position between the Physcia- and Physconia-types, lumina with a narrow canal in immature spores. Mature spores are darkly pigmented with a well-developed torus.
The structure and type of spores make it possible to confuse R. sibirica with R. ar-chaea (Ach.) Arnold (thin lumina channels in young spores and a persistent, slightly apical thickening in mature spores), but R. sibirica is characterized by darker pigmentation of the spore walls and torus, and there is always a stage of development with Physcia-type lumina with a pronounced apical thickening. Rinodina archaea is more common on wood (lignicolous), and R. sibirica prefers tree bark and is less common on wood (corticolous) (Sheard, 2010).
Rinodina sibirica has a morphology similar to R. cinereovirens (Vain.) Vain. but the thallus of that species is typically light grey and never with the strong brown color of R. sibirica. The spores of R. sibirica are smaller than those of R. cinereovirens although of a similar broadly ellipsoid shape and also differ from the latter in belonging to the Physconia- rather than the Physcia-type. The lack of crystals (sphaerophorin) in the medulla of R. sibirica is also definitive. Sheard (2010) compared R. sibirica with corticolous forms of R. turfacea (Wahlenb.) Korb., all of which subsequently have been shown to belong to R. cinereovirens (Sheard et al, 2017).
Another species that can be confused with R. sibirica is R. laevigata (Ach.) Malme. In addition, they can be found together, for example, in southern Siberia. Rinodina laevigata has a less developed thallus, grayish color, smaller spores and although they are of the same type of Physcia-Physconia, their lumina is more rounded and less elongated. Apothecia in R. laevigata are also more broadly attached to the thallus, the discs
Plate II. The structure of apothecia and spores of Rinodina sibirica.
1 — section through the apothecium with a well-formed thalline margin with a brown cortex; 2 —
mature Physcia-Physconia-type spores of R. sibirica with rounded lumina and darkly pigmented walls, and tori; 3 — mature spores with darkly pigmented walls and mostly thin apical walls
(Physconia-type); 4 — ascus with asynchronous development spores of R. sibirica (a) and a second
ascus with colorless young spores (b) showing type A development (no apical thickening before
septum development); 5 — immature spores with angular Physcia-type lumina and narrow canal;
6 — red-brown epihymenium.
Scale bars: 1 — 50 |im; 2-6 — 10 |im.
are typical plane rather than convex as in R. sibirica. Also, R. laevigata usually has a thick lower apothecial cortex with intricate hyphae.
The spores R. sibirica might be confused with those of R. orculata Poelt ex M. Steiner, but the latter has smaller spores, (12.5)16.0-16.5(20.0) x (6.0)8.0-8.5(9.5) jm and the thallus is usually gray and its areoles are mostly convex.
Specimens examined are listed in the Supplement1.
Distribution of Rinodina sibirica in Eurasia. All known localities of the species in Eurasia are shown at the map (Fig. 1).
Fig. 1. Distribution of Rinodina sibirica in Eurasia. Localities of studied samples are marked with black circles, asterisks indicate the localities according to published data, and triangles shows specimens identified by J. W. Sheard in H.
Despite the fact that the species is widespread in Russia, it is found mainly in the boreal zone of the Asian part of Russia, including almost the entire Far East (except for the Primorye Territory and Chukotka Peninsula), Republic of Sakha (Yakutia), Republic of Buryatia, Krasnoyarsk, and Trans-Baikal Territories, Irkutsk, Tyumen, and Tomsk regions. The northernmost localities of the species in Eurasia are: in the west, Yenisei River [Dudinka village (69°40'N) and Nikandrovsky Island (70°20'N)]; in the east, Adycha River (68°25'N). How the species is distributed further in western Russia and in Europe remains to be explored. The westernmost records in Russia are according to the literary data in the Tyumen Region (Kataeva et al., 2005, as R. granulans; Kotlov, 2008), as well as in the southwest in Altai (Sedel-
1 The Supplement is available at the end of the article page on the journal website (https://doi.Org/10.31111/nsnr/2021.55.2.393).
nikova, 1990, 2001, 2007). The southernmost finds of the species in Mongolia are in the Khentei mountain-taiga and Mongolian-Dahurian mountain-forest-steppe zones (Golubkova, 1981), in Russia these are localities in the Trans-Baikal Territory and, in the Far East, in Khabarovsk Territory. In the Trans-Baikal Territory, R. sibirica is most often found on the bark of Betula spp. and here it is the most frequent and abundant species of the genus Rinodina. In the Khabarovsk Territory, the species is rare.
When moving north the species is often found in floodplain forests, on various tree species, whereas Magnusson (1947) noted, Rinodina sibirica becomes less common northwards. In the central part of the Republic of Sakha (Yakutia) the species is very common and abundant, in the Magadan Region it occurs less often. But it should be noted that Magadan Region requires additional research to understand how far this species goes to the north on the Far East. It should also be noted that, like in North America, the species does not occur in the upper Arctic of the Far East and of the Republic of Sakha (Yakutia) (Andreev et al., 1996; Sheard, 2010). In North America (Thomson, 1997, as R. granulans, see Sheard, 2010), the species is found in the forest-tundra and tundra (subarctic) in Alaska but not in the high latitudes of the Arctic or east of Hudson Bay.
According to Sheard (2010) distribution of Rinodina sibirica may correspond to either the bilateral radians of northern Beringia or the Arctic-Pacific group of Hulten (1937) but very much truncated from the limits of these ranges. Despite this, we see a significant distribution of the species in Siberia along the continental corridor, leaving for the territory of Mongolia. Sheard (2010) believes that R. sibirica is unlikely to have survived the Ice Age in refugia north of the ice cap because of its reliance on low Arctic arboreal and shrub substrates.
Analyzing the current distribution of Rinodina sibirica, we confirm the opinion of Sheard (2010) that the species spread from the west after the ice sheet. Based on all the data we believe that R. sibirica is an American-Asian species with a wide Amphi-beringian range, only slightly extending to Europe in the Ural Mts.
Acknowledgments
We are very grateful for the consultation and assistance in identifying samples to Dr. J. W. Sheard, the monographer of the genus Rinodina in North America. We are also grateful to Teuvo Ahti for help during our work in the Herbarium of the University of Helsinki and an interesting story about the origin of the name of the R. sibiri-ca. The study of E. A. Davydov, A. K. Ezhkin, I. A. Galanina, and L. S. Yakovchenko was partly supported by RFBR (grant 18-04-00098). The study of S. V. Chesnokov, D. E. Himelbrant, E. S. Kuznetsova, and I. S. Stepanchikova was supported by RFBR (grant 18-05-60093) and carried out within the framework of the institutional research project "Flora and taxonomy of algae, lichens and bryophytes in Russia and phytogeo-graphically important regions of the world" (no. 121021600184-6) of the Komarov Botanical Institute RAS. The work of S. V. Chesnokov and L. A. Konoreva was carried
out in the frame of the institutional research project "Cryptogamic biota of Pacific Asia: taxonomy, biodiversity, species distribution" of the Botanical Garden-Institute of Far Eastern Branch of the Russian Academy of Sciences. The research was carried out within the state assignment of Ministry of Science and Higher Education of the Russian Federation (AAAA-A21-121012190038-0, AAAA-A17-117122590002-0, and AAAA-A17-117062710098-4). The work of T. M. Kharpukhaeva was carried out in the frame of the institutional research project "Biota of terrestrial ecosystems of the Baikal region: composition, structure, ecological and geographical features" 0271-2021-0001 (FWSM-2021-0001) of the IGEB SB RAS and was partly supported by RFBR project 18-04-01068-a. The work of L. N. Poryadina was carried out within the framework of the institutional research project "Vegetation cover of the cryolitho-zone of taiga Yakutia: biodiversity, habitat-forming functions, protection and rational use" 0297-2021-0023 (AAAA-A21-121012190038-0).
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