The Lichen Genus Rinodina (Physciaceae, Caliciales) in North-Eastern Asia
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The Lichenologist 49(6): 617–672 (2017) © British Lichen Society, 2017 doi:10.1017/S0024282917000536 The lichen genus Rinodina (Physciaceae, Caliciales) in north-eastern Asia John W. SHEARD, Alexander K. EZHKIN, Irina A. GALANINA, Dmitry HIMELBRANT, Ekaterina KUZNETSOVA, Akira SHIMIZU, Irina STEPANCHIKOVA, Göran THOR, Tor TØNSBERG, Lidia S. YAKOVCHENKO and Toby SPRIBILLE Abstract: Rinodina is a widespread, polyphyletic genus of crustose Physciaceae with c. 300 species worldwide. A major missing link in understanding its global biogeography has been eastern Asia where the genus has never been systematically revised. Here we review specimen and literature records for Rinodina for north-eastern Asia (Russian Far East, Japan and the Korean Peninsula) and recognize 43 species. We describe two species, R. hypobadia and R. orientalis,asnewtoscience.Rinodina hypobadia is distinguished by its pigmented hypothecium, Dirinaria-type ascospores and pannarin in both thallus and epihymenium. Rinodina orientalis is characterized by its erumpent apothecia that remain broadly attached, with discs sometimes becoming convex and excluding the thalline margins, ascospores belonging to the Physcia-type and secondary metabolites absent. Nine other species are reported from the region for the first time. These include R. dolichospora, R. freyi, R. metaboliza, R. sicula, R. subminuta and R. willeyi.Of particular biogeographical interest are three additional new records that have western North American– eastern Asian distributions: the corticolous species R. endospora, R. macrospora and R. megistospora.Six species have the better known eastern North American–eastern Asian distributions: R. ascociscana (syn. R. akagiensis, R. melancholica), R. buckii, R. chrysidiata, R. subminuta, R. tenuis (syn. R. adirondackii)and R. willeyi, and two have eastern North American–eastern Asian–European distributions: R. excrescens and R. moziana (syn. R. destituta, R. vezdae). Our study begins to close one of the largest gaps in our knowledge of circumboreal species distributions in Rinodina and, together with previous studies in North America and Europe, provides new insights into circumboreal crustose lichen biogeography. Rinodina cinereovirens (syn. R. turfacea var. cinereovirens) is also reported as new to North America. Keywords: biogeography, disjunctions, lichenized Ascomycetes, new species, synonyms, vicariance Accepted for publication 3 June 2017 Introduction boreal forest. Eastern Asia has long con- No crustose lichen genus in the Holarctic has stituted a large knowledge gap in the biogeo- been revised in every major sector of the graphy of lichens, especially crustose lichens. J. W. Sheard: Department of Biology, University of A. Shimizu: Graduate School of Environmental Earth Saskatchewan, 112 Science Place, Saskatoon, SK S7N Science, Hokkaido University, Hokkaido, 060-0810, 5E2, Canada. Email: [email protected] Japan. A. K. Ezhkin: Institute of Marine Geology and Geo- G. Thor: Department of Ecology, Swedish University physics, Nauki Str., 1B, Yuzhno-Sakhalinsk, 693022, of Agricultural Sciences, P.O. Box 7044, SE-750 07 Russia. Uppsala, Sweden. I. A. Galanina and L. S. Yakovchenko: Federal Scien- T. Tønsberg: Department of Natural History, tific Center of East Asian Terrestrial Biodiversity, Far University Museum, University of Bergen, Allégaten Eastern Branch of the Russian Academy of Sciences, 41, P.O. Box 7800, N-5020 Bergen, Norway. Vladivostok Str. 100-leta, Vladivostok 159, 690024, Russia. T. Spribille (co-corresponding author): Institute of D. Himelbrant, E. Kuznetsova and I. Stepanchikova: Plant Sciences, Karl-Franzens-Universität Graz, Department of Botany, St. Petersburg State University, Holteigasse 6, A-8010 Graz, Austria; current address: Universitetskaya Emb. 7–9, St. Petersburg 199034, Biological Sciences CW405, University of Alberta, Russia; Laboratory of Lichenology and Bryology, Edmonton, AB T6G 2E9, Canada. Email: spribill@ Komarov Botanical Institute RAS, Professor Popov ualberta.ca St. 2, St. Petersburg 197376, Russia. Downloaded from https://www.cambridge.org/core. University of Montana, on 15 Nov 2017 at 02:01:46, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0024282917000536 618 THE LICHENOLOGIST Vol. 49 This is all the more unfortunate due to the oreina (Ach.) Norman, which the authors central role that eastern Asia is known to have included in Rinodina) reported in the only played in the evolution of higher plants two Korean checklists published to date (Wen 1999; Xiang et al. 2000; Qian 2002), (Hur et al. 2005; Moon 2013). Rinodina and the need for testable hypotheses in lichen chrysidiata and R. buckii were added to the list biogeography that include eastern Asia. of known lichen biota by Lendemer et al. Rinodina (Ach.) Gray is one of the better (2012) and Sheard et al. (2012), respectively. understood genera across the Holarctic, Joshi et al. (2013) reported R. oleae Bagl. as modern monographic treatments being new to South Korea. Kondratyuk et al. available for the Iberian Peninsula (Giralt (2013) reported R. fimbriata Körb., R. oleae 2001), Scandinavia (Mayrhofer & Moberg (as new to Korea a second time), R. polyspora 2002), North America (Sheard 2010) and the Th. Fr., R. pyrina (Ach.) Arnold, R. sophodes saxicolous species of the Old World (Mayr- (Ach.) A. Massal. and R. teichophila (Nyl.) hofer 1984). Though polyphyletic (Helms Arnold as new to South Korea. Aptroot & et al. 2003; Kaschik 2006; Nadyeina et al. Moon (2014) reported a further four 2010; Resl et al. 2016), species-level concepts species: R. cinereovirescens (Harm.) Zahlbr. R. within Rinodina are becoming increasingly laevigata (Ach.) Malme, R. placynthielloides well established and clear outlines of species Aptroot and R. xanthomelana Müll. Arg. groups have begun to emerge in conjunction Finally, Kondratyuk et al. (2016) reported with a refined understanding of crustose- R. xanthophaea (Nyl.) Zahlbr., making a total macrolichen transitions. Particularly clear of 14 species for Korea. patterns of biogeography have come into focus Twenty-three species of Rinodina are with the availability of molecular data from accepted as occurring in the three main sectors different parts of the world (Resl et al. 2016). of the Russian Far East (arctic, northern and The present paper is a first attempt at an southern forested) in the most recent Russian inclusive treatment of the genus in north- checklist (Urbanavichus & Andreev 2010). eastern Asia. However, this tally missed or discounted pre- The Japanese lichen biota has long been 2010 reports from the “far eastern” sectors of one of the most studied in Asia but its crustose Russia of R. gennarii Bagl. (Bredkina et al. lichen biota is still poorly known. Reports of 1992; Chabanenko 2002), R. polyspora and 25 and 22 species of Rinodina, respectively, in R. trevisanii (Hepp) Körb. (Chabanenko two earlier checklists (Kurokawa 2003; 2002). Two further species, R. kozukensis Harada et al. 2004) were trimmed back in the (Vain.) Zahlbr. and R. teichophila,wereadded synonymized list of Kurokawa & Kashiwadani by Skirina (2010), while Galanina et al. (2011) (2006), who collated data showing that reported R. excrescens Vain. from numerous several taxa belong in other genera (e.g. localities in the Russian Far East, Sheard et al. Buellia) and accepted 18 species of Rinodina (2012) reported R. buckii from the region, for Japan. Two new corticolous species have Lendemer et al. (2012) included Russian recently been added to the lichen biota of localities in their description of R. chrysidiata, Japan, R. chrysidiata Sheard (Lendemer et al. and Urbanavichene & Palice (2016) reported 2012) and R. buckii Sheard (Sheard et al. R. efflorescens from the Stanovoye Nagor’e 2012), bringing the total to 20 species. This Mountains, making a total of 32 previously compares to 33 species for the British Isles reported species. (Giavarini et al. 2009), to which should be added R. intermedia Bagl. (Mayrhofer et al. 2001), making a total of 34 species for a Materials and Methods comparable maritime archipelago only 65% of the area of Japan. This study is based primarily on collections by the authors, with the exception of JWS. Type specimens of Fewer species have been reported from all corticolous species described from Japan have been Korea. Rinodina membranifera (Hue) Zahlbr. examined. For Japan, the northern island of Hokkaido was the one Rinodina (except Dimelaena has been the region most densely sampled. In far eastern Downloaded from https://www.cambridge.org/core. University of Montana, on 15 Nov 2017 at 02:01:46, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0024282917000536 2017 Rinodina in north-eastern Asia—Sheard et al. 619 Russia, collections have primarily been made from Of these, we record nine species for the first Zabaikalskiy Krai, Khabarovskiy Krai, Primorskiy Krai, time from the region: R. dolichospora Malme, Sakhalin Island and the Kamchatka Peninsula. The vast R. endospora Sheard, R. freyi H. Magn., majority of collections examined were newly made. Unfortunately, it was not possible to freely borrow R. macrospora Sheard, R. megistospora Sheard material that is the basis for previous reports in the & H. Mayrhofer, R. metaboliza Vain., literature, such as in Chabanenko (2002) or Skirina R. sicula H. Mayrhofer & Poelt, R. subminuta (2010), because of onerous restrictions on the mailing of H.