J. Jpn. Bot. 89: 346–354 (2014)

Nephroma squamigerum (, Lichenized ) Is a Distinct Species

a, b c Katja Fedrowitz *, Andreas Frisch , Ulla Kaasalainen b and Yoshihito Ohmura

aDepartment of Ecology, Swedish University of Agricultural Sciences (SLU), P.O. 7044, 750 07, Uppsala, SWEDEN; bDepartment of Botany, National Museum of Nature and Science, 4-1-1, Amakubo, Tsukuba, 305-0005 JAPAN; cDepartment of Biosciences, University of Helsinki, P.O. 65, 00014, Helsinki, FINLAND *Corresponding author: [email protected]

(Accepted on May 10, 2014)

We evaluated the taxonomic status of squamigerum in Japan, a taxon currently ascribed as a forma to N. bellum. Using an extensive data set of herbarium and freshly collected specimens we confirm that N. squamigerum differs morphologically, chemically and genetically from N. bellum and other Nephroma species, and should be treated as a distinct species. Nephroma squamigerum is morphologically similar to N. bellum and to glabrous morphotypes of N. helveticum and may have been frequently overlooked, especially in other parts of eastern Asia. However, the absence of substances in N. squamigerum always allows a clear identification. Also, N. squamigerum can be clearly distinguished from all other Nephroma species based on the sequences of ITS rDNA and mtSSU rDNA. Neotypification of N. squamigerum is made. A key to the Japanese taxa of Nephroma is provided.

Keywords: Asia, ITS rDNA, mtSSU, neotype, , triterpenoids.

The lichen Nephroma Ach. White and James 1988), the morphological (Nephromataceae, lichenized Ascomycota) is delimitations and of some variable almost cosmopolitan in its distribution. The and widespread taxa like N. bellum, N. highest species diversity can be found especially helveticum, N. laevigatum, N. resupinatum, and in oceanic to boreal-montane areas of both N. tropicum are unresolved to date (Asahina hemispheres (James and White 1987, White 1962, Lohtander et al. 2002, Piercey-Normore and James 1988, Smith et al. 2009). The genus et al. 2006, Louwhoff 2009, Sérusiaux et al. includes 36 species worldwide (Kirk et al. 2008), 2011, Fedrowitz et al. 2012a). Such inadequate and nine species including four infraspecific taxa knowledge of species may mask species’ have been reported from Japan (Kurokawa and endangered status and put them at extinction Kashiwadani 2006). risk since their occurrences cannot be reliably Despite extensive research on the genus monitored (Stork 1993). Nephroma species are Nephroma (e.g., Inumaru 1940, Wetmore sensitive to sulfur dioxide pollution and often 1960, Asahina 1962, James and White 1987, negatively affected by intensive forestry, and the

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2314. of the Environment and Finnish Environment Institute, Inumaru S. 1940. Studia Licheneum Japoniae II. Helsinki. Peltigeraceae Japoniae II. Nephroma Acharius Grex 2. Sérusiaux E., Villarreal A. J. C., Wheeler T. and Goffinet Glabrae. Acta Phytotax. Geobot. 9: 51– 63. B. 2011. Recent origin, active speciation and dispersal James P. W. and White F. J. 1987. Studies on the genus for the lichen genus Nephroma (Peltigerales) in Nephroma I. The European and Macaronesian species. Macaronesia. J. Biogeogr. 38: 1138–1151. Lichenologist 19: 215–268. Smith C. W., Aptroot A., Coppins B. J., Fletcher A., Gilbert Kirk M., Cannon P. F., Minter D. W. and Stalpers J. A. O. L., James P. W. and Wolseley P. A. (eds.) 2009. The (eds.) 2008. Ainsworth & Bisby’s Dictionary of the of Great Britain and Ireland. London: The Fungi, 10th Edition. CAB International, Wallingford. British Lichen Society. Kurokawa S. and Kashiwadani H. 2006. Checklist of Stork N. E. 1993. How many species are there? Biodivers. Japanese Lichens and Allied Fungi. National Science Conserv. 2: 215–232. Museum Monographs No. 33. National Science Vilgalys R. and Hester M. 1990. Rapid genetic Museum, Tokyo. identification and mapping of enzymatically amplified Kuusinen M. 1996. Cyanobacterial macrolichens on ribosomal DNA from several Cryptococcus species. J. Populus tremula as indicators of forest continuity in Bacteriol. 172: 4238–4246. Finland. Biol. Cons. 75: 43–49. Vitikainen O. 2007. Nephromataceae. Nordic Lichen Flora Lohtander K., Oksanen I. and Rikkinen J. 2002. A 3: 91–95. Uddevalla. phylogenetic study of Nephroma (lichen-forming Wetmore C. 1960. The lichen genus Nephroma in North Ascomycota). Mycol. Res. 106: 777–787. and Middle America. Publ. Mus. Michigan State Univ., Louwhoff S. H. J. J. 2009. Nephromataceae. In: McCarthy Biol. Ser. 1: 369–452. P. M., Flora of Australia vol. 57. Lichens 5. pp. 423– White F. J. and James P. W. 1988. Studies on the Genus 427. ABRS and CSIRO Publishing, Canberra. Nephroma II. The Southern Temperate Species. Nitare J. (ed.) 2000. Indicator Species for Assessing the Lichenologist 20: 103–166. Nature Conservation Value of Woodland Sites: A White T. J., Bruns T., Lee S. and Taylor J. 1990. Flora of Selected Cryptogams. Skogsstyrelsen Förlag, Amplification and direct sequencing of fungal Jönköping. ribosomal RNA genes for phylogenetics. In: Innis M. Orange A., James P. W. and White F. J. 2001. A., Gelfand D. H., Sninsky J. J. and White T. J. (eds.), Microchemical Methods for the Identification of PCR Protocols: A Guide to Methods and Applications. Lichens. 101 pp. British Lichen Society. pp. 315–322. Academic Press, San Diego. Piercey-Normore M. D., Coxson D., Goward T. and Wirth V. 1995. Die Flechten Baden-Württembergs, Teil 1 & Goffinet B. 2006. Phylogenetic position of a Pacific 2. Eugen Ulmer GmbH & Co., Stuttgart. Northwest North American endemic cyanolichen, Zoller S., Scheidegger C. and Sperisen C. 1999. PCR Nephroma occultum (Ascomycota, Peltigerales). primers for the amplification of mitochondrial Lichenologist 38: 441–456. small subunit ribosomal DNA of lichen-forming Rassi P., Hyvärinen E., Juslén A. and Mannerkoski I. (eds.) ascomycetes. Lichenologist 31: 511–516. 2010. The 2010 Red List of Finnish Species. Ministry

K. Fedrowitza,A. Frischb,U. Kaasalainenc,大村嘉人 b: コモチウラミゴケ(ウラミゴケ科,地衣化子嚢菌)は独 立種である コモチウラミゴケ Nephroma squamigerum は,地衣 核リボゾーム遺伝子 ITS 領域および mtSSU に基づく分 体腹面にトメンタがなく,小裂片を地衣体背面および周 子系統解析を行った.コモチウラミゴケは単一クレード 縁につけ,子器を有する裂片には小裂片ができるかまた を形成し,ナメラウラミゴケと姉妹群となったが,ウラ は全縁となる.ナメラウラミゴケ N. bellum やウラミゴ ミゴケモドキとの直接的な系統関係は示されなかった. ケモドキ N. helveticum と形態的に中間的な個体もしば 以上,形態的,化学的,分子系統学的結果より,コモチ しば見られるが,それらの種からはテルペン類などの地 ウラミゴケは独立種として扱うのが妥当であると結論 衣成分が含まれないことで区別できる.コモチウラミゴ づけた. ケは独立種とする見解がある一方で,形態的連続性から (a スウェーデン農科大学生態学部, ナメラウラミゴケの品種とする見解もある.本研究で b 国立科学博物館植物研究部, は,コモチウラミゴケの分類学的位置を明らかにするた c フィンランド・ヘルシンキ大学生物科学部) めに,関連種との形態および化学成分の比較に加えて,