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A Taxonomic Study of Astragalus Sikokianus with a Disjunct Distribution Between Northwestern China and the Korea-Japan Region

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A Taxonomic Study of Astragalus Sikokianus with a Disjunct Distribution Between Northwestern China and the Korea-Japan Region J. Jpn. Bot. 91 Suppl.: 217–230 (2016) A Taxonomic Study of Astragalus sikokianus with a Disjunct Distribution between Northwestern China and the Korea-Japan Region In-Su CHOI, Dong-Pil JIN and Byoung-Hee CHOI* Department of Biological Sciences, Inha University, Incheon 22212, KOREA *Corresponding author: [email protected] (Accepted on January 9, 2016) Astragalus sikokianus has been known as endemic to the Korea-Japan region but is taxonomically confused with A. koraiensis from Korea and A. bhotanensis from Bhutan and western China. To clarify its taxonomic boundary and distribution, we examined external morphological characters and nrITS sequence data. Morphologically, A. koraiensis and A. sikokianus are hardly distinguishable from each other but they differ from A. bhotanensis of Bhutan and southwestern China in their leaves, leaflets, peduncles, corollas, calyx, and legumes. In contrast, the diagnostic characters of plants known as A. bhotanensis from northwestern China match those of A. sikokianus. Our TCS network and ML tree based on ITS sequence variations also recognized the plants from northwestern China, Korea, and Japan as being an independent evolutionary lineage that is distinct from the other group of A. bhotanensis from southwestern China. Therefore, this morphological and molecular evidence leads us to conclude that A. koraiensis is synonymized with A. sikokianus and, as delimited here, the latter species also includes plants from northwestern China. It is also possible that this disjunct distribution of newly circumscribed A. sikokianus can be explained by historical geographic changes and long-distance dispersal in East Asia. Key words: Astragalus koraiensis, Astragalus sikokianus, Astragalus bhotanensis, disjunct distribution, taxonomy. Many species of Astragalus L. (Fabaceae; 136 sections. However, the taxonomic status of Papilionoideae; Galegeae) are narrow endemics some regionally endemic species is still unclear. that often prefer marginal habitats or specialized Especially, the Astragalus in East Asia has been substrates (Sanderson and Wojciechowski 2000). poorly examined since that region is peripheral Although this diversity largely arises from to its center of species diversity. those narrow endemics, some of those species Astragalus sikokianus Nakai has been are often merged with others as synonyms or known as a perennial herb distributed in coastal varieties. Therefore, the number of species areas of Korea and Japan (Kim et al. 2003). belonging to Astragalus ranges from 2,000 to This species was first described based on a 3,000 (Lock and Simpson 1991, Mabberley type specimen collected from Naruto City, 1997, Maassoumi 1998). Work by Podlech and Tokushima Prefecture, Shikoku District, Japan Zarre (2013) has led to a revision of Old World (Nakai 1953). However, the population in that Astragalus as having 2,398 taxa that belong to locality has become extinct and other natural —217— 218 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue habitats have not been found in Japan. Since taxonomic examinations (S. Y. Kim 2004, then, this species has been considered a Japanese Podlech and Zarre 2013, Song and Heo 2014, endemic but naturally extinct species (Ohashi Choi et al. 2015) are also congruent with close 1982, 2001) and is now designated as an extinct affinity among plants from western China, species in the wild on the Japanese Plant Red Korea, and Japan. Nevertheless, the taxonomic List (Japanese Ministry of the Environment circumscription and systematic position of these 2012). In Korea, however, Kim et al. (2003) species remain controversial. have reported its disjunct distribution within The internal transcribed spacer (ITS) some costal populations. Therefore, because of regions of nuclear ribosomal DNA have its classification as extinct in Japan but its newly been used in phylogenetic investigations discovered presence in Korea, the natural habitat of Astragalus (Wojciechowski et al. 1999, of A. sikokianus was thought to be restricted to Kazempour Osaloo et al. 2003, 2005) because Korea. they show higher evolutionary rates and are For taxonomic purposes, S. Y. Kim (2004) easy to amplify. In addition, the 1,250 ITS hypothesized that the Korean endemic species sequence accessions (http://www.ncbi.nlm. A. koraiensis Y. N. Lee, first described from nih.gov/nuccore/?term=Astragalus) belonging mountainous area of Gangwon-do (Lee 1981), to approximately 400 species of Astragalus should be merged to A. sikokianus as a synonym represent the most abundant collection of and that the latter (including A. koraiensis) sequences among DNA regions that can be has close affinity to A. bhotanensis Baker, applied to the phylogeny of this genus (see Zarre which is distributed from Bhutan to western and Azani 2013). Therefore, we selected the China. The closeness of A. sikokianus with A. ITS region for conducting the first phylogenetic bhotanensis and A. koraiensis has also been re- investigation of A. sikokianus. confirmed from a morphological study based Taxonomic study of endangered species is primarily on fruit characteristics (Song and Heo a prerequisite for their conservation. Moreover, 2014). However, the taxonomic revision of Old the disjunct distributions proposed for such World Astragalus (Podlech and Zarre 2013) taxonomically problematic species have not has separately assigned A. sikokianus to sect. previously been confirmed through molecular Uliginosi A. Gray as an independent species analysis. Thus, our research objective was to and A. koraiensis to sect. Brachycephalae N. delimit the taxonomic boundary of this species D. Simpson as a synonym of A. bhotanensis. by evaluating several morphological characters According to this treatment, A. bhotanensis and and by developing a molecular phylogeny based A. sikokianus coexist in Korea but also have on ITS regions relevant to A. bhotanensis, A. disjunct distributions from the Korean Peninsula koraiensis, and A. sikokianus. to western China and Shikoku of Japan. However, in our recent taxonomic study of Materials and Methods Korean Astragalus, we found no morphological Morphological observations and differences between Korean and Japanese distributional surveys plants nor between mountain and coastal Samples from various Astragalus species plants (i.e., A. koraiensis and A. sikokianus) were collected from 2002 to 2015. Voucher (Choi et al. 2015). Furthermore, we detected specimens are held in the Herbarium of morphological similarities between specimens Inha University (IUI). To investigate their identified as A. bhotanensis from northwestern morphological characters and geographical China and A. sikokianus (incl. A. koraiensis) in distribution, we also examined specimens of A. the Korea–Japan region. The results of various bhotanensis, A. koraiensis, and A. sikokianus December 2016 Choi et al.: Astragalus sikokianus 219 Table 1. Details of Astragalus samples sequenced for ITS. Numbers in the parentheses indicate the number of individuals Taxon Geographic origin Ribotype A. uliginosus Mt. Baekdu, Korea AULI (1) A. schelichowii Mt. Baekdu, Korea ASCH (1) A. sikokianus Tokushima Pref., Shikoku, Japan A (3) A. sikokianus Pohang-si, Gyeongsangbuk-do, Korea A (1) A. koraiensis Jeongseon-gun, Gangwon-do, Korea A (1), B (2) A. sikokianus Samcheok-si, Gangwon-do, Korea B (1) A. koraiensis Taebaek-si, Gangwon-do, Korea B (2) A. sikokianus Isl. Ganghwado, Incheon, Korea B (1) A. sikokianus Mao County, Sichuan, China C (1) A. sikokianus Jinchuan County, Sichuan, China C (1) A. bhotanensis Zhaojue County, Sichuan, China D (1) A. bhotanensis Weining County, Guizou, China E (1) A. bhotanensis Dafang County, Guizou, China E (1) from the following herbaria: Chonnam National from the A. sikokianus and related species from University (CNU); National Institute of eastern Asia (Table 1). All PCRs were conducted Biological Resources, Korea (KB); Kyungpook with a GeneAmp® PCR System 2700 Thermal National University (KNU); Kangwon National Cycler (Applied Biosystems). Each reaction University (KWNU); Korea National Arboretum mixture contained 200 μM dNTPs (GeneCraft), (KH); Sungkyunkwan University (SKK); Seoul 1x PCR buffer with 1.5 mM MgCl2, 1 U of Taq National University, College of Agriculture DNA polymerase (TaKaRa), 10 ng of DNA, Life Sciences (SNUA); Tohoku University and an appropriate concentration of primers in a (TUS); the University of Tokyo (TI); The New total volume of 50 μL. Conditions included an York Botanical Garden (NY); and The Chinese initial denaturation at 94°C for 2 min; followed Academy of Sciences, Beijing (PE). We also by 35 cycles at 94°C for 30 s, 52°C for 45 s, and studied photograph of Chinese specimens from 72°C for 1 min; with a final extension at 72°C the herbaria of Chengdu Institute of Biology for 10 min. The PCR products were visualized (CDBI); South China Botanical Garden (IBSC); on 2% agarose gels, purified by PCRquick- Kunming Institute of Botany (KUN); Northwest spinTM (Intron), and sequenced with an ABI Institute of Plateau (QTPMB); Chongqing 3100 Genetic Analyzer and an ABI BigDyeTM Municipal Academy of Chinese Materia Medica Terminator Cycle Sequencing Ready Reaction (SM); Northwestern Institute of Botany (WUK) Kit (Applied Biosystems). through the Chinese Virtual Herbarium (www. cvh.org.cn). Our analytical methods followed Phylogenetic analyses those we have previously described (Choi et al. All sequences determined in this study 2015). were aligned by using ClustalW software (Thompson et al. 1994) implemented in DNA extraction,
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