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DNA Barcode and Phylogenetic Study of the Tribe Desmodieae (Fabaceae) in Korea

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DNA Barcode and Phylogenetic Study of the Tribe Desmodieae (Fabaceae) in Korea pISSN 1225-8318 − Korean J. Pl. Taxon. 49(3): 224 239 (2019) eISSN 2466-1546 https://doi.org/10.11110/kjpt.2019.49.3.224 Korean Journal of RESEARCH ARTICLE Plant Taxonomy DNA barcode and phylogenetic study of the tribe Desmodieae (Fabaceae) in Korea Dong-Pil JIN, Jong-Won PARK, Jong-Soo PARK and Byoung-Hee CHOI* Department of Biological Sciences, Inha University, Incheon 22212, Korea (Received 12 August 2019; Revised 24 September 2019; Accepted 26 September 2019) ABSTRACT: Species identification for the Korean tribe Desmodieae was conducted using the DNA barcoding genes rbcL, matK (from chloroplast DNA) and ITS (from nuclear ribosomal DNA). A total of 25 taxa (n = 75) in five genera were sequenced, and neighbor-joining trees were constructed using different combinations of DNA barcodes. When comparing these phylogenetic trees, a tree with all loci combined (rbcL + matK + ITS) showed the highest rate of identification success (72%). On this tree, two subtribes and five genera within the tribe were supported as monophyletic. In the Desmodiinae clade, Desmodium and Hylodesmum were more closely related to each other than to Ohwia. In the Hylodesmum clade, H. oldhamii was found to be a sister to H. podocarpum complex, and all taxa within the complex were identified successfully. Subsp. fallax, regarded as a variety of subsp. oxyphyllum, is closely clustered with subsp. podocarpum. Although var. mandshuricum has been regarded as a synonym of var. oxyphyllum, this taxon is supported as a distinct variety. For the Lespe- dezinae clade, all species of Kummerowia were monophyletic, while nine of 16 Lespedeza taxa were identified successfully. In particular, the resolution of Macrolespedeza (28.5%) was lower than that of Junceae (77.8%). Among the Lespedeza taxa, L. cuneata was distinguishable from L. lichiyuniae, despite morphological similar- ities. It has been suggested that both L. maritima and L. inschanica are hybrids. The former is thought to be an independent species. While it is difficult to determine whether the latter originated via hybridization, this study showed that it is closely related to L. juncea. Keywords: Desmodieae, species identification, DNA barcode, rbcL, matK, ITS DNA 바코드(DNA barcode)는 orthologous DNA 구간들 (Kress and Erickson, 2008; Meusnier et al., 2008). 하지만 의 짧은 염기서열을 이용하여, 형태적으로 식별하기 어 고등식물의 COI 유전자는 동물의 COI 유전자에 비해 진 렵거나 분류학적으로 문제가 있는 종들을 동정하는 기법 화속도가 느려 DNA 바코드 마커로 사용되기 어렵다 이다(Kress et al., 2005). DNA 바코드는 식물이나 균류 등 (Kress et al., 2005). 이러한 이유로 식물군에서는 다른 다양한 분류군들의 동정과 계통학적 연구에 이용되고 있 DNA 마커들이 탐색되었고, 일반적으로 엽록체 DNA의 4 다(Purty and Chatterjee, 2016). 이상적인 DNA 바코드 구간 개 구간(matK 유전자, rbcL 유전자, ycf1 유전자, psbA- 은 근연종의 식별을 해결할 수 있을 정도로 충분히 가변 trnH spacer)과 핵 ribosomal DNA의 internal transcribed 적이고 저렴한 비용으로 실험을 할 수 있을 정도로 짧아 spacer (ITS)가 마커로 추천되었다(Edwards et al., 2008; 야 하며, 여러 분류군들에 적용될 수 있어야한다(Dong et Hollingsworth et al., 2011; Dong et al., 2015; Vassou et al., al., 2015). 최근 20여년간 각 생물군에 적용할 수 있는 이 2015). 엽록체 DNA 중에서 rbcL과 matK 구간은 DNA 상적인 DNA 바코드 마커를 찾고자 많은 연구들이 수행되 barcode의 기본 마커로 쓰이는 구간이다(CBOL Plant 어 왔다(Hebert et al., 2003; Kress et al., 2005; CBOL Plant Working Group., 2009). rbcL 구간은 polymerase chain Working Group, 2009; Dong et al., 2012, 2015). 동물군의 경 reaction (PCR) 성공률이 높고 염기서열분석의 안정성이 우에는 DNA 바코드 마커로 미토콘드리아의 cytochrome c 좋으나, 종간 해상력이 떨어지는 단점이 있다(Renner, oxidase I (COI) 유전자가 일반적으로 이용되고 있다 1999; Salazar et al., 2003). 반대로 matK 구간은 PCR 성공 *Author for correspondence: [email protected] http://e-kjpt.org, © 2019 the Korean Society of Plant Taxonomists. This is an open-access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited. 224 DNA barcode of the tribe Desmodieae in Korea 225 률과 시퀀싱의 안정성은 상대적으로 떨어지나 종식별률 유무 등으로 도둑놈의갈고리아족(subtribe Desmodiinae)과 이 높은 것으로 알려져 있다(Hollingsworth, 2011). ycf1 구 싸리아족(subtribe Lespedezinae)으로 구별되는 것으로 알 간의 경우에는 높은 염기서열 변이를 보이지만 이로 인 려져 있다(Ohashi et al., 1981; Jabbour et al., 2018). 도둑놈 해 primer를 제작하는 데 어려움이 있다(Dong et al., 2015). 의갈고리족 식물들은 사료, 토양의 비옥화, 사면안정화 psbA-trnH 구간은 non-coding 구간으로 길이가 짧으며 시 등을 목적으로 심어지고, 일부 종들은 약용으로 사용되 퀀싱 안정성이 높고 rbcL이나 matK와 조합하였을 때 일 기도 하여(Ohashi, 2005), 식물자원으로 가치가 높다. 본 부 분류군에서 높은 종식별률을 보였다(Vassou et al., 족의 식물들을 포함한 콩과 식물의 분자계통학적 연구는 2015). ITS 구간은 식물의 분자계통학적 연구 및 DNA 바 여러 차례 수행된 바 있다(Kajita et al., 2001; Legume 코드 연구에 가장 많이 이용되는 구간 중 하나로(e.g., Phylogeny Working Group, 2013, 2017). 이러한 분자계통 Riahi et al., 2011; Youm et al., 2016; Choi et al., 2018; 학적 연구 결과들은 본 족이 콩아과(Papilionoideae) 내에 Jabbour et al., 2018), 엽록체 DNA에 비해 진화속도가 빠 단계통임을 잘 지지했으며, 두 개의 아족들도 각각 단계 르기 때문에 구간의 길이에 비해 계통학적으로 유용한 통을 형성하였다. 최근 도둑놈의갈고리족의 많은 속이 염기서열의 비율이 많은 것으로 알려져 있다(Wolfe et al., 포함된 분자계통학적 연구(Jabbour et al., 2018)에서 속간 1987). 유연관계와 분류체계에 대해 논의된 바 있다. 싸리아족 도둑놈의갈고리족(tribe Desmodieae)은 콩과(family 의 경우, 싸리속(Lespedeza Michx.)을 중심으로 형태분류 Fabaceae)에 속하고 32속 530여종으로 이루어진 것으로 학적 연구(Lee, 1965; Hatusima, 1967; Akiyama, 1988)가 수 알려져 있으나(Ohashi, 2005), 최근 형태 및 분자계통학적 행되었고, 최근 분자계통학적 연구는 싸리속내 종간 유 연구 결과에 따르면 39속으로 인식된 바 있다(Ohashi et 연관계 및 근연속들과의 유연관계 등에 대해 분석하였다 al., 2018a, 2018b). 본 족의 식물들은 초본부터 교목까지 (Han et al., 2010; Nemoto et al., 2010; Xu et al., 2012). 한편 도 다양한 생활형을 보이고(Ohashi, 2005), 이들은 주로 열대, 둑놈의갈고리아족은 도둑놈의갈고리속(Desmodium Desv.) 아열대, 온대지역에서 분포하나 일부 종들은 동아시아와 을 중심으로 분류학 및 계통학적 연구들이 수행되어 왔다 북아메리카의 한대와 아한대 지역에서 자란다(Ohashi, (Ohashi, 1973; Kajita and Ohashi, 1994; Kajita et al., 2001; 2005). 도둑놈의갈고리족은 열매의 형태와 엽병 아래에 Jabbour et al., 2018; Ohashi et al., 2018a, 2018b). 도둑놈의 달리는 탁엽의 유무, 기판 기부에 귀모양 부속체(auricle)의 갈고리속은 계통수상에서 다계통으로 확인되어왔는데, Fig. 1. Six genera of tribe Desmodieae in Korea. A. Desmodium heterocarpon, photo by W. B. Cho, 29 Aug 2009, Sallok road, Harye-ri, Jeju- do. B. Hylodesmum podocarpum subsp. podocarpum, photo by D. P. Jin, 30 Sep 2016, Isl. Oeyeon, Chungcheongnam-do. C. Ohwia caudata, photo by J. S. Park, 21 Aug 2017, Dongbaekdongsan, Jeju-do. D. Campylotropis macrocarpa, photo by D. P. Jin, 15 Sep 2017, Paljoryeong, Cheongdo-gun, Gyeongsangbuk-do. E. Kummerowia striata, photo by D. P. Jin, 4 Aug 2017, Buldang-ri, Gwangju-si, Gyeonggi-do. F. Lespedeza cuneata, photo by J. S. Park, 10 Aug 2018, Sorae Marsh Ecological Park, Incheon-si. Korean Journal of Plant Taxonomy Vol. 49 No. 3 (2019) 226 Dong-Pil JIN et al. 이를 반영하여 현재 본 속내 많은 분류군들이 새로운 속 애기도둑놈의갈고리[subsp. oxyphyllum var. mandshuricum 으로 분류되었다(e.g., Ohashi, 1999; Ohashi and Mill, 2000; (Maxim.) H. Ohashi & R. R. Mill]는 일부 개체들에서 심한 Ohashi and Ohashi, 2012, 2018a, 2018b; Ohashi et al., 2018a, 형태적 변이를 보이고 있다(Choi, 1991). 또한 엽록체 유 2018b). 전체의 제한효소 절편길이 다형성 분석에 기반한 계통학 한반도에는 2아족 6속의 도둑놈의갈고리족 식물 즉 도 적 연구에서 개도둑놈의갈고리 종내분류군간 한계가 뚜 둑놈의갈고리아족 내에 도둑놈의갈고리속, 갈고리속 렷하지 않은 것으로 나타났다(Kajita and Ohashi, 1994). (Hylodesmum H. Ohashi & R. R. Mill), 된장풀속(Ohwia H. 따라서 본 연구는 한국산 도둑놈의갈고리족 식물들을 Ohashi) 3속, 싸리아족 내에 꽃싸리속(Campylotropis 대상으로 DNA 바코드를 이용한 종식별과 이들의 계통 Bunge), 매듭풀속(Kummerowia Schindl.), 싸리속 3속이 분 학적 유연관계 대해서 논의하고자 하였다. 포하는 것으로 알려져 있다(Choi, 2007) (Fig. 1). 한편 최 근 잔디갈고리의 학명은 Desmodium heterocarpon (L.) 재료 및 방법 DC.에서 Grona heterocarpa (L.) H. Ohashi & K. Ohashi로 변경된 바 있다(Ohashi and Ohashi, 2018a). 이들 중 싸리속 재료 수집 과 갈고리속은 상대적으로 많은 분류군들을 포함하고 있 본 연구에 사용된 도둑놈의갈고리족 식물은 국내에 자 으며 종간 분류에 어려움이 있다. 한국에 자라는 싸리속 생하고 있는 종을 대상으로 선정하여 총 25종 75개체가 식물들은 땅비수리절[section Junceae (Maxim.) H. Ohashi 수집되었다(Appendix 1). 수집된 잎은 신선한 잎을 선별 & T. Nemoto]과 싸리절(section Macrolespedeza Maxim.)로 하여 DNA 추출을 위해 실리카겔에 건조시켰다. 본 연구 구분되는데(Ohashi and Nemoto, 2014), 땅비수리절 식물 에서 포함된 분류군들은 개체 및 집단 간 염기서열에 의 들은 개방화와 폐쇄화가 모두 달리는데 비해 싸리절 식 한 차이와 무관하게 계통수상에서 분류군 별로 동정이 물들은 개방화만 달리는 특징으로 구분된다. 싸리절 식 가능한지 보기 위해, 분류군 당 3개체씩 서로 다른 곳에 물들은 주로 녹화용으로 쓰이며, 땅비수리절 식물들은 서 수집하였다. 일부 같은 장소에서 채집된 분류군들의 비료나 거름 등으로 쓰인다. 땅비수리절에 속하는 비수 경우에는 10 m 이상 거리를 띄어 같은 개체가 채집되는 리[L. cuneata (Dum. Cours.) G. Don]는 야관문이라는 약용 것을 피했다. 채집된 재료는 국립생물자원관 관속식물 명으로도 알려져 있는데, 최근 민간의 관심을 받으며 식 표본 수장고(KB)와 인하대학교 식물표본관(IUI)에 수장 품이나 약재자원으로 연구되고 있다(Chung and Cheong, 하였다. 2016). 하지만 본 속의 식물들은 큰 형태적 변이폭을 보이 며 간혹 종간 한계가 명확하지 않다. 이러한 다양한 형태 DNA 추출 및 PCR 적 변이는 학자에 따라 많은 종내분류군이나 독립종으로 염기서열 분석을 위해 채집된 식물들의 건조된 잎에서 인식되었고(Nakai, 1927; Lee, 1965, 1980; Hatusima, 1967; DNA를 추출하였으며, DNA 추출에는 DNeasy Plant Mini Akiyama, 1988), 청비수리[L. inschanica (Maxim.) Schindl.] Kit (Qiagen, Seoul, Korea)를 이용하였다. 추출된 DNA는 와 해변싸리(L. maritima Nakai) 등이 잡종으로 처리되었 2% agarose gel에 전기영동하여 유무와 상태를 확인하였 다(Lee, 1965, 1980). 따라서 외부형태에 따라 본 속의 식 고, DNA의 농도를 NanoDrop ND-1000 (NanoDrop 물들을 동정하는 것은 어려운 것으로 알려져 있다.
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