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Taxonomic Study of Korean Scirpus L. s.l. (Cyperaceae) II: Pattern of Phenotypic Evolution Inferred from Molecular Phylogeny

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Abstract

Scirpus L. s.l. is well known as one of the polyphyletic groups in Cyperaceae. Recent molecular phylogenetic studies clearly suggested that Scirpus s.l. should be separated into several independent genera. In this study, we intend to present the morphological variations and patterns of phenotypic evolution based on molecular phylogeny of Korean Scirpus s.l. Five genera, including 21 taxa from Korean Scirpus s.l., were examined: three species of Bolboschoenus, three species of Schoenoplectus, eight species of Schoenoplectiella including one hybrid, five species of Scirpus, and two species of Trichophorum. For morphological analyses, 23 and 48 characters were selected from vegetative and reproductive organs, respectively. Molecular phylogeny was inferred from a nuclear ribosomal internal transcribed spacer, the chloroplast rbcL gene, and the chloroplast trnL-F region. Distinct characteristics and quantitative variation was presented for identification of the five genera and their species. A pronounced pattern of morphological character change was reduction, although many other character states seem to be homoplastic. We suggest that the reduction of phenotypic characteristics has been expressed in terms of condensation of internodes, reduction of leaf blade, and simplification of inflorescence among five genera of Korean Scirpus s.l.

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Acknowledgments

We thank the curator at the Sungshin Women’s University herbarium for permitting the examination of specimens. This work was supported by National Research Foundation of Korea grant funded by the Korean Government (MEST; NRF-2009-0088457) and is also a part of a Ph.D. dissertation for the first author in the Graduate School of Ajou University.

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Correspondence to Hong-Keun Choi.

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Appendices

Appendix 1

Specimens used in investigation of morphological characters and molecular analysis. Acronym of herbarium is showed only at the specimens from herbarium of Sungshin Women’s University (SWU). The rest specimens were deposited in herbarium of Ajou University (AJOU). Asterisks indicate materials for total genomic DNA extraction. GenBank accession numbers (nrITS/rbcL/trnL-F) are provided in parenthesis. Previous published sequences of nrITS and rbcL are marked in italic (Jung and Choi 2010).

Bolboschoenus fluviatilis KOREA. Chungcheongbuk-do: Jecheon-si, 7 May 2008, J. Jung 805004; 27 June 2008, J.Jung 806053 * (GQ130340/GQ130358/JF313190); Boryeong-si, 23 June 2005, H.-K.Choi s.n.; 15 June 2008, J. Jung 806030, 806035; Taean-gun, 11 Aug. 2008, J.Jung 808002; Gangwon-do: Goseong-gun, 15 July 2008, H.R.Na 80211; 27 Aug. 2008, J.Jung 808308; 4 Sep. 2009, J.Jung 909119; Yanggu-gun, 25 June 2009, J.Jung 906035; Yangyang-gun, 26 May 2001, H.-K.Choi s.n.; Gyeonggi-do: Anseong-si, 19 May 2007, H.-K.Choi s.n.; Hwaseong-si, 9 July 2004, H.-K.Choi s.n.; 12 June 2008, J.Jung 806027; Suwon-si, 25 May 2003, H.R.Na s.n.; Yeoju-gun, 25 July 2007, H.R.Na s.n.; Gyeongsangnam-do: Changnyeong-gun, 25 July 2008, H.R.Na 80300; 23 July 2009, H.R.Na 90087; Incheon: Ganghwa-gun, 11 June 2008, J.Jung 806002, 806015; Jeollabuk-do: Gimje-si, 28 June 2007, H.R.Na s.n.; Jeongeup-si, 27 June 2007, H.R.Na s.n.; Jeollanam-do: Boseong-gun, 29 May 2008, H.R.Na 80038.

Bolboschoenus planiculmis KOREA. Chungcheongnam-do: Asan-si, 3 June 2007, H.R.Na s.n.; Taean-gun, 26 June 2006, H.-K.Choi s.n.; Gangwon-do: Gangneung-si, 2 July 2007, C.Kim s.n.; Goseong-gun, 3 July 2007, C.Kim s.n.; 15 July 2008, H.R.Na 80191; Yangyang-gun, 1 July 2006, H.-K.Choi s.n.; Gyeonggi-do: Ansan-si, 12 June 2008, J.Jung 806022; Hwaseong-si, 12 June 2008, J.Jung 806024; Siheung-si, 10 July 2004, H.-K.Choi s.n.; Gyeongsangbuk-do: Gunwi-gun, 24 July 2008, H.R.Na 80264; Gyeongsangnam-do: Geoje-si, 29 June 2006, H.-K.Choi s.n.; Hadong-gun, 27 July 2008, H.R.Na 807139; 11 Aug. 2009, C.Kim & J.Jung 908022, 908035; Incheon: Ganghwa-gun, 11 June 2008, J.Jung 806005 * (GQ130341/GQ130359/JF313192), 806014; Ongjin-gun, 12 June 2008, J.Jung 806025; Jeju-do: Bukjeju-gun, 9 July 2008, J.Jung 807062, 807063; Jeollanam-do: Goheung-gun, 29 May 2008, H.R.Na 80022; Haenam-gun, 27 June 2006, H.-K.Choi s.n.; 28 June 2006, H.-K.Choi s.n.; Yeongam-gun, 21 Aug. 2008, J.Jung 808032.

Bolboschoenus maritimus KOREA. Gyeonggi-do: Paju-si, 26 June 2002, C.Kim s.n.; Incheon: Ganghwa-gun, 11 June 2008, J.Jung 806004, 806008 * (JF313179/JF313185/JF313191); Seoul: Hangang, 30 Oct. 1983, Y.N.Lee s.n. (SWU).

Schoenoplectus nipponicus KOREA. Gangwon-do: Goseong-gun, 16 Aug. 1996, S.H.Park s.n. (SWU); 27 Aug. 2008, J.Jung 808332 * (JF313183/GQ130364/JF313209); 5 Sep. 2009, J.Jung 909127; Hwacheon-gun, 25 June 2009, J.Jung 906024; Jeju-do: Bukjeju-gun, 20 Aug. 2007, Y.R.Lee s.n.

Schoenoplectus tabernaemontani KOREA. Chungcheongnam-do: Taean-gun, 5 June 2008, H.-K.Choi 806001; 11 Aug. 2008, J.Jung 808003, 808005; Gangwon-do: Goseong-gun, 15 July 2008, H.R.Na 80190, 80206; Yangyang-gun, 14 July 2008, H.R.Na 80180; Gyeonggi-do: Yeoju-gun, 26 Apr. 2003, C.M.Ahn s.n.; Gyeongsangbuk-do: Mungyeong-si, 23 July 2008, H.R.Na 80245; Gyeongsangnam-do: Haman-gun, 25 July 2008, H.R.Na 80304; 9 Oct. 2008, J.Jung 810011; Incheon: Ganghwa-gun, 11 June 2008, J.Jung 806013, 806016 * (GQ130349/GQ130365/JF313210); Ongjin-gun, 12 June 2008, J.Jung 806026; Jeju-do: Bukjeju-gun, 8 July 2008, J.Jung 807088; 9 July 2008, J.Jung 807073, 807079; Jeollabuk-do: Gimje-si, 23 Sep. 2009, J.Jung 909174; Gochang-gun, 23 Sep. 2009, J.Jung & C.Kim 909161, 909171; Gunsan-si, 21 July 2009, H.R.Na 90044; Ulsan: Ulju-gun, 29 Aug. 2008, J.Jung 808337.

Schoenoplectus triqueter KOREA. Gangwon-do: Goseong-gun, 17 Aug. 2001, C.Kim s.n.; 27 Aug. 2008, J.Jung 808316, 808328; 4 Sep. 2009, J.Jung et al. 909113; Hwacheon-gun, 26 Aug. 2008, J.Jung 808303; 6 Sep. 2007, Y.R.Lee s.n.; Gyeongsangbuk-do: Gunwi-gun, 24 July 2008, H.R.Na 80265; Mungyeong-si, 14 Oct. 2004, H.-K.Choi s.n.; Uiseong-gun, 24 July 2009, H.R.Na 90139, 90148; Yecheon-gun, 24 July 2008, H.R.Na 80258 * (GQ130351/GQ130367/JF313211); Yeongcheon-si, 16 Sep. 2004, H.-K.Choi s.n.; Gyeongsangnam-do: Changnyeong-gun, 29 Aug. 2008, J.Jung 808334; Hadong-gun, 11 Aug. 2009, J.Jung 908036; Miryang-si, 21 Oct. 2004, H.-K.Choi s.n.; Incheon: Ongjin-gun, 4 Sep. 2008, J.Jung 809001; Jeollabuk-do: Gochang-gun, 23 Sep. 2009, J.Jung, C.Kim 909165; Ulsan: Ulju-gun, 29 Aug. 2008, J.Jung 808338.

Schoenoplectiella mucronata KOREA. Gyeongsangnam-do: Hadong-gun, 11 Aug. 2009, J.Jung 908034; Incheon: Ganghwa-gun, 4 Sep. 2008, J.Jung 809002 * (GQ130348/GQ130363/JF313200); 11 July 2007, H.R.Na s.n.; Jeollanam-do: Gangjin-gun, 22 Aug. 2008, J.Jung 808035; Yeongam-gun, 21 Aug. 2008, J.Jung 808027, 808028, 808029; Yeosu-si, 12 Aug. 2009, J.Jung 908052.

Schoenoplectiella triangulata KOREA. Chungcheongnam-do: Boryeong-si, 15 June 2008, J.Jung 806034 * (GQ130350/GQ130366/JF313201); Taean-gun, 11 Aug. 2008, J.Jung 808001; Gangwon-do: Goseong-gun, 27 Aug. 2008, J.Jung 808327; 4 Sep. 2009, J.Jung et al. 909114; 5 Sep. 2009, J.Jung et al.909128; Hwacheon-gun, 26 Aug. 2008, J.Jung 808302; Gyeongsangbuk-do: Gunwi-gun, 24 July 2009, H.R.Na 90135; Seongju-gun, 23 July 2009, H.R.Na 90112; Gyeongsangnam-do: Hadong-gun, 11 Aug. 2009, J.Jung 908040; Jeju-do: Bukjeju-gun, 8 July 2008, J.Jung 807087; 9 July 2008, J.Jung 807074, 807080; Jeollanam-do: Gangjin-gun, 22 Aug. 2008, J.Jung 808036; Gokseong-gun, 28 July 2008, H.R.Na 807142.

Schoenoplectiella hotarui KOREA. Chungcheongnam-do: Boryeong-si, 4 Sep. 2008, J.Jung 809003 * (GQ130345/GQ130360/JF313196); Gangwon-do: Goseong-gun, 26 Aug. 2006, H.R.Na s.n.; 27 Aug. 2008, J.Jung 808329; 5 Sep. 2009, J.Jung 909130, 909131; Gyeonggi-do: Ansan-si, 5 Sep. 2008, J.Jung 809006; 6 Aug. 2009, J.Jung 908004, 908006; Gyeongsangbuk-do: Mungyeong-si, 22 Sep. 2009, J.Jung 909212; Seongju-gun; Gyeongsangnam-do: Hadong-gun, 11 Aug. 2009, J.Jung 908032; Hapcheon-gun, 23 July 2009, H.R.Na 90102; Jeju-do: Bukjeju-gun, 20 Aug. 2007, H.R.Na s.n.; Jeollanam-do: Gangjin-gun, 22 Aug. 2008, J.Jung 808038; 29 Aug. 2006, H.-K.Choi s.n.

Schoenoplectiella juncoides KOREA. Chungcheongbuk-do: Jincheon-gun, 25 Sep. 2009, J.Jung et al. 909153; Gangwon-do: Goseong-gun, 27 Aug. 2008, J.Jung 808333; Hwacheon-gun, 25 June 2009, J.Jung 906025; 26 Aug. 2008, J.Jung 808301; Gyeonggi-do: Ansan-si, 9 July 2004, H.-K.Choi s.n.; Suwon-si, 1 July 2008, J.Jung 807020 * (GQ130346/GQ130361/JF313197); 5 Sep. 2008, J.Jung 809008; Gyeongsangbuk-do: Mungyeong-si, 22 Sep. 2009, J.Jung 909213; Yecheon-gun, 24 July 2008, H.R.Na 80257; Jeju-do: Bukjeju-gun, 7 Aug. 2009, H.R.Na 908066; Jeollabuk-do: Gimje-si, 23 Sep. 2009, J.Jung 909163; Gochang-gun, 23 Sep. 2009, J.Jung 909172; Jangsu-gun, 29 July 2008, H.R.Na 807144; Jinan-gun, 29 July 2008, H.R.Na 807145; 30 Aug. 2005, H.-K.Choi s.n.; Jeollanam-do: Boseong-gun, 29 Aug. 2005, H.-K.Choi s.n.; Gangjin-gun, 22 Aug. 2008, J.Jung 808034; 22 Aug. 2008, J.Jung 808051; 29 Sep. 2006, H.-K.Choi s.n.; Yeongam-gun, 21 Aug. 2008, J.Jung 808030; 21 Aug. 2008, J.Jung 808031.

Schoenoplectiella wallichii KOREA. Chungcheongbuk-do: Okcheon-gun, 20 Aug. 1968, M.S.Yuk s.n. (SWU); Gangwon-do: Goseong-gun, 23 Aug. 1997, E.J.Ham & Y.H.Lee s.n. (SWU); Gyeonggi-do: Gwacheon-si, 17 Sep. 1977, B.Y.Chang s.n. (SWU); Jeollabuk-do: Namwon-si, 7 July 2005, H.-K.Choi s.n. * (JF313182/JF313188/JF313203).

Schoenoplectiella komarovii KOREA. Gangwon-do: Hwacheon-gun, 6 Sep. 2007, H.R.Na s.n. * (JF313180/JF313186/JF313198).

Schoenoplectiella lineolata KOREA. Chungcheongbuk-do: Jecheon-si, 7 July 2009, J.Jung 907445, 907446; Chungcheongnam-do: Buyeo-gun, 24 Sep. 2009, J.Jung 909190; Gangwon-do: Chuncheon-si, 26 Aug. 2008, J.Jung 808310; Hwacheon-gun, 26 Aug. 2008, J.Jung 808306, 808321; Jeollanam-do: Gangjin-gun, 22 Aug. 2008, J.Jung 808041 * (GQ130347/GQ130362/JF313199); 30 Oct. 2008, J.Jung 810015; Gwangyang-si, 11 Aug. 2009, C.Kim & J.Jung 908045.

Schoenoplectiella x trapezoidea KOREA. Gangwon-do: Goseong-gun, 27 Aug. 2008, J.Jung 808330, J.Jung 808331 * (JF313181/JF313187/JF313202); 5 Sep. 2009, J.Jung et al.909132, J.Jung 909133.

Scirpus wichurai KOREA. Chungcheongbuk-do: Jincheon-gun, 25 Sep. 2009, J.Jung et al.909143; Gangwon-do: Goseong-gun, 14 July 2008, H.R.Na 80182; 27 Aug. 2008, J.Jung 808322 * (GQ130357/GQ130373/JF313208); 27 Aug. 2008, J.Jung 808325; 4 Sep. 2009, J.Jung et al. 909115; Gyeonggi-do: Ansan-si, 1 July 2008, J.Jung 807018; 5 Sep. 2008, J.Jung 809005; 6 Aug. 2009, J.Jung 908002; Paju-si, 23 Oct. 2007, H.R.Na s.n.; Suwon-si, 11 Oct. 2009, J.Jung 910001; Gyeongsangbuk-do: Mungyeong-si, 22 Sep. 2009, J.Jung 909193; Gyeongsangnam-do: Hadong-gun, 12 Aug. 2009, J.Jung 908055.

Scirpus karuisawensis KOREA. Chungcheongbuk-do: Jincheon-gun, 25 Sep. 2009, J.Jung et al. 909142; Gangwon-do: Goseong-gun, 27 Aug. 2008, J.Jung 808324; 4 Sep. 2009, J.Jung et al. 909116; Yangyang-gun, 5 Sep. 2009, J.Jung et al. 909045; Gyeonggi-do: Ansan-si, 1 July 2008, J.Jung 807017 * (GQ130353/GQ130369/JF313204); 5 Sep. 2008, J.Jung 809004; 6 Aug. 2009, J.Jung 908001; Gyeongsangbuk-do: Mungyeong-si, 22 Sep. 2009, J.Jung 909194.

Scirpus mitsukurianus KOREA. Chungcheongbuk-do: Taean-gun, 11 Aug. 2008, J.Jung 808007, 808008; Daejeon: Yuseong-gu, 25 Sep. 2009, J.Jung et al.909148; Gangwon-do: Chuncheon-si, 11 Oct. 2006, H.R.Na s.n.; 22 Sep. 2006, C.Kim s.n.; 26 Aug. 2008, J.Jung 808309; Hwacheon-gun, 26 Aug. 2008, J.Jung 808304 * (GQ130354/GQ130370/JF313205); Gyeonggi-do: Yeoju-gun, 25 July 2007, H.R.Na s.n.; 26 Sep. 2003, C.M.Ahn s.n.; Gyeongsangnam-do: Haman-gun, 9 Oct. 1985, D.Chung s.n.; 9 Oct. 2008, J.Jung 810010; Jeollanam-do: Gokseong-gun, 28 July 2008, H.R.Na 807143; 5 Oct. 2005, H.-K.Choi s.n.; Jangheung-gun, 28 July 2008, H.R.Na 807141; 10 Aug. 2006, C.Kim s.n.; Ulsan: Ulju-gun, 29 Aug. 2008, J.Jung 808336.

Scirpus radicans KOREA. Chungcheongbuk-do: Chungju-si, 9 July 1987, Y.C.Oh et al. s.n. (SWU); 14 Aug. 2002, C.Kim s.n.; Yeongdong-gun, 27 May 2005, H.-K.Choi s.n.; Chungcheongnam-do: Boryeong-si, 25 May 2005, H.-K.Choi s.n.; 15 June 2008, J.Jung 806032 * (GQ130356/GQ130372/JF313206), 806033; Gangwon-do: Hoengseong-gun, 30 July 2009, J.Jung et al. 907455; Hwacheon-gun, 24 June 2009, J.Jung 906012; 25 June 2009, J.Jung 906013; Pyeongchang-gun, 27 June 2008, J.Jung 806039, 806040; 24 June 2009, J.Jung 906007; Yanggu-gun, 25 June 2009, J.Jung 906034; Gyeonggi-do: Yeoju-gun, 29 May 2003, C.M.Ahn s.n.; 4 July 2008, J.Jung 807027; Jeollabuk-do: Namwon-si, 26 May 2005, H.-K.Choi s.n.; Jeollanam-do: Gokseong-gun, 26 May 2005, H.-K.Choi s.n.

Scirpus sylvaticus var. maximowiczii KOREA. Gangwon-do: Pyeongchang-gun, 27 June 2008, J.Jung 806038 * (GQ130355/GQ130371/JF313207); 24 June 2009, J.Jung 906008; Yanggu-gun, 25 June 2009, J.Jung 906032. CHINA. Jilin: Wonji, 25 July 2007, H.-K.Choi 808016–808019.

Trichophorum dioicum KOREA. Gangwon-do: Donggang, 17 Apr. 2008, J.Jung 804015 * (FJ797641/FJ797640/JF313213), 80401680427; 27 June 2008, J.Jung 806045, 806046.

Trichophorum alpinum, CHINA. Jilin: Mt. Baekdusan, 9 July 1993, Y.N.Lee s.n. (KB), 6 July 2010, J.Jung 1007071 * (JF313184/JF313189/JF313212), 10070721007075, 1007089.

Eleocharis congesta KOREA. Chungcheongbuk-do: Jecheon-si, 28 Aug. 2007, H.R.Na s.n.; Jincheon-gun, 25 Sep. 2009, J.Jung et al. 909150; Buyeo-gun, 24 Sep. 2009, J.Jung 909186; Gangwon-do: Goseong-gun, 5 Sep. 2009, J.Jung et al. 909125; Yanggu-gun, 25 June 2009, J.Jung 906033; Gyeonggi-do: Ansan-si, 23 May 2008, H.R.Na 80090; 23 May 2008, J.Jung 805083 * (GQ130343/GQ130375/JF313194); 1 July 2008, J.Jung 807015; Gyeongsangbuk-do: Mungyeong-si, 22 Sep. 2009, J.Jung 909205; Namhae-gun, 11 Aug. 2009, C.Kim & J.Jung 908011; Jeju-do: Bukjeju-gun, 8 July 2008, J.Jung 807075; 20 Aug. 2007, H.R.Na s.n.; 22 Aug. 2007, H.R.Na s.n.

Fimbristylis squarrosa KOREA. Chungcheongnam-do: Taean-gun, 11 Aug. 2008 J.Jung 808010; Gangwon-do: Yanggu-gun, 24 Aug. 2010 J.Jung 1008001; Yangyang-gun, 17 Aug. 2001 C.Kim et al. s.n.; 14 Sep. 2001 C.Kim et al. s.n.; Gyeonggi-do: Icheon-si, 4 July 2008, J.Jung 807041 * (GQ130344/GQ130376/JF313195); Gyeongsangbuk-do: Ulju-gun, 5 Oct. 2010 J.Jung 1010071.

Cladium chinense KOREA. Jeju-do: Seogwipo-si, 14 July 2006, H.-K.Choi et al. s.n. * (GQ130342/GQ130374/JF313193); 3 Nov. 2010, J.Jung 1011040.

Appendix 2

Table 5 Taxon and coded character state matrix

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Jung, J., Choi, HK. Taxonomic Study of Korean Scirpus L. s.l. (Cyperaceae) II: Pattern of Phenotypic Evolution Inferred from Molecular Phylogeny. J. Plant Biol. 54, 409–424 (2011). https://doi.org/10.1007/s12374-011-9181-8

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