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Korean J. Pl. Taxon > Volume 52(1); 2022 > Article
PARK, KIM, CHUNG, KIM, SON, and JANG: Clematis pseudotubulosa (Ranunculaceae), a new species from Korea

Abstract

Clematis pseudotubulosa (Ranunculaceae), a new species from Gyeonggi-do in the Republic of Korea, is described and illustrated. The new species is morphologically similar to C. takedana but differs in its leaves, which are always ternate with shallowly lobed leaflets (vs. 1-pinnate with deeply lobed three to five leaflets), and its oblong-shaped flower buds (vs. ovoid), larger flowers (vs. smaller flowers), and pantoporate pollen (vs. tricolpate). A detailed species description, geographical distribution, and an identification key for all species of Clematis that occur in Korea are also provided.

INTRODUCTION

The genus Clematis L. is one of the largest genera in the buttercup family (Ranunculaceae). It consists of approximately 300 species of herbaceous or woody vines and rarely erect shrubs or perennial herbs, which are common in temperate regions, including northern Europe, Siberia, and the Far East, whereas other species occur in tropical regions (Tamura, 1955, 1967, 1987, 1995; Wang and Bartholomew, 2001; Wang and Li, 2005; Kadota, 2006; Chang 2007; Kim, 2017).
This genus consists of 18 sections based on both morphological characters such as the leaf shape, flower structure, and achenes, as well as recent molecular phylogenies (Tamura, 1967, 1987; Jensen et al., 1995; Johnson, 2001; Kim, 2003; Wang and Li, 2005; Miikeda et al., 2006; Wang et al., 2009; Xie et al., 2011; Xie and Li, 2012; Lehtonen et al., 2016; Jiang et al., 2017; Kim, 2017; Ghimire et al. 2020; Park, 2021; Park et al., 2021). Among them, members of Clematis L. sect. Tubulosae Decaisne include approximately nine species, which are mainly distributed in East Asia (Decaisne, 1881; Kitagawa, 1937; Shi, 2003; Wang and Xie, 2007); these plants have trifoliate leaves, stems without tendrils, linear calyces that are tubular in shape with recurved upper parts, 12–24 stamens, persistent styles, and ovate-shaped fruits (Decaisne, 1881; Kitagawa, 1937; Wang and Xie, 2007). On the Korean Peninsula, two species have been reported in this section (Park, 2021), of which one species, C. urticifolia Nakai ex Kitag., is endemic to the Korean Peninsula.
During a recent floristic survey, one species of Clematis was collected from areas near Lake Sihwa and Namhansanseng Fortress in Gyeonggi-do, Republic of Korea. Clematis individuals from Lake Sihwa were once reported as C. takedana Makino (Moon et al., 2013; Chung, 2018), a natural hybrid of C. apiifolia DC. (sect. Clematis) and C. stans Siebold & Zucc. (sect. Tubulosae). Its habit, phyllotaxis, and sepal shapes are similar to those of C. apiifolia, whereas the features of its reproductive organs are similar to those of C. stans, a species endemic to Japan (Makino, 1907; Kitagawa, 1937; Ohwi, 1965; Kadota, 2006; Wang and Xie, 2007; Xie and Li, 2012). However, the putative parent species C. stans is not distributed in South Korea (Kadota, 2006); hence, the taxonomic identity of the Clematis individuals from Lake Sihwa and Namhansanseng Fortress is unclear. After careful examinations of various flora and herbarium specimens from Korea and adjacent countries, we concluded that the collected specimens could be proposed as a new species with the name Clematis pseudotubulosa.

MATERIALS AND METHODS

Morphological examination

We collected 34 fresh samples of C. pseudotubulosa and prepared a voucher specimen. By referring to the relevant protologues, flora, and monographs (Wang and Bartholomew, 2001; Kadota, 2006; Kim, 2017), we determined four target congeneric taxa for examination. Photographs in the field were captured using a digital camera (Nikon D3X, D7200 and Coolpix P530, Tokyo, Japan). To compare the morphological characteristics of the new species, we took loan of 132 herbarium specimens included C. tubulosa, deposited in the Korea National Arboretum (KH), Andong National University (ANH), Kangwon National University (KWNU), Kongju University (KNH), and National Institute of Biological Resources (KB) (Appendix 1). Additionally, voucher specimens image of C. takedana and C. stans deposited in the National Science Museum (TNS) and Kyoto University (KYO) were studied (Appendix 1). Measurements of the morphological characters were taken with digital Vernier calipers (CD-15APX, Sakado, Mitutoyo) and from data derived from field notes. Morphological observations of all plant parts were conducted, focusing primarily on the shapes of the leaves and floral parts, as well as the features of the reproductive organs, under an Olympus dissecting stereo microscope (SZX16, Tokyo, Japan). Microscopic floral parts such as sepals and pistils were dissected when required. Various leaf-associated characters (leaflet number and size, petiole type) and flower-related features (size and shape of the sepals, bracts, and stamens, and shape and number of carpels) were assessed (Table 1).

Pollen morphological analysis

To observe the pollen, we used mature anthers after the anthesis of C. pseudotubulosa flowers. Pollen was prepared by a slightly modified standard acetolysis method (Erdtman, 1971; Radford et al., 1974). Cut anthers were put into a 15 mL tube and an acetolysis mixture (acetic anhydride:sulfuric acid = 9:1) was added, followed by incubation in a hot water bath at 90°C for about 10 min. After 10 min of centrifugation at 1,500 rpm, the supernatant was removed and washed twice with glacial acetic acid and once each with distilled water and 30%, 50%, and 70% ethanol. The supernatant was then stored in 70% ethanol. For scanning electron microscopic observations, some acetolyzed pollen grains were sprinkled onto carbon tape attached to a slide glass and coated with gold (thickness: 200–250 Å) by ion sputtering. A scanning electron microscope (S-2500C; Hitachi, Japan; 15 kV; working distance, 35 mm) was used to observe and photograph the shape and surface of the pollen. Meanwhile, data on the pollen of C. takedana and C. stans were provided based on data reported by Wang and Xie (2007) and Xie and Li (2012).

TAXONOMIC TREATMENT

Clematis pseudotubulosa B. K. Park, sp. nov. (Figs. 13).— TYPE: KOREA. Gyeonggi-do: Hwaseong-si, Namyang-eup, Munho-ri, 27 Aug 2019. J.-S. Kim kjs19046 (holotype: KB (NIBRVP0000824262); isotype: KH (NIBRVP0000824263)).
Woody vine, deciduous. Rhizomes stout, brown or dark brown; roots fibrous, brown to reddish brown. Woody stems, grayish brown to dark brown, peeling; herbaceous stems node 4.3–8.2mm diam., yellowish green to purple, densely pubescent; internodes terete, 16.8–23.4 cm × 1.6–5.7 mm, yellowish green to purple, densely pubescent, grooved. Leaves opposite, ternate; petiole 6.7–12.0 cm × 1.5–2.7 mm, yellowish green, green, dark green, or purple, densely pubescent; terminal petiolules 3.5–8.9 cm × 1.1–1.9 mm; lateral petiolules 0.8–3.0 cm × 1.3–2.1 mm; leaflets 3, ovate or widely ovate, 2–3-lobed, apex acute or acuminate, base subcordate, rounded, or truncate, margin dentate, biserrate, or rarely entire, adaxial surface green or dark green, abaxial surface pale green or yellowish green, pubescent on both surfaces, papery; terminal leaflets 7.3–12.3 × 6.9–11.1 cm, lateral leaflet 6.5–11.8 × 5.0–8.4 cm. Inflorescences raceme-cyme, terminal and axillary, 13–37-flowered; bract 1-paired, foliaceous, not lobed or 2–3-lobed, lanceolate, ovate, or widely ovate, 2.8–19.6 × 1.0–7.1 mm, yellowish green, green, or dark green, densely pubescent; bracteoles lanceolate to widely ovate, 1.6–4.2 mm long, green, dark green, or purple, densely pubescent; peduncle 2.8–14.9 cm × 1.2–2.8 mm, pedicel 1.1–4.4 cm × 0.5–0.8 mm, peduncle and pedicel yellowish green, green, or reddish green, densely tomentose. Flower bisexual, incomplete, 2.1–4.0 cm width; sepals 4 or rarely 5, valvate, linear-oblong, oblong, or spatulate, 1.5–2.3 cm × 3.7–6.1 mm, apex mucronate, acute, rounded, or truncate, reflexed at apex, white to pale purple, adaxial surface glabrous, abaxial surface densely pubescent; stamens 32–41, filament 4.9–9.1 × 0.4–0.6 mm, white, pubescent at apex; anther linear to ellipsoid, 3.6–4.6 × 0.5–0.8 mm, mucronate at apex, yellow, pubescent at apex; pistils 20–32; ovary ovoid or ellipsoid, 0.4–0.7 × 0.3–0.7 mm, yellowish green or green, densely pubescent, style 4.9--5.8 mm long, green, plumose. Achenes ovoid, 2.4–4.4 × 1.6–2.4 mm, green or dark green, dark brown at maturity, persistent style 2.3–3 cm long, white plumose. Seed ovoid or ellipsoid, 1.9–2.4 × 1.1–1.4 mm, yellowish brown. Pollen pantoporate, spheroidal to subprolate, 13.7–22.2 × 11.1–20.7 μm, colpus orbicular, exine sculpture spinulose, conical, 0.15–0.31 μm long.
Etymology: The specific epithet “pseudotubulosa” is derived from a combination of “pseudo” and “tubular,” because the flowers are tubular-shaped in the beginning of flowering, but they spread upon full flowering.
Vernacular name: Deong-gul-jo-hui-pul (덩굴조희풀)
English colloquial name: Creeping hyacinth-flower clematis
Phenology: Flowering August to September; fruiting September to October
Distribution: Clematis pseudotubulosa is restricted to the Korean Peninsula (South Korea, including Hwaseong-si and Gwangju-si, Gyeonggi-do) (Fig. 4).
Taxonomic note: Clematis pseudotubulosa is morphologically similar to C. takedana with regard to its habit, leaf size and shape, and flower structures (Kadota, 2006; Wang and Xie, 2007). Despite these similarities, there are clear differences between these two species, such as a tendril-like petiole, hairs on the surface of the petiole, the number of leaflets and the shape of the bracts (Figs. 2, 3, Table 1), as well as shape of the flower buds (oblong for C. pseudotubulosa vs. ovoid for C. takedana) and pollen type (pantoporate for C. pseudotubulosa vs. tricolpate for C. takedana) (Fig. 5, Table 1) (Wang and Xie, 2007). Clematis tubulosa, which is found in Korea and China, and C. stans, which is found in Japan as an endemic species, also display tendril-like petioles, ternate leaves, hairy leaflets, and pantoporate pollen (Wang and Xie, 2007; Xie and Li, 2012). However, the new species differs from them by its habit, texture of its leaves, sexuality of the flower, and from the latter by its absence of a staminode (Figs. 3, 4, Table 1). Therefore, the Korean individual that Moon et al. (2013) reported as C. takedana was misidentified. Currently, only a few C. pseudotubulosa individuals grow in the two identified habitats. The habitats are threatened by destruction due to nearby development and ongoing work to repair castle walls. Hence, there is a pressing need to protect this species.
Meanwhile, considering the habitat of C. pseudotubulosa, this species may have arisen from natural hybridization between C. apiifolia and C. tubulosa. In addition, morphologically, the petiole type, leaflet and sepal shapes, stamen trichomes, pollen type are similar to those of C. tubulosa, whereas the features of its habit, phyllotaxis, and achene shapes are similar to those of C. apiifolia.
Additional specimens examined: KOREA. Gyeonggi-do: Gwangju-si, Namhansanseong-myeon, Sanseong-ri, Namhan-sanseong, 9 Aug 2019, B. K. Park Namhansanseong-190809-012 (KH); Hwaseong-si, Namyang-eup, Munho-ri, 9 Aug 2014, J. D. Lee & K. D. Jung Lee JD et al. 14352 (2 sheets, KB); same locality, 16 Sep 2014, B. K. Park Sihwaho-140916-010--020, (KH); same locality, 1 Aug 2016 H. J. Jo Sihwaho-160801-001--005 (KH); same locality, 19 Jul 2019, B. K. Park Sihwaho-190719-001--004 (KH); same locality, 9 Aug 2019, B. K. Park Sihwaho-190809-001--017 (KH); same locality, 27 Aug 2019, J.-S. Kim kjs19047 (4 sheets, KB); same locality, 21 Sep 2019, B. K Park. Sihwaho-190921-001--015 (KH); same locality, 18 Aug 2020, B. K. Park Sihwaho-200818-001--003, (KH); same locality, 7 Oct 2020, J.-S. Kim and J.-H. Kim kjs20037 (3 sheets, KB); same locality, 18 Aug 2021, J.-S. Kim FS-PS-20210067 (KB); same locality, 26 Aug 2021, B. K. Park & K.-H. Lee Sihwaho-210826-001 (KH); Lakeshore of Shihwaho, 7 Sep 2008, Jang et al. 20080907-001–003 (KNH); same locality, 26 Jul 2010, Jang et al. 20100726-001, 002 (KNH); same locality, 7 Aug 2011, Jang et al. 20110807-001 (KNH); same locality, 20 Aug 2011, Jang et al. 20110820-001 (KNH); same locality, 31 Aug 2011, Jang et al. 20110831-001--004 (KNH); same locality, 18 Sep 2011, Jang et al. 20110918-001 (KNH).

An identification key to Korean Clematis species

  • 1. Leaf margin serrate.

    • 2. Flowers downwards at flowering, sepals campanulate.

      • 3. Internode villous; inflorescences cyme; bract present; sepals hairy on adaxial surface; staminodes absent ················· ························································································································································· C. serratifolia 개버무리

      • 3. Internode pubescent; inflorescences solitary; bract absent; sepals glabrous on adaxial surface; staminodes present.

        • 4. Peduncles pubescent; sepal caruncles present ······································································· C. koreana 세잎종덩굴

        • 4. Peduncles villous; sepal caruncles absent.

          • 5. Leaves 2-ternate, texture papery ································································ C. alpina var. ochotensis 자주종덩굴

          • 5. Leaves ternate, texture coriaceous ··················································································· C. calcicola 바위종덩굴

    • 2. Flowers upwards at flowering, sepals spreading, tubular or urceolate.

      • 6. Petioles tendril-like; peduncles pubescent; flowers white or yellowish green; stamens glabrous; pollen grain 3-colpate.

        • 7. Nodes villous; leaves pinnate or rarely ternate, leaflets 5; inflorescence cyme; peduncles villous; flowers 3; achenes glabrous ···································································································································· C. trichotoma 할미밀망

        • 7. Nodes pubescent; leaves ternate or 2-ternate, leaflets 3--9; inflorescence panicle-cyme; peduncles pubescent; flowers many; achenes pubescent.

          • 8. Leaves ternate, leaflets ovate or elliptic ··············································································· C. apiifolia 사위질빵

          • 8. Leaves 2-ternate; leaflets lanceolate ·········································································· C. brevicaudata 좀사위질빵

      • 6. Petioles not tendril-like; peduncles tomentose; flowers blue or purple; stamens pubescent; pollen grain pantoporate.

        • 9. Plants scandent; flowers bisexual ·············································································· C. pseudotubulosa 덩굴조희풀

        • 9. Stems erect; flowers andromonoecious or androdiecious.

          • 10. Plants andromonoecious; sepals ovate, glabrous on adaxial surface ···························· C. urticifolia 병조희풀

          • 10. Plants androdioecious; sepals spatulate, pubescent on adaxial surface ······················ C. tubulosa 자주조희풀

  • 1. Leaf margin entire.

    • 11. Flowers downwards at flowering; sepals purple or brown, fleshy; stamens pubescent.

      • 12. Sepal white hairy ············································································································ C. fusca var. violacea 종덩굴

      • 12. Sepal brown to black hairy.

        • 13. Plant climbing, stems remnant of the shoot in winter season ······························· C. fusca var. fusca 검종덩굴

        • 13. Plant erect, stems not remnant of the shoot in winter season ························ C. fusca var. flabellate 요강나물

    • 11. Flowers upwards at flowering; sepals white or yellowish green, papery; stamens glabrous.

      • 14. Internodes and petioles villous; inflorescences solitary; bract absent; sepals more than 3 cm long; achene margin thin ··· ························································································································································· C. patens 큰꽃으아리

      • 14. Internodes and petioles pubescent; inflorescences cyme; bract present; sepals less than 3 cm long; achene margin thick.

        • 15. Stems erect; leaf texture coriaceous; sepals pubescent on adaxial surface ······ C. hexapetala 좁은잎사위질빵

        • 15. Stems scandent; leaf texture papery; sepals glabrous on adaxial surface.

          • 16. Petioles not tendril-like; peduncles pubescent; sepals glabrous on adaxial surface and margin; achenes winged; persistent styles less than 1.9 cm long ···················································· C. brachyura 외대으아리

          • 16. Petioles tendril-like; peduncles pubescent; sepals pubescent on adaxial surface and margin; achenes not winged; persistent styles more than 1.9 cm long.

            • 17. Persistent styles trichomes yellow to brown ·················································· C. taeguensis 대구으아리

            • 17. Persistent styles trichomes white.

              • 18. Stems not remnant of the shoot in winter season; leaflets projections on veins of abaxial surface ······ ·························································································································· C. mandshurica 으아리

              • 18. Stems remnant of the shoot in winter season; leaflets smooth on veins of abaxial surface ··········· ···························································································································· C. terniflora 참으아리

Acknowledgments

This study was supported by a grant from the Korea National Arboretum (KNA1-1-18, 15-3) and National Institute of Biological Resources (NIBR202203102). We would like to thank Hee Soo Kim for preparing the line drawings.

NOTES

CONFLICTS OF INTEREST
The authors declare that there are no conflicts of interest.

Fig. 1.
Holotype specimen of Clematis pseudotubulosa B. K. Park (27Aug 2019. J.-S. Kim kjs19046 KB).
kjpt-52-1-35f1.jpg
Fig. 2.
Illustration of Clematis pseudotubulosa B. K. Park. Illustrations by Hee Soo Kim. A. Plant. B. Adaxial surface of the leaflet. C. Abaxial surface of the leaflet. D. Polar view of the flower. E. Equatorial view of the flower. F. Sepal. G. Stamen. H. Pistil. I. Fruit.
kjpt-52-1-35f2.jpg
Fig. 3.
Photographs of Clematis pseudotubulosa B. K. Park. A. Plant. B. Lignified stem. C. Herbaceous stem. D–F. Leaf (E, adaxial surface; F, abaxial surface). G. Inflorescence. H. Peduncle. I. Pedicel. J. Flower. K–L. Sepal (L, adaxial surface; M, abaxial surface). N. Stamen. O–P. Pistil. Q. Aggregated achenes. R. Achene. S. Seed.
kjpt-52-1-35f3.jpg
Fig. 4.
Distribution of Clematis pseudotubulosa B. K. Park in South Korea.
kjpt-52-1-35f4.jpg
Fig. 5.
Pollen grains of Clematis pseudotubulosa B. K. Park and related taxa. A, B. C. pseudotubulosa. C, D. C. takedana (Wang and Xie, 2007). E, F. C. tubulosa. G, H. C. stans (Wang and Xie, 2007). Equatorial view: A, C, E, G; Exine sculpture of a pollen grain: B, D, F, H. Scale bars = 3 μm (A, C, E, G), 1 μm (B, D, F, H).
kjpt-52-1-35f5.jpg
Table 1.
Comparison of characteristics of Clematis pseudotubulosa and its related species.
Character C. pseudotubulosa C. takedana C. tubulosa C. stans
Habit Woody vine Woody vine or woody perennial herb Subshrub Perennial herb or subshrub
Growing type Scandent Scandent Erect or ascending Erect or ascending
Leaf Ternate Ternate or rarely 5 pinnate Ternate Ternate
  Petiole Not tendril-like Tendril-like Not tendril-like Not tendril-like
    Surface Tomentose Glabrous Tomentose Strigose
  Leaflet size (cm) 6.5–12.3 × 5.0–11.1 5–9.8 × 3.0–8.5 7.8–19.0 × 4.9–9.7 4.1–17.3 × 3.6–10
    Adaxial surface Pubescent Sparsely appressed puberulous Pubescent Subglabrous
    Abaxial surface Pubescent Puberulous on veins Pubescent Tomentose on veins
  Texture Papery Papery Papery Coriaceous
Inflorescence bract Oblanceolate, oblong, obovate or elliptic Lanceolate, triangular or elliptic Oblanceolate, narrowly oblong, obovate or elliptic Foliiform or linear
Flower Bisexual Bisexual Androdiecious Polygamous
  Flower bud shape Oblong or ellipsoid Ovoid Narrowly ovoid or narrowly oblong Narrowly ovoid or narrowly oblong
  Sepal size 1.5–2.3 cm × 3.7–6.1 mm 0.9–1.6 cm × 2.0–3.6 mm 1.3–2.3 cm × 2.8–5.3 mm 0.8–1.8 cm × 1.6–2.8 mm
    Shape Lanceolate, oblanceolate or oblong, spatulate Oblanceolate or narrowly obovate-oblong Oblanceolate or spatulate Narrowly oblong or narrowly obovate-oblong
    Adaxial surface White, pinkish white, bluish white or purple Pink or purple White, whitish blue to violet Purple
    Recurve Not recurved or recurved at the apex Not recurved or recurved at the apex Recurved at the middle Recurved to circinate at the middle
  Staminode Absent Absent Absent Present
  Filament length (mm) 4.9–9.1 5.3–10.3 2.7–4.0 2.2–4.8
    Shape Linear or lanceolate Linear, lanceolate or oblanceolate Linear to narrowly oblong Linear
    Surface Pubescent at the upper part Villous or pilose at the upper part or glabrous Pubescent at the upper part Pilose or glabrous
  Anther length (mm) 3.6–4.6 1.8–3.3 1.3–6.1 2.5–5.0
    Shape Linear, lanceolate or oblong Linear Linear to lanceolate Linear or linear ovate
  Style length (mm) 4.9–5.8 3.1–6.3 3.2–3.8 4–5
Fruit size (mm) 2.5–3.3 × 1.6–2.1 2.3–3.3 × 1.3–2.0 2.9–3.6 × 1.8–2.3 2.0–5.5 × 1.2–2.2
  Shape Ovoid to ellipsoid Ovoid Ovoid or widely ellipsoid Obovoid or ellipsoid
  Surface Pubescent Pubescent Villous Puberulous
  Persistent style Plumose Plumose Plumose Plumose
    Length (cm) 2.3–3.0 1.8–2 1.5–2.0 1.5–2.2
  Color White Mature unknown White White
  Pollen type Pantoporate Tricolpate Pantoporate Pantoporate
    Size (µm) 13.7–22.2 × 11.1–20.7 20.1–24.5 × 15.4–19.1 14.1–19.9 × 14.5–20.1 20.5–21.8 × 20.5–21.8
    P/E ratio 1.07 1.36 0.98 1.00
    Shape Spheroidal, subprolate Subprolate Spheroidal, suboblate Spheroidal
    Colpus Orbicular Linear Orbicular Orbicular

LITERATURE CITED

Chang, CS. 2007. Clematis L. The Genera of Vascular Plants of Korea. Flora of Korea Editorial Committee (ed.), Academy Publishing Co, Seoul. Pp. 191-195.

Chung, GY. 2018. Clematis takedana Makino In Silvics of Korea. 2: Oh, B-U. Oh, SH (eds.), Korea National Arboretum, Pocheon. Pp. 21-29.

Decaisne, MJ. 1881. Revision des clématites du groupe tubule-uses cultivées au Muséum. Nouvelles archives du Muséum d’Histoire Naturelle (Paris), Serie 2 4: 195-214.

Erdtman, G. 1971. Pollen Morphology and Plant Taxonomy (Angiosperm). Hafner Publishing Co., New York. Pp. 539.

Ghimire, B. Park, BK. Son, DC and Oh, S-H. 2020. Achene morphology and anatomy of Clematis L. (Ranunculaceae) in Korea and its taxonomic implications. Plants 9: 1279.
crossref pmc
Jensen, U. Hoot, SB. Johansson, JT and Kosuge, K. 1995. Sys tematics and phylogeny of the Ranunculaceae: a revised family concept on the basis of molecular data. Plant Systematics and Evolution 9(Suppl): 273-280.

Jiang, N. Zhou, Z. Yang, J-B. Zhang, S-D. Guan, K-Y. Tan, Y-H and Yu, W-B. 2017. Phylogenetic reassessment of tribe Anemoneae (Ranunculaceae): non-monophyly of Anemone s. l. revealed by plastid datasets. PLoS One 12: e0174792.
crossref pmid pmc
Johnson, M. 2001. The Genus Clematis . Plantskola, Stockholm. Pp. 896.

Kadota, Y. 2006. Clematis L. Flora of Japan. Vol. II a. Angio-spermae, Dicotyledoneae, Archichlamydeae. Iwatsuki, K. Boufford, DE. Ohba, H (eds.), Kodansha, Tokyo. Pp. 298-308.

Kim, JS. 2003. Phylogenetic relationship of genus Clematis L. in Korea. M.S. thesis. Kyungpook National University, Daegu, Korea. Pp. 45 (in Korean).

Kim, JS. 2017. Clematis L. In Flora of Korea. 2a: Magnoliidae. Flora of Korea Editorial Committee and National Institute of Biological Resources (ed.), National Institute of Biological Resources, Incheon. Pp. 69-76.

Kitagawa, M. 1937. On the species of Clematis sect. Tubulosae . The Journal of Japanese Botany 13: 343-360.

Lehtonen, S. Christenhusz, MJM and Falck, D. 2016. Sensitive phylogenetics of Clematis and its position in Ranunculaceae. Botanical Journal of the Linnean Society 182: 825-867.

Makino, T. 1907. Observations on the flora of Japan. Botanical Magazine (Tokyo) 21: 86-88.
crossref
Miikeda, O. Kita, K. Handa, T and Yukawa, T. 2006. Phylogenetic relationships of Clematis (Ranunculaceae) based on chloroplast and nuclear DNA sequences. Botanical Journal of the Linnean Society 152: 153-168.
crossref
Moon, A.-R.. Han, J-E. Lee, B-Y. Park, J-M and Jang, C-G. 2013. An unrecorded species of genus Clematis (Ranunculaceae) from Korea. Journal of Asia-Pacific Biodiversity 6: 415-418.
crossref
Ohwi, J. 1965. Clematis. In Flora of Japan. Meyer, FG. Walker, EH (eds.), Smithsonian Institution, Washington, DC. Pp. 440-442.

Park, BK. 2021. A systematic study on the genus Clematis L. (Ranunculaceae) in Korea. PhD dissertation,. Hannam University, Daejeon, Korea. Pp. 241 (in Korean).

Park, BK. Son, DC and Ghimire, B. 2021. Taxonomic relevance of petiole anatomical and micro-morphological characteristics of Clematis L. (Ranunculaceae) taxa from South Korea. PeerJ 9: e11669.
crossref pmid pmc
Radford, AE. Dickinson, WC. Massey, JR and Bell, CR. 1974. Vascular Plant Systematics. Harper and Row, New York. Pp. 891.

Shi, JH. 2003. A preliminary study of the hybrid origin of Clematis pinnata Maxim (Ranunculaceae) MS thesis, Beijing Institute of Botany, Chinese. Academy of Sciences, Beijing, China.

Tamura, M. 1955. Systema clematidis Asiae Orientalis. Science Reports. 4: College of General Education, Osaka University, Osaka. Pp. 43-55.

Tamura, M. 1967. Morphology, Ecology and Phylogeny of the Ranunculaceae VII. Science Reports. 16: College of General Education, Osaka University, Osaka. Pp. 21-43.

Tamura, M. 1987. A classification of genus Clematis . Acta Phytotaxonomica et Geobotanica 38: 33-44.

Tamura, M. 1995. Clematis. In Die Natürlichen Pflanzenfamilien. Zweite Aufl. 17a. IV: Hiepko, P (ed.), Dunker & Humbolt, Berlin. Pp. 368-387.

Wang, WT and Bartholomew, B. 2001. Clematis L. In Flora of China. 6: Caryophyllaceae through Lardizabalaceae. Wu, ZY. Raven, PH. Hong, DY (eds.), Science Press, Beijing and Missouri Botanical Garden Press, St Louis, MO. Pp. 333-386.

Wang, W-T and Li, L-Q. 2005. A new system of classification of the genus Clematis (Ranunculaceae). Acta Phytotaxonomica Sinica 43: 431-488.

Wang, W. Lu, A-M. Ren, Y. Endress, ME and Chen, Z-D. 2009. Phylogeny and classification of Ranunculales: evidence from four molecular loci and morphological data. Perspectives in Plant Ecology Evolution and Systematics 11: 81-110.
crossref
Wang, W-T and Xie, L. 2007. A revision of Clematis sect. Tubulosae (Ranunculaceae). Acta Phytotaxonomica Sinica 45: 425-457.
crossref
Xie, L and Li, L-Q. 2012. Variation of pollen morphology, and its implications in the phylogeny of Clematis (Ranunculaceae). Plant Systematics and Evolution 298: 1437-1453.
crossref
Xie, L. Wen, J and Li, LQ. 2011. Phylogenetic analyses of Clematis (Ranunculaceae) based on sequences of nuclear ribosomal ITS and three plastid regions. Systematic Botany 36: 907-921.
crossref

APPENDICES

Appendix 1.

A list of specimens of C. takedana, C. tubulosa, and C. stans examined in this study

Clematis takedana Makino (Photo): JAPAN. Nagano Pref. : Jul 1927, T. Makino s.n. (OSA); Komagatake, 22 Aug 1937, A. Kobayashi 26 (KYO); Mt. Hakubayama, Aug 1920, G. Koidozumi s.n. (KYO); Yahatadaira Street, 20 Aug 1828, ? (KYO); Mt. Shiroumayama, Iriguchi, Aug 1920, G. Koidozumi s.n. (KYO); 2 Aug 1917, Kitayazumo-gun, Houjyou-cho, G. Koidozumi s.n. (KYO); Kitaazuoi-gun, Mt. Shiroumayama, Iriguchi, Aug 1920, T. Makino 95711 (KYO); Ise Tajita Valley, 4 Sep 1959, K. Iba s.n. (KYO). Aomori Pref.: Aomori-shi, 6 Sep 2007, Y. Kadota 74321 (TNS); same locality, 29 Sep 2010, K. Yonekura 17550 (TNS). Iwate Pref.: Iwaizumi-cho, 2 Sep 1970, J. Haginiwa JH000326--332, JH000334--338, JH002013, JH041996 (TNS); Shizukuishi-cho 25 Aug 1960, K. Hosoi s.n. (TNS). Nagano Pref.: Hakuba-mura, 21 Aug 1957, J. Haginiwa JH000325, JH000333, JH002017 (TNS); same locality, 22 Aug 1963, J. Haginiwa JH000324, JH000339, JH000340, JH041995 (TNS); Hokujo-mura 27 Aug 1905, T. Hisayoshi s.n. (TNS); same locality, 4 Aug 1913, Y. Yazawa s.n. (TNS); same locality, 12 May 1916, H. Koidzumi Herb. H. Koidzumi 67170 (TNS); same locality, 15 Aug 1922, H. Koidzumi 47267 (TNS); same locality, 23 Aug 1949, Y. Hayashi s.n. (TNS); same locality, 28 Aug 1949, Y. Hayashi s.n. (TNS); Miwa-mura, Aug 1937, K. Hisauchi s.n. (TNS). Yamagata Pref.: Higashine-machi, 16 Aug 1952, S. Okuyama 21866 (TNS); Oguni-machi, 24 Aug 1960, D. Shimizu s.n. (TNS); Togo-mura, 16 Aug 1952, S. Okuyama 21866 (TNS); Yonezawa-shi, 20 Aug 1963, D. Shimizu s.n. (TNS).
Clematis tubulosa Turcz.: KOREA. Gyeonggi-do: Dongducheon-si, Sangbongam-dong, Mt. Soyosan, 28 Jul 2007, G. Y. Chung ANH-01080581 (KH); Hwaseong-si, Munho-ri, Sihwa lake, 21 Sep 2021. B. K Park. Sihwaho-140916-001009 (KH); same locality, 9 Aug 2019, B. K Park. Sihwaho-190809-018024 (KH); same locality, 21 Sep 2019, B. K Park. Sihwaho-190921-016026 (KH); same locality, 18 Aug 2020, B. K Park. Sihwaho-200818-004--007 (KH). Incheon-si: Ganghwa Is., Sep 1967, I. M. Kim s.n. (KH); Seonjae Is., 24 Sep 2008, S.-H. Park ParkSH81981 (KH); Yeongheung Is., Sipripo beach, 14 Sep 2010, Y. J. Chung NAPI-0899 (KH). Gyeonggi-do: Gwangju-si: Sanseong-ri, Namhansanseong, 19 Jul 2019, B. K. Park Namhansanseong-190719-001004 (KH); 9 Aug 2019, B. K Park Namhansanseong-190809-001008 (KH); Namyangju-si, Mt. Cheonmasan, 6 Aug 2012, W. K. Baek DJU-2013-036 (KH); Gagok-ri, Mt. Cheonmasan, 2 Aug 2007, J.-H. Kim & C.-Y. Yoon Cheonmasan070802-101 (KH); Pocheon-si, Eoryong-dong, Mt. Wangbangsan, 3 Aug 2007, W. K. Paik Wangbangsan70803-656 (KH); Jikdong-ri, Soribong, 20 Aug 2004, J. M. Jeong 04080122 (KH); Mamyeong-ri, Mamyeongro, 4 Aug 2009, S. H. Park et al. L90054 (KH); Uiwang-si, Wanggok-dong, Mt. Baekunsan, 26 Jul 2007, G.Y. Chung ANH-01070064 (KH). Chungcheongnam-do: Seosan-si, Sangga-ri, Mt. Gayasan, 9 Sep 2007, J. I. Jeon & H.-S. Lee Jeon11693 (KH).
Clematis stans Turcz. (Photo): JAPAN. Akita Pref.: Kyowa-machi, 21 Aug 1990, I. Kenji s.n. (TNS). Aomori Pref.: Aomori-shi, 10 Aug 1960, H. Ohashi s.n. (TNS); Iwasaki-mura 28 Jul 1961, S. Okuyama 12108 (TNS). Fukushima Pref.: Ichinoki-mura, Aug 1951, J. Haginiwa JH041978 (TNS); Ryozen-machi, 23 Sep 1941, J. Yukio 11049 (TNS); Tamura-shi, 27 Aug 2011, Y. Yuzawa 24813, 24814 (TNS); Yakmakami-mura, 24 Jul 1938, M. Hayashi 5743 (TNS). Gifu Pref.: Gero-cho, 24 Sep 1933, Y. Araki 1992 (TNS); Kasuga-mura, 27 Oct 1984, H. Takahashi 1030 (TNS); Kiyomi-mura, 7 Sep 1998, K. Okuda and M. Makiko s.n. (TNS); Miyama-cho, 4 Jul 1982, S. Mitsuta 12134 (TNS); same locality, 14 Sep 1982, H. Takahashi 6683 (TNS); Shirakawa-mura, 28 Sep 2003, Y. Kadota 35119 (TNS); same locality, 1 Oct 2006, H. Takahashi 22379 (TNS); Takane-mura, 13 Aug 2003, N. Naruhashi 3081301 (TNS). Gunma Pref.: Agatsuma-machi, 22 Sep 1968, K. Masuda 2460 (TNS); Haruna-machi, 12 Oct 1957, S. Okuyama 24114 (TNS); Myogi-machi, 20 Aug 1967, J. Yukio 32704 (TNS); Numata-shi, 31 Aug 1956, S. Okuyama 19244 (TNS); same locality, 26 Jul 1974, M. Haruo 285 (TNS); Takasaki-shi, 7 Jun 1964, K. Matsunaga 144 (TNS). Hiroshima Pref.: Hiroshima-shi, 21 Aug 1977, K. Oka 42767 (TNS). Hyogo Pref.: Onsen-cho, 11 Aug. 1993, S. Fujii 3782 (TNS); Yoka-cho, 11 Sep 1993, N. Fukuoka et al. 6369 (TNS). Ishikawa Pref.: Oguchi-mura, 30 Jul 1992, K. Deguchi 16719 (TNS); same locality, 2003 Sep 28, Y. Kadota s.n. (TNS); Yamanaka-machi, 6 Oct 1986, S. Tsugaru 7190 (TNS). Iwate Pref.: Matsuo-mura, 31 Aug 1962, H. Koyama 1493 (TNS); Shizukuishi-cho, 3 Sep 1989, J. Haginiwa JH000309, JH000310, JH039451 (TNS). Kanagawa Pref.: Hadano-shi, 13 Oct 1975, Y. Kadota 971 (TNS); Hakone-machi, 14 Sep 1998, N. Tanaka 63 (TNS). Miyagi Pref.: Kawasaki-mura, 10 Aug 1937, S. Okuyama 9316 (TNS). Nagano Pref.: Akaho-mura, 1 Aug 1922, H. Koidzumi 67025 (TNS); Ariake-mura, 22 Jul 1921, H. Koidzumi 67024 (TNS); Hakuba-mura, 28 Aug 2005, O. Miikeda Miikeda_05-808--811 (TNS); Hase-mura, 24 Jul 1960, Masuda Shinya 3 (TNS); same locality, 12 Aug 1962, S. Masuda 1837 (TNS); same locality, 12 Aug 1971, M. Wakabayashi 29 (TNS); Hokujo-mura, 21 Aug 1936, K. Hisauchi 1764 (TNS); Iriyamabe-mura, 23 Jul 1926, J. Yukio 48 (TNS); Kami-mura, 24 Sep 1932, M. Muramatsu 984 (TNS); Karuizawa-machi, 5 Sep 1983, J. Haginiwa JH035037, JH035038 (TNS); Kiso-machi, 23 Aug 2014 M. Muramatsu 27818 (TNS); Kitaaiki-mura, 17 Aug 1986, J. Murata 21601 (TNS); Minamiaiki-mura, 16 Aug 1982, H. Takahashi 6475 (TNS); Mitake-mura, 3 Aug 1927, J. Yukio 91 (TNS); Oshika-mura, 29 Aug 1948, M. Muramatsu 1219 (TNS); same locality, 29 Aug 1963, K. Iwatsuki & H. Koyama 9 (TNS). Niigata Pref.: Tochio-shi, 19 Aug 1962, F. Konta 846 (TNS). Shiga Pref.: Azai-cho, 10 Oct 1969, G. Murata 20832 (TNS); Taga-cho, 20 Oct 1962, H. Koyama 1611 (TNS). Shizuoka Pref.: Gotenba-shi, 28 Oct 2000, O. Miikeda Miikeda_00-1002 (TNS); same locality, 11 Nov 2007, O. Miikeda 71103 (TNS); Kamiide-mura, 4 Sep 1924, Y. Yabe s.n. (TNS); Misakubo-cho, 15 Sep 1980, T. Yamada 24 (TNS). Tochigi Pref. Awano-machi, 2 Oct 1969, S. Okuyama 12599a, 12599b (TNS). Tottori Pref.: Yodoe-cho, 2 Aug 1995, S. Fujii 4368 (TNS). Yamagata Pref.: Asahi-machi, 10 Aug 1965, K. Iwatsuki 591 (TNS); Nishikawa-machi, 27 Aug 1986, T. Takahashi 309 (TNS); Yawata-machi, 29 Sep 2004, Y. Kadota 44129 (TNS); Yonezawa-shi, 17 Aug 1962, H. Koyama 1315 (TNS). Yamanashi Pref.: Ashiwada-mura, 6 Sep 1968, F. Konta 7155 (TNS); Fujiyoshidashi, 24 Aug 1976, F. Konta 11474 (TNS); Fujiyoshida-shi, 11 Oct 1977, K. Murata 1633 (TNS); Hayakawa-cho, 4 Aug 1961, M. Shinya 474 (TNS); Kamikuishiki-mura, 9 Sep 2004, J. Makino s.n. (TNS); Nanbu-cho, 26 Oct 1975, Y. Kadota 750 (TNS).
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