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1 Korean J. Pl. Taxon. 45(4): (2015) ISSN Korean Journal of Plant Taxonomy A taxonomic study of the genus Dioscorea L. (Dioscoreaceae) in Korea based on morphological characters Dae Hee Chung and Gyu Young Chung* Department of Medicinal Plant Resources, Andong National University, Andong , Korea (Received 12 November 2015; Revised 27 November 2015; Accepted 9 December 2015) 한국산마속 ( 마과 ) 의외부형태형질에의한분류학적연구 정대희 정규영 * 안동대학교생약자원학과 ABSTRACT: The morphological characters were observed in six taxa in the genus Dioscorea of Korea (D. polystachya, D. japonica, D. nipponica, D. quinqueloba, D. tenuipes, and D. dokoro). The underground structure and the formation of bulbil were useful characters for separating sect. Enanatiophyllum from sect. Stenophora. The important characters for discriminating taxonomic entities were as follows: the position of the anther and the shape of the pistil in the microstructural characters, including the protuberance of the base of the petiole, the shape and color of the flower, the shape of the fruit, and the shape of the seed wing. Based on the results, a key to Korean Dioscorea is provided and the distribution of these species in Korea is described. Keywords: Dioscorea, Dioscoreaceae, morphology, key, distribution 적요 : 한국산마속 (Dioscorea L.) 6분류군 ( 마 ; D. polystachya, 참마 ; D. japonica, 부채마 ; D. nipponica, 단풍마 ; D. quinqueloba, 각시마 ; D. tenuipes, 도꼬로마 ; D. dokoro) 을대상으로외부형태학적형질을관찰하였다. 지하부의유형, 주아의형성유무등이마절 (Enanatiophyllum) 과부채마절 (Stenophora) 을구분하는유용한형질로관찰되었으며, 엽병기부의돌기유무, 꽃의형태및색, 열매의형태, 종자날개의형태등을비롯한약의위치와암술형태와같은미세구조적형질등이종을식별하는중요한형질로판단되었다. 또한상기식별형질을바탕으로검색표를제시하였고, 한반도내마속식물의분포를확인하였다. 주요어 : 마속, 마과, 외부형태, 검색표, 분포 마속 (Dioscorea L.) 은백합목 (Liliales), 마과 (Dioscoreaceae) 에포함되는분류군으로주로열대지역, 동아시아, 지중해연안및남 북아메리카등에약 600 종이분포한다 (Coursey, 1969; Li et al., 1978; Ohwi, 1984; Im, 2000; Lee, 2006). 마속은 Linnaeus (1753) 에의해잎의형태적차이로처음기재된후분류학적한계설정에많은이견과혼동이있어왔다. Burkill (1960) 은마과를자웅동주와자웅이주의형질로 Paradioscreeae 와 Dioscoreeae 의 2 족으로나누고, 지하부의 *Author for correspondence: gychung@andong.ac.kr Copyright 2015 the Korean Society of Plant Taxonomists 유형, 열매및종자의형태를바탕으로마속을 23 개절로구분하였다. 또한 Beccari (1870) 는자방의 1 실속에포함되어있는종자를바탕으로마속을 2 개의종자가포함된마속과그이상인 Stenomeris 속 (Stenomeris Planch.) 으로구분하였으며, Knuth (1850) 는종자에발달되는날개의특징으로종자를원형으로둘러싼형태인 Eudioscorea 아속 (Eudioscorea Kunth) 과한쪽으로발달하는 Helmia 아속 (Helmia Kunth) 으로구분하였다. 그리고 Engler and Prantl (1897) 는지하부의유형, 잎의형태, 꽃의형태및종자의형태등의형질을바탕으로 39 개절로구분하였으며, 이의분류체계는현재널리적용되고있다. 이외에도 Matsuno (1918), Maekawa et al. (1962) 등의다수학자들은잎의형태, 잎차례, 주아의유무, 380

2 A taxonomic study of the genus Dioscorea in Korea 381 엽병기부형태, 꽃의형태및색, 종자의형태등을속내분류체계를결정하는주요형질로취급하였으며, 최근지하부와열매의색소체유전자를이용한계통분석 (Wilkin et al., 2005) 및 DNA염기서열을이용한분자생물학적연구가이루어지면서분류군간의유연관계와진화양상이논의되고있다 (Gao et al., 2008; Hsu et al., 2013; Gawande et al., 2015). 한국산마속에대한연구는 Palibin (1901) 의각시마 (D. tenuipes Franch. & Sav.) 에대한기록을시작으로 Nakai (1911, 1952) 는 7종, Mori (1921) 는 8종 1변종, Park (1949) 은 7종 1변종을기록하고있으며, 현재는마 (D. batatas Decne.), 참마 (D. japonica Thunb.), 부채마 (D. nipponica Makino.), 단풍마 (D. quinqueloba Thunb.), 각시마 (D. tenuipes Frnach. & Sav.), 도꼬로마 (D. tokoro Makino.), 국화마 (D. septemloba Thunb.), 둥근마 (D. bulbifera L.) 등 8종이기록되고있다 (Lee, 2003; Lee, 2006; Oh and Lee, 2007). 하지만이는단순한학명의기록또는도감의기재로서마속분류군의형태적유사성및개체내심한변이의폭으로인하여분류학적처리에어려움이있는실정이다. 최근 Kang et al. (1992) 과 Oh et al. (1995, 1996) 에의해한국산마속의분류학적연구가진행되었으나, Kang et al. (1992) 은참마의재료가생략된상황에서문헌상의식별형질로만참마를구분하여마와참마를혼동한채연구를수행하였고, Oh et al. (1995, 1996) 의연구는분류군간의뚜렷한식별형질을제시하지못하였으며, 특히현재국내에서마의학명을 D. batatas (Decaisne, 1854) 와 D. polystachya (Turczaninow, 1837) 를혼용하여사용하는등최근의분류학적연구결과를반영하지못하고있는실정이다. 따라서본연구의목적은한국산마속의전반적인외부형태및미세구조의형질을재검토하고, 분류군간의유용한식별형질을파악하여종의명확한한계를구분하는데있다. 적형질을관찰 ( 해부현미경 ; Olympus Ax-70, Japan; Olympus DP2-BSW, Germany), 측정및촬영하였고, 주요형질을도해하였다. 미세구조관찰을위하여 FAA 에고정되어있는성숙개체의암술과수술을취하여 1.5% glutaraldehyde 에전처리후 0.1 M phosphate buffer (ph 6.8) 로 2 회세척, % ethanol 의단계적인탈수과정을거처 isoamylacetate 에 2 회치환하였다. 이를 critical point dryer (HCP-2) 로건조후 ion sputter 로 gold coating (thickness: A) 한후주사전자현미경 (Hitach; S-2500C, Japan; 20 Kv; 80 W; working distance 15 mm) 으로관찰및촬영하였다. 결 과 1. 지하부구조마절 (Enanatiophyllum) 의마와참마는수직으로곧게뻗는괴근의형태로발달하며, 부채마절 (Stenophora) 의부채마, 단풍마, 각시마, 도꼬로마는수평으로퍼지는근경의형태로발달한다 (Fig. 1, Table 1). 2. 주아마절의마와참마는엽액에주아가형성되고, 부채마절의부채마, 단풍마, 각시마, 도꼬로마는주아가형성되지않았다 (Fig. 2, Table 1). 재료및방법 재료실험재료는 2010 년 5 월부터 2015 년 10 월까지개화기와결실기를중심으로채집된성숙한개체를 FAA (Formalinacetic acid-ethyl alcohol) 에고정한액침표본과안동대학교생약자원학과 (ANH), 국립수목원 (KH), 서울대학교산림과학부 (SNUA) 에소장되어있는건조표본을사용하였다. 또한일부는안동대학교실험포장에이식하여재배한재료를사용하였으며, 관찰및실험과분포확인을위해사용된표본은 Appendix 1 에기재하였다. 또한현재까지관찰및채집이이루어지지못한국화마 (D. septemloba) 및둥근마 (D. bulbifera) 는본연구에서제외하였다. Fig. 1. The types of root of Korean Dioscorea L. A. Vertical tuberous root; B. Horizontal rhizome 방법동일집단에서성숙개체를대상으로건조표본, 액침표본, 생체를이용하여영양기관및생식기관의정성 정량 Fig. 2. The types of bulbil of Korean Dioscorea L. A. Bulbil; B. Non-Bulbil

3 Korean J. Pl. Taxon, Vol. 45, No. 4 Table 1. Measurement on external morphology of genus Dioscorea in Korea. Underground part Inflorescence Bulbil Petiole base Leaf Periacth Fruit Seed Type Male Female Present or Wing Form Shape Type Tepal color Tepal shape Shape Shape Absent position D. polystachya tuberous spike spike Present non-protuberance deltate closed white widely-ovate obovate-globose domed all round D. japonica tuberous spike spike Present non-protuberance narrowly-triangular closed white widely-ovate broadly ovate globose all round D. nipponica rhizomatous raceme raceme Absent non-protuberance deltate-ovate campaniform yellowish obovate Oblong oblong one side D. quinqueloba rhizomatous raceme raceme Absent protuberance deltate-ovate campaniform orange obovate broadly ovate globose all round D. tenuipes rhizomatous raceme raceme Absent protuberance triangular-cordiform reflexed yellowish-green linear broadly ovate globose all round D. dokoro rhizomatous raceme raceme Absent non-protuberance cordiform campaniform ivory ovate broadly ovate domed all round Leaf (cm) Male flower (mm) Petiole Blade Outer tepal Inner tepal * * L/W D. polystachya 2.6 (3.4) (5.6) (4.8) (1.9) (1.9) (1.7) (1.4) (1.3) (1.1) 1.3 D. japonica 1.7 (3.4) (8.7) (4.3) (2.4) (1.8) (2.4) (1.7) (1.6) (1.4) 1.6 D. nipponica 3.3 (5.8) (8.3) (6.9) (2.6) (2.5) (2.2) (1.5) (2.2) (1.4) 1.6 D. quinqueloba 3.8 (4.9) (9.6) (8.7) (1.0) (3.6) (1.8) (0.9) (1.6) (0.8) 1.0 D. tenuipes 2.0 (3.1) (7.2) (4.4) (1.3) (3.8) (1.6) (0.5) (1.5) (0.5) 0.6 D. dokoro 5.3 (6.7) (7.2) (5.7) (1.9) (3.6) (2.3) (1.3) (2.2) (1.3) 1.5 Male flower (mm) Female flower (mm) Stamen Anther Outer tepal Inner tepal * D. polystachya 0.9 (1.1) (0.4) (0.5) (0.25) (1.5) (2.3) (1.5) (1.4) (1.4) (1.3) 1.5 D. japonica 0.7 (0.8) (0.3) (0.5) (0.2) (1.7) (2.1) (1.5) (1.4) (1.2) (1.5) 1.6 D. nipponica 0.6 (0.7) (0.2) (0.5) (0.2) (1.4) (2.8) (1.6) (0.8) (1.5) (0.8) 1.0 D. quinqueloba 0.6 (0.7) (0.1) (0.3) (0.1) (1.9) (3.9) (2.0) (1.3) (1.8) (1.2) 1.6 D. tenuipes 0.6 (0.6) (0.8) (0.3) (0.1) (0.9) (3.6) (1.8) (0.4) (1.8) (0.4) 0.5 D. dokoro 0.7 (0.8) (0.1) (0.5) (0.2) (2.3) (2.8) (1.7) (1.0) (1.7) (1.0) 1.2 Female flower (mm) Fruit (mm) Seed (mm) Seed Wing (mm) Pistil * * * * * * * * D. polystachya 0.3 (0.4) (0.9) (14.4) (17.1) (3.9) (3.5) (9) (13.6) 16.5 D. japonica 0.4 (0.4) (0.9) (17.1) (27.4) (4.2) (4.2) (13.5) (15.8) 17.3 D. nipponica 1.2 (1.4) (1.0) (19.9) (17.8) (5.6) (3.4) (13.6) (6.6) 7.5 D. quinqueloba 0.6 (0.7) (0.9) (13.1) (15.9) (4.5) (3.5) (8.2) (10.4) 11.3 D. tenuipes 0.4 (0.5) (0.1) (16.0) (20.0) (5.0) (4.1) (9.4) (10.3) 11.4 D. dokoro 0.7 (0.8) (0.8) (23.5) (27.6) (5.2) (4.2) (9.8) (15.1) 16.1 * min (average) max 382 Dae Hee Chung and Gyu Young Chung

4 A taxonomic study of the genus Dioscorea in Korea 잎 1) 엽병엽병의길이는도꼬로마가평균 6.7 cm 로가장길고, 각시마가평균 3.1 cm 로가장짧게관찰되었다. 또한단풍마, 각시마의엽병기부에한쌍의작은다육성돌기가관찰된반면마, 참마, 부채마와도꼬로마는관찰되지않았다 (Fig. 3, Table 1). 2) 잎의형태잎의길이는단풍마가평균 9.7 cm 로가장길고, 마는평균 5.6 cm 로가장짧게관찰되었고, 너비는단풍마가평균 8.7 cm 로가장넓고, 각시마는평균 4.4 cm 로가장좁게관찰되었다. 또한마는삼각형, 참마는비율이 2:1 미만인좁은삼각형, 부채마와단풍마는삼각상난형, 각시마는삼각상심장형, 도꼬로마는심장형으로관찰되었으며, 마와참마는 0 1 쌍의측열편이갈라지고, 부채마와단풍마는 1 4 쌍의측열편이깊게갈라지며, 각시마와도꼬로마는갈라짐이관찰되지않았다 (Fig. 4, Table 1). 부채마와단풍마의잎을중앙열편 (middle-lobe), 1 측열편 (first-lobe), 2 측열편 (second- lobe) 으로구분하여비교하였다 (Fig. 5). 중앙열편의선단의형태는부채마와단풍마모두예두로관찰되었고, 1 측열편과 2 측열편의선단의형태는부채마가예두, 단풍마는둔두로관찰되었다. 중앙열편의길이는부채마가평균 5.4 cm, 단풍마는평균 5.3 cm, 너비는부채마가평균 3.3 cm, 단풍마는평균 3.9 cm 로관찰되었다. 1 측열편의길이는부채마가평균 2.1 cm, 단풍마는평균 2.0 cm, 너비는부채마가평균 1.4 cm, 단풍마는평균 2.0 cm 로관찰되었다. 2 측열편의길이는부채마가평균 1.6 cm, 단풍마는평균 1.3 cm, 너비는부채마가평균 1.4 cm, 단풍마는평균 1.7 cm 로관찰되었다. 또한중앙열편과 1 측열편사이의각은부채마가평균 57 o, 단풍마는평균 63 o 로관찰되었고, 1 측열편과 2 측열편사이의각은부채마가평균 77 o, 단풍마는평균 70 o 로관찰되었다 (Fig. 5, Table 2). 3) 부채마와단풍마잎의비교 Fig. 3. The petiole base of Korean Dioscorea L. A. Protrude papillae; B. Non-protrude papillae. Fig. 5. The comparison of leaf of D. nipponica and D. quinqueloba. A. D. nipponica; B. D. quinqueloba (1: of middle-lobe; 2: of middle-lobe; 3: of First-lobe; 4: of First-lobe; 5: of Second-lobe; 6: of Second-lobe; 7: First-angle; 8: Second-angle). Fig. 4. The shapes of leaf of Korean Dioscorea L. A. D. polystachya; B. D. japonica; C. D. nipponica; D. D. quinqueloba; E. D. tenuipes; F. D. tokoro.

5 384 Dae Hee Chung and Gyu Young Chung Table 2. The comparison between of D. nipponica and D. quinqueloba. Leaf Middle-lobe First-lobe apex * * apex * * D. nipoonica acute 4.1 (5.4) (3.3) 4.0 acute 1.0 (2.1) (1.4) 1.9 D. quinqueloba acute 2.7 (5.3) (3.9) 5.8 obtuse 1.0 (2.0) (2.0) 2.6 Leaf Second-lobe Angle ( ) apex * * first-angle second-angle D. nipoonica acute 0.5 (1.6) (1.4) (57) (77) 110 D. quinqueloba obtuse 0.8 (1.3) (1.7) (63) (70) 110 * min (average) max, scale cm Fig. 7. The shapes of tepal of Korean Dioscorea L. A. D. polystachya, D. japonica; B. D. nipponica, D. quinqueloba; C. D. tokoro; D. D. tenuipes. Fig. 6. The inforescence of Korean Dioscorea L. A. D. polystachya; B. D. japonica; C. D. nipponica, D. quinqueloba; D. tenuipes; D. tokoro. 4. 꽃단성화이며, 3 장씩의내 외화피로구성된다. 개화하는시기는 5 9 월이다. 1) 화서마와참마는자 웅화모두수상화서의형태로관찰되었으나, 마의자화경은지그재그형태로발달하였고, 참마는곧게발달하였다. 부채마, 단풍마, 각시마와도꼬로마는자 웅화모두총상화서로관찰되었다 (Fig. 6, Table 2). 2) 화피의형태개화시화피가펼쳐지지않는마와참마, 종형으로개화하는부채마, 단풍마및도꼬로마, 화피가뒤로젖혀지는각시마로관찰되었으며, 마와참마는화피편의형태가넓은난형, 도꼬로마는난형, 부채마와단풍마는도란형, 각시마는선형으로관찰되었다 (Figs. 7 8, Table 1). 3) 화피의색마와참마는꽃의색이모두흰색으로관찰되었으며, 부채마는옅은황색, 단풍마는짙은황색, 각시마는연녹색, 도꼬로마는상아색으로관찰되었다 (Table 1). 4) 약의위치마와참마및부채마는약이수술의전면, 단풍마, 도꼬로마는약이수술의측면, 각시마는수술의정단에위치하는것으로관찰되었다 (Fig. 9). 5) 암술의형태각시마는암술대의기부에서 3 갈래로갈라져분지하였고, 다른분류군은암술대의정단부에서 3 갈래로분지하는것으로관찰되었다 (Fig. 9). 5. 열매및종자 1) 열매열매의길이는도꼬로마가평균 23.5 mm 로가장길고, 단풍마는평균 13.1 mm 로가장짧게관찰되었고, 너비는도꼬로마가평균 27.6 mm 로가장넓고, 부채마는평균 15.9 mm 로가장좁게관찰되었다. 또한열매의형태는마는도란상구형, 부채마는장타원형, 참마및단풍마, 각시마, 도꼬로마는넓은도란형의형태로관찰되었다 (Fig. 10, Table 1). 2) 종자종자의길이는부채마가평균 5.6 mm 로가장길고, 마는

6 A taxonomic study of the genus Dioscorea in Korea 385 Fig. 8. The shape of flower of Korean Dioscorea L. Aa. Ab. D. polystachya; Ba. Bb. D. japonica; Ca. Cb. D. nipponica; Da. Db. D. quinqueloba; Ea. Eb. D. tenuipes; Fa. Fb. D. tokoro. Fig. 9. The shapes of stamen and pistil of Korean Dioscorea L. A. Stamen (a, b. D. polystachya; c, d. D. japonica; e, f. D. nipponica; g. D. quinqueloba; h. D. tenuipes; i. D. tokoro), B. Pistil (a. D. polystachya; b. D. japonica; c. D. nipponica; d. D. quinqueloba; e. D. tenuipes; f. D. tokoro).

7 386 Dae Hee Chung and Gyu Young Chung Fig. 10. The shapes of fruit and seed of Korean Dioscorea L. Aa. Ab. D. polystachya; Ba. Bb. D. japonica; Ca. Cb. D. nipponica; Da. Db. D. quinqueloba; Ea. Eb. D. tenuipes; Fa. Fb. D. tokoro. 평균 3.9 mm 로가장짧게관찰되었고, 너비는참마가평균 4.2 mm 로가장넓고, 부채마는평균 3.4 mm 로가장좁게관찰되었다. 종자는주변에막질의날개를가지고있으며, 날개의길이는부채마가평균 13.6 mm 로가장길고, 단풍마는평균 8.2 mm 로가장짧게관찰되었다. 너비는참마가평균 15.8 mm 로가장넓고, 부채마는평균 6.6 mm 로가장좁게관찰되었다. 또한마와도꼬로마는반구형 (domed), 부채마는장타원형, 참마및단풍마, 각시마는구형의형태로관찰되었으며, 종자가막질날개의기부에위치하는부채마와중심부에위치하는그외분류군으로관찰되었다 (Fig. 10, Table 1). 이상의결과를바탕으로작성한한국산마속의검색표는다음과같다. 한국산마속의검색표 1. 지하부는수직으로뻗는괴근이며, 주아가형성되고, 수상화서이다. 2. 잎은삼각형, 자화경은지그재그형태로발달하며, 열매의너비는 mm 이다 마 D. polystachya 2. 잎은좁은삼각형, 자화경은곧게발달하며, 열매의너비는 mm 이다 참마 D. japonica 1. 지하부는수평으로뻗는근경이며, 주아가형성되지않고, 총상화서이다. 3. 잎은 1 4 쌍의측열편이갈라진다. 4. 엽병의기부에는다육성돌기가발달하고, 잎의측열편은둔두, 종자는막질날개의중앙에위치한다 단풍마 D. quinqueloba 4. 엽병의기부에는다육성돌기가발달하지않고, 잎의측열편은예두, 종자는막질날개의기부에위치한다 부채마 D. nipponica 3. 잎은측열편이갈라지지않는다. 5. 엽병의기부에는다육성돌기가발달하고, 잎은삼각상심장형, 화피는뒤로젖혀지며, 약은수술대의정단에위치하고, 암술은기부에서갈라진다 각시마 D. tenuipes 5. 엽병의기부에는다육성돌기가발달하지않고, 잎은심장형, 화피는뒤로젖혀지며, 약은수술대의측면에위치하고, 암술은정단에서갈라진다 도꼬로마 D. tokoro 고 찰 한국산마속 (Dioscorea L.) 분류군에대한외부형태적형질인지하부의유형, 주아, 잎의형태및잎차례, 엽병기부의돌기유무, 꽃의형태및색, 암 수술의형태, 열매의형태, 종자및종자날개의형태등을비롯한미세구조인수술과암술의형태를본연구에서재검토하였다. 일반적으로마속내절을분류하는형질중지하부의유형 (Engler and Prantl, 1897; Burkill, 1960) 과주아의유무 (Matsuno, 1918; Maekawa, 1962) 는수직으로곧게뻗는괴근 (tuber) 의형태를가지고, 주아가형성되는마절 (Enantiophyllum) 과수평으로뻗는근경 (rhizome) 의형태를가지고, 주아가형성되지않는부채마절 (Stenophora) 로관찰되어절을분류하는형질로서의가치가있으나, 그이외의외부형태적형질인꽃의형태, 열매및종자의형태등 (Engler and Prantl, 1897; Burkill, 1960) 은한국산마속에서는절간의뚜렷한식별형질을확인할수없었다. 마절 (Enanatiophyllum) 에속하는마 (D. polystachya) 와참마 (D. japonica) 는매우유사한분류군이며, 잎의형태와잎차례, 줄기및엽병의색등이두분류군을식별하는형질로알려져있다 (Matsuno, 1918; Ohwi, 1984). 하지만 Kang et al. (1992) 은열악한생육환경에서자생하는마와

8 A taxonomic study of the genus Dioscorea in Korea 387 참마는식별형질인잎의형태나잎차례에차이가없었고, 마의줄기와엽병은색을띄지않아종을식별하는형질로생각하기에는어려움을있다고보고하였으며, Oh et al. (1995) 는마와참마는같은집단및동일개체내에서도잎의변이가다양하여잎의형태로마와참마를구별하는것이적합하지않다고보고한바있다. 마와참마는선행되어진연구에서의형질인잎차례나줄기및엽병의색등 (Matsuno, 1918; Ohwi, 1984) 에서종간의식별형질이관찰되지않았고, 그외의꽃의형태, 내 외화피의형태및크기, 종자의형태및크기등의형질에서도미세한양적형질의차이만이관찰되었을뿐종을식별하는분류형질로서타당성을확인하지못했다. 하지만잎의형태가마는삼각형 (deltate), 참마는길이와너비의비율이 2:1 이상인좁은삼각형 (narrowly-triangular) 으로확인되었고, 열매의너비가마는 mm, 참마는 mm 로확인되어잎의형태로두종을분류한 Maekawa (1962) 와열매의크기로두종을분류한 Tominari (1984) 의언급과도일치하였다. 또한화경의형태가마의자화경이지그재그로발달하여, 곧게발달하는참마와차이점을확인하였다. 부채마절 (Stenophora) 에속하는부채마 (D. nipponica), 단풍마 (D. quinqueloba), 각시마 (D. tenuipes), 도꼬로마 (D. dokoro) 는잎의형태, 엽병기부의돌기유무, 꽃의형태및색, 열매및종자날개의모양등의외부형태에의해분류되고있으며 (Lee, 2003; Lee, 2006), 잎의측열편이깊게갈라지는부채마와단풍마, 측열편이갈라지지않는각시마와도꼬로마로구분된다 (Oh et al., 1995). 잎의측열편이갈라지는부채마와단풍마는잎의형태적유사성과큰폭의변이로인해종의식별에어려움이있다. 이에부채마와단풍마의잎을각열편으로구분지어관찰하였으며, 각열편의길이와너비, 열편사이각등의양적형질에대한차이는미세하였으나, 측열편선단의형태가부채마는예두인데반해단풍마는둔두의형태를띠어두종간의차이점을확인하였다. 또한단풍마는엽병의기부에돌기가발달하여그렇지않은부채마와차이점을확인하였으며, 꽃의색, 종자의형태등의형질에서두분류군간의차이점을확인하였다. 잎의측열편이갈라지지않는각시마와도꼬로마는잎의형태가삼각상심장형인각시마와심장형인도꼬로마로구분되었으며, 각시마는단풍마와같이엽병기부에돌기가발달하여그렇지않은도꼬로마와차이점을확인하였다. 또한개화시화피의형태는각시마는선형이며뒤로젖혀지고, 도꼬로마는난형이며종형으로개화하는차이점을확인하였다. 미세구조인암술과수술의형태는마절의마와참마두분류군모두암술대의정단에서 3 갈래로분지하고, 약이수술대의전면에위치하는등두종간의형태적차이점은확인되지않았다. 반면부채마절의분류군에서는암술과수술의다양한형태를확인하였다. 각시마는암술대정단에서 3 갈래로분지하는다른분류군반해기부에서 3 갈래 로분지하였다. 또한약이수술대의정면에위치하는부채마, 측면에위치하는단풍마와도꼬로마, 상단부에위치하는각시마로관찰되어종을분류하는유용한형질이라판단된다. 현재국내와일본에서는마의학명을 Decaisne (1854) 가발표한 D. batatas 를사용하고있는반면중국에서는 Turczaninow (1837) 가발표한 D. polystachya 를사용하고있다. 두학명은지하부유형, 주아의존재, 잎의형태및열매의형태등의외부형태적형질을바탕으로발표된합법명이나, 선취권이앞선 D. polystachya Turcz 를사용하는중국의학명처리가타당한것으로판단된다. 상기식별형질을바탕으로한반도내마속의분포를확인하였다. 마는한반도전역에고루분포하는반면, 참마는제주도및남해안일대섬지역에서한정적으로분포하며, 부채마와도꼬로마는강원, 경기, 충북, 충남, 경북등의중부지역, 단풍마와각시마는전남, 경남, 제주도등의남부지역에분포하는것으로확인되었다 (Fig. 11). 단풍마와참마는한반도전역에분포한다고알려져있으나 (Park, 1949; Lee, 2003; Lee, 2006; Oh and Lee, 2007), 이는참마의실체를배재한체연구가수행되어졌거나 (Park, 1949) 마속의각분류군에대한식별형질의부재 (Lee, 2003; Lee, 2006; Oh and Lee, 2007) 로인해정확한동정이이루어지지않았기때문이라판단된다. 또한국화마 (D. septemloba) 는한반도의남해안지역과일본의 Sinkoku, Fig. 11. Distribution map of Korean Dioscorea L. D. polystachya; D. japonica; D. nipponica; D. quinqueloba; D. tenuipes; D. tokoro.

9 388 Dae Hee Chung and Gyu Young Chung Kyushu 지역에, 둥근마 (D. bulbifera) 는한반도전역을비롯한중국과일본의 Honshu, Sinkoku, Kyushu 지역의분포가보고되고있다 (Ohwi, 1984; Ding and Gilbert, 2000). 그러나국화마는잎의측열편이깊게갈라지며, 열매가도란상구형인특징, 둥근마는잎이심장형, 화피편의형태가피침형, 열매가장타원형인특징으로동속분류군과구분되어지며, 본연구를통하여 ANU, KH, SNUA 등에소장되어진국화마와둥근마로동정된표본을확인한결과, 잎의변이로인해측열편이깊게갈라진부채마와단풍마가국화마로오동정되어있었다. 또한마의어린개체가둥근마로오동정되어있었으며, 일본에서채집된표본만이둥근마로확인되었을뿐이다. 따라서본연구에서는국화마와둥근마의자생지를확인할수없었고, 따라서두종의국내분포여부에대해서는재고되어야할것으로판단된다. Literature Cited Beccari, O Nuovo Giornale Botanico Italiano Publicato da O. Beccari. Stabilimento di G. Pellas, Firenze. V Burkill, I. H The organography and the evolution of the Dioscoreaceae, the family of the yams. Botanical Journal of the Linnean Society 56: Coursey, D. G The future of the yam in world agriculture: an appraisal of some problems. Tropical Root and Tuber Crops Newsletter 2: 4 8. Decaisne, J Revue Horticole. Librairie agricole de la maison rustique, Paris. Ser. IV, iii. P Ding, Z. and M. G. Gilbert Dioscoreaceae. In: Wu, Z. and P. H. Raven. (eds), Flora of China. Flagellariaceae through Marantaceae. Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis. 24: Engler, A. and K. Prantl Die natürlichen Pflanzenfamilien V. 2(5). Wilhelm Engelmann, Leipzig. Pp Gao X., Y. P. Zhu, B. C. Wu, Y. M. Zhao, J. Q. Chen and Y. Y. Hang Phylogeny of Dioscorea sect. Stenophora based on the chloroplast matk, rbcl and trnl-f sequences. Journal of Systematics and Evolution 46: Gawande, P. A., N. V. Nimbhorkar and V. Thakare Distribution and morphodiversity analysis of genus Dioscorea from India with special reference to Satpura Hilly Ranges and Western Ghat of Maharashtra state, India. International Journal of Basic and Applied Sciences 4: Hsu, K. M., J. L. Tsai, M. Y. Chen, H. M. Ku and S. C. Liu Molecular phylogeny of Dioscorea (Dioscoreaceae) in East and Southeast Asia. Blumea 58: Im, R. J Flora Koreana. Vol. 9. The Science and Technology Publishing House. Pyongyang, Korea. Pp Kang, F. H., C. S. Lee and Y. C. Oh A systematic study of sect. Enantiophyllum and Stenophora taxa (Dioscorea, Dioscoreaceae) in Korea. Korean Journal of Plant Taxonomy 22(4): Knuth, C. S Dioscorineae. In: Enumeratio Plantarum. V. 5. Sumtibus J. G. Cottae, Stutgardiae et Tubingae. Pp Lee, T. B Coloured Flora of Korea. Vol.. Hayangmunsa, Seoul. Pp (in Korean). Lee, Y. N New Flora of Korea. Vol.. Kyohak Publising Co. Seoul. Pp (in Korean). Li, H. L., T. S. Liu, T. C. Huang, T. Koyama and C. E. De Flora of Taiwan. Epoch Publishing Co., Ltd. Vol. 5 Pp Linnaeus. C Diosocrea In: Species Plantarum. V. 2. Impensis Laurentii Salvii, Holmiae. Pp Maekawa, F., H. Hara and T. Tuyama Makino s New Illustrated Flora of Japan. Hohuryukan. Co. Ltd., Tokyo. Pp Matsuno, G Dioscorea in Japan. Journal of Japanese Botany 1: Mori, T An Enumeration of Plants hitherto Known from Corea. The government of Chosen, Seoul. Pp Nakai, T Flora Koreana. Pars secunda. The Journal of the College of Science, Imperial University of Tokyo. 31: 235. Nakai, T A synoptical Sketch of Korean Flora. Bulletin of the National Science Museum. Tokyo. 31: 148. Oh, Y. C., C. S. Lee and H. J. Lee A systematic study on Dioscorea sect. Enantiophyllum and Stenophora in Korea: Microstructure and chemistry. Korean Journal of Plant Taxonomy 25(1): Oh, Y. C., C. S. Chang and K. A. Lee Morphometric analyses of Dioscorea. Korean Journal of Plant Taxonomy 26(2): Oh, Y. C. and C. S. Lee The Genera of Vascular Plants of Korea. Academy Publishing Co., Seoul. Pp (in Korean). Ohwi, J Flora of Japan. Smithsonian Institution, Washington DC. Pp Palibin, J Conspectus Florae Koreae. Pars tertia. Trudy Imperatorskago S.-Peterburgskago Botaniceskago Sada. 19: Park, M. K An Enumeration of Korean Plants. Pp (in Korean). Tominari, T., S. Wanteri, S. Kitamura, J. Ohwi and Y. Satake Wild Flowers of Japan. Heibonsya, Tokyo. Pp Turczaninow, P. K. N. S Bulletin de la Societe Imperiale des Naturalistes de Moscou. 7: 158. Wilkin, P., P. Schols, M. W. Chase, K. Chayamarit, C. A. Furness, S. Huysmans, F. Rakotonasolo, E. Smets and C. Thapyai A Plastid gene phylogeny of the yam genus, Dioscorea: Roots, Fruits and Madagascar. Systematic Botany 30:

10 A taxonomic study of the genus Dioscorea in Korea 389 Appendix 1. Specimens examined (experimental materials was indicated bold and * ) Dioscorea polystachya Turcz. Gangwon-do, Mt. Mukbang, 18 Sep. 2009, C. S. Chang, (SNUA), Mt. Ballon, 14 Oct. 2010, J. W. Jang et al., J (KH), Mt. Ballon, 14 Oct. 2010, J. W. Jang et al., J (KH). Gyeonggi-do, Mt. Gwanggyo, 5 Sep. 2002, S. H. Ju et al., (SNUA), Mt. Taebong, 20 Aug. 2002, Y. Jeon et al., (SNUA), Mt. Hwanggeum, 22 Jul. 2004, H. N. Park et al., (SNUA), Mt. Wangbang, 9 Sep. 2006, K. S. Chang, (SNUA), Mt. Sigung, 27 Jul. 2007, G. Y. Chung et al., ANH (ANH) *, Mt. Yangja, 2 Aug. 2007, G. Y. Chung et al., ANH (ANH), Mt. Jungwon, 2 Aug. 2007, G. Y. Chung et al., ANH (ANH), Yeongjongdo, 15 Sep. 2009, S. H. Park et al., ParkSH94268 (KH). Gyeongsangbuk-do, Geumsu-myeon, 24 Aug. 2001, G. Y. Chung et al., EN (ANH), Pohang-si, 25 Sep. 2002, G. Y. Chung et al., EN (ANH), Mt. Omi, 22 Sep. 2006, G. Y. Chung et al., ANH (ANH), Mt. Eorae, 12 Aug. 2006, G. Y. Chung et al., ANH (ANH), Ulleung-gun, 14 Jul. 2008, G. Y. Chung et al., ANH (ANH), Mt. Seondal, 30 Jun. 1998, G. Y. Chung et al., s. n. (ANH), Mt. Jangjae, 13 Jul. 2000, G. Y. Chung et al., s. n. (ANH), Mt. Agu, 10 Aug. 2000, G. Y. Chung et al., s. n. (ANH), Mt. Bibong, 10 Aug. 2009, G. Y. Chung et al., ANH (ANH), Mt. Geumjang, 14 Aug. 2007, G. Y. Chung et al., ANH (ANH), Mt. Yuhak, 14 Jul. 2006, G. Y. Chung et al., ANH (ANH), Mt. Gapjang, 9 Oct. 2009, G. Y. Chung et al., ANH (ANH) *,Yeongdeok-gun, 1 Jul. 2010, G. Y. Chung et al., ANH (ANH) *. Gyeongsangnam-do, Mt. Sohak, 2 Oct. 2007, S. C. Ko et al., HNHM-A-1059 (KH), Mt. Jakdae, 7 Aug. 2003, G. Y. Chung et al., EN (ANH), Mt. Eungbong, 13 Aug. 2003, G. Y. Chung et al., (ANH) *, Mt. Gujae, 30 Jul. 2004, C. S. Chang et al., (SNUA). Chungcheongbuk-do, Mt. Dorak, 5 Sep. 1998, G. Y. Chung et al., s. n. (ANH), Eumseong-gun, 25 Jul. 2010, G. Y. Chung et al., ANH (ANH) *, Mt. Cheontae, 9 Oct. 2006, S. H. Park et al., ParkSH (KH). Chungcheongnam-do, Mt. Jillak, 15 Sep. 2001, E. S. Jeon, gwang10168 (KH), Mt. Cheonbang, 30 Aug. 2006, E. S. Jeon, ESJeon61311 (KH), Mt. Nam, 28 Jul. 2009, E. S. Jeon, ESJeon90739 (KH). Jeollabuk-do, Mt. Sangyeo, 7 Oct. 2004, G. Y. Chung et al., Buan(Sangyeobong) (ANH), Mt. Uisang, 30 Jun. 2004, G. Y. Chung et al., Buan(Uisangbong) (ANH), Mt. Yeonggu, 2 Jul. 2004, W. G. Baek, Jangsugun(younggusan) (KH), Munyeodo-ri, 18 Jul. 2012, S. H. Park et al., Parksh (KH). Jeollanam-do, Hwawon-myeon, 10 Oct. 2007, Y. H. Cho et al., WR (KH), Hwangjeon-myeon, 16 Sep. 2009, Y. H. Cho et al., WR (KH), Seondudo, 11 Jun. 2002, H. T. Im, (SNUA), Mt. Mabok. 12 Aug. 2003, M. J. Park et al., Goheung (KH), Mt. Baegun, 24 Aug. 2010, Chung et al., ANH (ANH) *, Mt. Baengnyeon, 11 Jul. 2004, J. H. et al., Najusi (Baengnyeonsan) (KH), Mt. Goseong, 31 Jul. 2004, H. T. Im, Jangseong (KH), Soryangdo, 23 Jul. 2008, Y. H. Cho et al., WR (KH). Dioscorea japonica Thunb. Jeollanam-do, Mt. Sanghwangbong, 23 Jul. 2010, S. C. Ko et al., HNHM (KH). Jeju-do, Jeju-si, 3 Aug. 2010, G. Y. Chung et al., ANH (ANH) *. Dioscorea nipponica Makino. Gangwon-do, Sindong-myeon, 5 Aug. 1958, T. B. Lee, (SNUA), Mt. Odae, 27 Jul. 2008, G. Y. Chung et al., ANH (ANH), Mt. Cheongtae, 4 Oct. 2008, G. Y. Chung et al., ANH (ANH), Mt. Hwangbyeong, 30 Aug. 2008, G. Y. Chung et al., ANH , ANH , ANH (ANH), Mt. Nochu, 29 Aug. 2008, G. Y. Chung et al., ANH (ANH) *, Mt. Maengdong, 23 Jun. 2008, G. Y. Chung et al., ANH (ANH), Mt. Cheongok, 30 Jun. 2008, G. Y. Chung et al., ANH (ANH), Mt. Seongnyong, 23 Jun. 2008, C. S. Jang, Seongnyongsan (KH), Mt. Biryong, 2 Aug. 2010, G. Y. Chung et al., ANH (ANH) *. Gyeonggi-do, Gwangneung, 25 Jun. 1961, T. B. Lee, (SNUA), Mt. Wonjeok, 21 Jun. 2007, G. Y. Chung et al., ANH (ANH). Gyeongsangbuk-do, Hyeongje-bong, 26 Jun. 1999, J. I. Jeon, (SNUA), Mt. Yuhak, 6 Jun. 2001, G. Y. Chung et al., EN (ANH), Mt. Irwol, 16 Jul. 2006, G. Y. Chung et al., ANH (ANH) *, Mt. Galla, 14 Jun. 2008, G. Y. Chung et al., ANH (ANH), Mt. Jacho, 13 Jul. 2010, G. Y. Chung et al., ANH (ANH) *.

11 390 Dae Hee Chung and Gyu Young Chung Chungcheongbuk-do, Mt. Baegun, 10 Jul. 2011, B. U. Oh et al., Baegunsan (CBU). Chungcheongnam-do, Mt. Gwangdeok, 24 Jul. 2005, Y. D. Kim et al., Cheonansi (Gwangdeoksan) (HUH). Dioscorea quinqueloba Thunb. Gyeongsangnam-do, Mt. Hogu, 13 Aug. 2003, G. Y. Chung et al., (ANH), Mt. Mangun, 19 Sep. 2003, G. Y. Chung et al., , (ANH), Mt. Eungbong, 13 Aug. 2003, G. Y. Chung et al., (ANH) *, Mt. Geum, 14 Aug. 2003, G. Y. Chung et al., (ANH), Mt. Geumo, 15 Aug. 2003, G. Y. Chung et al., (ANH), Mt. Jakdae, 7 Aug. 2003, G. Y. Chung et al., EN (ANH) *. Mt. Baegyang, 17 Aug. 1970, T. B. Lee et al., (SNUA), Mt. Baegun, 24 Aug. 2010, G. Y. Chung et al., ANH (ANH) *. Jeollabuk-do, Mt. Moak, 29 Aug. 2004, G. Y. Chung et al., Gimje (Moaksan) , Gimje(Moaksan) (ANH), Wido, 27 Aug. 2004, G. Y. Chung et al., Buan(Wido) (ANH), Sebong, 21 Jul. 2004, G. Y. Chung et al., Buan(Sebong) (ANH). Jeollanam-do, Goheung-gun, 23 Aug. 1970, T. B. Lee, (SNUA), Geomundo, 18 Aug. 1964, T. B. Lee, (SNUA), Jindo, 30 Aug. 1970, T. B. Lee, (SNUA), Imjado, 25 Aug. 2001, H. T. Im, (SNUA), Soando, 23 Sep. 1993, T. B. Lee et al., (SNUA), Odongdo, 15 Oct. 2005, C. S. Chang, (SNUA), Jindo, 30 Aug. 1970, T. B. Lee, (SNUA). Jeju-do, Mt. Bonggolmae, 24 Sep. 2013, S. H. Park, P (KH). Dioscorea tenuipes Franch. & Sav. Gyeongsangnam-do, Mt. Yeonhwa, 12 Aug. 2010, G. Y. Chung et al., ANH (ANH) *, Mt. Gujaebong, 30 Jul. 2004, H. Kim et al., Hadonggun(Gujaebong) (KH), Saengnim-myeon, 06 Aug. 2000, B. W. Han et al., skk0543 (KH). Jeollanam-do, Mt. Baegun, 24 Aug. 2010, G. Y. Chung et al., s. n. (ANH) *, Mt. Mudeung, 11 Sep. 2012, S. Y. Jung et al., P (KH), Mt. Seungdal, 22 Oct. 2007, Y. H. Cho et al., nam (KH), Mt. Sanghwangbong, 20 Aug. 2010, S. C. Ko et al., HNHM (KH), Mt. Suin, 18 Jul, 2007, Y. H. Cho et al., WR (KH). Dioscorea tokoro Makino. Gangwon-do, Cheongnyeongpo, 13 Aug. 2004, J. M. Chung et al., (KH), Mt. Yonghwa, 29 May. 2008, H. J. Lee, yonghwasan (KH), Mt. Gari, 15 Jun. 2011, K. O. Yoo et al., koyoo0118 (KH). Gyeonggi-do, Mt. Paldal, 31 May. 1967, T. B. Lee, (SNUA), Mt. Maok, 26 May. 2007, G. Y. Chung et al., ANH (ANH). Gyeongsangbuk-do, Mt. Maengdong, 28 Aug. 2006, G. Y. Jeong, Jeong210 (KH) *, Mt. podo, 7 Jul. 2007, G. Y. Chung et al., ANH (ANH) *, Mt. Galla, 6 Jun. 2008, G. Y. Chung et al., ANH (ANH) *, Mt. Jacho, 13 May. 2010, G. Y. Chung et al., ANH (ANH) *, Mt. Podo, 1 Oct. 2007, G. Y. Chung et al., ANH (ANH). Chungcheongbuk-do, Mt. Sosoki, 13 May. 2005, B. H. Oh et al., eumseonggun (sosogisan)-0505 (KH), Mt. Geumdan, 22 May. 1999, S, C, Chung et al., (CBU), Mt. Duta, 5 May. 2002, B. U. Oh et al., (CBU). Chungcheongnam-do, Sanan-ri, 11 Jun. 2006, E. S. Jeon, esjeon60608 (KH).