한국산 모시풀속(쐐기풀과)의 분류학적 연구 = A taxonomic study of the genus Boehmeria (Urticaceae) in Korea

국문 초록 (Abstract)

본 연구는 미기록 3종과 실체가 불분명한 것으로 여겨지는 한국특산 4종을 포함한 총 16분류군의 한국산 모시풀속을 대상으로 수행되었다. 외부형태학적, 해부학적 연구를 통해 주요 식별형질과 각 형질별 변이 폭을 검토하였고, 원기재문과 기준표본을 바탕으로 그 동안 실체가 혼동되어 온 한국산 모시풀속의 분류군을 규명하였다. 분자생물학적 연구와 전형질 분석으로 분류군 간의 유연관계를 파악하였고, 주요 식별형질에 근거하여 분류체계를 정립하였다. 또한, 기존에 알려진 주요 식별형질과 새로운 식별형질에 대한 분계분석을 통해 진화 경향성을 추론하였다. 이와 같은 결과를 토대로 분류학적 처리를 수행하여 정확한 학명을 부여하였고, 분류군 수를 정리하였다.
절의 구분 형질로는 탁엽의 부착위치, 탁엽의 합생 또는 이생 여부, 잎 이면의 백색 모용의 유무, 엽서의 종류, 개화시기, 줄기의 수(pith) 및 수과의 날개 유무가 유용한 것으로 나타났다.
열 또는 종을 구분하는 형질로는 생육지, 생활형, 줄기의 홈, 줄기의 색, 줄기의 모용, 탁엽의 젖혀짐 여부, 탁엽의 형태, 중부잎의 형태, 엽연의 형태, 거치의 균일성, 엽선의 갈라짐, 잎 이면의 주맥과 세맥의 모용, 화서의 형태, 수꽃화피 부속체의 유무, 수꽃화피의 모용, 단집과서의 배열 및 수과의 모용이 관찰되었다. 또한, 줄기의 해부학적 형질에서는 횡단면의 형태, 표피의 층 수, 인피 섬유 조직의 너비, 인피 섬유 세포의 두께 및 수의 형태가 종을 구분하는 형질로 나타났다. 엽병의 해부학적 형질에서는 횡단면의 형태, 사출수유관속의 유무, 인피 섬유의 유무 및 수의 유무가 종을 구분하는데 유용하였다. 잎의 해부학적형질에서는 향축면 중륵의 형태, 책상유조직의 수 및 중륵 유관속의 형태가 종의 구분형질로 관찰되었다.
한국산 모시풀속의 핵DNA의 ITS구간과 엽록체DNA의 ndhF, trnK에 대한 분자생물학적 연구결과는 절과 열의 분류체계에서 잘 지지되는 것으로 확인되었다. 모시풀절(sect. Tilocnide)과 왜모시풀절(sect. Duretia) 등 2개의 절로 분기하였고, 모시풀절 내에는 왜모시풀열(ser. Longispicae), 긴잎모시풀열(ser. Sieboldianae), 그리고 좀깨잎나무열(ser. Spicatae) 등 3개의 열로 다시 구분되었다. 그러나 38개의 외부형태학적 및 해부학적 형질에 근거하여 수행된 전형질 분석 결과는 절 수준에서는 분자생물학적 연구결과와 동일하게 나타났지만 왜모시풀열과 긴잎모시풀열은 같은 유집군을 형성하였다.
주요 형질별 분계분석에서는 생활형, 줄기의 모용, 엽서, 잎 이면의 모용, 암화서, 수꽃화피의 부속체, 수꽃화피의 모용 및 수과의 모용에 대한 진화경향성을 파악하여 원시형질과 파생형질을 추론하였다.
한국산 모시풀속의 분류학적 처리 결과, 기존 ‘var. tenacissima’로 취급되었던 섬모시풀의 학명은 종 수준에서 선취권을 갖는 B. nipononivea로 학명을 정리하였다. 거북꼬리(B. tricuspis)는 개모시풀(B. platanifolia)에 이명 처리되었으며, 개모시풀의 학명은 B. platanifolia (Maxim.) C. H. Wright로 취급하였다. 한국 특산종으로 알려진 제주긴잎모시풀(B. nakaiana), 섬거북꼬리(B. taquetii)은 긴잎모시풀(B. sieboldiana)에 이명 처리하였고, 털긴잎모시풀(B. hirtella)과 제주모시풀(B. quelpaertensis)은 실체가 확인되었으나 일본에도 분포하는 것으로 확인되었다.
결과적으로 한국산 모시풀속을 2절 3열 총 13분류군으로 분류하였다. 첫 번째 모시풀절에는 모시풀, 섬모시풀이 해당되고, 두 번째 왜모시풀절 내의 왜모시풀열에는 왕모시풀(B. holosericea), 왜모시풀(B. longispica), 개모시풀, 제주모시풀이 속하며, 긴잎모시풀열에는 털긴잎모시풀, 제주톱모시풀(B. nakashimae), 긴잎모시풀이 해당되고, 이 절은 본 연구에서 새롭게 열 계급으로 격하시켰다. 마지막으로 좀깨잎나무열은 톱거북꼬리(B. gracilis), 풀거북꼬리(B. paraspicata), 참거북꼬리(B. silvestrii), 좀깨잎나무(B. spicata)가 속한다. 본 연구에서 각 분류군에 대한 분류학적 처리, 기재문, 도판, 기준표본 및 검색표를 제시하였다.

다국어 초록 (Multilingual Abstract)

This study was conducted on 16 taxa of Korean Boehmeria including three unrecorded species and four endemic species whose taxonomic identities are unclear. Through external morphological and anatomical studies, the key characteristics and range of variation of each characteristic were reviewed. The taxa of the Korean Boehmeria, whose identities have been unclear, were confirmed based on the original descriptions and type specimens. Molecular and phenetic analyses were used to identify the relationships among taxa, and a classification system was established based on key characteristics and molecular results. Additionally, evolutionary trends were inferred from the results of cladistic analysis of previously known and new key characteristics. Based on these results, accurate scientific names were assigned by taxonomic treatment, and the number of taxa was determined.
The attachment and connation of the stipule, white trichome on the leaf abaxial surface, phyllotaxis, flowering initiation, pith of the stem, and wing of the achene were useful in classifying the section.
The habitat, life form, groove, color, trichome of the stem, the, stipule shape, shape, margin, tooth, apex of the middle leaf, trichome, distribution of the trichome, basal cell on the leaf abaxial surface, inflorescence type, trichome, distribution of the trichome, basal cell of the staminate flower perianth, glomerule shape of the infructescence, and trichome of the achene were useful in classifying the series or species. Regarding the anatomical characteristics of the stem, the shape of the transverse section, the depth of the groove, the number of epidermal layers, the width and thickness of the bast fiber cell, and the presence and shape of the pith were useful characteristics at the species level. Among the anatomical characteristics of the petiole, the shape of the transverse section and the presence of the medullary bundle, bast fiber, and pith were useful for distinguishing the species. Regarding the anatomical characteristics of the leaf, the midvein shape of the adaxial surface, the number of palisade parenchyma, and the vascular bundle shape of the midvein were useful characteristics among the species. Regarding the anatomical characteristics of the achene, the presence of the wing and the number of endocarp layers were useful for understanding the relationships among the species.
The analytical results based on nrDNA (ITS) and cpDNA (ndhF, trnK) from Korean Boehmeria were well supported at the section and series levels. Korean Boehmeria is divided into two sections: sect. Tilocnide and sect. Duretia. Sect. Duretia was separated into three series: Longispicae, Sieboldianae, and Spicatae. However, phenetic analysis derived from 38 morphological and anatomical characteristics showed that the classification at the section level was the same as the results of the molecular study, but ser. Longispicae and ser. Sieboldianae clustered in the same clade.
The primitive and derived characteristics of life form, phyllotaxis, shape of the pistillate inflorescence, appendage of the staminate flower perianth, trichomes of the stem, leaf abaxial surface, staminate perianth, and achene were confirmed by cladistic analysis.
‘Seom-mo-si-pul’ treated as B. nivea var. tenacissima was recognized as B. nipononivea based on its external morphological characteristics and molecular characteristics. B. tricuspis was treated as a synonym of B. platanifolia; I suggest the use of B. platanifolia (Maxim.) C. H. Wright, according to Art. 46.6 of the Shenzhen code, for B. platanifolia. The taxonomic treatment of the four endemic species was as follows: B. nakaiana and B. taquetii were individuals with continuous variation in B. sieboldiana. Although the identities of B. hirtella and B. quelpaertensis were confirmed, they were not treated as endemic species because they are also distributed in Japan.
In conclusion, Korean Boehmeria was classified into a total of thirteen species in two sections and three series. Taxonomic treatment, descriptions, photographs, type specimens, and a key for each taxon are provided in this study.

목차 (Table of Contents)

• Abstract ⅰ
• List of Tables ⅳ
• List of Figures ⅴ
• List of Appendices ⅸ
• Ⅰ. 서론 1
• Ⅱ. 재료 및 방법 11
• 1. 재료 11
• 2. 방법 12
• 가. 외부형태학적 연구 12
• 나. 해부학적 연구 12
• 다. 분자생물학적 연구 14
• 라. 전형질 분석 21
• 마. 주요 형질별 분계분석 21
• Ⅲ. 관찰 및 결과 22
• 1. 외부형태학적 연구 22
• 2. 해부학적 연구 46
• 3. 분자생물학적 연구 59
• 4. 전형질 분석 73
• 5. 주요 형질별 분계분석 79
• Ⅳ. 분류군의 기재 85
• Ⅴ. 고찰 157
• Ⅵ. 적요 175
• Ⅶ. 인용문헌 177
• Appendix 193

참고문헌 (Reference)

1. Flora Coreana, Im, R. J., Vol. 1 Pp. 260–268in Korean, , 1996

2. Flora of Japan, Ohwi, J., Pp. 440–443in Japanese, , 1953

3. Plant Systematics, Simpson, M. G., 3rd ed 774 pp, , 2019

4. New Flora of Korea, Lee, Y. N., Vol. I. Pp. 255–257in Korean, , 2006

5. Coloured Flora of Korea, Lee, T. B, Vol. I. Pp. 236 –239.in Korean, , 2003

6. Urtica frutescens THUNB, Koidzumi, G., 10: 223–224, , 1941

7. Lineamenta Florae Koreae, Lee, W. T., Pp. 196–200in Korean, , 1996

8. Plant resources of Korea, Lee, T. B., 20: 89–228, , 1969

9. Revisio Boehmeriae Sinicae, Wang, W. T., 3: 307–328.in Chinese, , 1981

10. Seed dispersal by adhesion, Sorensen, A. E., 17: 443–463, , 1986

11. Revisio Boehmeriae Sinicae (cont.), Wang, W. T., 3: 401–416in Chinese, , 1981

12. A new rank of Boehmeria (Urticaceae), Wang, W. T., 20: 204–204. (in Chinese, , 1982

13. Notulae ad Plantas Japoniae & Koreae, Nakai, T., XXXIV 41: 501–522, , 1927

14. Chromosomal Numbers of Flowering Plants, Fedorov, A. A, 254 pp, , 1969

15. GBIF Secretariat GBIF Backbone Taxonomy, Boehmeria Jacq in, Retrieved 2023 from https://doi. org/10.15468/39omei, , 2022

16. Cuticular studies as an aid to plant taxonomy, Stace CA, 4: 3–78, , 1965

17. Conspectus Florae Koreae.Ericaceae-Salicaceae), Palibin, J, Pars II. 18: 147–198, , 1901

18. Coloured Standard Illustrations of Korean Plants, Lee, W. T., Pp. 66–67in Korean, , 1996

19. Unraveling heteroplasmy patterns with NOVOPlasty, Dierckxsens, N., P. Mardulyn, G. Smits, 2: 1–10, , 2020

20. An enumertion of plants hitherto known from Corea, Mori. T., 174 pp, , 1922

21. MRBAYES: Bayesian inference of phylogenetic trees, F. Ronquist, Huelsenbeck, J. P., 17: 754–755, , 2001

22. The Sex Phenotype and Evolution in Boehmeria Jacq, G. G. Zang, Zhao, L. N., J. H. Cheng, Y. J. Li, 25: 209–212, , 2003

23. Magnoliophyta Fossilia URSS Ulmaceae– Betulaceae, Takhtajan, A, Vol. 2 216 pp. (in Russian, , 1982

24. Quantitative phyletics and the evolution of anurans, Kluge, A. G., J. S. Farris, 18: 1–32, , 1969

25. A phylogenetic study of the genus Boehmeria in Korea, Liang, S. A, 90 pp, , 2009

26. Evaluation of 10 plant barcodes in Bryophyta (Mosses), X. J. Ge, Liu, Y., T. Cao, H. F. Yan, 48: 36–46, , 2010

27. The retention index and the rescaled consistancy index, Farris, J. S., 5: 417–419, , 1989

28. Evolutionary Trends in the Inflorescence of Angiosperms, Stebbins, G. L., 162: 501–528, , 1973

29. A taxonomic study on Boehmeria tricuspis complex in Korea, Cho, H. J, 99 pp, , 2018

30. Evolution of agamospermous races in Boehmeria and Eupatorium, Yahara, T., 5: 183–196, , 1990

31. NTSYS-pc: Numerical Taxonomy and Multivariate Analysis System, Rohlf, F. J, Version 2.2, , 2005

32. The genera of the Urticaceae in the southeastern United States, Miller, N. G., 52: 40–68, , 1971

33. The effect of cuticular hardening on the form of epidermal cells, Watson, R. W., 41: 223–229, , 1942

34. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods), Swofford, D. L., Version 4.0a169, , 2002

35. Chemical composition of different parts of ramie (Boehmeria nivea), Y. M. Kim, Cho, S., J. Lee, S. Lee, Y. S. Jung, H. B. Kim, E. J. Cho, 44: 95–103, , 2017

36. Contribution a l'etude cytotaxonomique des moracees et des urticacees, Le Coq, C., 70: 385–426, , 1963

37. La ramie. Ètudes morphologique et taxonomique en vue de la sélétion, Rabéchault, H., 242 pp, , 1951

38. Phylogeny of genus Boehmeria Jacq. reconstructed by RAPD fingerprinting, Guo, A. P., 82 pp. (in Chinese, , 1999

39. Genetic diversity and population structure of ramie (Boehmeria nivea L.), A. G. Zhu, J. H. Chen, X. F. Wang, Ni, J. L., Z. M. Sun, Y. Xu, M. B. Luan, 115: 340–347, , 2018

40. New insights into the functions of carbon–calcium inclusions in plants, D. Nikolopoulos, P. Bresta, Karabourniotis, G., H. T. Horner, G. Liakopoulos, 228: 845–854, , 2020

41. Trichomes and stomata, and their taxonomic significance in the Urticales, J. A. Inamdar, Gangadhara, M., 127: 121–137, , 1977

42. A karyological study on five species in three sections of genus Boehmeria, Zang, G. G., 1–6, , 1993

43. A revision of Boehmeria sect. Duretia ser. Spicatae (Urticaceae) of China, Wang, W. T., 36: 805–814.in Chinese, , 2016

44. Classification, Evolution and Phylogeny of the Families of Monocotyledons, Goldberg, A, 314 pp, , 1989

45. Systematics and population genetics of the Boehmeria platanifolia complex, Hwang, S. H., 117 pp, , 2020

46. An Illustrated Flora of Nippon, with the Cultivated and Naturalized Plants, Makino, T., 1070 pp.in Japanese, , 1940

47. A taxonomic revision of the Boehmeria spicata complex (Urticaceae) in Korea, Jo, H. J., G. Y. Chung, S. Jeong, J. Y. Kim, Yuri Lee, S. H. Park, M. J. Kwon, 51: 115–132, , 2021

48. Gapped BLAST and PSI-BLAST: A new generation of protein database search programs, W. Miller, D. J. Lipman, A. A. Schaffer,, T. L. Madden,, Altschul, S. F.,, Z. Zhang, J. Zhang, 25: 3389–3402, , 1997

49. Relationships among Chinese Boehmeria species and the evolution of various clade, M. B. Luan, X. R. Chen, W. C. Liu, Z. M. Sun, Liang, Y. K., J. H. Chen, Q. Tang, 148: 112092, , 2020

50. Robinson a century on: the nomenclatural relevance of Roxburgh's Hortus Bengalensis, Turner, I. M., 62: 152–172, , 2013

51. On the origin of the fig: phylogenetic relationships of Moraceae from ndhF sequences, G. D. Weiblen, Datwyler, S. L., 91: 767–777, , 2004

52. Primer3_masker: integrating masking of template sequence with primer design software, L. Kaplinski, Koressaar, T., K. Raime, R.reson, M. Lepamets, M. Remm, 34: 1937–1938, , 2018

53. SSR markers associated with fiber yield traits in ramie (Boehmeria nivea L. Gaudich), X. F. Wang, Z. M. Sun, J. H. Chen, C. C. Liu, Y. Xu, Luan, M. B., 107: 439–445, , 2017

54. Studies on the karyotype and giemsa C-banding pattern on wild resources of Boehmeria, Y. C. Chen, Yang, R. F., Q. G. Guo, L. H. Wang, H. l. Sun, 22: 6–11, , 2000

55. Apomixis in the genus Boehmeria (cytological studies of the Apomixis in Angiosperms I), Okabe, S., 29: 207–215, , 1963

56. The phylogenetic significance of stomata and trichomes in the Labiatae and Verbenaceae, Cantino, P. D., 71: 323–370, , 1990

57. Universal primers for the amplification of the plastid trnK/matK region in land plants, D. Quandt, Wicke, S., 66: 285–288, , 2009

58. The Old World species of Boehmeria (Urticaceae, tribus Boehmerieae): A taxonomic revision, Wilmot-Dear, C. M., I. Friis, 58: 85– 216, , 2013

59. Function and evolution of a minimal plastid genome from a nonphotosynthetic parasitic plant, J. D. Palmer, C. W. Morden, Wolfe, K. H., 89: 10648–10652, , 1992

60. MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform, Katoh, K., K. I. Kuma, T. Miyata, K. Misawa, 30: 3059–3066, , 2002

61. Phylogenetic signal in matK vs. trnK: A case study in early diverging eudicots (angiosperms), C. Black, J. G. Burleigh, D. Diouf, Hilu, K. W., 48: 1120–1130, , 2008

62. Testing Darwin's transoceanic dispersal hypothesis for the inland nettle family (Urticaceae), C. J. Chen, J. Liu, Y. H. Luo, B. S. Amorim, R. I. Milne, M. W. Cadotte, J. Provan, H. Wang, D. Z. Li, Wu, Z. Y., 21: 1515–1529, , 2018

63. Why some stems are red: cauline anthocyanins shield photosystem II against high light stress, H. S. Neufeld, Gould, K. S., D. A. Dudle, 61: 2707–2717, , 2010

64. The genera of flowering plants (Angiospermae) Based Principally on the Genera Plantarum of G., Hutchinson, J, Vol. 2 659 pp, , 1967

65. The influence of water stress and temperature on leaf pubescence development in Encelia farinosa, Ehleringer, J., 69: 670–675, , 1982

66. The New World species of Boehmeria and Pouzolzia (Urticaceae, tribus Boehmerieae): A taxonomic revision, Wilmot-Dear, C. M., I. Friis, 129: 1–103, , 1996

67. Evolution of trichome types and its systematic significance in the genus Phlomoides (Lamioideae–Lamiaceae), Y. Salmaki, Khosroshahi, E. E., 37: 1–14, , 2019

68. Laxative and antioxidant effects of ramie (Boehmeria nivea L.) leaf extract in experimental constipated rats, E. J. Choi, J. -J. Lee, Lee, H. -J, S. Park, 8: 3389–3401, , 2020

69. Tropical Asian Origin, boreotropical migration and long-distance dispersal in Nettles (Urticeae, Urticaceae), T. Deng, N. Lin, M. J. Moore, H. Sun, Huang, X., H. Wang, Z. Yusupov, Z. Li, Y. Wang, K. S. Tojibaev, 137: 190–199, , 2019

70. Pouzolzia rugulosa transferred from Boehmeria, and the distinction between Boehmeria and Pouzolzia (Urticaceae), Wilmot-Dear, C. M., T. I. Kravtsova, N. Acharya, I. Friis, 66: 51–64, , 2009

71. Component Content, ACE and AChE Inhibitory Activity of Ramie (Leaf Boehmeria nivea M.) according to Harvest Time, Lee, J. -K., Y. Choi, J. -K. Kim, H. -J. Cho, J. Lee, 31: 213–219, , 2018

72. Distribution of sexual and agamospermous populations of Boehmeria sylvestrii and its three relatives (Urticaceae), Yahara, T., 19: 121–132, , 1986

73. Genomic analyses provide comprehensive insights into the domestication of bast fiber crop ramie (Boehmeria nivea), H. Wang, Wang, Y., T. Zhang, L. Yan, F. Li, T. Liu, Z. Zeng, D. An, Z. Bao, S. Zhu, Q. He, 107: 787-800, , 2021

74. Cytological investigations of the microsporogenesis in male-sterile ramie (Boehmeria nivea Gaud.) and its offspring, R. M. Baroni, Pierozzi, N. I., R. Benatti, 73: 21–31, , 2008

75. Comparative plastid genomics of four Pilea (Urticaceae) species: insight into interspecific plastid genome diversity in Pilea, J. Yu, S. Zeng, Li, J., J. Tang, F. Han, J. Yuan, 21: 1–13, , 2021

76. Molecular phylogeny of the nettle family (Urticaceae) inferred from multiple loci of three genomes and extensive generic sampling, A. K. Monro, H. Wang, Wu, Z. Y., T. S. Yi, J. Liu, D. Z. Li, R. I. Milne, 69: 814–827, , 2013

77. Taxonomic studies of the Urticaceae 1. The genus Boehmeria in Thailand A recognition of Boehmeria nakashimae, sp. nov. (Urticaceae), Yahara, T., Yahara, T., 32: 1–21 58: 82–89. (in Japanese), , 1981

78. Leaf trichomes of Dendrobium species (epiphytic orchids) in relation to foliar water uptake, leaf surface wettability, and water balance, J. D. M. Schreel, W. Guo, Y. -P. Li, Pan, Z. -L., S. -J. Yang, J. -Y. Gao, Y. -J. Zhang, 190: 104568., , 2021

79. Note on the nomenclature and infraspecific classification of Boehmeria japonica (L. f.) Miq. and B. spicata (Thunb.) Thunb. (Urticaceae), Yahara, T., 59: 129–142, , 1984

80. The chloroplast tRNALys (UUU) gene from mustard (Sinapis alba) contains a class II intron potentially coding for a maturase-related polypeptide, Neuhaus, H., G. Link, 11: 251–257, , 1987

81. Paraphyly of the genus Boehmeria (Urticaceae): a response to Liang et al Relationships among Chinese Boehmeria species and the evolution of various clade, I. Friis, O. Maurin, C. M. Wilmot-Dear, S. Dodsworth, Monro, A. K., L. F. Fu, 307: 1–3, , 2021

82. Glandular trichomes as a barrier against atmospheric oxidative stress: relationships with ozone uptake, leaf damage, and emission of LOX products across a diverse set of species, P. C. Harley, Ü. Niinemets, M. Grosberg, T. Tosens, Li, S., K. Jaamets, A. Kanagendran, Y. Jiang, 41: 1263–1277, , 2018

83. Genetic Diversity among Wild Resources of the Genus Boehmeria Jacq. from West China Determined Using Inter-simple Sequence Repeat and Rapid Amplification of Polymorphic DNA Markers, B. Wang, Liu, L. J., Z. Q. Meng, J. Y. Yang, X. X. Wang, D. X. Peng, 12: 88–96, , 2009