臺灣博碩士論文加值系統

本論文以植物體外部形態、花粉形態、染色體核型及地理分布等資料為分類依據，針對台灣產細辛屬（Asarum L.）植物進行分類研究。研究結果顯示在外部形態上，綜合花柱形態、柱頭位置、萼片形態及萼筒內紋飾為區別台灣產本屬植物之有用特徵。植物周年生長形式及萼筒形態等特徵亦有助於分類群間的區別。在花粉形態上，台灣產種類中包括無孔型和4-6孔型兩種，花粉外壁紋飾有助於區分台灣各分類群。染色體方面顯示，台灣產本屬植物染色體除上花細辛（Asarum epigynum Hayata）為2n=12外，其餘種類皆為2n=24。其中，除A. epigynum核型為不對稱外，其餘分類群的核型對性極高且頗為相似，皆具2對大型染色體及1對小型染色體；台灣產分類群的染色體核型主要可分為三種類型，符合一般對署以下分類的結果。由地理分布可知，大部分的種類為局部性分布，其中分布海拔最低的種類集中在南北兩端恆濕性氣候區中，且以東北氣候區中具有最多的限域分布物種。
根據研究結果將台灣細辛屬植物處理為13個分類群，此13個分類群如下：白斑細辛 (A. albomaculatum Hayata)，罈花細辛 (A. ampulliflorum C. T. Lu)，薄葉細辛 (A. caudigerum Hance)，插天山細辛 (A. chatienshanianum C. T. Lu)，鴛鴦湖細辛 (A. crassusepalum S. F. Huang, T. H. Hsieh & T. C. Huang)，上花細辛 (A. epigynum Hayata)，下花細辛 (A. hypogynum Hayata)，大花細辛 (A. macranthum Hook)，太平山細辛 (A. taipingshanianum S. F. Huang, T. H. Hsieh & T. C. Huang)，大屯細辛 (A. taitonense Hayata)，太魯閣細辛 (A. tarokoense C. T. Lu)，大武細辛 (A. tawushanianum C. T. Lu)及神秘湖細辛 (A. villisepalum C. T. Lu)。其中除將被Huang et al. (1995)併入大花細辛 (A. macranthum Hook)的白斑細辛 (A. albomaculatum Hayata)及大屯細辛 (A. taitonense Hayata)加以分開；將裡紫細辛 (A. infrapurpureum Hayata)併入大屯細辛 (A. taitonense Hayata)。另外發現5個新種：罈花細辛 (A. ampulliflorum C. T. Lu)，插天山細辛 (A. chatienshanianum C. T. Lu)，(A. tarokoense C. T. Lu)，大武細辛 (A. tawushanianum C. T. Lu)及神秘湖細辛 (A. villisepalum C. T. Lu)。

The genus Asarum (Aristolochiaceae) in Taiwan was taxonomically revised based on morphological, palynological, cytotaxonomical and phytogeographical studies. Styles morphology, stigma position, sepal morphology, and ornamentation of the inner wall of the calyx-tube are shown to be valuable for the classification of Taiwan species. Annual growth module and calyx-tube shape also provide useful information. The pollen grains show variable on the micromorpholoy of the exine sculpture in some, but not all taxa. All taxa of Taiwan, except Asarum epigynum (2n=12), have the same chromosome numbers (2n=24). Karyotype of most Taiwanese taxa except A. epigynum, are with very similar symmetrical characterized by two pair large and one pair small ones. Geographically, most species in Taiwan are narrowly distributed in local range. Most of them are found in the northeast climate area.
As a result of above studies, thirteen taxa, including five species (A. ampulliflorum C. T. Lu, A. chatienshanianum C. T. Lu, A. tarokoense C. T. Lu and A. villisepalum C. T. Lu), are recognized. This treatment differs from the revised of Huang et al. (1995) in the following: A. albomaculatum Hayata and A. taitonense Hayata that were previously reduced to synonym of A. macranthum Hook, are now treated as distinct. A. infrapurpureum Hayata is now treated as the synonym of A. taitonense Hayata.

附表目次…………..II
附圖目次…………..III
中文摘要…………..V
英文摘要…………..VI
一、前言……………………………………………1
二、材料與方法……………………………………8
三、形態特徵……………………………………...12
四、花粉形態………………………………………41
五、細胞分類………………………………………52
六、地理分布………………………………………78
七、分類處理………………………………………84
八、參考文獻……………………………………..147
附表目次
表1. 台灣產細辛屬植物分類群學名沿革一覽表…………………………………….6
表2. 台灣產細辛屬植物分類群一覽表……………………………………….13
表3. 歷年研究者對細辛屬植物周年生長模組的研究結果比較……………………14
表4. 台灣產細辛屬植物周年生長模組之比較………………………………………17
表5. 台灣產細辛屬植物各分類群花部特徵之比較…………………………………22
表6. 台灣產細辛屬植物各分類群之萼裂片表面被毛種類比較……………………27
表7. 台灣產細辛屬植物種子形態研究材料及證據標本資料………………………39
表8. 台灣產細辛屬植物花粉形態研究材料及證據標本資料………………………42
表9. 台灣產細辛屬植物花粉形態之比較……………………………………………44
表10. 台灣產細辛屬植物染色體數目報導與證據標本資料………………………..53
表11. 台灣產細辛屬植物之核型……………………………………………………..64
表12. 白斑細辛 (A. albomaculatum) 之核型參數…………………………………..65
表13. 罈花細辛 (A. ampulliflorum) 之核型參數……………………………………66
表14. 薄葉細辛 (A. caudigerum) 之核型參數………………………………………67
表15. 插天山細辛 (A. chatienshanianum) 之核型參數…………………………….68
表16. 鴛鴦湖細辛 (A. crassusepalum) 之核型參數………………………………...69
表17. 上花細辛 (A. epigynum) 之核型參數………………………………………...70
表18. 下花細辛 (A. hypogynum) 之核型參數………………………………………71
表19. 大花細辛 (A. macranthum) 之核型參數……………………………………..72
表20. 太平山細辛 (A. taipingshanianum) 之核型參數……………………………..73
表21. 大屯細辛 (A. taitonense) 之核型參數………………………………………..74
表22. 太魯閣細辛 (A. tarokoense) 之核型參數…………………………………….75
表23. 大武細辛 (A. tawushanianum) 之核型參數………………………………….76
表24. 神秘湖細辛 (A. villisepalum) 之核型參數…………………………………...77
表25. A. caudigerum及A. epigynum的比較…………………………………………86
表26. A. hypogynum及A. tawushanianum之比較……………………………………87
表27. A. ampulliflorum及A. macranthum之比較…………………………………….88
表28. A. albomaculatum, A. crassusepalum, A. taipingshanianum及 A. tarokoense之比較…………………………………………………………………………….89
表29. A. chatienshanianum, A. taitonense及A. villisepalum之比較…………………90
附圖目次
圖1. 台灣產細辛屬植物之周年生長模組……………………………………………15
圖2. 台灣產細辛屬植物之生長形態…………………………………………………19
圖3. 台灣產細辛屬植物之葉子形態…………………………………………………20
圖4. 台灣產細辛屬植物之萼裂片形態………………………………………………24
圖5. 台灣產細辛屬植物萼裂片表面之毛絨形態……………………………………26
圖6. 台灣產細辛屬植物之萼筒形態…………………………………………………29
圖7. 台灣產細辛屬植物之萼筒內紋飾形態…………………………………………30
圖8. 台灣產細辛屬植物之退化花瓣 (假雄蕊) 形態……………………………….32
圖9. 台灣產細辛屬植物之雌蕊形態…………………………………………………34
圖10. 台灣產細辛屬植物種子表面之掃描式電子顯微鏡照片……………………..40
圖11. 台灣產細辛屬植物之花粉形態 (A. caudigerum; A. epigynum)………………47
圖12. 台灣產細辛屬植物之花粉形態 (A. albomaculatum; A. tarokoense;
A. villisepalum)…………………………………………………………………48
圖13. 台灣產細辛屬植物之花粉形態 (A. taitonense; A. macranthum)…………….49
圖14. 台灣產細辛屬植物之花粉形態 (A. ampulliflorum; A. tawushanianum;
A. hypogynum; A. chatienshanianum)………………………………………….50
圖15. 台灣產細辛屬植物之花粉形態 (A. crassusepalum; A. taipingshanianum)….51
圖16. 台灣產細辛屬植物之染色體 (A. caudigerum; A. epigynum; A. albomaculatum;
A. ampulliflorum; A. chatienshanianum; A. crassusepalum; A. hypogynum;
A. macranthum)………………………………………………………………...54
圖17. 台灣產細辛屬植物之染色體 (A. taipingshanianum; A. taitonense;
A. tarokoense; A. tawushanianum; A. villisepalum)……………………………55
圖18. 台灣產細辛屬植物之核型圖………………………………………………58, 59
圖19. 台灣產細辛屬植物之核型模式圖…………………………………………60-63
圖20. 台灣產細辛屬植物之地理分布 ( A. caudigerum; A. epigynum)……………..82
圖21. 台灣產細辛屬植物之地理分布 (A. hypogynum; A. tawushanianum)………..82
圖22. 台灣產細辛屬植物之地理分布 (A. ampulliflorum; A. macranthum)….……..83
圖23. 台灣產細辛屬植物之地理分布 (A. albomaculatum; A. chatienshanianum;
A. crassusepalum; A. taipingshanianum; A. taitonense; A. tarokoense; A. villisepalum)…………………………………………………………………....83
圖24. Asarum albomaculatum Hayata 白斑細辛……………………………………98
圖25. Asarum ampulliflorum C. T. Lu 罈花細辛…………………………………..102
圖26. Asarum caudigerum Hance 薄葉細辛……………………………………….107
圖27. Asarum chatienshanianum C. T. Lu 插天山細辛……………………………110
圖28. Asarum crassusepalum . S. F. Huang, T. H. Hsieh & T. C. Huang 鴛鴦湖細辛
………………………………………………..……………………………….113
圖29. Asarum epigynum Hayata 上花細辛…………………………………………117
圖30. Asarum hypogynum Hayata 下花細辛……………………………………….121
圖31. Asarum macranthum Hook. 大花細辛………………………………………126
圖32. Asarum taipingshanianum S. F. Huang, T. H. Hsieh & T. C. Huang 太平山細辛
……………………………………………………………………………...130
圖33. Asarum taitonense Hayata 大屯細辛………………………………………...135
圖34. Asarum tarokoense C. T. Lu 太魯閣細辛……………………………………138
圖35. Asarum tawushanianum C. T. Lu 大武細辛…………………………………142
圖36. Asarum villisepalum C. T. Lu 神秘湖細辛………………………………….146

吳照美、黃錦雲、周水發、蔡理里、那琦 1974. 韓產細辛及臺灣杜蘅類之生藥研究。 北醫學報 第六卷 pp. 1- 46 私立台北醫學院編印
林鴻文 2000 台灣產佛甲草屬植物之分類研究。 國立台灣師範大學生物學研究所碩士論文。
桑于雯 1995 台灣產沿階草族 (百合科) 之分類研究。 pp. 8-9, 40-67 國立台灣師範大學生物學研究所碩士論文。
陳又君 1998 台灣產鴨跖草科的分類研究。 國立台灣師範大學生物學研究所碩士論文。
黃增泉、謝長富、黃星凡、楊琇玉 1993 台灣大學植物系標本館日據時期之台灣植物標本採集地點目錄。 台灣省立博物館年刊 36：117- 174。
賴明洲 1973. 台灣大花草科、蛇菰科及馬兜鈴科之訂正。 國立台灣大學森林所碩士論文。
蘇鴻傑 1992. 台灣之植群：山地植群帶與地理氣候區。 中央研究院植物研究所專刊第11號 pp. 39- 53。
Araki, Y. 1937. The species of Asarum in the Santan District. Acta Phytotax. Geobot. 6: 122- 135.
Araki, Y. 1953. Systema generis Asari. Acta Phytotax. Geobot. 15: 33- 36.
Cheng, C. Y. and C. S. Yang. 1983. A synopsis of the Chinese species of Asarum (Aristolochiaceae). J. Arn. Arb. 64: 565- 597.
Cheng, C. Y. and C. S. Yang. 1988. Asarum. In H. S. Kiu & Y. R. Ling (eds.), Fl. Reip. Pop. Sinica 24: 161-196. Science Press.
Endo, J., T. Nakamura and M. Nagasawa. 1976. Geographical variation of Asarum caulescens Maxim. J. Jap. Bot. 51: 209- 219.
Erdtman, G. 1966. Pollen morphology and plant taxonomy: angiosperm. Hafner Pub. Co. pp. 16-19, 61-62.
Hatusima, S. 1968. Five new species of Asarum from southern Kyushu and Ryukyu. J. Jap. Bot. 43: 429- 434.
Hatusima, S and T. Amano. 1994. Flora of the Ryukyus, south of Amani Island. Second edition p. 41. The Biology Society of Okinawa.
Hayata, B. 1911. Materials for a Flora of Formosa. J. Coll. Sci. Univ. Tokyo 30 (1): 234.
Hayata, B. 1915. Asarum. In Iconum Plantanum Formosaranum. Ⅴ. pp.139-150. Bureau of forestry. Taihoku (Taipei). Taiwan.
Hayata, B. 1916. Suppl. Icon. Pl. Form. Ⅵ.: General Index Fl. Form. p. 61. Bureau of forestry. Taihoku (Taipei). Taiwan.
Henry, A. 1896. A list of plants from Formosa. Trans. Asiat. Soc. Japan Suppl. 24: 77.
Hooker, J. D. 1888. Asarum macranthum. Curtis’ Bot. Mag. t. 7022.
Hooker, J. D. 1890. Asarum caudigerum. Curtis’ Bot. Mag. t. 7126.
Hooker, J. D. 1891. Asarum geophilum. Curtis’ Bot. Mag. t. 7168.
Hsiao, S. C. and C. I. Peng. 1990. Cytotaxonomic notes on Asarum senus lato (Aristolochiaceae) in Taiwan. (unpublished)
Huang, S. F. 1996. Asarum. In Huang, T. C. et. al. (eds.), Fl. Taiwan 2nd ed. Vol. 2. pp. 642- 651.
Huang, S. F., T. H. Hsieh, and T. C. Huang. 1995. Notes on the Flora of Taiwan (21) --- The Genus Asarum L. (Aristolochiaceae). Taiwania 40 (2): 91- 120.
Huang, T. C. 1972. Asarum. Pollen Flora of Taiwan. p. 64. Pl.16: 20-22. Dept. of Bot., NTU. Taipei.
Huber, H. 1993. Aristolochiaceae. In K. Kubitzki, J. G. Rohwer and V. Bittrich (eds.), The families and genera of vascular plants. pp. 129-137. Berlin: Springer-Verlag.
Ikuse, M. 1956. Aristolochiaceae. In Pollen grains of Japan. pp. 64-65. Hirokawa Pub. Co. Tokyo.
Iwanaga, M. and Y. Momotani. 1976. Inter- and intrapopulational variations in the peroxidase isozyme of Asarum nipponicum F. Maekawa. Bot. Mag. 89: 93- 98.
Liu, T. S. and M. J. Lai. 1976. Aristolochiaceae. In Li, H. L., T. S. Liu, T. C. Huang, C. E. DeVol and T. Koyama (eds.), Flora of Taiwan. Vol. 2. pp. 571- 581. Epoch Pub. Co. Taipei.
Kawakami, T. 1910. Aristolochiaceae. In A list of plants of Formosa. p. 93.
Kelly, L. M. 1997. A cladistic analysis of Asarum (Aristolochiaceae) and implications for the evolution of herkogamy. Amer. J. Bot. 84 (12): 1752- 1765.
Kelly, L. M. 1998. Phylogenetic relationships in Asarum (Aristolochiaceae) based on morphology and ITS sequences. Amer. J. Bot. 85 (10): 1454- 1467.
Kelly, L. M. 2001. Taxonomy of Asarum section Asarum (Aristolochiaceae). Syst. Bot. 26 (1): 17-53.
Kudo, Y. and G. Masamune. 1932. Genera Plantarum Formosanarum. Ann. Rep. Bot. Gard. Taihoku Imp. Univ. 2: 46-47.
Ma, S. J. 1990. The geographical distribution and the system of Aristolochiaceae. Acta Phytotax. Sinica 28 (5): 345-355.
Maekawa, F. 1933. Japanese Asaraceae (Ⅰ)-(Ⅶ). J. Jap. Bot. 9: 39-49, 96-103, 174-179, 241-246, 281-285, 364-370, 505-512.
Maekawa, F. 1934. Japanese Asaraceae (Ⅷ). J. Jap. Bot. 10: 358-363.
Maekawa, F. 1936a. Japanese Asaraceae (Ⅸ). J. Jap.Bot. 12: 28-35.
Maekawa, F. 1936b. Asiasarum. In T. Nakai (eds.), Flora sylvatica Koreana, vol. 21. pp. 17-18. Keijyo: Government General of Chosen.
Maekawa, F. 1953. Geohistorical distribution of east Asiatic Asaraceae. Ⅶ Pacific Science Congress, Proceedings 5: 217- 219.
Maekawa, F. 1963. Reduction of chromosomes and major polyploidy: their bearing on plant evolution. J. Fac. Sci. Univ. Tokyo Ⅲ, 8: 377- 398.
Maekawa, F. 1977. Notes on Asarum epigynum Hayata. J. Jap. Bot. 52: 231- 232.
Maekawa, F. 1978. Notes on Asarum and Heterotropa from Taiwan. J. Jap. Bot. 53: 289- 299.
Maekawa, Y. 1988. A new species of Heterotropa (Aristolochiaceae) from Honshu, Japan. J. Jap. Bot. 63: 33-37.
Masamune, G. 1940. Miscellaneous notes on the Flora of the eastern Asia ⅩⅥ. Trans. Nat. Hist. Soc. Form. 30: 36.
Masamune, G. 1954. Aristolochiaceae. In A List of Vascular Plants of Taiwan. pp. 48- 49.
Matsumura, J. and B. Hayata. 1906. Enumeratio Plantarum Formosanarum. J. Coll. Sci. Imp. Univ. Tokyo. 22: 343.
Mi, Q. W. and C. S. Yang. 1991. Pollen morphology of Asarum in China. Acta Phytotax. Sinica 29 (2): 164- 171.
Morren, C. and J. Decaisne. 1834. Observations sur la Flore du Japon. Annales des Sciences Naturalles 2: 314-315.
Nakai, T. 1936. Japonasarum. In T. Nakai (ed.), Flora sylvatica Koreana, Vol. 21. Keijyo: Government General of Chosen. pp. 17-18.
Nakamura, T. and M. Nagasawa. 1987. Variation in pollen grains of Japanese Asiasarum (Aristolochiaceae). J. Jap. Bot. 62: 134- 138.
Nakamura, T., J. Endo and T. Hamada. 1982. Variation of Asiasarum dimidiatum. J. Jap. Bot. 57: 366- 375.
Ohwi, T. 1965. Flora of Japan. pp. 397- 401. Smithsonian Institution Press, Washington, D.C.
Ono, M. 1960. Studies on Heterotropa and its related genera with special reference to their karyo-morphology and phylogeny. J. Fac. Sci. Univ. Tokyo Ⅲ, 7: 473- 499.
Sasaki, S. 1928. List of plants of Formosa. pp. 166-167.
Sasaki, S. 1930. A catalogue of the Government Herbarium. Dep. For. Gov. Resear. Inst. pp. 188-189.
Shimabuku, K. 1997. Check list vascular flora of the Ryukyu Islands. Revised edition. pp. 194- 198. Kyusyu University.
Stace, C. A. 1989. Plant taxonomy and biosystematics. 2nd ed. Edward Arnold.
Su, H. J. 1984a. Studies on the climate and vegetation types of the natural forest in Taiwan (Ⅰ): Analysis of the variations in climate factors. Q. Jour. Chin. For. 17 (3): 1- 14.
Su, H. J. 1984b. Studies on the climate and vegetation types of the natural forest in Taiwan (Ⅱ): Altitudinal vegetation zones in relation to temperature gradient. Q. Jour. Chin. For. 17 (4): 57- 73.
Su, H. J. 1985. Studies on the climate and vegetation types of the natural forest in Taiwan (Ⅲ): A scheme of geographical climatic regions. Q. Jour. Chin. For. 18 (3): 33- 44.
Sugawara, T. 1981. Taxonomic studies of Asarum sensu lato Ⅰ. karyotype and c-banding pattern in Asarum s. str., Asiasrum and Heterotropa. Bot. Mag. 94: 225-238.
Sugawara, T. 1982. Taxonomic studies of Asarum sensu lato Ⅱ. karyotype and c-banding pattern in two species of Hexastylis and Asarum epigynum. Bot. Mag. 95: 295- 302.
Sugawara, T. 1985. A note on Heterotropa psudosavatieri and its allied species (Aristolochiaceae). J. Jap. Bot. 60: 202- 212.
Sugawara, T. 1987. Taxonomic studies of Asarum sensu lato Ⅲ. Comparative floral anatomy. Bot. Mag. 100: 335- 348.
Sugawara, T. 1992. A taxonomic study of Heterotropa nipponica and H. savatieri ssp. pseudosavatieri (Aristolochiaceae) in the Shima Peninsula. Acta Phytotax. Geobot. 43 (1): 15- 26.
Sugawara, T. 1996. A new species of Asarum (Aristolochiaceae) from Goto Island, Japan. Acta Phytotax. Geobot. 47 (2): 135- 141.
Sugawara, T. 1998. A taxonomic study of Asarum megacalyx F. Maek. and related taxa (Aristolochiaceae) distributed in Niigata prefecture and adjacent areas of Japan. Acta Phytotax. Geobot. 49 (1): 1- 17.
Sugawara, T. and M. Ogisu. 1986. Karyotype analysis of five species of Asarum (Aristolochiaceae) in Sichuan. J. Jap. Bot. 61: 104- 111.
Sugawara, T. and M. Ogisu. 1992. Karyomorphology of 11 species of Aasrum (Aristolochiaceae) from Taiwan and Mainland China. Acta Phytotax. Geobot. 43: 89- 96.
Sugawara, T., M. Ogisu and C. Y. Cheng. 1990. Asarum yunnanense, a new species of Asarum (Aristolochiaceae) in Shichuan, China. Acta Phytotax. Geobot. 41: 7-13.
Suzuki, T. 1936. Aristolochiaceae. In Masamune, Short Flora of Formosa. pp. 53- 54. Edit. Dep. Kudoa.
Walker, E. H. 1976. Aristolochiaceae. In: Flora of Okinawa and the southern Ryukyu islands. pp. 422- 423. Smithsonian Institution, Washington, D.C.
Walker, J. W. 1974a. Evolution of exine structure in the pollen of primitive angiosperms. Amer. J. Bot. 61 (8): 891- 902.
Walker, J. W. 1974b. Aperture evolution in the pollen of primitive angiosperms. Amer. J. Bot. 61 (10): 1112- 1136.
Ying, S. S. 1990. Miscellaneous note on the Flora of Taiwan ⅩⅢ. Mem. Coll. Agr., Natl. Taiwan Unvi. 30 (2): 53- 72.
Ying, S. S. 1991. Miscellaneous note on the Flora of Taiwan ⅩⅣ. Mem. Coll. Agr., Natl. Taiwan Unvi. 31 (1): 18- 39.
Yinger, B. R. 1983. A horticultural monograph of the genus Asarum senus lato, in Japan. Master’s thesis, University of Delaware. Newark, DE.