台灣產羊耳蒜屬植物之分類研究A Taxonomic Study of Liparis LC Rich

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台灣產羊耳蒜屬植物之分類研究A Taxonomic Study of Liparis LC Rich ⊕ 國立中山大學生物科學系 碩士論文 台灣產羊耳蒜屬植物之分類研究 A Taxonomic Study of Liparis L. C. Rich. (Orchidaceae) of Taiwan 研究生:楊智凱 指導教授:楊遠波 博士 中華民國九十五年六月 恩師楊遠波博士的教誨,嚴謹的治學態度及談吐中分享著多年研究經驗,在 研究上放心且信賴的任我盡情揮灑,生活上的關心與照顧,學生獲益匪淺。而遇 到瓶頸以及問題時老師總能一語道破,使論文進展更順利,在論文初稿撰寫期間 更不厭其煩多次費心的細心批改,讓其黑髮一夜髮白,心中萬分的感謝實在非三 言兩語可表達。任何事情保持著 `懷疑’的研究精神,更深深地烙印在學生心裡。 文稿初成,承蒙台灣大學森林環境暨資源學系蘇鴻傑教授在觀念與文獻上的 細心提點,成功大學生命科學系郭長生教授在百忙中撥空指教,論文之缺失不足 之處因兩位師長的指導而趨於完整,在此謹致由衷的謝意。 感謝英國皇家邱植物園標本館(K)Dr. Emma 協助拍攝大量之珍貴模式標 本;波蘭格但斯克大學植物分類與植物地理研究室 Dr. Margonska 提供寶貴文 獻;中央研究院植物標本館(HAST)彭鏡毅博士、林業試驗所標本館(TAIF) 邱文良博士、台灣大學植物標本館(TAI)郭城孟博士、台灣大學森林學系標本 館( NTUF)應紹順博士、台灣師範大學生物系植物標本館(TNU)王震哲博士、 自然科學博物館植物標本館(TNM)王秋美博士、屏東科技大學森林學系標本 館(PPI)楊勝任博士與特有生物研究保育中心植物標本館(TESRI)惠予借閱 標本並提供諸多協助以及林業試驗所森林保護組黃倩容學姐協助電子顯微鏡的 拍攝工作。 感謝林業試驗所鐘詩文博士諸多關照,慷慨提供日本東京大學植物標本館 (TI)的珍貴模式標本照片並熱心介紹日本筑波植物園蘭科專家 Dr. Yukawa , 其提供許多日本的寶貴文獻並告知研究成果,對我有很大助益,在此一併感謝。 研究得以順利進行都因為師長與好友們的幫忙。感謝系上昆蟲系統分類與演 化研究室顏聖紘博士,於研究初期幫忙影印大量的國外相關文獻並細心教導系統 分類學實務及支序地理學上的觀念。感謝研究室學長姐:怡姍、美珠、冠儀、東 哥、施兄、玉鳳、小旻、麗珍的指導與鼓勵,特別是小柯學長幫助我熟悉這個陌 生的環境;俊奎學長在生物繪圖上的教導;淑枝學姐於研究期間經費的安排與調 度;默詩學姐協助拍攝大量印尼茂物標本館(BO)標本;以誠學長協助拍攝中 國國家標本館(PE)模式標本;阿三哥在研究上及生活上的激勵以及關心以及 阿財學長在東部採集的細心照料。另外很感謝小胖幫忙鑑定昆蟲、怡玲、建炘等 諸位學弟妹幫忙協助實驗室計畫。 在植物分類及植物生態上的啟蒙特別感謝中興大學森林學系歐辰雄博士、呂 金誠博士、曾彥學博士,不僅在大學及研究所期間於學業與生活上的關心,歐老 師及呂老師更是在我遭遇困難時大力支持。生命科學系劉思謙博士,亦師亦友的 他在不僅在生活及研究上提供鼓勵與支持,創新的想法更給了我驚異激盪的空 間。也感謝森林學系森林植物分類暨生態研究室的學長姐,鴻志、朱哥、立彥、 志強、志銓、玄武、建志、政人、靖融、清安、俊凱、喜哥、芒果、國銘、傑鈞、 錦聰、家銘、秋瑩、詩婷、敏君的指導與鼓勵,讓我這段路走的更紮實,特別是 歐師母黃玲玉小姐,不管在生活上以及課業上都將我視如己出細心照顧,在此想 說一聲「謝謝你,歐師母。」 屏東科技大學森林學系謝光普同學從台中高農時期即互相砥礪與鼓勵,一起 堅持著我們走的路,沒有膽怯,這段同袍情誼將永遠留藏我心。 更感謝大學時期的摯友盈如、哀力、育揚、偉青、雅慧、碩容等的鼓勵和協 助;高中時期的摯友明益、老劉、原彰、小呈、麗芬、佳霖、以親等的陪伴。臺 灣師範大學生命科學系王震哲老師實驗室的學長姐,淑娟、和明、于芯、以及同 儕小量、惠如,讓我在北部研究期間時有個家,生活及交通上也有了協助與照料, 在此一併感謝,特別是小量。此外感謝在研究這段期間不時稍來關心與充當垃圾 桶的好朋友,東華大學自然資源管理所的吳郁娟同學,使我保持著動力。 台灣大學演化生態研究所許天銓,提供寶貴的研究材料,並熱切與我討論相 關的分類問題,雷達般的眼睛,更使我受益良多,再次感謝。 研究期間承蒙許多師長的指導和協助,首先感謝台中高農森林科詹益順老師 的鼓勵和提攜;屏科大生命科學系許秋容博士在解剖構造觀念上以及學業上提點 與鼓勵;恆春工商郭明裕老師,在屏東等地採集的全力支持與幫忙,才找到高士 佛羊耳蒜,恆春三寶洋蔥、西瓜、瓊麻;郭老師三寶檳榔、過山蝦、真性情;台 大實驗林鍾年鈞博士親自帶領前往清水溝以及溪頭採集,更在談話過程中,毫無 保留的提供許多蘭科訊息,獲益匪淺;感謝麗蓉姊及秀齡姊在野外生活上以及研 究上的細心照顧以及關心,如果沒有你們視野不會那麼開闊;呂碧鳳老師及余勝 焜老師,提供了許多有關羊耳蒜的寶貴訊息,於研究上及生活上更給了許多關心 和照料,更多次前往深山協尋研究材料,讓研究得以更順利進行,點滴在心頭。 感謝同儕崑琦、思穎、南瑛、仁義、惠如、菱春的鼓勵與支持,我們都熬過 來了,特別是崑琦,謝謝你在論文最後衝刺期間每天晚上相伴說笑回宿舍,並帶 著我吃遍美食,從鼓山區一直到鹽埕區;思穎在寒冬中總會找我跟崑琦吃火鍋, 大熱天喝啤酒;南瑛老是被我隨堂考等,謝謝你們,研究所的生活多采多姿。系 辦雅慧姊,初到中山時你的一番話,更讓我下定決心兩年畢業,謝謝妳。 高醫生物系阿昌、阿平、蝦米、見歡,你們的出現使寫論文的這段期間多了 許多歡樂,謝謝。 從事植物研究的開始,父母和姊姊們只知道我常往山裡跑,回家像是吃飯店 睡旅社,更不用說是陪伴家人了。電視總上演著台灣探險隊、大冒險家、MIT 台灣誌,「爸媽電視上演的太誇張了不要相信阿!」。最後要特別感謝我母親,這 一路走來會這麼堅持,是因為您做人處事的態度深深影響著我,讓我遇到更陡峻 的山、更險惡的天氣,咬著牙也要撐過去的,未來我會更堅定我的志向,只因有 著您的鼓勵,我會更放心的展開翅膀在天空中遨翔。 謹致於西子灣 2006 年仲夏 目次 附表目次-----------------------------------------------------------------------------------------II 附圖目次----------------------------------------------------------------------------------------III 中文摘要----------------------------------------------------------------------------------------IV 英文摘要-----------------------------------------------------------------------------------------V 壹、前言------------------------------------------------------------------------------------------1 貳、前人研究------------------------------------------------------------------------------------2 參、材料與方法-------------------------------------------------------------------------------12 肆、形態特徵----------------------------------------------------------------------------------16 伍、分類處理----------------------------------------------------------------------------------72 陸、討論---------------------------------------------------------------------------------------150 柒、參考文獻---------------------------------------------------------------------------------152 捌、附錄----------------------------------------------------------------------------------------159 I 附表目次 表 1. 羊耳蒜屬下的分類系統之一覽表--------------------------------------------------4 表 2. Jones & Clements(2005)將台灣產羊耳蒜屬新組合植物名錄對照表--------3 表 3. 歷年台灣產羊耳蒜屬植物新分類群以及新紀錄種一覽表--------------------6 表 4. 各學者對台灣產羊耳蒜屬植物各分類群採用學名表------------------------7-8 表 5. 台灣產羊耳蒜屬染色體已報導種類-----------------------------------------------9 表 6. 台灣產羊耳蒜屬植物鹼已報導種類----------------------------------------------11 表 7. 台灣產羊耳蒜屬植物分類群一覽表----------------------------------------------16 表 8. 台灣產羊耳蒜屬生活型、假球莖及葉部形態綜合特徵-----------------------18 表 9. 台灣產羊耳蒜屬植物葉部顯微形態研究材料及證據標本-------------------26 表 10. 電子顯微鏡下羊耳蒜屬植物葉表皮特徵----------------------------------------27 表 11. 台灣產羊耳蒜屬植物種子形態研究材料及證據標本-------------------------42 表 12. 台灣產羊耳蒜屬種子顏色以及相關數值參數表-------------------------------45 表 13. 台灣產羊耳蒜屬植物物候表-------------------------------------------------------51 表 14. 台灣地理氣候區所包含的羊耳蒜種類-------------------------------------------53 表 15. 台灣羊耳蒜屬植物亦產於鄰近地區之分類群類統計表----------------------58 表 16. 台灣羊耳蒜屬植物亦產於鄰近地區之比率-------------------------------------58 表 17. 台灣產沼蘭族形態特徵數據矩陣-------------------------------------------------69 表 18. 長穗羊耳蒜 (L. elongata) 、日本羊耳蒜 (L. japonica)與牧野氏羊耳蒜(L. makinoana)之特徵比較-------------------------------------------------------------87 表 19. 寶島羊耳蒜(L. formosana)與紅花羊耳蒜(L. nervosa)之特徵比較----------90 表 20. 齒唇羊耳蒜 (L. henryi)、大花羊耳蒜 (L. nigra) 與紅花羊耳蒜 (L. nervosa) 形態特徵比較------------------------------------------------------------------------95 表 21. 插天山羊耳蒜 (L. sootenzanensis)與大花羊耳蒜 (L. nigra)之特徵比較 109 表 22. 長腳羊耳蒜(L. condylobulbon)與淡綠羊耳蒜(L. viridiflora 之特徵比較-123 表 23. 長葉羊耳蒜(L. nakaharai)與川上氏羊耳蒜(L. kawakamii)之特徵比較--134 表 24. 叢生羊耳蒜(L. cespitosa)與小花羊耳蒜(L. laurisilcatica)形態特徵比較 ---------------------------------------------------------------------------------------------------137 II 附圖目次 圖 1. 台灣產羊耳蒜屬植物之生長形態-------------------------------------------------17 圖 2. 台灣產羊耳蒜屬植物葉之數目----------------------------------------------------19 圖 3. 台灣產羊耳蒜屬植物之葉形-------------------------------------------------------20 圖 4. 台灣產一葉羊耳蒜及小花羊耳蒜葉片長寬之關係圖-------------------------21 圖 5. 台灣產羊耳蒜屬葉之摺疊方式----------------------------------------------------22 圖 6. 台灣羊耳蒜屬植物類群間氣孔密度與氣孔長寬比之比較-------------------23 圖 7∼ 15. 台灣產羊耳蒜屬植物葉表皮細胞之電子顯微鏡(SEM)照片------28-36 圖 16. 台灣產羊耳蒜屬植物受訪昆蟲種類----------------------------------------------40 圖 17. 台灣產羊耳蒜屬果實形態----------------------------------------------------------41 圖 18. 台灣產羊耳蒜屬種子形式----------------------------------------------------------42 圖 19. 台灣產羊耳蒜屬種子長度及寬度之關係----------------------------------------46 圖 20. 台灣產羊耳蒜屬種子長度對應著種皮細胞的數目----------------------------46 圖 21∼ 24. 台灣產羊耳蒜屬種子之電子顯微鏡(SEM)照片---------------------47-50 圖 25. 台灣產羊耳蒜屬植物與台灣植群帶對應圖-------------------------------------57 圖 26∼ 45. 台灣產羊耳蒜屬植物之地理分布---------------------------------------58-68 圖 46. 形態特徵分析出 396 棵最簡約的嚴格共同樹----------------------------------70 圖 47. 形態特徵在嚴格共同樹上之分布圖----------------------------------------------71 圖. 48. Liparis auriculata Blume ex Miq. --------------------------------------------------77 圖. 49. Liparis cordifolia Hook. -------------------------------------------------------------83 圖. 50. Lipais formosana Reichb. f. ---------------------------------------------------------91 圖. 51. Lipairs henryi Rolfe ------------------------------------------------------------------96 圖. 52. Liparis krameri Franch. & Sav. var. sasakii (Hayata) T. Hashim---------------99 圖. 53. Liparis nervosa (Thunb.) Lindl. ---------------------------------------------------102 圖. 54. Liparis nigra Seidenf. ---------------------------------------------------------------106 圖. 55. Liparis sootenzanensis Fukuy. -----------------------------------------------------110 圖. 56. Liparis bootanensis Griff. ----------------------------------------------------------120 圖. 57. Liparis condylobulbon Reichb. f. ------------------------------------------------- 124 圖. 58. Liparis elliptica Wight --------------------------------------------------------------129 圖. 59. Liparis grossaReichb.f --------------------------------------------------------------132 圖. 60. Lipairs laurisilvatica Fukuy. -------------------------------------------------------138 圖. 61. Liparis nakaharai Hayata ----------------------------------------------------------144 圖. 62. Liparis viridifolia (Blume) Lindl. -------------------------------------------------149 III 摘要 本研究利用外部形態、生態及地理分布等資料,針對台灣產羊耳蒜屬(Liparis L. C. Rich.)植物進行分類研究。在外部形態方面,假球莖的有無、葉的數目、唇瓣及 蕊柱為區別台灣產羊耳蒜屬植物之重要的特徵。葉部顯微及種子顯微形態的特徵 可用在劃分亞屬及節的有用特徵,亦可作為種間分類的輔助特徵。地理分布方 面,楠櫧林帶內分布 11 種羊耳蒜,其中的高士佛羊耳蒜(Liparis somai)、 彎柱 羊耳蒜(L. campylostalix)、鬚唇羊耳蒜(L. waryi)、齒唇羊耳蒜(L. henryi),分布狹 窄。台灣產 20 種羊耳蒜中有 6 特有種 1 特有變種。台灣產沼蘭族(Malaxideae) 植物支序分析結果顯示,羊耳蒜屬植物並非單系群,而是一個並系群。根據上述 研究結果,將台灣產羊耳蒜屬植物處理為 22 個分類群,包含了一個變種及 2 個 存疑種,其中長穗羊耳蒜(L. japonica)之學名訂正為 L. elongata、小花羊耳蒜(L. cespitosa)之學名訂正為 L. laurisilvatica;原先併入長葉羊耳蒜(L. nakaharai)的川 上氏羊耳蒜(L. kawakamii) 確認其為獨立種;亦將原先併入長穗羊耳蒜(L. japonica)的德基羊耳蒜(L. derchiensis)確認為獨立種;而白花羊耳蒜(L. amabilis) 及明潭羊耳蒜(L. hensoaensis),因資料不足暫處理為存疑種。 IV ABSTRACT The genus Liparis in Taiwan was taxonomically revised based on morphological, phytogeographical and ecological evidences. Pseudobulb articulation, leaf number, lip and column are the most valuable characters for the classification within the genus in Taiwan. The morphology of leaf epidermis and seed coat also provide useful information in subgenus or section level. Geographically, eleven species are distributed in Machilus-Castanopsis zone, in which the species L. somai Hayata, L. campylostalix Rchb. f., L. wrayi Hook. f., and L. henryi Rolfe have strict distributional range. According to cladistic analysis, the result reveals that Liparis is a paraphyletic group rather than a monophyletic group. As a result of above studies, twenty-two taxa, including one variety and two uncertain species, are recongnized. L. japonica (Miq.) Maxim. and L. cespitosa (Thouars) Lindl. are synonyms of L. elongata Fukuy. and L. laurisilvatica Fukuy., respectively. L. kawakamii Hayata and L. derchiensis
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