Taxonomy of the Lichen Genus Usnea in Taiwan

本文研究台灣地區的松蘿屬地衣，依形態、化學、培養、分子分析探討松蘿的分類學與生物學。自 2006 到 2008 年於中央山脈、雪山山脈、玉山山脈、阿里山山脈、海岸山脈，超過 300 筆採集紀錄中，共區分出 22 個物種，分別為 Usnea aciculifera、U. baileyi、U. bismolliuscula、U. ceratina、U. cornuta、U. diffracta、U. fragilescens、U. glabrata、U. himalayana、U. longissima、U. malmei、U. masudana、U. nidifica、U. orientalis、U. pangiana、U. pectinata、U. pseudogatai、U. rubicunda、U. rubrotincta、U. shimadai、U. sinensis、U. trichodeoides。經由高效液相層析 (high performance liquid chromatography; HPLC) 分析，發現 40 種以上不同的地衣化學物質，瞭解松蘿的化學組成有助於鑑定與分類。另外，U. longissima 在本研究的所有物種中，松蘿酸含量最高 (3.31%)。培養 U. orientalis 以 20℃ 生長最佳 (1.14-1.87 mm /月)、在洋菜膠濃度 4% 的生長速率大於 2%、添加松蘿酸不影響 U. orientalis 的生長。分析松蘿核糖體的 ITS (internal transcribed spacer) 核酸序列，支持將 U. rubicunda 與 U. rubrotincta 劃分為兩個不同的物種；Usnea himalayana 與 U. nidifica 由於序列相似度高並發現形態中間型，可以歸為同一個物種；而 U. orientalis-U. pygmoidea 可能是採用有性生殖及無性生殖策略之地衣物種對 (species pairs)。松蘿最常附生於松科植物上 (22%)，其次為薔薇科 (15%) 與樺木科 (12%)。台灣地區松蘿的海拔分佈範圍界於 800 至 3400 m，其中 U. himalayana、U. longissima、U. trichodeoides 屬於高海拔物種，U. baileyi、U. bismolliuscula、U. pectinata、U. rubicunda、U. rubrotincta 屬於低海拔物種。Taxonomy and biology of the lichen genus Usnea in Taiwan were studied based on morphological, chemical, cultural traits, and molecular analyses. We studied over 300 specimens collected from the Central Mountain Range, Snow Mountain Range, Jade Mountain Range, Ali Mountain Range and Coastal Mountain Range from 2006 to 2008. Twenty-two species were recognized, including U. aciculifera, U. baileyi, U. bismolliuscula, U. ceratina, U. cornuta, U. diffracta, U. fragilescens, U. glabrata, U. himalayana, U. longissima, U. malmei, U. masudana, U. nidifica, U. orientalis, U. pangiana, U. pectinata, U. pseudogatai, U. rubicunda, U. rubrotincta, U. shimadai, U. sinensis, and U. trichodeoides. More than 40 lichen substances were found by high performance liquid chromatography (HPLC). The chemical constitutions supported the identification of Usnea species. It was found that U. longissima had the highest usnic acid contents (3.31%). In cultural studies, the best temperature for the growth of U. orientalis was 20℃ (1.14-1.87 mm / month), and it grew better on 4% agar than on 2% agar. Adding usnic acid to the cultural media did not affect the growth of U. orientalis. Molecular analyses based on sequences of ITS (internal transcribed spacer) of ribosomal RNA genes supported the separation of U. rubicunda and U. rubrotincta. Usnea himalayana and U. nidifica could be considered conspecific due to the fact that their intermediate forms exist among our collections and that their DNA sequences were highly similar. The results also showed that U. orientalis-U. pygmoidea probably represent species pairs of teleomorph-anamorphs connection. Usnea species in Taiwan frequently grow on plants of Pinaceae (22%), Rosaceae (15%) and Betulaceae (12%). Their altitudinal distribution was within the range 800-3400 m. Usnea himalayana, U. longissima, and U. trichodeoides are distributed at higher elevations, whereas U. baileyi, U. bismolliuscula, U. pectinata, U. rubicunda, and U. rubrotincta are at lower elevations.謝辭 I文摘要 II文摘要 III一章 緒論 1.1 地衣與松蘿 1.2 台灣松蘿之研究 2.3 松蘿之特徵與分類 3二章 材料與方法 6.1 標本取樣與資料管理 6.2 形態分類 7.3 化學分析 7.4 培養研究 10.5 ITS序列分析 11.6 生態學資料分析 15三章 結果 16.1 形態特徵 16.2 物種分類 21.3 地衣化學物質 81.4 培養結果 82.5 分子分析 82.6 生態學調查 83四章 討論 84五章 參考文獻 91六章 附錄 101.1 藥品與培養基 101.2 松蘿特徵描述記錄表 105.3 台灣地區松蘿物種列表 106.4 圖 107.5 表 160.6 松蘿在中華文化與台灣的故事 16