台灣產莎草科薹屬植物表面微細構造與分類之研究

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(1)行政院國家科學委員會專題研究計畫成果報告台灣產莎草科薹屬植物表面微細構造與分類之研究 Micromorphological studies and Taxonomy of Carex (Cyperaceae) in Taiwan 計畫編號：NSC-88-2311-B006-002-B27 執行期限：89 年 8 月 1 日至 90 年 7 月 31 日主持人：郭長生 E-mail: [email protected] 執行單位：國立成功大學生物學系一、中文摘要. delimitation of their taxonomic status especially at the Section level. Meanwhile, some dubious taxa or species complex such as C. filicina susp. pseudo-filicin, C. rafflesiana subsp. scaberrima , C. cruciata subsp. rubro-brunnea , C. perakensis and C. morii, C. bilateralis, C. sacrosancta , C. hattoriana , C. brunnea and C. gentilis, C. manca subsp. takasagoana , C. ligata subsp. formosensis, , C. breviculmis and subsp. fibrillosa , C. makinoensis, C. alopecuroides, C. alopecuroides subsp. subtransversa and C. liuii will be revised by field studies as well as principal components analysis and analyses of variance of specimen measurements.. 本計劃擬透過 SEM（掃描式電子顯微鏡）觀察，研究臺灣產薹屬(Carex)植物囊果、瘦果等表面微細構造，以探討屬內、種間、種內之變異作為分類及鑑定依據，特別是分節問題的釐清。此外尚有一些疑問種或復合種群，包括 C. filicina susp.. pseudo-filicin, C. rafflesiana subsp. scaberrima , C. cruciata subsp. rubro-brunnea , C. perakensis and C. morii, C. bilateralis, C. sacrosancta , C. hattoriana , C. brunnea and C. gentilis, C. manca subsp. takasagoana , C. ligata subsp. formosensis, , C. breviculmis and subsp. fibrillosa , C. makinoensis, C. alopecuroides, C. alopecuroides subsp. subtransversa and C. liuii 等，也一併經由田野調查、主群體分析及變異數分析處理以確認其分類地位。. Keywor ds: Cyperaceae,. Carex, Seed coat,. Utricle, SEM 二、緣由與目的薹屬是莎草科中最大的屬約有 2,000 種 (Nelmes, 1951) 。本屬為溫帶地區濕地生態系中最重要的屬，可提供野生動物棲所、糧草材料、水土保持及維護各種溼地功能 (Yen, 1999)。在台灣也是最大的屬(T. Koyama, 1978)，於中高海拔濕地及草原生態系中也有同樣重要地位。是研究台灣物種多樣性和生態多樣性時不可忽略的題材。但是這一龐大的屬，分類及鑑定較一般植物困難。薹屬植物小穗構造複雜但種間外形極相似，可供鑑定和探討親緣之特徵相對於其他植物而言就特別少。多數分類主要依據為花序形態和細胞學染色體特. 關鍵詞：莎草科、薹屬、囊果、瘦果、微細構造、掃描式電子顯微鏡。 Abstr act. The comparative micromorphological studies among species of Carex in Taiwan will be studied. The fine structures of utricle and seed coat will be studied with scanning electron microscopy, and will be documented for identification and classification. The variation of these characteristics within the genus, species and intraspecies will be presented for 1.

(2) 性，因所用特徵有限各家說法不ㄧ，是難度極高ㄧ群 (Yen, 1999)。近年由於生物多樣性議題受到重視，這ㄧ群經濟上看似不重要，但生態上極為重要的屬逐漸成為熱門題材，各類研究紛紛出籠︰嘗試應用新特徵如表面超微構造或利用分析方法如主群體分析重新評估舊有分類系統或應用分子生物方法探討親緣演化 (Yen, 1999)。本人目前負責台灣植物誌第二版莎草科之編寫，且有頗具潛力之研究生已投入薹屬研究。且有一些預實驗初步結果，故擬定三年計畫長期就台灣該屬植物進行系統分類研究。分別就表面超微構造之應用、主群體分析探討復合群分類、釐清各疑問種，並綜合建立整屬完整分類架構。. C. chinensis. Leong 771 Yang & Hsieh 5443. 宜蘭觀音瀑布. C. cryptostachys. 台北忠治. Leong 2185 T. Susuki 8367. 台北龜山至火燒樟. 三、材料與方法. C. hoozanensis. 屏東南仁山. C. hsinchuensis. 新竹清泉. Leong 2227. C. lateralis. 台北烏來. Leong 773. C. lanyuensis. 台東蘭嶼. Leong 1965. C. mancrandrolepis. 高雄南橫. Kuoh 14676. C. manca subsp. takasagoana. 台中武陵農場. H. E Chang 10847. C. mitrata. (日本). Koyama & Kao 8613. Makino s. n. (HAST 48047). 材料：借自台灣各標本館及野外採集所得之薹屬標本。目前觀察台灣產薹屬宿柱薹節植物瘦果表面特徵材料如表一。. C. orthostemon. 屏東姑子崙山. Leong 1414. C. pisiformis. 高雄南橫埡口. Leong 1896. C. rhynchachaenium var. rhynchachaeni um. 屏東浸水營. Leong 1439. 表一、台灣產薹屬宿柱薹節植物瘦果表面微細特徵研究材料及. 台東達仁林場. Leong 2092.1 Chen s. n. May 2001. 南投蕙蓀林場. 證據標本. Taxa. 台北大桶山. 地點. 標本編號. C. arisanensis. 台北大桶山. Leong 1163. C. wahuensis. (日本). Makino s. n. (HAST 52653). C. breviculmis var. breviculmis. 嘉義阿里山. Kuoh 11020. C. breviculmis var. fibrillosa. 台北石門. Kao &Chuang 3347. C. breviculmis var. morrisonicola. 南投合歡山. Leong 1368. C. breviscapa. 台北象山. Leong 1112. C. rhynchachaenium var. formosensis. 嘉義阿里山. Chen s. n. Sep. 2000. C. rhynchachaenium var. kelongensis. 南投草屯. Kuoh s. n. Mar. 2001. C. transalpine. 台北大桶山. Leong 1162 Leong 2236. 台北拔刀爾山. Leong 1536. 台北南港. 2. C. tristachya var. pocilliformis. 新竹蓮花寺. Kuoh 16401. C. truncatigluma. 宜蘭福山植物園. Peng 13857.

(3) Carex sp.. 高雄南橫埡口. 所有細胞均不具矽質體的種類有 C.. Leong 2203.1 Leong 2246. breviscapa 、C. orthostemon、C. tristachya. 宜蘭思源埡口. var. pocilliformis 和 C. truncatigluma 4 種 (表二、圖 1e, f, 90a, c, d)，而 C. hoozanensis 和 C. manca subsp. takasagoana 僅約 1/10. 研究方法：挑選標本館中具有成熟果實之標本，記錄標籤上的資料，並取下所需果實。由於欲觀察之瘦果表面矽質體被細胞表層覆蓋，所得之乾燥或新鮮果實，須參考 Schuyler (1971)之方法稍微修改，將瘦果置於九份冰醋酸加一份濃硫酸製成之混合液中，浸泡十二至二十四小時後，以超音波震盪器震盪至細胞表層脫落，露出矽質體為止，再用 70%酒精沖洗，風乾四十八小時後貼上鋁台，以電子顯微鏡觀察、照相，比較個體內、族群內、族群間及各層級分類群間的差異。. 的細胞具有矽質體 (表二、圖 4)，另外 C.. hsinchuensis 約有 1/5 的細胞不具矽質體 (圖 3d)；具中央矽質體的種類又可依照矽質體頂端形狀區分，大部分種類頂端圓鈍，其中 C. lateralis 的矽質體巨大，幾乎佔據整個細胞表面 (表二、圖 5a-88c)；頂端明顯尖銳的種類有 C. arisanensis 及 C.. pisiformis， (表二、圖 1a, b, 90b)有時 C. rhynchachaenium 也出現尖銳頂端 ( 圖 6a)；部分種類的矽質體頂端具有不同程度. 四、結果與討論. 突起的附屬物，包括 C. brevuculmis var. 依瘦果表面矽化細胞大小，可將宿柱薹節. morrisonicola 、 C.. 植物分為兩群：細胞較大的直徑通常在. rhynchachaenium 和 C. transalpina 均有發. 40-60 ìm 之間，包括 C. arisanensis、C.. 現此現象 (表二、圖 2e, 89b-89c)，有時甚. wahuensis、C. breviscapa、C. hoozanensis、. 至形成分枝狀的長刺 (表二、圖 2c, d, 89d,. C. manca subsp. takasagoana 、C. lateralis. 90e)。. 和 C. macrandrolepis 7 種 (圖 1, 87, 88)，. chinensis 、 C.. (2) 衛星矽質體 (satellite). 其中 C. hoozanensis 的細胞最大，一般直徑僅出現在少數種類中，並可分成以下. 50-60 ìm，最大可達 70 ìm (表二、圖 4a, d,. 性狀：. f)；其他細胞較小的種類直徑一般在 15-30 ìm 之間 (圖 2, 86, 89, 90)。不同種類的矽. 瘤狀 (verrucose)：整個細胞表面散佈. 質細胞可明顯區別 (表二、圖 1-90)，其分. 瘤狀突起；可見於 C. breviculmis var.. 類特徵共有以下幾點：. breviculmis 和 C. breviculmis var. fibrillosa (表二、圖 2a, b,. (1) 中央矽質體 (central silica body). 3. )，有時也出現於 C..

(4) hsinchuensis、C. lanyuensis、C. transalpina. transalpina (表二、圖 7e)。. 和 C. truncatigluma 4 種 (表二、圖 3d, e,. 圓鈍：C. macrandrolepis 僅分布於邊. 90d, f)。. 緣 (peripheral)，較大 (表二、圖 5d-f)；. 五、參考文獻. C. cryptostachys 分布於邊緣與中間. Akiyama, S. 1955. Carices of the Far Eastern region of Asia. 1-257. Barthlott, W. 1981 Nord. J. Bot. 1: 345-355 Epidermal and seed surface characters of plants: systematic applicability and some evolutionary aspects. Brisson, J. D. and R. L. Peterson 1977 Ill. Inst. Techn. Res. Inst./SEM/1976/II: 697-712 The scanning electron microscope and X-ray microanalysis in the study of seeds: A bibliography of the period 1967-1976 Bruederle, L. P., D. E. Fairbrothers and s. L. Hanks 1989 Amer. J. Bot. 76: 124-132 A systematic circumscription of Carex mitchelliana (Cyperaceae) with reference to taxonomic status Canne, J. M. 1979 Syst. Bot. 4: 281-296 A light and scanning electron microscope study of seed morphology in Agalinis (Scrophulariaceae) and its taxonomic significance. Canne, J. M. 1980 Syst. Bot. 5: 241-252 Seed surface feature in Aureolaria, Brachystigma, Tomanthera, and certain South American Agalinis (Scrophulariaceae). Chance, G. D. and J. D. Bacon. 1984 Amer. J. Bot. 71: 829-842 Systematic implications of seed coat morphology in Nama (Hydrophyllaceae) Crins, W. J. and P. W. Ball 1988 Brittonia 40: 38-47 Sectional limits and phylogenetic considerations in Carex section Ceratocystis (Cyperaceae) Denton, M. L. 1983Syst. Bot. 8: 250-262 Echlin, P. 1968 J. Roy. Microscop. Soc. (London) 88: 407-418 The use of the scanning reflection electron microscope in the study of plant and microbial material Hayata, B. 1916. Icon. Pl. Form. 6: 122.. (peripheral & intermediate)，有時和中央矽質體相似(表二、圖 3a-c)。密佈分枝狀長刺(branching spine)：僅出現在 C. breviculmis var. morrisonicola ，與其矽質體頂端之長刺相似(表二、圖 2c, d)。細胞表面平臺 (platform)：表面下陷 (concave) 或隆起 (convex)的現象皆普遍 (表二)，C. lateralis 由於巨大的圓形中央矽質體佔據整個表面，使平臺顯得強烈上凸 (strongly convex) (圖 5a-c)。平臺邊緣通常筆直並緊鄰 (appressed) 周圍平臺，或邊緣彎曲並和周圍平臺保持一定距離 (nonappressed)，唯 C. lanyuensis 的邊緣筆直，但垂周壁呈蜂巢狀 (honeycombed)，因而造成平臺間的距離 (表二、圖 94-90)；又 C. orthoetemon 的平臺邊緣彎曲，但平臺間仍緊鄰 (表二、圖 7a)。平臺轉角通常圓滑 (round) ，明顯具角 (angular) 的種類有 C. arisanensis 、 C.. wahuensis 以及 C. hoozanensis (表二、圖 1a-84d, 87a, d, f)。垂周壁：筆直和波浪狀皆普遍 (表二 ) ，而呈蜂巢狀的種類計有 C. 4.

(5) Heywood, V. H. 1969 Micron 1: 1-14 Scanning electron microscopy in the study of plant materials Hoshino, T. 1984. Scanning electron microscopic observation of the surface pattern of achene in Carex. Bull. Hiruzen Res. Inst., Okayama Univ. Sci. 10: 59-71 Kern, J. H. and H.P. Nooteboom. 1979. Cyperaceae. II. Flora Malesianan Series I, Vol. 9: 107-187. Koyama, T. 1978. Cyperaceae In: Li, H. L. et al. Flora of Taiwan. 5: 191-372. Kukenthal, G. 1909. Cyperaceae-Caricoideae. Pp. 1-824. in Das Pflanzenreich. IV. Vol. 20. heft 38, Editor A. Engler. Leipzig: Wilhelm Engelmann. Menapace, F. J. and D. E. Wujek. 1987 Brittonia 39: 278-283 The systematic significance of achene micromorphology in Carex retrorsa (Cyperaceae) Menapace, F. J. and D. E. Wujek. 1991 Can. J. Bot. 69: 1533-1541 A preliminary micro-morphological analysis of Eleocharis (Cyperaceae) achenes for systematic potential Naczi, R. F. C., A. A. Reznicek, and B. A. Ford 1998. Morphological, geographical, and ecological differentiation in the Carex willdenowii complex (Cyperaceae). Amer. J. Bot. 85(3): 434-447. Nelmes, E. 1951. The genus Carex in Maleysia. Reinwardtia 1: 221-450. Ohwi, J. 1936. Mem. Coll. Sci. Kyoto Univ. B, 11: 461. Rettig, J. H. 1989 Sida 13: 449-452 Nomenclatural changes in the Carex pensylvanica group (section Acrocystis, Cyperaceae) Rothrock, P. E. 1991. The identity of Carex albolutescens, C. festucaea, and C. longii (Cyperaceae). Rhodora, 93(873): 51-66. Schuyler, A. E. 1971 Proc. Acad. Nat.. Sci. Philadelphia 123: 29-52 Scanning electron microscopy of achene epidermis in species of Scirpus (Cyperaceae) and related genera Standley, L. A. 1986 Amer. J. Bot. 73: 787 Anatomical studies of the Carex lenticularis complex in the New World Timonen, T. and H. Toivonen 1979. Gross and micromorphological comparison of Carex furva and C. lachenalii Ann. Bot. Fenn. 16:11-17 Toivonen, H. 1980. Carex canescens x mackenziei. A comparative study of two Carex species and their spontaneous hybrid. Ann. Bot. Fenn. 17: 91-123 Toivonen, H. and T. Timonen 1976. Perigynium and achene epidermis in some species of Carex subgenus Vignea (Cyperaceae) studied by SEM. Ann. Bot. Fenn. 13: 49-59 Varma, S. K., A. K. Pandey and A. K. Sinha 1989 Curr. Sci. 58: 1374-1377 Epidermal surface patterns of achene in Eleocharis R. Br. (Cyperaceae) Walter, K. S. 1975. A preliminary study of the achene epidermis of certain Carex (Cyperaceae) using SEM. Michigan Bot. 14: 67-72 Yen A. C.-T. 1999. Molecular Systematics of Cyperaceae Tribe Cariceae and Genus Carex. Doctor dissertation, University of Washington. 六、計畫成果自評研究內容與原計畫大致相符、初步達成預期目標之資料部分、大量資料尚待分析，研究成果可供莎草學分類與發育之學術上參考價值、已準被在學術期刊上發表。. 表二、台灣產薹屬宿柱薹節植物瘦果表面微細特徵 5.

(6) Taxa. Centr al silicon body. Satellite. Sur face of Mar gins of platfor m platfor m. Cor ner s of platfor m Anticlinal wall. C. arisanensis. large, acute at apex. absent. concave. straight and appressed. angular. straight. C. breviculmis var. breviculmis. large, round at apex. verrucose. convex. curved, nonappressed. round. sinuous. C. breviculmis var. fibrillosa. round. verrucose. concave. curved, nonappressed. round. sinuous. C. breviculmis var. morrisonicola. a large head with branching spines. dense, branching convex spines. curved, nonappressed. round. sinuous. C. breviscapa. absent. absent. straight and appressed. round. straight. C. chinensis. round or a head with absent a few irregular spines. concave to curved, convex nonappressed. round. sinuous. C. cryptostachys. round at apex. convex, irregular. curved, nonappressed. round. sinuous. C. hoozanensis. round, usually absent absent. concave. straight, appressed. angular. traight. C. hsinchuensis. round at apex, sometimes absent. convex. curved, nonappressed. round. sinuous and honeycombed. C. lateralis. very large, occupying absent the whole platform. strongly convex. straight, appressed. round. straight. C. lanyuensis. round at apex. absent. convex. straight, but nonappressed. round. straight, honeycombed. C. macrandrolepis. large, round at apex. peripheral, large, concave round at apex. straight, appressed. round. straight. C. manca subsp. takasagoana. small, round at apex, absent usually absent. straight, appressed. round. straight. C. mitrata. round at apex, usually intermediate, convex absent, size variable round, very few. curved, nonappressed. round. slightly sinuous. C. orthostemon. absent. absent. concave. curved, but appressed. round. curved but not sinuous. C. pisiformis. acute at apex. absent. convex. curved, nonappressed. round. curevd, seriously sinuous. C. rhynchachaenium var. rhynchachaenium. acute to a large head absent with branching spines. concave to straight, convex appressed. round to angular. straight to slightly sinuous. C. rhynchachaenium var. formosensis. a large head with branching spines. absent. convex. straight, appressed. round. straight. C. rhynchachaenium var. kelongensis. acute to a small verrucose head. absent. slightly concave. straight, appressed. round. slightly sinuous. C. transalpine. round or a large head absent or with branching spines verrucose. convex. curved, nonappressed. round. sinuous and honeycombed. C. tristachya var. pocilliformis. absent. absent. convex. straight, appressed. round. straight. C. truncatigluma. absent. absent. slightly concave. curved, nonappressed. round. sinuous and honeycombed. C. wahuensis. round at apex. absent. concave. straight and appressed. angular. straight. convex. peripheral and intermediate, round. absent. concave. 6.

(7) 圖 1. 台灣產薹屬宿柱薹節植物瘦果表面特徵示意圖. 63.

(8) 圖 2. 台灣產薹屬宿柱薹節植物瘦果表面之電子顯微鏡照片 a, b. C. arisanensis; c, d. C. wahuensis; e. f. C. breviscapa; bar = 10 ìm.. 64.

(9) 圖 3. 台灣產薹屬宿柱薹節植物瘦果表面之電子顯微鏡照片 a. C. breviculmis var. breviculmis; b. C. breviculmis var. fibrillosa ; c, d. C. breviculmis var. morrisonicola ; e. C. chinensis (Leong 771); f. C. chinensis (Yang & Hsieh 5443); bar = 10 ìm.. 65.

(10) 圖 4. 台灣產薹屬宿柱薹節植物瘦果表面之電子顯微鏡照片 a, b. C. cryptostachys (Leong 2185); c. C. cryptostachys (T. Susuki 8367); d. C. hsinchuensis; e. C. lanyuensis; f. C. mitrata ; bar = 10 ìm.. 66.

(11) 圖 5. 台灣產薹屬宿柱薹節植物瘦果表面之電子顯微鏡照片 a, c, e. C. manca subsp. takasagoana ; b, d, f. C. hoozanensis; bar = 10 ìm.. 67.

(12) 圖 6. 台灣產薹屬宿柱薹節植物瘦果表面之電子顯微鏡照片 a, b, c. C. lateralis; d, e, f. C. macrandrolepis; bar = 10 ìm.. 68.

(13) 圖 7. 台灣產薹屬宿柱薹節植物瘦果表面之電子顯微鏡照片 a. C.rhynchachaenium var. rhynchachaenium (Leong 2092.1); b. C.rhynchachaenium var. rhynchachaenium (Leong 1439); c. C.rhynchachaenium var. rhynchachaenium (Chen s. n. May 2001); d. C.rhynchachaenium var. formosensis; e, f. C. rhynchachaenium var. kelungensis; bar = 10 ìm.. 69.

(14) 圖 8. 台灣產薹屬宿柱薹節植物瘦果表面之電子顯微鏡照片 a. C. orthostemon; b. C. pisiformis; c. C. tristachya var. pocilliformis; d. C. truncatigluma ; e. C. transalpina (Leong 2236); f. C. transalpina (Leong 1162); bar = 10 ìm.. 70.

(15)