臺灣博碩士論文加值系統

花的形態常與傳粉媒介共演化而呈現多樣變化，不同的花部形態會吸引不同的傳粉者而造成物種間的生殖隔離，因此在植物的繁衍上扮演著重要角色。樟科植物是構成臺灣常綠闊葉林的重要組成樹種，以往在臺灣樟科植物雖已有多位學者進行過系統分類研究，但對於花部形態的基礎研究卻仍不完整，甚至有形態描述不一致之情形。本研究針對臺灣產樟科植物選擇具代表性之類群進行花部形態及開花週期之探討，以釐清臺灣產樟科植物屬間及屬內各類群花部形態特徵與開花週期的異同，研究結果將可供授粉生物學及保育生物學等科學研究的背景。本研究觀察臺灣產之樟屬(Cinnamomum)的樟樹(Ci. camphora)、土肉桂(Ci. osmophloeum)，厚殼桂屬(Cryptocarya)的厚殼桂(Cr. chinensis)，釣樟屬(Lindera)的大葉釣樟(Lin. megaphylla)，木薑子屬(Litsea)的山胡椒(Lit. cubeba)，楨楠屬(Machilus)的大葉楠(M. japonica var. kusanoi)、紅楠(M. thunbergii)、香楠(M. zuihoensis)及雅楠屬(Phoebe)的臺灣雅楠(P. formosana)，除觀察野外個體的花部物候現象外，同時採集樣本攜回實驗室，每隔2小時觀察記錄一次開花狀態，並以相機進行縮時攝影協助記錄。
研究結果發現，單性花類群(大葉釣樟及山胡椒)之花朵開放後維持1至5日不等，並無夜間閉合之兩次開花特性；而兩性花各類群(樟樹、土肉桂、厚殼桂、大葉楠、紅楠、香楠及臺灣雅楠等)均為雌雄異熟(dichogamy)，且為雌先熟(protogynous)。大多數具晚間閉合之兩次開花特性，記錄到之開花週期共有五種類型：(A)首日上午開放而雌蕊熟，午後閉合，次日下午再度開放而雄蕊熟，晚間閉合；(B)首日下午開放而雌蕊熟，晚間閉合，次日上午再度開放而雄蕊熟，午後閉合；(C)首日上午開放而雌蕊熟，晚間閉合，次日上午再度開放而雄蕊熟，晚間閉合；(D)首日晚間開放而雌蕊熟，夜間閉合，次日上午再度開放而雄蕊熟，午後閉合；(E)不具兩次開花特性，花開時雄蕊已熟；其中類型A及B與前人研究結果相似；此外，前人研究中指出同一植株之花均屬同一類型且開放時間一致性高，但本研究觀察到部分個體同時具有兩種類型的花，單一植株不同花朵開放時間並非完全一致。

Floral morphologies are diversified because of the coevolution with pollinators. Different morphologies can attract different pollinators and lead to reproductive isolations. As a result, flowers are the most important feature of plant reproduction. Lauraceae is an important component of evergreen broad-leaved forest in Taiwan, however most studies of Lauraceae in Taiwan focus on systematics, few studies paying attention to floral biology. Therefore the description of floral morphologies and floral phenology for some taxa are incomplete or inconsistent. In this study, my aims are to identify and compare the floral morphologies and floral phenology between genera and between taxa of Lauraceae in Taiwan. The results of this study can provide basic information for pollination biology and conservation biology. We choose Cinnamomum camphora, Ci. osmophloeum, Cryptocarya chinensis, Lindera megaphylla, Litsea cubeba, Machilus japonica var. kusanoi, M. thunbergii, M. zuihoensis, and Phoebe formosana for observation. In addition to field observation of floral phenology, we cut the branches with floral buds and inserted them into water bottles in laboratory for observations. Checking the floral condition once every two hours, and recording with time-lapse photography. In my observations, unisexual flowers (Lindera megaphylla, and Litsea cubeba) maintain opening for 1 to 5 days, and these flowers neither open twice nor close in the evening. Bisexual flowers (Cinnamomum camphora, Ci. osmophloeum, Cryptocarya chinensis, Machilus japonica var. kusanoi, M. thunbergii, M. zuihoensis, and Phoebe formosana) are dichogamy and protogynous, and most of them open twice and close in the evening. The floral phenology of bisexual flowers can be categorized into the following types: (a) first day flowers open as female in the morning and close in the afternoon until next afternoon open again as male, then close again in the evening; (b) first day flowers open as female in the afternoon and close in the evening until next morning open again as male, then close again in the afternoon; (c) first day flowers open as female in the morning and close in the afternoon until next morning open again as male, then close again in the afternoon; (d) first day opening as female in the evening and close at night until next morning open again as male, then close again in the afternoon; (e) without dianthesis, flowers open as male. In addition, the flowers of the same branch we observed don’t entirely show the phenomenon of synchronous dichogamy, and some of them had two types of the floral phenology.

目錄
摘要-------------------------------------------------------2
一、研究背景及目的------------------------------------------6
二、材料與方法---------------------------------------------17
三、結果
(一)花部形態----------------------------------------------21
(二)開花週期----------------------------------------------59 (三)蜜腺分泌時間-------------------------------------------77 (四)樟科植物之開花現象-------------------------------------89 四、討論
(一)花部形態----------------------------------------------90 (二)開花週期----------------------------------------------95 (三)蜜腺分泌時間------------------------------------------104 (四)樟科植物的花部特徵演化與生殖模式探討--------------------106
(五)未來發展方向------------------------------------------112 五、參考文獻---------------------------------------------113

呂勝由。2008。台灣樟科楨楠屬植物系統分類之研究。國立中山大學生物科學系博士論文。
呂勝由、陳舜英。1996。香楠及霧社楨楠地理分布與分類之研究。台灣林業科學11(3): 239-244。
呂福原。1969。臺灣產楨楠屬(樟科)之訂正。中華林學季刊2(3): 12-36。
呂福原、歐辰雄、陳運造、祁豫生、呂金誠。2000。臺灣樹木圖誌第一卷。臺中：國立中興大學森林學系森林植物與生態研究室。
李學勇。1984。台灣經濟植物生殖機制之研究-三、楨楠屬植物之生殖機制。林業特刊 7: 20-30。
范義彬。2006a。台灣樟科植物開花授粉生物學之研究。國立臺灣大學昆蟲學研究所博士論文。
范義彬。2006b。樟樹與山胡椒開花授粉生物學之研究。國立臺灣博物館學刊 59(1): 75-90。
張慶恩。2005。台灣樟科植物之研究綜述。中華林學季刊 38(4): 133-145。
陳國章、張乃航。1999。台灣樟科新木薑屬植物之訂正。台灣林業科學14(2): 141-163。
陳國章、潘富俊。1997。臺灣樟科新木薑子屬植物之新見。台灣林業科學12(2): 155-166。
廖日京。1987。臺灣樟科植物學名綜談(II)。中華林學季刊 20(3): 73-77。
劉棠瑞。1960。台灣木本植物圖誌(上)。臺北：國立臺灣大學農學院。
劉業經、呂福原、歐辰雄。1988。臺灣樹木誌。臺中：國立中興大學農學院。
應紹舜。1985。台灣高等植物彩色圖誌第一卷。新店：淑馨出版社。
應紹舜。1985。臺灣產樟科植物之訂正。國立臺灣大學農學院研究報告 25: 83-117。
蔡永信。2011。臺灣檫樹開花行為與微衛星體基因座之特性分析。國立臺灣大學森林環境暨資源學研究所碩士論文。
Arpaia, M. L., A. E. Fetscher., and R. Hofshi. 2001. Avocado flowering basics. http://ucavo.ucr.edu/Flowering/FloweringBasics.html
Balthazar, M., and P. K. Endress. 1998. Floral bract function, flowering process and breeding systems of Sarcandra and Chloranthus (Chloranthaceae). Plant Systematics and Evolution 218(3-4): 161-178.
Bessey, C. E. 1915. The phylogenetic taxonomy of flowering plants. Annals of the Missouri Botanical Garden 2(1/2): 109-164.
Bolin, J. F., E. Maass, and L. J. Musselman. 2009. Pollination biology of Hydnora africana Thunb. (Hydnoraceae) in Namibia: brood-site mimicry with insect imprisonment. International Journal of Plant Sciences 170(2): 157-163.
Burgess, K. S., J. Singfield, V. Melendez, and P. G. Kevan. 2004. Pollination biology of Aristolochia grandiflora (Aristochiaceae) in Veracruz, Mexico. Annals of the Missouri Botanical Garden 91(2): 346-356.
Chanderbali, A. S., H. Werff, and S. S. Renner. 2001. Phylogeny and historical biogeography of Lauraceae: Evidence from the chloroplast and nuclear genomes. Annals of the Missouri Botanical Garden 88(1): 104-134.
Chang, C.-E. 1976. Lauraceae. In: H.-L. Li et al. [eds.], Flora of Taiwan, 1st edition, vol. 2., 406-468. Epoch Publishing, Taipei, Taiwan, ROC.
Chaw, S.-M. 1992. Pollination, breeding syndromes, and systematics of Trochodendron aralioides Sieb. &; Zucc. (Trochodendraceae), a relictual species in Eastern Asia. In Phytogeography and Botanical Inventory of Taiwan (Peng, C.-I., ed.), pp. 63-77, Institute of Botany, Academia Sinica Monograph Series 12.
Chiou, C.-R., C.-F. Hsieh, J.-C. Wang, M.-Y. Chen, H.-Y. Liu, C.-L. Yeh, S.-Z. Yang, T.-Y. Chen, Y.-J. Hsia, G.-Z. M. Song. 2009. The first national vegetation inventory in Taiwan. Taiwan Journal of Forest Science 24(4): 295-302.
Chung, K.-F., H. Werff, and C.-I. Peng. 2010. Observations on the floral morphology of Sassafras randaiense (Lauraceae). Annals of the Missouri Botanical Garden 97(1): 1-10.
Davenport, T. L. 1986. Avocado flowering. Horticultural Reviews 8: 257-289.
Dommée, B., J.-L. Bompar, and N. Denelle. 1990. Sexual tetramorphism in Thymelaea hirsute (Thymelaeaceae): Evidence of the pathway from heterodichogamy to dioecy at the infraspecific level. American Journal of Botany 77(11): 1449-1462.
Dommée, B., A. Biascamano, N. Denelle, J.-L. Bompar, and J. D. Thompson. 1995. Sexual tetramorphism in Thymelaea hirsute (Thymelaeaceae): Morph ratios in open-pollinated progeny. American Journal of Botany 82(6): 734-740.
Endress, P. K. 1984. The flowering process in the Eupomatiaceae (Magnoliales). Botanische Jahrbücher für Systematik 104: 297–319.
Endress, P. K. 2010. The evolution of floral biology in basal angiosperms. Philosophical Transactions of the Royal Society B: Biological Sciences 365(1539): 411-421.
Endress, P. K., and D. H. Lorence. 2004. Heterodichogamy of a novel type in Hernandia (Hernandiaceae) and its structural basis. International Journal of Plant Sciences 165(5): 753-763.
Galil, J, and M. Zernoi. 1967. On the pollination of Zizyphus spina-christi (L.) Willd. in Israel. Israel Journal of Botany 16(2): 71-77.
Gleeson, S. K. 1982. Heterodichogamy in walnuts: Inheritance and stable ratios. Evolution 36(5): 892-902.
Ho, K.-Y., and T.-Y. Hung. 2011. Cladistic relationships within the genus Cinnamomum (Lauraceae) in Taiwan based on analysis of leaf morphology and inter-simple sequence repeat (ISSR) and internal transcribed spacer (ITS) molecular markers. African Journal of Biotechnology 10(24): 4802-4815.
Jong, P. C. 1976. Flowering and sex expression in Acer L.: A biosystematic study. Mededeelingen van de Landbouwhoogeschool te Wageningen 76-2: 1-201.
Kanehira, R. 1936. Formosan Trees Indigenous to the Islands, 2nd ed. Department of Forestry, Government Research Institute, Taihoku (In Japanese, with English notes).
Kubitzki, K., and H. Kurz. 1984. Synchronized dichogamy and dioecy in Neotropical Lauraceae. Plant Systematics and Evolution 147(3-4): 253-266.
Li, H.-L. 1963. Woody flora of Taiwan. Livingston Publishing Company, Narberth, Pennsylvania, USA.
Li, H.-W., J. Li, P.-H. Huang, F.-N. Wei, H.-P. Tsui, and H.Werff. 2008. Lauraceae. In: Z.-Y. Wu et al. [eds.], Flora of China, vol. 7, 102-254. Missouri Botanical Garden Press, St. Louis, MO.
Li, J., D. C. Christophel, J. G. Conran, and H.-W. Li. 2004. Phylogenetic relationships within the ‘core’ Laureae (Litsea complex, Lauraceae) inferred from sequences of the chloroplast gene matK and nuclear ribosomal DNA ITS regions. Plant Systematics and Evolution 246(1-2): 19-34.
Li, L., J. Li, J. G. Conran, and X.-W. Li (H.-W. Li). 2007. Phylogeny of Neolitsea (Lauraceae) inferred from Bayesian analysis of nrDNA ITS and ETS sequences. Plant Systematics and Evolution 269(3-4): 203-221.
Li, Q.-J., Z.-F. Xu, Y.-M. Xia, L. Zhang, X.-B. Deng, and J.-Y. Gao. 2001. Study on the flexistyly pollination mechanism in Alpinia plants (Zingiberaceae). Acta Botanica Sinica 43(4): 364-369.
Li, Q.-J., Z.-F. Xu, W. J. Kress, Y.-M. Xia, L. Zhang, X.-B. Deng, J.-Y. Gao, and Z.-L. Bai. 2001. Pollination: Flexible style that encourages outcrossing. Nature 410(6827): 432.
Li, Q.-J., W. J. Kress, Z.-F. Xu, Y.-M. Xia, L. Zhang, X.-B. Deng, and J.-Y. Gao. 2002. Mating system and stigmatic behavior during flowering of Alpinia kwangsiensis (Zingiberaceae). Plant Systematics and Evolution 232(1-2): 123-132.
Liao, J.-C. 1988. The taxonomic revisions of the family Lauraceae in Taiwan. National Taiwan University, Taipei, Taiwan, ROC.
Liao, J.-C. 1996. Lauraceae. In: T.-C. Huang et al. [eds.], Flora of Taiwan, 2nd edition, vol. 2, 433-499. National Taiwan University, Taipei, Taiwan, ROC.
Lyrene, P. M.1983. Flowering and fruiting of Chinese jujubes in Florida. HortScience 18(2): 208-209.
Miller, H. 1875. Flowering of Hazel. Nature 12(1875): 26.
Nie, Z.-L., J. Wen, and H. Sun. 2007. Phylogeny and biogeography of Sassafras (Lauraceae) disjunct between eastern Asia and eastern north America. Plant Systematics and Evolution 267(1-4): 191-203.
Nirody, B. S. 1922. Investigations in avocado breeding. California Avocado Association Annual Report 6: 65-78.
Norman, E. M., and D. Clayton. 1986. Reproductive biology of two Florida pawpaws: Asimina obovata and A. pygmaea (Annonaceae). Bulletin of the Torrey Botanical Club 113(1): 16-22.
Papademetriou, M. K. 1976. Some aspects of the flower behaviour, pollination and fruit set of avocado (Persea americana Mill.) in Trinidad. California Avocado Society Yearbook 60(1976): 106-152.
Pendleton, R. L., D. C. Freeman, E. D. McArthur, and S. C. Sanderson. 2000. Gender specialization in heterodichogamous Grayia brandegei (Chenopodiaceae): evidence for an alternative pathway to dioecy. American Journal of Botany 87(4): 508-516.
Pendleton, R. L., E. D. McArthur, D. C. Freeman, and A. C. Blauer. 1998. Heterodichogamy in Grayia brandegei (Chenopodiaceae): report from a new family. American Journal of Botany 75(2): 267-274.
Polito, V. S., and K. Pinney. 1997. The relationship between phenology of pistillate flower organogenesis and mode of heterodichogamy in Juglans regia L. (Juglandaceae). Sexual Plant Reproduction 10(1): 36-39.
Raju, M. V. S. 1961. Morphology and anatomy of the Saururaceae. I. Floral anatomy and embryology. Annals of the Missouri Botanical Garden 48(2): 107-124.
Rama Devi, K., J. B. Atluri, and C. Subba Reddi. 1989. Pollination ecology of Zizyphus mauritiana (Rhamnaceae). Proceedings of the Indian Academy of Sciences. Plant Sciences 99(3): 223-239.
Renner, S. S. 2001. How common is heterodichogamy? Trends in Ecology and Evolution 16(11): 595-597.
Rohwer, J. G. 2000. Toward a phylogenetic classification of the Lauraceae: Evidence from matK sequences. Systematic Botany 25(1): 60-71.
Rohwer, J. G. 2009. The timing of nectar secretion in staminal and staminodial glands in Lauraceae. Plant Biology 11: 490-492.
Rohwer, J. G., J. Li, B. Rudolph, S. A. Schmidt, H. Werff, and H.-W. Li. 2009. Is Persea (Lauraceae) monophyletic? Evidence from nuclear ribosomal ITS sequences. Taxon 58(4): 1153-1167.
Rohwer, J. G., P. L. R. De Moraes, B. Rudolph, and H. Werff. 2014. A phylogenetic analysis of the Cryptocarya group (Lauraceae), and relationships of Dahlgrenodendron, Sinopora, Triadodaphne, and Yasunia. Phytotaxa 158(2): 111-132.
Schneider, E. L., and J. M. Jeter. 1982. Morphological studies of the Nymphaeaceae. XII. The floral biology of Cabomba caroliniana. American Journal of Botany 69(9): 1410-1419.
Schneider, E. L., and L. A. Moore. 1977. Morphological studies of the Nymphaeaceae. VII. The floral biology of Nuphar lutea subsp. macrophylla. Brittonia 29(1): 88-99.
Schneider, E. L., and T. Chaney. 1981. The floral biology of Nymphaea odorata (Nymphaeaceae). The Southwestern Naturalist 26(2): 159-165.
Shivaprasad, D., C. N. Prasannakumar, R. K. Somashekar, and B. C. Nagaraja. 2015. Reproductive biology of Cinnamomum sulphuratum Nees. from wet evergreen forest of Western Ghats in Karnataka. Proceedings of the International Academy of Ecology and Environmental Sciences 5(1): 7-15.
Simpson, M. G. 2010. Plant Systematics, 2nd edition. Academic Press, US.
Skutch, A. F. 1945. The behavior of the flowers of the aguacatillo (Persea caerulea). Torreya 45(4): 110-116.
Soltis, P. S., D. E. Soltis, M. J. Zanis, and S. Kim. 2000. Basal lineage of angiosperms: relationships and implications for floral evolution. International Journal of Plant Sciences 161(6 Suppl.): S97-S107.
Soltis, P. S., S. F. Brockington, M. J. Yoo, A. Piedrahita, M. Latvis, M. J. Moore, A. S. Chanderbali, and D. E. Soltis. 2009. Floral variation and floral genetics in basal angiosperms. American Journal of Botany 96(1): 110-128.
Stout, A. B. 1922-1923. A study in cross-pollination of avocados in southern California. California Avocado Association Annual Report 7: 29-45.
Stout, A. B. 1927. The flower behavior of avocados. Journal of the New York Botanical Garden 7: 145-203.
Stout, A. B. 1933. The pollination of avocados. University of Florida Agricultural Experiment Station Bulletin 257.
Stout, A. B., and E. M. Savage. 1925. The flower behavior of avocados with special reference to interplanting. Proceedings of the Florida State Horticultural Society 38: 80-91.
Thien, L. B. 1974. Floral biology of Magnolia. American Journal of Botany 61(10): 1037-1045.
Thien, L. B., D. A. White, and L. Y. Yatsu. 1983. The reproductive biology of a relict-Illicium floridanum Ellis. American Journal of Botany 70(5): 719-727.
Thompson, T. E., and L. D. Romberg. 1985. Inheritance of heterodichogamy in pecan. Journal of Heredity 76(6): 456-458.
Troll, W. 1964. Die Infloreszen, Typologie und Stellung im Aufbau des Vegetationskoerpers: Deskriptive Morphologie der Indloreszenzen - Typologie der Infloreszenzen. Lubrecht and Cramer, Limited.
Trujillo, C. G., and A. N. Sérsic. 2006. Floral biology of Aristolochia argentina (Aristolochiaceae). Flora 201(5): 374-382.
Waster, P. J. 1910. Pollination experiments with Anonas. Bulletin of the Torrey Botanical Club 37(11): 529-539.
Watanabe, S., N. Noma, and T. Nishida. 2015. Flowering phenology and mating success of the heterodichogamous tree Machilus thunbergii Sieb. et Zucc (Lauraceae). Plant Species Biology doi: 10.1111/1442-1984.12078.
Wiersema, J. H. 1988. Reproductive biology of Nymphaea (Nymphaeaceae). Annals of the Missouri Botanical Garden 75(3): 795-804.
Worboys, S. J., and B. R. Jackers. 2005. Pollination processes in Idiospermum ausrtaliense (Calycanthaceae), an arborescent basal angiosperm of Australia’s tropical rain forests. Plant Systematics and Evolution 251(2-4): 107-117.
Werff, H., and H. G. Richter. 1996. Toward an improved classification of Lauraceae. Annals of the Missouri Botanical Garden 83(3): 409-418.
Wu, S.-H., C.-Y. Hwang, T.-P. Lin, J.-D. Chung, Y.-P. Cheng, and S.-Y. Hwang. 2006. Contrasting phylogeographical patterns of two closely related species, Machilus thunbergii and Machilus kusanoi (Lauraceae), in Taiwan. Journal of Biogeography 33(5): 936-947.
Zietsman, P. C., A. E. Wyk, and F. C. Botha. 1989. Vegetative and reproductive phenology of Ziziphus mucronata subsp. mucronata (Rhamnaceae). South African Journal of Botany 55(6): 564-573.