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姓名 許秋容(Chiuo-Rong Sheue) 電子郵件信箱 d8725805@student.nsysu.edu.tw
畢業系所 生物科學系研究所(Biological Sciences)
畢業學位 博士(Ph.D.) 畢業時期 91學年第2學期
論文名稱(中) 東方紅樹林紅樹科植物之形態解剖比解
論文名稱(英) The Comparative Morphology and Anatomy of the Eastern Mangrove Rhizophoraceae
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    摘要(中) 紅樹林為一群具不同親緣關係,生長於熱帶及亞熱帶河口與海岸地區的維管束植物,主要為喬木及灌木所構成之森林。紅樹科的紅樹族包含四個屬,紅茄苳屬(Bruguiera)、細蕊紅樹屬(Ceriops)、水筆仔屬(Kandelia)及紅樹屬(Rhizophora),均為紅樹林之成員,故亦稱為紅樹林紅樹科植物(mangrove Rhizophoraceae),全世界計有17種。其中的14種分布於東非和太平洋西岸之間,屬於東方紅樹林種類。
    本研究即以東方紅樹林紅樹科之13種植物做系統性之形態及解剖比較,包括托葉、葉部、花部、果實和胎生苗等器官。另外,紅樹科的3屬4種陸生型植物亦包含於托葉與葉部的比較研究中。
    研究結果顯示所有紅樹科的植物之托葉向軸面基部有若干個至數百個指狀腺體(colleters)呈列狀排列聚集。本文首次提出同種植物的該腺體之聚集形狀、列數、數目及個別形態相當穩定,可做為紅樹科植物屬或種之分類鑑別特徵。並且本科的紅樹林植物較其陸生型植物具有大型托葉暨多數且大型的指狀腺體,可能意涵此托葉及腺體構造在本科紅樹林植物具重要之選擇適應性。三種形態的厚壁異形細胞在細蕊紅樹屬及紅樹屬的托葉中發現,但於紅茄苳、水筆仔屬則無。
    本研究結果釐清先前存在之葉部解剖構造的諸多矛盾及爭議處,對葉結構之塑性、厚壁異形細胞的類型及結晶等特徵亦首次提出。紅樹林紅樹科植物的葉厚具有厚角質層、環型且下陷的氣孔具顯著的內、外角質層突起、大細胞的下皮層具二層至八層、末稍管胞明顯增大,具厚壁異形細胞、含豐富的丹寧細胞和晶簇形結晶。而陸生型紅樹科的植物葉薄、微披角質層、不定型位葉表的氣孔角質層突起不顯著、無下皮層及增大之末稍管胞,亦無異形細胞及丹寧細胞,但同具束狀和晶簇形結晶。本族的四屬植物之上下葉表均有出現木栓疣的可能,但只在紅樹屬的葉下表規則地出現。
    多樣化的花部及果實特徵,提供許多重要且信實可靠的分類依據,尤其是花序、花瓣及雄蕊等特徵。本研究提供了本族植物詳盡的花部及果實(含胎生苗)特徵比較。若干獨特且有趣的現象,如2至4枚子葉或聯合子葉和子葉筒、外溢胚乳、紅茄苳屬的下胚軸具胚根鞘等,提供了紅樹科紅樹林植物胚胎學研究重要的的資訊。胎生苗懸掛於母株的方式有二種, 一為向下; 一為向上至斜下, 但後者於先前並未受到注意。
    經由本研究,確認兩個新分類群,其中之一為分布在南中國海以北的水筆仔,定名為Kandelia obovata C. R. Sheue, H. Y. Liu and J. W. H. Yong。Ceriops australis (C. T. White) E. R. Ballment, T. J. Smith III & J. A. Stoddart之多項形態及解剖特徵也首次於此描述及確認其分類位階。全世界系統性的紅樹林紅樹科研究仍然迫切需要,尤其是細蕊紅樹屬及紅樹屬。
    摘要(英) Mangroves are a group of the intertidal plants with unrelated phylogenetic affinity distributed in regions of estuaries, riverbanks or along the coastlines of tropical and subtropical areas. The four genera of tribe Rhizophoreae of Rhizophoraceae are important members of mangroves, namely Bruguiera, Ceriops, Kandelia and Rhizophora and also known as mangrove Rhizophoraceae, with a wide distribution range. Fourteen of 17 species of mangrove Rhizophoraceae distributed in the areas between the Eastern Africa to the Pacific West Coast, belong to the eastern mangrove Rhizophoraceae.
    Thirteen species of the eastern mangrove Rhizophoraceae from different populations were sampled for the studies in comparative morphology and anatomy on stipules, leaves, flowers, fruits and viviparous seedlings. In contrast, four species of three genera belong to the inland Rhizophoraceae were also sampled. Such works were emphasized on stipules and leaves.
    Three kinds of sclereids were found in the stipules of Ceriops and Rhizophora, but none in Kandelia and Bruguiera. Several to hundreds colleters arranging in several series occur at the adaxial base of a stipule in Rhizophoraceae. The aggregated forms, series number, number and individual morphology of colleters are generic or specific, and could be an aid for taxonomic identification. The number and size of colleters found in the mangrove Rhizophoraceae are more and larger than those of the inland species. Additionally, the size of stipule in mangrove Rhizophoraceae is larger than those of the inland Rhizophoraceae. These characters might imply an adaptive selection of stipule and colleters in Rhizophoreae.
    In this study, many of the previous discrepancies of leaf anatomical features were clarified. Meanwhile, the plasticity of leaf structure, features of sclereid ideoblasts and crystals were not previously reported. The leaves of mangrove Rhizophoraceae present thick leaves and cuticular wax, sunken and cyclocytic stomata with distinct inner and outer ridges, 2 to 8 cell-layered hypodermis, enlarged terminal tracheids, four types of sclereid ideoblasts and abundant tanniferous cells and drused crystals. All species of the mangrove Rhizophoraceae has potential to produce cork warts on both sides, but only Rhizophora has the warts mainly on abaxial side regularly. Detailed stomatal features could serve as diagnostic character in species level. On the contrary, the inland species has thin leaves with less cuticular wax, superficial anomocytic stomata, no hypodermis and enlarged terminal tracheids, drused and raphid crystals.
    The diverse morphological features in flowers and fruits, especially in inflorescences, petals and stamens, provided valuable and reliable taxonomic characteristics. A detailed survey for flower and fruit parts (including the viviparous seedlings) were provided in this study. Some unique and curious phenomenon, including multi-cotyledons to united cotyledon and cotyledon collar, endosperm overflow, the present of coleorhiza in the hypocotyl of Bruguiera, provided interesting and valuable embryological information for the mangrove Rhizophoraceae. Downward and upward orientation seedlings on the mother trees were observed, however the latter were not noticed previously.
    Two new taxa were recognized based on many lines of evidences in this study. A new species, Kandelia obovata C. R. Sheue, H. Y. Liu and J. W. H. Yong, which distributed northern from the South China Sea, was named and described. The detailed morphological and anatomical features of Ceriops australis (C. T. White) E. R. Ballment, T. J. Smith III & J. A. Stoddart were firstly described, which confirmed its taxonomic status. A systematic approach with a global view on the mangrove Rhizophoraceae is still needed, especially for Ceriops and Rhizophora.
    關鍵字(中)
  • 解剖
  • 紅樹林
  • 紅樹科
  • 形態
  • 關鍵字(英)
  • Morphology
  • Rhizophoraceae
  • Mangroves
  • Anatomy
  • 論文目次 Table of Contents
    Chapter 1. Introduction 1 ~14
    Chapter 2. Materials and Methods 15 ~22
    Introduction 15
    Materials 16
    Methods 17
    Chapter 3. Stipules and Colleters 23 ~36
    Introduction 23
    Results 25
    Stipule 25
    Morphology, and vernation 25
    Anatomy 26
    Colleter 27
    Aggregated shape, series and number--------- 27
    Individual shape and size 28
    Anatomy 29
    Ultrastructure 30
    Discussion 33
    Chapter 4. Leaf Morphology and Anatomy 51 ~115
    Introduction 51
    Results 54
    Morphological features of leaf 54
    Shape and size 54
    Lateral vein 55
    Leaf scars 56
    Vernation 56
    Stomata complex 57
    Cork warts 58
    Anatomical features of leaf 64
    Leaf structure 64
    Cuticular wax 65
    Epidermis 66
    Hypodermis 66
    Photosynthetic mesophyll 67
    Vein and terminal tracheid 68
    Mucus cell 69
    Sclereid ideoblast 70
    Crystal 71
    Preliminary study of ultrastructure 72
    Discussion 75
    Chapter 5. Flower and Pollen 116 ~175
    Introduction 116
    Results 119
    Inflorescence 119
    Bracteole and colleter 121
    Perianth 121
    Calyx morphology 122
    Sepal anatomy 123
    Petal morphology 123
    Petal anatomy 125
    Androecium 126
    Stamen number and position 126
    Stamen morphology 127
    Sporangiate number 129
    Dehiscence pattern 129
    Gynoecium 129
    Ovary position 129
    Carpel number 130
    Placentation and ovules 130
    Style and stigma 130
    Pollen 131
    Discussion 136
    Chapter 6. Fruit and Viviparous seedling 176 ~207
    Introduction 176
    Results 178
    Fruit 179
    Morphological features 179
    Anatomical features 180
    Viviparous seedling 182
    Orientation of seedling on the mother tree--- 182
    Morphological features of seedlings 183
    Detaching from mother tree 184
    Discussion 188
    Chapter 7. Conclusion --------------------------------------------------208~212
    Character comparison among taxa --------------------------- 208
    Characters applied to taxonomy ------------------------------ 210
    Perspectives ------------------------------------------------------ 211
    Literature Cited 213 ~228
    List of Figures
    Fig. 2.1. The distribution map of collection localities in this study 22
    Fig. 3.1. Stipule morphology and vernation of the mangrove Rhizophoraceae
    and two inland species of the Rhizophoraceae 38
    Fig. 3.2. The aggregated form of colleters at the adaxial base of stipule of the mangrove Rhizophoraceae 39
    Fig. 3.3. Individual morphology of colleters in stipules of the Rhizophoraceae 40
    Fig. 3.4. Stipules and colleters of the mangrove Rhizophoraceae 41
    Fig. 3.5. Anatomical features in stipule of the Rhizophoraceae 43
    Fig. 3.6. Detailed view of colleters in stipule of the mangrove Rhizophoraceae 45
    Fig. 3.7. Anatomical features of stipule and colleters of the mangrove
    Rhizophoraceae 47
    Fig. 3.8. Ultrastructure of colleter and stipule of Rhizophora apiculata 49
    Fig. 4.1. Leaf shapes of the genus Bruguiera 86
    Fig. 4.2 Leaf shapes of the genus Ceriops 88
    Fig. 4.3 Leaf shapes of the genus Kandelia 88
    Fig. 4.4. Leaf shapes of the genus Rhizophora 89
    Fig. 4.5. Leaf shapes of some inland species of the Rhizophoraceae s. l. 90
    Fig. 4.6. Leaf scar of the mangrove Rhizophoraceae 91
    Fig. 4.7. Stomata type and transverse section of guard cell of the Rhizophoraceae 92
    Fig. 4.8. The arrangements of subsidiary cells in the mangrove Rhizophoraceae 92
    Fig. 4.9. Detailed surface view of guard cells treated with clearing method from
    the lower surfaces of leaves in the mangrove Rhizophoraceae 93
    Fig. 4.10. Four types of sclereid ideoblasts in leaves of Rhizophora 94
    Fig. 4.11. Leaf scar and leaf surface features of the Rhizophoraceae, including mangrove and inland species 96
    Fig. 4. 12. Stomata of the mangrove Rhizophoraceae and two inland species 98
    Fig. 4.13. A series sections of a cork wart in leaves of Rhizophora stylosa 100
    Fig. 4.14. Leaf transverse sections of the genus Bruguiera 102
    Fig. 4.15. Leaf transverse sections of the genera Kandelia and Ceriops 104
    Fig. 4.16. Leaf transverse sections of the genus Rhizophora 106
    Fig. 4.17. Features of ending vein in the Rhizophoraceae 108
    Fig. 4.18. Sclereid ideoblast and mucus cell of genus Rhizophora 110
    Fig. 4.19. Crystal types and leaf transverse section of inland species of the Rhizophoraceae s. l. 112
    Fig. 4.20. Leaf ultrastructure of the mangrove Rhizophoraceae 114
    Fig. 5.1. The patterns and variation of inflorescence of the mangrove
    Rhizophoraceae 144
    Fig. 5.2. The detailed inflorescence of Ceriops tagal 145
    Fig. 5.3. The petal morphology of Bruguiera and petal variation of Bruguiera sexangula 145
    Fig. 5.4. Illustration of Bruguiera sexangula 146
    Fig. 5.5. Illustration of Ceriops australis 148
    Fig. 5.6. Illustration of Kandelia obovata 150
    Fig. 5.7. Detailed morphology of flower parts in the genus Rhizophora 152
    Fig. 5.8. Detailed morphology of flower parts of Rhizophora × lamarckii 153
    Fig. 5.9. Some characters of inflorescence of the mangrove Rhizophoraceae 154
    Fig. 5.10. Floral characters of large-flowered Bruguiera 156
    Fig. 5.11. Floral characters of small-flowered Bruguiera and petal section of B. sexangula 158
    Fig. 5.12. Flowers and petals of Ceriops 160
    Fig. 5.13. SEM view of trichomes on petals of Ceriops 162
    Fig. 5.14. Floral characters and sections of Ceriops 164
    Fig. 5.15. Floral characters of Kandelia candel and Kandelia obovata 166
    Fig. 5.16.Cross section of flower parts of Kandelia candel and Kandelia obovata
    168
    Fig. 5.17. Floral characters of Rhizophora 170
    Fig. 5.18. Pollen grains of the mangrove Rhizophoraceae 172
    Fig. 5.19. SEM view of pollen grains of Ceriops 174
    Fig. 6.1. The viviparous seedlings of the genus Bruguiera 192
    Fig. 6.2. The viviparous seedlings of the genus Ceriops, fruits with cotyledon
    collars only 193
    Fig. 6.3. The vivipary seedlings of Ceriops and their orientations on the mother
        trees 194
    Fig. 6.4. The viviparous seedlings of the genus Kandelia 195
    Fig. 6.5. The viviparous seedlings of Rhizophora 196
    Fig. 6.6. Some features of viviparous seedling of Bruguiera gymnorhiza at
    different stages 197
    Fig. 6.7. Some features of viviparous seedlings of Bruguiera parviflora and
        Bruguiera cylindrica at different stages 198
    Fig. 6.8. Some features of viviparous seedling of Ceriops at different stages 199
    Fig. 6.9. Some features of viviparous seedling of Kandelia obovata in
    longitudinal view at different stages 200
    Fig. 6.10. Some embryonic features of viviparous seedling in Rhizophora
    stylosa 201
    Fig. 6.11. Some features of viviparous seedling of the mangrove
    Rhizophoraceae 202
    Fig. 6.12. Some features of viviparous seedling of Kandelia and the
    reproductive organs of Bruguiera gymnorihza 204
    Fig. 6.13. Some embryonic features of viviparous seedling in Rhizophora
    stylosa 206
    List of Tables
    Table 2-1. Details of collection data 19
    Table 3.1. Morphological characters of stipules and colleters of mangrove Rhizophoraceae and four inland species 32
    Table 3.2. Morphometric characters of colleters of the mangrove Rhizophoraceae
    and two inland species 33
    Table 4.1. Morphological characters of leaves of the mangrove Rhizophoraceae
    and some inland species 61
    Table 4.2. Stomata complex characters of leaves of the mangrove Rhizophoraceae
    and three inland species 62
    Table 4.3. Size of guard cell of the mangrove Rhizophoraceae and one inland
    species 63
    Table 4.4. Variation of cork warts diameter in leaves of the genus Rhizophora
    from different populations 63
    Table 4.5. Features of leaf structure of the mangrove Rhizophoraceae and two
    inland species 73
    Table 4.6. Features of terminal tracheid, sclereid ideoblast and crystal type
    in leaves of the mangrove Rhizophoraceae and two inland species 74
    Table 5.1. Comparison of floral morphological features of Bruguiera 132
    Table 5.2. Comparison of floral morphological characters of Ceriops 133
    Table 5.3. Comparison of floral morphological characters between Kandelia
    candel and Kandelia obovata 134
    Table 5.4. Comparison of floral morphological characters of Rhizophora 135
    Table 5.5. Pollen characters of some species of the mangrove Rhizophoraceae 136
    Table 6.1. The Comparison of morphological features of fruit in the mangrove Rhizophoraceae 185
    Table 6.2. Comparison of anatomical character of viviparous seedling of
    the mangrove Rhizophoraceae 187
    參考文獻 Abeysinghe, P. D., L. Triest, B. de Greef, N. Koedam and S. Hettiarachi. 2000. Genetic and geographic variation of the mangrove tree Bruguiera in Sri Lanka. Aquat. Bot. 67: 131- 141.
    Areschoug, F. W. C. 1902. Untersuchungen
    口試委員
  • 劉和義 - 召集委員
  • 邱少婷 - 委員
  • 鄒稚華 - 委員
  • 黃玲瓏 - 委員
  • 楊遠波 - 指導教授
  • 口試日期 2003-06-24 繳交日期 2003-07-28

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