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. |
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 |