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。

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.

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

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