臺灣博碩士論文加值系統

中越邊境喀斯特岩溶地貌蘊含了十分豐富的植物多樣性及特有度，為了解石
灰岩植物多樣性的生成機制，本論文以分布於中越邊境石灰岩地貌的秋海棠來探
討遺傳分化。中越邊境石灰岩秋海棠包含側膜組秋海棠、棒果組秋海棠以及四種
秋海棠組秋海棠，物種數高達七十種。因大部分的石灰岩物種傳播能力不佳且在
高度破碎化的石灰岩生境多呈現狹域分布的格局，推論基因交流的受限可能為造
成其遺傳分化之主因；形態差異主要在營養器官且與環境無明顯相關性，暗示形
態的變異可能為非適應性變化。本研究利用兩段葉綠體 DNA 片段 (trnC-ycf6、
petB-petD) 重建 61 種中越邊境石灰岩秋海棠的親緣關係，並根據親緣關係分析結
果選取四組近緣種支序 (A-D 支序) 以及三種廣泛分布種進行遺傳、地理以及形態
距離三方的相關性檢測。研究結果顯示，中越邊境石灰岩秋海棠形成一支持度不
高的單系群，其內包括十一支亞支序群；種內和種間層級的遺傳距離與地理距離
之相關性檢測，除了 D 支序外，皆具有顯著相關性；葉部形態變異與遺傳及地理
距離的相關性檢測在所有的檢測單位皆不具相關性；花部形態距離與遺傳及地理
距離的相關性檢測在所有的檢測，除了 C 支序外，皆不具相關性。根據分析結果
推論，中越邊境石灰岩秋海棠的高度物種多樣性為輻射演化的結果，傳播能力的
受限以及地理上的隔離為造成種化的重要推手，形態變異依據目前檢測結果傾向
為天擇造成，推論此結果可能與物種因具有棲位保守性相關。

V

Abstract
The Karst terrains across the Sino-Vietnamese border are known for its extremely
high plant diversity and endemism. To understand mechanisms that had generated its
remarkable biodiversity, we study patterns of genetic diversification of the
Sino-Vietnamese limestone Begonia. About 70 Begonia species are found exclusively in
the limestone habitats of the region, including all members of sect. Coelocentrum and
sect. Leprosae and four species of sect. Diploclinium. Because most of these limestone
species are poor dispersers and have narrow and restricted distributions on the highly
fragmented limestone habitats, limited gene flow might have been the major contributor
to their genetic diversification. Morphological variation of Sino-Vietnamese limestone
Begonia is mainly manifested in vegetative traits that have no apparent correlation to
their environments, implying that morphological variation could have been primarily
non-adaptive. We collected DNA sequences of two chloroplast intergenic spacers
(trnC-ycf6 and petB-petD) of 61 Sino-Vietnamese limestone Begonia species to
reconstruct their phylogenetic relationships. Mantel tests between genetic, geographical,
and morphological distances were performed for three widespread species and four
groups (clade A-D) of closely related species revealed by the phylogenetic trees. Results
show that Sino-Vietnamese limestone Begonia group forms a poorly supported clade
composed of 11 subclades. A significant positive correlation between genetic and
geographic distance was observed in both inter- and intraspecific tests except for clade
D. Correlations between morphological distance of leaf traits and genetic distance were
not significant in all tests. Correlations between leaf morphological distance and
geographic distance were also not significant in all tests. Except for clade C,
VI

correlations between floral morphological distance and genetic distance were not
significant. Correlations between morphological distance of floral traits and geographic
distance were not significant except for clade C. Based on these analyses and inferences,
the high species diversity of Sino-Vietnamese limestone Begonia appears to be results
of species radiation. Limited dispersability and geographic isolation should have played
an important role in spawning its species diversity. Morphological variation of these
limestone Begonia species appears to be resulted from natural selection, likely related to
the strong niche conservatism to their habitats.

i

目錄
國立臺灣大學碩博士學位論文口試委員會審定書 .................................... I
誌謝 .............................................................................................................. II
中文摘要 ..................................................................................................... IV
Abstract ........................................................................................................ V
圖目錄 .......................................................................................................... iv
表目錄 ......................................................................................................... vii
1. 前言 ........................................................................................................ 1
1.1. 廣西喀斯特石灰岩地區生物多樣性特性 .............................. 1
1.2. 中越邊境石灰岩秋海棠概述 .................................................. 2
1.3. 秋海棠屬植物親緣關係研究 .................................................. 3
1.4. 秋海棠屬植物種化機制假說 .................................................. 4
1.5. 「非」適應性輻射演化 .......................................................... 9
1.6. 生物避難所 ............................................................................ 15
1.7. 研究目的 ................................................................................ 18
2. 材料與方法 .......................................................................................... 19
2.1. 材料取樣 ................................................................................ 19
2.2. 植物全基因組 DNA 之萃取法 ............................................. 19
ii

2.2.1. CTAB DNA 萃取法 ........................................................ 19
2.2.2. Qiagen DNA 萃取組萃取 DNA..................................... 20
2.3. PCR 擴增及定序 .................................................................... 21
2.3.1. 聚合酶連鎖反應 ............................................................. 21
2.3.2. 洋菜膠電泳分析及 PCR 產物純化 ................................ 23
2.3.3. 定序 ................................................................................. 23
2.4. 譜系分析 ................................................................................ 23
2.4.1. 序列整理以及資料矩陣建立 ......................................... 23
2.4.2. 最大簡約性分析 ............................................................. 24
2.4.3. 最大似然性分析 ............................................................. 25
2.4.4. 貝葉氏導出式分析 ......................................................... 25
2.5. 相關性檢測 ............................................................................ 26
3. 結果 ...................................................................................................... 27
3.1. 葉綠體基因片段同質性分析 ................................................ 27
3.2. 葉綠體親緣關係分析 ............................................................ 28
3.3. 相關性檢測 ............................................................................ 35
3.3.1. 地理距離矩陣與遺傳距離矩陣相關性分析 ................. 35
3.3.2. 遺傳距離矩陣與形態距離矩陣相關性分析 ................. 38
3.3.2.1. 總體形態特徵檢測............................................ 38
3.3.2.2. 葉部形態特徵檢測............................................ 38
3.3.2.3. 花部形態特徵檢測............................................ 39
3.3.3. 形態距離矩陣與地理距離矩陣相關性分析 ................. 39

iii

3.3.3.1. 總體形態特徵檢測............................................ 39
3.3.3.2. 葉部形態特徵檢測............................................ 39
3.3.3.3. 花部形態特徵檢測............................................ 39
4. 討論 ...................................................................................................... 47
4.1. 中越邊境石灰岩秋海棠物種多樣性是否為輻射演化？ 47
4.2. 中越邊境石灰岩秋海棠的遺傳分化是否符合距離隔離模
式？ ................................................................................................ 52
4.3. 中越邊境石灰岩秋海棠的形態變異是否由基因漂變所產
生？ ................................................................................................ 56
4.4. 中越邊境石灰岩秋海棠的種化機制 .................................... 57
4.5. 未來研究方向 ........................................................................ 58
附錄一、取樣物種基本資料列表 ............................................................. 70
附錄二、基本形態資料 (Ku, 2006) ......................................................... 77
附錄三、基本形態性狀編碼表 (Ku, 2006) ............................................. 86
附錄四、相關性檢測詳細取樣資料 ......................................................... 92

59

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