台灣在不同南島語族遷徙理論中,地位有極大的差異,其中一個原因是考古 學和語言學證據分別適用於不同的人類活動面向,語言擴散指涉到文化的影響,
例如族群間的語言採借與改變,而人類基因的擴散則涉及族群人口的大量流動或 族群通婚(Cavalli-Sforza, Menozzi, and Piazza, 1994)。所以探討南島文化的傳播與南 島語族人的傳播路徑會有相似但不完全相同的結果 (陳柏楨, 1998)。由於樹皮布是 南島語族重要的文化成分之一 (Howard, 2006; Cameron, 2008),所以追查構樹的傳 播能讓我們進一步了解南島語族文化擴張的過程。也因為探討的是文化的傳播,
所以我會以語言學證據為輔,體質人類學證據為主與構樹的傳播做比較。
35 發生的重要證據。而Diamond (2000)與 Moodley 等人 (2009) 所提到的第二次長期 停留,雖然因為近大洋洲取樣地點及數量等原因,並無法斷定近大洋洲與台灣或
36
完全排除慢船說發生的可能性。但是目前的證據明確指出,台灣在南島文化傳播 中的地位,可部分印證快車說中對於南島文化起源與傳播中的論述。
37
5 結論
1. 結合不同的分析資料,構樹族群間的基因交流程度相當低,顯示其傳播能力有 限,長距離天然傳播不易發生,所以結構中顯示的族群分布擴張與長距離傳播 可能都是人為因素的影響。
2. 東亞地區內構樹存有明顯的遺傳差異,且差異程度大致與距離相關,而大洋洲 中夏威夷群島的構樹因為多次人為傳播而呈現複雜的結構。
3. 遠大洋洲地區構樹的遺傳變異顯示至少有兩次構樹傳入,其中一次侷限於夏威 夷,另一次的影響廣布遠大洋洲,其中影響範圍較廣的事件與台灣東部及南部 高度相關。
4. 越南的構樹雖然無法完全排除與大洋洲構樹族群間的關係,但相對於台灣或是 中國大陸,關係比較不重要。
5. 菲律賓、近大洋洲、留尼旺群島出現的構樹應該是近代商業繁殖傳播的結果,
而且來源可能是中國或是大陸東南亞。
6. 構樹的傳播大致能與快車說推定的南島語族拓殖路線相互對照,但是快車說所 提及在島嶼東南亞的傳播與在近大洋洲的停留,還需要更多證據才能確定。
38
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表 1、PCR 引子表
分子標記 引子 引子序列 來源
ITS ITS4 ITS5B
5’GCTTA AACTC AGCGG GTACG 3’
5’TCGCG AGAAG TCCAC TGAA 3’
ndhF-rpl32 rpl32-R ndhF
ndhF-rpl32 Denature Denature
50
表 3、構樹各地區族群個體數 (N) 、ITS 片段核苷酸歧異度 (nucleotide diversity: π)、基因歧異度 (haplotype diversity: h )以及族群個 體中性檢測 (Tajima’s D test)
地理區 個體(N) 核苷酸歧異度(π) 基因歧異度(h) Tajima’s D
台灣 197 0.00062 0.35573 -0.32173
中國 234 0.0008 0.42348 -0.6489
福建 14 0.00027 0.15385 -1.14915
廣東 123 0.00061 0.34684 -0.08198
廣西 17 0.00138 0.61579 0.93886
雲南 68 0.00138 0.42362 0.0834
四川 4 0.00088 0.5 0.0834
上海 4 0 0 ~
北京 3 0 0 ~
海南 1 ~ ~ ~
日本 3 0 0 ~
越南 41 0.00069 0.37317 -0.29025
菲律賓 12 0.00093 0.5303 0.48516
印尼 16 0 0 ~
大洋洲 62 0.00182 0.5542 -0.67449
近大洋洲 2 0.00174 1 ~
遠大洋洲 60 0.00175 0.52373 1.12099
留尼旺 2 0 0 ~
Total 567 0.00134 0.61551 -0.67449
51
表 4、構樹各地區族群個體數 (N) 、葉綠體 ndhF-rpl32 片段核苷酸歧異度 (nucleotide diversity: π)、基因歧異度 (haplotype diversity: h ) 以及族群個體中性檢測 (Tajima’s D test),*表示 P<0.005
地理區 個體(N) 核苷酸歧異度(π) 基因歧異度(h) Tajima’s D
台灣 169 0.00264 0.81866 -0.42358
中國 206 0.0022 0.85544 -1.10547
福建 14 0.00152 0.70513 1.33302
廣東 115 0.00208 0.74003 0.47284
廣西 17 0.00232 0.86275 0.16514
雲南 48 0.00184 0.79255 0.84213
四川 3 0.00057 0.66667 ~
上海 4 0 0 ~
北京 4 0.0017 0.5 -0.78012
海南 1 ~ ~ ~
日本 3 0 0 ~
越南 46 0.00122 0.70808 0.6437
菲律賓 14 0.00089 0.61538 0.36212
印尼 16 0 0 ~
大洋洲 58 0.00381 0.53297 2.5323*
近大洋洲 2 0 0 ~
遠大洋洲 56 0.0037 0.49935 -1.10547
留尼旺 2 0 0 ~
Total 514 0.00318 0.92017 -1.14614
52
表 5 東亞與大洋洲構樹 ITS 片段兩兩族群間遺傳分化與基因交流指數。對角線下方為 Fst值,對角線上方為 Nm值。
Mn
台灣 福建 廣東 廣西 雲南 四川 北京 上海 日本 越南 菲律賓 印尼 近大洋洲 遠大洋洲 留尼旺 台灣 0.00 1.61 1.07 0.42 1.00 -1.55 1.12 1.12 1.12 0.12 0.31 1.12 0.11 0.35 0.04 福建 0.13 0.00 6.08 0.53 1.33 0.11 0.27 0.00 0.08 0.37 0.02 廣東 0.19 0.04 0.00 1.22 1.14 2.49 1.03 1.03 1.03 0.23 0.31 1.03 0.11 0.51 0.05 廣西 0.38 0.32 0.17 0.46 0.53 0.32 0.32 0.32 1.55 0.26 0.32 0.13 0.70 0.08 雲南 0.20 0.16 0.18 0.35 0.00 1.95 0.91 0.91 0.91 0.14 1.40 0.91 0.12 0.37 0.10
四川 -0.19 0.00 0.09 0.32 0.11 0.00 0.15 0.40 0.14 0.41 0.06 北京 0.18 0.00 0.20 0.44 0.22 0.00 0.00 0.07 0.21 0.05 0.29 0.00 Fst 上海 0.18 0.00 0.20 0.44 0.22 0.00 0.00 0.00 0.07 0.21 0.05 0.14 0.00
日本 0.18 0.00 0.20 0.44 0.22 0.00 0.00 0.00 0.00 0.07 0.21 0.05 0.29 0.00 越南 0.67 0.69 0.53 0.14 0.64 0.62 0.79 0.79 0.79 0.00 0.11 0.07 0.05 0.38 0.03 菲律賓 0.45 0.48 0.44 0.49 0.15 0.38 0.55 0.55 0.55 0.69 0.00 0.21 0.11 0.26 0.44 印尼 0.18 0.00 0.20 0.44 0.22 0.00 0.00 0.00 0.00 0.79 0.55 0.00 0.05 0.29 0.00 近大洋洲 0.70 0.76 0.70 0.66 0.67 0.65 0.82 0.82 0.82 0.80 0.70 0.82 0.00 0.72 0.03 遠大洋洲 0.42 0.40 0.33 0.26 0.40 0.38 0.47 0.65 0.47 0.40 0.49 0.47 0.26 0.00 0.10 留尼旺 0.85 0.93 0.85 0.77 0.72 0.80 1.00 1.00 1.00 0.90 0.36 1.00 0.90 0.72 0.00
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表 6 東亞與大洋洲構樹葉綠體 ndhF-rpl32 片段兩兩族群間遺傳分化與基因交流指數。對角線下方為 Fst值,對角線上方為 Nm值。
Nm
台灣 福建 廣東 廣西 雲南 四川 北京 上海 日本 越南 菲律賓 印尼 近大洋洲 遠大洋洲 留尼旺 台灣 0.00 1.24 1.07 1.07 0.85 0.33 0.85 0.45 0.21 0.85 0.73 0.46 0.27 1.77 0.27 福建 0.29 0.00 1.65 1.65 1.16 0.26 0.73 0.80 0.13 1.32 1.07 0.12 0.19 0.69 0.19 廣東 0.32 0.23 0.00 7.24 4.49 1.11 0.61 0.56 0.21 3.53 1.59 0.14 0.88 0.73 0.88 廣西 0.32 0.23 0.06 0.00 4.81 0.75 0.64 0.51 0.29 6.55 1.87 0.16 0.61 0.82 0.61 雲南 0.37 0.30 0.10 0.09 0.00 0.83 0.51 0.36 0.15 14.59 1.31 0.12 0.63 0.60 0.63 四川 0.61 0.65 0.31 0.40 0.38 0.00 0.19 0.06 0.03 0.45 0.24 0.03 0.30
北京 0.37 0.41 0.45 0.44 0.49 0.72 0.00 0.25 0.11 0.49 0.42 0.13 0.14 0.57 0.14 Fst 上海 0.53 0.39 0.47 0.50 0.58 0.90 0.67 0.00 0.00 0.29 0.20 0.00 0.00 0.34 0.00 日本 0.70 0.79 0.70 0.63 0.76 0.94 0.82 1.00 0.00 0.11 0.08 0.00 0.00 0.38 0.00 越南 0.37 0.27 0.12 0.07 0.03 0.53 0.51 0.63 0.81 0.00 1.38 0.09 0.31 0.59 0.31 菲律賓 0.41 0.32 0.24 0.21 0.28 0.67 0.55 0.72 0.86 0.27 0.00 0.06 0.15 0.54 0.15 印尼 0.52 0.81 0.78 0.76 0.81 0.95 0.80 1.00 1.00 0.85 0.89 0.00 0.00 0.82 0.00 近大洋洲 0.65 0.72 0.36 0.45 0.44 0.00 0.78 1.00 1.00 0.61 0.77 1.00 0.00 0.26
遠大洋洲 0.22 0.42 0.41 0.38 0.45 0.63 0.47 0.59 0.57 0.46 0.48 0.38 0.66 0.00 0.26 留尼旺 0.65 0.72 0.36 0.45 0.44 0.00 0.78 1.00 1.00 0.61 0.77 1.00 0.00 0.66 0.00
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表 7 構樹 ITS 片段的 Nested Clade Analysis (NCA)的推論順序及結果表。Clade 名稱如圖 4,推論依據 Temploton (2004)。
1-step clade Total Cladogram Clade 1-1
1-2-11-17-4 NO Χ 2=10.0807
Restricted gene flow with isolation by distance (restricted dispersal by distance in non-sexual species).
Clade 2-1 1-2-11-12 NO Χ 2=920.876*
Contiguous range expansion.
Clade 1-2 1-2-11-12 NO Χ 2=957.3979*
Contiguous range expansion.
Clade 1-3 1-2-3-4 NO Χ 2=80.3174*
Restricted gene flow with isolation by distance (restricted dispersal by distance in non-sexual species).
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表 8 構樹葉綠體 ndhF-rpl32 片段的 Nested Clade Analysis (NCA)的推論順序及結果表。Clade 名稱如圖 5。
表 8 構樹葉綠體 ndhF-rpl32 片段的 Nested Clade Analysis (NCA)的推論順序及結果表。Clade 名稱如圖 5。