植物保護學會會刊 45：1-15, 2003 1
寡毛實蠅亞科（Diptera: Tephritidae: Dacinae）果實蠅之親緣關係
張弘毅 1* 曾經洲 2 蔡偉皇 3 高穗生 2 曾義雄 4
1. ︽現皘笰穨〆穦笆从 浪 2. 铭畃秏 ︽現皘笰穨〆穦笰穨媚 瑀 刚喷┮ 3. 瓣ミ芖厩挛厩╯┮ 4. 竒蕾 夹非浪喷 だ
( ら戳 地チ瓣 91 12 る 18 ら)
篕 璶
眎 驾* 竒瑆 罦ネ 竡 2003 硅を龟们ㄈDiptera: Tephritidae: Dacinae狦龟们 克絫闽玒 从玂穦 45：1－15
ノ 穝 の玭щ 秏 稲秏蹦眔 狦龟们夹 琁篈 ﹚秈︽ 16S rDNA ﹚だ猂侯浪癚篈だ摸籔だ疭紉 克絫闽玒ゑ癸だ猂璉硅を龟们妮(Bactrocera)璉硅を龟们ㄈ妮 狥よ狦龟们 B. (B.) dorsalis︹龟们B. (B.) ferruginea㎝硅を龟们ㄈ妮 Zeugodacus 灿盿龟们B. (Z.) ciliferaナ龟们B. (Z.) cucurbitae堵兵龟们B. (Z.) diaphora堵葶狾龟们B. (Z.) scutellata 龟们B. (Z.) synnephes玭ナ 龟们B. (Z.) tau硅を龟们妮Dacus 龟们ㄈ妮Callantra 芖 龟们D. (C.) formosanus 9 贺硅を龟们ㄈDacinae龟们のㄤ 10 贺 妮ㄤ狦龟们ㄈ 竤龟们 16S rDNA 戈 贺 框肚崩 阶硅を龟们ㄈ の籔ㄤㄈ狦龟们 克絫闽玒陪ボ┮蹦ノ だだ摸 狦 籔篈だ摸╰参 狦璓 虑 阶靡 1) 盢︹龟们の狥よ狦龟们粄 ﹚ 贺 眔坝篹2) 龟们妮Coelotrypes 纔龟们妮 Euphranta 縒ミ 妮3) 盿龟们壁Gastrozonini籔硅を龟们ㄈ ﹏ゝ竤闽玒 络睲4) 龟们ㄈ Trypetinae 龟们壁 Trypetini のぶを龟们壁Adramini 虫 竤
(闽龄迭狦龟们硅を龟们ㄈ16S rDNA克絫闽玒) * 硄癟E-mail: [email protected] 2 植物保護學會會刊 第 45 卷 第 1 期 2003
狐 摸 だだ摸戈 禸 癚 硅を龟们ㄈ贺 克絫闽玒崩阶ㄤ 狦龟们Tephritidae硄嘿 狦龟 籔狦龟们ㄤㄈ贺摸 克絫闽玒 们Fruit fly笰穨 蛮ヘ 竒蕾 ┦程 璶 ㄤ 甡狦ナ 籔よ猭 摸 贺摸ㄤ 挛盚ネ狦龟 璓ㄏ狦 龟籊年 狦礚猭 癸狦攫 ネ 狦龟们夹 籤 硑 腨 パㄤ 挛肩旅 芖 穝 狦龟 矱 秈闽浪 眖 の玭щ 秏らる兼 ┺ 狦龟 芠浪琩ㄓ ㄤ竒 跋の ┺ 稲秏铭 ノヒ 蕾甡 腨 ┦瓣悔 琌獶跋瓣產 膀 猳methyl eugeol们cuelure 碭 盢狦龟们 璶 浪甡挛 炳狾の独フ屡厚︹贺翲 狦龟们Ceratitis capitata狥よ狦 竚λ狶馒れ狶皐 狶 よ秈 龟们Bactrocera dorsalis ナ 龟们 ︽狦龟们夹 ﹚戳蹦 ┮ 莉 狦龟 Bactrocera cucurbitae孽狦龟们 们 挛 皐 狸夹 ㄌ沮篈疭 Bactrocera tryoni ﹁狦龟们 紉 ﹚だ摸盢 だ 狦龟们 Anastrepha ludens の 狦狦龟们 挛 95 %皊弘┪ 狸玂 称 16S Rhagoletis pomonella τㄤ rDNA ﹚だ猂ノ 讽 贺摸玒妮硅を龟们ㄈ 璉硅を 龟们妮Bactorcera ぶ计妮赣 夹 ﹚ 硅を龟们妮Dacus 贺摸ヘ玡 狦龟们皐 夹 篈だ摸 ﹚ 祇 硅を龟们ㄈ贺摸笷 7 κ 璶玒ㄌ沮祇 闽浪だ摸篈 贺ㄤ 璉硅を龟们妮 5 κ贺硅を龟 磞瓃(2, 3, 17, 21, 31, 32, 34, 35, 36)㎝だ摸╰参 们妮 2 κ贺芖祇 璉硅を龟们 魁(1, 27) だ摸 妮贺摸 31 贺硅を龟们妮 3 贺(27, 35)璉 硅を龟们妮 狦龟们妮 贺摸计程 だ猂 妮瞶だ 璶だ 狥よ跋 蹦莉 狦龟们虫 狦龟们皊弘┪ の緿瑆跋ㄤネ篈聪┮だて妓┦ 狸玂 挛妓 睪瘁硉縤 璶盚ネ从 狦龟 笛 て DNA LR-J-12883 の 罙秆狦龟们贺摸 克絫闽玒惠 LR-N-13770 ま糤碩采絬砰 16S rDNA ヵ苦ㄤ篈ネ 疭┦瞶だ よ 900 荒琿 砰筿猘だ瞒竒 戈だ猂 よ 讽 ╯(9, パ恨琖 て LR-J- 12883 10, 11, 12, 22, 26, 36)计 琌篈 疭 LR-J-13323LR-N-13398LR-N-13770 紉 璶 だ猂ㄌ沮 穝 だネм 荒ま秈︽虫 舼 干 荒 砃 ノら 約獂(7, 14, 18, 20)ㄤ 采絬砰 PCR 糤碩は の猘笆秆 だ猂(7, 13, 14, 19, 26, 16S rDNA だ荒だ猂盽砆ノㄓゑ 30) ﹍计沮家览筿猘酚の 癸克絫闽玒パ采絬砰 DNA 琌ダ╰框 Chromas 硁砰癸 狦 荒だ 肚 ︽膀 舱ㄤ rDNA 跋琿 猂硁砰DNASIS® for Windows (Hitachi) 程ㄣ 玂┦ 続贺 克絫闽玒 秈︽ はの琿硈 だ猂 妓 (4, 16) ゅ沽刚ノ芖 狦龟们贺 珇 16S rDNA 荒戈畐 絋粄 寡毛實蠅亞科（Diptera: Tephritidae: Dacinae）果實蠅之親緣關係 3
籔戈畐 箇 荒 篈だ摸の盚 猵 甅杆硁砰Vector NTI Suite (v.6.0) 刚喷┮ㄏノ 狦龟们竒篈 ﹚ (InforMax, Inc.)DNASIS® (Hitachi)の 19 贺狦龟们珹硅を龟们ㄈ 2 妮 3 MEGA2 秈︽荒ゑ癸だ猂(24)ゑ癸 ㄈ妮 9 贺 竤ゑ癸 狦龟们珹 贺 框肚のだ摸克絫闽玒 您龟们ㄈCeratitidinae滇龟们ㄈ Phytalmiinae ㎝ 龟们ㄈ 狦籔癚阶 Tephritinae 2 妮 2 贺の 龟们 ㄈTrypetinae4 妮 4 贺
19 贺ㄑ刚狦龟们 だ摸 Table 1. Classification status of the 19 tested fruit flies Taxon Chinese name Subfamily Dacinae 硅を龟们ㄈ Tribe Dacini 硅を龟们壁 Bactrocera (Bactrocera) dorsalis (Hendel) 狥よ狦龟们 B. (B.) ferruginea (Fabricius) ︹龟们 B. (Zeugodacus) cilifera (Hendel) 灿盿龟们 B. (Z.) cucurbitae (Coquillet) ナ龟们 B. (Z.) diaphora (Hendel) 堵兵龟们 B. (Z.) scutellata (Hendel) 堵葶狾龟们 B. (Z.) synnephes (Hendel) 龟们 B. (Z.) tau (Walker) 玭ナ龟们 Dacus (Callentra) formosanus Tseng & Chu 芖 龟们 Outgroup 竤 Subfamily Ceratitidinae 您龟们ㄈ Tribe Gastrozonini 盿龟们壁 Acrotaeniostola sexvittata Hendel 兵龟们 Gastrozona fasciventris (Macquart) 盿龟们 Subfamily Phytalmiinae 滇龟们ㄈ Tribe Acanthonevrini 龟们壁 Acanthonevra speciosa (Hendel) 翴龟们 Phorelliosoma hexachaeta Hendel を 风龟们 Subfamily Tephritinae 龟们ㄈ Tribe Tephritini 龟们壁 Spathulina acroleuca (Schiner) 狠フ巴 龟们 Tribe Schistopterini 吊龟们壁 Rhabdochaeta formosana (Shiraki) 腳畄次を龟们 Subfamily Trypetinae 龟们ㄈ Tribe Adramini ぶを龟们壁 Euphranta apicalis Hendel 狠 纔龟们 Coelotrypes latilimbata Enderlein 溜絫 龟们 Tribe Trypetini 龟们壁 Carpophthoracidia matsumotoi Shiraki 猀 龟们 Fusciludia ensifera (Ito) 糃 フ葶龟们 4 植物保護學會會刊 第 45 卷 第 1 期 2003
瓃狦龟们 盚絛瞅の祇ネ よ latiflorus Munro λ Phyllostachys 硅を龟们ㄈ璉硅を龟们妮 璉硅を makinoi Hayata λPhyllostachys nigra 龟们ㄈ妮 贺摸よ 狥よ狦龟们 ㄈび (Lodd.) Munro ﹕﹙λPhyllostachys 跋 筂祇ネ 贺摸璶甡狦攫 狦龟 pubescens Mazel ex. ll. de Leh (31)滇 ㄤ盚从 禬筁 250 贺 狦攫の 垫狦 龟们ㄈ 翴龟们の を 风龟们 龟ㄤ 39 117 贺竒筁箎▅絋粄(6)︹ 盚 玡λ狶耕 莉妮 龟们 だ摸籔狥よ狦龟们 ┺跋 笆 贺摸度﹗ だ摸厨盢ㄤ籔狥よ狦龟们 ┺λ狶祇瞷(8) 龟们ㄈ 狠 纔龟 贺 ㄤ盚从 璶 狦攫璉硅を龟 们玒盚ネ 递Aeginitia indica Roxb 们妮硅を龟们ㄈ妮 贺摸よ 灿盿龟们 讽(Orobanchaceae) 掺 (31)溜絫 龟 璶甡れ Momordica cochinchinensis 们の糃 フ葶龟们 盚 猀 龟们 (Lour.) Spreng. (Cucurbitaceae) 盚 沮ㄤ 恨 崩代 肩 璫(6, 31)ナ龟们 甡 の璛 盚ネ 贺摸 龟们ㄈ 狠フ巴 龟们 Solanaceae从 の狦龟程腨 狦 璶甡碘从 繷 妮肩矱 龟们 盚笷 125 贺ㄤ 12 璫 贺摸(28)腳畄次を龟们 盚从 ︺ 42 贺竒筁箎▅絋粄(6) 甡 耕 腨 垫 Blumea sp. 碘 (Compositae)(33) 珹玭ナ妮 Cucurbita ナ妮 Luffa 妮Lagenaria ナ妮Cucumis だだ摸 璚ナ妮 Momordicaナ妮 Benincasa 竒采絬砰 16S rDNA 戈ゑ癸 19 ﹁ナ妮Citrullus珹加妮Trichosanthes 贺狦龟们摸絫闽玒 Vector NTI Suite だ の 璛Cyphomandra betacea (Cav.) 猂硁砰ゑ耕瓜 DNASIS® 硁砰だ Sendt.堵兵狦龟们 盚从 堵葶 猂 ネ荒秈て闽玒だや 瓜瓜 狾龟们 盚从 瓣玭ナCitrullus Kimura-2-parameter 笲衡家 (23) moschata (Duchesne ex Lam.) のナ Neighbor-join 竤籈よ ―眔克絫闽 Trichosanthes cucumeroides Maxim. 玒瓜瓜 maxumum parsimony 璫(21, 32) 龟们 璶盚 ナ 瞶币祇 穓碝莉眔跋办┦程ㄎ や Luffa cylindrica (L.) Roem芖 瓜瓜竒パ瓃甅杆硁砰だ猂 Bryonopsis laciniosa Naud. の 蛮 近 ナ 狦 陪ボ ㄈ狦龟们夹 だだ摸 Diplocylos palmatus (L.) C. Jeffrey玭ナ 狦籔篈だ摸 パだだ摸 龟们 盚絛瞅珹 ナ摸璛のぶ 狦睲 groupingㄌ篈だ 计狦攫 狦龟ㄤ 9 34 贺竒筁箎▅絋 摸┮跋だ 硅を龟们ㄈの您龟们ㄈ滇 粄(5, 6) 龟们ㄈ 龟们ㄈぶを龟们壁 您龟们ㄈ 兵龟们の 盿龟们 龟们壁 龟们ㄈ ㄈの 壁 芖 λ狶跋祇ネ 筂玡 盚 Tribe 硅を龟们ㄈ DNA 从 盚从 狵λ 畉 絛瞅 0.8 %12.4 %ㄤ Bambusa dolichoclada Hayata 弟λ ㄈの壁 ┮ノ 2 贺狦龟们 畉 絛瞅 Bambusa multiplex (Lour.) Raeuschel だ 4.8 % 14.9 % 陪ボ λBambusa tuldoides Munroを厚λ 刚喷┮ノ DNA ┮陪ボ だ戈 Bambusa oldhamii Munro踠λBambusa 籔篈戈 跋狦龟们 贺┪ stenostachya Hackel陈λDendrocalamus 碞贺 克絫闽玒
贺狦龟们 畉 Table 2. Nucleotide divergence among 19 species of fruit fly Taxon1) 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Bs 1
Bt 2 0.008 寡
Bci 3 0.045 0.041 毛 實
Bsc 4 0.035 0.032 0.039 蠅 亞
Bdi 5 0.044 0.040 0.053 0.029 科 （
Bdo 6 0.054 0.047 0.052 0.054 0.055 D i p t
Bc 7 0.027 0.024 0.056 0.045 0.056 0.063 e r a :
Bf 8 0.061 0.058 0.062 0.054 0.062 0.052 0.063 T e p
Df 9 0.113 0.108 0.115 0.092 0.104 0.116 0.124 0.117 h r i t i d
Rf 10 0.127 0.119 0.135 0.131 0.123 0.132 0.143 0.143 0.172 a e :
Sa 11 0.144 0.136 0.150 0.146 0.135 0.127 0.155 0.157 0.186 0.109 D a c i
Gf 12 0.079 0.078 0.087 0.082 0.083 0.084 0.090 0.092 0.138 0.123 0.132 n a e As 13 0.082 0.079 0.096 0.085 0.082 0.078 0.092 0.099 0.137 0.130 0.118 0.048 ） 果
Ph 14 0.104 0.103 0.114 0.111 0.113 0.101 0.116 0.104 0.160 0.144 0.148 0.120 0.111 實 蠅
Asp 15 0.133 0.145 0.154 0.126 0.157 0.147 0.156 0.153 0.194 0.189 0.184 0.147 0.129 0.102 之 親
Fe 16 0.129 0.137 0.158 0.134 0.145 0.138 0.150 0.151 0.205 0.187 0.174 0.165 0.153 0.152 0.193 緣
Cm 17 0.125 0.139 0.146 0.119 0.136 0.139 0.147 0.142 0.192 0.171 0.189 0.137 0.137 0.142 0.185 0.149 關 係
Cl 18 0.110 0.124 0.143 0.095 0.121 0.130 0.142 0.132 0.178 0.151 0.167 0.139 0.119 0.122 0.165 0.188 0.142 Ea 19 0.113 0.111 0.125 0.113 0.109 0.119 0.128 0.132 0.166 0.161 0.180 0.138 0.133 0.138 0.153 0.203 0.145 0.079 1) Bs: B. synnephes, Bt: B. tau, Bci: B. cilifera, Bsc: B. scutellata, Bdi: B. diaphora, Bdo: B. dorsalis, Bc: B. cucurbitae, Bf: B. ferruginea, Df: D. formosanus, Rf: R. formosana, Sa: S. acroleuca, Gf: G. fasciventris, As: A. sexvittata, Ph: P. hexachaeta, Asp: A. speciosa, Fe: F. ensifera, Cm: C. matsumotoi, Cl: C. latilimbata, Ea: E. apicalis.
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6 植物保護學會會刊 第 45 卷 第 1 期 2003
克絫闽玒 瓜瓜の瓜 祇瞷妮 篈疭紉 克絫闽玒 癚よ パ 硅を龟们ㄈ妮 堵葶狾龟们籔堵兵龟 刚喷┮蹦ノ 硅を龟们 贺摸计ぶ度 们 だ疭紉耕ㄤ DNA 畉 度 珹 2 妮 3 ㄈ妮 9 贺珿沮 篈戈 2.9 %パ瓜瓜 陪ボ だ猂ㄤ妮ㄈ妮┪贺 克絫闽玒琌 妮璉硅を龟们ㄈ妮 狥よ狦龟们籔盽 籔だ戈 ゑ癸White(37)ノ 砆 籔ㄤ贺 ︹龟们 克絫闽 37 篈疭紉秈︽硅を龟们壁 44 贺硅を龟 玒ㄤ DNA 畉 5.2 % 们ㄤ 3 贺籔 刚喷 克絫闽 τ 18S+ITS PCR-RFLP 玒ナ龟们籔玭ナ龟们 克絫耕ㄤ RsaI 籔 AluI 芭ち 挛 ネ 狥よ狦龟们 ㄤ 龟们ㄈ妮 芭ち琿瓜眯 竒瑆 祇戈 贺摸 刚喷┮莉眔 だ戈よ パ 赣ㄢ贺 陪 だ疭紉畉 瓜瓜の瓜 祇瞷妮硅をㄈ ヘ玡计度綼ㄤ篈疭紉 妮 ナ龟们玭 ナ 龟们の 龟们 克絫闽 粄﹚ 贺 ﹚ 穝浪 玒ㄤ 龟们籔玭ナ龟们 癚Tseng (35) 盢 ㄢ贺龟们ㄌㄤ篈 DNA 畉 度 0.8 % 疭紉跋だ︹龟们 们 瓜 陪ボ 龟们籔玭ナ龟们 克絫闽 癸们癬は 籔狥よ狦龟们癸ヒ膀 玒 ㄤ籔ナ龟们 猳癬は 陪畉 玒妮 ㄤ龟パ瓃 3 贺狦龟们 篈疭紉 贺 ╯赣挛だ疭紉 陪 璝虫縒ノ篈疭紉龟 畉 ョやㄤ猭 螟皐癸ㄤ 克絫闽玒﹚ 灿盿龟们 だ疭紉瓜瓜 だ疭紉 ︴靡癸玭ナ龟 の瓜 陪ボㄤ籔 龟们玭ナ龟们 们の 龟们 篈克絫┦ 籔ナ龟们 克絫闽玒耕 τㄤ籔堵葶 秆睦ㄒ篈だ摸葶狾 狾龟们堵兵龟们 克絫闽玒ョ会 膀を 礚琌盢璉硅を龟们妮跋だ 璉 ㄤ ┺箄 眔 硅を龟们ㄈ妮の硅を龟们ㄈ妮 璶疭 琌篈だ摸 芠翴厩 癸灿盿龟 紉ナ龟们礚 を┮ 戳砆耴璉 们╯ 妮璉硅を龟们ㄈ妮┪硅を龟们 硅を龟们ㄈ妮τ玭ナ龟们の 龟们玥 ㄈ妮 ┮ (25, 35)パ瓜瓜 ㄣ を耴硅を龟们ㄈ妮 陪ボ赣贺克絫闽玒 籔硅を龟们 琌 ㄒ 琌玡 だ摸盡產(2, 36)の ㄈ妮 贺摸耕珿ョ︴靡ㄤ 硅 祇瞷ㄤ龟赣疭紉 跑 ナ を龟们ㄈ妮耕続讽 龟们 夹 ぶ计 夹 ㄣ 赣を 芖 龟们 だ疭紉瓜の瓜 硑 だ摸 耑 筁繦帝だ 陪ボㄤ克絫闽玒玒 璉硅を龟们 摸疭紉 浪癚の秈˙ヘ玡赣 2 ㄈ妮 だ 妮贺摸籔ㄤㄈ狦龟们 瓜玥 摸疭紉эノ ┦ 挛ネ崔 Ю璉 陪ボㄤ 璉硅を龟们ㄈ妮籔硅を龟们 耕祏の 材 5 狾絫 瞏 ㄈ妮 瓜 玥陪ボㄤ籔 ㄈ V 璉硅を龟们ㄈ妮疭紉は ㄤ贺摸 克絫闽玒耕 竤 玡耕度 睱 硅を龟们ㄈ 您龟们ㄈ 盿龟们壁 兵龟们㎝ 妮疭紉ㄌㄤだ疭紉戈и粄 ㄌ 盿龟们 篈だ摸盢ㄤだ 妮 疭紉盢ナ龟们灿盿龟们の堵兵龟们耴 硅を龟们妮 龟们ㄈ妮 妮続 硅を龟们ㄈ妮 だ猭ゑ耕瞶 讽ㄤ DNA 畉 11.7 % 寡毛實蠅亞科（Diptera: Tephritidae: Dacinae）果實蠅之親緣關係 7
篈だ摸赣妮 璶疭紉珹 材 1 竊 疭 て 璊噶 灿竬 璉狾 材 23 竊 耕 篈耕疭 璴紉
瓜 19 贺狦龟们采絬砰 16S rDNA Vector NTI Suite 硁砰だ猂 克絫闽玒笲 ノ Neighbor Joining method (NJ)よ猭だ猂克絫攫(24) Fig. 1. Relationships of 19 fruit flies, analyzed with the software, Vector NTI Suite. The phylogenetic tree in the Vector NTI Suite was built using the Neighbor-joining (NJ) method(24). The NJ method works on a matrix of distances between all pairs of sequences to be analyzed. These distances are related to the degree of divergence between the sequences. The phylogenetic tree was calculated after the sequences were aligned. 8 植物保護學會會刊 第 45 卷 第 1 期 2003
瓜 19 贺狦龟们采絬砰 16S rDNA DNASIS® 硁砰だ猂 克絫闽玒笲ノ UPGMA よ猭(24)秈︽ Higgins-Sharp ゑ癸ㄤ 把计 砞﹚ㄤ﹍ Gap penalty: 5, No. of top diagonals: 5, Fixed gap penalty: 10, Floating gap penalty: 10, K-tuple: 4, Window size: 5 Fig. 2. Relationships of 19 fruit flies, analyzed with the software, DNASIS® . The program takes as input a dendrogram produced by applying the UPGMA method to a matrix of similarity scores for all aligned sequences(24). The Higgins-Sharp branch-scoring operation used these default parameters (gap penalty: 5, no. of top diagonals: 5, fixed gap penalty: 10, floating gap penalty: 10, K-tuple: 4, window size: 5).
竤 だ猂よ パ瓜瓜 龟们ㄈ 龟们壁 狠フ巴 龟们㎝吊 ョ祇瞷 龟们ㄈ 龟们壁 龟龟们壁 腳畄次を龟们您龟们ㄈ 猀 龟们㎝糃 フ葶龟们ぶを龟们壁 兵龟们㎝ 盿龟们 竤ㄈ 溜絫龟们㎝狠 纔龟们滇龟们ㄈ 壁 竤籔硅を龟们 龟们壁 翴龟们㎝ を 风龟们 ㄈ 贺摸陪 瞒陪ボ赣ㄇ狦龟们摸 寡毛實蠅亞科（Diptera: Tephritidae: Dacinae）果實蠅之親緣關係 9
だ疭紉┮陪ボ 克絫闽玒籔篈だ 们㎝糃 フ葶龟们 14.9 %溜絫龟们㎝狠 摸 ョパ瓜の瓜 ㄤㄢㄢ 纔龟们 7.9 %翴龟们㎝ を 风 耕 ┺箄 玥陪 龟们 10.2 %狠フ巴 龟们㎝腳畄次を龟 ボㄤㄢㄢ DNA 畉 だ 猀 龟 们 10.9 % 兵龟们㎝ 盿龟们 4.8 %
瓜 贺狦龟们采絬砰 16S rDNA Kimura-2-parameter 笲衡家 (23)竒 Neighbor-joining 竤籈よ猭┮崩阶 克絫闽玒 Fig. 3. Phylogenetic analysis from 962 nucleotide sequences of the partial 16S rDNA of 19 species of fruit fly, constructed by Kimura 2-parameter distances(23) under the Neighbor-joining cluster method. One thousand bootstrap replications were used. 10 植物保護學會會刊 第 45 卷 第 1 期 2003
瓜 贺狦龟们采絬砰 16S rDNA 程稽 稽瞶の币祇 穓碝┮崩 阶 程ㄎや瓜 Fig. 4. Cladogram from 16S rDNA sequences of 19 species of fruit fly, constructed by maximum parsimony methods based on a heuristic search under the close-neighbor-interchange (CNI) method. One thousand bootstrap replications were used, and only consensus values greater than 50 are shown.
璝だ猂瓃竤籔硅を龟们ㄈ 瓜 盿龟们壁 赣 2 贺龟们籔 克絫闽玒パ瓜の瓜 祇瞷您龟 璉硅を龟们妮 克絫闽玒临ゑ硅を龟们 们ㄈ 盿龟们壁 兵龟们㎝ 盿龟们 妮 芖 龟们 贺 狦ョや 籔硅を龟们ㄈ 贺摸程 だ摸厩ㄤ ㄣ 2 猅纗弘舗 寡毛實蠅亞科（Diptera: Tephritidae: Dacinae）果實蠅之親緣關係 11
Spermatheca 疭紉 ㄤ 狦龟们 だや瓜(15)ョ陪ボ妮 ㄈ 3 猅τ盢您龟们ㄈㄖ硅を龟 龟们壁 Trypeta の Euleia 硂 2 妮 们壁 Dacini 盿龟们ㄈ壁 贺摸籔硅を龟们ㄈ 贺摸 克絫闽玒耕 Gastrozonina だ猭(27)ㄤ龟刘妮您 τぶを龟们壁 贺摸籔ㄤ耕 龟们ㄈ 2 壁珹 盿龟们壁の 眔 琌 龟们妮Coelotrypes 狦龟们┮妮 您龟们壁 ㄓㄤ篈疭紉籔纔龟们妮 Ceratitidini篈のだ疭紉ョ Euphrantaだ摸厩 粄 陪ボㄤ籔硅を龟们摸 耕 克絫闽玒(15, ㄢ ㄖ 妮(29) ╯ 16, 36) 猵 荷礛眖瓜の瓜 だ戈陪ボ赣 2 贺龟们 DNA 祇瞷 兵龟们㎝ 盿龟们 籔狠フ 畉 7.9 %珿盢ㄤ縒ミ 巴 龟们㎝腳畄次を龟们 龟们ㄈ 龟们妮ョ礚 讽 よ 盢ㄓゑ耕赣 克絫闽玒ョ瘤礛ㄤ┺箄 ㄢ妮 贺摸 だ戈络睲 珿璝ㄌ だ疭紉 だ猂 狦ョ ┮ 竤 璝 盢 戈 癸盢您龟们ㄈ 縒ミ ㄈ だ猭 衡ㄈ┮贺摸籔硅を龟们ㄈ 8 贺 ︴靡 秈˙ 络睲╯ 龟们 DNA 畉 祇瞷 龟们ㄈ ㄢ贺だ摸よ 耕続讽 惠だ 龟们壁 2 贺龟们 13.3 %-15.0 % 猂 DNA 戈 龟们ㄈ 13.3 %-14.5 % 龟们ㄈ 龟们ㄈ 龟们壁 克絫闽 ぶを龟们壁 13.3 %-13.6 %滇龟们ㄈ 玒よ Han(12) ノ 35 篈疭紉秈︽ 12.1 %-15.5 % 盿龟们壁 9.4 %-9.6 赣壁 33 妮 龟们 克絫闽玒╯粄 % 刚喷 狦籔 Han and 赣壁だ 2 ㄈ壁 龟们ㄈ壁 McPheron(15)┮厨旧妓だ猂 16S rDNA Trypetina の 龟们ㄈ壁 ┮莉眔 狦龟们 克絫闽玒 狦 Chetostomatina 刚喷 ┮ㄏノ 糃 パだ戈 靡沮祇瞷 龟们ㄈ ぶ 龟们妮Fusciludia の 絫 龟们妮 を龟们壁籔硅を龟们ㄈ耕 τㄤ Carpophthoracidia 龟们ㄈ壁 龟们壁玥耕 赣ㄢ壁だ摸 贺摸瓃厨 碞 龟们壁 虫 竤竤 您龟们ㄈ 盿 克絫闽玒 癚パ瓜 祇 龟们壁籔硅を龟们ㄈ 闽玒程 瞷赣ㄈ ぶを龟们壁の 龟们壁 だ克絫闽玒篈だ摸盢ㄤ耴 阶 ㄈ 妮瞶琌瓜瓜 の瓜 玥陪ボぶを龟们壁 贺摸 狦龟们 だ摸ノ篈疭紉 だ だ克絫闽玒玒 翴龟们㎝ を 摸パ贺 畉 ㄓㄤ 风龟们滇龟们ㄈ籔 龟们壁 だ摸疭紉硋 绰 灿稬 篶糤 贺摸 ㄤ龟ネ篈の盚絛瞅疭紉 ﹚ 螟硓筁だ戈 ┦τ 粄 ぶを龟们 贺摸籔 ミ 度 ど ﹚ 硉の非絋 滇龟们ㄈ 贺摸 翴籔ノ采絬 ┦ノだ戈秈︽狦龟们摸克絫闽玒 砰 赁砰 DNA ┮ 狦龟们 だ ╯ョミ讽 狦 竒パだ や瓜(15) 狦琌璓 珿 粄 疭紉 癸阶靡狦龟们琘ㄇ 龟们ㄈㄓ獶虫 竤 篈疭紉 だ摸疭紉 瞶┦ョ ノ采絬砰 赁砰 DNA ┮ 讽 痲 ゅ┮秈︽だ戈だ猂 12 植物保護學會會刊 第 45 卷 第 1 期 2003
狦龟们贺摸度琌赣贺摸 ぶ计硓 5. 糂ド 狶键2000 玭ナ龟们 筁 贺疭紉 癸崩﹚狥よ狦龟们 (Bactrocera tau)蛮ヘ狦龟们 贺 贺 篈の絋粄 龟们妮 篈祇▅关㏑のㄤユЮ︽ だ摸 祇 地挛 20: 311-25 ㄓ硓筁闽戈 ミの籔瓣闽 6. Allwood, A. J., Chinajariyawong, A., ╯虫 ユ瑈癸 ╯狦龟们ㄈ妮 Drew, R. A. I., Hamacek, E. L., Hancock, ┪妮 克絫闽玒籔络睲ㄤ篈だ摸 D. L., Hengsawad, C., Jipanin, J. C., 瞺繴箇戳盢穦 Jirasurat, M., Kong, C., Krong, C., Kritsaneepaiboon, S., Leong, C. T. S., 谅 勉 and Vijaysegaran, S. 1999. Host plant records for fruit flies (Diptera: ┯籜芖厩挛╰砛瑌紋ゅ Tephritidae) in South East Asia. Raffles 毙甭旧の 蝴の㏄ゅネ蹦 Bull. Zool. Suppl. 7, 92 pp. 夹 洛厩厩 ゅ賧瞶毙甭 7. Armstrong, K. F., Cameron, C. M., and 旧笰〆笰穨媚 瑀 刚喷┮ Frampton, E. R. 1997. Fruit fly (Diptera: 睶﹋蹦 夹 のド礮 Tephritidae) species identification: a ﹋だ猂略 ㄖ粁谅 rapid molecular diagnostic technique for quarantine application. Bull. Entomol. まノゅ膍 Res. 87: 111-118. 8. Chang, H. Y., Hsu, T. C., and Wu, W. J. 1. 眎 驾2001芖 λ狶 笆 2001. Species diversity and seasonal 狦龟们 贺摸妓┦のㄤ﹗竊跑笆 fluctuation of fruit flies (Diptera: 蛮ヘ狦龟们瓣ミ芖厩 Tephritidae) in bamboo stands in Taipei. 从 痜挛甡厩╯┮痴阶ゅ Formosan Entomol. 21: 47-64. 362 9. Drew, R. A. J., and Hancock, D. L. 1994. 2. 竡 狶穝1974芖狦龟们 The Bactrocera dorsalis complex of fruit 糵﹚(I)狦龟们竒蕾 坝珇浪喷 flies (Diptera: Tephritidae) in Asia. Bull. 浪喷╯盡 56 Entomol. Res. Suppl. 2, 68 pp. 3. 竡 眎 驾1998硅を狦龟们ㄈ 10. Drew, R. A. J., and Hancock, D. L. 1999. 瞶だ の从 浪 璶 Phylogeny of the tribe Dacini (Dacinae) 贺摸痹瓃芖狦龟们 獀м砃癚穦 based on morphological, distributional, 盡70-129 糂ド 琇秜 絪 and biological data. pp. 491-533. In: 胯瓣ミ 厩挛厩╰絪 Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior. CRC Press LLC. 4. ゅ賧 ヲ瓜砛 猭1999ㄌ采 944 pp. 絬砰 16S 赁砰赁荒だ 11. Fletcher, B. S. 1987. The biology of 猂您峦羆摸絫闽玒挛だ摸の秈て dacine fruit flies. Annu. Rev. Entomol. 癚穦盡57-82 耙 絪胯 32: 115-144. 瓣ミ 厩挛厩╰芖ミ痴 12. Han, H. Y. 1992. Classification of the 繻絪 tribe Trypetini (Diptera: Tephritidae: 寡毛實蠅亞科（Diptera: Tephritidae: Dacinae）果實蠅之親緣關係 13
Trypetinae). Ph.D. thesis, The between strains of the Mediterranean Pennsylvania State University, 274 pp. fruit fly, Ceratitis capitata detected by 13. Han, H. Y. 2000. Molecular phylogenetic DNA fingerprinting. Genome 35: study of the tribe Trypetini (Diptera: 528-533. Tephritidae), using mitochondrial 16S 20. Haymer, D. S., Tanaka, T., and Teramae, ribosomal DNA sequences. Biochem. Syst. C. 1994. DNA probes can be used to Ecol. 28: 501-513. discriminate between Tephritid species 14. Han, H. Y., and McPheron, B. A. 1994. at all stages of the life cycle (Diptera: Phylogenetic study of selected tephritid Tephritidae). J. Econ. Entomol. 87: flies (Insecta: Diptera: Tephritidae) 741-746. using partial segments of the nuclear 18S 21. Ito, S. 1983-85. Die Japanischen ribosomal DNA. Biochem. Syst. Ecol. 22: Bohrfliegen. Maruzen Co., Ltd., Osaka. 447-457. 352 pp. 15. Han, H. Y., and McPheron, B. A. 1997. 22. Iwaizumi, R., Kaneda, M., and Iwahashi, Molecular phylogenetic study of O. 1997. Correlation of length of Tephritidae (Insecta: Diptera) using terminalia of males and females among partial sequences of the mitochondrial nine species of Bactrocera (Diptera: 16S ribosomal DNA. Mol. Phylogenet. Tephritidae) and difference among Evol. 7: 17-32. sympatric species of B. dorsalis complex. 16. Han, H. Y., and McPheron, B. A. 2000. Ann. Entomol. Soc. Am. 90: 664-666. Nucleotide sequence data as a tool to test 23. Kimura, M. 1980. A simple method for phylogenetic relationships among higher estimating evolutionary rate of base groups of Tephritidae: a case study using substitutions through comparative mitochondria ribosomal DNA. pp. studies of nucleotide sequences. J. Mol. 115-131. In: Fruit Flies (Tephritidae): Evo. 16: 111-120. Phylogeny and Evolution of Behavior. M. 24. Kumar S., Tamura K., Jakobsen I. B., Aluja and A. L. Norrbom, eds. CCRC and Nei, M. 2001. MEGA2: molecular Press. Boca Raton, Florida. evolutionary genetics analysis software. 17. Hardy, D. E. 1973. The fruit flies Bioinformatics 17: 1244-1245. (Tephritidae-Diptera) of Thailand and 25. Liang, G. Q., Hancock, D. L., Xu, W., bordering countries. Pacific Insects and Liang, F. 1993. Notes on the Monograph 31: 1-353. Dacinae of Southern China (Diptera: 18. Haymer, D. S. 1995. Genetic analysis of Tephritidae). J. Aust. Entomol. Soc. 32: laboratory and wild strains of the melon 137-140. fly (Diptera: Tephritidae) using random 26. McPheron, B. A., and Han, H. Y. 1997. amplified polymorphic DNA-polymerase Phylogenetic analysis of North American chain reaction. Ann. Entomol. Soc. Am. Rhagoletis (Diptera: Tephritidae) and 88: 705-710. related genera using mitochondria DNA 19. Haymer, D. S., McInnis, D. O., and sequence data. Mol. Phylogenet. Evol. 7: Arcangeli, L. 1992. Genetic variation 1-16. 14 植物保護學會會刊 第 45 卷 第 1 期 2003
27. Norrbom, A. L., Carroll, L. E., and 33. Sueyoshi, M. 1999. Immature stages of Friedberg, A. 1998. Status of knowledge. three Oriental species of the genus pp. 9-48. In: F. C. Thompson ed. Fruit Rhabdochaeta de Meijere (Diptera: Fly Expert Identification System and Tephritidae), with brief biological notes. Systematic Information Database. Myia Entomol. Sci. 2: 217-230. 9. Leiden, Netherlands. 534 pp. 34. Tseng, Y. H., and Chu, Y. I. 1983. A 28. Palacio, I. P., Adorada, J. L., and Mora, new fruit fly Callentra formosana from J. G. 1992. Tephritids infesting Eclipta Taiwan (Tephritidae, Diptera). Chinese J. alba (L.) Hassk. The Phillipine Entmol. Entomol. 3: 119-122. 8: 1189-1192. 35. Tseng, Y. H., Chen, C. C., and Chu, Y. I. 29. Permkam, S., and Hancock, D. L. 1992. The fruit flies, genus Dacus 1995. Australian Trypetinae (Diptera: Fabricius of Taiwan (Diptera: Tephritidae). Invertebr. Taxon. 9: Tephritidae). J. Taiwan Mus. 45: 15-91. 1047-1209. 36. Wang, X. J. 1996. The fruit flies 30. Sheppard, W. S., Steck, G. J., and (Diptera: Tephritidae) of the East Asian McPheron, B. A. 1992. Geographic region. Acta Zootaxon. Sinica 21, populations of the medfly may be Supplement. 338 pp. differentiated by mitochondrial DNA 37. White, I. M. 1999. Morphological variation. Experientia 48: 1010-1013. features of the tribe Dacini (Dacinae): 31. Shiraki, T. 1933. A systematic study of their significance to behavior and Trypetidae in the Japanese Empire. Mem. classification. pp. 505-533. In: F. C. Fac. Sci. Agric. Taihoku Imp. Univ. 8. Thompson ed. Fruit Fly Expert 509 pp. Identification System and Systematic 32. Shiraki, T. 1968. Fruit flies of Ryukyu Information Database. Myia 9. Leiden, Islands. U.S. Natl. Mus. Bull. 263: Netherlands. 534 pp. 1-104. 寡毛實蠅亞科（Diptera: Tephritidae: Dacinae）果實蠅之親緣關係 15
ABSTRACT
Chang, H. Y.1*, Tzeng, C. C.2, Tsai, W. H.3, Kao, S. S.2, and Tseng, Y. H.4 2003. Phylogenetic study of the Dacinae (Diptera: Tephritidae) using partial sequences of mitochondrial 16S ribosomal DNA. Plant Prot. Bull. 45: 1- 15. (1Bureau of Animal and Plant Health Inspection and Quarantine, Council of Agriculture, Taipei, Taiwan 100, ROC; 2Department of Bio-pesticide, Taiwan Agricultural Chemicals and Toxic Substances Research Institute, COA, Wufeng, Taichung, Taiwan 413, ROC; 3Department of Entomology, National Taiwan University, Taipei, Taiwan 106, ROC; 4Kaohsiung Branch, Bureau of Standards, Metrology and Inspection, MOEA, Kaohsiung, Taiwan 802, ROC)
Nineteen species of fruit flies trapped from northern and central Taiwan were examined, including 9 species of the tribe Dacini, Bactrocera dorsalis, B. ferruginea, B. cilifera, B. cucurbitae, B. diaphora, B. scutellata, B. synnephes, B. tau, and Dacus formosonus, and 10 outgroup taxa from the other 4 subfamilies of the Tephritidae, Acrotaeniostola sexvittata, Gastrozona fasciventris, Acanthonevra speciosa, Phorelliosoma hexachaeta, Euphranta apicalis, Coelotrypes latilimbata, Carpophthoracidia matsumotoi, Fusciludia ensifera, Spathulina acroleuca, and Rhabdochaeta formosana. Sequence data from about 900 bp within the mitochondrial 16S ribosomal DNA were amplified by polymerase chain reaction and sequenced using the dideoxy chain determination method. Based on sequence comparisons, phylogenetic relationships among the included taxa were inferred using Neighbor-joining and maximum parsimony methods. Trees were congruent with the morphological classification and suggested the following phylogenetic relationships: (1) monophyly of the tribe Dacini including Bactrocera and Dacus; (2) a sister group relationship between the tribes Gastrozonini and Dacini; (3) respective monophyly of the tribe Adramini, tribe Trypetini, and tribe Acanthonevrini, and a possible relationship between them and the tribe Dacini remain to be clearified; and (4) the non-monophyletic nature of the Trypetinae between the tribes Adramini and Trypetini. From 16S ribosomal DNA data, we also propose that (1) B. ferruginea is a definite species, and its previous combination as a synonym of B. dorsalis should be revised; and (2) Coelotrypes can be maintained as a separate genus distinct from Euphranta.
(Key words: Tephritidae, Dacinae, 16S rDNA, phylogeny)
*Corresponding author. E-mail: [email protected]