台灣產銀腹蛛屬(蜘蛛目：長腳蛛科)蜘蛛之系統分類研究

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(1)國立台國立台灣師範大學生命科學系碩士論文灣師範大學生命科學系碩士論文. 台灣產銀腹蛛屬(蜘蛛目台灣產銀腹蛛屬蜘蛛目：蜘蛛目：長腳蛛科)蜘蛛長腳蛛科蜘蛛系統分類研究之系統分類研究 A Taxonomic Study on the Spider Genus Leucauge (Aranea: Tetragnathidae) of Taiwan. 研究生：劉錫軒 Hsi-hsuan Liu 指導教授：陳世煌博士 Shyh-Hwang Chen. 中華民國九十八年一月.

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(4) 致謝. 在本論文付梓之際，要感謝的人很多，以此文表示我的誠摯謝意。承蒙恩師陳世煌博士給我的悉心指導，引導我從陌生到熟悉蜘蛛多樣性的世界，也從老師身上學習到嚴謹的治學態度及科學研究精神。感謝口試委陳順其老師及徐堉峰老師在忙碌的研究之中，費心審閱本論文，提供許多寶貴的意見，使本論文能更趨完整而嚴謹。論文研究期間，感謝蜘蛛實驗室的文俊學長、宏仁學長、鈞漢學長、鈺婷學姊、天勻、英元、雅惠及宸瑜在野外工作上提供許多的協助，並且陪伴我到各地採集，典諺、嘉容、珞璿及奕儂在我口試時辛苦地為我準備器材；感謝王穎老師及實驗室同學玉潔、詩涵、士清、家宏等提供登山器材的協助；感謝王震哲老師及實驗室長澤學長、和明學長、淑娟學姊、怡靜、鈴雅、杏倩在器材及演化軟體上的幫助，依恆的爸爸幫忙登山的接駁；感謝徐堉峰老師實驗室的嘉龍學長在野外的照顧，至堅學長、立偉學長給我的建議，育綺幫忙寫標本的標籤。感謝好友至軒、霆翰、明聰、明福、伯寬學長及培鈞學長在生活中給我的鼓勵並且常陪我吃飯，煒婷辛苦地幫忙標本的打字工作。實驗室外，感謝台北靈糧堂的朋友淑惠姊、如婷、佑庭、娟娟、Fancie、Jennifer、 Ivy 等好友的支持打氣；公舘長老教會的張朝棟牧師及龔師母在生活上給予的關心；文山長老教會的郁屏及契友們的陪伴，及蘭陽靈糧堂五二小組弟兄姊妹的支持與代禱。最後，由衷地感謝我的父母親及家人的和鼓勵，給我機會唸書及經濟上無條件的支持。論文完稿之際，謹在此將它獻給最疼愛我的天父。. 衷心感謝所有關心及愛護我的人，對你們致上最深的祝福！.

(5) 目錄目錄............................................................................................................i 圖目次….…………………………………………………….………….iii 表目次………………………………………………………….……..….v 附錄目次………………………………………………………….….….vi 中文摘要………………….……………………………………….……vii 英文摘要…..…………..……………….………………………………viii 壹、前言…………………………………..………………………...…….1 貳、研究材料與方法…………………..………….……………………...7 一、研究材料…………………..……….….………………………...7 二、研究方法…………………..………….………………………..13 三、支序分析…………………..………….………………………..15 參、親緣關係分析…………………..……………………….………….18 一、類群..…………...………………………………………………18 二、特徵描述…...………………………………...………………...19 三、結果…...………..………………………………………………26 四、討論…...…………………..……………………………………27 肆、分類處理…...…………………………………………………...…31 Leucauge argentina (Hasselt, 1882) 雪銀腹蛛…..…….…..….....35 Leucauge blanda (L. Koch, 1878) 肩斑銀腹蛛……………..…...38 Leucauge celebesiana (Walckenaer, 1842) 西里伯銀腹蛛…..…..44 Leucauge crucinota (Bösenberg et Strand, 1906) 十字銀腹蛛….54 Leucauge decorata (Blackwall, 1864) 尖尾銀腹蛛…………...…58 Leucauge subblanda Bösenberg et Strand, 1906 小肩斑銀腹蛛...63 Leucauge taiwanensis sp. nov. 台灣銀腹蛛………………..……68 i.

(6) Leucauge tessellata (Thorell, 1887) 方格銀腹蛛………………..72 Leucauge wulingensis Song et Zhu, 1992 武陵銀腹蛛…….…....79 伍、參考文獻……………………………………………………………83 附圖…….….……………………………………………………………95 附表…...….……………………………………………………………128 附錄…….……………………………………………………………129. ii.

(7) 圖目次. Figure 1. External features of genus Leucauge…………………………95 Figure 2. External features and copulatory organs of genus Leucauge....96 Figure 3. Variations of male left palp of Leucauge from Taiwan……….97 Figure 4. Variations of female epigynum of Leucauge from Taiwan…...98 Figure 5. The most parsimonious cladograms……….….…………..…..99 Figure 6. The most parsimonious cladograms with bootstrap value......100 Figure 7. Leucauge argentina (Hasselt, 1882), female……...……...…101 Figure 8. Leucauge argentina (Hasselt, 1882), male…………………102 Figure 9. Leucauge blanda (L. Koch, 1878), female………………….103 Figure 10. Leucauge blanda (L. Koch, 1878), male………………..….104 Figure 11. Leucauge celebesiana (Walckenaer, 1842), female……..…105 Figure 12. Leucauge celebesiana (Walckenaer, 1842), male…….....…106 Figure 13. Leucauge crucinota (Bösenberg et Strand, 1906), female....107 Figure 14. Leucauge crucinota (Bösenberg et Strand, 1906), male...…108 Figure 15. Leucauge decorata (Blackwall, 1864), female….............…109 Figure 16. Leucauge decorata (Blackwall, 1864), male..............…..…110 Figure 17. Leucauge subblanda Bösenberg et Strand, 1906, female….111 Figure 18. Leucauge subblanda Bösenberg et Strand, 1906, male....…112 Figure 19. Leucauge taiwanensis sp. nov., female..…...………………113 Figure 20. Leucauge taiwanensis sp. nov., male..……………………..114 Figure 21. Leucauge tessellata (Thorell, 1887), female……………….115 Figure 22. Leucauge tessellata (Thorell, 1887), male…………...…….116 Figure 23. Leucauge wulingensis Song et Zhu, 1992, female…...…….117 Figure 24. Leucauge wulingensis Song et Zhu, 1992, male……..…….118 iii.

(8) Figure 25. Collecting locations of Leucauge argentina in Taiwan….…119 Figure 26. Collecting locations of Leucauge blanda in Taiwan…….…120 Figure 27. Collecting locations of Leucauge celebesiana in Taiwan….121 Figure 28. Collecting locations of Leucauge crucinota in Taiwan….…122 Figure 29. Collecting locations of Leucauge decorata in Taiwan..……123 Figure 30. Collecting locations of Leucauge subblanda in Taiwan...…124 Figure 31. Collecting locations of Leucauge taiwanensis in Taiwan….125 Figure 32. Collecting locations of Leucauge tessellata in Taiwan….…126 Figure 33. Collecting locations of Leucauge wulingensis in Taiwan.…127. iv.

(9) 表目次. Table 1. Materials used in cladisitc analysis..……..…………………...128 Table 2. Data matrix used for cladistic analysis......…………………...130. v.

(10) 附錄目附錄目次. Appendix 1. Collect Locations and Collectors in Chinese and in English... .…………………………………………………………131 Appendix 2. Checklist of Genus Leucauge of Taiwan………..…….…137 Appendix 3. Diagnosis of Leucauge species of Taiwan……………….138 Appendix 4. Color Pictures of Genus Leucauge of Taiwan………….139. vi.

(11) 摘要. 本論文共報告台灣產銀腹蛛屬蜘蛛 9 種，其中新種有台灣銀腹蛛 (Leucauge taiwanensis sp. nov. )；新紀錄種有 4 種：西里伯銀腹蛛(L. celebesiana)、十字銀腹蛛(L. crucinota)、小肩斑銀腹蛛(L. subblanda)、武陵銀腹蛛(L. wulingensis)。文中將大銀腹蛛(L. magnifica)處理為西里伯銀腹蛛的同物異名；擬方格銀腹蛛(L. subtessellata)處理為方格銀腹蛛(L. tessellata)的同物異名。脈斑銀腹蛛(L. venusta)，處理為排除種。並以 19 種銀腹蛛亞科類群為內群，長腳蛛屬為外群，以 45 個形態特徵進行銀腹蛛屬的支序分析，台灣產銀腹蛛屬可分為兩個亞群，其中十字銀腹蛛與雪銀腹蛛(L. argentina)成為一單系群，可能屬於未知的新屬，而隨蛛屬(Opadometa)蜘蛛與其餘的台灣產銀腹蛛屬蜘蛛被歸入另一個亞群，其姊妹群為天星蛛屬(Mesida)。結果顯示現有的台灣產銀腹蛛屬為一複系群。本研究結果將可增進台灣銀腹蛛屬蜘蛛資源和物種多樣性之知識基礎。. 關鍵詞：蜘蛛目、長腳蛛科、銀腹蛛屬、系統分類、親緣、新種、新記錄種、台灣。. vii.

(12) Abstract. Nine spiecies of Leucauge spiders are reviewed from Taiwan in this study, among which one species, L. taiwanensis sp. nov is new to the science. L. celebesiana (Walckenaer, 1842), L. crucinota (Bösenberg & Strand, 1906), L. subblanda Bösenberg & Strand, 1906, and L. wulingensis Song & Zhu, 1992, are newly recorded from Taiwan. L. venusta was excluded from the spider fauna of Taiwan. Forty-five morphological characters of 19 leucaugines species and Tetragnatha as Outgroup were included in a phylogentic analysis. The results showed that Leucauge can be divided into two subgroups. L. argentina and L. crucinota belong to a monophyletic group, which might belong to a new genus. The other Leucauge species and Opadometa form another subgroup with Mesida as a sister group. The results suggest that genus Leucauge is a polyphyletic assamblage. A further classification study is needed. The present study will provide a basic knowledge for the diversity conservation of spiders in Taiwan.. Key Words: Araneae, Tetragnathidae, orchid spider, Leucauge, taxonomy, phylogenetics, systematics, new species, new record, Taiwan.. viii.

(13) 壹、前言. 蜘蛛在全世界目前大約有 109 科，3694 屬，40000 種，而長腳蛛科(Tetragnathidae)目前記錄有 48 屬 938 種(Platnick, 2008)，其中銀腹蛛屬(Genus Leucauge)蜘蛛，中國大陸又稱做銀鱗蛛，為全世界廣泛分布、種類最豐富的類群之一，目前記錄有 173 種(Platnick, 2008)。長腳蛛科的蜘蛛經常倒掛在平面空心圓網的中心，或棲息在蛛網附近的植被，多在溪河、農田或是樹林等較潮溼的植被上結網。其空心圓網具有少許的輻絲及螺旋狀的橫絲，與圓蛛科(Araneidae)蜘蛛所結的圓網類似，但長腳蛛科雄性觸肢之中突(median apophysis)闕如及引導器(conductor)包覆插入器(embolus)呈螺旋狀(Hormiga et al., 1995) 等特徵可以和圓蛛科區別。銀腹蛛為長腳蛛科中常被研究的類群之一，其成員的第 IV 對步足腿節背側具有二列細長的聽毛(trichobothria) 為鑑定本屬最重要的特徵(Comstock, 1940; Fitch, 1963; Levi, 1980; Hormiga et al., 1995)；腹部背面常有葉脈狀深色的縱線，並帶有銀白色金屬光澤，具有吸引昆蟲的功能(Tso et al, 2006)。雌蛛的受精囊 (spermatheca)呈薄壁狀，大多具有三個腔室，但少數種類，如十字銀腹蛛(L. crucinota Bösenberg et Strand, 1906)僅具一個腔室；雄蛛的觸肢構造簡單，主要由包覆盤曲精導管的球狀盾片(tegulum)、副盾片. 1.

(14) (subtegulum)及基部血囊(basal hematodocha)所構成，其插入器通常被引導器所包圍 (Levi, 1980; Tanikawa, 2001; Zhu et al., 2003)。長腳蛛科最早由 Menge 在 1866 年建立(Menge, 1866; Levi, 1986)。法國蜘蛛學家 Simon (1892)在 Historie Naturelle de Araignées 中，將長腳蛛科中的各屬歸在金蛛科(Argiopidae)下的長腳蛛亞科 (Tetragnathinae)和絡新婦蛛亞科(Nephilinae)之中。Roewer(1942)的蜘蛛名錄中，長腳蛛亞科被獨立為科，但將后蛛亞科(Metinae)、和絡新婦蛛亞科放在圓蛛科(Araneidae)之下。Kaston (1948)，以腿節上聽毛 (trichobothria)的存在、螯肢上的外踝(boss)消失或退化、及巨大的螯肢等特徵，將長腳蛛科的各屬從圓蛛科中分別出來，但后蛛屬(Meta) 仍被歸於圓蛛科中。Comstock (1940)在金蛛科(Argiopidae)之下，以胃外溝(epigastric furrow)之彎曲或平直，將長腳蛛亞科(Tetragnathinae) 和后蛛亞科(Metinae)分成二個亞科，其中銀腹蛛屬因其存有退化的外踝(rudimentary external condyle)、胃外溝平直等特徵，被放在后蛛亞科之中。Bonnet (1956, 1957, 1958, 1959)將所有長腳蛛科的各屬放入金蛛科。Locket et al. (1974)將歐洲產的后蛛屬蜘蛛放入長腳蛛科。 Levi (1980)將北美的長腳蛛分為后蛛亞科和長腳蛛亞科，但仍歸在圓蛛科中。Brignoli (1983)將長腳蛛處理為后蛛科(Metidae)及長腳蛛科 (Tetragnathidae)二個類群，並把絡新婦蛛亞科歸於圓蛛科。Levi (1986) 2.

(15) 和 Coddington (1986)的研究認為圓網是一個原始的特徵，因此不再把圓蛛科和長腳蛛科合併。Davies (1988)以檢索表的方式將長腳蛛科分為 6 個亞科，包含：長腳蛛亞科、坊螯蛛亞科(Phonognathinae)、長螯蛛亞科(Dolichognathinae)、銀腹蛛亞科(Leucauginae)、后蛛亞科、和絡新婦亞科。Coddington (1990)依據圓蛛總科(Araneoidae)15 個類群雄性觸肢的特徵，利用支序分析(cladistics)的原理和方法，以四個可能的共衍徵(synapomorphies)：后蛛類觸肢(Metine palp)的存在、副跗舟 (paracymbium)的形狀、盾片精導管呈之字形(switchback)和中突 (median apophysis)消失等，支持長腳蛛科為一單系群。Hormiga et al. (1995)選用 60 個特徵，包含行為、蛛網和紡器等特徵，結果同樣支持長腳蛛科為單系群，並支持長腳蛛亞科 + 絡新婦蛛亞科為單系群，且認為后蛛亞科(Metinae)為並系群(paraphyletic)，建議將銀腹蛛屬從后蛛亞科中分出來，成為銀腹蛛亞科(Leucauginae)。Griswold et al. (1998)選用 93 個特徵，在圓蛛總科的 12 個科中選取 31 個代表屬進行支序分析，以具有引導器(conductor)包圍插入器(embolus)、插入器 -盾片膜(embolus-tegulum membrane)的存在和中突的消失等可能的共衍徵，支持長腳蛛科為單系群，包含長腳蛛亞科、后蛛亞科及絡新婦蛛亞科等三亞科。而銀腹蛛屬、后蛛屬、長腳蛛屬(Tetragnatha)和骨螯蛛屬(Glenognatha)，則被歸在長腳蛛亞科之內。谷川明男(Tanikawa, 3.

(16) 2001)以 20 個形態特徵，認為銀腹蛛屬的姊妹群為沖繩蛛屬 (Okileucauge) + 高腹蛛屬(Tylorida) + 天星蛛屬(Mesida)。朱明生等 (Zhu et al., 2003)，選取中國的長腳蛛科 20 屬的 45 個特徵進行支序分析，將中國的長腳蛛科分為六個亞科：桂齊蛛亞科(Guizygiellinae)、后蛛亞科、絡新婦蛛亞科、長腳蛛亞科、銀腹蛛亞科和鋸螯蛛亞科 (Dyschiriognathinae)。其中銀腹蛛亞科代表了受精囊薄壁化的演化方向，包括臥龍蛛屬(Wolongia)、長蹠蛛屬(Metleucauge)、高腹蛛屬、波斑蛛屬(Orsinome)、隨蛛屬(Opadometa)、銀腹蛛屬、天星蛛屬和沖繩蛛屬等八屬。Kuntner (2006)選取 32 個屬，197 個特徵，對絡新婦蛛亞科進行分析，結果顯示絡新婦蛛亞科為金蛛科的姊妹群，因此將絡新婦蛛亞科獨立為絡新婦科。Pan et al. (2004)利用 rRNA and MaSp1 基因的 DNA 序列對絡新婦屬(Nephila)親緣關係進行分析，認為絡新婦屬與圓蛛科的親緣關係較長腳蛛科來得近。Álvarez-Padilla (2007) 針對長腳蛛科的 38 個類群，以 105 個形態和行為的特徵進行支序分析，將傳統上的后蛛亞科(metines)重新畫分為兩個類群：「銀腹蛛類群(Leucaugines)」與「嚴格的后蛛類群(metines sensu stricto)」，其中銀腹蛛類群包含：阿齊爾蛛屬(Azilia)、波斑蛛屬、高腹蛛屬、銀腹蛛屬、Metabus、天星蛛屬和隨蛛屬等六屬，並以四個共衍徵：受精囊的薄壁化且有膨大的現象、精導管呈之字形、盾片與副盾片易位 4.

(17) (tegulum ectally displaced)，支持銀腹蛛類群為單系，但隨蛛屬的支持度很低，因此仍不是穩定的分類群。有關於台灣地區銀腹蛛屬的研究歷史，最早的記錄是日人中島利重(Nakajima, 1921) 於台北新公園(現址：台北市二二八紀念公園)蜘蛛名錄中記載了肩斑銀腹蛛(L. blanda L. Koch, 1878)。齋藤三郎(Saito, 1933) 記述分布於台灣北部的脈斑銀腹蛛(L. venusta Walckenaer, 1842)，但僅有外雌器的外部構造，並無身體或觸肢的詳細構造以及描述。由於脈斑銀腹蛛目前只分布於美洲地區從美國到巴拿馬一帶 (Platnick, 2008; Zhu et al., 2003)，Song et al. (1999)已將其視為一疑問種。萱嶋泉(Kayashima, 1943)首次描繪了大銀腹蛛(L. magnifica Yaginuma, 1954)，但僅有身體的外形圖，沒有拉丁學名。李長林(Lee, 1964)重新描述台灣地區的肩斑銀腹蛛、脈斑銀腹蛛和大銀腹蛛。 Chang (1970)亦描述了肩斑銀腹蛛和脈斑銀腹蛛之身體特徵，但脈斑銀腹蛛並沒有附圖。朱耀沂與大熊千代子(Chu and Okuma, 1970)首次記錄台灣的尖尾銀腹蛛(L. decorata Blackwall, 1864)，並重新描述肩斑銀腹蛛。陳世煌(1996)《A Checklist of Spiders in Taiwan》中，共記載了四種：肩斑銀腹蛛、尖尾銀腹蛛、大銀腹蛛和脈斑銀腹蛛。此外，陳世煌(Chen, 1997)首次記錄台灣的端斑銀腹蛛(L. termisticta Song et Zhu, 1992)，但後來經 Zhu et al. (2003)修訂為方格銀腹蛛(L. tessellata 5.

(18) Thorell, 1887)的異名。卓逸民與谷川明男(Tso and Tanikawa, 2000)首次記錄雪銀腹蛛(L. argentina Hasselt, 1882)。Zhu et al. (2003)在《中國動物誌》中，認為卓逸民與谷川明男(Tso and Tanikawa, 2000)所記述的方格銀腹蛛為鑑定錯誤，重新命名為擬方格銀腹蛛(L. subtessellata Zhu, Song et Zhang, 2003)；此外，Zhu et al. (2003)並認為大銀腹蛛可能為西里伯銀腹蛛(L. celebesiana Walckenaer, 1842)之同種異名，但仍暫定為有效種名。而 Zhu et al. (2003)也認為千國安之輔(Chikuni, 1989) 在《Pictorial Encyclopedia of Spiders in Japan》書中所提供的大銀腹蛛的外雌器和觸肢的圖片，可能為西里伯銀腹蛛。銀腹蛛屬在中國目前記錄有 21 種(Zhu et al., 2003)，而日本有 6 種(Tanikawa, 2007)，台灣目前記錄也有 6 種(Zhu et al., 2003)。但根據平時的野外採集及檢視國立台灣師範大學生命科學系蜘蛛研究室保存的標本，均發現許多前人文獻中所未記錄的種類。因此本論文主要目的為整理前人文獻，加上個人採集及檢視研究室標本，對台灣的銀腹蛛屬種類進行整理修訂，描述其形態特徵並重新繪圖與測量，以了解台灣產銀腹蛛屬的物種多樣性，並以支序分析的方式，探討銀腹蛛屬種內的親緣關係，建立台灣地區銀腹蛛屬各物種間的親緣關係及其地理分布，可作為日後相關研究和蜘蛛保育之基礎。. 6.

(19) 貳、材料與方法. 一、研究材料：本研究所使用的新鮮材料，主要參考館藏的採集紀錄及相關文獻後，至全省各地採集，採集時以徒手為主，以目視搜尋樹葉樹叢間結網的銀腹蛛，發現蜘蛛則以空底片盒捕捉；標本以 70%酒精固定保存，存放於國立臺灣師範大學生命科學系標本館(NTNUB-Ar)。此外並檢視國立台灣師範大學生命科學系館藏蜘蛛標本。 (一)、生活習性：銀腹蛛在台灣從平地到高海拔山區皆可發現，所結的網為水平或稍傾斜的空心圓網，廣泛地分布在稻田、灌叢、和樹林等較潮溼的地方。平時大多棲息在蛛網的中心，或是附近的植被。蛛網上若有落葉或是食物殘屑，銀腹蛛會將之清除。. (二)、形態特徵：銀腹蛛的身體可區分為頭胸部(cephalothorax)與腹部(abdomen)兩部分，也有學者把頭胸部稱為前體部(prosoma)，把腹部稱為後體部 (opisthosoma)。兩者以腹柄(pedicel)相連接，腹柄由腹部第一節特化而成(Zhu et al., 2003)，體表上有許多的毛或棘刺。 (1) 頭胸部 (Figs. 1, 2) 7.

(20) 頭胸部具幾丁質的外骨骼，背面稱為背甲(carapace)，腹面稱為胸板(sternum) (Figs. 1A, B)。在胸板與背甲之間著生六對附肢，第一對稱為螯肢(chelicerae)，第二對為觸肢(palp)，第三至六對為步足，分別稱為第一、第二、第三及第四對步足。銀腹蛛的背甲通常呈淡綠色或黃褐色，表面有細毛。背甲前端為頭區(cephalic region)，後端為胸區 (thoracic region)，頭區與胸區以頸溝(cervical groove)為界，胸區中央凹陷部分稱為中窩(thoracic groove; fovea)，中窩兩側具有放射溝 (radial furrow)。頭區具有八枚單眼(Fig. 2A)，前四眼排成一列，稱為前眼列(anterior eye row, AER)，後四眼稱為後眼列(posterior eye row, PER); 前眼列中央之兩眼稱為前中眼(anterior median eye, AME)，側邊之兩眼稱為前側眼(anterior lateral eye, ALE)；後眼列中央之兩眼稱為後中眼(posterior median eye, PME)，側邊之左右兩眼稱為後側眼 (posterior lateral eye, PLE)，前後側眼在銀腹蛛屬通常會相接觸(Fig. 2A)；前後中眼圍成的區域稱為中眼域(median ocular area, MOA)；前中眼前緣至背甲前緣之間的區域稱為額(clypeus)。螯肢(chelicera)上具有毒牙(fang)，毒牙平時收在基節的牙溝內 (fang furrow)，在牙溝前後兩側具有許多小突起，前側稱為前牙堤齒 (promarginal teeth)；後側稱為後牙堤齒(retromarginal teeth)(Fig. 2B)，牙堤齒的排列、數目、形態均可作為分類的特徵。台灣產銀腹蛛屬前 8.

(21) 牙堤齒的數目皆為 3 齒，但後牙堤齒的數目變化則較大：具 5 齒的為雪銀腹蛛及十字銀腹蛛之雌蛛(Figs. 7C, 8B, 13C)；具 4 齒的則較為常見，如肩斑銀腹蛛之雌蛛、西里伯銀腹蛛、尖尾銀腹蛛、小肩斑銀腹蛛、台灣銀腹蛛之雄蛛、方格銀腹蛛、武陵銀腹蛛和十字銀腹蛛之雄蛛(Figs. 9C, 11C, 12B, 14B, 15C, 16B, 17C, 18B, 20B, 21C, 22B, 23C, 24B)；具 3 齒的只見於肩斑銀腹蛛之雄蛛與台灣銀腹蛛之雌蛛(Figs. 10B, 19C)。觸肢有 6 節位於螯肢的後方，分別為基節(coxa)、轉節 (trochanter)、腿節(femur)、膝節(patella)、脛節(tibiae)和跗節(tarsus)。觸肢基節變形成為下顎(maxillae)(Fig. 1B)，又稱為顎葉(endite)。下唇 (labium)(Fig. 1B)位於胸板的正前方，銀腹蛛屬蜘蛛通常長大於寬。步足有 4 對，每一步足由 7 節所組成，即基節(coxa)、轉節(trochantor)、腿節(femur)、膝節(patella)、脛節(tibia)、蹠節(metatarsus)與跗節 (tarsus)。步足相對長度可以足式(leg formula)表示，銀腹蛛屬蜘蛛足式為 I > II > IV > III，最長的步足為第一對步足。銀腹蛛屬蜘蛛的共同特徵是在第四對步足的腿節上有成列的聽毛(trichobothria)(Fig. 1A)。有些銀腹蛛，例如肩斑銀腹蛛、方格銀腹蛛和武陵銀腹蛛等，在雄蛛的第一對及第二對脛節和蹠節腹面，常有小疣齒(denticles) (Figs. 10F, 20F, 22F, 24F)，可做為鑑別的參考。此外，有些銀腹蛛屬 9.

(22) 的雌蛛，例如方格銀腹蛛、武陵銀腹蛛和台灣銀腹蛛等，其第四對步足的脛節末端帶有瓶刷狀的密毛(tufts of setae)(Figs. 19G, 21D, 23D)。 (2) 腹部(abdomen)(Fig. 1B) 銀腹蛛屬蜘蛛的腹部通常呈卵形，背面常具有金屬光澤的斑點，而腹部上方通常還有許多成對凹陷的筋點(sigilla)，是肌肉附著的地方。腹部花紋隨種類而異，主要是呈葉脈狀的花紋，例如西里伯銀腹蛛、肩斑銀腹蛛和尖尾銀腹蛛；但是有些種類則散布銀白色的斑點，例如雪銀腹蛛和十字銀腹蛛。腹部腹面前方具有一條胃外溝 (epigastric furrow)，兩側為書肺(book lungs)之開口，即書肺氣門(lung slit)；生殖孔(genital opening)位於腹部的腹面前端。雌蛛在生殖孔及胃外溝上方通常呈現角質化(sclerotized)，稱為外雌器(epigynum)；雄蛛的生殖孔亦位於腹部的腹面前端，胃外溝前方的中央，但其腹部缺乏交配器官，也無角質化的現象，因此並不明顯；腹面末端具有三對絲疣(spinnerets)，即前側絲疣(anterior lateral spinneret, ALS)、後中絲疣(posterior midian spinneret, PMS)和後側絲疣(posterior lateral spinneret, PLS)，絲疣的後面有一個肛丘(anal tubercle)，而在絲疣的前方還有一退化的舌狀體(colulus)。 (3) 生殖構造生殖結構是鑑別蜘蛛種類很重要的依據，有關於銀腹蛛屬蜘蛛之雌、 10.

(23) 雄生殖構造敘述如下：雄性生殖構造(Fig. 2C)：雄蛛腹部具有一對精巢(testes)，精子經輸精管輸送至儲精囊，但雄蛛腹部並無交配器官，因此需藉由觸肢末端之特化之觸肢器(palpal organ)進行交配，銀腹蛛屬蜘蛛的觸肢器構造較其他種類蛛蜘簡單，雄蛛觸肢由兩部分所構成，杯葉(cymbium)及其內部構造-生殖球 (genital bulb)，而生殖球主要有盾片(tegulum)、小盾片(subtegulum)、導引器(conductor)、插入器(embolus)等構造。雄蛛在交配前，會先織成”Y” 字形的精網，隨後將精子排在其上，利用觸肢器上的導引器和插入器末端吸取精液(Zhu et al., 2003)。不同種類的蜘蛛觸肢器差異很大，可作為蜘蛛分類的主要依據(Fig. 3)。台灣產銀腹蛛屬蜘蛛雄蛛觸肢器依據其脛節的相對長度及杯葉基部突起(cymbial basal process, CBP)的有無可分為以下三大類： 1. 脛節較短且杯葉基部無突起(short palpal tibia without CBP)：脛節較杯葉短，且無杯葉基部突起。如雪銀腹蛛(Fig. 8E)和十字銀腹蛛(Fig. 24E)。 2. 脛節短且杯葉基部具突起(short palpal tibia with CBP)：脛節較杯葉短或等長，並具有杯葉基部突起。如方格銀腹蛛(Fig. 20E)、肩斑銀腹蛛(Fig. 10E)及西里伯銀腹蛛(Fig. 12E)。 11.

(24) 3. 脛節長並杯葉基部具突起(long palpal tibia with CBP)：脛節較杯葉長，並具有杯葉基部突起。如小肩斑銀腹蛛(Fig. 18E)、台灣銀腹蛛(Fig. 20E)、武陵銀腹蛛(Fig. 24C)與尖尾銀腹蛛(Fig. 16E)。. 雌性生殖構造：(Figs. 2D-E) 胃外溝中央前緣為生殖腺的開口(genital opening)，通常呈角質化，稱為外雌器(epigynum)，位於雌蛛腹部腹面靠近前端部分，有些種類的銀腹蛛在外雌器前端具有垂兜(hood)的構造，可作為分類的參考。外雌器外部具有成對的交配孔(copulatory opening)，經由交配管 (copulatory duct)連接到內部的受精囊(spermatheca)。銀腹蛛交配時，精子會從交配孔進入經交配管到受精囊的第一室，第一室與第二室及第三室相連通，經過交配後，第一室會膨大，第三室的一端連接受精管(fertilization duct)，可將精子送到子宮與陰道交接處與卵子結合，子宮左右兩邊各接一條輸卵管，分別通向一邊之卵巢(Eberhard and Huber, 1998)。台灣產銀腹蛛屬蜘蛛雌蛛依據外雌器的形態及垂兜的有無可分為三大類(Fig. 4)： 1. 中央凹陷且無垂兜(epigynum central-depressed without hood)：外雌器中央凹陷，無垂兜。如雪銀腹蛛和十字銀腹蛛(Figs. 4A, B)。 2. 中央平坦且無垂兜(epigynum central-flattened without hood)：外雌. 12.

(25) 器的中央平坦，且不具垂兜。如小肩斑銀腹蛛(Fig. 4C)、肩斑銀腹蛛(Fig. 4D)和武陵銀腹蛛(Fig. 4E)。 3. 中央平坦且具垂兜(epigynum central-flattened with hood)：外雌器的中央平坦，且具垂兜。如方格銀腹蛛(Fig. 4F)、台灣銀腹蛛(Fig. 4G)、西里伯銀腹蛛(Fig. 4H)和尖尾銀腹蛛(Fig. 4I)。. 二、研究方法：形態鑑定將全省各地採集的標本及台灣師範大學生命科學系館藏標本，根據蜘蛛的生殖構造特徵，作為最主要的分類依據。雄蛛觸肢器以細剪剪下，分別觀察觸肢器各部分骨片的詳細構造；雌蛛除觀察外雌器外部形態外，並將外雌器以細剪剪下，在 10% KOH 溶液加熱處理(Chen et al., 2003)，溶解蛋白質等雜質後，觀察幾丁質的受精囊(spermatheca) 與受精管(fertilizing tube)等內部生殖構造，並觀察比對蜘蛛身體其他各部位，如眼區、背甲、腹部、下唇、下顎、胸板、牙堤齒、步足和絲疣等外部構造和特徵(Figs. 1 and 2)加以計數或描述，顏色的描述主要依據浸泡在酒精中的標本顏色，並以野外觀察或拍照來輔助活體的描述。標本置於解剖顯微鏡(Leica Wild M3Z)下觀察繪圖，並以目鏡測微器測量下列形質(測量單位為公釐，精確至小數點第二位)：體長. 13.

(26) (頭胸部長加上腹部長)；背甲長(頭胸部前緣量至後緣之最長處)、背甲寬(取最寬處測量)、背甲高(取胸板至中窩的高度)；腹長(腹部最前緣量至肛丘末端，不包含絲疣的長度)、腹寬(取最寬處測量)、腹高(從胃外溝量起之垂直高度)；步足長為腿節(femur)、膝節(patella) + 脛節 (tibiae)、蹠節(metatarsus)與跗節(tarsus)長度之和、雄觸肢長為腿節 (femur)、膝節(patella)、脛節(tibia)與杯葉(cymbium)長度之和；前絲疣長(anterior spinneret, Asp)與後絲疣長 (posterior spinneret, Psp)自基部量起的長度。八個單眼在背甲前端呈二列，前後眼列各具四(Figs. 1A, 2A)，眼徑為各眼角膜之最長直徑，但後中眼則測量角膜水平方向最長距離。中眼域(median ocular area, MOA)為由前中眼及後中眼外所圍成的區域，中眼域長(length of MOA, MOA-L)由前中眼前緣量至後中眼後緣之長度；中眼域前寬(anterior width of MOA, MOA-AW)為量取兩前中眼外緣距離；中眼域後寬(posterior width of MOA, MOA-PW)為量取兩後中眼外緣距離。額高(Fig. 2A)(clypeus)為前中眼下緣量至背甲邊緣最凹處之距離。前中眼間距 (anterior medium eyes interval, AMI)為前中眼間最短距離; 後中眼距(posterior medium eyes interval, PMI)為後中眼間最短距離; 前中側眼距(anterior medium and lateral eyes interval, AMLI)為前中眼與前側眼間最短距離; 後中側眼. 14.

(27) 距(posterior medium and lateral eyes interval, PMLI)為後中眼與後側眼間最短距離。. 三、支序分析根據 Hormiga et al. (1995)的研究，銀腹蛛屬的姊妹群為長腳蛛屬 (Tetragnatha) + 骨螯蛛屬(Glenognatha) + 粗螯蛛屬(Pachygnatha)；谷川明男(Tanikawa, 2001)，提出銀腹蛛屬的姊妹群為沖繩蛛屬 (Okileucauge) + 高腹蛛屬(Tylorida) + 天星蛛屬(Mesida)；朱明生等 (Zhu et al., 2003)，將中國的長腳蛛科分為六個亞科：桂齊蛛亞科 (Guizygiellinae)、后蛛亞科、絡新婦蛛亞科、長腳蛛亞科、銀腹蛛亞科和鋸螯蛛亞科(Dyschiriognathinae)。其中銀腹蛛亞科包括臥龍蛛屬 (Wolongia)、長蹠蛛屬(Metleucauge)、高腹蛛屬、波斑蛛屬(Orsinome)、隨蛛屬(Opadometa)、銀腹蛛屬、天星蛛屬、和沖繩蛛屬等八屬。因此本研究以台灣地區野外採集的銀腹蛛標本及國立台灣師範大學生命科學系館藏蜘蛛標本為內群，包含十種銀腹蛛屬蜘蛛：雪銀腹蛛 (Leucauge argentina Hasselt, 1882)，肩斑銀腹蛛(L. blanda L. Koch, 1878), 西里伯銀腹蛛(L. celebesiana Walckenaer, 1842), 十字銀腹蛛(L. crucinota Bösenberg et Strand, 1906)，尖尾銀腹蛛(L. decorata Blackwall, 1864), 小肩斑銀腹蛛(L. subblanda Bösenberg et Strand,. 15.

(28) 1906), 台灣銀腹蛛(L. taiwanensis sp. nov.)，方格銀腹蛛(L. tessellata Thorell, 1887), 武陵銀腹蛛(L. wulingensis Song et Zhu,1992), 與採自美國的脈斑銀腹蛛(L. venusta Walckenaer, 1842)，其他的內群成員包含裝飾天星蛛(Mesida gemmae Hasselt, 1882), 印氏天星蛛(M. yini Zhu, Song et Zhang, 2003), Metabus ocellatus (Keyserling, 1864) (形態特徵資料取自 Álvarez-Padilla, 2007), 佐賀長蹠蛛(Metleucauge kompirensis Bösenberg et Strand, 1906), 佐佐木沖繩蛛(Okileucauge sasakii Tanikawa, 2001a), 喜隨蛛(Opadometa grata Guérin, 1838) (形態特徵資料取自 Zhu et al., 2003), 麥氏波斑蛛(Orsinome vethi Hasselt, 1882) (形態特徵資料取自 Zhu et al., 2003), 條紋高腹蛛(Tylorida striata Thorell, 1877)，及橫帶高腹蛛(T. ventralis Thorell, 1877)，並參考各文獻中所列的類群及特徵(Hormiga et al., 1995; Coddington, 1990; Tanikawa, 2001; Zhu et al., 2003; Álvarez-Padilla, 2007; Kuntner et al., 2008)以台灣產長腳蛛屬的華麗長腳蛛 Tetragnatha nitens (Audouin, 1826)為外群(Table 1)，加上本研究所觀察到共 45 個特徵，利用支序分析軟體 NONA version 2.0 (Goloboff et al., 2004)與 PAUP* version 4.0 beta 10 (Swofford, 2002)做本屬親緣關係的最儉約分析(parsimony analyses)，並以 Ratchet Search(通常能搜尋最短儉約樹)(Nixon, 1999) 與分枝-限定搜尋法(Branch and Bound Search)，找出最儉約樹並分析. 16.

(29) 結果。以 Winclada 1.00.08 (Nixon, 2002)編輯資料矩陣(Table 2)和支序樹圖，所有多態性的特徵值(multistate characters)皆設定為非加成性 (non-addictive)，Bootstrap 值(Felsenstein, 1985)以 Winclada 的預設值 mult*10(重覆取樣 100 次)，並分析其步長(Tree length)、一致指數 (consistency Index, CI)值、及保留指數(retension index, RI)值(Farris, 1969)與支持度 Bremer support (Bremer, 1995)。. 17.

(30) 參、親緣關係分析 Phylogenetic analysis 一、Taxa (類群類群) 類群 Twenty taxa were used in the phylogenetic analysis to test the monophyly of genus Leucauge (Table 1). Judging from the cladogram presented by Hormiga et al. (1995), Leucauge is the sister group of the clade Tetragnatha + Pachygnatha + Glenognatha. Sister group of Leucauge presented by Tanikawa (2001) is the clade Okileucauge + Tylorida + Mesida. The Leucauginae presented by Zhu et al. (2003) included genera Wolongia, Metleucauge, Tylorida, Orsinome, Opadometa, Leucauge, Mesida, and Okileucauge. Álvarez-Padilla (2007) presented “leucaugines” included the genera Azilla, Orsinome, Tylorida, Leucauge, Metabus, Mesida, and Opadometa. I choose genera Tetragnatha with the exemplar Tetragnatha nitens (Audouin, 1826) as an out-group for the present cladistic analysis. Nine species of Leucauge from Taiwan and one species from North America are selected as in-group species: Leucauge argentina (Hasselt, 1882), L. blanda (L. Koch, 1878), L. celebesiana (Walckenaer, 1842), L. crucinota (Bösenberg et Strand, 1906), L. decorata (Blackwall, 1864), L. subblanda Bösenberg et Strand, 1906, L. taiwanensis sp. nov., L. tessellata (Thorell, 1887), L. wulingensis Song et Zhu,1992, and L. venusta (Walckenaer, 1842) (from Miami, USA). In addition, other genera of the subfamily Leucaugine are also chosen as in-group species: Mesida gemmae (Hasselt, 1882), M. yini Zhu et al., 2003, Metleucauge kompirensis (Bösenberg et Strand, 1906), 18.

(31) Okileucauge sasakii Tanikawa, 2001, Tylorida striata (Thorell, 1877), T. ventralis (Thorell, 1877), Metabus ocellatus (Keyserling, 1864) (morphological data from Álvarez-Padilla, 2007), Opadometa grata (Guérin, 1838) (morphological data from Zhu et al., 2003), and Orsinome vethi (Hasselt, 1882) (morphological data from Zhu et al., 2003).. 二、Characters descriptions (特徵描述特徵描述) 特徵描述 Characters scoring are presented in Table 2. The character matrix contains 45 characters: 25 characters derived from female somatic morphology, 7 from female copulatory organs, 1 from male somatic morphology, 12 from male copulatory organs. Characters used in previous phylogenetic studies are cited at the end of each character using a code. The codes of references are used: Álvarez-Padilla, 2007 = A07; Kuntner et al., 2008 = K08; Tanikawa, 2001 = T01; Zhu et al., 2003 = Z03. Original character numbers of the reference are following the code. Character 1. Female body size: 0 = large, larger than 4 mm, 1 = small, equal or smaller than 4 mm. Lecauge crucinota and L. argentina differ from the other congeners by having a small body size. Character 2. Female thoracic groove, fovea (T01-2, K08-6): 0 = shallow, bottom visible; 1 = deep, bottom invisible. Mesida, Okileucauge and some tetragnathids have a shallow thoracic groove. Character 3. Female cephal-area with black stripes: 0 = absent, 1 = present. The cephal-area of geuns Metleucauge, Tylorida, Orsinome, and some tetragnathids have a black stripe on it.. 19.

(32) Character 4. Female retrolateral cephal-area depressions: 0 = absent, 1 = present. Genus Opadometa, and Leucauge argentina (Fig. 7A) have a pair of depressions on the retrolateral cephal-area. Character 5. Female lateral eyes (T01-12, Z03-0): 0 = separated, 1 = touching. The lateral eyes of Tetragnatha are separated, and the other taxa are touching. Character 6. Female AME-ALE separation (K08-13, modified): 0 = wide, wider than one AME width, 1 = narrow, equal or narrow than one AME. Okileucauge sasakii, Tylorida striata, T. ventralis, Orsinome vethi, Leucauge argentina, and L. crucinota have a narrow separation of AME-ALE. Character 7. Female PLE-PME separation (K08-13 modified): 0 = wide, more than one PME width, 1 = narrow, equal or less than one PME width. Leucauge crucinota, Metleucauge kompirensis, Tylorida striata, and Okileucauge sasakii have a narrow separation of PLE-PME. Character 8. PME tapetum (Z03-2, A07-23, K08-17): 0 = absent, 1 = present. The tapetum of PME in genus Tetragnatha is absent. Character 9. PLE tapetum (A07-24, K08-18): 0 = absent, 1 = present. The tapetum of PLE in genus Tetragnatha is absent. Character 10. Female clypeus height (T01-7, Z03-2, A07-25, K08-19): 0 = low, less than one AME width, 1 = median, more than one AME, but less than two AME width, 2 = high, equal or more than two AME width. The clypeus height of Leucauge argnetina, L. crucinota, and genus Mesida are low. Character 11. Labium (Z03-5): 0 = longer than wide, 1 = wider than long. Genus Tetragnatha has a long labium. 20.

(33) Character 12. Dorsal femoral trichobothria (T01-1, Z03-8, A07-27, K08-47): 0 = absent, 1 = present. Okileucauge, Metleucauge, and Metabus lack the femoral trichobothria, present in the other taxa. Character 13. Femur IV trichobothria arrangements (A07-28, K08-48): 0 = scattered, 1 = in rows. In Leucauge, Opadometa, Mesida, Tylorida, and Orsinome the trichobothira are arranged in a paired row on prolateral femur IV. Character 14. Femur IV trichobothria rows: 0 = less than 9 rows, 1 = 10-11 rows, 2 = 13-15 rows, 3 = 19-21 rows, 4 = more than 25 rows. In Opadometa the trichobothria are more than 25 rows, but Tylorida and Mesida have less than 9 rows. The rows of trichobothria in Leucauge are various. Character 15. Femur IV trichobothria extension (A07-30, modified): 0 = less than 3/5 femur length, 1 = more than 3/5 femur length. In Opadometa the extension of trichobohthria is usually more than 3/5 femur length. Character 16. Femur IV Trichobothria morphology (1) (A07-29 modified): 0 = smooth, 1 = branched. Tetragnatha, Tylorida, and Orsinome vethi have the smooth trichobothria on femur IV. Character 17. Femur IV Trichobothria morphology (2) (A07-29, modified): 0 = short branched, 1 = long branched. Femur IV trichobothria of Mesida are long branched. Character 18. Female leg I relatively longer than the other legs: 0 = absent, 1 = present. Legs I of Tylorida is obviously longer than the others legs. 21.

(34) Character 19. Female tibia IV tufts (A07-31, K08-51): 0 = absent, 1 = present. Opadometa species, Leucauge taiwanensis, L. tessellata, and L. wulingensis (Figs. 19G, 21D, 23D) possess dense tibial setae on legs IV. Character 20. Female metatarsus IV tufts: 0 = absent, 1 = present. Leucauge taiwanensis, and L. wulingensis possess dense setae on metatarsus IV (Figs. 19G, 23D). Character 21. Female abdomen anterior extension: 0 = absent, 1 = present. Anterior abdomen of Opadometa is strong extension. Character 22. Female abdominal humps (K08-60, 61, modified): 0 = none or inconspicuous, 1 = one pair, 2 = two pairs or more. One pair of abdominal hump is found in Lecuage blanda and L. decorata (Figs. 9A-B, 15A-B), and two pairs or more in L.argentina, and L. crucinota (Figs. 7A-B, 9A-B). Character 23. Female dorsal abdomen silver spots (T01-05, A07-02, K08-82): 0 = absent, 1 = present. Conspicious abdominal silver spots, typical in Leucauge, Okileucauge, Mesida, is not found in Metleucauge. Character 24. Female dorsal abdomen leaf vein-liked pattern: 0 = absent, 1 = present. Dorsal abdomen with leaf vein-liked pattern is typically found in some Leucauge species (e.g. L. celebesiana, Fig. 11A) Character 25. Ventral abdomen with orange pigments in life: 0 = absent, 1 = present. Orange pigments are found on the ventral abdomen of Mesida, and some Leucauge species in life (e.g. L. crucinta, and L. venusta). The orange pigments will be vanished when the specimen stored in alcohol.. 22.

(35) Character 26. Epigynum (T01-20, Z03-37, A07-79, K08-87): 0 = not sclerotized, 1 = sclerotized. Tetragnathines (Tetragnatha,Glenognatha, and Pachygnatha) lack the epigynum plate (Kuntner et al., 2008). Character 27. Median plate of epigynium: 0 = flattened, 1 = depressed with weak-sclerotized border, 2 = depressed with anterior well-sclerotized ridge, 3 = swelled. The median plate of epigynum in Tylorida (Zhu et al., 2003: figs. 169G, 172F), Okileucauge (Tanikawa, 2001) are depressed with weak-sclertoized border. In Lecauge argentina and L. crucinota, the median plate of epigynium is depressed with anterior well-sclertoized ridge. Metleucage has a swelled median plate of epigynum (Zhu et al., 2003: fig. 151F). Character 28. Epigynal cuticle hood: 0 = absent, 1 = present. The epigynal cuticle hood is semitranslucent (e.g. Leucauge celebesiana, Figs. 11D-F). Character 29. Copulatory opening position (K08-103): 0 = ventral, 1 = caudal. In Metleucauge, Opadometa, Leucauge argentina, and L. crucinota, the position of copulatory openings is caudal (Figs. 7E, 13E). Character 30. Copulatory ducts: 0 = straight, 1 = u-looped, 2 = n-looped. The copulatory ducts are straight in geuns Tetragnatha, Metleucauge, Opadometa, Okileucauge, and Tylorida species. In most species of Leucauge the copulatory ducts are u-looped, except for L. argentina and L. crucinota that are n-looped (Figs. 7E, 13E). Character 31. Fertilization ducts (A07-80, K08-110): 0 = absent, 1 = present. Tetragnathines lack fertilization ducts (Kuntner et al., 2008) Character 32. Chambers of spermatheca (Z03-38): 0 = one, 1 = two, 2 = three. Metleucauge and L. crucinota (Fig. 13E) have only one 23.

(36) chamber of spermatheca. Two chambers of spermatheca are in Opadometa, Tylorida, and Orsinome, and three are in Metabus, Mesida, and most Leucauge. Character 33. Male cheliceral clasping spurs (T01-10, K08-128): 0 = absent, 1 = present. Male cheliceral clasping spurs are found in Mesida Opadometa grata (Zhu et al., 2003: fig. 161A) and Tetragnatha (Zhu et al., 2003: Figs. 86A-C, 145B). Character 34. Male palpal trochanter (K08-134): 0 = short (twice the width or less), 1 = long (more than twice the width). Male palpal trochanter of Leucauge crucinota is short. Character 35. Male palpal patella macrosetae (T01-18, A07-36, K08-136): 0 = none, 1 = one. Male palpal patella macrosetae of Leucauge argentina and L. crucinota are absent (Figs. 8C, 14C). Character 36. Male palpal tibia length (T01-19, K08-137, modified): 0 = very short (tibia/cymbium length less than 0.4, e.g. Leucauge crucinota, Fig. 14C), 1 = short, (tibia/cymbium length equal 0.5 to 1, e.g. L. blanda, Fig. 10C), 2 = long (tibia/cymbium length equal 1 to 1.5, e.g. L. wulingensis, Fig. 24C), 3 = very long (tibia/cymbium length more than 1.5, e.g. L. decorata, Fig. 16C). Character 37. Cymbial basal process (CBP) (A07-41, K08-142): 0 = absent or reduced, 1 = present. CBP is present in Opadometa, Mesida, Tylorida, Okileucauge, some Metabus species and most Leucauge (e.g. L. blanda, Fig. 10E). Character 38. Cymbium (K08-138): 0 = constricted, 1 = entire. In Tetragnatha and Opadometa grata the cymbium is constricted (Zhu et al., 2003: figs. 161C, D). 24.

(37) Character 39. Paracymbium morphology (A07-53, K08-147): 0 = longer than wide, straight, 1 = short basal structure, hook-shaped. The paracymbium of Tetragnatha is long and straight (Zhu et al., 2003: figs. 86A). Character 40. Position of subtegulum: 0 = located under the tegulum, 1 = ectal displacement. In Tetragnatha and Opadometa the position of subtegulum is located under the tegulum. Character 41.Size of tegulum: 0 = normal, 1 = enlarged. Tegulum of Tylorida is enlarged in size (Zhu et al., 2003: figs. 170B-D, 177I-J). Character 42. Reservoir course (sperm duct) (T01-13, A07-60, K08-155): 0 = spiraled, 1 = with switchbacks. Sperm duct turns and twist within the tegulum in Leucauge, Metabus, Mesida, Okileucauge, and Tylorida. Character 43. Conductor and embolus (T01-15, A07-62, K08-168): 0 = separate, 1 = conductor wraps embolus. Character 44. Measurement of subtegulum/cymbium area, ventral view: 0 = small (subtegulum/cymbium area less than 0.15, e.g. Leucauge argentina Fig. 8D), 1 = median (subtegulum/cymbium area equal 0.2-0.5, e.g. L. blanda Fig. 10D), 2 = large (subtegulum/cymbium area more than 0.6, e.g. Leucauge crucinota, Figs. 14D-F). Character 45. Conductor secondary apophysis (CSA) (K08-169): 0 = absent, 1 = present. The secondary apophysis of conductor is absent in Tetrangatha, and Leucauge crucinota (Figs. 14D-F).. 25.

(38) 三、Results (結果結果) 結果 Ratchet searches under equal weights analyses of the complete data set (Table 2) in NONA (Goloboff et al., 2004) obtained the most parsimonious tree (Fig. 5, L = 96, CI = 0.58, RI = 0.61) with unambiguous optimization. By branch and bound searches under equal weights analyses in PAUP (Swofford, 2002), 96 most parsimonious trees were obtained (L = 96, CI = 0.58, RI = 0.61), and the 96 trees differ in ingroup relations by having low support values except for the assemblage of L. crucinota and L. argentina whose support values is high (Fig. 6). As the result, genus Leucauge can be divided into two main groups. The Leucauge group I included L.argentina and L. crucinota is a monophyletic group and is sister to the other leucaugines. Six synapomorphies support the Leucauge gruop I: female with low clypeus (10-0), femur IV trichobothria branched (16-1), female dorsal abdomen with two or more pairs of humps (22-2), copulatory ducts n-looped (30-2), male palpal trochanter short (35-0), and subtegulum area depression (42-1). Group I have high support values (Bootstrap 93%, Fig. 33) and far away from the Leucauge group II. These reasons imply that Leucauge group I may belong to a new genus. The rest species of Leucauge and Opadometa form a second clade, the Leucauge group II. Mesida is the sister group of Leucauge group II. Three synapomorphies support the monophyly of Leucauge gruop II: i.e., female thoracic groove deep (2-1), femur IV with 19-21 rows of trichobothria (14-2), and the median plate of epigynium flattened (27-0). The tibita IV in female with tufts of setae (19-0) diagnoses (Opadometa 26.

(39) grata (L. tessellata (L. taiwanesis, L. wulingensis))) as a subgroup of the Leucauge gruop II, and the rest species of Leucauge members into another subgroup. However, nodes within Leucauge group II have only a low support values (Bootstrap < 50%). A further study of electron microscope in those of spinnerets and the behavior characters are necessary to support the Leucauge group II. In sum, the results suggest that genus Leucauge is probably a polyphyletic group as previously proposed by Zhu et al. (2003). Yet, the poor support values of many nodes within the cladogram (Fig. 6) in this article, the author adopts the previous study of Leucauge species without changing their genus position.. 四、Discussion (討論討論) 討論 Although the studys of tetragnathids phylogeny are well promotive in recent years (Álvarez-Padilla, 2007, Kuntner et al., 2008, Zhu et al., 2003). The species number of Leucauge from Southeastern Asia is made up 29 % of the world (Zhu and Wang, 2004). However, the phylogenetic relationships within subfamily Leucauginae remain unknown, and the genus status of some known species is doubtful. For instance, L. nanshan Zhu et al. (2003: 236, fig. 128) having three pairs of humps on the dorsum of abdomen and with a low clypeus probably belongs to the Leucauge group I (Fig. 5), and L. xiuying Zhu et al. (2003: 256, fig. 142), having only one chamber of spermatheca in female, palpal conductor without secondary apophysis and tegulum without a switchback in male, maybe belong to an unnamed genus. These evidences imply that many 27.

(40) genera of Leucauginae remain unknown in the present taxonomy. Six synapomorphies support the clade Leucauge gruop I (Fig. 5) such as female with a low clypeus (10-0), dorsum of abdomen with two or more pairs of humps (22-2), and n-looped copulatory ducts (30-2), and male with a short palpal trochanter (35-0). The Leucauge group I, a basal clade of subgenus Leucauge is a sister group of the other genera of Leucaugines. L. nanshan Zhu et al. (2003: 236, fig. 128) from China having three pairs of humps on the dorsum of abdomen and with a low clypeus probably belongs to the group I. High support value implies group I may not be a part of the genus Leucauge, but an unknown genus. By having many autapomorphies, such as tegulum with a shield and conductor edge with many minute denticles, L. crucinota is quite different from L. argentina. In addition, the chambers of spermatheca seems various within group I, since the spermatheca of L. argentina has three chambers and that of L. crucinota as well as L. nanshan each has only one chamber. These characters suggest that L. crucinota and L. argentina may belong to a second group within group I. However, lacking the male characters of L. nanshan, the phylogenetic relations among Leucauge group I is still problematic. Zhu et al. (2003) included genera Wolongia, Metleucauge, Tylorida, Orsinome, Opadometa, Leucauge, Mesida, and Okileucauge in the subfamily Leucauginae. Mesida and Okileucauge are the sister groups of Leucauge. Mesida is the sister group of present Leucauge group II (Fig. 5) which is congruent with the cladogram present by Zhu et al. (2003). However, making an accurate definition of genus Leucauge is difficult, since the type specimen of the type species, Leucauge argyrobata (White, 28.

(41) 1841), are lost and the identity of the species is not know (Levi, 1980). In the present study, Lecuauge venusta, a common species came from Norther America was grouped into Leucauge group II, and was classified as a species of genus Leucauge by Cambridge (1903). One synapomorphy, trichobothria on femur IV of female with 10 pairs or more, supports the clade of genus Leucauge by Tanikawa (2001). Álvarez-Padilla (2007) gives two synapomorphies to support the monophyly of genus Leucauge by having clypeus height equal or larger than one AME diameter and paracymbium distal margin with a triangular sclerotized growth. However, the nodes within the clade of Leucaugines also have low support values (Álvarez-Padilla, 2007). These evidences imply that group II is probably closer to the traditional concept of genus Leucauge by having the female with a deep thoracic groove (2-1), femur IV with 19 to 21 pairs of trichobothria (14-2), and the median plate of epigynium flattened (27-0) in the phylogenetic tree (Fig. 5). Unlike Leucauge group I, the chambers of spermatheca in group II are stable by having three chambers, except for Opadometa which has two chambers. Genus Opadometa is also grouped in the Lecuage group II. Opadometa is a small genus, currently with only two speices and eight subspecies (Platnick, 2008). Opadometa resembles Leucauge by having the dense tibial setae on female legs IV and a pair of long trichobothria on femur IV, but can be distinguished from the latter by having anterior abdomen extension, two chambers of spermatheca in female, and a basal apophysis of palpal conductor in male (Zhu et al., 2003: figs. 160, 161C-E). Kuntner et al. (2008: fig.18) presented Opadometa + Leucauge is sister group of Tetragnathinae by having the following synapomorphies: 29.

(42) wide sternum, femoral IV trichobothria in rows, central light pigmented spot on venter, slit-shaped copulatory openings, membranous spermathecae, aggregate spigots embracing flagelliforms, conductor secondary apophysis, unsclerotized embolus. It deserves to be mentioned that the conductor secondary apophysis (CSA) of Opadometa grata is strong reduced and forming a short hook, which differs from Leucauge species. Zhu et al. (2003) considered genus Opadometa was valid, a sister group of the clade (Leucauge (Mesida, Okileucauge)), that is not congruent with the result of my study. Since I have not obtained the male of O. fastigata, another species of genus Opadometa, it is difficult to make any conclusion on the systematic of Opadometa in this study. By having the low support values (Fig. 6), group II is still an unstable clade, and needs more characters to support it. In short, the cladogram shows that present genus Leucauge is a polyphyletic assamblage. Further detailed investigations into the genus is needed that may improve the generic level of phylogenetic analyses successfully.. 30.

(43) 肆、分類處理. 本論文共報告台灣產銀腹蛛屬蜘蛛 9 種，其中新種有 1 種，台灣銀腹蛛(Leucauge taiwanensis sp. nov.)；新紀錄種有 4 種：西里伯銀腹蛛(L. celebesiana)、十字銀腹蛛(L. crucinota)、小肩斑銀腹蛛(L. subblanda)、武陵銀腹蛛(L. wulingensis)。文中將大銀腹蛛(L. magnifica) 處理為西里伯銀腹蛛的同物異名；擬方格銀腹蛛(L. subtessellata)處理為方格銀腹蛛(L. tessellata)的同物異名。齋藤三郎(Saito, 1933)記錄分布於台灣北部的脈斑銀腹蛛(L. venusta)處理為排除種。. SYSTEMATICS. Genus Leucauge White, 1841 銀腹蛛屬銀腹蛛屬 Type species: Leucauge argyrobapta (White, 1841). Type species by monotypy Linyphia (Leucauge) argyrobapta White from Rio De Janeiro (里約熱內盧). The name is feminine (Levi, 1980).. Diagnosis Genus Leucauge were grouped based on two synapomorphies, the presence of dorsal femoral trichobothria and of posterior gut caeca (Hormiga et al., 1995). Leucauge most resembles Mesida, Tylorida, Opadometa, and Okileucauge within the Leucauginae by having the 31.

(44) abdominal silver pigment, but can be distinguished by the following characteristics. Leucauge, Mesida, Tylorida, and Opadometa differ from Okileucauge by the presence of long, feathered trichobothria on the anterior surface of the femur IV (Tanikawa, 2001). However, Leucauge differs from Mesida by the absence of a spur on the anterior surface of male chelicerae. Leucauge differs from Tylorida by the absence of the swollen tegulum of male palp and the long complex-curved sperm duct (Tanikawa, 2004). Leucauge differs from Opadometa by the absence of the apophysis on the base of male conductor, and the anterior stretched of female abdomen (Zhu et al., 2003).. 32.

(45) Key to the species of genus Leucauge from Taiwan. 1. Male……...…………………………………………………………….2 Female...…………………….………………………………………..10 2. Chelicera armed with 5 retromarginal teeth………......……L. argentina Chelicera armed with 3 or 4 retromarginal teeth…...………………….3 3. First promarginal tooth minute……………….……………....L. blanda First promarginal tooth normal ………………………….…………….4 4. Palpal tibiae very long ( > = 1.6 times cymbium)……...…...L. decorata Palpal tibiae not very long ( < 1.5 times cymbium)……..…...………..5 5. Tip of embolus elongated………..………….………..….…………… 6 Tip of embolus short…………….……..……………….……………...7 6. Palpal tibiae longer than cymbium………….…….…......L. taiwanensis Palpal tibiae shorter than cymbium………………...…...….L. tessellata 7. Palpal tibiae longer than cymbium……………….………………..…..8 Palpal tibiae shorter or equal to cymbium……………….…………….9 8. Base of conductor swollen……………...…………………L. subblanda Base of conductor flattened…………......….……………L. wulingensis 9. Body length smaller than 4 mm………...…………….……L. crucinota Body length larger than 4 mm……...………………….…L.celebesiana 10. Chelicera armed with 5 retromarginal teeth………...………..……..11 Chelicera armed with 3 or 4 retromarginal teeth……………………12 11. Spermatheca with one chamber………………………..…L. crucinota Spermatheca with three chambers……………………..…L. argentina 12. Chelicera armed with 3 retromarginal teeth………...….L. taiwanensis Chelicera armed with 4 retromarginal teeth…...…………..………..13 33.

(46) 13. Epigynum without hood…………………………………………….14 Epigynum with hood……….……………………………………….16 14. Tibia IV without tufts of satea………………………………………15 Tibia IV with tufts of satea…………………………..…L. wulingensis 15. Copulatory openings arch-shaped……………..………..…...L. blanda Copulatory openings round-shaped……...…………...….L. subblanda 16 .Abdomen caudal-elongated……………………….……….L. decorata Abdomen oval-shaped……………………………..………………..17 17. Tibia IV without tufts of satea………………………….L. celebesiana Tibia IV with tufts of satea……………..……………........L. tessellata. 34.

(47) Leucauge argentina (Hasselt, 1882) 雪銀腹蛛 (Figs. 7, 8, 25). Theridion argentinum Hasselt, 1882: 34, pl. 2, f. 5. Argyroepeira argentina: Thorell, 1890: 199; Workman, 1896: 54, pl. 54. Leucauge argentina: Barrion and Litsinger, 1995: 543, f. 336a-e; Tso and Tanikawa, 2000: 126, f. 1-6; Zhu et al., 2003: 217, f. 115A-G.. Specimens examined TAITUNG Co.:1♂, 3♀, Lanyu, Sankungtung, alt. 10 m, 3 Feb. 2001, coll. W.-J. Huang (NTNUB-Ar 12007-12010); 4 ♀, Lanyu, Chung-ai Bridge, alt. 10 m, 4 Feb. 2001, coll. W.-J. Huang (NTNUB-Ar 12964-12967); 1 ♀, Lanyu, Chung-ai Bridge, alt. 10 m, 2 May 2001, coll. W.-J. Huang (NTNUB-Ar 12998).. Diagnosis Leucauge argentina differs from the other congeners by having 5 retromarginal teeth in both female and male (Figs. 7C, 8B). The female of L. argentina resembles L. crucinota by having a similar dorsal abdominal pattern, but can be distinguished from the latter by having three chambers of spermatheca rather than one (Fig. 7E).. Description Female (NTNUB- Ar 12998). Total length 4.63: cephalothorax length 1.88, width1.44, height 1.13; 35.

(48) abdomen length 3.13, width 2.00, height 2.06. Measurements of palpus, legs, and spinnerets: palpus 1.84 (0.63, 0.25, 0.34, 0.63); leg I 10.50 (3.13, 3.44, 2.94, 1.00); II 8.75 (2.63, 2.81, 2.50, 0.81); III 3.88 (1.25, 1.13, 1.00, 0.50); IV 7.00 (2.25, 2.06, 2.00, 0.69); ALS 0.08; PLS 0.08. Carapace yellowish brown in alcohol, greenish in life, with a gray margin on each side and black eye margins; cephalic region higher than thoracic region with a pair of depression posteriorly; thoracic groove deep, trifid posteriorly (Fig. 7A). Diameters of eyes in ratio, AME: ALE: PME: PLE = 0.13: 0.13: 0.10: 0.10. Length of MOA in ratio, MOA-L: MOA-AW: MOA-PW = 0.33: 0.28: 0.28. AMI: AMLI: PMI: PMLI = 0.05: 0.08: 0.08: 0.13. Clypeus height 0.23 times diameter of AME. Chelicerae brown; fang dark brown; promargin of fang groove armed with 3 robust triangular teeth and retromargin with 5 teeth (Fig. 7C). Endite brown, longer than wide. Labium reddish brown, longer than wide. Sternum brownish, heart-shaped. Palpus and legs yellowish brown, with black spines and hairs. Femur IV with two fringe of trichobothria (11 pairs) on prolateral surface of proximal half. Order of leg length I > II > IV > III. Abdomen oval with three pairs of humps; dorsum silvery, posterior of abdomen with two black stripes (Figs. 7A-B); venter dark brown with two parallel longitudinal white bands and silver spots. Epigynum brownish; central region rectangle with depression; posterior end of median septums wider than anterior (Fig. 7D). Spermatheca thin-walled with one large and two small chambers, the third one is multi-lobed (Fig. 7E).. Male (NTNUB- Ar12007). 36.

(49) Similar to female in shape and coloration (Fig. 8A). Total length 3.13: cephalothorax length 1.50, width 1.13, height 0.63; abdomen length 1.69, width 1.25, height 1.25. Measurements of palpus, legs, and spinnerets: palpus 2.88 (1.06, 0.44, 0.31, 1.06); leg I 10.75 (3.44, 3.38, 3.00, 0.94); II 8.56 (2.50, 2.69, 2.50, 0.88); III 3.75 (1.25, 0.94, 0.94, 0.63); IV 6.44 (2.25, 1.88, 1.69, 0.63); ALS 0.21; PLS 0.18. Diameters of eyes in ratio, AME: ALE: PME: PLE = 0.12: 0.10: 0.08: 0.10. AMI: AMLI: PMI: PMLI = 0.05: 0.05: 0.05: 0.10. Clypeus height 0.67 times diameter of AME. Promargin of fang groove armed with 3 teeth, retromargin with 5 teeth with the first one strongest (Fig. 8B). Subtegulum concave; without cymbial basal process; paracymbium finger-shaped; conductor inflatable, hook-shaped on the tip; palpal tibiae shorter than cymbium (Figs. 8C-E).. Variations Five females from Lanyu, Taitung Co. were measured (with the mean in parentheses). Total length 3.38-4.88 (4.20); carapace length 1.88-2.00 (1.94); width 1.25-1.63 (1.45); abdomen length 1.75-3.38 (2.64), width 1.25-2.56 (1.86).. Distribution Singapore, Sumatra, Philippines, and Taiwan.. Remarks Leucauge argentina was first recorded from Lanyu, Taiwan by Tso and Tanikawa (2000). It only occurs at Lanyu island in Taiwan (Fig. 25). 37.

(50) Leucauge blanda (L. Koch, 1878) 肩斑銀腹蛛 (Figs. 9, 10, 26). Meta blanda L. Koch, 1878: 743, pl. 15, f. 5. Meta japonica Thorell, 1881: 126. Leucauge blanda: Bösenberg and Strand, 1906: 182, pl. 3, f. 8, pl. 15, f. 394; Nakatsudi, 1942: 309, f. 22; Yaginuma, 1953: 28, f. 1; Yaginuma, 1954: 2, f. 3-4, 8, 12; Saito, 1959: 111, f. 134a-d; Yaginuma, 1960: 70, f. 66.2, 4; Yaginuma, 1960: 70, f. 66.2, 4; Lee, 1964: 55, f. 18e-g; Chang, 1970:16-4, f. 7; Chu and Okuma, 1970: 72, f. 3A-C; Yaginuma, 1971: 70, f. 66.2, 4; Yin, 1976: 120, pl. I.1-12; Song, 1980: 114, f. 58a-g; Hu, 1984: 136, f. 138.1-8; Yaginuma, 1986: 127, f. 68.2; Song, 1987: 181, f. 140; Chikuni, 1989: 91, f. 5; Feng, 1990: 104, f. 79.1-10; Chen and Gao, 1990: 77, f. 96a-f; Chen and Zhang, 1991: 120, f. 114.1-10; Zhao, 1993: 266, f. 126a-d; Kim et al., 1999: 46, f. 3, 28, 50; Song et al., 1999: 213, f. 121A-B, Q-R, 130G-H; Hu, 2001: 591, f. 401.1-4; Namkung, 2002: 223, f. 18.10a-b; Zhu et al., 2003: 220, f. 117A-I, 118A-D; Namkung, 2003: 225, f. 18.10a-b. Leucauge blanda japonica Bösenberg et Strand, 1906: 184. Leucauge szechuensis Schenkel, 1936: 93, f. 33.. Specimens examined KEELUNG City: 1♀, Cidu, Madong, alt. 50 m, 21 Oct. 2006, coll. H.-H. Liu (NTNUB-Ar34176). TAIPEI City: 1♀, 1♂, Wenshan, Hsienchihyien, alt. 100 m, 24 Feb. 2006, coll. H.-H. Liu 38.

(51) (NTNUB-Ar29586-29587); 11♀, 1♂, Wenshan, Chihnankung, alt. 300 m, 30 Mar. 2006, coll. H.-H. Liu (NTNUB-Ar29497- 29508). TAIPEI Co.: 1♀, Chungho, Yuantungszu, alt. 170 m, 9 Mar. 2006, coll. H.-H. Liu (NTNUB-Ar29144); 4♀, Shihting, Mt. Erhko, alt. 450 m, 11 Mar. 2000, coll. S.-H. Chen (NTNUB-Ar14037-14040); 2♀, Shihting, Yinghotung, alt. 400 m, 1 May 1982, coll. S.-H. Chen (NTNUB-Ar13575- 113576). HSINCHU Co.: 10♀, Guansi, Shenkengz, alt. 170 m, 9 Feb. 2006, coll. H.-H. Liu (NTNUB-Ar34089- 34098). MIAOLI Co.: 5♀, 1♂, Tunglo, Changshu, alt. 300 m, 13 Oct. 2006, coll. H.-H. Liu (NTNUB-Ar2912329128); 4♀, 4♂ Chunan, alt. 20 m, 16 Mar. 2007, coll. H.-H. Liu (NTNUB-Ar34194- 34201). TAICHUNG City: 13♀, Peitun, Dakeng, alt. 300 m, 31 Mar. 2006, coll. H.-H. Liu (NTNUB- Ar29554- 29566). CHANGHUA Co.: 3♀, 3♂, Tienchung, Neiwan, alt. 300 m, 13 May 2006, coll. H.-H. Liu (NTNUB-Ar29155- 29160). NANTOU Co.: 2♀, Caotun, Jioujioufeng, alt. 200 m, 19 Jul. 2006, coll. H.-H. Liu (NTNUB-Ar29912, 29913); 2♀, 1♂, Yuchih, Kongciyueyuan, alt. 750 m, 30 Jan. 2006, H.-H. Liu (NTNUB-Ar29891-29893); 5♀, 2♂, Nantou , Jongsingsincun, alt. 150 m, 21 Jun. 2006, coll. H.-H. Liu (NTNUB-Ar34050-34056); 2♀, Mingchien, Chingshuiwei, alt. 150 m, 2 Apr. 2003, coll. S.-H. Chen (NTNUB-Ar16471-16472). CHIAYI Co.: 1♀, Tapu, Sanjiaonanshan, alt. 500 m, 8 Feb. 2006, coll. S.-H. Chen (NTNUB-Ar29271). TAINAN Co.: 1♀, Kunmiao, Kaoling Bridge, alt. 50 m, 2 Nov. 2001. coll. S.-H. Chen (NTNUB-Ar13958); 1♀, 2♂, Nansi, Tsengwenshuiku, 100 m, 30 Aug. 2006, coll. H.-H. Liu (NTNUB-Ar29532- 29534). TAINAN City: 2♀, 1♂, Anping, Chiumaoyuan, alt. 5 m, 16 Apr.1983, coll. S.-H. Chen (NTNUBAr12791- 12793). KAOHSIUNG Co.: 3♀, Jiasian, Hean, alt. 400 m, 6 39.

(52) Oct. 2006, coll. H.-H. Liu (NTNUB- Ar29849-29851); 3♀, Meinung, Chutzmen, alt. 50 m, 4 Apr. 2003, coll. S.-H. Chen (NTNUB-Ar1646016462). KAOHSIUNG City: 1♀, 1♂, Kushan, Chaishan, alt. 100 m, 16 Sep. 2006, coll. T.-Y. Cheng leg. (NTNUB-Ar34338- 34339). PINTUNG Co.: 1♀, Shihtzu, Shuangliu, alt. 300 m, 6 Apr. 2007, coll. H.-H. Liu (NTNUB- Ar34337). ILAN Co.: 1♀, Suao, Yongle, alt. 200 m, 6 Apr. 2007, coll. H.-H. Liu (NTNUB-Ar29765); 1♀,Tatung, Hanchi, alt. 50 m, 13 May 2000, coll. S.-H. Chen (NTNUB-Ar604); 1♀, Chiaochi, Paoma Trail, alt. 300 m, 27 Oct. 1999, coll.W.-J. Huang (NTNUB-Ar490); 13♀, 2♂, Shanhsing, changpihu, alt. 200 m, 6 Apr. 2006, coll. H.-H. Liu (NTNUB- Ar29414- 29429). HUALIEN Co.: 1♀, Shoufeng, Liyutan, alt. 150 m, 3 Mar. 1996, coll.W.-J. Huang (NTNUB-Ar13034); 6♀, 2♂, Soufeng, lingting, alt. 50 m, 17 Feb. 1998, coll. S.-H. Chen (NTNUBAr9425-9432); 2♀, Yuli, Antung, alt. 150 m, 26 Aug. 2001, coll. S.-H. Chen (NTNUB-Ar10479-10480); 3♀, 1♂, Juisui, 3.5 Km SE Fuyuan, alt. 100 m, 18 Feb. 1998, coll. S.-H. Chen (NTNUB-Ar9415-9418). TAITUNG Co.: 1♀, Peinan, Chiafeng, alt. 150 m, 27 Aug. 2001, coll. S.-H. Chen (NTNUB- Ar10952); 2♀, 1♂, Tungho, Tulan, alt. 100 m, 27 Jun. 2003, coll. W.-J. Huang (NTNUB-Ar18327-18329); 1♀, 1♂, Chihshang, 2 Km S Wanan, alt. 250 m, 23 Jun. 2002, coll. S.-H. Chen (NTNUB-Ar11759-11760). Diagnosis Leucauge blanda resembles L. celebesiana by having similar leafvein-liked pattern of dorsal abdomen (Figs. 9A-B), but can be distinguished from the latter by having many brown denticles on the 40.

(53) venter of femora I, II, and tibiae I, II in male (Fig. 10F) that are absent in the latter. Opening of female copulatory duct sclerotized with an arch-shaped ridge is also diagnosized (Fig. 9D).. Description Female (NTNUB- Ar 29765). Total length 7.20: cephalothorax length 2.70, width 1.90, height 1.50; abdomen length 4.85, width 2.40, height 2.50. Measurements of palpus, legs, and spinnerets: palpus 0.26 (0.08, 0.03, 0.06, 0.09); leg I 17.90 (5.00, 5.90, 5.50, 1.50); leg II 13.70 (4.00, 4.20, 4.20, 1.30); leg III 7.0 (2.30, 1.90, 1.90, 0.90); leg IV 11.60 (3.80, 3.90, 3.40, 0.50); ALS 0.46; PLS 0.38. Carapace yellowish brown in alcohol, greenish in life, with a gray margin on each side and black eye margins; cephalic region slightly higher than thoracic region; thoracic groove deep, trifid posteriorly (Fig. 9A). Diameters of eyes in ratio, AME: ALE: PME: PLE = 0.12: 0.10: 0.12: 0.10. Length of MOA in ratio, MOA-L: MOA-AW: MOA-PW = 0.40: 0.36: 0.36. Clypeus height 1.4 times diameter of AME. Chelicerae brown; fang dark brown; promargin of the fang groove armed with 3 robust triangular teeth and retromargin with 4 teeth (Fig. 9C). Endite brown, longer than wide. Labium reddish brown, longer than wide. Sternum brownish, heart-shaped. Palpus and legs yellowish brown, with black spines and hairs. Femur IV with 18 pairs of trichobothria on prolateral surface of proximal half. Order of leg length I > II > IV > III. Abdomen oval with a pair of anterior humps, each with a large black spot; dorsum silvery with three main longitudinal black stripes and some oblique stripes origined from midline, posterior end of abdomen with two 41.

(54) black spots (Figs. 9A, B); venter yellowish brown with two parallel longitudinal silvery bands and having silver specking between the two longitudinal bands. Epigynum yellowish; without hood; copulatory opening arch-shaped, brownish and heavily sclerotized; posterior end of median septum wider than that of anterior (Fig. 9D). Spermatheca thin-walled with one large and two small chambers (Fig. 9E).. Male (NTNUB- Ar29766). Similar to female in shape and coloration (Fig. 10A). Total length 5.32: cephalothorax length 2.40, width 1.68, height 1.16; abdomen length 3.32, width 1.52, height 1.16. Measurements of palpus, legs, and spinnerets: palpus 2.75 (1.20, 0.34, 0.45, 0.76); leg I 24.08 (6.36, 8.08, 7.76, 1.88); leg II 17.28 (4.96, 5.84, 5.36, 1.12); leg III 7.68 (2.60, 2.16, 2.04, 0.88); leg IV 13.32 (4.28, 3.72, 4.20, 1.12); ALS 0.23; PLS 0.18. Diameters of eyes in ratio, AME: ALE: PME: PLE = 0.10: 0.12: 0.13: 0.10. Clypeus height 2.1 times diameter of AME. Promargin of fang groove armed with 3 teeth; first one minute and far from the others; retromargin with 4 teeth (Fig. 10B). Venter of abdomen with two silver longitudinal bands, lack silver speckling between the two longitudinal bands. Venter of femora I, II, and tibiae I, II, with many brown denticles, denticles of femur II less than those of femur I (Fig. 8F). Order of leg length I > II > IV > III. Tegulum wide equal to long; cymbial basal process short; paracymbium finger-shaped; conductor inflatable, pellet-shaped and apically projected; palpal tibiae shorter than cymbium (Figs. 10C-E). 42.

(55) Variations Five females and five males from Taipei Co. (Chungho) were measured. Variations among females are followed by those of males (with the mean in parentheses). Total length 6.48-8.42 (7.56) and 4.41-5.45 (4.73); carapace length 2.48-3.07 (2.67) and 1.81-2.48 (2.15), width 1.74-2.32 (1.98) and 1.47-1.83 (1.63); height 1.32-1.68 (1.50) and 1.01-1.82 (1.35); abdomen length 4.26-6.41 (5.39) and 2.67-3.40 (3.06), width 2.21-3.39 (2.85) and 1.32-1.75 (1.47), height 2.47-3.38 (2.98) and 1.30-1.67 (1.45).. Distribution Russia, China, Japan, Korea, and Taiwan.. Remarks Leucauge blanda was recorded from Taiwan by Nakajima (1921) for the first time. It is one of the most common and widely distributed species in the lowlands of Taiwan (Fig. 26).. 43.