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Insecta:Hymenoptera: Vespidae:Vespinae

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Insecta:Hymenoptera: Vespidae:Vespinae JapaneseJapaneseSociety Society ofSystematicZoologyof Systematic Zoology Species Diversity,2011, 16, 65-74 Phylogenetic Analysis and Biogeography of the Nocturnal Hornets, Provespa (Insecta: Hymenoptera: Vespidae: Vespinae) Fuki Saitoi:2 and Jun-ichi KQjima2 iIlostcloctoral IJlellow of the .Jtipan Society for the Promotion of Seience E-mail:[email protected] 21Victtural liistory Laboratoro,, decutty of Science, ibaraki [1itiversity, Mito, 31a-a512 Jctpan (Received 30 March 2010; Aecepted 21 September 2010) Relationships among the three species of nocturnal hornet of the genus Provespa Ashmead, 1903 are cladistically analyzed based on adult morpho- logical characters and mitochondrial DNA sequence data. Monophyly of Provespa is supported and the species relationships are expressed as (P. barthelempi+(P, anomala+P. nocturna)). A"ovespa bartheleras)i (Du Buyssen, 1905) is distributed mainly in the southeastern part of the Asian continent from easterri lndia to Indochina, while P. nocturna Vecht, 1935 and P. anomala (Saussure, 1854) occur mainly in Sumatra, Borneo, and the southern part of the Malay Peninsula, [[1ie speciation and biogeography of Provespa are briefly discussed, with reference te a supposed vicariance event around the Isthmus of Kra. Key Words: Vespidae, Vespinae, Provespa, phylogeny, biogeography, South- east Asia. Introduction Wasps of the vesptne genus Provespa Ashmead, 1903 are nocturnal and found new colonies as a swarm of workers accompanied by a single queen (Matsuura 1991). This genus, consisting of the three species P. anomata (Saussure, 1854), P. barthelernyi (Du Buysson, 1905), and P. nocturna Vecht, 1935, is distributed in southern Asia from eastern India in the west, through indochina and southern China, to Sumatra, Borneo, and Java in the east, Archer (2000) analyzed three morphological characters (apex of the aedeagus, tyloides of the male antenna, and anterior margin of the clypeus) cif the three species of Provespa and proposed on this basis that the relationship among them could be expressed as (P, bartheleTnyi+(R anomalatP. noctuma)). These three "were characters were those that found whose character states could be polarized by outgroup comparison" (Archer 2000: 127), but hrcher did not specifY any ouV group. He did summarize the distribution records of the three species and stated, "the refemring to their phylogenetic relationships, that ancestor of PTovespa first split tnto a mainland species (P barthelempi) and an island species which later split into two species (P. anonzala and P. nocturna)." In the present paper we conduct a cladistic analysis of Provespa based on botih molecular and morphological data. Referring to these results and the historical ge- NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology 66 F, Saito and J. Kojima ography of the regions inhabitect by these wasps, we also briefiy reconsider the bio- geography of the genus, Materiais and Methods Specimens from which legs were taken for DNA extraetion are deposited in the Natural History Collection of Ibaraki University, Mito, Japan (abbreviated as "ruNH"). Dried pinned specimens used in coding the morphological characters and those examined to add distribution records [we give only their localities and depositories here] are deposited in the IUNH, the Museum Zoologicum Bogoriense, Bogor, Indonesia (MZB), and the National Institute fbr Agro-Environmental Sci- ences, Tsukuba, Japan (NIAES). The speeies included in our cladistic analyses were the three mentioned species of Provespa; four species of three other vespine genera (Vk?spa analis Fabri- cius, 1775, V basagts Smith, 1852, Dolichovespula media (Retzius, 1783), and Vlespula shiclai Ishikawa, Yamane and Wagner, 1980); and two species of Polistes (Poltstes chinensts antennalis Perez, 1905 and P. jmponicus Saussure, 1858) in the Polistinae, which according to Pickett and Carpenter (2010) is the sister group of the Vespinae. Polistes chinensts antennalts was used to root the trees. Morphological data The merphological characters given below, together with their states and codes (Table 1), were extracted from Caxpenter (1987), Archer (1994, 20oo), Carpen- ter and Perera (20os), and Pickett and Carpenter (2010). We observed their states on pinned and dried specimens under a stereoscopic microscope, except that the male genital characters were observed on dissected-out genitalia in glycerin under a stereoscopic microscope andlor observed wiih a scamiing electron microscope (ABT 32, Topcon Technohouse) after coating with osmium by an osmium coater "non-additive" (Neoc-st, Meiwafbsis). All characters were coded as additive unless is specified in parentheses after the character name, Ilorewing 1. Prestigma length: prestigma shorter than pterostigma=O; prestigma longer than pterostigma, but less than 2,5 times longer=1; prestigma approximately three times or more longer than pterostigma:=2 2. Length of first submarginal ceil: shorter than distance from apex of cell to apex of wing=O; longer than this distance=1 3 Base of seeond submarginal cell: more or less angled, M oriented obliquely with respect to m-cul=O; barely angled, M oriented vertically=1 4 Placement of meu2: elose to r-m2=O; distantly separated from r-m2=1 Hindwing 5 Hamuli placement: beginning basad ef fork of Rl and RS=O; beginning at this fork:=1 6 Jugal lobe: present=O; absent:=1 7 Axillary incision: present=O; at)sent==1 Male antenna 8 Tyloids (non-additive): narrow ridge or flattened rectangle==O; two on each fiagellomere==1; absent==2 NII-Electronic Library Service Japanese SooietySociety of SystematicSystematio Zoology Phylogeny and biogeography of Provespα 67 お HHOOHHHH 目 霧 oo 目 Hco ooo 一 0 う 〇 O 劇っ 寸 Φ 9 う 崘 寸 守 寸 專 H 鬥 OOoHoeo 爵 ゆ の ゆ 蕊 蕊 蕊 QD ゆ 寸 ゆ 爵 鎔 露 Q つ 03 σ9 o っ QD 器 路 g コ 田 α ⊃ N OOH 同 HNH 祠 H 旨 恥 の 鴫 ゆ ゆ 玲 頃 口 【m ［m 国 m 国 餡 m 偉員 ロロ ＜ ＜ く 〈 く く ぐ く く 鵠 H 鬥 oo 蔚 ⇔ ooo OOHH ゆ 寸 a り N 崘 H ooN 国 囚 一 赳 目 鵠 oo 一 目 鬥 oooo ． O ⇔っ 目 N oo 州 HH 目 目 H 目 口 鴎 ト oコ OO oっ OQ 切 ゆ ゆ 靄 お 囲 自 讒 O 8 ． 〇 OOoOH 閣 ooo の 一 ， 鴎 瞼 零 留 靄 爵 頃 爵 紹 OD OD QQ QQ ◎∋ Q20 g っ 旨 』 リワ ＜ 頃 の ゆ の 嶋 ゆ の 崘 リDOOO 蝸 鬥 祠 目 Z 国 餡 田 m 箇 m NO oH oooo o ρ 国 舅 頸 く く く ＜ ＜ く 〈 く ＜ ooo 鬥 囚 N ＜ 自 ooooOaH 自 目 咽O の ◎3H の oo − HHN 国 凶 N OO g ワ 』 卜 OoOOOOH 鬥 H 〇 H 邸 一 〇 図 ト 為 QQ 鋤 O H 国 ◎つ ぐ ゆ ゆ ゆ の 〇 H 目 OoH 肖 ooo 口 雪 HOQ ま 霹 蕊 ま 鴇 』 蝿 め 鴫 鳴 ゆ の 而 仁口 09 瑟 場 駆 ◎Q 9◎ 鴇 ◎つ ◎コ 謂 旨 OOOOOOHHH 口 鴫 問 め ゆ 頃 ゆ 頃 玲 ゆ 〔ワ ρq ρ自 凶 自亀 ［m 【m 自q ［m ρ籍 Q ＜ く く く ＜ ＜ ＜ ＜ く コ OOH − ooooo ＼ qm o つ 1ooIOOHHooo ロ ， 同 o 噛 ≧ 一 国 q ロ 鬥 HHH 圏 N ＼ 隔 ｛ 腎 σ5q H 自 oo 囲 HHH 目 目 H q 目 ∩ 邸 O ∩ O oo σ『 − HHHH 郭 目 眉 毫 （ ぢ 同 自 笛 oo 目 鬥 HH 闔 H 臣 Φ Od 頸 ごpO 魯 肩 絹 り o コ oo 目 H 囚 H HN 鬥 薯 ＞ 鳶 詳 防 O 謂 嵩 一昏 』 邸 O 鵠 ト OOHH 州 HHH 同 山 角 q 台 鎌 も ← 冒 日 喟 o 謂 ¢ h 蜀 20h 罷 あ 器 ‘ Φ OOHHH 目 H 鬥 H ⇔ 8q 図 β 脚 謀 蠶国 【O 自 〇 齧 ．「〉 肩 一 顰 2 圏 軌 、 肩 崘 oooo 目 HH 鬥 一 』 頒 円 羣 匿 薯 O 隅 客 の 轟 噂 O OO 、 ． ． 軌 ． の Φ 謂 藹 o 弓 祠 HoHH 一 口 q d9 寸 oo 目 ← ） 冩 Oq 窪 蠧 5 冪昆靂 も q 曾 巴呂 一 冨 の o っ oooo 目 鬥 HHH d 蠶 留 扁 』 弓 h 鐚 乞 哨 鐚 日 〉 屠 団 剰 旨 O 朝 OOOOOoH 同 H 邸 屑 』 邸 ε 一 OON 製 HH 目 HH の 月 （ OHQO 凵昏 卜 円 稔 8N 。う 亳 鴨 煽 ） Φ ミ 一自 削§ 日 ミ 唱 § 菴 』 Φ 幅 ρ § 穐 N 醤 趨 毬 ｝ 穏 唱 』 心 漣 唱 鬥 き 眉 む 邸 り ミ 始 8 翰 §む 。， 謹 的 鎗 蒿 霆 ミ ミ 口 Φ 6 惹 驀 仁 総 認 ミ 黛 輪 の k O 稔 N o “ ミ ゜。 邸 ち ミ N o 橇 N ◎ 選 遣 の § 匿 篷 N §粒 。 転 遒 鴨 。 ミ 葺 恥 黛 O 」 〇 O 摘 選 暮 遣 も 防 嵩 90 一 ミ ミ 彑 O 超 毫 尊 篳 o 芯 竃 齪 OQO 〇 身 ミ 魯 e Q 霆 尊 陵 §ミ リ 帽 ◎ 湧 ミ 魯 竃 、 む 冨 身 § リ 軸 画 畧 o 竃 、 建 鳶 む 恥 § oS ∩ 急 の § ミ 剰q ． 劬 吻 弓 忘 O 旨 鋤 魯 ミ ． 遣 越 鼕勉 § H ミ ミ ミ 野 勢 毬 葛 O o 註 恥 軸 き 幅 高 物 稔 自 OO 島 N 鵡 § 8 O § 堯 稔 〇 Φ 一 凝 、 魯 』 O Ok 一 P 遠 ρ Oq ← § ρ 母 馬 § 量量q ミミ乱 σコ £ £ 〉 量§ ミ螽へ 邸 【〔 臼 臼 尸 一 NII-ElectronicN 工 工 Eleotronio Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology ce F. Saito and J. Kojima Male head 9 Apicai margin of clypeus: convex=O; nearly straight or very shailowly emar- ginate medially=1; distinctly emarginate medially=2 Iilemate hend le Interantennal space: broad, rounded=O; triangular=1; strongly raised==2 11 Dorsa:L margin of clypeus: nearly straight=:O; distinctly bisinuate=1 12 Apex of clypeus (Figs 1, 2): pointed==O; shallowly emarginate=1; deeply emar- ginate=2 13 Clypeal lateroapical angles: blunt, rounded=O; triangular=il 14 Vertex length: ocelloccipital distance shorter than length of ocellar triangle==O; ocelloecipital distance longer than Iength of oeellar triangle=1 15 Ocelli: diameter less tihan distance between posterior ocellus and eye=O; diam- eter distinctly greater than this distance=1 16 Occipital carina (ventrally): reaching to base of mandible==O; effaced near mandibular base:=1 17 Occipital carina (dorsally): present dorsolaterally==O; absent dorsolaterally=1 18 Mandibular teeth: pointed==e; with elongate cutting edge of less than twice length of apical part==1; with strongly elongate cutting edge of twice or more length of apical part=2 Mesosoma 19 Pronotal carina: complete==O; dorsally reduced=1; present only as lateral rem- nant=2; absent=3 20 Pretegular carina: complete=O; ventrally ethced=1; absent==2 21 Female scutum: distinctiy longer than wide==O; nearly as wide as long==1 22 Scutal lamella: present=O; absent=1 23 Metanotai lobe: absent.=O; present posteromedially=1 24 Hind coxal carina: absent=O; weak or as traces= 1; present, strong==2 Male metasoma 25 Seventh tergum apex: rounded to straight=O; sltghtly emarginate==1 Male genitalia 26 Aedeagus apex (non-additive): little differentiated= O; transverse, projecting laterally=1; triangular=2; elengate==3; narrow==4; subcircular=5 27 Aedeagal apical indentation: deep =O; shallow=1; absent=2 28 Aedeagus width (non-additive): narrew threughout=O; as wide as or wider api- caily than at midiength=1; narrower apically than at midlength=2 29 Aedeagal medial lobes: absent=O; present==1 30 Paramere precess (non-additive): absent==O; broadly infiected=1; pointed=2; finger-like=3 31 Digital apex prqjections: present=O; absent=1 Figs 1, 2.
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