Out-of-Africa again: A phylogenetic hypothesis of the genus Charaxes (Lepidoptera: Nymphalidae) based on five gene regions文獻來源： Aduse-Poku K, Vingerhoedt E, Wahlberg N. 2009. Out-of-Africa again: A phylogenetic hypothesis of the genus Charaxes (Lepidoptera: Nymphalidae) based on five gene regions. Molecular Phylogenetics and Evolution 53 :463–478
雖然螯蛺蝶屬（Charaxes）長期在採集者與研究者之間受到歡迎，其演化歷史大多仍然未知。近期被接受的種群和屬內近緣種是專斷地以成蟲形態與生活史 為基礎，作者在本研究中以來自五個基因區域（1粒線體基因＋4核基因）共 4167 bp序列資料檢驗螯蛺蝶屬內的單系性與演化上的親緣關係，並探討其近緣屬Euxanthe, Polyura（雙尾蝶屬）與 Palla之間的關係。在此提議的親緣關係架構中，作者評估屬級的分化年代並重建其歷史上的生物地理。作者採用包括所有非洲與亞洲的已知種群、所有已知的 Euxanthe與 Palla物種和兩個Polyura的代表種作為研究對象代表，研究結果發現螯蛺蝶屬（Charaxes）為一個駢系群、與Euxanthe 和Polyura有關聯，此結果和以往所認為的單系群假設相反。作者發現以形態定義的16個種群 （超過一個物種）中有13個為有強力支持的單系分支，Charaxes nichetes為其他Charaxes物種的姊妹群。 Polyura與Zoolina 和 Pleione 種群形成一個有良好支持度的支系，而Euxanthe則與Lycurgus 種群歸在一群。作者的結果指出Charaxes的共同祖先是在始新世中期（約4500萬年前）於非洲（中部）由Palla的共同祖先分化而來，並在1500萬年之後開始分化岀其他物種成員。此屬大部份的多樣性在漸新世晚期與中新世間、全球氣候發生強烈的波動時開始 增加。相當大量的衍生物種於上新世時從其姊妹種演化而來。播遷-割據分析推測在其衍生物種的分布過程中、播遷事件的發生多於割據事件。 Polyura與印澳區的Charaxes最可能是非洲的Charaxes於中新世在亞洲的三個獨立拓殖之結果。作者將以下兩屬處理為Charaxes的 同物異名：Polyura (syn. nov.)與Euxanthe (syn. nov.)，並將新定義的 Charaxes劃分為五個亞屬以反應其親緣關係。
Abstract
Despite the long popularity of Charaxes among collectors and researchers, their evolutionary history is largely unknown. The current and accepted species groupings and relationships within the genus are based exclusively on adult morphology and life histories. Here, we examine the monophyly and evolutionary affinities of the species-groups within the genus Charaxes and explore how they relate to members of their closest genera (Euxanthe, Polyura and Palla) using 4167 bp of sequence data from five (1 mitochondrial and 4 nuclear) gene regions. Within the proposed phylogenetic framework, we estimate ages of divergence within the genus and also reconstruct their historical biogeography. We included representatives of all known species-groups in Africa and Asia, all known species of Euxanthe and Palla and two exemplar species of Polyura. We found the genus Charaxes to be a paraphyletic group with regard to the genera Polyura and Euxanthe, contrary to the earlier assumption of monophyly. We found that 13 out of 16 morphologically defined species-groups with more than one species were strongly supported monophyletic clades. Charaxes nichetes is the sister group to all the other Charaxes. Polyura grouped with the Zoolina and Pleione species-groups as a well-supported clade, and Euxanthe grouped with the Lycurgus species-group. Our results indicated that the common ancestor of Charaxes diverged from the common ancestor of Palla in the mid Eocene (45 million years ago) in (Central) Africa and began diversifying to its extant members 15 million years later. Most of the major diversifications within the genus occurred between the late Oligocene and Miocene when the global climates were putatively undergoing drastic fluctuations. A considerable number of extant species diverged from sister species during the Pliocene. A dispersal–vicariance analysis suggests that many dispersal rather than vicariance events resulted in the distribution of the extant species. The genus Polyura and the Indo-Australian Charaxes are most likely the results of three independent colonizations of Asia by African Charaxes in the Miocene. We synonymize the genera Polyura (syn. nov.) and Euxanthe (syn. nov.) with Charaxes, with the currently circumscribed Charaxes subdivided into five subgenera to reflect its phylogeny.
Despite the long popularity of Charaxes among collectors and researchers, their evolutionary history is largely unknown. The current and accepted species groupings and relationships within the genus are based exclusively on adult morphology and life histories. Here, we examine the monophyly and evolutionary affinities of the species-groups within the genus Charaxes and explore how they relate to members of their closest genera (Euxanthe, Polyura and Palla) using 4167 bp of sequence data from five (1 mitochondrial and 4 nuclear) gene regions. Within the proposed phylogenetic framework, we estimate ages of divergence within the genus and also reconstruct their historical biogeography. We included representatives of all known species-groups in Africa and Asia, all known species of Euxanthe and Palla and two exemplar species of Polyura. We found the genus Charaxes to be a paraphyletic group with regard to the genera Polyura and Euxanthe, contrary to the earlier assumption of monophyly. We found that 13 out of 16 morphologically defined species-groups with more than one species were strongly supported monophyletic clades. Charaxes nichetes is the sister group to all the other Charaxes. Polyura grouped with the Zoolina and Pleione species-groups as a well-supported clade, and Euxanthe grouped with the Lycurgus species-group. Our results indicated that the common ancestor of Charaxes diverged from the common ancestor of Palla in the mid Eocene (45 million years ago) in (Central) Africa and began diversifying to its extant members 15 million years later. Most of the major diversifications within the genus occurred between the late Oligocene and Miocene when the global climates were putatively undergoing drastic fluctuations. A considerable number of extant species diverged from sister species during the Pliocene. A dispersal–vicariance analysis suggests that many dispersal rather than vicariance events resulted in the distribution of the extant species. The genus Polyura and the Indo-Australian Charaxes are most likely the results of three independent colonizations of Asia by African Charaxes in the Miocene. We synonymize the genera Polyura (syn. nov.) and Euxanthe (syn. nov.) with Charaxes, with the currently circumscribed Charaxes subdivided into five subgenera to reflect its phylogeny.
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