根據序列與形態分析顯示尖翅翠蛺蝶與E. jpona是同種, E. phemius降為尖翅翠蛺蝶的一個亞種

文獻來源: Yago M, Yokochi T, Kondon M, Braby MF, Yahya B, Peggie D, Wang M, William M, Morita S, Ueshima R. 2012. Revision of the Euthalia phemius complex (Lepidoptera: Nymphalidae) based on morphology and molecular analyses. Zoological Journal of the Linnean Society 164(1): 304-327.

Abstract
Adults of the Euthalia phemius complex, which is composed of three South-East Asian nymphalid species, Euthalia phemius, Euthalia ipona, and Euthalia euphemia, were genetically analysed by examining mitochondrial and nuclear genes. The E. phemius complex was also examined morphologically, with particular emphasis on wing markings and male genitalia. No significant differences amongst the three species in the complex were detected with respect to either genetic distance or genital morphology. We therefore conclude that the three currently recognized Euthalia species belong to a single species. Accordingly, E. ipona is synonymized with E. phemius. Euthalia euphemia is treated as a subspecies of E. phemius. Type specimens of all taxa and a synonymic list for the E. phemius complex are also given. In addition, we briefly discuss the evolution and biogeography of the species complex.

由形態特徵重建新熱帶美洲蛺蝶Hamadryas屬之親緣關係並著重發聲結構的失去與其他替代性別辨識機制產生的討論

文獻來源：Garzo-Orduna, I. J. 2011. Phylogenetic evidence for loss of sound production and a shift in sexual recognition signals in Hamadryas butterflies (Nymphalidae: Biblidinae). Systematic Entomology. DOI: 10.1111/j.1365-3113.2011.00599.x

簡介

分布於新熱帶美洲的Hamadryas屬為蛺蝶科Biblidinae亞科類群，共計有20個物種，當中約有近半數具有前翅發聲結構、少數具有雌雄翅紋二型性與雄蟲具發香鱗片的種類。由過去的生物學顯示此屬辨識特徵尚有翅背面如花部狀紋路、吸食腐敗果實、停棲時頭下尾上以及飛行路徑飄忽不定。

過去30年間雖然有此屬的屬內歸群處理，然而尚未有以支序學所建立的親緣關係研究，本文章即首次以形態特徵重建親緣，並討論以下議題：(1) Hamadryas是否為一單系枝系，具有哪些衍徵來支持？(2) 此屬姊妹群為何? (3) 評估過去所建立亞屬之單系性；(4) 演化上如發聲、二型性與發香鱗等特徵出現與先後的判定與討論。

結果討論顯示

(1) Hamadryas屬為單系群；

(2) 此屬姊妹群為Ectima屬；

(3) 除了H. laodamia枝系外其餘過去之屬內亞屬歸群皆非單系群；

(4) 在發聲、二型性與發香鱗等特徵的演化上，發聲結構在特徵分析中被拆解為獨立的四個前翅翅脈特徵，然而能夠發聲的行為僅產生過一次(單一起源)，該枝系當中的H. laodamia與其近緣種則再次失去此發聲結構。雖然H. laodamia枝系失去發聲能力，此支系卻擁有雄蟲發香鱗與翅型/翅紋雌雄二型性的特徵。作者因此認為若發聲行為是作為物種辨識用，則失去發聲行為的枝系則以發香鱗與二型性等特徵替代辨識。目前尚不清楚此三特徵組的出現順序，待後續研究。

(5) 藉由親緣關係樹所得之雌雄二型性特徵分佈，作者也比對此結果是否符合過去三個雌雄二型性起源假說(註)之一。由於雌雄二型性枝系H. laodamia的雌蟲翅紋為祖徵，而雄蟲翅紋為衍徵，因此雖然沒有直接的證據支持，此結果仍較符合達爾文所提出的雌蟲偏好特定雄蟲特徵的假說；

(6) 由於所重建之親緣關係樹整體樹型支持度不高，後續仍有待幼蟲形態資訊以及分子證據的加入，此外對於雌雄蟲辨識所利用特徵的方式，則需要更詳盡的野外觀察與網室試驗來解釋。

註：過去三個主要之雌雄二型性起源假說

(1) 達爾文 (1971) - 性擇假說，由雌性偏好與特定雄性產生後代的選汰方式達成；

(2) 華萊士 (1889) - 天擇假說：雌性儘可能產生隱蔽自身的形狀、斑紋等(相對於祖先型者)來增加適存值；

(3) Silberglied (1984)- 雄性間互動假說：雄性藉由視覺來辨識其他雄性個體的物種異同，並用以對同種雌性進行展示。

Abstract

The neotropical butterfly genus Hamadryas Hubner comprises 20 species that exhibit an intriguing variation in their natural history traits. Although revised in 1983, no phylogenetic hypothesis was presented: the first phylogenetic hypothesis is estimated here based on 93 characters and including species from the three other genera in the tribe Ageroniini. The phylogeny is used to test the monophyly of the genus, establish the sister group of Hamadryas and identify its apomorphies. The tree allows the inference of patterns of character change in sound production and sexual dimorphism. Implied weights show that Hamadryas is monophyletic and corroborate Ectima Doubleday as a sister genus. Previously suggested subgenera for Hamadryas were non-monophyletic, with the exception of the laodamia clade, supported by the presence of a complete sterigma. Sound production is inferred to be a derived condition in Hamadryas that has been lost in the laodamia clade. This, plus the presence of androconial organs and sexual dimorphism in the laodamia clade, suggests a shift in sexual recognition signalling. Furthermore, the phylogeny indicates that the colour pattern of males in the laodamia clade is novel, supporting a Darwinian origin of sexual dimorphism.

以全球紅蛺蝶探討廣域分佈類群與其侷限分佈姊妹群之種化形成

Photo by Dumbou

文獻來源: Wahlberg N. and D. Rubinoff. 2011. Vagility across Vanessa (Lepidoptera: Nymphalidae): mobility in butterfly species does not inhibit the formation and persistence of isolated sister taxa. Systematic Entomology 36(2): 362–370. [Abstract]

簡介

異域種化假設了族群間受地理隔離而無法進行基因交流，最終形成兩個獨立的分類群，因此對於廣域且活動能力強的物種來說此機制似乎無法形成隔離且侷限分佈的姊妹種。蛺蝶科紅蛺蝶屬(Vanessa)包含了許多世界性廣域分佈物種，卻也擁有不少局限分佈種，因次此屬成為測試此假說理想的材料之一。此研究藉由涵蓋全球紅蛺蝶屬與其姊妹群的取樣，使用粒腺體COI與8段核基因序列，進行親緣關係的重建與分子鐘枝系分歧的時間推估，所獲得的結果為: 1. 各基因所重建的親緣關係樹見少有衝突，且樹型符合過去形態證據所劃分的烏個紅蛺蝶種群; 2. 外群Antanartia成為駢系群，其中改屬兩物種在此研究中與Vanessa形成單系群，故以此將其轉屬，現用組合名為V. hippomene以及V. dimorphica，現今紅蛺蝶共計22種; 3. 由生物地理分析的結果呈現多數的侷限分佈種，其姊妹種為廣域分布種，因而顯示擴散分布與異域種化的存在。然而就分子定年的結果顯示這些分枝分化時間都相當久(大於500萬年)，此結果支持了長時間的隔離分化。然而作者認為紅蛺蝶屬的高活動力與異域隔離在此研究中，並無法完全解釋此屬在系統發生上的成因。

Abstract

Allopatric speciation is thought to occur in the absence of gene flow, thereby suggesting that widespread vagile species might be less likely to generate restricted sister taxa because of a lack of isolation. The butterfly genus Vanessa provides an ideal test of this concept, as it contains some of the most cosmopolitan and vagile species of butterflies on the planet, as well as some highly restricted taxa. Given the age of these groups, this arrangement offers a special opportunity to examine the relationship between vagility and phylogeny in generating novel taxa; specifically, does the vagility of some lineages impede allopatric speciation, leaving restricted clades more speciose? A phylogenetic hypothesis is proposed for all species belonging to the butterfly genus Vanessa based on DNA sequences from one mitochondrial and eight nuclear gene regions. The resulting topology shows very little conflict among gene regions, with five well-supported clades corresponding to morphologically consistent species groups. The data very strongly indicate a polyphyletic genus Antanartia, and thus to preserve monophyly two species previously assigned to Antanartia are transferred to Vanessa, Vanessa hippomene comb.n. and Vanessa dimorphica comb. n., resulting in a total of 22 species placed in Vanessa. A biogeographical analysis shows that in many cases the most geographically restricted species are sister to geographically widespread species, suggesting dispersal and allopatric speciation. Surprisingly, in almost all cases the divergences between widespread and restricted species are quite old (>5 Ma), suggesting long-term isolation and stability of both vagile and sedentary species, despite the high (even intercontinental) vagility of many extant species and, by extension, ancestral species. The biogeography of Vanessa suggests that species vagility and allopatry do not fully explain the forces governing cladogenesis in this remarkable genus.

棋盤格蛺蝶生態上分化之族群間的後生殖隔離

Fig. 3D圖顯示純種與雜交個體的偏好，圖片引用自http://www.plosbiology.org

文獻來源: McBride CS. and Singer MC. 2010. Field Studies Reveal Strong Postmating Isolation Between Ecologically Divergent Butterfly Population. PLoS Biology 8(10): 1~17 (full article)

簡介

外在的後生殖隔離機制(extrinsic postzygotic isolation, EPI)被認為是物種種化必經的一個起始過程，但相較於研究詳盡的前生殖隔離機制(prezygotic isolation)，因EPI出現的時間較短(註一)以及局限研究方式(註二)，使得人們對於EPI所知並不多。在McBride & Singer (2010)的研究中，藉由穩定適應不同寄主植物且異域的棋盤格蛺蝶(Checkerspot butterfly, Euphydryas editha)進行"野外"的實驗，透過雜交個體在自然環境下的適應情況來探索EPI，但不同於以往監測整體的適存度(比如存活率)，作者們先推測雜交個體應該在各方面都位居於兩異域的偏好棋盤格蛺蝶之間，而野外觀測到雜交個體的幼蟲寄主偏好、取食位置的偏好、產卵的位置偏好以及單一次產卵量等，也都如預期在兩個異域的棋盤格蛺蝶間，再藉由野外測定生長速率、卵的孵化率以及整體的存活率，作者們發現雜交個體都相對於純種來的差。更進一步將純種與雜交個體拓殖到所適應的寄主植物之外，發現純種以及雜交種在新拓殖的植物在各方面並無差異，藉以證實雜交確實是受到EPI的影響而非內在生理的影響。作者們亦選用了三個因子繪製成模擬棲位(niche)的3D圖(見上圖)，在所構成的平面上兩純種有各自聚集成兩個頂點，而雜交種則是如同預期般散布在兩純種間。雖然目前不能很清楚了解什麼原因導致雜交個體在自然環境適存度降低，但EPI的確在自然界影響非淺。

註一、雜交個體可能因為較不適應環境而滅絕。

註二、大多的實驗都在實驗室進行或採用改良式的田野實驗，因此會造成很多因子被忽略，比如說天敵的因子可能就會被排除，因此在相關研究通常會低估或忽略EPI的影響力。

印度-太平洋區的雙尾蝶屬種類之演化歷史與系統分類

文章來源:Muller CR, Wahlberg N, Beheregaray B. (2010) 'After Africa': the evolutionary history and systematics of the genus Charaxes Ochsenheimer (Lepidoptera: Nymphalidae) in the Indo-Pacific region. Biological Journal of Linnean Society, 100: 457-481 [abstract]

相關閱讀：Aduse-Poku K, Vingerhoedt E, Walhberg 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. [abstract]

雙尾蝶屬(Charaxes)是一個高多樣性且吸引眾多鱗翅學家的蝴蝶類群，至目前為止約包含250種分為5個亞屬，其中約220種分佈在非洲區，31種分佈在印澳區。本篇針對分佈在印澳區的31種，基於COI, ND5與wingless三個基因分析其親緣關係與其生物地理歷史。根據以最大簡約法與貝氏分析法的結果顯示除了C. solon之外，印度太平洋(Indo-Pacific region)的種類為一單系群。分析中確認了印度太平洋區種類的八個主要的支系，包含solon + African, elwesi, harmodius, amycus, mars, eurialus, latona, nitebis與bernardus支系。除了其他基於主要以後翅型態為基礎的研究，這是第一個分析非洲區以外的雙尾蝶親緣關係。生物地理的分析顯示，印度太平洋地區的種類在非洲起源後，在這個區域的演化歷史，特別是在華萊士區，與其地質事件與氣候歷史有密切的關係。本研究推測所有華萊士區以東的雙尾蝶(除了C. solon)的起源是在華萊士區。最早的印度太平洋區支系從就大陸分化的時約為一千三百萬年前。這個區域的雙尾蝶的分化主要與Pliocene的氣候變遷有關，並且可能近至Pleistocene。雖然播遷與隔離在這個區域雙尾蝶扮演同樣重要的角色，後者對於華萊士區的雙尾蝶意義較為重大。

斑豹蛺蝶屬之分類綜述- 物種描述、模式標本影像與相關文獻

文獻來源：JC Dunford. 2009. Taxonomic overview of the greater fritillary genus Speyeria Scudder and the atlantis - hesperis species complexes, with species accounts, type images, and relevant literature (Lepidoptera: Nymphalidae). INSECTA MUNDI 0090: 1-74. [full text PDF]

本文為斑豹蛺蝶屬 Speyeria Scudder (= Argynnis Fabricius) 之綜述，並說明其置於蛺蝶科(Nymphalidae)毒蝶亞科(Heliconiinae)之原因。文中對於16個已被命名的斑豹蛺蝶屬物種以及 Speyeria atlantis (Edwards) 與 Speyeria hesperis種群下的(Edwards)25個亞種，均有物種描述、模式標本影像與相關文獻。本文指出過去命名上的錯誤，並更新分類處理、生活史與分布資訊，同時對模式標本產地資訊進行討論。最後，原有一亞種Speyeria atlantis hanseni Emmel, Emmel, and Matoon, 1998, 被處理至另一物種之下，故有新學名組合產生：Speyeria hesperis hanseni Emmel, Emmel, and Matoon.

Abstract.

A taxonomic overview for the fritillary genus Speyeria Scudder (= Argynnis Fabricius) and its placement within the Heliconiinae (Nymphalidae) is presented. Taxonomic accounts, type images, and relevant literature for the 25 subspecies within the Speyeria atlantis (Edwards) and Speyeria hesperis (Edwards) complexes and the 16 nominate Speyeria species are included. Errors in nomenclature are identified, taxonomic, life history, and distributional information are updated, and type locality information is discussed. Images of primary type specimens for all 16 Speyeria species and the 25 subspecies in the atlantis - hesperis complexes appear together in color here for the first time. One new combination of a species-subspecies is created: Speyeria hesperis hanseni Emmel, Emmel, and Matoon, 1998, new combination, which was previously Speyeria atlantis hanseni Emmel, Emmel, and Matoon.

Key words. Argynnis, bibliography, fritillary, Lepidoptera, North America, Nymphalidae, Speyeria, synonymy, taxonomy, types

最新的證據顯示蛺蝶科的多樣化起源自接近白堊紀/第三紀的過渡時期

文獻來源：Wahlberg, N., Leneveu, J., Kodandaramaiah, U., Pena, C., Nylin, S., Freitas, A. V. L. and Brower, A. V. Z. (2009) Nymphalid butterflies diversify following near demise at the cretaceous/tertiary boundary. Proc. R. Soc. B. doi: 10.1098/rspb.2009.1303

簡介

蛺蝶科分布廣泛，在生態系中為重要的植食性昆蟲，也同時是除了果蠅外的重要模式物種，在演化發育、擬態生物學等領域扮演重要角色。由於蛺蝶科為泛世界型分布的種類，其分化年代在本文章之前，僅有針對亞科級或是屬級進行定年，因此對於蛺蝶科的分化年代仍未有定論。本文章利用10個基因以及235型態特徵，重建蛺蝶科的親緣關係，包含400個分布於蛺蝶科主要支系的屬。作者們的分析顯示蛺蝶科的起源約為9000萬年前的白堊紀，而10-12個支系在新熱帶區與東方區的祖先則逃過了白堊紀末期的大絕滅災難。其分化的格局顯示其他支系的滅絕年代在白堊紀/第三紀的邊緣，以及在第三紀時種化效率較為快速。

Abstract
The butterfly family Nymphalidae contains some of the most important non-drosophilid insect model systems for evolutionary and ecological studies, yet the evolutionary history of the group has remained shrouded in mystery. We have inferred a robust phylogenetic hypothesis based on sequences of 10 genes and 235 morphological characters for exemplars of 400 of the 540 valid nymphalid genera representing all major lineages of the family. By dating the branching events, we infer that Nymphalidae originated in the Cretaceous at 90 Ma, but that the ancestors of 10–12 lineages survived the end-Cretaceous catastrophe in the Neotropical and Oriental regions. Patterns of diversification suggest extinction of lineages at the Cretaceous/Tertiary boundary (65 Ma) and subsequent elevated speciation rates in the Tertiary.