利用植物標本館檔案及老DNA片段追蹤歐洲入侵性植食者- 七葉樹潛蛾的入侵起源

資料來源: DC Lees, HW Lack, R Rougerie, A Hernandez-Lopez, T Raus, ND Avtzis, S Augustin & Lopez-Vaamonde. 2011. Tracking origins of invasive herbivores through herbaria and archival DNA: the case of the horse-chestnut leaf miner. Frontiers in Ecology and the Environment doi:10.1890/100098

簡介
七葉樹潛蛾（Cameraria ohridella）1986年首度在歐洲被描述之後，在歐洲成為一個擴散路徑和起源地具有爭議性的入侵物種，2009年8月在本部落格曾介紹過下列文章: 粒線體與微衛星體DNA標記顯示具高入侵性的七葉樹潛蛾起源自巴爾幹半島，該文章報導此種潛蛾的COI單體型‭ (‬haplotype‭)‬（不計入觀賞用樹木上者）多為特有型或是侷限分布，唯有單體型A是廣泛優勢的，不僅遍佈全歐洲而且在巴爾幹半島殘餘的七葉樹族群分布地點上也能找到此型。故「此種潛蛾為何是在最近40年才從巴爾幹半島擴張出去？又是如何擴散到其它地方？」仍然未清楚，而單體型A為何會在擴及歐洲各地的七葉樹潛蛾COI遺傳組成中占優勢則成為本研究想要探討的問題，作者提出兩個假說‭ ‬H0‭:‬‭ ‬單體型A在歷史空間上原本就是廣布，且在自然組群中占優勢，因此就有較高的機率隨著景觀植物的種植被散播出去，H1‭: ‬過去的歷史中單體型A和其他單體型一樣稀少而且局限分布在巴爾幹半島，直到晚近才變得具有入侵性，甚至入侵到殘存的天然七葉樹族群中。

2011年6月此研究團隊利用植物標本館內寄主植物的館藏獲得這些潛蛾的歷史分布資料，‭ ‬最早可追溯自1879年採集自希臘中部的天然族群之標本，‭ ‬可作為推測此蛾擴散年代之參考，也由於歷史標本的各野生採集地非常難以到達，作者認為現代道路系統的發展是促成潛蛾擴散的原因，因為快速的汽車交通能在無意中將落葉中的幼蟲和渡冬蛹運送到另外一的個地點。

從標本檔案的葉片內留存的幼蟲或蛹的乾燥樣本可用統計受感染趨勢，甚至從這些樣本中擴增出短片段的核DNA與粒線體DNA作為遺傳單體型分析以族群遺傳學方式判斷擴散起源，這些檔案序列不僅確認了C‭.‬‭ ‬ohridella的身分更把其在歐洲的歷史回溯至少100年，雖然作者的結果無法充分的排除H0但是較支持他的‭ ‬H１，即單型體Ａ是在1970年間道路系統發展後才擴散到其它地區的。本文強調植物標本館的重要性長期受到忽略，本研究的結果顯示研究入侵物種的擴散歷史和原因時，少數的植物標本館是唯一能追蹤過去空間歷史的僅存資訊，並能提供優質的資源得到接近真實的推測，提供入侵種管理策略之參考。

Abstract
Determining the native geographic range or origin of alien invasive species is crucial to developing invasive species management strategies. However, the necessary historical dimension is often lacking. The origin of the highly invasive horse-chestnut leaf-mining moth Cameraria ohridella has been controversial since the insect was first described in 1986 in Europe. Here, we reveal that herbarium collections across Europe indicate a Balkan origin for C ohridella. We successfully amplified nuclear DNA and mitochondrial DNA barcode fragments from larvae pressed within leaves of herbarium samples collected as early as 1879. These archival sequences confirm an identity of C ohridella and set back its history in Europe by more than a century. The herbarium samples uncovered previously unknown mitochondrial haplotypes and locally undocumented alleles, showing local outbreaks of C ohridella back to at least 1961 and dynamic frequency changes that may be associated with road development. This case history demonstrates that herbaria are greatly underutilized in studies of insect–plant interactions, herbivore biodiversity, and invasive species’ origins.

更多相關資訊
doi:10.1890/100098 – zum Artikel (Full text available via http://dx.doi.org)
www.bgbm.org/bgbm/pr/Archiv/pressimages/press_images.HTM#Miniermotte – for press photos
http://eolspecies.lifedesks.org/pages/8675 – about the horse-chestnut leaf miner

以標準化流程進行不同地理區的熱帶雨林蝶類多樣性比較

文獻來源: Y Basset, R Eastwood, L Sam, DJ Lohman, V Novotny, T Treuer, SE Miller, GD Weiblen, NE Pierce, S Bunyavejchewin, W Sakchoowong, P Kongnoo and MA Osorio-Arenas. 2011. Comparison of rainforest butterfly assemblages across three biogeographical regions using standardized protocols. The Journal of Research on the Lepidoptera 44: 17-2.

摘要

與大多數其他生物一樣，昆蟲的多樣性在熱帶區域較溫帶區域為高，然而熱帶各區域之間的標準化比較研究相當稀少。要解開影響群聚多樣性的生態，演化與生物地理因子需要標準化的作業流程和長期研究。作者在低地雨林鬱閉樹冠的底層以Pollard walk穿越線計數法比較巴拿馬的巴羅科羅拉多島(BCI)，泰國的拷秋(KHC)與新幾內亞的瓦那(WAN)共三個地點的蝴蝶豐富度和多樣性。作者在上述三地分別以231、231、120次穿越線調查，觀察到1792、 2797、 3331隻蝴蝶個體，各有128、 131、134個物種。分別對穿越線長度和調查時間長度進行數據校正後，結果顯示蝴蝶數量與物種豐富度分別在WAN和KHC最高。雖然在WAN的蝴蝶豐富度看來似乎並非由於人工方法所造成，此觀察的生物學上意義仍然是模糊的。WAN地區的植物多樣性與KHC相當，但是所擁有的蝴蝶霧種多樣性卻較KHC低。此現象強調了在植物多樣性之外的因素，例如生物地理歷史，對於解釋造成蝶類多樣性的成因的重要性。KHC特別多樣的蝴蝶相可能是由於其地理位置位在印度中國區和巽它大陸區的交會點。相反的，WAN則穩固位在澳洲地理區，且其較低的物種多樣性可能導因自島嶼生物地理過程。這三個地點的常見物種皆有幾個特色：食果和食蜜者平均出現，超過一半的常見種幼蟲時期取食附生植物或藤蔓類，且他們的翅長近似。由這些觀察推測， Pollard walk穿越線計數法在不同的熱帶雨林樣區指出了常見種的相似集合，並且是一個熱帶雨林蝴蝶組成長期監測的有用工具。

Abstract. Insects, like most other organisms, are more diverse in tropical than in temperate regions, but standardized comparisons of diversity among tropical regions are rare. Disentangling the effects of ecological, evolutionary, and biogeographic factors on community diversity requires standardized protocols and long-term studies. We compared the abundance and diversity of butterflies using standardised ‘Pollard walk’ transect counts in the understory of closed-canopy lowland rainforests in Panama (Barro Colorado Island, BCI), Thailand (Khao Chong, KHC) and Papua New Guinea (Wanang, WAN). We observed 1792, 1797 and 3331 butterflies representing 128, 131 and 134 species during 230, 231 and 120 transects at BCI, KHC and WAN, respectively. When corrected for length and duration of transects, butterfly abundance and species richness were highest at WAN and KHC, respectively. Although high butterfly abundance at WAN did not appear to result from methodological artefacts, the biological meaning of this observation remains obscure. The WAN site appeared as floristically diverse as KHC, but supported lower butterfly diversity. This emphasizes that factors other than plant diversity, such as biogeographic history, may be crucial for explaining butterfly diversity. The KHC butterfly fauna may be unusually species rich because the site is at a biogeographic crossroads between the Indochinese and Sundaland regions. In contrast, WAN is firmly within the Australian biogeographic region and relatively low species numbers may result from island biogeographic processes. The common species at each of the three sites shared several traits: fruit and nectar feeders were equally represented, more than half of common species fed on either epiphytes or lianas as larvae, and their range in wing sizes was similar. These observations suggest that Pollard walks in different tropical rainforests target similar assemblages of common species, and, hence, represent a useful tool for long-term monitoring of rainforest butterfly assemblages.

Keywords: Barro Colorado Island, Center for Tropical Forest Science, Lepidoptera, tropical rainforest, Panama, Papua New Guinea, Pollard walks, Smithsonian Institution Global Earth Observatories, Thailand.

Received: 22 December 2010
Accepted: 24 January 2011
Published online at www.lepidopteraresearchfoundation.org on 4 May 2011

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.

氣候變遷以及物種分布界線上的演化適應

文獻來源: Jane K. Hill, Hannah M. Griffiths and Chris D. Thomas, 2010. Climate Change and Evolutionary Adaptations at species' Range margins. Annu. Rev. Entomol. 56: 143-159 [abstract]

氣候變遷影響生物的分布其實不是個最近才有的新議題，在離現在最近的冰河範圍最廣的盛冰期(Last Glacial Maximum簡稱LGM)之前，早就經歷過數次氣候變遷以及冰河時期。現今的物種分布與最後盛冰期的生物避難所(Refugia又稱冰河種遺區)息息相關，甚至從許多物種的分布擴展可以看出從盛冰期到現今的氣候變遷。冰河時期結束後，物種從冰河時期的避難所播遷到新棲地，但隨著逐漸暖化的氣候，物種也不斷的往高緯度以及高海拔遷移，更有了適應環境的演化，但氣候暖化除了分布改變以及演化適應外，同時也造成瓶頸效應，使得物種的遺傳多樣性降低(註一)，Dr. Hill 等人針對這些現象，在2010回顧了近年來氣候的變遷以及生物分布，文章主要分成三個部分，分別從(1)氣候變遷對生物分布的影響、(2)生物對於新拓展的邊界環境的演化適應、以及(3)探討生物對於未來氣候暖化的適應的生物潛能，三個不同面相的探討，除了佐以許多的案例外，Dr. Hill等人也從生物遺傳多樣性的角度去探討氣候變遷所造成的影響。

以下是回顧的總結

1. 近年來氣候暖化造成許多物種往高海拔以及高緯度遷移，而且在往高緯度的分布界線擴展的物 種有演化適應的現象。
2. 在冰河時期後的許多廣布性物種，其遺傳多樣隨著分布緯度越高遺傳多樣性的降低，可以反映 出從2500前的盛冰期到現在的氣候暖化現象。
3. 遺傳多樣性的降低可能是由少數長距離播遷到新棲地的個體，因為等位基因的缺失以及異型核子的降低所造成。
4. 冰河時期後，瓶頸效應所遺留下的基因會加速或限制現今的分布是不得而知，但可以確定的是現今的棲地破碎化是會導致基因多樣性的降低(註二)。
5. 增加播遷能力是在分布邊界最常被觀察到的演化適應，(如:關於飛行能力的翅型型態改變以及代謝率增加)，當然也有演化適應是改變在與棲地的關聯性。
6. 在擴展新分布邊界時，常會在飛行能力與繁殖力間取捨，通常會藉由降低繁殖力來提高播遷力。
7. 了解生物適應環境的廣度以及速度，對於未來氣候改變反映在生物多樣性以及生態系統是很重要的關鍵。
   

註一、除了氣候之外，人類的開發造成棲地破碎化也會造成族群間的基因交流率降低，導致遺傳多樣性的降低。

註二、後冰河時期的瓶頸效應的影想就像雙面刃一樣，選汰的結果可能會使適應新環境的基因被穩定下來，幫助環境的適應，或者是相反，導致族群快速滅絕。

表型可塑性和族群分佈範圍的關係

Fig.2 Modified individuals obtained from the Fukaura area(http://www.biomedcentral.com/1471-2148/10/252/figure/F2)

文獻來源: Otaki JM, Hiyama A, Iwata M, Kudo T. 2010. Phenotypic plasticity in the range-margin population of the lycaenid butterfly Zizeeria maha. BMC Evolutionary Biology 10: 252. doi: 10.1186/1471-2148-10-252.

近期研究顯示在歐洲, 日本與美洲都有許多蝶類具有棲地範圍北移或擴大分布範圍的趨勢，而氣候暖化被認為是造成這個現象的因素之一，這些物種在適應新環境時表型甚至與其在原分布範圍有些差異。這篇文章的作者們以以灰蝶科的酢醬灰蝶(Zizeeria maha)之翅紋為例，探索在調控溫度的狀態下是否能夠證實物種表型的可塑性與溫度間的關聯。

Zizeeria maha主要分布在亞洲，成蟲展翅約25 mm，在腹面的前後翅上具有黑色斑點但不易區分性別，若以背面翅色可區分性別。幼蟲取食Oxalis corniculata葉片，不耐低溫，在低於12℃時不易進行蛻皮及化蛹，即使東部溫暖區域在11月底也不易觀察到野外族群，幼蟲會躲藏在下位葉、落葉堆或小石頭堆中化蛹，蛹期春季約14-20天，夏季約7天，成蟲期一年6~7代以夏季族群量最大。

物種分佈實驗的部份，作者在日本本洲北端青森縣(Aomori Prefecture)之深浦町(Fukaura: latitude 40°38.7’N and longitude 139°55.9’E)內選取9個樣區，在2000年至2004年間的夏季進行抽樣調查，再配合溫度變化去做翅紋分析，另外也將部份雌性個體帶回實驗室飼育，進行表型變化的研究。物種處理的部份是先將選育的表型基因轉入幼蟲體內並飼養10代後再將正常個體釋放，釋放族群和野生族群間可能產生基因上的交流，結果具選育基因的個體和一般個體相較之下具有較佳的耐寒性，因而產生族群的大量擴張，也使表型爆發的產生。

由此研究得知，表型和環境溫度及壓力有關，族群分佈範圍和遺傳組成有關，其次是族群擴大分佈的利益，這些不只顯示出表型的可塑性，也是未來演化研究的新方向。

由颱風引起的蛾類長距離傳播- 以東南亞大燕蛾為例

<此生態影像為泰國南部個體>

文章出處: Tokeshi, M. & M. Yoko-o. 2007. New record of the tropical swallowtail moth Lyssa zampa (Butler) (Lepidoptera: Uraniidae) from mainland Japan. Entomological Science 10(1): 103-106(4)

簡介

對於蛾類藉由風力的被動長距離傳播現象，過去雖然有被觀察過，然而超過1000公里以上的傳播並不多見且不容易被證實。本文對於日本西九州西所新採獲的大燕蛾(Lyssa zampa (Butler))個體，藉由氣象局所記錄的颱風路徑與形成資訊，提出單一個體由菲律賓受颱風傳播的可能性。此文中亦討論到台灣大燕蛾族群的來源，由於作者認為(1)台灣所採獲標本多不完整且多為偶發紀錄；(2)再加上此東南亞廣為分布的物種於該分布地區僅取食大戟科的Endospermum屬植物，而Yen et al. (1995)推測台灣大燕蛾族群可能的替代寄主是 Endospermum malaccense (黃桐)(但這個觀點顯然是Tokeshi & Yoko-o誤讀Yen et al.的文章所致, Yen et al.已經指出Endospermum不產於台灣, 因此其它大戟科植物或許可能被在地發生個體所利用)，然作者對此推測有所保留，而推測台灣族群可能亦由颱風因子而由菲律賓傳播而來，然此說法尚待台灣是否能採獲幼蟲與寄主紀錄的發現，以及族群遺傳學的資訊來佐證或反駁。同樣的，目前分布於台灣而被認為屬於菲律賓系統或南方海島起源的昆蟲類群，颱風與其他氣流的被動傳播方式的確是比須考量的因素之一。除此之外，對於地區性甚至全島的資源普查與物候發生資料蒐集，也是探討較大尺度生物地理議題時所不可或缺的資訊。(註: 燕蛾的長距離播遷事實上並非新聞, 中美洲的Urania是著名的遷徙性物種, 而印尼產的Lyssa可被海上作業漁船燈光誘引也曾被記錄過)

關於氣流傳播的文章尚有:

Owada, M. 1989. A migrant noctuid moth, Hypocala violacea Butler, obtained at Itoigawa on the Japan Sea coast, central Honshu. Japan Heterocerist's J. 152: 27.

Abstract

The tropical swallowtail moth Lyssa zampa (Butler) (Lepidoptera: Uraniidae) was recorded for the first time from mainland Japan. A male specimen of this species was captured in Amakusa, western Kyushu, on the night of 19 May 2006. An analysis of typhoon data suggests that this individual is likely to have reached the western coast of Kyushu from the Philippines, being transported over a distance of approximately 3000 km by typhoon no. 1 of 2006.

科西嘉島與薩丁尼亞島的眼蝶亞科具有萬花筒般複雜的種內生物地理學

文獻來源：L DAPPORTO. 2010. Satyrinae butterflies from Sardinia and Corsica show a kaleidoscopic intraspecific biogeography (Lepidoptera, Nymphlidae). Biological Journal of the Linnean Society. 100: 195–212.

科西嘉島與薩丁尼亞島等地中海島嶼因具有大量的特有種而受人矚目，蝶類更是備受關注。然而，科西嘉島與薩丁尼亞島的蝶類並未與鄰近大陸或西西里島所產的族群進行廣泛的研究。本研究選取11個分布於科西嘉島與薩丁尼亞島之眼蝶亞科物種，和其產於鄰近大陸及西西里島之族群，針對雄蟲生殖器結構進行幾何 形態矩陣的比較。利用相對扭曲計算、判別分析法、階層式分群法與交叉驗證測試，已鑑定包含島嶼和大陸族群的一致分布樣式。這十一個物種表現出多面向的分布樣式，但可歸納出以下三個重點：（１）來自北非和西班牙的族群一般而言和屬於義大利半島者不同；（2）來自薩丁尼亞島和西西里島的族群通常與北非/西班牙 的族群相似；（3）海洋屏障似乎是族群散佈的屏障，雖然其效力似乎與擴增無關。雖然科西嘉島與薩丁尼亞島之間的海峽和西西里島與義大利之間的海峽較短，但 在阻止動物相的交流上表現出強大的效力。而科西嘉島與北義大利之間、西西里島與突尼西亞之間的海域較廣，但卻各自表現出高度的相似性。這十一個分佈在島嶼 和大陸上的分類群之比較，有助於釐清關於地中海蝶類族群分布的歷史因素、避難所的動態和後冰河期（近代）遷徙等因子的複雜交互作用。

ABSTRACT
The Mediterranean islands of Sardinia and Corsica are known for their multitude of endemics. Butterflies inparticular have received much attention. However, no comprehensive studies aiming to compare populations of butterflies from Sardinia and Corsica with those from the neighbouring mainland and Sicily have been carried out. In the present study, the eleven Satyrinae species inhabiting Sardinia and Corsica islands were examined and compared with continental and Sicilian populations by means of geometric morphometrics of male genitalia. Relative warp computation, discriminant analyses, hierarchical clustering, and cross-validation tests were used to identify coherent distributional patterns including both islands and mainland populations. The eleven species showed multifaceted distributional patterns, although three main conclusions can be drawn: (1) populations from North Africa and Spain are generally different from those belonging to the Italian Peninsula; (2) populations from Sardinia and Sicily often resemble the North Africa/Spain ones; Corsica shows transitional populations similar to those from France; and (3) sea barriers represent filters to dispersal, although their efficacy appears to be unrelated to their extension. Indeed, the short sea straits between Sardinia and Corsica and between Sicily and the Italian Peninsula revealed a strong effectiveness with respect to preventing faunal exchanges; populations giving onto sea channels between Corsica and Northern Italy and between Sicily and Tunisia showed a higher similarity. A comparison of island and mainland distributions of the eleven taxa have helped to unravel the complex co-occurrence of historical factors, refugial dynamics, and recent (post-glacial) dispersal with respect to shaping the populations of Mediterranean island butterflies.

ADDITIONAL KEYWORDS: geometric morphometrics – glacial maxima – Mediterranean – refugia.

由於東亞特有的銀蛇目蝶切入Euptychiina亞族的親緣關係使得一個新的東亞-北美間斷分布案例被發現

圖片來源：ashung

文獻來源：Pen˜ a, C., Nylin, S., Freitas, A. V. L. & Wahlberg, N. (2010). Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae).—Zoologica Scripta, 39, 243–258.
doi:10.1111/j.1463-6409.2010.00421.x

蛺蝶科眼蝶亞科中具高多樣性的Euptychiina亞族過去被認為局限分布在美洲地區。然而，新的證據將東方區的銀蛇 目蝶（Palaeonympha opalina，又稱古眼蝶，分布於台灣和中國的中西部）歸入該亞族之下，使得該亞族的分布成為東亞-北美東部的間斷分布類型。這種東亞-北美東部分布類型雖然在其他動物群中已被發現，而在植物類群中也早在林奈時代之前就已知道，但是尚未在任何蝶類類群中被報導過。作者利用粒線體上的一個基因與四個細胞核 DNA片段，共計長度 4447 bp 的序列資料，分析102個 Euptychiina亞族下的分類群，以建立其親緣關係假說、評估其起源時間、主要分支之分化與生物地理學上的分析。Euptychiina亞族在 3100萬年前起源於南美洲。早期的Euptychiina亞族在始新世和漸新世之間、透過短暫的GAARlandia連接由北美洲擴散到南美洲。晚近之 東方區的銀蛇目蝶的間斷分布是由於南美洲一個支系朝北擴散、經由北美洲進入東亞。Palaeonympha屬的共祖與其姊妹群Megisto屬棲息在北亞和北美洲的帶狀森林中，過去曾以白令陸橋相連。而大約3100萬年前，白令陸橋的消失使得Palaeonympha屬和Megisto屬分開，且中新世晚 期（3150萬年前開始）的氣候巨變在北美洲西部造成嚴重的滅絕事件，故形成現在經典的「東亞-北美東部的間斷分布類型」。

本文章與台灣 產分類群相關者，是對銀蛇目蝶（Palaeonympha opalina）親緣關係地位之判斷，依據分子資料顯示，銀蛇目蝶應是Euptychiina亞族中某些局限分布在美洲之成員的姊妹群。此假說早在 1968年即由Miller提出，他發現Palaeonympha與Euptychiina亞族在形態上有某些相似性，這些發現促使他想要把這個東亞分布 的屬歸入原本僅有美洲產成員的Euptychiina亞族中，但是由於此奇特的間斷分布類型，最終Miller將銀蛇目蝶處理為分類地位未明的狀態。 Pen˜ a et al. (2006)的研究發現銀蛇目蝶和產於巴西東南方的Euptychiina亞族成員關係較密切，但是他並未全面地對產於北美洲和中美洲的 Euptychiina亞族成員進行採樣，故尚未發現經典的「東亞-北美東部的間斷分布類型」。本研究則進行較廣泛的採樣，進而重建並推論產於東方區的Palaeonympha的生物地理歷史，但本研究並未對每個屬中的每一個種作採樣，因此此親緣關係假說可能會受到較大屬的樣本增加而造成改變，此外，由 於只採樣銀蛇目蝶的其中一個產於台灣的亞種，因此無法確認銀蛇目蝶的每個亞種是否形成單系群。

Abstract
The diverse butterfly subtribe Euptychiina was thought to be restricted to the Americas. However, there is mounting evidence for the Oriental Palaeonympha opalina being part of Euptychiina and thus a disjunct distribution between it (in eastern Asia) and its sister taxon (in eastern North America). Such a disjunct distribution in both eastern Asia and eastern North America has never been reported for any butterfly taxon. We used 4447 bp of DNA sequences from one mitochondrial gene and four nuclear genes for 102 Euptychiina taxa to obtain a phylogenetic hypothesis of the subtribe, estimate dates of origin and diversification for major clades and perform a biogeographic analysis. Euptychiina originated 31 Ma in South America. Early Euptychiina dispersed from North to South America via the temporary connection known as GAARlandia during Eocene–Oligocene times. The current disjunct distribution of the Oriental Palaeonympha opalina is the result of a northbound dispersal of a lineage from South America into eastern Asia via North America. The common ancestor of Palaeonympha and its sister taxon Megisto inhabited the continuous forest belt across North Asia and North America, which was connected by Beringia. The closure of this connection caused the split between Palaeonympha and Megisto around 13 Ma and the severe extinctions in western North America because of the climatic changes of the Late Miocene (from 13.5 Ma onwards) resulted in the classic ‘eastern Asia and eastern North America’ disjunct distribution.

晚近擴張物種在第四紀冰河期的歷史與現代分布模式

文獻來源：C Kerdelhué et al.. 2009. Quaternary history and contemporary patterns in a currently expanding species. BMC Evolutionary Biology 9: 220 doi:10.1186/1471-2148-9-220

背景
第四紀的氣候振盪對物種的演化造成巨大的影響。在高緯度地區，族群必須在避難所區域度過嚴寒的冰河期，然後週期性地在間冰期拓殖回高緯度地帶。這種歷史通常會導致遺傳多樣性的損失；反之，未經歷過冰河時期的族群，可能會比其祖先具有較高的遺傳多樣性。這些特色強烈影響到現今的遺傳多樣性分佈狀況，甚至影響到這些物種如何因應當前的全球變遷。作者針對一種廣分佈的松舟蛾複合群(Thaumetopoea pityocampa/T. wilkinsoni complex, Notodontidae)進行粒線體遺傳多樣性的研究，該種是地中海盆地與南歐地區的森林性害蟲。此物種對近期氣候變遷的反應為快速的自然擴張其分佈區域，其中或許也包括偶然的人為遷移。本研究的目標是探討第四紀冰河期氣候振盪是否影響該物種的分佈範圍，以及判斷晚近拓殖地區的族群是否能鑑別岀近代遷移過程的遺傳足跡。

結果
作者鑑別岀三個具有地理結構的主要支系。在大部份的歐洲地區，其遺傳岐異度的樣式符合典型經過顯著冰河週期的物種。除了避難所以外，歐洲的各個族群的特色為只具有一種單倍基因型(haplotype)與基因多樣性極低，此特色被預期發生在氣候條件變好時受到物種快速地重新拓殖的地區。相反地，其他在地中海盆地周圍的次支系發生在受限的部分且有強烈的空間結構，如同受冰河影響有限的區域，在各個族群中保留其高度的遺傳多樣性，且大多數的單倍基因型是該地特有的。這種格局在地中海的遙遠離島（克里特島、塞普勒斯、科西嘉島）上格外顯著，亦具有特有的單倍基因型。近期引入的典型是會在地理分佈距離遙遠的族群間出現非常近似的單倍基因型，但是由於缺乏整體的遺傳結構、故在歐洲大部份的地區難以偵測。

結論
在冰川作用不明顯的地區，晚近的蛾類引入/擴張可由是否存在強烈的空間遺傳結構來偵測。反之，經歷第四紀冰河期強烈氣候振盪的區域，其天然族群則缺乏遺傳結構，而晚近的族群擴張樣式則尚無法偵測。

Abstract
Background Quaternary climatic oscillations had dramatic effects on species evolution. In northern latitudes, populations had to survive the coldest periods in refugial areas and recurrently colonized northern regions during interglacials. Such a history usually results in a loss of genetic diversity. Populations that did not experience glaciations, in contrast, probably maintained most of their ancestral genetic diversity. These characteristics dramatically affected the present-day distribution of genetic diversity and may influence the ability of species to cope with the current global changes. We conducted a range-wide study of mitochondrial genetic diversity in the pine processionary moth (Thaumetopoea pityocampa/T. wilkinsoni complex, Notodontidae), a forest pest occurring around the Mediterranean Basin and in southern Europe. This species is responding to the current climate change by rapid natural range expansion and can also be accidentally transported by humans. Our aim was to assess if Quaternary climatic oscillations had a different effect across the species' range and to determine if genetic footprints of contemporary processes can be identified in areas of recent introduction.
Results We identified three main clades that were spatially structured. In most of Europe, the genetic diversity pattern was typical for species that experienced marked glaciation cycles. Except in refugia, European populations were characterized by the occurrence of one main haplotype and by a strong reduction in genetic diversity, which is expected in regions that were rapidly re-colonized when climatic conditions improved. In contrast, all other sub-clades around the Mediterranean Basin occurred in limited parts of the range and were strongly structured in space, as is expected in regions in which the impact of glaciations was limited. In such places, genetic diversity was retained in most populations, and almost all haplotypes were endemic. This pattern was extreme on remote Mediterranean islands (Crete, Cyprus, Corsica) where highly differentiated, endemic haplotypes were found. Recent introductions were typified by the existence of closely-related haplotypes in geographically distant populations, which is difficult to detect in most of Europe because of a lack of overall genetic structure.
Conclusion In regions that were not prone to marked glaciations, recent moth introductions/expansions could be detected due to the existence of a strong spatial genetic structure. In contrast, in regions that experienced the most intense Quaternary climatic oscillations, the natural populations are not genetically structured, and contemporary patterns of population expansion remain undetected.

北美高山絹蝶在第四紀的氣候循環中的生物地理學研究

文獻來源：Schoville SD & Roderick GK. 2009. Alpine biogeography of Parnassian butterflies during Quaternary climate cycles in North America. Molecular Ecology. 18(1): 3471-3485. [Abstract]

更新世的冰河消長對於北美洲高山的地景，以及現居高山生態系的生物有深遠的影響。雖然物種豐度的模式偏離了島嶼生物地理學的預測，但在生物地理學上北美高山仍常被視為空中島嶼。原因之一即為受第四紀末期的地景變遷影響，造成了高山生物非平衡的群聚動態以及地理分布的變動，此可從遺傳資料中獲得證據。此研究利用粒線體DNA的COI片段以及核DNA的wingless與EF1a片段做分析、推測福布絹蝶（Parnassius phoebus）種複合群的演化歷史。作者對Parnassius smintheus與Parnassius behrii生物地理分布的變動歷史進行了多項測試，結果支持高山島嶼避難所模式，意即高山蝶類族群於冰河期縮減，並隨著間冰期的來到而擴張。雖然蝴蝶有能力於冰河期時在高山之間進行長距離的播遷，但事實上基因交流卻很少。此研究發現加州與科羅拉多州的高山棲地曾於間冰期形成連結，接著蝴蝶族群就於約兩萬年前快速擴張，進入上次冰河期後又縮減成小族群。研究譜系分枝的結果顯示P. smintheus與P. behrii的形態差異源自於更新世末期（約四萬八千年前），由於不順暢的基因交流造成分化所形成。此研究結果說明了空間的異質性與非平衡的族群動態影響了高山的生物多樣性。

Abstract
Growth of alpine glaciers during the Pleistocene had profound effects on montane landscapes in North America and the organisms now inhabiting alpine ecosystems. Biogeography of this region has often been viewed as a system of sky islands despite the fact that species richness patterns deviate from a strict island biogeographic model. One explanation is that alpine species are not in equilibrium because of late Quaternary geographic range shifts. Genetic data can provide evidence of nonequilibrium dynamics and the distributional shifts that occur during glaciation events in alpine landscapes. Using mitochondrial and nuclear sequence data, we examine the evolutionary history of butterflies in the Parnassius phoebus complex. We test explicit, alternative models of the biogeographic history of Parnassius smintheus and Parnassius behrii, including an equilibrium island model, ancestral radiation and fragmentation, an expanding alpine archipelago and an alpine archipelago refuge model. Our results support the alpine archipelago refuge model, in which alpine butterflies undergo population contraction during glacial climates followed by population expansion during interglacial phases. While butterflies can disperse between distant mountain ranges during glacial periods, gene flow is rare. We find evidence of recent connectivity between California and Colorado, population expansion events following deglaciation ∼20 000 years B.P., and small population sizes during the last glacial period. An analysis of lineage splitting suggests that morphological differences in P. smintheus and P. behrii are the result of late Pleistocene divergence (∼48 000 years B.P.) with limited gene flow. Our results demonstrate that spatially complex and nonequilibrium population dynamics influence alpine diversity patterns.

起源自安地斯山之透翅蛺蝶的物種多樣化格局

Photo by Pia's birdseye view

Out of the Andes: patterns of diversification in clearwing butterflies

Full text: PDF

Molecular Ecology (2009) 18, 1716–1729

M. ELIAS,*†‡ M. JORON,†§ K. WILLMOTT,¶ K. L . SILVA-BRANDÃO,**,†† V. KAISER,† C. F. ARIAS,‡‡ L. M. GOMEZ PIÑEREZ,§§ S . URIBE,§§ A. V. Z. BROWER,¶¶ A. V. L. FREITAS** and C . D. JIGGINS‡

安地斯山的熱帶森林擁有全球生物多樣性的高峰，其令人矚目的地理特色似乎可提供促進產生生物多樣性的機會，如地理割據與生態上的種化。然而，對於占生物多樣性最多比例的昆蟲，這些山脈所扮演的角色仍很少被使用親緣關係方法進行研究。作者在本文中探討安地斯山在新熱帶區高多樣性的昆蟲- 蛉蛺蝶亞科(the clearwing butterflies）演化上扮演的角色。作者使用定年的種級親緣關係去評估發生種化的年代以及推測兩個多樣化的屬的祖先分布海拔範圍。結果顯示兩個屬可能都起源自第三紀中新世中期的安地斯山中海拔地區，與對脊椎動物發表的研究成果大多起源自低海拔相反。儘管作者偵測到由上新世-中新世之間安地斯山脈隆起所造成的割據種化訊號，大多數的姊妹種都是鄰域分布且沒有明顯的地理屏障。結合整體上趨緩的種化速率，此結果推測生態上的種化與適應性輻射所扮演的角色比單純的地理割據更為重要。

Abstract
Global biodiversity peaks in the tropical forests of the Andes, a striking geological feature that has likely been instrumental in generating biodiversity by providing opportunities for both vicariant and ecological speciation. However, the role of these mountains in the diversification of insects, which dominate biodiversity, has been poorly explored using phylogenetic methods. Here we study the role of the Andes in the evolution of a diverse Neotropical insect group, the clearwing butterflies. We used dated species-level phylogenies to investigate the time course of speciation and to infer ancestral elevation ranges for two diverse genera. We show that both genera likely originated at middle elevations in the Andes in the Middle Miocene, contrasting with most published results in vertebrates that point to a lowland origin. Although we detected a signature of vicariance caused by the uplift of the Andes at the Miocene–Pliocene boundary, most sister species were parapatric without any obvious vicariant barrier. Combined with an overall decelerating speciation rate, these results suggest an important role for ecological speciation and adaptive radiation, rather than simple vicariance.

舊世界廣佈蝶種-波紋小灰蝶之親緣地理與遺傳多樣性

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Phylogeography and genetic diversity of a widespread Old World butterfly, Lampides boeticus (Lepidoptera: Lycaenidae) BMC Evolutionary Biology 2008, 8:301

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David J Lohman* 1, Djunijanti Peggie 2, Naomi E Pierce 3 and Rudolf Meier 1

Address: 1 Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore 117543, Republic of Singapore, 2 Division of Zoology, Research Centre for Biology-LIPI, Jl. Raya Jakarta-Bogor Km. 46, Cibinong-Bogor 16911, Indonesia and 3 Museum of Comparative Zoology, Harvard University, 26 Oxford St., Cambridge, Massachusetts 02138, USA

Abstract

Background: Evolutionary genetics provides a rich theoretical framework for empirical studies of phylogeography. Investigations of intraspecific genetic variation can uncover new putative species while allowing inference into the evolutionary origin and history of extant populations. With a distribution on four continents ranging throughout most of the Old World, Lampides boeticus (Lepidoptera: Lycaenidae) is one of the most widely distributed species of butterfly. It is placed in a monotypic genus with no commonly accepted subspecies. Here, we investigate the demographic history and taxonomic status of this widespread species, and screen for the presence or absence of the bacterial endosymbiont Wolbachia.
Results: We performed phylogenetic, population genetic, and phylogeographic analyses using 1799 bp of mitochondrial sequence data from 57 specimens collected throughout the species' range. Most of the samples (>90%) were nearly genetically identical, with uncorrected pairwise sequence differences of 0 – 0.5% across geographic distances > 9,000 km. However, five samples from central Thailand, Madagascar, northern Australia and the Moluccas formed two divergent clades differing from the majority of samples by uncorrected pairwise distances ranging from 1.79 – 2.21%. Phylogenetic analyses suggest that L. boeticus is almost certainly monophyletic, with all sampled genes coalescing well after the divergence from three closely related taxa included for outgroup comparisons. Analyses of molecular diversity indicate that most L.. boeticus individuals in extant populations are descended from one or two relatively recent population bottlenecks.
Conclusion: The combined analyses suggest a scenario in which the most recent common ancestor of L. boeticus and its sister taxon lived in the African region approximately 7 Mya; extant lineages of L. boeticus began spreading throughout the Old World at least 1.5 Mya. More recently, expansion after population bottlenecks approximately 1.4 Mya seem to have displaced most of the ancestral polymorphism throughout its range, though at least two early-branching lineages still persist. One of these lineages, in northern Australia and the Moluccas, may have experienced accelerated differentiation due to infection with the bacterial endosymbiont Wolbachia, which affects reproduction. Examination of a haplotype network suggests that Australia has been colonized by the species several times. While there is little evidence for the existence of morphologically cryptic species, these results suggest a complex history affected by repeated dispersal events.

[Article]Molecular systematics and biogeography of the genus Zizina

Zootaxa 1746: 15-38 (11 Apr. 2008) 25 plates; 86 references Accepted: 6 Mar. 2008
Molecular systematics and biogeography of the genus Zizina (Lepidoptera: Lycaenidae)
MASAYA YAGO (Japan), NORIO HIRAI (Japan), MARIKO KONDO (Japan), TETSUO TANIKAWA (Japan), MINORU ISHII (Japan), MIN WANG (P.R. China), MARK WILLIAMS (South Africa) & REI UESHIMA (Japan) Abstract & excerpt (PDF; 20KB) free | Full article (PDF; 770KB) open access