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

Archaeoattacus屬天蠶蛾隱蔽種的證據

圖: 左上為Ar. edeardsii 右上為 Ar. malayanus 圖下為Ar. staudingeri圖片引用自原發表文章

文獻來源: Wolfgang A. Nässig, Stefan Naumann, and Rodolphe Rougerie. 2010. Evidence for the existence of three species in the genus Archaeoattacus (Lepidoptera: Saturniidae). The Journal of Research on the Lepidoptera 43: 37-47.[full pdf]

文章簡介
Archaeoattcus屬的模式種為Ar. edwardsii (Watson, 1895)，同屬的另外一種Ar. staudingeri亦由Watson在1914年時所發表，但當時此兩種都被放在Attacus中，直到五十年後Seitz等人在Attacus下建立了Archaeoattacus這個亞屬，並同時描述了一個亞種Ar. edwardsii malayanus，但是這個亞種由於沒有穩定且令人信服的特徵，再加上它在非馬來地區的亞洲大陸廣泛分布，最後被當成Ar. edwardsii的同物異名，但是Nässig等人於2010年提出了新的證據，包含了以生命條碼技術以及一些細微的雄性生殖器穩定差異，證實了確實有第三種的存在。所以原被當成同物異名的亞種名，被復活使用且提升為種Ar. malayanus。除此之外，分子證據還支持Ar. malayanus與Ar. staudingeri較為接近，而在生殖器的形態上，兩者幾乎沒有差別。

從藍小灰蝶探討，物種生命條碼的差異(Barcoding gap)真的存在嗎?

圖為Agrodiaetus merhaba

圖片引用自: butterflies-moths-turkey.com

文章出處

Martin Wiemers and Konrad Fielder, 2007. Does the DNA barcoding gap exist?- A case study in blue butterflies (Lepidoptera: Lycaenidae) Frontier in Zoology 4: 8 [pdf]

文章簡介

生命條碼是利用COI的一段序列作為物種快速鑑定的構想，最初是從鳥類以及一部分的節肢動物開始做起，在這兩個類群都顯示出物種間有明顯的基因的歧異度上可以作為鑑識用也就是所謂的生命條碼，後這個構想也幫助隱蔽種(cryptic species)的發現，更有人應用於演化生物學以及系統分類學上，但是有很多的分類學家很不贊同這樣，其中一個原因在於界定物種的生命條碼的差異(Barcoding gap)，可能是人為的因素所造成，起因於物種取樣(不論在種間或種內)不完全所導致。Wiemers & Fielder在2007利用了藍小灰蝶重新檢視生命條碼的差異是否存在。他們所選的Agrodiaetus屬(或亞屬)的藍小灰蝶不同種間有染色體數目上的差異，所以雜交無法受孕(或後代不孕)，也間接提供一個物種鑑定的依據，所以Agrodiaetus屬(或亞屬)的藍小灰蝶，是一個很適合檢測生命條碼的差異是否存在的材料。分析的部分，將基因序列使用K2P model (以便與之前文章比較)所計算出來的遺傳距離(genetic distance)來看種間和種內的差異，或是跑neighbor joing的樹，都無法完全的將種內或種間完全區隔開，換句話說也就是所謂的基因條碼的差異真的只是人為的結果。但是生命條碼對於隱蔽種或新種的發現還是有其重要性在，只是在使用生命條碼時需要更加的嚴謹，除此之外，生命條碼還有一個很基本也很重要的問題要解決，那就是物種的鑑定，Wiemers & Fielder (2007)年文中提到基因庫(Genebank)中幾個物種鑑定錯誤的序列，類似的問題說不定也存在於其他的資料庫中，如果連最基本的問題都沒解決，那生命條碼的可性度就令人質疑了。

數十年來氣候暖化造成熱帶高山蛾類不對稱的分布界限位移

圖片來源：Bettaman, Kinabalu Mountain Range and its Flora on 3500m

文獻來源：I-C Chen, JK Hill, H-J Shiu, JD Holloway, S Benedick, VK Chey, HS Barlow and CD Thomas. 2010. Asymmetric boundary shifts of tropical montane Lepidoptera over four decades of climate warming. Global Ecology and Biogeography. DOI: 10.1111/j.1466-8238.2010.00594.x
www.blackwellpublishing.com/geb

本文之研究群曾於PNAS (2009) 發表過本研究的初步探討分析，詳見這篇部落格介紹文章：氣候暖化造成熱帶型高山的蛾類群落往較高海拔移動 ，本文為對其資料進行較深入的比較與分析。

摘要：
目標 評估物種因應氣候變遷而移動其分布界限移動（寒冷的海拔上限與溫暖的分布下限）之現象，是否是以相等的速度發生。作者首先針對熱帶昆蟲提出證據，測試山區蛾類海拔分布上限與下界之移動，同時也考慮到熱帶山地物種之威脅。

地點 馬來西亞沙巴州 京那巴魯山

方法 作者於2007年調查京那巴魯山的尺蛾，與1965年所作的調查資料進行比較。物種的海拔分布上限和下界均有改變，海拔廣度和範圍地區均有評估。作者隨 機在幾年間進行亞取樣以確保有可進行比較之資料。資料分析的評估針對特有種和廣布種進行比較，以及在不同海拔高度達到其分布界限之物種。

結果 分布上限在2500–2700 m的物種（n = 28）皆撤退其上限與下界，且平均分布有收縮的趨勢(垂直分布約減少300 m, 分布範圍之面積約減少45 km2)。這些消退可能與雲量的變化有關，以及其他地理障蔽的影響(如地質與植被的轉變)可能阻擋其往山頂移動。除了這組物種外，大部份的物種（n = 109, 占本調查約80%）拓展了其分布的上限(平均約152 m)，且較其撤退的下界(77m)更多。

主要結論 在沒有阻礙的情形下，往山頂遷移的前進邊緣挪動較下限的退縮快速，因此許多物種因而擴增了其海拔分布之廣度。然而此結果並不表示分布範圍區域有所增加， 原因是可使用的的陸地面積會隨著海拔的上升縮減。接近山區主要生態上/地質上交會帶之物種受到兩面包抄而縮減其分布範圍區域。儘管目前尚未分布未及的棲地 是可以獲得的，這些物種可能早在將分布提升到山頂之前、就已面臨了物種滅絕的危機。

ABSTRACT
Aim To estimate whether species have shifted at equal rates at their leading edges (cool boundaries) and trailing edges (warm boundaries) in response to climate change. We provide the first such evidence for tropical insects, here examining elevation shifts for the upper and lower boundaries shifts of montane moths. Threats to species on tropical mountains are considered.
Location Mount Kinabalu, Sabah, Malaysia.
Methods We surveyed Lepidoptera (Geometridae) on Mount Kinabalu in 2007, 42 years after the previous surveys in 1965. Changes in species upper and lower boundaries, elevational extents and range areas were assessed. We randomly subsampled the data to ensure comparable datasets between years. Estimated shifts were compared for endemic versus more widespread species, and for species that reached their range limits at different elevations.
Results Species that reached their upper limits at 2500–2700 m (n = 28 species, 20% of those considered) retreated at both their lower and upper boundaries, and hence showed substantial average range contractions (-300 m in elevational extent and -45 km2 in estimated range area). These declines may be associated with changes in cloud cover and the presence of ecological barriers (geological and vegetation transitions) which impede uphill movement. Other than this group, most species (n = 109, 80% of the species considered) expanded their upper boundaries
upwards (by an average of 152 m) more than they retreated at their lower boundaries (77 m).
Main conclusions Without constraints, leading margins shifted uphill faster than trailing margins retreated, such that many species increased their elevational extents. However, this did not result in increases in range area because the area of land available declines with increasing elevation. Species close to a major ecological/geological transition zone on the mountain flank declined in their range areas. Extinction risk may increase long before species reach the summit, even when undisturbed habitats are available.

雖有穩定的植物資源，非洲雨林中鱗翅目昆蟲的組成仍具頗高的季節性差異

大戟科木本植物 Neoboutonia macrocalyx Pax 圖片來源：Prelude Medicinal Plants Database

文獻來源：Sini Skippari, Timo Olavi Veteli, John Kasenene, Pekka Niemela and Heikki Roininen. 2009. High temporal variation in the assemblage of Lepidopteran larvae on a constant resource. African Journal of Ecology 47: 537–545

本文作者在非洲烏干達的吉貝利(Kibale)國家公園的中海拔熱帶雨林中，研究不同樹木植株之間的鱗翅目幼蟲物種多樣性與物種組成之時間變化。調查方式為在1995年1月～1996年12月間，每個月對大戟科樹木 Neoboutonia macrocalyx Pax 葉片上的鱗翅目幼蟲進行採樣。在研究其間總計採得1961個樣本，共有76個形態種的鱗翅目昆蟲。各個物種的標本數量差異非常大，範圍少至一種只有1個樣本、多至一種有707個樣本。不同植株上採得的物種組成非常相似且以尺蛾科幼蟲為主。植食者豐度的時間性變化很高，其個體量的高峰在1995年主要發生於雨季，但此現象在1996年並未出現，且在這兩年當中、豐度均未和降雨量有顯著的相關性。相反地，在這兩年中的觀察資料顯示：鱗翅目多樣性與降雨量反而呈現負相關，且似乎因此造成一個半年為期的循環，使得每年會出現一至二個物種多樣性的高峰。此外，物種組成的相似程度亦具有季節性，而且同一物種會在一年之中的特定時段出現在該組成當中。此研究顯示：短期的生物多樣性評估會對熱帶雨林中生物多樣性做出偏頗的結論。

Abstract
We studied variation in the assemblage of lepidopteran larvae between individual trees, and temporal variation in the diversity and species composition of the assemblage in a medium-altitude rainforest in Kibale National Park, Uganda. Monthly samples of lepidopteran larvae were collected from the leaves of Neoboutonia macrocalyx Pax. between January 1995 and December 1996. During this period, a total of 1961 specimens representing 76 lepi- dopteran morphospecies were found. The numbers of individuals within species varied enormously, ranging from one to 707 individuals. Assemblages of individual trees were highly similar and dominated by geometrid larvae. Temporal variation in herbivore abundance was high. The number of individuals peaked during the major rainy season in 1995 but not in 1996 and was not cor- related significantly with rainfall within these 2 years. In contrast, a negative correlation was found between lepi- dopteran diversity and rainfall that seems to cause a semi- annual trend in diversity with one or two peaks per year. Furthermore, there was seasonality in the similarity of the assemblage. Consequently, the same species were found in the assemblage during certain times of the year. Our study shows that short-term biodiversity assessments can give a skewed picture of the diversity of tropical forests.

Key words: diversity, herbivory, insects, resource avail- ability, seasonality, species richness

何者決定熱帶非洲尺蛾科的多樣性：植物多樣性，植被結構還是非生物因子？

本圖片由Tambako the Jaguar上載至flickr

Determinants of diversity in afrotropical herbivorous insects (Lepidoptera: Geometridae): plant diversity, vegetation structure or abiotic factors?

Full Text: HTML, PDF (Size: 289K)

Journal of Biogeography (J. Biogeogr.) (2009) 36: 337–349
Jan C. Axmacher1*, Gunnar Brehm2, Andreas Hemp3, Henry Tu¨nte4, Herbert V. M. Lyaruu5, Klaus Mu¨ller-Hohenstein4 and Konrad Fiedler
1UCL Department of Geography, University College London, London, UK, 2Institut fu¨r Spezielle Zoologie mit Phyletischem Museum, Friedrich-Schiller-Universita¨t Jena, Jena, 3Department of Plant Systematics, Universita¨t Bayreuth, Bayreuth, Germany, 4Department of Biogeography, Universita¨t Bayreuth, Bayreuth, Germany, 5Department of Botany, University of Dar es Salaam, Dar es Salaam, Tanzania and 6Department of Population Ecology, Vienna Ecology Centre, University of Vienna, Vienna, Austria

此研究於非洲第一高峰吉力馬札羅山在海拔1200到3150 m的範圍設置48個樣點400平方公尺的樣區，在各樣區中央針對植食性且具高多樣性的昆蟲類群 - 尺蛾科，以燈光誘集方式進行物種數量與物種更替率調查，並比較其與樣區內維管束植物種類、植被結構(包含樹冠分層、覆蓋度、葉片外形、分枝形式等資料)之關係。結果顯示尺蛾物種的多樣性與樣區內常見植物的科內物種多樣性有關(R2 = 0.49)，且與植物結構指數有關(R2 = 0.22)，但與整體的植物多樣性無顯著相關。尺蛾的物種更替率與植物科級多樣性之改變範圍有明顯關聯(40% explained variance)，與植被外觀的改變關聯較低(25%)，與整體的植物多樣性關聯更弱(5%)。然而上述大部分的比較缺乏直接關聯，透過溫度和溼度等環境因子則會同時影響植被與蛾類相，尺蛾科物種多樣性和潛在寄主植物間可能的直接連結顯得更弱。整體植物物種豐富度與尺蛾科多樣性缺乏顯著關聯，暗示熱帶的尺蛾可能對寄主植物並不太有選擇性。因此，植物多樣性與植食者的物種多樣性間的清楚關聯被認為是過於簡化的想法，維管束植物多樣性亦不能被普遍地通用為昆蟲的多樣性指標。

ABSTRACT
Aim This study was conducted to investigate the potential of predicting alpha diversity and turnover rates of a highly diverse herbivorous insect family (Geometridae) based on vascular plant species richness and vegetation structure.

Location The study was carried out on the south-western slopes of Mount Kilimanjaro within a wide range of habitats between 1200 and 3150 m elevation.

Methods The floristic and structural composition of the vegetation was recorded at 48 plots of 400 m2. Geometrid moths were sampled manually at light sources located at the plot centres. Principal components analysis, redundancy analysis and multiple linear regression were used to explore how alpha diversity and species turnover of geometrid moths are related to vegetation structure and plant species richness.

Results Alpha diversity of geometrid moths was significantly correlated with species diversity patterns in the
most common vascular plant families (R2 = 0.49) and with plant structural parameters (R2 = 0.22), but not with overall floristic diversity. Species turnover of geometrid moths was strongly linked to diversity changes in a range of plant families (40% explained variance), less strongly to changes in vegetation physiognomy (25%), and only weakly to overall floristic diversity (5%). Changes in elevation were a better predictor of both alpha diversity and species turnover of geometrid moths than any principal component extracted from the vegetation data.

Main conclusions Vegetation composition, diversity and structure all showed significant correlations with the diversity and species composition of geometrid moth assemblages. Nevertheless, in most cases relationships were indirect, via environmental parameters such as temperature and humidity, which influenced both vegetation and moth fauna. Possible direct links between geometrid diversity and potential food plants were much weaker. The lack of a significant correlation between overall plant species richness and geometrid diversity indicates that tropical geometrid moths may not be very selective in their food plant choice. Accordingly, a clear correlation between floral diversity and herbivore species richness must be regarded as overly simplistic, and the diversity of vascular plants cannot universally be used as a suitable biodiversity indicator for diverse insect taxa at higher trophic levels.

氣候暖化造成熱帶型高山的蛾類群落往較高海拔移動

Elevation increases in moth assemblages over 42 years on a tropical mountain

PNAS (2009) 105: 1479-1483. Full Text: PDF (Size: 870K)

I-Ching Chen^a, Hau-Jie Shiu^b, Suzan Benedick^c, Jeremy D. Holloway^d, Vun Khen Chey^e, Henry S. Barlow^f, Jane K. Hill^a and Chris D. Thomas^a

a Department of Biology, University of York, P.O. Box 373, York YO10 5YW, United Kingdom; b Graduate Institute of Environmental Sciences and Ecology, National University of Tainan, 33 Su-Lin Street, Section 2, Tainan 700, Taiwan; c S chool of Sustainable Agriculture, Universiti Malaysia Sabah, Aras 2, Blok B, Bangunan Sekolah Sains and Teknologi, Locked Bag 2073, 88999 Kota Kinabalu, Sabah, Malaysia; d Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom; e Entomology Section, Forest Research Centre of Sabah, Sepilok, P.O. Box 1407, Postcode 90715, Sandakan, Malaysia; and f P.O. Box 10139, Postcode 50704, Kuala Lumpur, Malaysia
簡介

以長時間的尺度來討論生物與環境間的議題，在國外有不少的文獻可供參考，這類研究除了對於標的生物(target species)進行族群動態的監測之外，其統計數據亦可用於瞭解其組成結構的環境因子是否有所改變，而大部分研究最關切的議題即是全球暖化。以2007年聯合國跨政府氣後變遷小組 IPCC (Intergovernmental Panel on Climate Change) 所出版的統計數據，顯示從1970年至2004年共有28,586件生物波動觀察與佐證，其中絕多數為溫帶地區所作的研究，此外熱帶地區僅有7件以鳥類、爬蟲類、兩生類等脊椎動物所進行的調查，而其中並沒有在無脊椎或昆蟲相上的數據。

要了解長期的物種或地域性的群聚波動，即需要不同年份與季節的數據資料、不因時空背景而淘汰且重覆性高的研究方法，甚至是經費上的有力支援。在台灣以長期動態為研究目標的計畫早期大多為植被相監測，例如南仁山、福山植物園與楠溪內所設的永久樣區，以最早1989年南仁山所設的三個樣區來說，至今近20年才開始進行第一次複查，而動物相與昆蟲相大致僅有福山植物園內已完成或進行中的大鱗翅群調查計畫，可見台灣的長期生物相調查仍處於初始資料的累積。

2008年由赴英國約克大學(York University)攻讀學位的台灣籍研究生陳一菁、台南大學環境生態研究所許皓捷博士以及他國際研究員等人提出了婆羅洲京那巴魯山區(Mt. Kinabalu)蛾類群落往較高海拔移動的證據。此研究比較了1965年與2007年兩次在此山區採集的鱗翅目樣本，經統計得到102種尺蛾科分類群之海拔梯度分布在42年間上升了67公尺。關於這樣的結果，研究群認為除了可能是氣候變遷所造成的直接影響，也可能導因於尺蛾的寄主植物受環境因子波動的間接結果。

副熱帶地區的昆蟲相是否受到全球氣候變遷而有所影像? 副熱帶高山帶是否可作為氣候暖化時，溫帶型物種或是地域性特有種的庇護所? 這兩個議題在介於熱帶與副熱帶的台灣都可以籌議長期的生物相調查。

相關報導與參考:

NHM- Climate change pushes moths up mountain

Sceience News- Global impact of climate change on biodiversity

Moths of Borneo

Images of Mt. Kinabalu and its flora (Bettaman)

ABSTRACT

Physiological research suggests that tropical insects are particularly sensitive to temperature, but information on their responses toclimate change has been lacking—even though the majority of all terrestrial species are insects and their diversity is concentrated in the tropics. Here, we provide evidence that tropical insect specieshave already undertaken altitude increases, confirming the global reach of climate change impacts on biodiversity. In 2007, we repeated a historical altitudinal transect, originally carried out in 1965 on Mount Kinabalu in Borneo, sampling 6 moth assemblagesbetween 1,885 and 3,675 m elevation. We estimate that the average altitudes of individuals of 102 montane moth species, inthe family Geometridae, increased by a mean of 67 m over the 42 years. Our findings indicate that tropical species are likely to be as sensitive as temperate species to climate warming, and we urge ecologists to seek other historic tropical samples to carry out similar repeat surveys. These observed changes, in combination with the high diversity and thermal sensitivity of insects, suggest that large numbers of tropical insect species could be affected by climate warming. As the highest mountain in one of the most biodiverse regions of the world, Mount Kinabalu is a globally important refuge for terrestrial species that become restricted to high altitudes by climate warming.