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標題: 台灣產鍬形蟲(鞘翅目:鍬形蟲科)的遺傳變異:高山特有亞種分化事件及紅圓翅鍬形蟲複合群之親緣地理
Genetic variation of stag beetles (Coleoptera, Lucanidae) in Taiwan: subspecific differentiation events of endemic montane stag beetles and phylogeography of Neolucanus swinhoei complex
作者: 蔡正隆
Cheng-Lung Tsai
關鍵字: 親緣地理學;更新世冰河期;山脈阻隔;避難所;鍬形蟲;紅圓翅鍬形蟲複合群;phylogeography;Pleistocene glaciations;mountain hindrance;refugia;stag beetles;Neolucanus swinhoei complex
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台灣生物相的演化史深受更新世冰河期氣候變動與中央山脈(Central Mountain Range, CMR)的地理隔離影響;在冰河事件與山脈阻隔雙重影響下,島上九種特有鍬形蟲的遺傳分化、種化及分類議題,備受注目。台灣產鍬形蟲之特有種與特有亞種比例高(約87%),部分爭議達數十年之久,其形態變異與局域適應相關;生物地理親緣研究涉及族群結構、遺傳分化、分子定年與遷徙路徑,可應用於解析島上鍬形蟲的演化歷程。本研究檢視台灣鍬形蟲的遺傳變異及闡明部分高山鍬形蟲的演化史,依據COI、16S rDNA及28S rDNA所建構的48種台灣鍬形蟲親緣關係顯示多數物種為單系群,兩個粒線體基因顯示種內遺傳距離多小於2%、種間變異大於2%,可有效鑑定多數物種。COI與16S rDNA之分析結果也確認黑腳深山鍬形蟲(Lucanus ogakii)、栗色深山鍬形蟲(L. kanoi)及金鬼鍬形蟲(Prismognathus davidis)的兩個亞種位階,與其地理分布相關,分化時間約發生於70-100萬年前。此外,黑腳深山鍬形蟲、栗色深山鍬形蟲與高砂深山鍬形蟲間,可能存在種間基因滲入/雜交事件。紅圓翅鍬形蟲複合群(Neolucanus swinhoei complex)由於形態鑑識特徵不明確、成蟲飛行能力弱及生活史約兩週,演化故事相形複雜、分類爭議近90年之久;本研究使用九個形態特徵與四個分子標誌(COI、16S rDNA、28S rDNA及wingless)來釐清其分類位階及遺傳分化;依據COI、16S rDNA及wingless之親緣關係顯示可分為一個東部與三個西部支系,後者分別為一個低海拔廣布支系與兩個山區支系;東、西部支系約於100萬年前分化,西部各支系約於50-75萬年前;考量這些鍬形蟲具共同的地理親緣史、形態鑑定特徵不明確及支系內不同族群間的DNA混雜狀況,這些圓翅鍬形蟲皆應視為紅圓翅鍬形蟲(Neolucanus swinhoei Bates, 1866)。為了進一步闡明此複合群的演化史,採集西北部山區各海拔及全台各地的族群;結果顯示中央山脈西北部(northwestern-CMR, NW-CMR)與廣布(Widespread)支系共存於許多西北山區海拔族群,廣布(Widespread)支系則遍布全島。定年時間顯示兩大支系起源於64萬年前,各支系分歧則在45-64萬年前。NW-CMR支系的四個分支有地理分布特性,分歧時間可追溯至25-48萬年前的里斯/民德冰期;同山區各高山族群遺傳組成相似且族群變動分析顯示有從低海拔往高海拔遷徙的趨勢,將有利於不同山頭族群間的遺傳交流。最後,了解廣布支系內的東部族群起源,將有助於釐清中央山脈的阻隔影響,COI與16S rDNA的親緣分析顯示廣佈支系具有南、北地域分化的趨勢且族群內變異大;同時,分子定年顯示主要分歧時間與里斯冰期有關,而隨後形成的分支則與最後一次冰河期相關;兩段基因顯示全台族群皆有的主基因單型,可能受祖先基因多型性及冰河期循環的族群擴增影響;儘管中央山脈在族群遷徙上扮演重要角色,但東部族群可藉由山脈南、北兩端低海拔山區遷徙過去。因此,山脈阻隔伴隨週期性冰河期是造成這些成蟲期短、遷移力弱、高山適應性鍬形蟲遺傳分化的主因。

Evolutionary history of Taiwanese biota has significantly affected by the climatic oscillation in Pleistocene and vicariance events of the Central Mountain Range (CMR). With the influence of glaciation events and the mountain hindrances, issues upon genetic differentiation, speciation, and taxonomic recognition of nine endemic stag beetles have been appealing in Taiwan. Several taxonomic debates have been carrying on for decades over Taiwanese stag beetles, which consist of a high proportion of endemic species and subspecies, i.e. about 87%, featuring morphological variations associated with local adaptation. Phylogeographic studies involving population structure, genetic differentiation, molecular dating, and dispersal routes of Taiwanese organisms could be applied to resolve the evolutionary processes of extant taxa. The study herein is to survey genetic variations of Taiwanese stag beetles and to unravel the evolutionary history of several montane stag beetles. Phylogenetic inferences based on COI+16S rDNA+28S rDNA of 48 Taiwanese lucanid species have confirmed most species are monophyletic groups; and the intraspecific (<2%) and interspecific (>2%) genetic distances of the two mitochondrial genes could be applied concordantly for taxonomic identification. COI and 16S rDNA have also confirmed the two subspecific status each for Lucanus ogakii, L. kanoi, and Prismognathus davidis which are congruent with their geographic distribution in this island and their subspecific split events might have occurred 0.7-1 million years ago (Mya). Moreover, genetic differentiation including introgression/hybridization events might have occurred among L. ogakii, L. kanoi, and L. maculifemoratus. About the more complicated evolutionary story in a 90-year controversy of Neolucanus swinhoei complex, a group of stag beetles with overlapping morphological characteristics, limited/weak flight capability, and two-week adult lifespan, nine morphological characteristics and four molecular amplicons, i.e. COI, 16S rDNA, 28S rDNA, and wingless, are exploited to address their taxonomic status and genetic differentiation. Phylogenetic inferences based on COI+16S rDNA+wingless showed one eastern and three western lineages, with the latter consisting of one low-hill Widespread and two montane lineages. The differentiation events of western and eastern lineages might have occurred ca. 1 Mya and the differentiation in western ones happened approximately 0.50-0.75 Mya. In consideration of these stag beetles have shared phylogeographical history, overlapping morphological characteristics, and intermingled DNA sequences from different populations within each lineage, all of these stag beetles should be regarded as Neolucanus swinhoei Bates, 1866. To further elucidate the evolutionary history of N. swonhoei complex, more individuals were collected from different altitudinal populations of the northwestern (NW) areas and across Taiwan Island. Results showed the coexistence of both NW-CMR and Widespread Lineage in many altitudinal populations, and Widespread Lineage existed all over Taiwan Island. Calibration dating showed that these two lineages might have originated prior to 0.64 Mya and differentiated during 0.45-0.64 Mya. In NW-CMR Lineage, four major clades are corresponding to their geographic distribution and their diversification events could be dated back to 0.25-0.48 Mya in Riss/Mindel glaciations. Similar genetic compositions are found among montane populations in the same muntain area and the demographic analysis showed a dispersal trend from hill to montane areas which would facilitate the genetic exchanges among different montane populations. Finally, to understand the origination of eastern populations of the Widespread Lineage would be helpful to elucidate the effect of the CMR hindrance. Phylogenetic inference, on the basis of COI and 16S rDNA, showed a tendency of north-south genetic differentiation and high variance components within populations. Meanwhile, molecular dating shown major diversifications were associated with Riss glaciations, while the subsequent forming lineages were in correlation to the Last Glacial Maximum. Both genes revealed that major haplotypes across all populations might be resulted by ancestral polymorphisms and the possible expansion events during glacial cycles. Although the CMR did play a vital effect on their dispersal, the eastern populations might have originated via lower mountain areas of northern and southern CMR. Thus, geological effects of mountain hindrance accompanied by periodical glaciations could have been vital impact leading to the genetic differentiation of these limited/weak flight capability, two-week adult lifespan, and montane-adaptive stag beetles.
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