| 题名 | 姬鼠属和白腹鼠属系统发育及谱系地理研究 |
| 作者 | 陈鹏 |
| 学位类别 | 博士 |
| 答辩日期 | 2012-05 |
| 授予单位 | 中国科学院研究生院 |
| 授予地点 | 北京 |
| 导师 | 蒋学龙 |
| 关键词 | 姬鼠属 白腹鼠属 东喜马拉雅-横断山地区 分子系统进化 系统地理学 |
| 其他题名 | Phylogeny and phylogeography of Genera Apodemus |
| 学位专业 | 动物学 |
| 中文摘要 | 姬鼠属主要分布在古北区，向南延伸分布到东洋区的边缘；而白腹鼠属为东洋区分布。该两属啮齿动物有多个物种在东喜马拉雅-横断山区重叠分布。本研究分别以两属的大多数现生种类为研究对象，展开属的分子系统进化研究和属内部分种群的谱系地理研究，以重建两个属的系统演化关系和探讨动物区系物种谱系地理格局形成的原因。同时利用形态学多变量分析方法结合分子系统发育对中华姬鼠复合种群（Apodemus draco complex）的分类地位进行厘定。 为厘清中华姬鼠复合种群的分类地位，我们采集了横断山地区和云南高原39个采集点中华姬鼠复合种群标本97号，对每一样本进行27项头骨特征测量并进行多变量分析；同时对这97号标本的组织样品进行线粒体cyt-b基因全序列测序及其中16号标本的组织样品进行核基因IRBP序列测序，结合姬鼠属11个物种的已有序列分别重建整个属的系统进化树。研究结果表明：头骨形态多变量分析、DNA遗传距离和分子系统发育关系表明澜沧江姬鼠（A. ilex）是一独立种，而长尾姬鼠（A. orestes）不是有效种，其长尾不是区分与澜沧江姬鼠或中华姬鼠的特征。 对分布于四川地区16个采集点93个中华姬鼠（A. draco）样品和分布于云南高原63个采集点的298个澜沧江姬鼠样品进行的cyt-b基因全序列进行分子系统地理分析，结果表明深切的河谷是造成中华姬鼠和澜沧江姬鼠种群东西隔离地理遗传结构的主要因素：中新世晚期，中华姬鼠与澜沧江姬鼠以金沙江为界产生分化，早上新世以来中华姬鼠以大渡河为界分化成东西两个大支，大渡河以东的支系在青藏运动以来相继分化出三个亚支系；大渡河以西的支系则以雅砻江为界继续分化成东西两个亚支系；澜沧江姬鼠在早上新世以来以澜沧江为界分化为东西两个大支系，西部种群以怒江为界分化成两个亚支系；东部种群相继分化为三个亚支系。此外，澜沧江姬鼠在金沙江两侧玉龙和哈巴雪山的群体没有发生明显的遗传分化，这可能与金沙江石鼓段更新世时期堰塞湖的形成以及哈巴雪山和玉龙雪山在第四纪前处于同一个夷平面有关。种群动态分析表明气候剧烈变化是推动种群扩张的动力，不同谱系同域分布是由于间冰期种群多次二次接触的结果。 我们一共扩增了约1740 bp的线粒体基因片段（cyt-b基因全序列和HVR基因序列）和约1952 bp的核基因片段（IRBP基因和FGB基因序列）以重构白腹鼠属系统进化关系。研究结果确定了马来白腹鼠（N. bukit）、N. huang种的分类地位；首次记录中国有南洋鼠（N. langbiani）、白腹鼠（N. niviventer）和马来白腹鼠分布；白腹鼠属内部仍然存在多个隐存种的可能；揭示了台湾社鼠（N. culturatus）和台湾白腹鼠（N. coninga）是姐妹群关系；白腹鼠（N. niviventer）与N. huang互为姐妹群关系。系统发育结合分子钟推算揭示白腹鼠属的演化格局：地质演化和气候的变迁是推动白腹鼠属物种扩散、分化的主要动力；分布于高海拔地区的白腹鼠种群在冰期向低海拔地区迁移，并因冰期-间冰期旋回体现出相应的反复扩散和退缩，相应的白腹鼠种群则以身体被扁平刺毛的多寡体现出对不同海拔的适应性进化。白腹鼠物种早上新世中期从喜马拉雅山南麓分别向东、向南辐射演化并最终广布于中国大陆、台湾岛、海南岛及东南亚地区，暗示冰期-间冰期旋回对白腹鼠属演化的驱动作用，同时揭示冰期陆连给白腹鼠种群向离岛迁移带来了可能。 基于系统进化的研究背景，我们使用白腹鼠属中的社鼠（N. confucianus）、刺毛鼠（N. fulvescens）和川西白腹鼠（N. excelsior）进行分子系统地理分析以探讨东喜马拉雅-横断山及邻近地区小型兽类的遗传地理格局： 社鼠的分子系统地理研究使用了来自于四川、云南和广西61个采集点的192个样品的线粒体cyt-b基因全序列及其中127个样品的HVR基因序列。研究表明社鼠的地理遗传格局与东喜马拉雅-横断山复杂的地形地貌以及更新世以来该地区多次的冰期和间冰期旋回有关，且气候变化对其的影响更明显。 刺毛鼠谱系地理分析使用了来自于东喜马拉雅-横断山、泰国、越南和海南等地区32个采集点的137个刺毛鼠样品的线粒体cyt-b基因全序列，以及其中84个样品的HVR序列。研究结果证实了刺毛鼠也是更新世起源的的观点；东喜马拉雅-横断山地区复杂的地质构造和气候变迁共同影响刺毛鼠的地理遗传格局，更新世多次冰期-间冰期旋回是推动其迁移扩散的主要诱因；刺毛鼠的适应性进化主要体现在身体被刺毛以及冰期中向低纬度地区的迁移上。 东喜马拉雅-横断山地区56个采集点的184个川西白腹鼠样品用于该物种的系统地理研究中，共获得145条cyt-b基因全序列和184条HVR基因序列。研究结果表明，早上新世到更新世的青藏高原及滇西北地区全面快速隆升和显著降温是导致川西白腹鼠金沙江两侧种群分化的主要动力，更新世以来的冰期-间冰期旋回是直接导致川西白腹鼠退居避难所和扩散的原因。 澜沧江姬鼠、中华姬鼠和白腹鼠属三个物种的系统地理研究结果表明，地质变迁和气候变化共同影响该地区的种群演化历史，但是高山纵谷对种群地理遗传格局的影响因不同物种所具有的相异的扩散能力和适应性而有所不同：金沙江两侧的中华姬鼠和澜沧江姬鼠种群地理遗传格局均呈现南北扩散、东西隔离的特点，显示与东喜马拉雅-横断山地区南北走向的山脉和河谷有着紧密联系，同时古气候的变化是驱动两物种各亚支系扩散和分化的重要因素；白腹鼠属三个物种的种群系统地理分析都显示出以金沙江及其沿线的干热河谷为界谱系分化、东西扩散的遗传地理结构，暗示对具有较大扩散能力和适应性的白腹鼠类群而言，相较地形，气候的变化对种群遗传地理格局的影响更大。另外，不同谱系同域分布表明该地区在冰期存在多个种群退缩的微避难所，而在间冰期它们存在广泛地二次接触。 |
| 英文摘要 | The genus Apodemus is mainly distributed in the Palaearctic Region and south edge of Oriental, while genus Niviventer is distributed in the Oriental Region. More than one species of the two rodents overlap in the eastern Himalayan and Hengduan Mountains. In this study, we reconstructed the phylogenies of the genus Apodemus and the genus Niviventer to infer their evolutionary history. We also analyzed the phylogeographic patterns of several species in these two genera. Meanwhile, the endemic Chinese species A. draco complex was evaluated in this study basing on the combination of morphometric characters and molecular data. We analyzed 97 specimens of the A. draco complex from 39 localities in the Hengduan Mountains and Yunnan plateau for distinguishing the A. draco complex. 27 morphometric variables of each sample were measured for multivariate analysis. All these 97 samples and 16 of them were examined to reconstruct phylogenetic trees using mitochondrial cyt-b and nuclear IRBP gene respectively. Another 11 species of Apodemus were included in these studies for Apodemus phylogenetic analyses. The results of multivariate analyses, Genetic distance of cyt-b and phylogenetic analyses confirmed that A. ilex is a valid species, but A. orestes is not. The tail length is not a valid diagnostic character for distinguishing A. draco and A. ilex. To assess the genetic diversity and biogeographic patterns of A. draco and A. ilex lineage, we sequenced the cyt-b gene from 93 samples of A. draco that were collected from 16 localities in Sichuan and 298 samples of A. ilex from 63 localities on the Yunnan plateau. The results indicated that the phylogenetic pattern of A. draco and A. ilex was strongly related to the existence of deep River valley systems in the Hengduan Mountains: The Jinsha River seemed to be the geographical barrier for divergence of A. draco and A. ilex in the Late Miocene. A. draco could be divided into eastern and western phylogroups by Dadu River, the eastern phylogroup was divided into three subclades following the strong tectonic uplift of the Qinghai-Tibet plateau and strong cooling climate. The western phylogroup was further divided into eastern and western subclades with the Yalong River as the geographical barrier. A. ilex could be divided into eastern and western clades with the Lancang River as a geographical barrier, the western clade was further divided into eastern and western sub-clades with the Nu River as the boundary. The eastern clade was further divided into three subclades. There was no significant difference between populaitons from Mt. Haba and Mt. Yulong that were separated by the Jinsha River, and this might be related to the damming of Shigu barrier lake in the Pleistocene when Mt. Haba and Mt. Yulong were connected. Population dynamic analyses indicated that lineage divergences and population expansions were strongly affected by climate fluctuation and multiple second contacts occurred to form current distributions. We employed phylogenetic analyses of ~1740 bp mitochondrial and ~1952 bp nuclear genes to reconstruct the phylogeny of Niviventer. The results confirmed the validity of N. bukit and N. huang. N. culturatus and N. coninga are sister species, the same as N. niviventer and N. huang. N. langbiani, N. niviventer and N. bukit were first recorded in China, and there might be existed some potential cryptic species. Phylogenetic analyses combined with molecular clock analyses revealed that the evolutionary history of Niviventer has been strongly affected by geological change and climate fluctuation: species in high elevations moved down during the glacial periods and recolonized during the interglacial periods. We infer that the evolutionary radiation of Niviventer derive from early Pliocene and resulted in the current geographical distribution. It implied that the glacial-interglacial periods stimulated evolution of Niviventer and Niviventer recolonized Taiwan, Hainan and other islands when the islands were connected by land bridges during glacial periods. We examined the phylogeographic patterns of three species of Niviventer in the eastern Himalayas and Hengduan Mountains: We first analysed the phylogeographic pattern of N. confucianus using 192 samples from 61 localities in Sichuan, Yunnan and Guangxi. 192 cyt-b sequences and 127 HVR sequences were employed for analyses. Our results revealed that the lineages of N. confucianus relate to mountain ridges, drainage systems and climate changes. Lineages were more strongly affected by climate fluctuation than topography. Sympatric populations clustered into different lineages and indicated a multi-refugia pattern. In N. fulvescens phylogeographic study, 137 cyt-b sequences and 84 HVR sequences were determined from 137 samples that were collected from the eastern Himalayas, Hengduan Mountains, Hainan, Thailand and Vietnam. Our results confirmed that N. confucianus originated from the Pleistocene. Lineage divergences and population expansions were strongly affected by climate fluctuation in the Pleistocene. Adaptive evolution of N. fulvescens was reflected by a move down to low altitudes and low latitudes. We employed phylogeographic analyses of N. excelsior using 184 samples from 56 localities in the eastern Himalayas and Hengduan Mountains. 145 cyt-b sequences and 184 HVR sequences were analysed. The results showed that the lineages were congruent with different phylogenetic structure: the phylogenetic pattern of N. excelsior was strongly related to a rapid uplift of the Tibetan plateau and “three river” region, especially the cooling climate after the uplift. In summary, the mountains and river valleys promoted the genetic diversity of species in the eastern Himalayas and Hengduan Mountains. The phylogeography of Apodemus and Niviventer was strongly related to the complex geographical topography and climate fluctuation. Animals with different dispersal abilities and different ecological adaptability were varying from their phylogeographic patterns: Apodemus were strongly related to the deep drain systems with dry-hot valley, while Niviventer lineages were more strongly affected by climate fluctuation than drainage systems. Sympatric populations clustered into different lineages indicating a multi-refugia pattern. |
| 语种 | 中文 |
| 公开日期 | 2012-09-25 |
| 内容类型 | 学位论文 |
| 源URL | [http://159.226.149.42:8088/handle/152453/7122]  |
| 专题 | 昆明动物研究所_兽类生态与进化 |
| 推荐引用方式 GB/T 7714 | 陈鹏. 姬鼠属和白腹鼠属系统发育及谱系地理研究[D]. 北京. 中国科学院研究生院. 2012. |
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