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3种菊头蝠科蝙蝠分子系统地理学研究

Research on Molecular Phylogeography of Three Rhinolophidae Bats

【作者】 许立杰

【导师】 冯江;

【作者基本信息】 东北师范大学 , 环境科学, 2010, 博士

【摘要】 菊头蝠科蝙蝠隶属于翼手目小蝙蝠亚目,该科包括两个亚科:蹄蝠亚科和菊头蝠亚科。大蹄蝠和中华菊头蝠/托氏菊头蝠复杂体分别是两个亚科在中国的广布种,且他们的分布区范围基本相似,均分布于中国南部和西南部。本文以这3种菊头蝠科蝙蝠为研究对象,研究大蹄蝠的系统地理学和种群遗传结构以及中华菊头蝠/托氏菊头蝠复杂体的遗传多样性和系统地理学。通过分析mtDNA控制区615 bp的序列变异评价中国大蹄蝠遗传结构的地理模式。分子变异分析揭示出大蹄蝠的5个区域(西南区、东部区、西部区、滇南区和海南区)间具有较强的遗传结构。NJ树、由TCS构建的单倍型网络图和MDS散点图都表明在5个区域之间存在显著的地理分化。在大蹄蝠内检测到的高遗传结构可能是较弱的扩散能力、当地的适应性或是显著的雌蝠归家冲动的结果。由高黎贡山和琼州海峡分隔成的3个区域之间缺乏遗传结构可能是由于不完全的世系分类。我们估计了大蹄蝠种群的分歧时间,暗示出较近期的分化。应该给予具有最高遗传多样性的滇南群体和具有最低遗传多样性的海南群体最优先的保护。虽然已经有研究表明大蹄蝠携带与人类疾病直接相关的病毒,但我们基本上没有发现种群混合的证据。因此我们建议为了减少病毒传播的潜在风险,尽量减少对蝙蝠栖息洞穴的干扰。中华菊头蝠和托氏菊头蝠在形态上十分相似,关于这两个种的分类地位一直存在很多争议。生物地理学研究可以提供更多关于物种进化模式和机制的有价值的观点,同时也可以提供更多关于物种分类学的观点,对物种的保护十分重要。论文通过分析mtDNA控制区部分序列465 bp的序列变异探讨了中华菊头蝠和托氏菊头蝠复杂体的遗传多样性和种群统计学历史。NJ树、ML树和MP树表明来自于4个区域(西南区、东部区、西部区、滇南区)的复杂体发生了显著的地理分化,形成了与地理区域划分一致的4个亚进化枝。在所有种群中以及4个区域中均检测到高水平的遗传多样性,反映了这个复杂体具有一个长期稳定的进化历史。它们具有共同祖先的时间大约是254 700年前,暗示了这个复杂体是一个古老的物种。根据分歧时间和进化树中各亚进化枝间的亲缘关系,我们推测来自于滇南区的群体可能是复杂体内的隐藏种,且相对于中华菊头蝠和托氏菊头蝠而言是一个更古老的物种。大约是在200 000年前滇南区群体与东部区群体最早发生分化,处于中更新世的晚期。随后西部区和西南区组成的群体与东部区群体和滇南区群体相继发生分化。所有样本大约在晚更新世的早期135600年前发生种群扩张。我们的数据暗示“[滇南区] [西南区] [西部区] [东部区]”这4个区域应该被看作4个管理单元(MU),由于它们在遗传上的唯一性,应该同时予以保护。由于普遍的平行演化和趋同现象,通过形态数据不一定能反映出分类单元之间的系统发育关系。然而,将形态和分子数据结合可以洞察生物体的进化以及潜在的相关因素。这里我们集中研究了分类关系不明确的中华菊头蝠和托氏菊头蝠。传统的形态测定方法很难将它们分开,而我们以Cyt b基因为基础的研究表明它们分化成3个强支持的亚进化枝。我们进一步使用以标志点为基础的几何形态测量法分析了属于这个物种复杂体的80个样本的头骨差异。头骨的质心大小和形状差异将它们界定成3个形态类群,与以分子数据为基础提出的分类指定相一致,因此我们推测来自于滇南区的种群应为中华菊头蝠/托氏菊头蝠复杂体内的隐藏种。

【Abstract】 The family Rhinolophidae belongs to the suborder Microchiroptera, which includes two subfamilies, namely, Hipposiderinae and Rhinolophinae. Hipposideros armiger and Rhinolophus sinicus/Rhinolophus thomasi complex including respectively to the two subfamilies are widely distributed in China. Their distribution area is similar and located in the southern and southwestern China. In this study, we assessed phylogeography and genetic structure of H. armiger and R. sinicus/R. thomasi complex.The geographical patterns of the genetic structure of H. armiger in China were assessed by analyzing sequence variation in the mtDNA control region. Analysis of molecular variance revealed a very strong genetic structure among five regions in H. armiger. A NJ tree, haplotype network construction by TCS and MDS plots all showed significant geographic differentiation among five regions. The high genetic structure detected in H. armiger could be a consequence of poor dispersal ability, local adaptation, or marked female philopatry. The lack of genetic structure among three regions separated by the Gaoligong Range and the Qiongzhou Strait could be due to incomplete lineage sorting. Our estimated times of divergence for H. armiger populations suggested a relatively recent split. The S Yunnan population with the highest genetic diversity and the Hainan population with the lowest genetic diversity should be equally given priority for conservation. Although H. armiger has been shown to carry viruses implicated in human disease, we find little evidence for population mixing. We thus suggest minimizing disturbance to bats’roosting caves for minimizing the potential risk of virus transmission.Owing to their similarity in morphology between R. sinicus and R. thomasi, there are much controversy in regard to their taxonomic status. Biogeography can provide more insight into patterns and mechanisms of evolution, as well as into taxa, and be critical to conservation. We analyzed sequence variation in mtDNA control region of 465 bp to assess genetic diversity and demographic history of the complex. NJ, ML and MP trees indicated that significant geographical differentiation was found in the complex species from 4 regions (Southwest, East, West and S Yunnan). A high level of genetic diversity was observed within all regions and each region, reflecting a long evolutionary history of a large, stable population. The time of the most recent common ancestor for the complex species was estimated to be 254 700 years ago, suggesting that this complex is an old species. According to the divergence times and the phylogenetic relationship between the species, we speculated that the populations from the S Yunnan region might be cryptic species within the complex species. The earliest differentiation in the complex is estimated to be approximately 200 000 years ago between S Yunnan region and East region during the later Middle Pleistocene. Subsequently, differentiation has occurred between the West/Southwest group and East group and between the West/Southwest group and S Yunnan group. The estimated time since expansion for all the samples was about 135 600 years ago during the early Later Pleistocene. Our data indicated that those regions of“[S Yunnan] [Southwest] [West] [East]”should be regards as 4 MUs. We suggested that the protection should be implemented simultaneously due to their genetic uniqueness.Phylogenetic relationships between taxa are not necessarily reflected by morphological data due to widespread homoplasy and convergence. However, combining morphological and molecular data provides insights into the evolution of biological forms and into the potential factors involved. Here we focus on R. sinicus and R. thomasi with unclear taxonomic affinities. Traditional morphometric methods were difficult to separate them, whereas our Cyt b gene-based studies suggested that they were divided into three strongly supported subclades. We further used landmark-based geometric morphometrics methods to analyze the skull variability of 80 specimens belonging to this species complex. Patterns of size and shape delimitate three morphological groups that are congruent with the proposed taxonomic assignments based on molecular data, and therefore support the existence of cryptic species from the S Yunnan population.

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