KIZ OpenIR
鼠兔(Ochotona)的多基因系统发育,行为及其与人类的相互作用
其他题名Multilocus Phylogeny, Behaviors of Pika (Genus Ochotona) and its Interaction with Human
NARAYAN PRASAD KOJU
学位类型博士
导师蒋学龙
2017-05
学位授予单位中国科学院大学
学位授予地点北京
学位名称理学博士
学位专业动物学
关键词多基因系统发育 隐存种 线粒体基因渗入 传统医学 人为干扰 Multilocus Phylogeny Cryptic Species Mtdna Introgression Traditional Medicine Anthropogenic Interference
摘要

鼠兔鼠(Ochotona)隶属于兔形目鼠兔科。目前,现生鼠兔在中国西南地区的多样性仍未探明,其系统发育关系亦存在许多疑窦。本研究就是利用分子和形态学手段探讨鼠兔属的系统发育关系,线粒体基因渗透,形态特征及其与人类的相互作用等。本次研究采用了来自23个地点的96个组织样品,分属3个亚属(Pika, Ochotona and Conothoa)的11个种。我们测定了2个线粒体基因(CYT B , COI) 和5个核基因(RAG1, RAG2, TTN, OXAIL和IL1RAPL1),共计7,031bp。通过最大似然法(RAxML)和贝叶斯法(BEAST)构建的系统发育树,我们得到鼠兔属的进化历史,并利用物种界定分析(BPP)探究了其属内的物种多样性。结果显示,O. syrinx (O. huangensis) 已作为独立的分支进化。目前亚属间的系统发育关系仍未得到完全解决,究其原因,可能是该属在中新世中期(~13.90 Ma)的快速分化。另外,基因树中出现的不一致性说明O. cansus 和O. curzoniae中存在线粒体基因渗透现象,但目前仍无证据表明分布在秦岭地区的鼠兔间存在基因流。在形态学分析中,我们用电子游标卡尺测量了来自中国、美国、加拿大、尼泊尔、俄罗斯和伊朗的226个完整头骨(22个种及2个假定种)的22个颅骨指标。所有测量数据进行了对数变换,并在SPSS v17.0中进行主成份分析(PCA)。形态分析的结果较好的支持结果中关于假定种与假定亚属的推论。一些鼠兔物种间在皮肤颜色和头骨形态上有很高的相似性,这在野外鉴定中需要谨慎辨别。目前,对于鼠兔行为学的研究仍较匮乏。我们利用焦点扫描采样法记录了位于尼泊尔喜马拉雅山中心的朗塘国家公园(LNP;85°15’ E,28°20’ N - 28°32’ N)中Ochotona roylii的生态行为。研究的海拔范围在2800至5200米之间,但真正记录了行为的仅在3500米左右。研究时间是在2012年4月至2014年1月间,分别在夏季和冬季进行了观察。我们记录了鼠兔的梳理、凝望、进食、觅食、追逐与鸣叫的行为,并用曼惠特尼U检验和方差分析进行了数据分析。结果显示,在不同季节和生境下,鼠兔的行为分类存在着显著的差异。在Ochotona roylii成年个体中,无论夏季 (31.75%) 或冬季 (66.49%) 凝望均是最主要的活动行为。我们未记录冬季亚成体的任何行为。而在夏季中,亚成体花费大量时间在觅食(32.75%)上,其次为进食(22.26%),鸣叫与追逐是最少的行为。无论是成体还是亚成体,它们在不同的季节和生境下,进行觅食与进食的行为主要与食物的剩余量(50%)相关。人类与野生动物的相互作用离不开人类的进化。在尼泊尔,两者的关系与种族划分和文化差异息息相关。本次研究关注于人类与鼠兔的相互作用。研究于2012年至2015年间,在尼泊尔的两个保护区域内进行——朗塘国家公园(LNP)和ApiNampa保护区(ANCA)。我们进行了9次野外工作,通过问卷调查,民间故事和对当地人、朝圣者、冬虫夏草的采集人等的生活实践与宗教仪式等的观察进行了数据收集。鼠兔在朗塘被认为是佛教喇嘛的化身而得到保护,但在戈桑昆达湖,其生境由于朝圣者的宗教活动而严重退化。在ApiNampa保护区内的鼠兔则因其医药功效而被食用,鼠兔能够间接的通过进食存贮毒性植物而使家畜免受毒害。目前生物化学和生物药学的研究已经批准了将鼠兔进行药用和食用。最后,本研究从分子和形态上论证了鼠兔属仍存在三个来自陕西、四川和云南的三个隐存种,表明在中国西南地区的“空中之岛”山区的物种多样性是被低估的。另外,鼠兔的生态行为和与人类作用是在尼泊尔的喜马拉雅山进行的,我们希望多维的人类与鼠兔关系研究能用于喜马拉雅地区鼠兔的利用与保护。本研究亦建议今后能采用更丰富的物种数和样本,并结合基因组学和更详细的形态学分析来修正分类学与系统发育的关系。

其他摘要

Multilocus phylogeny, mtDNA introgression, morphometric analysis, behavior of genus Ochotona (pika) and its interaction with human culture and practices were the main focus of the study. The phylogeny of pikas (Ochotonidae, Ochotona) remains obscure, and pika species diversity in southwestern China has not been well explored. In this study, 96 tissue samples from 11 valid species in three classified subgenera (Pika, Ochotona and Conothoa) from 23 locations were characterized using multilocus sequences of 7031bp. Two mitochondrial (CYT B and COI) and five nuclear gene segments (RAG1, RAG2, TTN, OXAIL and IL1RAPL1) were sequenced. Evolutionary histories were analyzed using maximum likelihood (RAxML) and Bayesian analyses (BEAST), and also used molecular species delimitation analyses (BPP) to explore species diversity. Our study supported O. syrinx (O. huangensis) as a distinct clade from all named subgenera. Relationships among subgenera were not fully resolved, which may be due to a rapid diversification in the middle Miocene (~13.90 Ma). Conflicting gene trees implied mitochondrial introgression from O. cansus to O. curzoniae but there is no evidence of gene flow between pika species sympatrically inhabit in Qinling Mountains Craniodental analysis was carried out to support molecular result, total of 226 skulls representing 24 species of pikas with 22 valid and two putative species from China, USA, Nepal, Russia, Iran and Canada were thoroughly observed. Total 22 craniodental measurements were taken using a digital caliper graduated. All measurements were log-transformed and analyzed using principal component analysis (PCA) implemented in SPSS v17.0. The morphometric analysis support the molecular result unveiling putative species and putative subgenus. The behavior of Ochotona roylii was recorded in Langtang National Park (LNP), Nepal using focal scan sampling method. LNP is located in the central Himalayas of Nepal between 85°15’ E to 86°0’E and 28°20’ N to 28°32’ N. The research was conducted above 3500 masl. The field study for behavior was conducted from 2013 to 2015. Study period was divided into two seasons: summer and winter. Six behaviors: groom, muse, feed, forage, chase, and call behaviors were observed as behaviors of pikas. The statistical tools Mann Whitney U-test an ANOVA test were used for data analysis. The result revealed that there is significant difference in pikas behaviors in different seasons and habitats. The muse was the major activity observed in adult Ochotona roylii both in winter (66.49%) and in summer (31.75%). Meanwhile juvenile pika spent its maximum time budget in forage (32.75%) which is followed by feed (22.26%) in summer. There were no observation of juvenile in winter. Call and chase were the least observed behavior (>1%). Combining forage and feed the behavior related to food had more than 50% total budget both in juvenile and adult pika in each season and habitat.Interaction between human and wildlife continuing from evolution of human. The relation between them in country like Nepal with diverse ethnicity and culture is very specific and important. The study on human and pika (Ochotona) interaction was conducted in two protected area of Nepal: Langtang National Park (LNP) and ApiNampa Conservation Area (ANCA) from 2013 to 2015. Total nine field works (3 in ANCA and 6 in LNP) were carried to collect local people’s perception toward pika. The study was carried out by questionnaire, folk stories and direct observation of practices and rituals of local people, herders, pilgrim and Yarsagumba (Cordyceps sinesis) collectors. Pikas in LNP respect as symbol of Buddhist monk ‘Lama’. The habitat of pika is degrading in Gosainkunda, LNP due to anthropogenic pollution and in ANCA pikas are consumed by local people as medicine. Pikas were observed consuming and storing toxic plants, indirectly protecting livestock from killing accidently. Finally, this study uncovered three cryptic species from Shaanxi, Sichuan and Yunnan with strong support, suggesting an underestimation of species diversity in the “sky-island” mountains of southwest China. The behavior and human–pika interaction were explored in Himalaya of Nepal. We recommend further for further study pika and human interaction from multiple dimension for use and conservation of pikas in Himalaya. This study suggest for further work in genomics and detail morphometric study increasing species number and sample area for taxonomical and phylogenetic revision. 

学科领域生物学
学科门类动物学
语种英语
文献类型学位论文
条目标识符http://ir.kiz.ac.cn/handle/152453/12474
专题昆明动物研究所
遗传资源与进化国家重点实验室
科研部门_兽类生态与进化(蒋学龙)
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NARAYAN PRASAD KOJU. 鼠兔(Ochotona)的多基因系统发育,行为及其与人类的相互作用[D]. 北京. 中国科学院大学,2017.
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