书虱种群遗传多样性及线粒体基因组进化研究
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摘要
书虱(booklice)隶属于啮虫目(Psocoptera)、书虱科(Liposcelididae)、书虱属(Liposcelis),是世界范围内广泛发生并造成严重经济损失的一类储藏物害虫。书虱还能携带、传播病菌和引起过敏反应,直接威胁人们的身体健康。由于书虱具有取食范围广、活动能力强、繁殖率高和繁殖方式多样的特点；在防治上长期不合理地使用化学药剂,使书虱对多种储粮保护剂及熏蒸剂产生了较高的忍耐力或抗药性,其防控难度很大。常见的书虱种常混合发生,因而对书虱进行准确的物种鉴定是书虱有效防控及开展后续相关研究的前提。目前,国内外书虱的研究主要集中在基础生物学、与环境胁迫相关酶系的生理生化及分子生物学、以及应用防治等方面,而对书虱种群遗传与进化的研究非常少见。深入研究书虱种群遗传结构,有利于从分子水平上理解书虱的生态适应及成灾机制,为制定该类害虫的综合防控策略提供遗传学信息。此外,在啮总目中,书虱与寄生虱亲缘关系的界定一直存在争议,用于书虱分子系统发育关系研究的分子标记都局限于个别基因片段的联合或单个基因序列,说服力不强,而基于线粒体基因组序列的系统发育关系的研究,其结果更为准确、可靠。迄今为止,在300多种已完成线粒体基因组测序的昆虫中,仅有一种皮虱属于啮虫目,而关于其它啮虫目昆虫的线粒体基因组至今未见报道。因此,测定书虱线粒体基因组序列,不仅可以丰富昆虫线粒体基因组数量,推动昆虫比较基因组学和分子系统学研究的发展,而且还可阐明啮虫目和虱目之间的亲缘关系,有利于认识和理解两侧对称动物线粒体基因组进化的特征。
本学位论文紧密围绕书虱种群遗传与进化这一主线,开展了一系列研究。在建立书虱常见种快速鉴定的分子技术的基础之上,重点解析书虱的种群遗传结构,测序获得书虱线粒体基因组序列,分析书虱线粒体基因组的进化属性和特点,重建基于线粒体基因组序列的啮总目昆虫系统发生关系,并构建书虱微卫星富集文库。本论文的主要研究结果如下：
1、在全国范围内采集获得了嗜卷书虱Liposcelis bostrychophila Badonnel、嗜虫书虱L. entomophila (Enderlein)、小眼书虱L. paeta Pearman、三色书虱L. tricolor Badonnel、无色书虱L. decolor (Pearman)和云南书虱L. yunnaniensis Li&Li等6种书虱的20个地理种群。对6种书虱100头个体的核糖体转录内间隔区ITS(ITS1-5.8S-ITS2)基因进行克隆测序发现,6种书虱ITS1序列的长度变异范围在349-390bp之间,而ITS2序列的长度变异范围在198～418bp之间。基于ITS数据集计算的6种书虱种间水平的核苷酸歧异度(0.4167～0.7143)远远高于其他昆虫属内的种间水平,甚至达到某些昆虫科内的属级水平。鉴于ITS2基因的长度变异更加剧烈,本研究以ITS2作为书虱属内种间鉴定的DNA条形码,建立了一种基于多重PCR技术的快速鉴定这6种书虱的分子方法。利用ITS数据集,采用最大似然法(ML)和邻接法(NJ)构建的系统发育树显示,6种书虱可以很好地分为3支,这与传统形态分类学的结果相一致,说明ITS基因序列也适宜作为书虱分子系统发育关系研究的分子标记。
2、为揭示书虱种内遗传多样性水平以及基因流的范围和方式,利用线粒体Cytb基因部分序列和核糖体ITS全序列对嗜卷书虱和嗜虫书虱共15个地理种群的种群遗传结构进行了比较研究。
(1)基于Cytb数据集,共测序获得了103头嗜卷书虱和148头嗜虫书虱的Cytb基因片段,分别得到40个和39个单倍型。综合贝叶斯系统发育树和单倍型网络进化图的分析结果发现,嗜卷书虱的40个单倍型分为两支,它与两大动物地理区划西南区(SWCR)和华中区(CCR)的分布相吻合。对于39个嗜虫书虱单倍型,贝叶斯系统发育树与网络进化图均显示,来自同一种群或同一地理区域的单倍型并未聚在一起,即无种群或区域特异支。多层次的分子变异(AMOVA)分析表明,两种书虱种群在地理区划间存在显著的遗传结构。在进行书虱种群历史动态分析时,中性检验和误配分析参数表明,书虱在近期经历了种群扩张事件。两种书虱成对种群间的固定指数FST值多数大于0.33,说明两种书虱的种群遗传分化程度很高。然而基因流的计算结果显示,在这些种群之间存在着较高水平的基因流。这种看似矛盾的现象是由于书虱独特的生活习性所致,因为书虱种群中的个体趋于与近邻的个体相交配,这是导致其在同一分布区域的两端种群之间的基因流显著降低的重要原因。
(2)基于ITS数据集,共测序获得了74头嗜卷书虱和124头嗜虫书虱的ITS序列,分别检测到59个和77个单倍型。贝叶斯系统发育树和单倍型网络进化图的分析结果均显示两种书虱无种群或地理区域特异支。AMOVA分析的结果表明,两种书虱在地理区划间不存在显著的遗传结构,这一结果与利用Cytb基因的研究有所不同,其差异的原因很可能是由于线粒体Cytb基因和核糖体ITS基因序列的进化速率不同,以及取样分布和样本量大小不一致所造成的。两种书虱成对种群间的固定指数FST值显示,高水平的种群遗传分化在两种书虱种群间普遍存在。
(3)综合Cytb和ITS两种数据集的结果可以得出以下结论,两种不同生殖模式的书虱均具有很高的遗传多样性,且营专性孤雌生殖的嗜卷书虱遗传多样性高于两性生殖的嗜虫书虱遗传多样性,这一研究结果为解释书虱具有极强的环境适应能力提供了遗传信息上的证据。两种书虱种群间具有显著的遗传分化,但基因流相对较高,这就佐证了书虱具有易于随人类活动而传播和容易产生抗药性的特点。通过中性检验和误配分布检验的分析发现,书虱在近期经历了种群扩张事件,说明书虱即使在经历种群萎缩后,仍可以由极少数个体迅速建立起庞大的种群。Mantel检测表明,种群间的遗传距离与地理距离无相关性,即地理隔离并不是造成遗传分化的主要原因。因而,书虱种群间的遗传分化可能更多的是由随机遗传漂变、近交效应或防治策略所导致。
3、对书虱属代表种嗜卷书虱的线粒体基因组序列进行测定和分析。结果发现,嗜卷书虱的线粒体基因组从典型动物的一个环状DNA分子裂化成了两个较小的环状DNA分子,分别命名为mt chromosome Ⅰ (8,530bp)和mt chromosome Ⅱ (7,933bp), GenBank登录号分别为JN645275和JN645276。其中mt chromosome Ⅰ包含22个基因,mt chromosome Ⅱ包含16个基因。此外,两环同时拥有一段长度为945bp的序列,且此序列包含了3个tRNA基因,因而,两环状染色体只包括典型动物线粒体基因组的35个基因,丢失了两个tRNA基因(trnH和trnN)。在两个环状线粒体基因组上均出现了蛋白质假基因,它们的存在说明线粒体小环是典型线粒体基因组在基因删除作用下进化而来的。定量PCR结果显示,mt chromosome Ⅰ和mt chromosome Ⅱ具有不相等的拷贝数,且前者是后者的两倍左右。PCR检测发现,这种特殊的线粒体基因组裂化方式在嗜卷书虱不同地理种群中稳定存在。基于线粒体基因组序列重建的啮总目系统发育树的结果表明,啮虫目和虱目(Phthiraptera)中的细角亚目(Ischnocera)均为并系群,且书虱和整个虱目互为姊妹群。在阐述两侧对称动物线粒体基因组进化趋势时,本论文引入了线粒体核型(mitochondrial karyotypes)的概念。分析发现两侧对称动物线粒体染色体有减小的趋势,其中线粒体基因组的裂化是这种趋势体现的重要策略之一。裂化后的环状线粒体基因组具有以下共同特征：1)不同的小环之间拥有一段相似度很高的序列作为转录和复制的信息位点；2)每个小环必须具有至少一个完整功能的基因。只有具备这样的条件,小环才能逐步取代典型的线粒体染色体大环。这也暗示,动物线粒体主要功能基因的“丢失”,很可能是由于它位于具有上述特征的其他线粒体染色体小环上,只是未被检测到而已。嗜卷书虱线粒体基因组的裂化现象,有助于理解动物线粒体的起源和进化。从其他书虱种(如小眼书虱)的线粒体基因组测序结果看,线粒体基因组的裂化在书虱属中可能是一种普遍的现象。书虱线粒体基因组的裂化和虱目中吸虱类线粒体基因组的多裂化现象很可能是相互独立进化的遗传学事件。
4、为了以后更为全面地解析书虱的种群遗传结构,对嗜卷书虱和嗜虫书虱基因组中微卫星位点进行了筛选。利用链霉亲和素与生物素之间的强亲和性原理,将链霉亲和素偶联的磁珠与生物素标记的微卫星探针(AC)12、(TC)12、(ATC)8、(ATG)8、(AAC)8、(ATAC)6及(GATA)6结合后,再退火杂交含接头和微卫星序列的单链书虱基因组DNA限制性酶切片段,经PCR扩增形成双链后进行克隆、建库。本论文成功构建了6个嗜卷书虱微卫星富集文库和7个嗜虫书虱微卫星富集文库；两种书虱13个文库中共随机检测5,218个克隆,其中嗜卷书虱2,542个克隆；嗜虫书虱2,676个克隆,其阳性克隆率在9.38%至100%之间。经检测发现了两种书虱的260条微卫星DNA序列。本研究的微卫星富集文库构建过程是FIASCO法和Bloor的微卫星磁珠富集法的结合。该方法省略了酶切产物胶回收纯化步骤,提高了微卫星富集的效率。对微卫星序列进行分析发现,书虱两碱基微卫星主要以多拷贝微卫星位点存在,且其中有相当一部分是以基因家族的形式存在(不同位点的一侧侧翼序列极为相似),这与鳞翅目昆虫的微卫星家族极为类似。书虱三碱基重复微卫星在最终得到的单拷贝多态性位点中,其所占比例要远高于二碱基微卫星,并具有较高的优化率。这两种书虱微卫星富集文库的建立,将为嗜卷书虱和嗜虫书虱的种群遗传与进化、基因连锁图谱构建和分子系统发育等后续研究提供大量分子遗传标记。
综上所述,本论文对6种书虱种间和2种书虱种内两个层次水平的遗传进化规律进行了深入分析；克隆得到了6种书虱的ITS全序列,并由此建立了一套快速鉴定常见书虱种的分子方法,为书虱DNA条形码的开发和应用提供了借鉴,同时也为书虱种间进化关系的研究提供了分子标记；联合使用线粒体基因Cytb和核基因ITS对两种书虱种群遗传结构进行比较解析,系统阐明了书虱现实的种群遗传多样性、基因流的范围和方式、遗传分化程度以及种群历史动态,为解释书虱极强的抗胁迫能力和环境适应能力提供了基础的遗传学信息；获得了嗜卷书虱的全线粒体基因组序列,为啮总目系统发生关系和两侧对称动物线粒体基因组进化模式提供了全新的视角；构建了两种书虱的13个微卫星富集文库,为后续多态性微卫星位点的筛选和书虱种群遗传结构研究奠定了坚实的基础。本研究结果,充实了比较基因组学生物进化的研究内容,从线粒体基因组序列阐明了书虱物种多样性的形成进化机制和书虱与环境之间的相互作用机制,也为书虱抗药性治理方案的制定提供了丰富的遗传学信息,具有重要的理论意义和实践价值。
The psocids (booklice), Liposcelis spp.(Psocoptera:Liposcelididae), are major pests of stored grain and commonly occur on a wide range of stored products. Increasingly, the genus of Liposcelis has gained recognition of their importance due to their feeding on stored grains, contaminating food, and agricultural commodities as well as transmitting harmful microorganisms, including fungi and bacteria. Intensive use of chemical insecticides for pest control has facilitated resistance development in psocids. Control of these pests has proven difficult because they do not respond to management tactics that have been developed for other stored-product pests. The psocids have close morphological similarities and often commix occur at the same ecosystems. Therefore, a first step necessary to further implement population study of one Liposcelis species, species-discriminating criteria based on molecular techniques are needed. Previous research has focused on psocid bionomics, physiology and biochemistry, molecular biology, and control measures in grain storage systems. However, the population genetic structure and gene flow of psocids have not been well categorized, which may be useful to understand the formation of resistance mechanisms of psocids to insecticides and helpful to plan appropriate strategies for the control of these pests. Therefore, a detailed understanding of psocid gene flow patterns, determining the geographical origins and dispersal of source populations and the resultant genetic structure among populations within China is also needed. Liposcelididae species has a very special position in the phylogeny of Psocodea. Thus far, complete mt genome sequences have been determined for more than300species of insects. Only one of these species, a barklouse, Lepidopsocid sp., however, is from the order Psocoptera and no booklice has been sequenced for complete mitochondrial genomes. It is necessary to reconstruct the phylogenetic relationships among the major lineages of Psocodea using mitochondrial genome sequences, and the characteristic of mitochondrial genome of psocid will also give us invaluable insights into the evolution of mitochondrial genomes in bilateral animals.
In this thesis, we focused on the scientific issue on population genetic and evolution of major psocid species and the research include establishing the molecular technology for quickly identification of Liposcelis spp.; studying of population genetics of two psocid species; sequencing the whole mitochondrial genome of representative psocid species; analyzing the new evolutionary properties and patterns of mitochondrial genome, and constructing the microsatellite enriched libraries of L. bostrychophila and L. entomophila. The main results as follows:
1. Nucleotide sequences of the nuclear rDNA internal transcribed spacer (ITS) region were determined in100individuals of six Liposcelis species (L. bostrychophila Badonnel, L. entomophila (Enderlein), L. paeta Pearman, L. tricolor Badonnel, L. decolor (Pearman) and L. yunnaniensis Li&Li) from16locations (20populations) of China. For these six psocid species, the lengths of ITS1sequences ranged from349to390bp and the lengths of ITS2sequences from198to418bp. We also calculate the nucleotide divergence for these six psocid species using the ITS sequence alignments according to p-distances and the result showed that the inter-specific nucleotide divergence (0.4167to0.7143) of these six Liposcelis species was much higher than other insects at the same level and it almost reach the inter-genus nucleotide divergence of other insects. The molecular phylogenetic tree of six Liposcelis species was constructed inferred from the ITS-5.8S-ITS2sequencetypes and was generated from Maximum Likelihood (ML) and Neighbor-joining (NJ) methods. From both the ML and NJ phylogenetic trees, these six species were well divided into three clades which are consistent to the accepted morphological classification. We also developed a molecular identification method for six Liposcelis species based on ITS2sequence. The six reverse primers was designed to bind to one of these six species only and worked with the same forward primer. The identification method involves a single PCR reaction using DNA from a psocid and the special reverse primer was only able to generate a PCR product from its complementary species. Above results demonstrate that ITS1-5.8S-ITS2sequences are useful tools for phylogeny estimation of Liposcelis species and the ITS2sequence can be an excellent tool for species discrimination to facilitate entomological studies of the six species and developed multiplex PCR method here is proved to be reliable when tested through different geographical populations. This study showed that ITS (ITS1and ITS2) sequences displayed considerable differentiation both in length and base composition within psocid species and which suggest that great genetic differentiation existed within these species.
2. The two psocids, L. bostrychophila and L. entomophila were sampled in15localities in China and analyzed for polymorphism at mitochondrial DNA (Cytb) and ITS (ITS1-5.8S-ITS2) regions to study genetic diversity in psocid populations at intra-specific levels.
(1) In the Cytb dataset, A433bp fragment of the Cytb genes were aligned and analyzed from103L. bostrychophila and148L. entomophila individuals. Totally,39haplotypes were detected in L. entomophila populations and40haplotypes in L. bostrychophila populations. When the Bayesian tree and median-joining network for haplotypes were combined, the40haplotypes of L. bostiychophila were divided into two clades that coincided with our predefined regions (SWCR and CCR region). However, no population-specific clustering pattern was revealed by the Bayesian and median-joining network analysis for the39haplotypes of L. entomophila. AMOVA results showed that there was significant genetic differentiation of L. bostiychophila and L. entomophila populations at various hierarchical levels (among regions, among populations within regions, and within populations), and comprehensive analysis of demographic history changes for L. bostrychophila and L. entomophila populations using two neutrality tests and mismatch distributions indicate a rapid demographic expansion for both species. For L. bostrychophila and L. entomophila, most of the pairwise FST values were statistically significant, which indicate that there was widespread population genetic differentiation within and between two species populations. However, the high levels of gene flow were also detected among these populations and this paradoxical phenomenon could be explained using the specific life habit of psocids.
(2) In the ITS dataset,74individuals of L. bostrychophila were sequenced and59haplotypes were detected whereas124individuals of L. entomophila were sequenced and77haplotypes were detected. The phylogenetic analyses and median-joining networks did not show clear pattern of structure for both species according to the same sampling localities, or predefined geographic region. In contrast to the AMOVAs result inferred from the Cytb dataset, several AMOVAs at different hierarchical levels inferred from ITS gene showed that there were no genetic structures of L. bostrychophila and L. entomophila populations according to our predefined regions, however pairwise estimates of FST among L. bostrychophila and L. entomophila populations revealed that the genetic differentiation existed among a large portion of populations. The neutrality tests and mismatch distribution test statistics for both psocid species populations indicated a rapid historically demographic expansion in the past.
(3) From above analysis, both Cytb and ITS sequences show high genetic diversity in these two species, and significant genetic structure and population differentiation were also detected. The high genetic diversity of L. bostrychophila and L. entomophila might explain why these species has a broad tolerance to environmental and habitat stresses and the fast mutational processes inherent in individuals, as well as populations, can enable these two Liposcelis species to successfully adapt to complex environments. Compared to sexually reproducing L. entomophila, asexually reproducing L. bostrychophila has a higher genetic diversity. The Mantel test indicated that for both species there was no evidence for isolation-by-distance (IBD). Therefore, the significant population differentiation detected in both psocids may be mainly due to other factors, such as genetic drift, inbreeding or control practices, and less by geographic distance. The neutrality test and mismatch distribution statistics revealed that the two species might have undergone population expansions in the past in the sampled areas.
3. In this study, we sequenced the mitochondrial (mt) genome of L. bostrychophila and found that the typical single mt chromosome of bilateral animals has fragmented into and been replaced by two medium-sized chromosomes; each of these chromosomes has about half of the genes of the typical mt chromosome of bilateral animals. These mt chromosomes are8,530bp (mt chromosome I) and7,933bp (mt chromosome II) in size. For mt chromosome I, six protein-coding genes and five tRNA genes were on one strand whereas one protein-coding gene, one rRNA gene and nine tRNA genes were on the other strand. For mt chromosome II, five protein-coding genes and four tRNA genes were on one strand whereas one protein-coding gene, one rRNA gene and five tRNA genes were on the other strand. There is an identical region (945bp) on both of the two mt chromosomes, and three tRNA genes and an extensive non-coding region that contains putative D-loop and control sequences were included in this region. Intriguingly, qPCR test indicated that mt chromosome I is twice as abundant as chromosome II. PCR test of multipartite mt genome are successful in other strains of L. bostrychophila, indicating that the2-chromosome structure is stable but with product size slightly change, which possibly due to deletion of some of non-coding regions. The mitochondrial gene order of L. bostrychophila differs radically from Lepidopsocidae sp.(Psocoptera) and the parasitic lice which are closest relatives of Liposcelididae. Totally, four putative pseudogenes on the two mitochondrial chromosomes demonstrated the formation of mini-circles was proceeding step by step with degeneration of the functional protein genes, and they are also powerful proofed that the recombination were happened between these two chromosomes. It appears that the selection pressure for compact mt genomes in bilateral animals favors small mt chromosomes when small mt chromosomes co-exist with the typical large mt chromosomes. Thus, small mt chromosomes may have selective advantages over large mt chromosomes in bilateral animals. Phylogenetic analyses of mt genome sequences of Psocodea (i.e. Psocoptera plus Phthiraptera) indicate that:1) the order Psocoptera (booklice and barklice) is paraphyletic; and2) the order Phthiraptera (the parasitic lice) is monophyletic. Within parasitic lice, however, the suborder Ischnocera is paraphyletic; this differs from the traditional view that each suborder of parasitic lice is monophyletic. From this study, we summarized that the mini-circle mt chromosomes which can replaced the typical animal single circle mt must have at least one functional gene and one non-coding region which has a high similarity across different mini-circles. We also speculated that the mt chromosomes in bilateral animals have the tendency of reducing their size and the fragmentation of mt genome is one of important strategies of processing of mt genome streaming. According to result of sequencing the mt genome of other Liposcelis species, we intend to conclude that multipartite mt genomes are common in Liposcelis species and the multipartite mt genome of Liposcelis species also give us invaluable insights into the origin and evolution of mitochondria in bilateral animals.
4. In the present paper, microsatellite-enriched libraries of L. bostrychophila and L. entomophila were constructed utilizing methodologies that exploit the strong affinity between biotin and the protein streptavidin. We propose a fast and easy protocol which is combination of two different published methods. Briefly, high-quality genomic DNA was digested by the restriction enzyme Msel and then ligated to designed adaptors.250-700bp microsatellite-containing DNA fragments were captured by streptavidin-coated magnetic beads. The beads affinity capture of microsatellite repeats using biotinylated oligonucleotide probes (AC)12,(TC)12,(ATC)8,(ATG)8,(AAC)8,(ATAC)6, and (GATA)6Subsequently, PCR was used to amplify the captured molecules for transferring single strand DNA to double strand DNA. The PCR products (enriched microsatellite DNA fragments) were ligated to pGEM-T Easy vector and transformed into Trans5a competent cells. In total,13microsatellite enriched libraries were constructed; six for L. bostrychophila, and seven for L. entomophila. A total of5,218clones were PCR screened for microsatellite content. The clones of L. bostiychophila and L. entomophila were2,542and2,676, respectively. The percentage of the microsatellite positive clones ranged9.38%-100%. The number of microsatellites detected for these two species was260. Comparative analysis of microsatellite sequences for these two psocids revealed that the microsatellite sequences exist in multiple copies in the psocid genome. In addition, they were found to have similar or almost identical flanking regions. From our present study, the tri-nucleotide libraries yielded greater results for determining gene flow.
In summary, in the present study, the genetic evolution patterns of psocids at both the inter-and intra-specific levels were analyzed systematically. Internal transcribed spacers (ITS) regions were cloned from six psocid species. Based on great variation of the ITS2sequences, a molecular identification method for six Liposcelis species was developed. The study here will provide potential molecular markers to investigate the evolutionary relationship among the species of genus Liposcelis and also will provide the DNA barcoding tools in psocids species. Population genetics of two asexually and sexually reproducing psocids, L. bostrychophila and L. entomophila were comparatively analyzed using the mitochondrial Cytb and nuclear ITS gene sequences and a thorough understanding of these two psocids population genetics, including gene flow patterns, genetic diversities and the resultant genetic structure among populations within China is useful for proposing successful integrated pest management tactics for these pests in grain storage systems. The completed mt genome of L. bostrychophila was sequenced and this multipartite mt genome provided new insights into the phylogeny of psocodea and the evolution of mitochondrial chromosomes in bilateral animals. We constructed13microsatellite enriched libraries for two psocids. Accumulation of multiple polymorphic microsatellite loci will greatly facilitate studies of individual identification and they are crucial to investigations of population genetics and evolution, gene linkage mapping, and elucidating the molecular phylogeny of psocids in China and internationally. Therefore, the studies included in this thesis have important theoretical and practical significance.
本学位论文紧密围绕书虱种群遗传与进化这一主线,开展了一系列研究。在建立书虱常见种快速鉴定的分子技术的基础之上,重点解析书虱的种群遗传结构,测序获得书虱线粒体基因组序列,分析书虱线粒体基因组的进化属性和特点,重建基于线粒体基因组序列的啮总目昆虫系统发生关系,并构建书虱微卫星富集文库。本论文的主要研究结果如下：
1、在全国范围内采集获得了嗜卷书虱Liposcelis bostrychophila Badonnel、嗜虫书虱L. entomophila (Enderlein)、小眼书虱L. paeta Pearman、三色书虱L. tricolor Badonnel、无色书虱L. decolor (Pearman)和云南书虱L. yunnaniensis Li&Li等6种书虱的20个地理种群。对6种书虱100头个体的核糖体转录内间隔区ITS(ITS1-5.8S-ITS2)基因进行克隆测序发现,6种书虱ITS1序列的长度变异范围在349-390bp之间,而ITS2序列的长度变异范围在198～418bp之间。基于ITS数据集计算的6种书虱种间水平的核苷酸歧异度(0.4167～0.7143)远远高于其他昆虫属内的种间水平,甚至达到某些昆虫科内的属级水平。鉴于ITS2基因的长度变异更加剧烈,本研究以ITS2作为书虱属内种间鉴定的DNA条形码,建立了一种基于多重PCR技术的快速鉴定这6种书虱的分子方法。利用ITS数据集,采用最大似然法(ML)和邻接法(NJ)构建的系统发育树显示,6种书虱可以很好地分为3支,这与传统形态分类学的结果相一致,说明ITS基因序列也适宜作为书虱分子系统发育关系研究的分子标记。
2、为揭示书虱种内遗传多样性水平以及基因流的范围和方式,利用线粒体Cytb基因部分序列和核糖体ITS全序列对嗜卷书虱和嗜虫书虱共15个地理种群的种群遗传结构进行了比较研究。
(1)基于Cytb数据集,共测序获得了103头嗜卷书虱和148头嗜虫书虱的Cytb基因片段,分别得到40个和39个单倍型。综合贝叶斯系统发育树和单倍型网络进化图的分析结果发现,嗜卷书虱的40个单倍型分为两支,它与两大动物地理区划西南区(SWCR)和华中区(CCR)的分布相吻合。对于39个嗜虫书虱单倍型,贝叶斯系统发育树与网络进化图均显示,来自同一种群或同一地理区域的单倍型并未聚在一起,即无种群或区域特异支。多层次的分子变异(AMOVA)分析表明,两种书虱种群在地理区划间存在显著的遗传结构。在进行书虱种群历史动态分析时,中性检验和误配分析参数表明,书虱在近期经历了种群扩张事件。两种书虱成对种群间的固定指数FST值多数大于0.33,说明两种书虱的种群遗传分化程度很高。然而基因流的计算结果显示,在这些种群之间存在着较高水平的基因流。这种看似矛盾的现象是由于书虱独特的生活习性所致,因为书虱种群中的个体趋于与近邻的个体相交配,这是导致其在同一分布区域的两端种群之间的基因流显著降低的重要原因。
(2)基于ITS数据集,共测序获得了74头嗜卷书虱和124头嗜虫书虱的ITS序列,分别检测到59个和77个单倍型。贝叶斯系统发育树和单倍型网络进化图的分析结果均显示两种书虱无种群或地理区域特异支。AMOVA分析的结果表明,两种书虱在地理区划间不存在显著的遗传结构,这一结果与利用Cytb基因的研究有所不同,其差异的原因很可能是由于线粒体Cytb基因和核糖体ITS基因序列的进化速率不同,以及取样分布和样本量大小不一致所造成的。两种书虱成对种群间的固定指数FST值显示,高水平的种群遗传分化在两种书虱种群间普遍存在。
(3)综合Cytb和ITS两种数据集的结果可以得出以下结论,两种不同生殖模式的书虱均具有很高的遗传多样性,且营专性孤雌生殖的嗜卷书虱遗传多样性高于两性生殖的嗜虫书虱遗传多样性,这一研究结果为解释书虱具有极强的环境适应能力提供了遗传信息上的证据。两种书虱种群间具有显著的遗传分化,但基因流相对较高,这就佐证了书虱具有易于随人类活动而传播和容易产生抗药性的特点。通过中性检验和误配分布检验的分析发现,书虱在近期经历了种群扩张事件,说明书虱即使在经历种群萎缩后,仍可以由极少数个体迅速建立起庞大的种群。Mantel检测表明,种群间的遗传距离与地理距离无相关性,即地理隔离并不是造成遗传分化的主要原因。因而,书虱种群间的遗传分化可能更多的是由随机遗传漂变、近交效应或防治策略所导致。
3、对书虱属代表种嗜卷书虱的线粒体基因组序列进行测定和分析。结果发现,嗜卷书虱的线粒体基因组从典型动物的一个环状DNA分子裂化成了两个较小的环状DNA分子,分别命名为mt chromosome Ⅰ (8,530bp)和mt chromosome Ⅱ (7,933bp), GenBank登录号分别为JN645275和JN645276。其中mt chromosome Ⅰ包含22个基因,mt chromosome Ⅱ包含16个基因。此外,两环同时拥有一段长度为945bp的序列,且此序列包含了3个tRNA基因,因而,两环状染色体只包括典型动物线粒体基因组的35个基因,丢失了两个tRNA基因(trnH和trnN)。在两个环状线粒体基因组上均出现了蛋白质假基因,它们的存在说明线粒体小环是典型线粒体基因组在基因删除作用下进化而来的。定量PCR结果显示,mt chromosome Ⅰ和mt chromosome Ⅱ具有不相等的拷贝数,且前者是后者的两倍左右。PCR检测发现,这种特殊的线粒体基因组裂化方式在嗜卷书虱不同地理种群中稳定存在。基于线粒体基因组序列重建的啮总目系统发育树的结果表明,啮虫目和虱目(Phthiraptera)中的细角亚目(Ischnocera)均为并系群,且书虱和整个虱目互为姊妹群。在阐述两侧对称动物线粒体基因组进化趋势时,本论文引入了线粒体核型(mitochondrial karyotypes)的概念。分析发现两侧对称动物线粒体染色体有减小的趋势,其中线粒体基因组的裂化是这种趋势体现的重要策略之一。裂化后的环状线粒体基因组具有以下共同特征：1)不同的小环之间拥有一段相似度很高的序列作为转录和复制的信息位点；2)每个小环必须具有至少一个完整功能的基因。只有具备这样的条件,小环才能逐步取代典型的线粒体染色体大环。这也暗示,动物线粒体主要功能基因的“丢失”,很可能是由于它位于具有上述特征的其他线粒体染色体小环上,只是未被检测到而已。嗜卷书虱线粒体基因组的裂化现象,有助于理解动物线粒体的起源和进化。从其他书虱种(如小眼书虱)的线粒体基因组测序结果看,线粒体基因组的裂化在书虱属中可能是一种普遍的现象。书虱线粒体基因组的裂化和虱目中吸虱类线粒体基因组的多裂化现象很可能是相互独立进化的遗传学事件。
4、为了以后更为全面地解析书虱的种群遗传结构,对嗜卷书虱和嗜虫书虱基因组中微卫星位点进行了筛选。利用链霉亲和素与生物素之间的强亲和性原理,将链霉亲和素偶联的磁珠与生物素标记的微卫星探针(AC)12、(TC)12、(ATC)8、(ATG)8、(AAC)8、(ATAC)6及(GATA)6结合后,再退火杂交含接头和微卫星序列的单链书虱基因组DNA限制性酶切片段,经PCR扩增形成双链后进行克隆、建库。本论文成功构建了6个嗜卷书虱微卫星富集文库和7个嗜虫书虱微卫星富集文库；两种书虱13个文库中共随机检测5,218个克隆,其中嗜卷书虱2,542个克隆；嗜虫书虱2,676个克隆,其阳性克隆率在9.38%至100%之间。经检测发现了两种书虱的260条微卫星DNA序列。本研究的微卫星富集文库构建过程是FIASCO法和Bloor的微卫星磁珠富集法的结合。该方法省略了酶切产物胶回收纯化步骤,提高了微卫星富集的效率。对微卫星序列进行分析发现,书虱两碱基微卫星主要以多拷贝微卫星位点存在,且其中有相当一部分是以基因家族的形式存在(不同位点的一侧侧翼序列极为相似),这与鳞翅目昆虫的微卫星家族极为类似。书虱三碱基重复微卫星在最终得到的单拷贝多态性位点中,其所占比例要远高于二碱基微卫星,并具有较高的优化率。这两种书虱微卫星富集文库的建立,将为嗜卷书虱和嗜虫书虱的种群遗传与进化、基因连锁图谱构建和分子系统发育等后续研究提供大量分子遗传标记。
综上所述,本论文对6种书虱种间和2种书虱种内两个层次水平的遗传进化规律进行了深入分析；克隆得到了6种书虱的ITS全序列,并由此建立了一套快速鉴定常见书虱种的分子方法,为书虱DNA条形码的开发和应用提供了借鉴,同时也为书虱种间进化关系的研究提供了分子标记；联合使用线粒体基因Cytb和核基因ITS对两种书虱种群遗传结构进行比较解析,系统阐明了书虱现实的种群遗传多样性、基因流的范围和方式、遗传分化程度以及种群历史动态,为解释书虱极强的抗胁迫能力和环境适应能力提供了基础的遗传学信息；获得了嗜卷书虱的全线粒体基因组序列,为啮总目系统发生关系和两侧对称动物线粒体基因组进化模式提供了全新的视角；构建了两种书虱的13个微卫星富集文库,为后续多态性微卫星位点的筛选和书虱种群遗传结构研究奠定了坚实的基础。本研究结果,充实了比较基因组学生物进化的研究内容,从线粒体基因组序列阐明了书虱物种多样性的形成进化机制和书虱与环境之间的相互作用机制,也为书虱抗药性治理方案的制定提供了丰富的遗传学信息,具有重要的理论意义和实践价值。
The psocids (booklice), Liposcelis spp.(Psocoptera:Liposcelididae), are major pests of stored grain and commonly occur on a wide range of stored products. Increasingly, the genus of Liposcelis has gained recognition of their importance due to their feeding on stored grains, contaminating food, and agricultural commodities as well as transmitting harmful microorganisms, including fungi and bacteria. Intensive use of chemical insecticides for pest control has facilitated resistance development in psocids. Control of these pests has proven difficult because they do not respond to management tactics that have been developed for other stored-product pests. The psocids have close morphological similarities and often commix occur at the same ecosystems. Therefore, a first step necessary to further implement population study of one Liposcelis species, species-discriminating criteria based on molecular techniques are needed. Previous research has focused on psocid bionomics, physiology and biochemistry, molecular biology, and control measures in grain storage systems. However, the population genetic structure and gene flow of psocids have not been well categorized, which may be useful to understand the formation of resistance mechanisms of psocids to insecticides and helpful to plan appropriate strategies for the control of these pests. Therefore, a detailed understanding of psocid gene flow patterns, determining the geographical origins and dispersal of source populations and the resultant genetic structure among populations within China is also needed. Liposcelididae species has a very special position in the phylogeny of Psocodea. Thus far, complete mt genome sequences have been determined for more than300species of insects. Only one of these species, a barklouse, Lepidopsocid sp., however, is from the order Psocoptera and no booklice has been sequenced for complete mitochondrial genomes. It is necessary to reconstruct the phylogenetic relationships among the major lineages of Psocodea using mitochondrial genome sequences, and the characteristic of mitochondrial genome of psocid will also give us invaluable insights into the evolution of mitochondrial genomes in bilateral animals.
In this thesis, we focused on the scientific issue on population genetic and evolution of major psocid species and the research include establishing the molecular technology for quickly identification of Liposcelis spp.; studying of population genetics of two psocid species; sequencing the whole mitochondrial genome of representative psocid species; analyzing the new evolutionary properties and patterns of mitochondrial genome, and constructing the microsatellite enriched libraries of L. bostrychophila and L. entomophila. The main results as follows:
1. Nucleotide sequences of the nuclear rDNA internal transcribed spacer (ITS) region were determined in100individuals of six Liposcelis species (L. bostrychophila Badonnel, L. entomophila (Enderlein), L. paeta Pearman, L. tricolor Badonnel, L. decolor (Pearman) and L. yunnaniensis Li&Li) from16locations (20populations) of China. For these six psocid species, the lengths of ITS1sequences ranged from349to390bp and the lengths of ITS2sequences from198to418bp. We also calculate the nucleotide divergence for these six psocid species using the ITS sequence alignments according to p-distances and the result showed that the inter-specific nucleotide divergence (0.4167to0.7143) of these six Liposcelis species was much higher than other insects at the same level and it almost reach the inter-genus nucleotide divergence of other insects. The molecular phylogenetic tree of six Liposcelis species was constructed inferred from the ITS-5.8S-ITS2sequencetypes and was generated from Maximum Likelihood (ML) and Neighbor-joining (NJ) methods. From both the ML and NJ phylogenetic trees, these six species were well divided into three clades which are consistent to the accepted morphological classification. We also developed a molecular identification method for six Liposcelis species based on ITS2sequence. The six reverse primers was designed to bind to one of these six species only and worked with the same forward primer. The identification method involves a single PCR reaction using DNA from a psocid and the special reverse primer was only able to generate a PCR product from its complementary species. Above results demonstrate that ITS1-5.8S-ITS2sequences are useful tools for phylogeny estimation of Liposcelis species and the ITS2sequence can be an excellent tool for species discrimination to facilitate entomological studies of the six species and developed multiplex PCR method here is proved to be reliable when tested through different geographical populations. This study showed that ITS (ITS1and ITS2) sequences displayed considerable differentiation both in length and base composition within psocid species and which suggest that great genetic differentiation existed within these species.
2. The two psocids, L. bostrychophila and L. entomophila were sampled in15localities in China and analyzed for polymorphism at mitochondrial DNA (Cytb) and ITS (ITS1-5.8S-ITS2) regions to study genetic diversity in psocid populations at intra-specific levels.
(1) In the Cytb dataset, A433bp fragment of the Cytb genes were aligned and analyzed from103L. bostrychophila and148L. entomophila individuals. Totally,39haplotypes were detected in L. entomophila populations and40haplotypes in L. bostrychophila populations. When the Bayesian tree and median-joining network for haplotypes were combined, the40haplotypes of L. bostiychophila were divided into two clades that coincided with our predefined regions (SWCR and CCR region). However, no population-specific clustering pattern was revealed by the Bayesian and median-joining network analysis for the39haplotypes of L. entomophila. AMOVA results showed that there was significant genetic differentiation of L. bostiychophila and L. entomophila populations at various hierarchical levels (among regions, among populations within regions, and within populations), and comprehensive analysis of demographic history changes for L. bostrychophila and L. entomophila populations using two neutrality tests and mismatch distributions indicate a rapid demographic expansion for both species. For L. bostrychophila and L. entomophila, most of the pairwise FST values were statistically significant, which indicate that there was widespread population genetic differentiation within and between two species populations. However, the high levels of gene flow were also detected among these populations and this paradoxical phenomenon could be explained using the specific life habit of psocids.
(2) In the ITS dataset,74individuals of L. bostrychophila were sequenced and59haplotypes were detected whereas124individuals of L. entomophila were sequenced and77haplotypes were detected. The phylogenetic analyses and median-joining networks did not show clear pattern of structure for both species according to the same sampling localities, or predefined geographic region. In contrast to the AMOVAs result inferred from the Cytb dataset, several AMOVAs at different hierarchical levels inferred from ITS gene showed that there were no genetic structures of L. bostrychophila and L. entomophila populations according to our predefined regions, however pairwise estimates of FST among L. bostrychophila and L. entomophila populations revealed that the genetic differentiation existed among a large portion of populations. The neutrality tests and mismatch distribution test statistics for both psocid species populations indicated a rapid historically demographic expansion in the past.
(3) From above analysis, both Cytb and ITS sequences show high genetic diversity in these two species, and significant genetic structure and population differentiation were also detected. The high genetic diversity of L. bostrychophila and L. entomophila might explain why these species has a broad tolerance to environmental and habitat stresses and the fast mutational processes inherent in individuals, as well as populations, can enable these two Liposcelis species to successfully adapt to complex environments. Compared to sexually reproducing L. entomophila, asexually reproducing L. bostrychophila has a higher genetic diversity. The Mantel test indicated that for both species there was no evidence for isolation-by-distance (IBD). Therefore, the significant population differentiation detected in both psocids may be mainly due to other factors, such as genetic drift, inbreeding or control practices, and less by geographic distance. The neutrality test and mismatch distribution statistics revealed that the two species might have undergone population expansions in the past in the sampled areas.
3. In this study, we sequenced the mitochondrial (mt) genome of L. bostrychophila and found that the typical single mt chromosome of bilateral animals has fragmented into and been replaced by two medium-sized chromosomes; each of these chromosomes has about half of the genes of the typical mt chromosome of bilateral animals. These mt chromosomes are8,530bp (mt chromosome I) and7,933bp (mt chromosome II) in size. For mt chromosome I, six protein-coding genes and five tRNA genes were on one strand whereas one protein-coding gene, one rRNA gene and nine tRNA genes were on the other strand. For mt chromosome II, five protein-coding genes and four tRNA genes were on one strand whereas one protein-coding gene, one rRNA gene and five tRNA genes were on the other strand. There is an identical region (945bp) on both of the two mt chromosomes, and three tRNA genes and an extensive non-coding region that contains putative D-loop and control sequences were included in this region. Intriguingly, qPCR test indicated that mt chromosome I is twice as abundant as chromosome II. PCR test of multipartite mt genome are successful in other strains of L. bostrychophila, indicating that the2-chromosome structure is stable but with product size slightly change, which possibly due to deletion of some of non-coding regions. The mitochondrial gene order of L. bostrychophila differs radically from Lepidopsocidae sp.(Psocoptera) and the parasitic lice which are closest relatives of Liposcelididae. Totally, four putative pseudogenes on the two mitochondrial chromosomes demonstrated the formation of mini-circles was proceeding step by step with degeneration of the functional protein genes, and they are also powerful proofed that the recombination were happened between these two chromosomes. It appears that the selection pressure for compact mt genomes in bilateral animals favors small mt chromosomes when small mt chromosomes co-exist with the typical large mt chromosomes. Thus, small mt chromosomes may have selective advantages over large mt chromosomes in bilateral animals. Phylogenetic analyses of mt genome sequences of Psocodea (i.e. Psocoptera plus Phthiraptera) indicate that:1) the order Psocoptera (booklice and barklice) is paraphyletic; and2) the order Phthiraptera (the parasitic lice) is monophyletic. Within parasitic lice, however, the suborder Ischnocera is paraphyletic; this differs from the traditional view that each suborder of parasitic lice is monophyletic. From this study, we summarized that the mini-circle mt chromosomes which can replaced the typical animal single circle mt must have at least one functional gene and one non-coding region which has a high similarity across different mini-circles. We also speculated that the mt chromosomes in bilateral animals have the tendency of reducing their size and the fragmentation of mt genome is one of important strategies of processing of mt genome streaming. According to result of sequencing the mt genome of other Liposcelis species, we intend to conclude that multipartite mt genomes are common in Liposcelis species and the multipartite mt genome of Liposcelis species also give us invaluable insights into the origin and evolution of mitochondria in bilateral animals.
4. In the present paper, microsatellite-enriched libraries of L. bostrychophila and L. entomophila were constructed utilizing methodologies that exploit the strong affinity between biotin and the protein streptavidin. We propose a fast and easy protocol which is combination of two different published methods. Briefly, high-quality genomic DNA was digested by the restriction enzyme Msel and then ligated to designed adaptors.250-700bp microsatellite-containing DNA fragments were captured by streptavidin-coated magnetic beads. The beads affinity capture of microsatellite repeats using biotinylated oligonucleotide probes (AC)12,(TC)12,(ATC)8,(ATG)8,(AAC)8,(ATAC)6, and (GATA)6Subsequently, PCR was used to amplify the captured molecules for transferring single strand DNA to double strand DNA. The PCR products (enriched microsatellite DNA fragments) were ligated to pGEM-T Easy vector and transformed into Trans5a competent cells. In total,13microsatellite enriched libraries were constructed; six for L. bostrychophila, and seven for L. entomophila. A total of5,218clones were PCR screened for microsatellite content. The clones of L. bostiychophila and L. entomophila were2,542and2,676, respectively. The percentage of the microsatellite positive clones ranged9.38%-100%. The number of microsatellites detected for these two species was260. Comparative analysis of microsatellite sequences for these two psocids revealed that the microsatellite sequences exist in multiple copies in the psocid genome. In addition, they were found to have similar or almost identical flanking regions. From our present study, the tri-nucleotide libraries yielded greater results for determining gene flow.
In summary, in the present study, the genetic evolution patterns of psocids at both the inter-and intra-specific levels were analyzed systematically. Internal transcribed spacers (ITS) regions were cloned from six psocid species. Based on great variation of the ITS2sequences, a molecular identification method for six Liposcelis species was developed. The study here will provide potential molecular markers to investigate the evolutionary relationship among the species of genus Liposcelis and also will provide the DNA barcoding tools in psocids species. Population genetics of two asexually and sexually reproducing psocids, L. bostrychophila and L. entomophila were comparatively analyzed using the mitochondrial Cytb and nuclear ITS gene sequences and a thorough understanding of these two psocids population genetics, including gene flow patterns, genetic diversities and the resultant genetic structure among populations within China is useful for proposing successful integrated pest management tactics for these pests in grain storage systems. The completed mt genome of L. bostrychophila was sequenced and this multipartite mt genome provided new insights into the phylogeny of psocodea and the evolution of mitochondrial chromosomes in bilateral animals. We constructed13microsatellite enriched libraries for two psocids. Accumulation of multiple polymorphic microsatellite loci will greatly facilitate studies of individual identification and they are crucial to investigations of population genetics and evolution, gene linkage mapping, and elucidating the molecular phylogeny of psocids in China and internationally. Therefore, the studies included in this thesis have important theoretical and practical significance.
引文
1. Turner BD. Liposcelis bostrychophila (Psocoptera:Liposcelididae), a stored product food pest in the UK. International Journal of Pest Management,1994,40:179-190.
2. 李法圣,李志红,彩万志,杨定.中国啮目昆虫分类研究.中国农业大学学报,2008,13：7-18.
3. 赵朔,李志红,秦萌.书虱及其分子生物学研究进展.植物保护,2009,35：17-21.
4. 程伟霞,王进军.嗜卷书虱和嗜虫书虱抗性及防治方法的研究概况.安徽农业科学,2008,36：6376-6379.
5. Nayak MK, Collins PJ, Pavic H. Long-term effectiveness of grain protectants and structural treatments against Liposcelis decolor Pearman Psocoptera:Liposcelididae, a pest of stored products. Pest Management Science,2002,58:1223-1228.
6. Ding W, Wang JJ, Zhao ZM, Tsai JH. Effects of controlled atmosphere and DDVP on population growth and resistance development by the psocid, Liposcelis bostrychophila Badonnel (Psocoptera:Liposcelididae). Journal of Stored Products Research,2002,38: 229-237.
7. Wang JJ, Zhao ZM, Li LS. Selection of resistance strains in Liposcelis bostrychophila Badonnel to CO2-enriched atmospheres. Entomologia Sinica,1999,6:45-52.
8. Nayak MK. Potential of piperonyl butoxide-synergised pyrethrins against psocids (Psocoptera:Liposcelididae) for stored-grain protection. Pest Management Science,2010,66: 295-300.
9. Behar A, McCormick LJ, Perlman SJ. Rickettsia felis infection in a common household insect pest, Liposcelis bostrychophila (Psocoptera:Liposcelidae). Applied and Environmental Microbiology,2010,76:2280-2285.
10. Lin Y, Chan M, Ko C, Hsieh M. Nail infestation by Liposcelis bostrychophila badonnel. Clinical and Experiment Dermatology,2004,29:620-621.
11. Ahmedani MS, Shagufta N, Aslam M, Hussnain SA. Psocid:A new risk for global food security and safety. Applied Entomology and Zoology,2010,45:89-100.
12. Yoshizawa K, Johnson KP. How stable is the "Polyphyly of Lice" hypothesis (Insecta: Psocodea)?:A comparison of phylogenetic signal in multiple genes. Molecular Phylogenetics and Evolution,2010,55:939-951.
13. Ishiwata K, Sasaki G, Ogawa J, Miyata T, Su ZH. Phylogenetic relationships among insect orders based on three nuclear protein-coding gene sequences. Molecular Phylogenetics and Evolution,2010,58:169-180.
14. Yoshizawa K, Lienhard C, Johnson KP. Molecular systematics of the suborder Trogiomorpha (Insecta:Psocodea:'Psocoptera'). Zoological Journal of the Linnean Society, 2006,146:287-299.
15. Lyal CHC. Phylogeny and classification of the Psocodea, with particular reference to the lice (Psocodea:Phthiraptera). Systematic Entomology,1985,10:145-165.
16. Grimaldi D, Engel MS. Fossil Liposcelididae and the lice ages (Insecta:Psocodea). Proceedings of the Royal Society B-Biological Sciences,2006,273:625-633.
17. Leong ECW, Ho S. Life cycle of Liposcelis entomophila (Psocoptera:Liposcelididae) and a culturing regime for liposcelids. Bulletin of Entomological Research,1995,85:501-506.
18. 王进军,赵志模,郭依泉,周亦红,周晨曦.温度、湿度对嗜虫书虱生长发育和繁殖的影响.植物保护学报,1996,23：147-151.
19. Wang JJ, Ren Y, Wei XQ, Dou W. Development, survival, and reproduction of the psocid Liposcelis paeta (Psocoptera:Liposcelididae) as a function of temperature. Journal of Economic Entomology,2009,102:1705-1713.
20. Green PWC. Insect-derived compounds affect the behaviour of Liposcelis bostrychophila: Effects of combination and structure. Journal of Stored Products Research,2011,47: 262-266.
21. Xiao LS, Dou W, Li Y, Wang JJ. Comparative studies of acetylcholinesterase purified from three field populations of Liposcelis entomophila (enderlein) (psocoptera:liposcelididae). Archives of Insect Biochemistry and Physiology,2010,75:158-173.
22. Dou W, Xiao LS, Niu JZ, Jiang HB, Wang JJ. Characterization of the purified glutathione S-transferases from two psocids Liposcelis bostiychophila and L. entomophila. Agricultural Sciences in China,2010,9:1008-1016.
23. Ren Y, Wei XQ, Wu SA, Dou W, Wang JJ. Comparison of acetylcholinesterase from three field populations of Liposcelis paeta Pearman (Psocoptera:Liposcelididae):Implications of insecticide resistance. Pesticide Biochemistry and Physiology,2008,90:196-202.
24. Cheng WX, Dou W, Chai YX, Wang JJ. Comparison of biochemical and toxicological characterizations of glutathione S-transferases and superoxide dismutase between Liposcelis bostrychophila Badonnel and L. entomophila (Enderlein) (Psocoptera:Liposcelididae). Pesticide Biochemistry and Physiology,2007,89:151-157.
25. Wu S, Li M, Tang PA, Felton GW, Wang JJ. Cloning and characterization of acetylcholinesterase 1 genes from insecticide-resistant field populations of Liposcelis paeta Pearman (Psocoptera:Liposcelididae). Insect Biochemistry and Molecular Biology,2010, 40:415-424.
26. Jiang HB, Tang PA, Xu YQ, An FM, Wang JJ. Molecular characterization of two novel deltamethrin-inducible P450 genes from Liposcelis bostrychophila Badonnel (Psocoptera: Liposcelididae). Archives of Insect Biochemistry and Physiology,2010,74:17-37.
27. Tang PA, Jiang HB, Xu YQ, An FM, Wang JJ. Molecular characterization of two nicotinic acetylcholine receptor subunits from Liposcelis bostrychophila Badonnel (Psocoptera: Liposcelididae). Archives of Insect Biochemistry and Physiology,2009,72:34-47.
28. Mikac KM, FitzSimmons NN. Genetic structure and dispersal patterns of the invasive psocid Liposcelis decolor (Pearman) in Australian grain storage systems. Bulletin of Entomological Research,2010,100:521-527.
29. Mikac KM, Clarke GM. Tracing the geographic origin of the cosmopolitan parthenogenetic insect pest Liposcelis bostrychophila (Psocoptera:Liposcelididae). Bulletin of Entomological Research,2006,96:523-530.
30. 丁伟,李隆术,赵志模.书虱综合防治技术研究进展.粮食储藏,2001,30：3-6.
31. 周世良,张方,王中仁.等位酶淀粉凝胶电泳技术中的几个应引起重视的问题.植物学通报,1998,15：68-72.
32. 王进军.嗜卷书虱种群生态及其对气调抗性的机理研究.博士学位论文,1997,重庆：西南农业大学.
33. Ali N, Turner B. Allozyme polymorphism and variability in permethrin tolerance in British population of the parthenogenetic stored product pest Liposcelis bostrychophila (Liposcelididae, Psocoptera). Journal of Stored Products Research,2001,37:111-125.
34. Mikac KM. PCR confirms multiple Wolbachia strain infection in Australian and international populations of the invasive stored-product psocid Liposcelis bostrychophila Badonnel. Journal of Stored Products Research,2007,43:594-597.
35. Qin M, Li ZH, Kucerova Z, Cao Y, Stejskal V. Rapid discrimination of the common species of the stored product pest Liposcelis (Psocoptera:Liposcelididae) from China and the Czech Republic, based on PCR-RFLP analysis. European Journal of Entomology,2008,105: 713-717.
36. Tautz D. Hypervariability of simple sequences as a general source for polymorphic DNA markers. Nucleic Acids Research,1989,17:6463-6471.
37. Selkoe KA, Toonen RJ. Microsatellites for ecologists:a practical guide to using and evaluating microsatellite markers. Ecology Letters,2006,9:615-629.
38. Mikac KM. Isolation and characterization of the first microsatellite loci from the order Psocoptera in the economically important pest insect Liposcelis decolor (Pearman) and cross-species amplification. Molecular Ecology Resources,2006,6:1102-1104.
39. 魏丹丹,袁明龙,王保军,朱礼明,王进军.两种书虱微卫星富集文库的构建及比较.生态学报,2011,31：4182-4189.
40. Behura SK. Molecular marker systems in insects:current trends and future avenues. Molecular Ecology,2006,15:3087-3113.
41. Yoshizawa K, Johnson KP. Phylogenetic position of Phthiraptera (Insecta:Paraneoptera) and elevated rate of evolution in mitochondrial 12S and 16S rDNA. Molecular Phylogenetics and Evolution,2003,29:102-114.
42. Johnson K, Mockford EL. Molecular systematics of Psocomorpha (Psocoptera). Systematic Entomology,2003,28:409-416.
43. Yoshizawa K. Molecular phylogeny of major lineages of Trichadenotecnum and a review of diagnostic morphological characters (Psocoptera:Psocidae). Systematic Entomology,2004, 29:383-394.
44. Yoshizawa K, Johnson KP. Molecular systematics of the barklouse family Psocidae (Insecta: Psocodea:'Psocoptera') and implications for morphological and behavioral evolution. Molecular phylogenetics and evolution,2008,46:547-559.
45. Wei DD, Yuan ML, Wang BJ, Zhou AW, Dou W, Wang JJ. Population genetics of two asexually and sexually reproducing psocids species inferred by the analysis of mitochondrial and nuclear DNA sequences. PLoS ONE,2012,7:e33883.
46. Li ZH, Kucerova Z, Zhao S, Stejskal V, Opit G, Qin M. Morphological and molecular identification of three geographical populations of the storage pest Liposcelis bostrychophila (Psocoptera). Journal of Stored Products Research,2011,47:168-172.
47. Arif M, Ochoa-Corona FM, Opit GP, Li ZH, Kucerova Z, Stejskal V, Yang QQ. PCR and isothermal-based molecular identification of the stored-product psocid pest Lepinotus reticulatus (Psocoptera:Trogiidae). Journal of Stored Products Research,2012,49:184-188.
48. Yang QQ, Kucerova Z, Li ZH, Kalinovic I, Stejskal V, Opit G, Cao Y. Diagnosis of Liposcelis entomophila (Insecta:Psocodea:Liposcelididae) based on morphological characteristics and DNA barcodes. Journal of Stored Products Research,2012,48:120-125.
49. Yuan ML, Wang BJ, Lu F, Hu CX, Wei DD, Dou W, Wang JJ. Evaluation of genetic diversity and population structure of Panonychus citri (Acari:Tetranychidae) in China using ribosomal internal transcribed spacer 1 sequences. Annals of the Entomological Society of America,2011,104:800-807.
50. Wei DD, Yuan ML, Wang ZY, Wang D, Wang BJ, Dou W, Wang JJ. Sequence analysis of the ribosomal internal transcribed spacers region in psocids (Psocoptera:Liposcelididae) for phylogenetic inference and species discrimination. Journal of Economic Entomology,2011, 104:1720-1729.
51. Lang BF, Gray MW, Burger G. Mitochondrial genome evolution and the origin of eukaryotes. Annual Review of Genetics,1999,33:351-397.
52. Papillon D, Perez Y, Caubit X, Parco YL. Identification of chaetognaths as protostomes is supported by the analysis of their mitochondrial genome. Molecular Biology and Evolution, 2004,21:2122-2129.
53. Gissi C, Iannelli F, Pesole G. Evolution of the mitochondrial genome of Metazoa as exemplified by comparison of congeneric species. Heredity,2008,101:301-320.
54. Dermauw W, Vanholme B, Tirry L, Leeuwen TV. Mitochondrial genome analysis of the predatory mite Phytoseiulus persimilis and a revisit of the Metaseiulus occidentalis mitochondrial genome. Genome,2010,53:285-301.
55. Hu M, Chilton NB, Gasser RB. The mitochondrial genome of Strongyloides stercoralis (Nematoda)-idiosyncratic gene order and evolutionary implications. International Journal for Parasitology,2003,33:1393-1408.
56. Burger G, Gray MW, Lang BF. Mitochondrial genomes:anything goes. TRENDS in Genetics,2003,19:709-716.
57. Yuan ML, Wei DD, Wang BJ, Dou W, Wang JJ. The complete mitochondrial genome of the citrus red mite Panonychus citri (Acari:Tetranychidae):high genome rearrangement and extremely truncated tRNAs. BMC Genomics,2010,11:597.
58. Shao R, Mitani H, Barker SC, Takahashi M, Fukunaga M. Novel mitochondrial gene content and gene arrangement indicate illegitimate inter-mtDNA recombination in the chigger mite, Leptotrombidium pallidum. Journal of Molecular Evolution,2005,60: 764-773.
59. Shao R, Barker SC, Mitani H, Aoki Y, Fukunaga M. Evolution of duplicate control regions in the mitochondrial genomes of metazoa:a case study with Australasian Ixodes ticks. Molecular Biology and Evolution,2005,22:620-629.
60. Cameron SL, Yoshizawa K, Mizukoshi A, Whiting MF, Johnson KP. Mitochondrial genome deletions and minicircles are common in lice (Insecta:Phthiraptera). BMC Genomics,2011, 12:394.
61. Watanabe KI, Bessho Y, Kawasaki M, Hori H. Mitochondrial genes are found on minicircle DNA molecules in the mesozoan animal Dicyema. Journal of Molecular Biology,1999,286: 645-650.
62. Awata H, Noto T, Endoh H. Differentiation of somatic mitochondria and the structural changes in mtDNA during development of the dicyemid Dicyema japonicum (Mesozoa). Molecular Genetics and Genomics,2005,273:441-449.
63. Suga K, Welch DBM, Tanaka Y, Sakakura Y, Hagiwarak A. Two circular chromosomes of unequal copy number make up the mitochondrial genome of the rotifer Brachionus plicatilis. Molecular Biology and Evolution,2008,25:1129-1137.
64. Armstrong MR, Blok VC, Phillips MS. A multipartite mitochondrial genome in the potato cyst nematode Globodera pallida. Genetics,2000,154:181-192.
65. Gibson T, Blok VC, Dowton M. Sequence and characterization of six mitochondrial subgenomes from Globodera rostochiensis:multipartite structure is conserved among close nematode relatives. Journal of Molecular Evolution,2007,65:308-315.
66. Sugiyama Y, Watase Y, Nagase M, Makita N, Yagura S, Hirai A, Sugiura M. The complete nucleotide sequence and multipartite organization of the tobacco mitochondrial genome: comparative analysis of mitochondrial genomes in higher plants. Molecular Genetics and Genomics,2005,272:603-615.
67. Burger G, Lang B F. Parallels in genome evolution in mitochondria and bacterial symbionts. IUBMB Life,2003,55:205-212.
68. Voigt O, Erpenbeck D, Worheide G. A fragmented metazoan organellar genome:the two mitochondrial chromosomes of Hydra magnipapillata. BMC Genomics,2008,9:350.
69. Fan J, Lee RW. Mitochondrial genome of the colorless green alga Polvtomella parva:two linear DNA molecules with homologous inverted repeat termini. Molecular Biology and Evolution,2002,19:999-1007.
70. Nosek J, Tomaska L, Fukuhara H, Suyama Y, Kovac L. Linear mitochondrial genomes:30 years down the line. TRENDS in Genetics,1998,14:184-188.
71. Marande W, Lukes J, Burger G. Unique mitochondrial genome structure in diplonemids, the sister group of kinetoplastids. Eukaryotic Cell,2005,4:1137-1146.
72. Gibson T, Blok VC, Phillips MS, Hong G, Kumarasinghe D, Riley IT, Dowton M. The mitochondrial subgenomes of the nematode Globodera pallida are mosaics:evidence of recombination in an animal mitochondrial genome. Journal of Molecular Evolution,2007, 64:463-471.
73. Shao R, Kirkness EF, Barker SC. The single mitochondrial chromosome typical of animals has evolved into 18 minichromosomes in the human body louse, Pediculus humanus. Genome Research,2009,19:904-912.
74. Melov S, Lithgow GJ, Fischer DR, Tedesco PM, Johnson TE. Increased frequency of deletions in the mitochondrial genome with age of Caenorhabditis elegans. Nucleic Acids Research,1995,23:1419-1425.
75. Fauron C, Casper M, Gao Y, Moore B. The maize mitochondrial genome:dynamic, yet functional. Trends in Genetics,1995,11:228-235.
76. Shao R, Barker SC. Chimeric mitochondrial minichromosomes of the human body louse, Pediculus humanus:Evidence for homologous and non-homologous recombination. Gene, 2011,473:36-43.
77. Dowton M, Austin AD. Evolutionary dynamics of a mitochondrial rearrangement "Hot Spot" in the Hymenoptera. Molecular Biology and Evolution,1999,16:298-309.
78. Mueller R, Boore J. Molecular mechanisms of extensive mitochondrial gene rearrangement in plethodontid salamanders. Molecular Biology and Evolution,2005,22:2104-2112.
79. Wei DD, Shao R, Yuan ML, Dou W, Barker SC, Wang JJ. The multipartite mitochondrial genome of Liposcelis bostiychophila:insights into the evolution of mitochondrial genomes in bilateral animals. PLoS ONE,2012,7:e33973.
80. Landweber LF. Why genomes in pieces? Science,2007,318:405-407.
81. Rand DM.'Why genomes in pieces?' revisited:Sucking lice do their own thing in mtDNA circle game. Genome Research,2009,19:700-702.
82. Kirkness E, Haas B, Sun W, Braig H, Perotti M, Clark J, Lee S, Robertson H, Kennedy R, Elhaik E, Gerlach D, Kriventseva E, Elsik C, Graur D, Hill C, Veenstra J, Walenz B, Tubio J, Ribeiro J, Rozas J, Johnston J, Reese J, Popadic A, Tojo M, Raoult D, Reed D, Tomoyasu Y, Krause E, Mittapalli O, Margam V, Li H, Meyer J, Johnson R, Romero-Severson J, Vanzee J, Alvarez-Ponce D, Vieira F, Aguade M, Guirao-Rico S, Anzola J, Yoon K, Strycharz J, Unger M, Christley S, Lobo N, Seufferheld M, Wang N, Dasch G, Struchiner C, Madey G, Hannick L, Bidwell S, Joardar V, Caler E, Shao R, Barker S, Cameron S, Bruggner R, Regier A, Johnson J, Viswanathan L, Utterback T, Sutton G, Lawson D, Waterhouse R, Venter J, Strausberg R, Berenbaum M, Collins F, Zdobnov E, Pittendrigh B. Genome sequences of the human body louse and its primary endosymbiont provide insights into the permanent parasitic lifestyle. Proceedings of the National Academy of Sciences of the United States of America,2010,107:12168-12173.
83. Korhonen JA, Pham XH, Pellegrini M, Falkenberg M. Reconstitution of a minimal mtDNA replisome in vitro. EMBO Journal,2004,23:2423-2429.
84. Johnson KP, Yoshizawa K, Smith VS. Multiple origins of parasitism in lice. Proceedings of the Royal Society B-Biological Sciences,2004,271:1771-1776.
85. Mockford EL. Some Psocoptera from the plumage of birds. Proceedings of the Entomological Society of Washington,1967,69:307-309.
86. Pearlman JV. Some African Psocoptera found on rats. Entomologist,1960,93:246-250.
87. Lienhard C. Revision of the western palaearctic species of Liposcelis Motschulsky (Psocoptera:Liposcelididae). Zoologische Jahrbucher. Abteilung fur Systematik, Okologie und Geographie der Tiere,1990,117:117-174.
88. Li YC, Korol AB, Fahima T, Beiles A, Nevo E. Microsatellites:genomie distribution, putative functions and mutational mechanisms:a review. Molecular Ecology,2002,11: 2453-2465.
89. Zane L, Bargelloni L, Patarnello T. Strategies for microsatellite isolation:a review. Molecular Ecology,2002,11:1-16.
90. Yu J, Won C, Jun J, Lim Y, Kwak M. Fast and cost-effective mining of microsatellite markers using NGS technology:an example of a Korean water deer Hydropotes inermis argyropus. PLoS ONE,2011,6:e26933.
91. Archak S, Meduri E, Sravana KP, Nagaraju J. InSatDb:A microsatellite database of fully sequenced insect genomes. Nucleic Acids Research,2007,35:D36-D39.
92. Aishwarya V, Grover A, Sharma PC. EuMicroSatdb:A database for microsatellites in the sequenced genomes of eukaryotes. BMC Genomics,2007,8:225.
93. Chen MH, Dorn S. Cross-amplification of microsatellites from the codling moth Cydia pomonella to three other species of the tribe Grapholitini (Lepidoptera:Tortricidae). Molecular Ecology Resources,2010,10:1034-1037.
94. Ma H, Chen S. Development of polymorphic microsatellite markers in barfin flounder (Verasper moseri) and spotted halibut(Verasper variegatus) by the cross-species amplification. Molecular Biology Reports,2010,38:4545-4551.
95. Barbara T, Palma-Silva C, Paggi G, Bered F, Fay M, Lexer C. Cross-species transfer of nuclear microsatellite markers:potential and limitations. Molecular Ecology,2007,16: 3759-3767.
96. Merkel A, Gemmell NJ. Detecting microsatellites in genome data:variance in definitions and bioinformatic approaches cause systematic bias. Evolutionary Bioinformatics,2008,4: 1-6.
97. Durand J, Bodenes C, Chancerel E, Frigerio JM, Vendramin G, Sebastiani F, Buonamici A, Gailing O, Koelewijn HP, Villani F, Mattioni C, Cherubini M, Goikoetxea P, Herran A, Ikaran Z, Cabane C, Ueno S, Alberto F, Dum PY, Guichoux E, Daruvar A, Kremer A, Plomion C. A fast and cost-effective approach to develop and map EST-SSR markers:oak as a case study. BMC Genomics,2010,11:570.
98. 张琳琳,魏朝明,廉振民,孔光耀.赤拟谷盗全基因组和EST中微卫星的丰度.昆虫知识,2008,45：38-42.
99. 刘玉娣,侯茂林.褐飞虱EST资源的微卫星信息分析.昆虫学报,2010,53：239-247.
100. Mikheyev AS, Tanya Vol BW, Singer MC, Parmesan C. Rapid microsatellite isolation from a butterfly by de novo transcriptome sequencing:performance and a comparison with AFLP-derived distances. PLoS ONE,2010,5:e11212.
101. 孙波,鲍毅新,赵庆洋,张龙龙,胡知渊.微卫星位点获取方法的研究进展.生态学杂志,2009,28：2130-2137.
102. Andres J, Bogdanowicz S. Isolating microsatellite Loci:looking back, looking ahead. Methods in Molecular Biology,2011,772:211-32.
103. Glenn TC, Schable NA. Isolating microsatelline DNA loci. Methods in Enzymology,2005, 395:202-222.
104. Kandpal RP, Kandpal G, Weissman SM. Construction of libraries enriched for sequence repeats and jumping clones, and hybridization selection for region specific markers. Proceedings of the National Academy of Sciences of the United States of America,1994,91: 88-92.
105. Techen N, Arias RS, Glynn NC, Pan Z, Khan IA, Scheffler BE. Optimized construction of microsatellite-enriched libraries. Molecular Ecology Resources,2010,10:508-515.
106. 邓欣,陈信波,龙松华,王孝纯,高原,何东锋,王进,王玉富.用磁珠富集法分离亚麻基因组微卫星分子标记.作物学报,2008,34：2100-2105.
107. Fisher P, Gardner R, Richardson T. Single locus microsatellites isolated using 5'anchored PCR. Nucleic Acids Research,1996,24:4369-4371.
108. Hayden MJ, Sharp PJ. Targeted development of informative microsatellite (SSR) markers. Nucletic Acids Research,2001,29:e44.
109. Shao R, Dowton M, Murrell A, Barker SC. Rates of gene rearrangement and nucleotide substitution are correlated in the mitochondrial genomes of insects. Molecular Biology and Evolution,2003,20:1612-1619.
110. Leeuwen TV, Vanholme B, Pottelberge SV, Nieuwenhuyse PV, Nauen R, Tirry L, Denholm I. Mitochondrial heteroplasmy and the evolution of insecticide resistance:Non-Mendelian inheritance in action. Proceedings of the National Academy of Sciences of the United States of America,2008,105:5980-5985.
111. 李法圣,中国啮目志.2002,北京：科学出版社.
112. Wang ZY, Wang JJ, Lienhard C. Two new species of Liposcelis (Psocoptera, Liposcelididae). Acta Zootaxonomica Sinica,2006,31:564-568.
113. Tautz D, Arctander P, Minelli A, Thomas RH, Vogler AP. DNA points the way ahead in taxonomy. Nature,2002,418:479.
114. Hebert PDN, Cywinska A, Ball SL, deWaard JR. Biological identifications through DNA barcodes. Proceedings of the Royal Society B-Biological Sciences,2003,270:313-321.
115. Hebert PDN, Ratnasingham S, de Waard JR. Barcoding animal life:cytochrome c oxidase subunit 1 divergences among closely related species. Proceedings of the Royal Society B-Biological Sciences,2003,270:S96-S99.
116. Galtier N, Nabholz B, Glemin S, Hurst GD. Mitochondrial DNA as a marker of molecular diversity:a reappraisal. Molecular Ecology,2009,18:4541-4550.
117. Yao H, Song J, Liu C, Luo K, Han J, Li Y, Pang X, Xu H, Zhu Y, Xiao P, Chen S. Use of ITS2 region as the universal DNA barcode for plants and animals. PLoS ONE,2010,5: e13102.
118. Li DZ, Gao LM, Li HT, Wang H, Ge XJ, Liu JQ, Chen ZD, Zhou SL, Chen SL, Yang JB, Fu CX, Zeng CX, Yan HF, Zhu YJ, Sun YS, Chen SY, Zhao L, Wang K, Yang T, Duan GW. Comparative analysis of a large dataset indicates that internal transcribed spacer (ITS) should be incorporated into the core barcode for seed plants. Proceedings of the National Academy of Sciences of the United States of America,2011,108:19641-19646.
119. Matsumoto Y, Yanase T, Tsuda T, Noda H. Characterization of internal transcribed spacer (ITS1)-ITS2 region of ribosomal RNA gene from 25 species of Culicoides biting midges (Diptera:Ceratopogonidae) in Japan. Journal of Medical Entomology,2009,46:1099-1108.
120. Oh HK, Yoon HJ, Kim MJ, Jeong HU, Kim SR, Hwang JS, Bae CH, Kim I. ITS2 ribosomal DNA sequence variation of the bumblebee, Bombus ardens (Hymenoptera:Apidae). Genes & Genomics,2009,31:293-303.
121. Wiemer M, Keller A, Wolf M. ITS2 secondary structure improves phylogeny estimation in a radiation of blue butterflies of the subgenus Agrodiaetus (Lepidoptera:Lycaenidae: Polyommatus). BMC Evolutionary Biology,2009,9:300.
122. Stephan A, Clausen PH, Bauer B, Steuber S. PCR identification of Culicoides dewulfi midges (Diptera:Ceratopogonidae), potential vectors of bluetongue in Germany. Parasitology Research,2009,105:367-371.
123. Rugman-Jones PF, Hoddle MS, Mound LA, Stouthamer R. Molecular identification key for pest species of Scirtothrips (Thysanoptera:Thripidae). Journal of Economic Entomology, 2006,99:1813-1819.
124. Farris RE, Ruiz-Arce R, Ciomperlik M, Vasquez JD, DeLeon R. Development of a ribosomal DNA ITS2 marker for the identification of the thrips, Scirtothrips dorsalis. Journal of Insect Science,2010,10:26.
125. Kent RJ, Deus S, Williams M, Savage HM. Development of a multiplexed polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) assay to identify common members of the Subgenera Culex(Culex) and Culex(Phenacomyia) in Guatemala. The American Journal of Tropical Medicine and Hygiene,2010,83:285-291.
126. Jia FX, Yang MS, Yang WJ, Wang JJ. Influence of continuous high temperature conditions on Wolbachia infection frequency and the fitness of Liposcelis tricolor (Psocoptera: Liposcelididae). Environmental Entomology,2009,38:1365-1372.
127. Keller A, Schleicher T, Schultz J, Muller T, Dandekar T, Wolf M.5.8S-28S rRNA interaction and HMM-based ITS2 annotation. Gene,2009,430:50-57.
128. Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG. The ClustalX windows interface:flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research,1997,24:4876-4882.
129. Librado P, Rozas J. DnaSP v5:a software for comprehensive analysis of DNA polymorphism data. Bioinformatics,2009,25:1451-1452.
130. Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S. MEGA5:molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution,2011,28:2731-2739.
131. Castresana J. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution,2000,17:540-552.
132. Posada D. jModelTest:phylogenetic model averaging. Molecular Biology and Evolution, 2008,25:1253-1256.
133. Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O. New algorithms and methods to estimate maximum-likelihood phylogenies:assessing the performance of PhyML 3.0. Systematic Biology,2010,59:307-321.
134. Page RDM. TreeView:an application to display phylogenetic trees on personal computers. Computer Applications in The Biosciences,1996,12:357-358.
135. Ullrich B, Reinhold K, Niehuis O, Misof B. Secondary structure and phylogenetic analysis of the internal transcribed spacers 1 and 2 of bush crickets (Orthoptera:Tettigoniidae: Barbitistini). Journal of Zoological Systematics and Evolutionary Research,2010,48: 219-228.
136. Kengne P, Antonio-Nkondjio C, Awono-Ambene HP, Simard F, Awolola TS, Fontenille D. Molecular differentiation of three closely related members of the mosquito species complex, Anopheles moucheti, by mitochondrial and ribosomal DNA polymorphism. Medical and Veterinary Entomology,2007,21:177-182.
137. Li JJ, Ledger TW, Tredici PD. Phylogenetics of Calycanthaceae based on molecular and morphological data with special reference to divergent paralogues of the nrDNA ITS region. Harvard Papers in Botany,2004,9:69-82.
138. Harpke D, Peterson A.5.8S motifs for the identification of pseudogenic ITS regions. Botany, 2008,86:300-305.
139. David J, Huber K, Failloux A, Rey D, Meyran J. The role of environment in shaping the genetic diversity of the subalpine mosquito, Aedes rusticus (Diptera, Culicidae). Molecular Ecology,2003,12:1951-1961.
140. Amos W, Harwood J. Factors affecting levels of genetic diversity in natural populations. Philosophical Transactions of the Royal Society B-Biological Sciences,1998,353:177-186.
141. Markert JA, Champlin DM, Gutjahr-Gobell R, Grear JS, Kuhn A, Jr TJM, Roth A, Bagley MJ, Nacci DE. Population genetic diversity and fitness in multiple environments. BMC Evolutionary Biology,2010,10:205.
142. Rua N, Stevens L, Dorn PL. High genetic diversity in a single population of Triatoma sanguisuga (LeConte,1855) inferred from two mitochondrial markers:cytochrome b and 16S ribosomal DNA. Infection, Genetics and Evolution,2011,11:671-677.
143. Collins F, Paskewitz S. A review of the use of ribosomal DNA (rDNA) to differentiate among cryptic Anopheles species. Insect Molecular Biology,1996,5:1-9.
144. Suomalainen E, Saura A, Lokki J, Teeri T. Genetic polymorphism and evolution in parthenogenetic animals. Theoretical and Applied Genetics,1980,57:129-132.
145. Weeks AR. Marec F, Breeuwer JAJ. A mite species that consists entirely of haploid females. Science,2001,292:2479-2482.
146. Simon JC, Delmotte F, Rispe C, Crease T. Phylogenetic relationships between parthenogens and their sexual relatives:the possible routes to parthenogenesis in animals. Biological Journal of the Linnean Society,2003,79:151-163.
147. Bengtsson BO. Genetic variation in organisms with sexual and asexual reproduction. Journal of Evolutionary Biology,2003,16:189-199.
148. Shreve S, Mockford E, Johnson K. Elevated genetic diversity of mitochondrial genes in asexual populations of bark lice ("Psocoptera":Echmepteryx hageni). Molecular Ecology, 2011,20:4433-4451.
149. Jiang HB, Liu YH, Tang PA, Zhou AW, Wang JJ. Validation of endogenous reference genes for insecticide-induced and developmental expression profiling of Liposcelis bostrychophila (Psocoptera:Liposcelididae). Molecular Biology Reports,2010,37:1019-1029.
150. Guedes RNC, Zhu KY, Opit GP, Throne JE. Differential heat shock tolerance and expression of heat-inducible proteins in two stored-product psocids. Journal of Economic Entomology,2008,101:1974-1982.
151. Simon C, Frati F, Beckenbach A, Crespi B, Liu H, Flook P. Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America,1994, 87:651-701.
152. Hall TA. BIOEDIT:a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series,1999,41:95-98.
153. Zuker M. Mfold web server for nucleic acid folding and hybridization prediction. Nucleic Acids Research,2003,31:3406-3415.
154. Wuyts H, De Rijk P, Van de Peer Y, Winkelmans T, De Wachter R. The European large subunit ribosomal RNA database. Nucleic Acids Research,2001,29:175-177.
155. Excoffier L, Laval G, Schneider S. Arlequin (version 3.0):an integrated software package for population genetics data analysis. Evolution Bioinformatics Online,2005,1:47-50.
156. Dupanloup I, Schneider S, Excoffier L. A simulated annealing approach to define the genetic structure of populations. Molecular Ecology,2002,11:2571-2581.
157. Jensen JL, Bohonak AJ, Kelley ST. Isolation by distance, web service. BMC Genetics,2005, 6:13.
158. Tajima F. Statistical method for testing the neutral mutation hypothesis by DNA polymorphism. Genetics,1989,123:585-595.
159. Fu YX. Statistical tests of neutrality of mutations against population growth, hitchhiking and background selection. Genetics,1997,147:915-925.
160. Rogers AR, Harpending H. Population growth makes waves in the distribution of pairwise genetic differences. Molecuar Biology and Evolution,1992,9:552-569.
161. Dsouli-Aymes N, Michaux J, Stordeur ED, Couloux A, Veuille M, Duvallet G. Global population structure of the stable fly(Stomoxys calcitrans) inferred by mitochondrial and nuclear sequence data. Infection, Genetics and Evolution,2011,11:334-342.
162. Xia X, Xie Z. DAMBE:software package for data analysis in molecular biology and evolution. The Journal of Heredity,2001,92:371-373.
163. Ronquist F, Huelsenbeck JP. MrBayes 3:Bayesian phylogenetic inference under mixed models. Bioinformatics,2003,19:1572-1574.
164. Huelsenbeck JP, Ronquist F, Nielsen R, Bollback JP. Bayesian inference of phylogeny and its impact on evolutionary biology. Science,2001,294:2310-2314.
165. Posada D, Crandall K. Intraspecific gene genealogies:trees grafting into networks. TRENDS in Ecology and Evolution,2001,16:37-45.
166. Bandelt HJ, Forster P, Rohl A. Median-Joining networks for inferring intraspecific phylogenies. Molecular Biology and Evolution,1999,16:37-48.
167. Szalanski AL, Austin JW, McKern JA, Steelman CD, Gold RE. Mitochondrial and ribosomal internal transcribed spacer 1 diversity of Cimex lectularius (Hemiptera: Cimicidae). Journal of Medical Entomology,2008,45:229-236.
168. Loxdale HD. Rapid genetic changes in natural insect populations. Ecological Entomology, 2010,35:155-164.
169. Mockford EL, Krushelnycky PD. New species and records of Liposcelis Motschulsky (Psocoptera:Liposcelididae) from Hawaii with first description of the male of Liposcelis bostrychophila Badonnel. Zootaxa,2008,1766:53-68.
170. Wang JJ, Dong P, Xiao LS, Dou W. Effects of removal of Cardinium infection on fitness of the stored-product pest Liposcelis bostrychophila (Psocoptera:Liposcelididae). Journal of Economic Entomology,2008,101:1711-1717.
171. Yusuf M, Turner B, Whitfield P, Miles R, Pacey J. Electron microscopical evidence of a vertically transmitted Wolbachia-like parasite in the parthenogenetic, stored-product pest Liposcelis bostrychophila Badonnel (Psocoptera). Journal of Stored Products Research, 2000,36:169-175.
172. Yu MZ, Zhang KJ, Xue XF, Hong XY. Effects of Wolbachia on mtDNA variation and evolution in natural populations of Tetranychus urticae Koch. Insect Molecular Biology, 2011,20:311-321.
173. Narita S, Nomura M, Kato Y, Fukatsu T. Genetic structure of sibling butterfly species affected by Wolbachia infection sweep:evolutionary and biogeographical implications. Molecular Ecology,2006,15:1095-1108.
174. Fu Y, Li. W. Statistical tests of neutrality of mutations. Genetics,1993,133:693-709.
175. Slatkin M. Isolation by distance in equilibrium and non-equilibrium populations. Evolution, 1993,47:264-279.
176. Douglas CW. Mitochondrial DNA mutations in disease and aging. Environmental and Molecular Mutagenesis,2010,51:440-450.
177. Boore JL. Animal mitochondrial genomes. Nucleic Acids Research,1999,27:1767-1780.
178. Murrell A, Barker SC. Multiple origins of parasitism in lice:phylogenetic analysis of SSU rDNA indicates that the Phthiraptera and Psocoptera are not monophyletic. Parasitology Research,2005,97:274-280.
179. Kim MJ, Kang AR, Jeong HC, Kim KG, Kim I. Reconstructing intraordinal relationships in Lepidoptera using mitochondrial genome data with the description of two newly sequenced lycaenids, Spindasis takanonis and Protantigius superans (Lepidoptera:Lycaenidae). Molecular Phylogenetics and Evolution,2011,61:436-445.
180. Cameron S, Lambkin C, Barker S, Whiting M. A mitochondrial genome phylogeny of Diptera:whole genome sequence data accurately resolve relationships over broad timescales with high precision. Systematic Entomology,2007,32:40-59.
181. Wei S, Shi M, Sharkey MJ, Achterberg Cv, Chen X. Comparative mitogenomics of Braconidae (Insecta:Hymenoptera) and the phylogenetic utility of mitochondrial genomes with special reference to Holometabolous insects. BMC Genomics,2010,11:371.
182. 龚鹏,杨效文,张孝羲,刘向东,陈晓峰.棉蚜(Aphis gossypii)种群寄主分化和季节分化的微卫星引物PCR研究.生态学报,2001,21：765-771.
183. Zhang DX, Hewitt GM. Assessment of the universality and utility of a set of conserved mitochondrial COI primers in insects. Insect Molecular Biology,1997,6:143-150.
184. Simon C, Buckley TR, Frati F, Stewart JB, Beckenbach AT. Incorporating molecular evolution into phylogenetic analysis, and a new compilation of conserved polymerase chain reaction primers for animal mitochondrial DNA. Annual Review of Ecology, Evolution, and Systematics,2006,37:545-79.
185. Hopp TP, Woods KR. Prediction of protein antigenic determinants from amino acid sequences. Proceedings of the National Academy of Sciences of the United States of America, 1981, 78: 3824-3828.
186. Lowe TM, Eddy SR. tRNAscan-SE: A program for improved detection of transfer RNA genes in genomic sequence. Nucleic Acids Research, 1997, 25: 955-964.
187. Laslett D, Canback B. ARWEN: a program to detect tRNA genes in metazoan mitochondrial nucleotide sequences. Bioinformatics, 2008, 24: 172-175.
188. Mathews DH. Predicting a set of minimal free energy RNA secondary structures common to two sequences. Bioinformatics, 2005, 21: 2246-2253.
189. Pfaffl MW. A new mathematical model for relative quantification in real-time RT-PCR. Nucleic Acids Research, 2001, 29: e45.
190. Cameron SL, Johnson KP, Whiting MF. The mitochondrial genome of the screamer louse Bothriometopus (Phthiraptera: Ischnocera): effects of extensive gene rearrangements on the evolution of the genome. Journal of Molecular Evolution, 2007, 65: 589-604.
191. Covacin C, Shao R, Cameron S, Barker SC. Extraordinary number of gene rearrangements in the mitochondrial genomes of lice (Phthiraptera: Insecta). Insect Molecular Biology, 2006, 15: 63-68.
192. Shao R, Campbell NJH, Barker SC. Numerous gene rearrangements in the mitochondrial genome of the wallaby louse, Heterodoxus macropus (Phthiraptera). Molecular biology and evolution, 2001, 18: 858-865.
193. 魏书军.膜翅目线粒体基因组的特征与进化及其在系统发育研究中的应用.博士毕业论文,2009,杭州：浙江大学.
194. Fearnley IM, Walker JE. Two overlapping genes in bovine mitochondrial DNA encode membrane components of ATP synthase. The EMBO Journal, 1986, 5: 2003-2008.
195. Ojala D, Montoya J, Attardi G. tRNA punctuation model of RNA processing in human mitochondria. Nature, 1981, 290: 470-474.
196. Giege R, Juhling F, Putz J, Stadler P, Sauter C, Florentz C. Structure of transfer RNAs: similarity and variability. WIREs RNA, 2012, 3: 37-61.
197. Liu B, Liu Y, Motyka SA, Agbo EEC, Englund PT. Fellowship of the rings: the replication of kinetoplast DNA. TRENDS in Parasitology, 2005, 21: 363-369.
198. Lukes J, Guilbride DL, Votypka J, Zikova A, Benne R, Englund PT. Kinetoplast DNA network: evolution of an improbable structure. Eukaryotic Cell, 2002, 1: 495-502.
199. Bogenhagen DF. Mitochondrial DNA nucleoid structure. Biochimica et Biophysica Acta, 2011: doi:10.1016/j.bbagrm.2011.11.005.
200. Johnson KP, Whiting MF. Multiple genes and the monophyly of Ischnocera. Molecular Phylogenetics and Evolution, 2002, 22: 101-110.
201. Lavrov DV. Key transitions in animal evolution: a mitochondrial DNA perspective. Integrative and Comparative Biology, 2007, 47: 734-743.
202. Helfenbein K, Fourcade H, Vanjani R, Boore J. The mitochondrial genome of Paraspadella gotoi is highly reduced and reveals that chaetognaths are a sister group to protostomes. Proceedings of the National Academy of Sciences of the United States of America,2004, 101:10639-10643.
203. Domes K, Maraun M, Scheu S, Cameron SL. The complete mitochondrial genome of the sexual oribatid mite Steganacarus magnus:genome rearrangements and loss of tRNAs. BMC Genomics,2008,9:532.
204. Hayashi JL, Ohta S, Kikuchi A, Takemitsu M, Goto Y, Nonaka I. Introduction of disease-related mitochondrial DNA deletions into HeLa cells lacking mitochondrial DNA results in mitochondrial dysfunction. Proceedings of the National Academy of Sciences of the United States of America,1991,88:10614-10618.
205. Kajander OA, Rovio AT, Majamaa K, Poulton J, Spelbrink JN, Holt IJ, Karhunen PJ, Jacobs HT. Human mtDNA sublimons resemble rearranged mitochondrial genoms found in pathological states. Human Molecular Genetics,2000,9:2821-2835.
206. Melov S, Hinerfeld D, Esposito L, Wallace DC. Multi-organ characterization of mitochondrial genomic rearrangements in ad libitum and caloric restricted mice show striking somatic mitochondrial DNA rearrangements with age. Nucleic Acids Research, 1997,25:974-982.
207. Palmer JD, Shields CR. Tripartite structure of the Brassica campestris mitochondrial genome. Nature,1984,307:437-440.
208. Wu Y, Li ZH, Wu JJ. Polymorphic microsatellite markers in the Melon Fruit Fly, Bactrocera cucurbitae (Coquillett) (Diptera:Tephritidae). Molecular Ecology Resources,2009,9: 1404-1406.
209. Bloor PA, Barker FS, Watts PC, Noyes HA, Kemp SJ. Microsatellite libraries by enrichment. 2001:http://www.genomics.liv.ac.uk/animal/Protocoll.html.
210. Weber JL. Informativeness of human (dC-dA)n · (dG-dT)n polymorphisms. Genomics, 1990,7:524-530.
211. Rassmann K, Schlotterer C, Tautz D. Isolation of simple sequence loci for use in polymerase chain reaction based DNA fingerprinting. Electrophoresis,1991,12:113-118.
212. Fagerberg AJ, Fulton RE, IV WCB. Microsatellite loci are not abundant in all arthropod genomes:analyses in the hard tick, Ixodes scapularis and the yellow fever mosquito, Aedes aegypt. Insect Molecular Biology,2001,10:225-236.
213. Steiner FM, Arthofer W, Schlick-Steiner BC, Crozier RH, Stauffer C. Twenty four new microsatellite markers in two invasive pavement ants, Tetramorium sp. E and T. tsushimae (Hymenoptera:Formicidae). Conservation Genetics,2008,9:757-759.
214. 樊勇,马雅军.中华按蚊多态微卫星DNA位点的筛选和特征研究.第四军医大学学报,2008,29：1537-1539.
215. 张国彦,翟保平.东方粘虫(Pseudaletia separata (Walker))微卫星富集文库的构建与分析.生态学报,2008,28：3860-3867.
216. Santana QC, Coetzee MPA, Steenkamp ET, Mlonyeni OX, Hammond GNA, Wingfield MJ, Wingfield BD. Microsatellite discovery by deep sequencing of enriched genomic libraries. Biotechniques,2009,46:217-223.
217. Zhang DX. Lepidopteran microsatellite DNA:redundant but promising. TRENDS in Ecology and Evolution,2004,19:507-509.
218. Katti MV, Ranjekar PK, Gupta VS. Differential distribution of simple sequence repeats in eukaryotic genome sequences. Molecular Biology and Evolution,2001,18:1161-1167.
219. Toth G, Gaspari Z, Jurka J. Microsatellites in different eukaryotic genomes:survey and analysis. Genome Research,2000,10:967-981.
2. 李法圣,李志红,彩万志,杨定.中国啮目昆虫分类研究.中国农业大学学报,2008,13：7-18.
3. 赵朔,李志红,秦萌.书虱及其分子生物学研究进展.植物保护,2009,35：17-21.
4. 程伟霞,王进军.嗜卷书虱和嗜虫书虱抗性及防治方法的研究概况.安徽农业科学,2008,36：6376-6379.
5. Nayak MK, Collins PJ, Pavic H. Long-term effectiveness of grain protectants and structural treatments against Liposcelis decolor Pearman Psocoptera:Liposcelididae, a pest of stored products. Pest Management Science,2002,58:1223-1228.
6. Ding W, Wang JJ, Zhao ZM, Tsai JH. Effects of controlled atmosphere and DDVP on population growth and resistance development by the psocid, Liposcelis bostrychophila Badonnel (Psocoptera:Liposcelididae). Journal of Stored Products Research,2002,38: 229-237.
7. Wang JJ, Zhao ZM, Li LS. Selection of resistance strains in Liposcelis bostrychophila Badonnel to CO2-enriched atmospheres. Entomologia Sinica,1999,6:45-52.
8. Nayak MK. Potential of piperonyl butoxide-synergised pyrethrins against psocids (Psocoptera:Liposcelididae) for stored-grain protection. Pest Management Science,2010,66: 295-300.
9. Behar A, McCormick LJ, Perlman SJ. Rickettsia felis infection in a common household insect pest, Liposcelis bostrychophila (Psocoptera:Liposcelidae). Applied and Environmental Microbiology,2010,76:2280-2285.
10. Lin Y, Chan M, Ko C, Hsieh M. Nail infestation by Liposcelis bostrychophila badonnel. Clinical and Experiment Dermatology,2004,29:620-621.
11. Ahmedani MS, Shagufta N, Aslam M, Hussnain SA. Psocid:A new risk for global food security and safety. Applied Entomology and Zoology,2010,45:89-100.
12. Yoshizawa K, Johnson KP. How stable is the "Polyphyly of Lice" hypothesis (Insecta: Psocodea)?:A comparison of phylogenetic signal in multiple genes. Molecular Phylogenetics and Evolution,2010,55:939-951.
13. Ishiwata K, Sasaki G, Ogawa J, Miyata T, Su ZH. Phylogenetic relationships among insect orders based on three nuclear protein-coding gene sequences. Molecular Phylogenetics and Evolution,2010,58:169-180.
14. Yoshizawa K, Lienhard C, Johnson KP. Molecular systematics of the suborder Trogiomorpha (Insecta:Psocodea:'Psocoptera'). Zoological Journal of the Linnean Society, 2006,146:287-299.
15. Lyal CHC. Phylogeny and classification of the Psocodea, with particular reference to the lice (Psocodea:Phthiraptera). Systematic Entomology,1985,10:145-165.
16. Grimaldi D, Engel MS. Fossil Liposcelididae and the lice ages (Insecta:Psocodea). Proceedings of the Royal Society B-Biological Sciences,2006,273:625-633.
17. Leong ECW, Ho S. Life cycle of Liposcelis entomophila (Psocoptera:Liposcelididae) and a culturing regime for liposcelids. Bulletin of Entomological Research,1995,85:501-506.
18. 王进军,赵志模,郭依泉,周亦红,周晨曦.温度、湿度对嗜虫书虱生长发育和繁殖的影响.植物保护学报,1996,23：147-151.
19. Wang JJ, Ren Y, Wei XQ, Dou W. Development, survival, and reproduction of the psocid Liposcelis paeta (Psocoptera:Liposcelididae) as a function of temperature. Journal of Economic Entomology,2009,102:1705-1713.
20. Green PWC. Insect-derived compounds affect the behaviour of Liposcelis bostrychophila: Effects of combination and structure. Journal of Stored Products Research,2011,47: 262-266.
21. Xiao LS, Dou W, Li Y, Wang JJ. Comparative studies of acetylcholinesterase purified from three field populations of Liposcelis entomophila (enderlein) (psocoptera:liposcelididae). Archives of Insect Biochemistry and Physiology,2010,75:158-173.
22. Dou W, Xiao LS, Niu JZ, Jiang HB, Wang JJ. Characterization of the purified glutathione S-transferases from two psocids Liposcelis bostiychophila and L. entomophila. Agricultural Sciences in China,2010,9:1008-1016.
23. Ren Y, Wei XQ, Wu SA, Dou W, Wang JJ. Comparison of acetylcholinesterase from three field populations of Liposcelis paeta Pearman (Psocoptera:Liposcelididae):Implications of insecticide resistance. Pesticide Biochemistry and Physiology,2008,90:196-202.
24. Cheng WX, Dou W, Chai YX, Wang JJ. Comparison of biochemical and toxicological characterizations of glutathione S-transferases and superoxide dismutase between Liposcelis bostrychophila Badonnel and L. entomophila (Enderlein) (Psocoptera:Liposcelididae). Pesticide Biochemistry and Physiology,2007,89:151-157.
25. Wu S, Li M, Tang PA, Felton GW, Wang JJ. Cloning and characterization of acetylcholinesterase 1 genes from insecticide-resistant field populations of Liposcelis paeta Pearman (Psocoptera:Liposcelididae). Insect Biochemistry and Molecular Biology,2010, 40:415-424.
26. Jiang HB, Tang PA, Xu YQ, An FM, Wang JJ. Molecular characterization of two novel deltamethrin-inducible P450 genes from Liposcelis bostrychophila Badonnel (Psocoptera: Liposcelididae). Archives of Insect Biochemistry and Physiology,2010,74:17-37.
27. Tang PA, Jiang HB, Xu YQ, An FM, Wang JJ. Molecular characterization of two nicotinic acetylcholine receptor subunits from Liposcelis bostrychophila Badonnel (Psocoptera: Liposcelididae). Archives of Insect Biochemistry and Physiology,2009,72:34-47.
28. Mikac KM, FitzSimmons NN. Genetic structure and dispersal patterns of the invasive psocid Liposcelis decolor (Pearman) in Australian grain storage systems. Bulletin of Entomological Research,2010,100:521-527.
29. Mikac KM, Clarke GM. Tracing the geographic origin of the cosmopolitan parthenogenetic insect pest Liposcelis bostrychophila (Psocoptera:Liposcelididae). Bulletin of Entomological Research,2006,96:523-530.
30. 丁伟,李隆术,赵志模.书虱综合防治技术研究进展.粮食储藏,2001,30：3-6.
31. 周世良,张方,王中仁.等位酶淀粉凝胶电泳技术中的几个应引起重视的问题.植物学通报,1998,15：68-72.
32. 王进军.嗜卷书虱种群生态及其对气调抗性的机理研究.博士学位论文,1997,重庆：西南农业大学.
33. Ali N, Turner B. Allozyme polymorphism and variability in permethrin tolerance in British population of the parthenogenetic stored product pest Liposcelis bostrychophila (Liposcelididae, Psocoptera). Journal of Stored Products Research,2001,37:111-125.
34. Mikac KM. PCR confirms multiple Wolbachia strain infection in Australian and international populations of the invasive stored-product psocid Liposcelis bostrychophila Badonnel. Journal of Stored Products Research,2007,43:594-597.
35. Qin M, Li ZH, Kucerova Z, Cao Y, Stejskal V. Rapid discrimination of the common species of the stored product pest Liposcelis (Psocoptera:Liposcelididae) from China and the Czech Republic, based on PCR-RFLP analysis. European Journal of Entomology,2008,105: 713-717.
36. Tautz D. Hypervariability of simple sequences as a general source for polymorphic DNA markers. Nucleic Acids Research,1989,17:6463-6471.
37. Selkoe KA, Toonen RJ. Microsatellites for ecologists:a practical guide to using and evaluating microsatellite markers. Ecology Letters,2006,9:615-629.
38. Mikac KM. Isolation and characterization of the first microsatellite loci from the order Psocoptera in the economically important pest insect Liposcelis decolor (Pearman) and cross-species amplification. Molecular Ecology Resources,2006,6:1102-1104.
39. 魏丹丹,袁明龙,王保军,朱礼明,王进军.两种书虱微卫星富集文库的构建及比较.生态学报,2011,31：4182-4189.
40. Behura SK. Molecular marker systems in insects:current trends and future avenues. Molecular Ecology,2006,15:3087-3113.
41. Yoshizawa K, Johnson KP. Phylogenetic position of Phthiraptera (Insecta:Paraneoptera) and elevated rate of evolution in mitochondrial 12S and 16S rDNA. Molecular Phylogenetics and Evolution,2003,29:102-114.
42. Johnson K, Mockford EL. Molecular systematics of Psocomorpha (Psocoptera). Systematic Entomology,2003,28:409-416.
43. Yoshizawa K. Molecular phylogeny of major lineages of Trichadenotecnum and a review of diagnostic morphological characters (Psocoptera:Psocidae). Systematic Entomology,2004, 29:383-394.
44. Yoshizawa K, Johnson KP. Molecular systematics of the barklouse family Psocidae (Insecta: Psocodea:'Psocoptera') and implications for morphological and behavioral evolution. Molecular phylogenetics and evolution,2008,46:547-559.
45. Wei DD, Yuan ML, Wang BJ, Zhou AW, Dou W, Wang JJ. Population genetics of two asexually and sexually reproducing psocids species inferred by the analysis of mitochondrial and nuclear DNA sequences. PLoS ONE,2012,7:e33883.
46. Li ZH, Kucerova Z, Zhao S, Stejskal V, Opit G, Qin M. Morphological and molecular identification of three geographical populations of the storage pest Liposcelis bostrychophila (Psocoptera). Journal of Stored Products Research,2011,47:168-172.
47. Arif M, Ochoa-Corona FM, Opit GP, Li ZH, Kucerova Z, Stejskal V, Yang QQ. PCR and isothermal-based molecular identification of the stored-product psocid pest Lepinotus reticulatus (Psocoptera:Trogiidae). Journal of Stored Products Research,2012,49:184-188.
48. Yang QQ, Kucerova Z, Li ZH, Kalinovic I, Stejskal V, Opit G, Cao Y. Diagnosis of Liposcelis entomophila (Insecta:Psocodea:Liposcelididae) based on morphological characteristics and DNA barcodes. Journal of Stored Products Research,2012,48:120-125.
49. Yuan ML, Wang BJ, Lu F, Hu CX, Wei DD, Dou W, Wang JJ. Evaluation of genetic diversity and population structure of Panonychus citri (Acari:Tetranychidae) in China using ribosomal internal transcribed spacer 1 sequences. Annals of the Entomological Society of America,2011,104:800-807.
50. Wei DD, Yuan ML, Wang ZY, Wang D, Wang BJ, Dou W, Wang JJ. Sequence analysis of the ribosomal internal transcribed spacers region in psocids (Psocoptera:Liposcelididae) for phylogenetic inference and species discrimination. Journal of Economic Entomology,2011, 104:1720-1729.
51. Lang BF, Gray MW, Burger G. Mitochondrial genome evolution and the origin of eukaryotes. Annual Review of Genetics,1999,33:351-397.
52. Papillon D, Perez Y, Caubit X, Parco YL. Identification of chaetognaths as protostomes is supported by the analysis of their mitochondrial genome. Molecular Biology and Evolution, 2004,21:2122-2129.
53. Gissi C, Iannelli F, Pesole G. Evolution of the mitochondrial genome of Metazoa as exemplified by comparison of congeneric species. Heredity,2008,101:301-320.
54. Dermauw W, Vanholme B, Tirry L, Leeuwen TV. Mitochondrial genome analysis of the predatory mite Phytoseiulus persimilis and a revisit of the Metaseiulus occidentalis mitochondrial genome. Genome,2010,53:285-301.
55. Hu M, Chilton NB, Gasser RB. The mitochondrial genome of Strongyloides stercoralis (Nematoda)-idiosyncratic gene order and evolutionary implications. International Journal for Parasitology,2003,33:1393-1408.
56. Burger G, Gray MW, Lang BF. Mitochondrial genomes:anything goes. TRENDS in Genetics,2003,19:709-716.
57. Yuan ML, Wei DD, Wang BJ, Dou W, Wang JJ. The complete mitochondrial genome of the citrus red mite Panonychus citri (Acari:Tetranychidae):high genome rearrangement and extremely truncated tRNAs. BMC Genomics,2010,11:597.
58. Shao R, Mitani H, Barker SC, Takahashi M, Fukunaga M. Novel mitochondrial gene content and gene arrangement indicate illegitimate inter-mtDNA recombination in the chigger mite, Leptotrombidium pallidum. Journal of Molecular Evolution,2005,60: 764-773.
59. Shao R, Barker SC, Mitani H, Aoki Y, Fukunaga M. Evolution of duplicate control regions in the mitochondrial genomes of metazoa:a case study with Australasian Ixodes ticks. Molecular Biology and Evolution,2005,22:620-629.
60. Cameron SL, Yoshizawa K, Mizukoshi A, Whiting MF, Johnson KP. Mitochondrial genome deletions and minicircles are common in lice (Insecta:Phthiraptera). BMC Genomics,2011, 12:394.
61. Watanabe KI, Bessho Y, Kawasaki M, Hori H. Mitochondrial genes are found on minicircle DNA molecules in the mesozoan animal Dicyema. Journal of Molecular Biology,1999,286: 645-650.
62. Awata H, Noto T, Endoh H. Differentiation of somatic mitochondria and the structural changes in mtDNA during development of the dicyemid Dicyema japonicum (Mesozoa). Molecular Genetics and Genomics,2005,273:441-449.
63. Suga K, Welch DBM, Tanaka Y, Sakakura Y, Hagiwarak A. Two circular chromosomes of unequal copy number make up the mitochondrial genome of the rotifer Brachionus plicatilis. Molecular Biology and Evolution,2008,25:1129-1137.
64. Armstrong MR, Blok VC, Phillips MS. A multipartite mitochondrial genome in the potato cyst nematode Globodera pallida. Genetics,2000,154:181-192.
65. Gibson T, Blok VC, Dowton M. Sequence and characterization of six mitochondrial subgenomes from Globodera rostochiensis:multipartite structure is conserved among close nematode relatives. Journal of Molecular Evolution,2007,65:308-315.
66. Sugiyama Y, Watase Y, Nagase M, Makita N, Yagura S, Hirai A, Sugiura M. The complete nucleotide sequence and multipartite organization of the tobacco mitochondrial genome: comparative analysis of mitochondrial genomes in higher plants. Molecular Genetics and Genomics,2005,272:603-615.
67. Burger G, Lang B F. Parallels in genome evolution in mitochondria and bacterial symbionts. IUBMB Life,2003,55:205-212.
68. Voigt O, Erpenbeck D, Worheide G. A fragmented metazoan organellar genome:the two mitochondrial chromosomes of Hydra magnipapillata. BMC Genomics,2008,9:350.
69. Fan J, Lee RW. Mitochondrial genome of the colorless green alga Polvtomella parva:two linear DNA molecules with homologous inverted repeat termini. Molecular Biology and Evolution,2002,19:999-1007.
70. Nosek J, Tomaska L, Fukuhara H, Suyama Y, Kovac L. Linear mitochondrial genomes:30 years down the line. TRENDS in Genetics,1998,14:184-188.
71. Marande W, Lukes J, Burger G. Unique mitochondrial genome structure in diplonemids, the sister group of kinetoplastids. Eukaryotic Cell,2005,4:1137-1146.
72. Gibson T, Blok VC, Phillips MS, Hong G, Kumarasinghe D, Riley IT, Dowton M. The mitochondrial subgenomes of the nematode Globodera pallida are mosaics:evidence of recombination in an animal mitochondrial genome. Journal of Molecular Evolution,2007, 64:463-471.
73. Shao R, Kirkness EF, Barker SC. The single mitochondrial chromosome typical of animals has evolved into 18 minichromosomes in the human body louse, Pediculus humanus. Genome Research,2009,19:904-912.
74. Melov S, Lithgow GJ, Fischer DR, Tedesco PM, Johnson TE. Increased frequency of deletions in the mitochondrial genome with age of Caenorhabditis elegans. Nucleic Acids Research,1995,23:1419-1425.
75. Fauron C, Casper M, Gao Y, Moore B. The maize mitochondrial genome:dynamic, yet functional. Trends in Genetics,1995,11:228-235.
76. Shao R, Barker SC. Chimeric mitochondrial minichromosomes of the human body louse, Pediculus humanus:Evidence for homologous and non-homologous recombination. Gene, 2011,473:36-43.
77. Dowton M, Austin AD. Evolutionary dynamics of a mitochondrial rearrangement "Hot Spot" in the Hymenoptera. Molecular Biology and Evolution,1999,16:298-309.
78. Mueller R, Boore J. Molecular mechanisms of extensive mitochondrial gene rearrangement in plethodontid salamanders. Molecular Biology and Evolution,2005,22:2104-2112.
79. Wei DD, Shao R, Yuan ML, Dou W, Barker SC, Wang JJ. The multipartite mitochondrial genome of Liposcelis bostiychophila:insights into the evolution of mitochondrial genomes in bilateral animals. PLoS ONE,2012,7:e33973.
80. Landweber LF. Why genomes in pieces? Science,2007,318:405-407.
81. Rand DM.'Why genomes in pieces?' revisited:Sucking lice do their own thing in mtDNA circle game. Genome Research,2009,19:700-702.
82. Kirkness E, Haas B, Sun W, Braig H, Perotti M, Clark J, Lee S, Robertson H, Kennedy R, Elhaik E, Gerlach D, Kriventseva E, Elsik C, Graur D, Hill C, Veenstra J, Walenz B, Tubio J, Ribeiro J, Rozas J, Johnston J, Reese J, Popadic A, Tojo M, Raoult D, Reed D, Tomoyasu Y, Krause E, Mittapalli O, Margam V, Li H, Meyer J, Johnson R, Romero-Severson J, Vanzee J, Alvarez-Ponce D, Vieira F, Aguade M, Guirao-Rico S, Anzola J, Yoon K, Strycharz J, Unger M, Christley S, Lobo N, Seufferheld M, Wang N, Dasch G, Struchiner C, Madey G, Hannick L, Bidwell S, Joardar V, Caler E, Shao R, Barker S, Cameron S, Bruggner R, Regier A, Johnson J, Viswanathan L, Utterback T, Sutton G, Lawson D, Waterhouse R, Venter J, Strausberg R, Berenbaum M, Collins F, Zdobnov E, Pittendrigh B. Genome sequences of the human body louse and its primary endosymbiont provide insights into the permanent parasitic lifestyle. Proceedings of the National Academy of Sciences of the United States of America,2010,107:12168-12173.
83. Korhonen JA, Pham XH, Pellegrini M, Falkenberg M. Reconstitution of a minimal mtDNA replisome in vitro. EMBO Journal,2004,23:2423-2429.
84. Johnson KP, Yoshizawa K, Smith VS. Multiple origins of parasitism in lice. Proceedings of the Royal Society B-Biological Sciences,2004,271:1771-1776.
85. Mockford EL. Some Psocoptera from the plumage of birds. Proceedings of the Entomological Society of Washington,1967,69:307-309.
86. Pearlman JV. Some African Psocoptera found on rats. Entomologist,1960,93:246-250.
87. Lienhard C. Revision of the western palaearctic species of Liposcelis Motschulsky (Psocoptera:Liposcelididae). Zoologische Jahrbucher. Abteilung fur Systematik, Okologie und Geographie der Tiere,1990,117:117-174.
88. Li YC, Korol AB, Fahima T, Beiles A, Nevo E. Microsatellites:genomie distribution, putative functions and mutational mechanisms:a review. Molecular Ecology,2002,11: 2453-2465.
89. Zane L, Bargelloni L, Patarnello T. Strategies for microsatellite isolation:a review. Molecular Ecology,2002,11:1-16.
90. Yu J, Won C, Jun J, Lim Y, Kwak M. Fast and cost-effective mining of microsatellite markers using NGS technology:an example of a Korean water deer Hydropotes inermis argyropus. PLoS ONE,2011,6:e26933.
91. Archak S, Meduri E, Sravana KP, Nagaraju J. InSatDb:A microsatellite database of fully sequenced insect genomes. Nucleic Acids Research,2007,35:D36-D39.
92. Aishwarya V, Grover A, Sharma PC. EuMicroSatdb:A database for microsatellites in the sequenced genomes of eukaryotes. BMC Genomics,2007,8:225.
93. Chen MH, Dorn S. Cross-amplification of microsatellites from the codling moth Cydia pomonella to three other species of the tribe Grapholitini (Lepidoptera:Tortricidae). Molecular Ecology Resources,2010,10:1034-1037.
94. Ma H, Chen S. Development of polymorphic microsatellite markers in barfin flounder (Verasper moseri) and spotted halibut(Verasper variegatus) by the cross-species amplification. Molecular Biology Reports,2010,38:4545-4551.
95. Barbara T, Palma-Silva C, Paggi G, Bered F, Fay M, Lexer C. Cross-species transfer of nuclear microsatellite markers:potential and limitations. Molecular Ecology,2007,16: 3759-3767.
96. Merkel A, Gemmell NJ. Detecting microsatellites in genome data:variance in definitions and bioinformatic approaches cause systematic bias. Evolutionary Bioinformatics,2008,4: 1-6.
97. Durand J, Bodenes C, Chancerel E, Frigerio JM, Vendramin G, Sebastiani F, Buonamici A, Gailing O, Koelewijn HP, Villani F, Mattioni C, Cherubini M, Goikoetxea P, Herran A, Ikaran Z, Cabane C, Ueno S, Alberto F, Dum PY, Guichoux E, Daruvar A, Kremer A, Plomion C. A fast and cost-effective approach to develop and map EST-SSR markers:oak as a case study. BMC Genomics,2010,11:570.
98. 张琳琳,魏朝明,廉振民,孔光耀.赤拟谷盗全基因组和EST中微卫星的丰度.昆虫知识,2008,45：38-42.
99. 刘玉娣,侯茂林.褐飞虱EST资源的微卫星信息分析.昆虫学报,2010,53：239-247.
100. Mikheyev AS, Tanya Vol BW, Singer MC, Parmesan C. Rapid microsatellite isolation from a butterfly by de novo transcriptome sequencing:performance and a comparison with AFLP-derived distances. PLoS ONE,2010,5:e11212.
101. 孙波,鲍毅新,赵庆洋,张龙龙,胡知渊.微卫星位点获取方法的研究进展.生态学杂志,2009,28：2130-2137.
102. Andres J, Bogdanowicz S. Isolating microsatellite Loci:looking back, looking ahead. Methods in Molecular Biology,2011,772:211-32.
103. Glenn TC, Schable NA. Isolating microsatelline DNA loci. Methods in Enzymology,2005, 395:202-222.
104. Kandpal RP, Kandpal G, Weissman SM. Construction of libraries enriched for sequence repeats and jumping clones, and hybridization selection for region specific markers. Proceedings of the National Academy of Sciences of the United States of America,1994,91: 88-92.
105. Techen N, Arias RS, Glynn NC, Pan Z, Khan IA, Scheffler BE. Optimized construction of microsatellite-enriched libraries. Molecular Ecology Resources,2010,10:508-515.
106. 邓欣,陈信波,龙松华,王孝纯,高原,何东锋,王进,王玉富.用磁珠富集法分离亚麻基因组微卫星分子标记.作物学报,2008,34：2100-2105.
107. Fisher P, Gardner R, Richardson T. Single locus microsatellites isolated using 5'anchored PCR. Nucleic Acids Research,1996,24:4369-4371.
108. Hayden MJ, Sharp PJ. Targeted development of informative microsatellite (SSR) markers. Nucletic Acids Research,2001,29:e44.
109. Shao R, Dowton M, Murrell A, Barker SC. Rates of gene rearrangement and nucleotide substitution are correlated in the mitochondrial genomes of insects. Molecular Biology and Evolution,2003,20:1612-1619.
110. Leeuwen TV, Vanholme B, Pottelberge SV, Nieuwenhuyse PV, Nauen R, Tirry L, Denholm I. Mitochondrial heteroplasmy and the evolution of insecticide resistance:Non-Mendelian inheritance in action. Proceedings of the National Academy of Sciences of the United States of America,2008,105:5980-5985.
111. 李法圣,中国啮目志.2002,北京：科学出版社.
112. Wang ZY, Wang JJ, Lienhard C. Two new species of Liposcelis (Psocoptera, Liposcelididae). Acta Zootaxonomica Sinica,2006,31:564-568.
113. Tautz D, Arctander P, Minelli A, Thomas RH, Vogler AP. DNA points the way ahead in taxonomy. Nature,2002,418:479.
114. Hebert PDN, Cywinska A, Ball SL, deWaard JR. Biological identifications through DNA barcodes. Proceedings of the Royal Society B-Biological Sciences,2003,270:313-321.
115. Hebert PDN, Ratnasingham S, de Waard JR. Barcoding animal life:cytochrome c oxidase subunit 1 divergences among closely related species. Proceedings of the Royal Society B-Biological Sciences,2003,270:S96-S99.
116. Galtier N, Nabholz B, Glemin S, Hurst GD. Mitochondrial DNA as a marker of molecular diversity:a reappraisal. Molecular Ecology,2009,18:4541-4550.
117. Yao H, Song J, Liu C, Luo K, Han J, Li Y, Pang X, Xu H, Zhu Y, Xiao P, Chen S. Use of ITS2 region as the universal DNA barcode for plants and animals. PLoS ONE,2010,5: e13102.
118. Li DZ, Gao LM, Li HT, Wang H, Ge XJ, Liu JQ, Chen ZD, Zhou SL, Chen SL, Yang JB, Fu CX, Zeng CX, Yan HF, Zhu YJ, Sun YS, Chen SY, Zhao L, Wang K, Yang T, Duan GW. Comparative analysis of a large dataset indicates that internal transcribed spacer (ITS) should be incorporated into the core barcode for seed plants. Proceedings of the National Academy of Sciences of the United States of America,2011,108:19641-19646.
119. Matsumoto Y, Yanase T, Tsuda T, Noda H. Characterization of internal transcribed spacer (ITS1)-ITS2 region of ribosomal RNA gene from 25 species of Culicoides biting midges (Diptera:Ceratopogonidae) in Japan. Journal of Medical Entomology,2009,46:1099-1108.
120. Oh HK, Yoon HJ, Kim MJ, Jeong HU, Kim SR, Hwang JS, Bae CH, Kim I. ITS2 ribosomal DNA sequence variation of the bumblebee, Bombus ardens (Hymenoptera:Apidae). Genes & Genomics,2009,31:293-303.
121. Wiemer M, Keller A, Wolf M. ITS2 secondary structure improves phylogeny estimation in a radiation of blue butterflies of the subgenus Agrodiaetus (Lepidoptera:Lycaenidae: Polyommatus). BMC Evolutionary Biology,2009,9:300.
122. Stephan A, Clausen PH, Bauer B, Steuber S. PCR identification of Culicoides dewulfi midges (Diptera:Ceratopogonidae), potential vectors of bluetongue in Germany. Parasitology Research,2009,105:367-371.
123. Rugman-Jones PF, Hoddle MS, Mound LA, Stouthamer R. Molecular identification key for pest species of Scirtothrips (Thysanoptera:Thripidae). Journal of Economic Entomology, 2006,99:1813-1819.
124. Farris RE, Ruiz-Arce R, Ciomperlik M, Vasquez JD, DeLeon R. Development of a ribosomal DNA ITS2 marker for the identification of the thrips, Scirtothrips dorsalis. Journal of Insect Science,2010,10:26.
125. Kent RJ, Deus S, Williams M, Savage HM. Development of a multiplexed polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) assay to identify common members of the Subgenera Culex(Culex) and Culex(Phenacomyia) in Guatemala. The American Journal of Tropical Medicine and Hygiene,2010,83:285-291.
126. Jia FX, Yang MS, Yang WJ, Wang JJ. Influence of continuous high temperature conditions on Wolbachia infection frequency and the fitness of Liposcelis tricolor (Psocoptera: Liposcelididae). Environmental Entomology,2009,38:1365-1372.
127. Keller A, Schleicher T, Schultz J, Muller T, Dandekar T, Wolf M.5.8S-28S rRNA interaction and HMM-based ITS2 annotation. Gene,2009,430:50-57.
128. Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG. The ClustalX windows interface:flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research,1997,24:4876-4882.
129. Librado P, Rozas J. DnaSP v5:a software for comprehensive analysis of DNA polymorphism data. Bioinformatics,2009,25:1451-1452.
130. Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S. MEGA5:molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution,2011,28:2731-2739.
131. Castresana J. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution,2000,17:540-552.
132. Posada D. jModelTest:phylogenetic model averaging. Molecular Biology and Evolution, 2008,25:1253-1256.
133. Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O. New algorithms and methods to estimate maximum-likelihood phylogenies:assessing the performance of PhyML 3.0. Systematic Biology,2010,59:307-321.
134. Page RDM. TreeView:an application to display phylogenetic trees on personal computers. Computer Applications in The Biosciences,1996,12:357-358.
135. Ullrich B, Reinhold K, Niehuis O, Misof B. Secondary structure and phylogenetic analysis of the internal transcribed spacers 1 and 2 of bush crickets (Orthoptera:Tettigoniidae: Barbitistini). Journal of Zoological Systematics and Evolutionary Research,2010,48: 219-228.
136. Kengne P, Antonio-Nkondjio C, Awono-Ambene HP, Simard F, Awolola TS, Fontenille D. Molecular differentiation of three closely related members of the mosquito species complex, Anopheles moucheti, by mitochondrial and ribosomal DNA polymorphism. Medical and Veterinary Entomology,2007,21:177-182.
137. Li JJ, Ledger TW, Tredici PD. Phylogenetics of Calycanthaceae based on molecular and morphological data with special reference to divergent paralogues of the nrDNA ITS region. Harvard Papers in Botany,2004,9:69-82.
138. Harpke D, Peterson A.5.8S motifs for the identification of pseudogenic ITS regions. Botany, 2008,86:300-305.
139. David J, Huber K, Failloux A, Rey D, Meyran J. The role of environment in shaping the genetic diversity of the subalpine mosquito, Aedes rusticus (Diptera, Culicidae). Molecular Ecology,2003,12:1951-1961.
140. Amos W, Harwood J. Factors affecting levels of genetic diversity in natural populations. Philosophical Transactions of the Royal Society B-Biological Sciences,1998,353:177-186.
141. Markert JA, Champlin DM, Gutjahr-Gobell R, Grear JS, Kuhn A, Jr TJM, Roth A, Bagley MJ, Nacci DE. Population genetic diversity and fitness in multiple environments. BMC Evolutionary Biology,2010,10:205.
142. Rua N, Stevens L, Dorn PL. High genetic diversity in a single population of Triatoma sanguisuga (LeConte,1855) inferred from two mitochondrial markers:cytochrome b and 16S ribosomal DNA. Infection, Genetics and Evolution,2011,11:671-677.
143. Collins F, Paskewitz S. A review of the use of ribosomal DNA (rDNA) to differentiate among cryptic Anopheles species. Insect Molecular Biology,1996,5:1-9.
144. Suomalainen E, Saura A, Lokki J, Teeri T. Genetic polymorphism and evolution in parthenogenetic animals. Theoretical and Applied Genetics,1980,57:129-132.
145. Weeks AR. Marec F, Breeuwer JAJ. A mite species that consists entirely of haploid females. Science,2001,292:2479-2482.
146. Simon JC, Delmotte F, Rispe C, Crease T. Phylogenetic relationships between parthenogens and their sexual relatives:the possible routes to parthenogenesis in animals. Biological Journal of the Linnean Society,2003,79:151-163.
147. Bengtsson BO. Genetic variation in organisms with sexual and asexual reproduction. Journal of Evolutionary Biology,2003,16:189-199.
148. Shreve S, Mockford E, Johnson K. Elevated genetic diversity of mitochondrial genes in asexual populations of bark lice ("Psocoptera":Echmepteryx hageni). Molecular Ecology, 2011,20:4433-4451.
149. Jiang HB, Liu YH, Tang PA, Zhou AW, Wang JJ. Validation of endogenous reference genes for insecticide-induced and developmental expression profiling of Liposcelis bostrychophila (Psocoptera:Liposcelididae). Molecular Biology Reports,2010,37:1019-1029.
150. Guedes RNC, Zhu KY, Opit GP, Throne JE. Differential heat shock tolerance and expression of heat-inducible proteins in two stored-product psocids. Journal of Economic Entomology,2008,101:1974-1982.
151. Simon C, Frati F, Beckenbach A, Crespi B, Liu H, Flook P. Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America,1994, 87:651-701.
152. Hall TA. BIOEDIT:a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series,1999,41:95-98.
153. Zuker M. Mfold web server for nucleic acid folding and hybridization prediction. Nucleic Acids Research,2003,31:3406-3415.
154. Wuyts H, De Rijk P, Van de Peer Y, Winkelmans T, De Wachter R. The European large subunit ribosomal RNA database. Nucleic Acids Research,2001,29:175-177.
155. Excoffier L, Laval G, Schneider S. Arlequin (version 3.0):an integrated software package for population genetics data analysis. Evolution Bioinformatics Online,2005,1:47-50.
156. Dupanloup I, Schneider S, Excoffier L. A simulated annealing approach to define the genetic structure of populations. Molecular Ecology,2002,11:2571-2581.
157. Jensen JL, Bohonak AJ, Kelley ST. Isolation by distance, web service. BMC Genetics,2005, 6:13.
158. Tajima F. Statistical method for testing the neutral mutation hypothesis by DNA polymorphism. Genetics,1989,123:585-595.
159. Fu YX. Statistical tests of neutrality of mutations against population growth, hitchhiking and background selection. Genetics,1997,147:915-925.
160. Rogers AR, Harpending H. Population growth makes waves in the distribution of pairwise genetic differences. Molecuar Biology and Evolution,1992,9:552-569.
161. Dsouli-Aymes N, Michaux J, Stordeur ED, Couloux A, Veuille M, Duvallet G. Global population structure of the stable fly(Stomoxys calcitrans) inferred by mitochondrial and nuclear sequence data. Infection, Genetics and Evolution,2011,11:334-342.
162. Xia X, Xie Z. DAMBE:software package for data analysis in molecular biology and evolution. The Journal of Heredity,2001,92:371-373.
163. Ronquist F, Huelsenbeck JP. MrBayes 3:Bayesian phylogenetic inference under mixed models. Bioinformatics,2003,19:1572-1574.
164. Huelsenbeck JP, Ronquist F, Nielsen R, Bollback JP. Bayesian inference of phylogeny and its impact on evolutionary biology. Science,2001,294:2310-2314.
165. Posada D, Crandall K. Intraspecific gene genealogies:trees grafting into networks. TRENDS in Ecology and Evolution,2001,16:37-45.
166. Bandelt HJ, Forster P, Rohl A. Median-Joining networks for inferring intraspecific phylogenies. Molecular Biology and Evolution,1999,16:37-48.
167. Szalanski AL, Austin JW, McKern JA, Steelman CD, Gold RE. Mitochondrial and ribosomal internal transcribed spacer 1 diversity of Cimex lectularius (Hemiptera: Cimicidae). Journal of Medical Entomology,2008,45:229-236.
168. Loxdale HD. Rapid genetic changes in natural insect populations. Ecological Entomology, 2010,35:155-164.
169. Mockford EL, Krushelnycky PD. New species and records of Liposcelis Motschulsky (Psocoptera:Liposcelididae) from Hawaii with first description of the male of Liposcelis bostrychophila Badonnel. Zootaxa,2008,1766:53-68.
170. Wang JJ, Dong P, Xiao LS, Dou W. Effects of removal of Cardinium infection on fitness of the stored-product pest Liposcelis bostrychophila (Psocoptera:Liposcelididae). Journal of Economic Entomology,2008,101:1711-1717.
171. Yusuf M, Turner B, Whitfield P, Miles R, Pacey J. Electron microscopical evidence of a vertically transmitted Wolbachia-like parasite in the parthenogenetic, stored-product pest Liposcelis bostrychophila Badonnel (Psocoptera). Journal of Stored Products Research, 2000,36:169-175.
172. Yu MZ, Zhang KJ, Xue XF, Hong XY. Effects of Wolbachia on mtDNA variation and evolution in natural populations of Tetranychus urticae Koch. Insect Molecular Biology, 2011,20:311-321.
173. Narita S, Nomura M, Kato Y, Fukatsu T. Genetic structure of sibling butterfly species affected by Wolbachia infection sweep:evolutionary and biogeographical implications. Molecular Ecology,2006,15:1095-1108.
174. Fu Y, Li. W. Statistical tests of neutrality of mutations. Genetics,1993,133:693-709.
175. Slatkin M. Isolation by distance in equilibrium and non-equilibrium populations. Evolution, 1993,47:264-279.
176. Douglas CW. Mitochondrial DNA mutations in disease and aging. Environmental and Molecular Mutagenesis,2010,51:440-450.
177. Boore JL. Animal mitochondrial genomes. Nucleic Acids Research,1999,27:1767-1780.
178. Murrell A, Barker SC. Multiple origins of parasitism in lice:phylogenetic analysis of SSU rDNA indicates that the Phthiraptera and Psocoptera are not monophyletic. Parasitology Research,2005,97:274-280.
179. Kim MJ, Kang AR, Jeong HC, Kim KG, Kim I. Reconstructing intraordinal relationships in Lepidoptera using mitochondrial genome data with the description of two newly sequenced lycaenids, Spindasis takanonis and Protantigius superans (Lepidoptera:Lycaenidae). Molecular Phylogenetics and Evolution,2011,61:436-445.
180. Cameron S, Lambkin C, Barker S, Whiting M. A mitochondrial genome phylogeny of Diptera:whole genome sequence data accurately resolve relationships over broad timescales with high precision. Systematic Entomology,2007,32:40-59.
181. Wei S, Shi M, Sharkey MJ, Achterberg Cv, Chen X. Comparative mitogenomics of Braconidae (Insecta:Hymenoptera) and the phylogenetic utility of mitochondrial genomes with special reference to Holometabolous insects. BMC Genomics,2010,11:371.
182. 龚鹏,杨效文,张孝羲,刘向东,陈晓峰.棉蚜(Aphis gossypii)种群寄主分化和季节分化的微卫星引物PCR研究.生态学报,2001,21：765-771.
183. Zhang DX, Hewitt GM. Assessment of the universality and utility of a set of conserved mitochondrial COI primers in insects. Insect Molecular Biology,1997,6:143-150.
184. Simon C, Buckley TR, Frati F, Stewart JB, Beckenbach AT. Incorporating molecular evolution into phylogenetic analysis, and a new compilation of conserved polymerase chain reaction primers for animal mitochondrial DNA. Annual Review of Ecology, Evolution, and Systematics,2006,37:545-79.
185. Hopp TP, Woods KR. Prediction of protein antigenic determinants from amino acid sequences. Proceedings of the National Academy of Sciences of the United States of America, 1981, 78: 3824-3828.
186. Lowe TM, Eddy SR. tRNAscan-SE: A program for improved detection of transfer RNA genes in genomic sequence. Nucleic Acids Research, 1997, 25: 955-964.
187. Laslett D, Canback B. ARWEN: a program to detect tRNA genes in metazoan mitochondrial nucleotide sequences. Bioinformatics, 2008, 24: 172-175.
188. Mathews DH. Predicting a set of minimal free energy RNA secondary structures common to two sequences. Bioinformatics, 2005, 21: 2246-2253.
189. Pfaffl MW. A new mathematical model for relative quantification in real-time RT-PCR. Nucleic Acids Research, 2001, 29: e45.
190. Cameron SL, Johnson KP, Whiting MF. The mitochondrial genome of the screamer louse Bothriometopus (Phthiraptera: Ischnocera): effects of extensive gene rearrangements on the evolution of the genome. Journal of Molecular Evolution, 2007, 65: 589-604.
191. Covacin C, Shao R, Cameron S, Barker SC. Extraordinary number of gene rearrangements in the mitochondrial genomes of lice (Phthiraptera: Insecta). Insect Molecular Biology, 2006, 15: 63-68.
192. Shao R, Campbell NJH, Barker SC. Numerous gene rearrangements in the mitochondrial genome of the wallaby louse, Heterodoxus macropus (Phthiraptera). Molecular biology and evolution, 2001, 18: 858-865.
193. 魏书军.膜翅目线粒体基因组的特征与进化及其在系统发育研究中的应用.博士毕业论文,2009,杭州：浙江大学.
194. Fearnley IM, Walker JE. Two overlapping genes in bovine mitochondrial DNA encode membrane components of ATP synthase. The EMBO Journal, 1986, 5: 2003-2008.
195. Ojala D, Montoya J, Attardi G. tRNA punctuation model of RNA processing in human mitochondria. Nature, 1981, 290: 470-474.
196. Giege R, Juhling F, Putz J, Stadler P, Sauter C, Florentz C. Structure of transfer RNAs: similarity and variability. WIREs RNA, 2012, 3: 37-61.
197. Liu B, Liu Y, Motyka SA, Agbo EEC, Englund PT. Fellowship of the rings: the replication of kinetoplast DNA. TRENDS in Parasitology, 2005, 21: 363-369.
198. Lukes J, Guilbride DL, Votypka J, Zikova A, Benne R, Englund PT. Kinetoplast DNA network: evolution of an improbable structure. Eukaryotic Cell, 2002, 1: 495-502.
199. Bogenhagen DF. Mitochondrial DNA nucleoid structure. Biochimica et Biophysica Acta, 2011: doi:10.1016/j.bbagrm.2011.11.005.
200. Johnson KP, Whiting MF. Multiple genes and the monophyly of Ischnocera. Molecular Phylogenetics and Evolution, 2002, 22: 101-110.
201. Lavrov DV. Key transitions in animal evolution: a mitochondrial DNA perspective. Integrative and Comparative Biology, 2007, 47: 734-743.
202. Helfenbein K, Fourcade H, Vanjani R, Boore J. The mitochondrial genome of Paraspadella gotoi is highly reduced and reveals that chaetognaths are a sister group to protostomes. Proceedings of the National Academy of Sciences of the United States of America,2004, 101:10639-10643.
203. Domes K, Maraun M, Scheu S, Cameron SL. The complete mitochondrial genome of the sexual oribatid mite Steganacarus magnus:genome rearrangements and loss of tRNAs. BMC Genomics,2008,9:532.
204. Hayashi JL, Ohta S, Kikuchi A, Takemitsu M, Goto Y, Nonaka I. Introduction of disease-related mitochondrial DNA deletions into HeLa cells lacking mitochondrial DNA results in mitochondrial dysfunction. Proceedings of the National Academy of Sciences of the United States of America,1991,88:10614-10618.
205. Kajander OA, Rovio AT, Majamaa K, Poulton J, Spelbrink JN, Holt IJ, Karhunen PJ, Jacobs HT. Human mtDNA sublimons resemble rearranged mitochondrial genoms found in pathological states. Human Molecular Genetics,2000,9:2821-2835.
206. Melov S, Hinerfeld D, Esposito L, Wallace DC. Multi-organ characterization of mitochondrial genomic rearrangements in ad libitum and caloric restricted mice show striking somatic mitochondrial DNA rearrangements with age. Nucleic Acids Research, 1997,25:974-982.
207. Palmer JD, Shields CR. Tripartite structure of the Brassica campestris mitochondrial genome. Nature,1984,307:437-440.
208. Wu Y, Li ZH, Wu JJ. Polymorphic microsatellite markers in the Melon Fruit Fly, Bactrocera cucurbitae (Coquillett) (Diptera:Tephritidae). Molecular Ecology Resources,2009,9: 1404-1406.
209. Bloor PA, Barker FS, Watts PC, Noyes HA, Kemp SJ. Microsatellite libraries by enrichment. 2001:http://www.genomics.liv.ac.uk/animal/Protocoll.html.
210. Weber JL. Informativeness of human (dC-dA)n · (dG-dT)n polymorphisms. Genomics, 1990,7:524-530.
211. Rassmann K, Schlotterer C, Tautz D. Isolation of simple sequence loci for use in polymerase chain reaction based DNA fingerprinting. Electrophoresis,1991,12:113-118.
212. Fagerberg AJ, Fulton RE, IV WCB. Microsatellite loci are not abundant in all arthropod genomes:analyses in the hard tick, Ixodes scapularis and the yellow fever mosquito, Aedes aegypt. Insect Molecular Biology,2001,10:225-236.
213. Steiner FM, Arthofer W, Schlick-Steiner BC, Crozier RH, Stauffer C. Twenty four new microsatellite markers in two invasive pavement ants, Tetramorium sp. E and T. tsushimae (Hymenoptera:Formicidae). Conservation Genetics,2008,9:757-759.
214. 樊勇,马雅军.中华按蚊多态微卫星DNA位点的筛选和特征研究.第四军医大学学报,2008,29：1537-1539.
215. 张国彦,翟保平.东方粘虫(Pseudaletia separata (Walker))微卫星富集文库的构建与分析.生态学报,2008,28：3860-3867.
216. Santana QC, Coetzee MPA, Steenkamp ET, Mlonyeni OX, Hammond GNA, Wingfield MJ, Wingfield BD. Microsatellite discovery by deep sequencing of enriched genomic libraries. Biotechniques,2009,46:217-223.
217. Zhang DX. Lepidopteran microsatellite DNA:redundant but promising. TRENDS in Ecology and Evolution,2004,19:507-509.
218. Katti MV, Ranjekar PK, Gupta VS. Differential distribution of simple sequence repeats in eukaryotic genome sequences. Molecular Biology and Evolution,2001,18:1161-1167.
219. Toth G, Gaspari Z, Jurka J. Microsatellites in different eukaryotic genomes:survey and analysis. Genome Research,2000,10:967-981.