Abstract Much progress in speciation research stems from documenting patterns of morphological and genetic variation in hybrid zones. Contrasting patterns of marker introgression in different sections of the contact can provide valuable insights on the relative importance of various evolutionary mechanisms maintaining species differences in the face of hybridization and gene flow and on hybrid zone temporal and spatial dynamics. We studied species interactions in the common toads Bufo bufo and B. spinosus in France and northwestern Italy using morphological and molecular data from the mitochondrial and nuclear genomes in an extensive survey, including two independent transects west and east of the Alps. At both we found sharp, coincident and concordant nuclear genetic transitions. However, morphological clines were wider or absent and mtDNA introgression was asymmetric. We discuss alternative, non-exclusive hypotheses about evolutionary processes generating these patterns, including drift, selection, long-distance dispersal and spatial shifts in hybrid zone location and structure. The distribution of intraspecific mtDNA lineages supports a scenario in which B. bufo held a local refugium during the last glacial maximum. Present day genetic profiles are best explained by an advance of B. spinosus from a nearby Iberian refugium, largely superseding the local B. bufo population, followed by an advance of B. bufo from the Balkans, with prongs north and south of the Alps, driving B. spinosus southwards. A pendulum moving hybrid zone, first northwards and then southwards, explains the wide areas of introgression at either side of the current position of the contact zones. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.

Plant phylogeographic studies in East Asia have provided support for the biogeographic hypothesis that the complex landforms and climate of this region have provided substantial opportunities for allopatric speciation. However, most of these studies have been based on maternally inherited chloroplast DNA (cpDNA) markers and were therefore unable to reveal the role of pollen-mediated gene flow in preventing population divergence. Here, we investigate the phylogeography of the Chinese walnut Juglans cathayensis, a temperate deciduous tree widely distributed across disjunct montane sites in subtropical China. We genotyped 19 populations using seven cpDNA fragments and ten nuclear microsatellite loci and modeled the ecological niche of J. cathayensis. CpDNA analysis identified a total of nine haplotypes, and each of the 19 sampled populations was fixed for a single haplotype, displaying a prominent phylogeographic structure. The results of ecological niche modeling indicated that J. cathayensis populations survived the last glaciation in situ, although they were probably more fragmented than today. In contrast, we detected a much weaker, but nonetheless clear, genetic structure based on nuclear microsatellite data. Our study demonstrates how extensive pollen flow can erase the genetic imprint of long-term refugial isolation in maternal lineages, effectively preventing population differentiation in temperate, particularly wind-pollinated, forest trees in subtropical China.

利用叶绿体基因（rbcl和psbA-trnH）对青藏高原地区的10个手参（Gymnadenia conopsea）种群进行谱系地理学研究，旨在揭示手参的遗传谱系分布格局及其演化过程。结果表明，共得到11个单倍型，分子方差分析显示65.93%的遗传变异来源于种群间，FST值为0.659（P<0.01），种群间存在显著的遗传分化；遗传分化系数NST（0.398）小于GST（0.626），说明青藏高原地区的手参没有明显的谱系地理结构；多峰的失配分析曲线和Tajima’s D中性检验（-1.455，P>0.1）表明手参种群近期没有经历扩张；系统发育分析和分歧时间估计表明，现代青藏高原上的手参为本地起源，手参单倍型在19.08 Mya的第三纪中新世时期开始分化，与青藏高原隆起关系密切，青藏高原上手参的遗传分布格局在第四纪冰期之前已基本形成。

利用叶绿体基因（rbcl和psbA-trnH）对青藏高原地区的10个手参（Gymnadenia conopsea）种群进行谱系地理学研究，旨在揭示手参的遗传谱系分布格局及其演化过程。结果表明，共得到11个单倍型，分子方差分析显示65.93%的遗传变异来源于种群间，FST值为0.659（P<0.01），种群间存在显著的遗传分化；遗传分化系数NST（0.398）小于GST（0.626），说明青藏高原地区的手参没有明显的谱系地理结构；多峰的失配分析曲线和Tajima’s D中性检验（-1.455，P>0.1）表明手参种群近期没有经历扩张；系统发育分析和分歧时间估计表明，现代青藏高原上的手参为本地起源，手参单倍型在19.08 Mya的第三纪中新世时期开始分化，与青藏高原隆起关系密切，青藏高原上手参的遗传分布格局在第四纪冰期之前已基本形成。

A phylogenetic analysis of the New World genus Cuphea was conducted employing sequences from the nuclear rDNA internal transcribed spacer (ITS) and chloroplast trnL-trnF spacer and rpl16 intron. The analysis expands the number of Cuphea species from 53 in an earlier ITS study to 70 and adds two chloroplast data sets in order to generate a more complete and robust phylogeny and to test a previous result that suggested the presence of a large North American clade. Results reaffirm the monophyly of Cuphea with Pleurophora as the sister genus and recover a basal divergence event that mirrors the two subgenera of the current classification. Phylogenies of the two chloroplast regions are largely unresolved beyond the initial dichotomy and some resolution at the terminal and subterminal nodes. Based on the ITS phylogeny, five major clades are recognized. Subgenus Cuphea (Clade 1), defined morphologically by the synapomorphic loss of bracteoles, is sister to the much larger subg. Bracteolatae (Clades 2鈥5). Clades 2 4, comprising the South American and Caribbean species, grade successively to Clade 5, an exclusively North American lineage of 29 species. Among the 12 sections included in the study, only section Trispermum, a subclade of Clade 4, is monophyletic. Sectoin Pseudocircaea is nested within Clade 3, which is largely equivalent to section Euandra. The North American endemic clade includes four sections, of which none are recovered as monophyletic in this study.

Abstract Although organelle inheritance is predominantly maternal across animals and plants, biparental chloroplast inheritance has arisen multiple times in the angiosperms. Biparental inheritance has the potential to impact the evolutionary dynamics of cytonuclear incompatibility, interactions between nuclear and organelle genomes that are proposed to be among the earliest types of genetic incompatibility to arise in speciation. We examine the interplay between biparental inheritance and cytonuclear incompatibility in Campanulastrum americanum, a plant species exhibiting both traits. We first determine patterns of chloroplast inheritance in genetically similar and divergent crosses, and then associate inheritance with hybrid survival across multiple generations. There is substantial biparental inheritance in C.0002americanum. The frequency of biparental inheritance is greater in divergent crosses and in the presence of cytonuclear incompatibility. Biparental inheritance helps to mitigate cytonuclear incompatibility, leading to increased fitness of F 1 hybrids and recovery in the F 2 generation. This study demonstrates the potential for biparental chloroplast inheritance to rescue cytonuclear compatibility, reducing cytonuclear incompatibility's contribution to reproductive isolation and potentially slowing speciation. The efficacy of rescue depended upon the strength of incompatibility, with a greater persistence of weak incompatibilities in later generations. These findings suggest that incompatible plastids may lead to selection for biparental inheritance. 0008 2016 The Authors. New Phytologist 0008 2016 New Phytologist Trust.

Phylogeography is a young, vigorous and integrative field of study that uses genetic data to understand the history of populations. This field has recently expanded into many areas of biology and also into several historical disciplines of Earth sciences. In this review, I present a numerical synthesis of the phylogeography literature based on an examination of over 3000 articles published during the first 20 years of the field (i.e. from 1987 to 2006). Information from several topics needed to evaluate the progress, tendencies and deficiencies of the field is summarized for 10 major groups of organisms and at a global scale. The topics include the geography of phylogeographic surveys, comparative nature of studies, temporal scales and major environments investigated, and genetic markers used. I also identify disparities in research productivity between the developing and the developed world, and propose ways to reduce some of the challenges faced by phylogeographers from less affluent countries. Phylogeography has experienced explosive growth in recent years fuelled by developments in DNA technology, theory and statistical analysis. I argue that the intellectual maturation of the field will eventually depend not only on these recent developments, but also on syntheses of comparative information across different regions of the globe. For this to become a reality, many empirical phylogeographic surveys in regions of the Southern Hemisphere (and in developing countries of the Northern Hemisphere) are needed. I expect the information and views presented here will assist in promoting international collaborative work in phylogeography and in guiding research efforts at both regional and global levels.

Micromeria Benth. (Lamiaceae, Nepetoideae) is a very common genus in the Mediterranean region. To test the monophyly of the genus and to elucidate its phylogenetic placement within subtribe Menthinae (Dumort) Endl. of tribe Mentheae Dumort we performed parsimony analysis of trnK intron sequence data of 51 accessions representing 15 genera of Nepetoideae and two genera of subfamily Ajugoideae. Tree topology reveals a well-supported "core group" indicating four distinct lineages. The first one comprises three species of Satureja L. s.str., the second one includes taxa of the genus Clinopodium L. from both the Old and the New World, paraphyletic with respect to Monarda L. and two species of Micromeria section Pseudomelissa Benth. A third group contains all samples of the remaining Micromeria species. Within this monophyly, a western lineage including taxa from NW Africa, the Balearic, and the Canary Islands, is sister to an eastern lineage with species distributed from SE Asia to the western Mediterranean. In a further clade the genera Thymbra L., Thymus L., and Origanum L. are grouped together. Combined analysis using a reduced dataset of trnK/trnL-F sequences increased support for the infrageneric resolution within Micromeria. Based on the phylogenetic reconstructions there is evidence that the genus as currently circumscribed is polyphyletic. Results are discussed in the context of morphology, karyology, and biogeography, outlining the necessity of removing section Pseudomelissa from Micromeria.

Aim The strawberry poison frog, Oophaga pumilio, has undergone a remarkable radiation of colour morphs in the Bocas del Toro archipelago in Panama. This species shows extreme variation in colour and pattern between populations that have been geographically isolated for < 10,000 years. While previous research has suggested the involvement of divergent selection, to date no quantitative test has examined this hypothesis.Location Bocas del Toro archipelago, Panama.Methods We use a combination of population genetics, phylogeography and phenotypic analyses to test for divergent selection in coloration in O. pumilio. Tissue samples of 88 individuals from 15 distinct populations were collected. Using these data, we developed a gene tree using the mitochondrial DNA (mtDNA) d-loop region. Using parameters derived from our mtDNA phylogeny, we predicted the coalescence of a hypothetical nuclear gene underlying coloration. We collected spectral reflectance and body size measurements on 94 individuals from four of the populations and performed a quantitative analysis of phenotypic divergence.Results The mtDNA d-loop tree revealed considerable polyphyly across populations. Coalescent reconstructions of gene trees within population trees revealed incomplete genotypic sorting among populations. The quantitative analysis of phenotypic divergence revealed complete lineage sorting by colour, but not by body size: populations showed non-overlapping variation in spectral reflectance measures of body coloration, while variation in body size did not separate populations. Simulations of the coalescent using parameter values derived from our empirical analyses demonstrated that the level of sorting among populations seen in colour cannot reasonably be attributed to drift.Main conclusions These results imply that divergence in colour, but not body size, is occurring at a faster rate than expected under neutral processes. Our study provides the first quantitative support for the claim that strong diversifying selection underlies colour variation in the strawberry poison frog.

Phylogeography in plants is hampered by lack of DNA-sequence regions that detect sufficient variation in intra-specific lineages to reveal historical patterns. We tested 13 putatively highly variable non-coding chloroplast regions in six species complexes, from four different angiosperm families, where phylogeographic patterns have previously been identified using restriction fragment length polymorphism analysis of the chloroplast genome. All regions tested amplified in most of the species. The intergenic spacer regions trnQ–rps16, trnS–trnG, psbA–trnH, psbD–trnT and ndhC–trnV were the five most promising regions for phylogeographic analysis in terms of variability, and petB and rpl16 were variable, given the utility of being amplified in a single reaction. The trnQ–rps16 and psbA–trnH intergenic spacer regions and the rpl16 D4-loop intron showed variation between known lineages in all species. The psbA–trnH intergenic spacer that has been suggested as a suitable barcoding gene for plants, generally showed a level of variation similar to that in other variable regions in the species investigated here, suggesting that some caution is required in the use of this region for barcoding applications. The present analysis identified a set of seven chloroplast regions that are a useful basis for informed selection of sequences for assessment of phylogeographic structure in plants.

DNA barcoding involves sequencing a standard region of DNA as a tool for species identification. However, there has been no agreement on which region(s) should be used for barcoding land plants. To provide a community recommendation on a standard plant barcode, we have compared the performance of 7 leading candidate plastid DNA regions (atpF-atpH spacer, matK gene, rbcL gene, rpoB gene, rpoC1 gene, psbK-psbl spacer, and trnH-psbA spacer). Based on assessments of recoverability, sequence quality, and levels of species discrimination, we recommend the 2-locus combination of rbcL+ matK as the plant barcode. This core 2-locus barcode will provide a universal framework for the routine use of DNA sequence data to identify specimens and contribute toward the discovery of overlooked species of land plants.

We have developed a diagnostic method to screen rapidly for plant species potentially capable of biparental inheritance of plastid DNA using the DNA fluorochrome 4',6-diamidino-2-phenyl-indole (DAPI) in conjunction with epifluorescence microscopy. Pollen shed from 235 plant species (including about 50 of agronomic importance) representing 80 families were screened. Putative plastid DNA was detected in the generative and/or sperm cells of pollen from 26 genera (43 species) representing 15 families. Plastid DNA was not detected in the generative or sperm cells of pollen from 192 plant species, thereby strongly suggesting that these species have only maternal inheritance. Our cytological diagnosis corroborated the known genetic evidence in 42 plant species and conflicted with the genetic reports in five species, which are discussed. The data suggest that biparental inheritance of plastids is rare; overall, it may occur in about 14% of flowering plant genera, examples of which are scattered among 19% of the families examined. This methodology also readily reveals whether pollen is bi- or trinucleate.

The isolation-with-migration (IM) model is commonly used to make inferences about gene flow during speciation, using polymorphism data. However, it has been reported that the parameter estimates obtained by fitting the IM model are very sensitive to the model’s assumptions—including the assumption of constant gene flow until the present. This article is concerned with the isolation-with-initial-migration (IIM) model, which drops precisely this assumption. In the IIM model, one ancestral population divides into two descendant subpopulations, between which there is an initial period of gene flow and a subsequent period of isolation. We derive a very fast method of fitting an extended version of the IIM model, which also allows for asymmetric gene flow and unequal population sizes. This is a maximum-likelihood method, applicable to data on the number of segregating sites between pairs of DNA sequences from a large number of independent loci. In addition to obtaining parameter estimates, our method can also be used, by means of likelihood-ratio tests, to distinguish between alternative models representing the following divergence scenarios: (a) divergence with potentially asymmetric gene flow until the present, (b) divergence with potentially asymmetric gene flow until some point in the past and in isolation since then, and (c) divergence in complete isolation. We illustrate the procedure on pairs ofDrosophilasequences from 6530,000 loci. The computing time needed to fit the most complex version of the model to this data set is only a couple of minutes. The R code to fit the IIM model can be found in the supplementary files of this article.

Evolutionary theory is primed to synthesize microevolutionary processes with macroevolutionary divergence by taking advantage of multilocus multispecies genomic data in the molecular evolutionary analysis of biodiversity. While coalescent theory bridges across timescales to facilitate this integration, it is important to appreciate the assumptions, caveats, and recent theoretical advances so as to most effectively exploit genomic analysis. Here I outline the connections between population processes and phylogeny, with special attention to how genomic features play into underlying predictions. I discuss empirical and theoretical complications, and solutions, relating to recombination and multifurcating genealogical processes, predictions about how genome structure affects gene tree heterogeneity, and practical choices in genome sequencing and analysis. I illustrate the conceptual implications and practical benefits of how genomic features generate predictable patterns of discordance of gene trees and species trees along genomes, for example, as a consequence of how regions of low recombination and sex linkage interact with natural selection and with the accumulation of reproductive incompatibilities in speciation. Moreover, treating population genetic parameters as characters to be mapped onto phylogenies offers a new way to understand the evolutionary drivers of diversity within and differentiation between populations. Despite a number of challenges conferred by genomic information, the melding of phylogenetics, phylogeography and population genetics into integrative molecular evolution is poised to improve our understanding of biodiversity at all levels. (C) 2013 Elsevier Inc. All rights reserved.

Under a coalescent model for within-species evolution, gene trees may differ from species trees to such an extent that the gene tree topology most likely to evolve along the branches of a species tree can disagree with the species tree topology. Gene tree topologies that are more likely to be produced than the topology that matches that of the species tree are termed anomalous, and the region of branch-length space that gives rise to anomalous gene trees (AGTs) is the anomaly zone. We examine the occurrence of anomalous gene trees for the case of five taxa, the smallest number of taxa for which every species tree topology has a nonempty anomaly zone. Considering all sets of branch lengths that give rise to anomalous gene trees, the largest value possible for the smallest branch length in the species tree is greater in the five-taxon case (0.1934 coalescent time units) than in the previously studied case of four taxa (0.1568). The five-taxon case demonstrates the existence of three phenomena that do not occur in the four-taxon case. First, anomalous gene trees can have the same unlabeled topology as the species tree. Second, the anomaly zone does not necessarily enclose a ball centered at the origin in branch-length space, in which all branches are short. Third, as a branch length increases, it is possible for the number of AGTs to increase rather than decrease or remain constant. These results, which help to describe how the properties of anomalous gene trees increase in complexity as the number of taxa increases, will be useful in formulating strategies for evading the problem of anomalous gene trees during species tree inference from multilocus data.

Flood et al. discover that strong selection on a chloroplast gene has extended to the nuclear genome, which has hitched a ride along with the selected chloroplast. This is the first description of organelle-mediated genetic draft and shows that selection on organelles can directly impact nuclear genetic diversity.

Empirical phylogeographic studies have progressively sampled greater numbers of loci over time, in part motivated by theoretical papers showing that estimates of key demographic parameters improve as the number of loci increases. Recently, next-generation sequencing has been applied to questions about organismal history, with the promise of revolutionizing the field. However, no systematic assessment of how phylogeographic data sets have changed over time with respect to overall size and information content has been performed. Here, we quantify the changing nature of these genetic data sets over the past 20 years, focusing on papers published in Molecular Ecology . We found that the number of independent loci, the total number of alleles sampled and the total number of single nucleotide polymorphisms (SNPs) per data set has improved over time, with particularly dramatic increases within the past 5 years. Interestingly, uniparentally inherited organellar markers (e.g. animal mitochondrial and plant chloroplast DNA) continue to represent an important component of phylogeographic data. Single-species studies (cf. comparative studies) that focus on vertebrates (particularly fish and to some extent, birds) represent the gold standard of phylogeographic data collection. Based on the current trajectory seen in our survey data, forecast modelling indicates that the median number of SNPs per data set for studies published by the end of the year 2016 may approach ~20 000. This survey provides baseline information for understanding the evolution of phylogeographic data sets and underscores the fact that development of analytical methods for handling very large genetic data sets will be critical for facilitating growth of the field.

The Amentotaxus argotaenia (Taxaceae) species complex is comprised of four relict and endangered gymnosperms. Amentotaxus argotaenia and A. yunnanensis are scattered throughout South and Central China, and A. formosana, and A. poilanei are restricted to Taiwan and Vietnam. In the present study, we examined genetic divergence and biogeographical patterns in the species complex based on chloroplast DNA atpI-atpH and rpS16-trnK intergenic spacers, the mitochondrial DNA nad1 intron, and microsatellite markers. Genealogical analyses revealed systematic inconsistencies between organellar DNA markers, with paraphyletic species inferred based on cpDNA, versus a monophyletic A. formosana, and clustering of A. poilanei within A. yunnanensis at mtDNA. AMOVA revealed that most organellar DNA variants resided among species and populations, a pattern also supported by clustering with STRUCTURE analyses on microsatellites. Higher levels of genetic diversity in the Yongxin population of A. argotaenia reflected features of a refugium. Most speciation events were rather recent, with divergence no earlier than 2.50 MYA, although many ancient lineages derived 4.12-13.84 MYA were maintained within species. Significant genetic structuring within A. argotaenia was detected among three geographical regions-a phylogeographical pattern likely attributable to past fragmentations. In contrast to paraphyletic A. argotaenia based on the maintenance of ancestral polymorphisms, interspecific hybridization likely contributed to polyphyly in A. yunnanensis. Furthermore, despite the low likelihood of recurrent gene flow due to geographical isolation, IM analyses revealed continued interspecific gene flow after species divergence, reflecting a parapatric speciation model.

We used chloroplast DNA sequences from matK and rbcL to infer the phylogeny for 101 of the approximately 111 species of Pinus (Pinaceae). At the level of subsection and above, the cpDNA tree is congruent with phylogenies based on nuclear DNA with one notable exception: cpDNA sequences from subsect. Contortae are sister to all other North American hard pines rather than occupying a more derived position in the same clade. We used the cpDNA tree plus evidence from nuclear ribosomal DNA and morphology to propose a new classification for the genus. The molecular phylogenies are symmetrical at the deepest branches of the genus, allowing for the delineation of two subgenera, each with two sections that form sister groups. Within sections, clades were slightly asymmetric and sometimes ambiguously resolved. To accomodate ambiguity in some interrelationships, avoid the creation of new ranks, and retain traditional names, we recognised up to three monophyletic subsections per section. Subgenus Pinus (the diploxylon, or hard pines) is divided into the predominantly Eurasian and Mediterranean section Pinus, composed of subsections Pinus and Pinaster, and the strictly North American section Trifoliae, composed of subsections Australes, Ponderosae, and Contortae. Subgenus Strobus (the haploxylon, or soft pines) is divided into the strictly North American section Parrya, composed of subsections Cembroides, Nelsoniae, and Balfourianae, and the Eurasian and North American section Quinquefoliae, composed of subsections Gerardianae, Krempfianae, and Strobus. Mapping of ten morphological and distributional characters indicates that two were diagnostic for infrageneric taxa: the number of vascular bundles per leaf distinguishes subgenus Pinus from subgenus Strobus, and a terminal-positioned umbo on the ovulate cone scale is diagnostic of subsect. Strobus.

Excoecaria agallochais one of the predominant mangrove species in the Indo-West Pacific (IWP) region with an extensive range of distribution. To infer the current geographical patterns of genetic...

半日花是半日花科的一种半灌木或灌木，间断分布于我国内蒙古和新疆地区，由于种群数量稀少，现已被列为国家二级保护植物，开展半日花种群遗传多样性和谱系地理结构的研究对于半日花的科学保护具有重要意义。本文基于叶绿体DNA片段trnc—ycf6序列分析和AMOYA发现，半日花种群的遗传变异主要存在于居群间（93％），且居群问遗传分化很高（Gst=0．71，Fst=0．93，Nst=0．92）；9个单倍型中4种为内蒙古特有，5种单倍型为新疆特有，有着显著的谱系地理学结构（Nst〉Gst）且居群问基因流较低（Nm=0．1），表明半日花种群存在地理隔离，这也是该物种现有单倍型分布格局形成的原因。单倍型最大简约树和网络状树分析，西鄂尔多斯高原可能是半Et花冰期“避难所”。

半日花是半日花科的一种半灌木或灌木，间断分布于我国内蒙古和新疆地区，由于种群数量稀少，现已被列为国家二级保护植物，开展半日花种群遗传多样性和谱系地理结构的研究对于半日花的科学保护具有重要意义。本文基于叶绿体DNA片段trnc—ycf6序列分析和AMOYA发现，半日花种群的遗传变异主要存在于居群间（93％），且居群问遗传分化很高（Gst=0．71，Fst=0．93，Nst=0．92）；9个单倍型中4种为内蒙古特有，5种单倍型为新疆特有，有着显著的谱系地理学结构（Nst〉Gst）且居群问基因流较低（Nm=0．1），表明半日花种群存在地理隔离，这也是该物种现有单倍型分布格局形成的原因。单倍型最大简约树和网络状树分析，西鄂尔多斯高原可能是半Et花冰期“避难所”。

目的研究我国自然地理分布范围内远志Polygala tenuifolia的分子谱系地理情况,揭示其地理分布格局的形成原因,推测该物种潜在的冰期避难所及冰期后的迁移扩散路线。方法使用叶绿体非编码片段(trn L内含子序列)对远志在我国自然地理分布区内的39个居群308个个体的遗传变异分布模式进行检测。结果共发现26个多态性位点,得到12种叶绿体单倍型。单倍型系统发育分析显示远志自然居群可划分为南、北2个地理组群:北方(包括中国东北、中部、西北地区)组群和南方组群,南北组群没有共享单倍型。居群遗传结构分析表明2个地理组群之间遗传分化较大(Gst=0.783,P0.001),物种水平遗传多样性较高(Ht=0.755),北方组群不存在明显的谱系地理结构。结论第四纪冰期时远志在中国北方和南方地区存在多个避难所;冰期后或间冰期,北方地区发生了明显的居群扩张事件;南、北组群的分化可能是由于长期的地理隔离所致。

目的研究我国自然地理分布范围内远志Polygala tenuifolia的分子谱系地理情况,揭示其地理分布格局的形成原因,推测该物种潜在的冰期避难所及冰期后的迁移扩散路线。方法使用叶绿体非编码片段(trn L内含子序列)对远志在我国自然地理分布区内的39个居群308个个体的遗传变异分布模式进行检测。结果共发现26个多态性位点,得到12种叶绿体单倍型。单倍型系统发育分析显示远志自然居群可划分为南、北2个地理组群:北方(包括中国东北、中部、西北地区)组群和南方组群,南北组群没有共享单倍型。居群遗传结构分析表明2个地理组群之间遗传分化较大(Gst=0.783,P0.001),物种水平遗传多样性较高(Ht=0.755),北方组群不存在明显的谱系地理结构。结论第四纪冰期时远志在中国北方和南方地区存在多个避难所;冰期后或间冰期,北方地区发生了明显的居群扩张事件;南、北组群的分化可能是由于长期的地理隔离所致。

Approximately 20 years ago, Avise and colleagues proposed the integration of phylogenetics and population genetics for investigating the connection between micro- and macroevolutionary phenomena. The new field was termed phylogeography. Since the naming of the field, the statistical rigor of phylogeography has increased, in large part due to concurrent advances in coalescent theory which enabled model-based parameter estimation and hypothesis testing. The next phase will involve phylogeography increasingly becoming the integrative and comparative multi-taxon endeavor that it was originally conceived to be. This exciting convergence will likely involve combining spatially-explicit multiple taxon coalescent models, genomic studies of natural selection, ecological niche modeling, studies of ecological speciation, community assembly and functional trait evolution. This ambitious synthesis will allow us to determine the causal links between geography, climate change, ecological interactions and the evolution and composition of taxa across whole communities and assemblages. Although such integration presents analytical and computational challenges that will only be intensified by the growth of genomic data in non-model taxa, the rapid development of ikelihood-free approximate Bayesian methods should permit parameter estimation and hypotheses testing using complex evolutionary demographic models and genomic phylogeographic data. We first review the conceptual beginnings of phylogeography and its accomplishments and then illustrate how it evolved into a statistically rigorous enterprise with the concurrent rise of coalescent theory. Subsequently, we discuss ways in which model-based phylogeography can interface with various subfields to become one of the most integrative fields in all of ecology and evolutionary biology.

This paper extends to hermaphrodite outcrossing plant populations the existing gene genealogy theories for a sample of genes randomly chosen from geographically discrete or continuously distributed populations. Three plant genomes ( nuclear, chloroplast and mitochondrial DNA) with different modes of inheritance are considered separately due to the difference in migration rates. It is shown that on certain assumptions, the previous coalescence theories can be applied to plant by appropiate reparamterization of the effective population size and migration rate specific to each genome. One result is that estimation of the ratio of pollen to seed flow from a sample of n (n 2) indicidual gene can be obtained in terms of the number of segregating sites between and within populations that are discretely distributed in space. Another result of theoretical interest is that in the duscrete model of population structure, mean coalescent time is the shortest for the paternally inherited genes (cpDNA in conifers) and, given certain conditions, is the longest for the maternally inherited genes (cpDNA in angiosperms and mtDNA in conifers and angiosperms). However, these results are diffcult to obtain in the model of population that is continuously distributed in space.

<p>一个植物种群体的空间分布大致可划分为两类:离散分布与连续分布。然而隐藏在这种表观物理分布内部的遗传变异是十分复杂的。这种变异是如何分布的呢?这种分布又是如何维持的呢?因此,认识一个种的群体遗传结构有助于我们理解该种的进化过程,并提供为遗传资源保护做出决策的重要信息。本文对三大经典遗传结构模型(岛屿模型、步石模型及距离隔离模型)与渐变群理论及它们的异体和发展进行了详细的分析和评述。目前获得的许多不同类型的遗传标记使这些理论的应用得到不同程度的实现,但由于这些理论结论涉及到许多假设,在实际应用时应十分慎重。作者认为许多已建立的适合于动物群体遗传结构理论不能简单地直接应用于植物群体上,今后一个重要的研究内容就是要建立适合于植物群体的遗传结构理论。</p>

<p>一个植物种群体的空间分布大致可划分为两类:离散分布与连续分布。然而隐藏在这种表观物理分布内部的遗传变异是十分复杂的。这种变异是如何分布的呢?这种分布又是如何维持的呢?因此,认识一个种的群体遗传结构有助于我们理解该种的进化过程,并提供为遗传资源保护做出决策的重要信息。本文对三大经典遗传结构模型(岛屿模型、步石模型及距离隔离模型)与渐变群理论及它们的异体和发展进行了详细的分析和评述。目前获得的许多不同类型的遗传标记使这些理论的应用得到不同程度的实现,但由于这些理论结论涉及到许多假设,在实际应用时应十分慎重。作者认为许多已建立的适合于动物群体遗传结构理论不能简单地直接应用于植物群体上,今后一个重要的研究内容就是要建立适合于植物群体的遗传结构理论。</p>

Previous theories show that tension and ecological zones are indistinguishable in terms of gene frequency clines. Here I analytically show that these two types of zones can be distinguished in terms of genetic statistics other than gene frequency. A two-locus cline model is examined with the assumptions of random mating, weak selection, no drift, no mutation, and multiplicative viabilities. The genetic statistics for distinguishing the two types of zones are the deviations of one- or two-locus genotypic frequencies from Hardy einberg equilibrium (HWE) or from random association of gametes (RAG), and the deviations of additive and dominance variances from the values at HWE. These deviations have a discontinuous distribution in space and different extents of interruptions in the ecological zone with a sharp boundary, but exhibit a continuous distribution in the tension zone. Linkage disequilibrium enhances the difference between the deviations from HWE and from RAG for any two-locus genotypic frequency.

Neutral alleles can eventually pass a hybrid zone and their initial clines generated by a pure diffusion process dissipate with time, irrespective of the presence or absence of physical barriers. However, the transient neutral clines at the nuclear or organelle sites can be reinforced by the cytonuclear disequilibrium generated by diploid seed and haploid pollen dispersal. In this study, the spread of a neutral allele in an ecological zone of hermaphrodite plants is examined under three cytonuclear systems for genomes with contrasting modes of inheritance (paternal, maternal, and biparental inheritance). The results show that the transient neutral clines can exhibit the spatial pattern similar to the selective clines from separate genomes although discordance between them exists. The spread of a neutral allele is not only related to the vectors of seed and pollen dispersal but also to the mode of its inheritance. Pollen dispersal facilitates the direct effects of the selective organelle sites with paternal inheritance on the spread of a neutral nuclear allele. It also enhances its indirect effects on the spread of a neutral organelle allele with maternal inheritance via modifying the cytonuclear disequilibrium. A positive relationship exists between the barriers to the spread of selective nuclear (or organelle) and neutral organelle (or nuclear) alleles. An asymmetric barrier to the spread of the neutral alleles exists on the two sides of the physical barrier, given the presence of symmetric barrier to the spread of the selective alleles. These theoretical predictions highlight the effects of cytonuclear disequilibrium on the spread of a neutral allele and draw attention to our empirical cline analysis with neutral markers.

Crowet al.(1990) and Barton (1992) have examined the critical migration rate for swamping selection in the nuclear system. Here, I use the same methodology to examine the critical migration rate in the cytonuclear system for hermaphrodite plants with a mixed mating system. Two selection schemes for a nuclear gene (heterozygote disadvantage and directional selection) and the directional selection scheme for organelle genes are considered. Results show that under random mating, the previous results are applicable to plant species by appropriate re-parameterization of the migration rate for nuclear and paternal organelle genes. A simple complementary relationship exists between seed and pollen flow in contributing to the critical migration rate. Under the mixed mating system, the critical migration rate of seeds and pollen for nuclear and paternal organelle genes can be changed due to the effects of selection and the cytonuclear linkage disequilibrium generated by migration and inbreeding. A negative but not complementary relationship exists between seed and pollen flow in contributing to the critical migration rate, varying with the mating system. Partial selfing can also adjust the critical seed flow for the maternal organelle gene, with a small critical migration rate for species of a high selfing rate. Both concordance and discordance among cytonuclear genes can occur under certain conditions during the process of swamping selection. This theory predicts the presence of various contributions of seed versus pollen flow to genetic swamping for plants with diverse mating systems.

Understanding mating system as one of reproductive isolating barriers remains important although this barrier is classified in a different sense from behavioral, ecological, and mechanical isolating barriers. Selfing enhances incipient speciation while outcrossing facilitates species integrity. Here, I study how mating system affects gene exchanges between genetically diverging species in a hybrid zone. Results show that a predominant selfing species has a greater barrier to selective gene flow than does a predominant outcrossing species. Barrier to neutral gene flow convexly changes with the selfing rate due to linkage disequilibrium, with a maximum at around intermediate selfing rate. Asymmetric transient or steady-state barriers to neutral gene flow occur between two sides of a hybrid zone when the neutral gene is affected by its linked selective gene whose alternative alleles are adaptive to heterogeneous habitats. Selfing interacts with both a physical barrier and a density-dependent ecological regulation (a logarithmic model) to strengthen the barriers to neutral and selective gene flow. This theory helps to interpret incipient speciation driven by selfing or to explain the asymmetric gene flow or unequal genomic mixtures between closely related species caused by their asymmetric mating systems in natural hybrid zones.

The disproportionately large involvement of the X-chromosome in the isolation of closely related species (the large-X effect) has been reported for many animals, where X-linked genes are mostly hemizygous in the heterogametic sex. The expression of deleterious recessive mutations is thought to drive the frequent involvement of the X-chromosome in hybrid sterility, as well as to reduce interspecific gene flow for X-linked genes. Here, we evaluate the role of the X-chromosome in the speciation of two closely related plant species the white and red campions (Silene latifolia and S. dioica) that hybridize widely across Europe. The two species evolved separate sexes and sex chromosomes relatively recently (~107 years), and unlike most animal species, most X-linked genes have intact Y-linked homologs. We demonstrate that the X-linked genes show a very small and insignificant amount of interspecific gene flow, while gene flow involving autosomal loci is significant and sufficient to homogenise the gene pools of the two species. These findings are consistent with the hypothesis of the large-X effect in Silene and comprise the first report of this effect in plants. Non-hemizygosity of many X-linked genes in Silene males indicates that exposure of recessive mutations to selection may not be essential for the occurrence of the large-X effect. Several possible causes of the large-X effect in Silene are discussed.

A general analytical formula is derived, which predicts the effects of background selection on population differentiation at a neutral locus as a result of its linkage with selected loci of deleterious mutations. The theory is based on the assumptions of random mating, multiplicative fitness, and weak selection in hermaphrodite plants in the island model of population structure. The analytical results show that F st at the neutral locus increases as a result of the effects of background selection, regardless of the dependence or independence among linked background selective loci. The increment in F st is closely related to the magnitude of linkage disequilibria between the neutral locus and selected loci, and can be estimated by the ratio of F st with background selection to F st without background selection minus one. The steady-state linkage disequilibrium between a neutral locus and a selected locus in subpopulations, primarily attained by gene flow, decreases with the recombination rate, and can be enhanced when there are dependence among linked selected loci. Monte Carlo computer simulations with two- and three-locus models show that the analytical formulae perform well under general conditions. Application of the present theory may aid in analyzing the genome-wide mapping of the effect of background selection in terms of F st.

The relationships between seed and pollen flow and cline discordance/concordance between cytoplasmic and nuclear genes, with the incorporation of the effects of natural selection, are formulated for one locus with two alleles, under assumptions of random mating, no drift and no mutation. Results show that under certain conditions, the relative roles of seed and pollen flow in shaping cline discordance/ concordance are very similar to their roles in influencing population differentiation for selectively neutral markers with different modes of inheritance. Where the disequilibria between cytoplasmic and nuclear genes are of the order similar to selection coefficient, cline discordance/concordance can be predicted from the relative values of the ratio of pollen to seed flow and the ratio of selection coefficients. Where the disequilibria attained by seed and pollen flow are significant, the integrated cytonuclear data are recommended for cline analysis. In both cases, the relative rates of selection coefficients between cytoplasmic and nuclear genes can be roughly estimated according to their characteristic length.

Copy-number-variable (CNV) loci differ from single nucleotide polymorphic (SNP) sites in size, mutation rate, and mechanisms of maintenance in natural populations. It is therefore hypothesized that population genetic divergence at CNV loci will differ from that found at SNP sites. Here, we test this hypothesis by analysing 856 CNV loci from the genomes of 1184 healthy individuals from 11 HapMap populations with a wide range of ancestry. The results show that population genetic divergence at the CNV loci is generally more than three times lower than at genome-wide SNP sites. Populations generally exhibit very small genetic divergence (G65=650.0565±650.049). The smallest divergence is among African populations (G65=650.008165±650.0025), with increased divergence among non-African populations (G65=650.021765±650.0109) and then among African and non-African populations (G65=650.032465±650.0064). Genetic diversity is high in African populations (~0.13), low in Asian populations (~0.11), and intermediate in the remaining 11 populations. Few significant linkage disequilibria (LDs) occur between the genome-wide CNV loci. Patterns of gametic and zygotic LDs indicate the absence of epistasis among CNV loci. Mutation rate is about twice as large as the migration rate in the non-African populations, suggesting that the high mutation rates play dominant roles in producing the low population genetic divergence at CNV loci.

Abstract . A genealogical species is defined as a basal group of organisms whose members are all more closely related to each other than they are to any organisms outside the group ("exclusivity'), and which contains no exclusive group within it. In practice, a pair of species is so defined when phylogenies of alleles from a sample of loci shows them to be reciprocally monophyletic at all or some specified fraction of the loci. We investigate the length of time it takes to attain this status when an ancestral population divides into two descendant populations of equal size with no gene exchange, and when genetic drift and mutation are the only evolutionary forces operating. The number of loci used has a substantial effect on the probability of observing reciprocal monophyly at different times after population separation, with very long times needed to observe complete reciprocal monophyly for a large number of loci. In contrast, the number of alleles sampled per locus has a relatively small effect on the probability of reciprocal monophyly. Because a single mitochondrial or chloroplast locus becomes reciprocally monophyletic much faster than does a single nuclear locus, it is not advisable to use mitochondrial and chloroplast DNA to recognize genealogical species for long periods after population divergence. Using a weaker criterion of assigning genealogical species status when more than 50% of sampled nuclear loci show reciprocal monophyly, genealogical species status depends much less on the number of sampled loci, and is attained at roughly 4 7 N generations after populations are isolated, where N is the historically effective population size of each descendant. If genealogical species status is defined as more than 95% of sampled nuclear loci showing reciprocal monophyly, this status is attained after roughly 9 12 N generations.

Phylogenetic relationships of the aquatic moss genus Platyhypnidium and terrestrial species of Rhynchostegium were explored using a phylogeny based on three DNA sequence regions, the nuclear ITS1-5.8S-ITS2, and the plastid trnL-F and trnD-T. This is the first study using trnD-T in the phyogeny of mosses. Platyhypnidium appeared polyphyletic and most of its species were found within a Rhynchostegium clade including species from four different genera: Rhynchostegium, Eriodon, Eurhynchiella and Platyhypnidium. Use of the name Rhynchostegium requires conservation against Eriodon. Within Rhynchostegium s.l., aquatic Rhynchostegium species (i.e., Platyhypnidium) were resolved in four geographically delimited groups: an Afro-European group, an American ast Asiatic group, a Malesian-Australian group and a group comprising the South American P. fuegianum together with Eriodon and Eurhynchiella. Topologies from independent analyses of plastid and nuclear data did not show conflict, and the occurrence of hybridzation events in the clade was therefore not supported. The phylogeny indicates numerous habitat shifts between terrestrial and aquatic habitats, as well as between different terrestrial (epiphytic and epigeic) habitats, which may have affected taxonomic complexity in Rhynchostegium. New morphological characters, such as a striate leaf lamina, axillary hairs composed of relatively short cells, and clearly differentiated cells of the costa, separate Platyhypnidium species within Eurhynchioideae from those in the Oxyrrhynchium-Donrichardsia clade (Helicodontioideae). Two morphologically unique specimens, Platyhypnidium muellerii from Hawaii and Platyhypnidium sp. from Northern Australia, are described as Donrichardsia bartramii and Rhynchostegium brevinerve. The new genus Hedenaesia (Helicodontioideae) is established to accommodate Platyhypnidium austrinum.

Previous phylogeographic studies of alpine plants in Japan have inferred that populations in central Honshu persisted during the Pleistocene climatic oscillations and suggested interglacial survival in high mountains. However, Arcterica nana (Maxim.) Makino (Ericaceae) exhibits a homogenous genetic structure throughout Japan and may therefore have a unique phylogeographic history. This inconsistency could have resulted from insufficient resolution of previously analyzed chloroplast DNA sequences. Therefore, we conducted a phylogeographic investigation based on amplified fragment length polymorphisms. Using 176 individuals from 21 populations, the relationships among individuals and populations were determined by principal coordinate analysis and a neighbor-joining tree, respectively. In addition, genetic differentiation was estimated using analysis of molecular variance and spatial autocorrelation analysis. These analyses demonstrate a homogenous structure throughout the entire Japanese range, supporting the previous cpDNA phylogeography. Although this genetic structure is inconsistent with those of other alpine plants, it is difficult to postulate that pre-existing genetic differentiation was swamped exclusively within A . nana . Therefore, this homogenous genetic structure may have been caused by the distinct history of populations of A. nana . Specifically, the southern-ward migration and the subsequent continuous populations enabled gene flow throughout the Japanese archipelago during the last glacial period. Thus, our data suggest that alpine plants in the Japanese archipelago did not always experience a shared distribution change following climatic oscillations.

Discordant phylogeographical patterns among species with similar distributions may not only denote specific biogeographical histories of different species, but also could represent stochastic variance of genealogies in applied genetic markers. A multilocus investigation representing different genomes can be used to address the latter concern, allowing robust inference to biogeographical history. In the present study, we conducted a multilocus phylogeographical analysis to re-examine the genetic structuring of Phyllodoce nipponica, in which chloroplast (cp)DNA markers exhibited a discordant pattern compared to those of other alpine plants. The geographical structure of sequence variation at five nuclear loci was not consistent with that of cpDNA and showed differentiation between the northern and southern parts of the range of this species. Its demographic history inferred from the isolation-with-migration model suggests that the north090009south divergence originated from Pleistocene vicariance. In addition, the demographic parameters showed a lack of chloroplast-specific gene flow, suggesting that stochastic variance in genealogy resulted in the discordant geographical structure. Thus, P.090009nipponica probably experienced Pleistocene vicariance between its southern and northern range parts in concordance with other alpine plants in the Japanese archipelago. The findings of the present study demonstrates the importance of using a multilocus approach for inferring population dynamics, as well as for reconciling discordant phylogeographical patterns among species. 0008 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110, 214090009226.

Members of the phlox family (Polemoniaceae) serve as useful models for studying various evolutionary and biological processes. Despite its biological importance, no family-wide phylogenetic estimate based on multiple DNA regions with complete generic sampling is available. Here, we analyze one nuclear and five chloroplast DNA sequence regions (nuclear ITS, chloroplast , intron plus trnL–trnF intergeneric spacer, and the trnS–trnG, trnD–trnT, and psbM–trnD intergenic spacers) using parsimony and Bayesian methods, as well as assessments of congruence and long branch attraction, to explore phylogenetic relationships among 84 ingroup species representing all currently recognized Polemoniaceae genera. Relationships inferred from the ITS and concatenated chloroplast regions are similar overall. A combined analysis provides strong support for the monophyly of Polemoniaceae and subfamilies Acanthogilioideae, Cobaeoideae, and Polemonioideae. Relationships among subfamilies, and thus for the precise root of Polemoniaceae, remain poorly supported. Within the largest subfamily, Polemonioideae, four clades corresponding to tribes Polemonieae, Phlocideae, Gilieae, and Loeselieae receive strong support. The monogeneric Polemonieae appears sister to Phlocideae. Relationships within Polemonieae, Phlocideae, and Gilieae are mostly consistent between analyses and data permutations. Many relationships within Loeselieae remain uncertain. Overall, inferred phylogenetic relationships support a higher-level classification for Polemoniaceae proposed in 2000.

Relationships within the Polygonaceae have been recently examined using rbcL sequences, with an emphasis on Polygonum and its segregates. Here we test these results with respect to Polygonum (sensu lato) with an expanded dataset, including additional species and gene regions. Specifically, we focus on inferring the relationships of Eupersicaria (Polygonum sect. Persicaria in many prior treatments), using the chloroplast genes rbcL, trnL090009F, trnK intron090009matK, and psbA090009trnH IGS, and nuclear ribosomal ITS sequences. We conclude that Eupersicaria is monophyletic and most closely related to Tovara and Echinocaulon. In turn, this clade is most closely related to Cephalophilon. The sister group of this entire Persicaria clade contains Bistorta and a clade including Aconogonon and Koenigia, which supports the monophyly of the Persicarieae. Within Eupersicaria there appears to be a deep split between P. amphibia and the remaining species, and there is strong conflict regarding the placement of P. punctata. These results set the stage for a more detailed phylogenetic analysis of Eupersicaria.

The opposite modes of chloroplast DNA (cpDNA) inheritance were found to operate in the reciprocal crossings of Scots pine (Pinus sylvestris L.) and mountain dwarf pine (Pinus mugo Turra). The crossing

The distribution of a rare chloroplast-DNA structural mutation, the loss of a large inverted repeat, has been determined for 95 species representing 77 genera and 25 of the 31 tribes in the legume subfamily Papilionoideae. This mutation, which is regarded as a derived feature of singular origin within the subfamily, marks a group comprising six temperate tribes, the Galegeae, Hedysareae, Carmichaelieae, Vicieae, Cicereae, and Trifolieae, an assemblage traditionally considered to be monophyletic. This mutation also occurs in the chloroplast genome of Wisteria, a member of the tropical tribe Millettieae whose other members so far surveyed lack the mutation. These new DNA data, together with traditional evidence, support the hypothesis that Wisteria is an unspecialized member of a lineage that gave rise to the temperate tribes marked by the chloroplast-DNA mutation; the probable paraphylesis of Millettieae is revealed. Two other tribes, Loteae and Coronilleae (traditionally regarded as a derived element of the aforesaid temperate tribes) do not possess this chloroplast-DNA structural mutation and, therefore, presumably represent a distinct temperate lineage. This hypothesis is supported by additional evidence from pollen, inflorescence, and root-nodule morphology that suggests that the Loteae and Coronilleae share a more recent ancestry with tropical tribes such as Phaseoleae and Millettieae than with other temperate tribes.

The drivers of species diversification and persistence are of great interest to current biogeography, especially in those global biodiversity otspots harbouring most of Earth's animal and plant life. Classical multispecies biogeographical work has yielded fascinating insights into broad-scale patterns of diversification, and DNA-based intraspecific phylogeographical studies have started to complement this picture at much finer temporal and spatial scales. The advent of novel next-generation sequencing (NGS) technologies provides the opportunity to greatly scale up the numbers of individuals, populations and species sampled, potentially merging intraspecific and interspecific approaches to biogeographical inference. Here, we outline these prospects and issues by using the example of an undisputed hotspot, the Cape of southern Africa. We outline the current state of knowledge on the biogeography of species diversification within the Cape, review the literature for phylogeographical evidence of its likely drivers and mechanisms, and suggest possible ways forward based on NGS approaches. We demonstrate the potential of these methods and current bioinformatic issues with the help of restriction-site-associated DNA (RAD) sequencing data for three highly divergent species of the Restionaceae, an important plant radiation in the Cape. A thorough understanding of the mechanisms that facilitate species diversification and persistence in spatially structured, species-rich environments will require the adoption of novel genomic and bioinformatic tools in biogeographical studies.

物种形成过程是生物多样性形成的基础,长期以来一直是进化生物学的中心议题之一。传统的异域物种形成理论认为,地理隔离是物种分化的主要决定因子,物种形成只有在种群之间存在地理隔离的情况下才能发生。近年来,随着种群基因组学的发展和溯祖理论分析方法的完善,种群间存在基因流情况下的物种形成成为进化生物学领域新的研究焦点。物种形成过程中是否有基因流的发生?基因流如何影响物种的形成与分化?基因流存在条件下物种形成的生殖隔离机制是什么?根据已发表的相关文献资料,作者综述了当前物种形成研究中基因流的时间和空间分布模式、基因流对物种分化的影响以及生殖隔离机制形成等问题,指出基因流存在条件下的物种形成可能是自然界普遍发生的一种模式。

物种形成过程是生物多样性形成的基础,长期以来一直是进化生物学的中心议题之一。传统的异域物种形成理论认为,地理隔离是物种分化的主要决定因子,物种形成只有在种群之间存在地理隔离的情况下才能发生。近年来,随着种群基因组学的发展和溯祖理论分析方法的完善,种群间存在基因流情况下的物种形成成为进化生物学领域新的研究焦点。物种形成过程中是否有基因流的发生?基因流如何影响物种的形成与分化?基因流存在条件下物种形成的生殖隔离机制是什么?根据已发表的相关文献资料,作者综述了当前物种形成研究中基因流的时间和空间分布模式、基因流对物种分化的影响以及生殖隔离机制形成等问题,指出基因流存在条件下的物种形成可能是自然界普遍发生的一种模式。

生物自然居群间的基因流不但可以阻止遗传分化以维持物种的完整性，而且也能积极响应生物多样 化的进程。理解与基因流相关的适应性进化及其内在机理将有助于我们更好地认识生物物种形成和多样化的原始动力以及真正原因。该文通过对植物种内和种间居群 基因流动态进行讨论，阐述了近年来有关植物基因流动态的一些重要理论观念和研究进展，以期为相关领域动态及趋势研究提供参考。

生物自然居群间的基因流不但可以阻止遗传分化以维持物种的完整性，而且也能积极响应生物多样 化的进程。理解与基因流相关的适应性进化及其内在机理将有助于我们更好地认识生物物种形成和多样化的原始动力以及真正原因。该文通过对植物种内和种间居群 基因流动态进行讨论，阐述了近年来有关植物基因流动态的一些重要理论观念和研究进展，以期为相关领域动态及趋势研究提供参考。

Chloroplasts were once free-living cyanobacteria that became endosymbionts, but the genomes of contemporary plastids encode only ≈5-10% as many genes as those of their free-living cousins, indicating that many genes were either lost from plastids or transferred to the nucleus during the course of plant evolution. Previous estimates have suggested that between 800 and perhaps as many as 2,000 genes in the Arabidopsis genome might come from cyanobacteria, but genome-wide phylogenetic surveys that could provide direct estimates of this number are lacking. We compared 24,990 proteins encoded in the Arabidopsis genome to the proteins from three cyanobacterial genomes, 16 other prokaryotic reference genomes, and yeast. Of 9,368 Arabidopsis proteins sufficiently conserved for primary sequence comparison, 866 detected homologues only among cyanobacteria and 834 other branched with cyanobacterial homologues in phylogenetic trees. Extrapolating from these conserved proteins to the whole genome, the data suggest that ≈4,500 of Arabidopsis protein-coding genes (≈18% of the total) were acquired from the cyanobacterial ancestor of plastids. These proteins encompass all functional classes, and the majority of them are targeted to cell compartments other than the chloroplast. Analysis of 15 sequenced chloroplast genomes revealed 117 nuclear-encoded proteins that are also still present in at least one chloroplast genome. A phylogeny of chloroplast genomes inferred from 41 proteins and 8,303 amino acids sites indicates that at least two independent secondary endosymbiotic events have occurred involving red algae and that amino acid composition bias in chloroplast proteins strongly affects plastid genome phylogeny.

Abstract Hybridization between species is known to greatly affect their genetic diversity and, therefore, their evolution. Also, within species, there may be genetic clusters between which gene flow is limited, which may impact natural selection. However, few studies have looked simultaneously at the influence of among-species and within-species gene flow. Here, we study the influence of hybridization between Populus balsamifera and P. trichocarpa on population structure and adaptation in P. balsamifera. We did this by sampling a total of 1,517 individuals from across the ranges of these two species, and by genotyping them using a combination of 93 nuclear and 17 cpDNA SNPs. We found that hybridization is mostly limited to the contact zone where the species' distributions overlap. Within P. balsamifera, we found multiple levels of population structure. Interestingly, the border between the Eastern and Central clusters is very sharp, while the border between the Central and Western clusters is diffuse. Outlier analysis revealed that three loci associated with the sharp border were also associated with climate. We hypothesize that the observed clusters derive from three refugia during the Pleistocene ice ages. Between the Central and Western clusters, postglacial long-distance gene flow has led to the diffusion of their border. In the Eastern cluster, we hypothesize that endogenous genomic barriers have developed, leading to the sharp border and a spurious climate association. We conclude that the large-scale genetic structure of P. balsamifera is mostly shaped by historical factors and the influence of hybridization is limited. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.

由于青藏高原的地理效应,第四纪冰期气候的反复变化应对现今该地区生物的地理分布及其居群遗传结构产生重大影响。肋果沙棘Hippophae neurocarpa是青藏高原地区的一个特有种,根据叶上的附属物(星状鳞毛或者鳞片状鳞毛)分为两个亚种:肋果沙棘亚种subsp.neurocarpa和密毛肋果沙棘亚种subsp.stellatopilosa。依据母系遗传的叶绿体DNA片段对该物种谱系地理学进行研究不仅能阐明该物种冰期避难所,而且对于理解两个亚种的母系分化也具有重要意义。共对两个亚种14居群的70个个体的trnL-F序列进行了测序,共发现8种单倍型,其中5种单倍型出现在肋果沙棘亚种中,4种出现在密毛肋果沙棘亚种中,两个亚种共享一种单倍型。种内谱系分化与两个亚种形态上的分化不一致。嵌套进化分支把8种单倍型分为三支:一支为肋果沙棘亚种,其他两支中两个亚种单倍型嵌套组成,且肋果沙棘亚种处于进化末端。我们的研究结果还表明肋果沙棘在冰期可能在高海拔地区存在多个避难所,并且存在来自避难所的邻域扩张。

由于青藏高原的地理效应,第四纪冰期气候的反复变化应对现今该地区生物的地理分布及其居群遗传结构产生重大影响。肋果沙棘Hippophae neurocarpa是青藏高原地区的一个特有种,根据叶上的附属物(星状鳞毛或者鳞片状鳞毛)分为两个亚种:肋果沙棘亚种subsp.neurocarpa和密毛肋果沙棘亚种subsp.stellatopilosa。依据母系遗传的叶绿体DNA片段对该物种谱系地理学进行研究不仅能阐明该物种冰期避难所,而且对于理解两个亚种的母系分化也具有重要意义。共对两个亚种14居群的70个个体的trnL-F序列进行了测序,共发现8种单倍型,其中5种单倍型出现在肋果沙棘亚种中,4种出现在密毛肋果沙棘亚种中,两个亚种共享一种单倍型。种内谱系分化与两个亚种形态上的分化不一致。嵌套进化分支把8种单倍型分为三支:一支为肋果沙棘亚种,其他两支中两个亚种单倍型嵌套组成,且肋果沙棘亚种处于进化末端。我们的研究结果还表明肋果沙棘在冰期可能在高海拔地区存在多个避难所,并且存在来自避难所的邻域扩张。

Annu Rev Genet. 1985;19:325-54. Comparative Study; Review

Coalescence theory predicts when genetic drift at nuclear loci will result in fixation of sequence differences to produce monophyletic gene trees. However, the theory is difficult to apply to particular taxa because it hinges on genetically effective population size, which is generally unknown. Neutral theory also predicts that evolution of monophyly will be four times slower in nuclear than in mitochondrial genes primarily because genetic drift is slower at nuclear loci. Variation in mitochondrial DNA (mtDNA) within and between species has been studied extensively, but can these mtDNA data be used to predict coalescence in nuclear loci? Comparison of neutral theories of coalescence of mitochondrial and nuclear loci suggests a simple rule of thumb. The "three-times rule" states that, on average, most nuclear loci will be monophyletic when the branch length leading to the mtDNA sequences of a species is three times longer than the average mtDNA sequence diversity observed within that species. A test using mitochondrial and nuclear intron data from seven species of whales and dolphins suggests general agreement with predictions of the three-times rule. We define the coalescence ratio as the mitochondrial branch length for a species divided by intraspecific mtDNA diversity. We show that species with high coalescence ratios show nuclear monophyly, whereas species with low ratios have polyphyletic nuclear gene trees. As expected, species with intermediate coalescence ratios show a variety of patterns. Especially at very high or low coalescence ratios, the three- times rule predicts nuclear gene patterns that can help detect the action of selection. The three-times rule may be useful as an empirical benchmark for evaluating evolutionary processes occurring at multiple loci.

The chloroplast genomes of some species of legumes lack the large inverted repeat (IR) that is a trademark of most land-plant chloroplasts. Our analysis of chloroplast genes in legume species that have an IR shows that the synonymous (silent) substitution rate in IR genes is 2.3-fold lower than in single-copy (SC) genes, which is largely in agreement with earlier findings. Given that all genes in species that lack the IR are single-copy, what level of synonymous substitution exists in these genes? We report a uniform substitution rate in IR-less genomes, and moreover, we find this rate to be at the level otherwise reserved for SC genes. In other words, the synonymous substitution rate has accelerated in the remaining copy of the duplicate region. We propose that this acceleration is a direct result of the decrease in the copy number of the sequence, rather than an intrinsic property of the genes normally located in the IR.

The genus Ipomopsis (Polemoniaceae) encompasses about 29 species and 24 subspecies generally divided into three sections: sect. Ipomopsis, sect. Microgilia, and sect. Phloganthea. We employed maximum likelihood and Bayesian inference of DNA sequences from the nuclear ribosomal ITS region (ITS1, 5.8S ribosomal subunit, ITS2) and the chloroplast trnL090009F region (trnL intron + trnL090009trnF intergenic spacer) to estimate phylogenetic relationships within this genus and its placement among other genera of Polemoniaceae. The chloroplast and combined sequences provide support for the monophyly of Ipomopsis, but only if four species previously included in the genus are removed: Ipomopsis havardii, I. sonorae, Microgilia minutiflora (= I. minutiflora), and Loeseliastrum depressum (= I. depressa). Of the three sections, two are conditionally supported as being monophyletic. Section Microgilia (with 11 species and 11 infra-specific taxa) is supported as monophyletic if I. polycladon, I. sonorae, I. depressa, and I. minutiflora (the type of the section) are removed. This clade is treated here as section Elaphocera. Section Ipomopsis is inferred to be monophyletic with the inclusion of several members of sect. Phloganthea (I. multiflora, I. pinnata, and I. polyantha). There is no support for monophyly or paraphyly of sect. Phloganthea. The Giliopsis group (I. effusa, I. guttata, and I. tenuifolia) is supported as monophyletic by both data sets, and the cpDNA sequences place it as sister to the remainder of Ipomopsis. This clade is treated as a new section, Giliopsis. Nuclear data place Giliopsis in a clade with Ipomopsis havardii, I. sonorae, Microgilia minutiflora, Loeseliastrum depressum, Eriastrum spp., Langloisia, and Dayia grantii. Using the Eocene fossil Gilisenium hueberii to calibrate the most recent common ancestor of tribe Gilieae, we estimate that Ipomopsis has its origin 28.2 00± 0. 4009000939.0 00± 1.14 MYA (trnL090009F and ITS, respectively). Using this same relaxed clock, the node (or coalescent event) that defines the I. aggregata complex is dated at 16.2 00± 0.38 and 27.1 00± 0.83 MYA (trnL090009F and ITS, respectively). The deep divergence of the I. aggregata complex suggests that reticulation, rather than lineage sorting, is the source of conflict among phylogenetic markers used to infer the placement of I. macrosiphon.

The nonrecombinant, uniparentally inherited nature of organelle genomes makes them useful tools for evolutionary studies. However, in plants, detecting useful polymorphism at the population level is often difficult because of the low level of substitutions in the chloroplast genome, and because of the slow substitution rates and intramolecular recombination of mtDNA. Chloroplast microsatellites represent potentially useful markers to circumvent this problem and, to date, studies have demonstrated high levels of intraspecific variability. Here, we discuss the use of these markers in ecological and evolutionary studies of plants, as well as highlighting some of the potential problems associated with such use.

The Sino-Japanese Floristic Region(SJFR) of East Asia harbors the most diverse of the world's temperate flora,and was the most important glacial refuge for its Tertiary representatives('relics') throughout Quaternary ice-age cycles.A steadily increasing number of phylogeographic studies in the SJFR of mainland China and adjacent areas,including the Qinghai-Tibetan-Plateau(QTP) and Sino-Himalayan region,have documented the population histories of temperate plant species in these regions.Here we review this current literature that challenges the oft-stated view of the SJFR as a glacial

Molecular phylogenetic reconstructions have indicated that Haworthia is not monophyletic. Here we show, using considerably expanded datasets of both chloroplast (trnL-trnF and psbA-trnH intergenic spacers) and nuclear (ITS1) markers that the issue of a polyphyletic Haworthia is more complicated than previously reported. Both parsimony and bayesian analyses of cpDNA and ITS1 produced poorly resolved phylogenies, with little or no support for deeper nodes. Species of Haworthia are placed in three of the four main lineages in the cpDNA phylogeny, while in the ITS1 phylogeny Haworthia species were placed in two of the four main lineages retrieved. The cpDNA phylogeny was incongruent with that obtained from the ITS1 data, and the topology of the combined dataset reflected that obtained from the ITS1 dataset. As species of H. subg. Hexangulares and subg. Robustipedunculares have been reportedly involved in intergeneric hybridizations with other Alooideae, it is postulated that a history of ancient hybridization may account for a polyphyletic Haworthia. Hybridization is also ongoing between species within H. subg. Haworthia, and may be responsible for results indicating that some species in this subgenus are not monophyletic. This hybridization may be facilitated by the lack of effective reproductive barriers. We hypothesize that Haworthia is undergoing a contemporary and explosive radiation in southern Africa, driven in part by local adaptation to relatively fine-grained ecological heterogeneity of soils and microclimate.

Background Hybridization events are relatively common in vascular plants. However, the frequency of these events is unevenly distributed across the plant phylogeny. Plant families in which individual species are pollinated by specific pollinator species are predicted to be less prone to hybridization than other families. However, exceptions may occur within these families, when pollinators shift host-plant species. Indeed, host shifts are expected to increase the rate of hybridization events. Pollinators of Ficus section Galoglychia are suspected to have changed host repeatedly, based on several cases of incongruence between plant phylogeny and taxonomy, and insect phylogeny and taxonomy. We tracked cyto-nuclear discordance across section Galoglychia as evidence for hybridization. To achieve a proper global view, we first clarified the monophyly of section Galoglychia as it had been questioned by recent phylogenetic studies. Moreover, we investigated if fig size could be a factor facilitating host shifts. Results Phylogenetic chloroplast and nuclear results demonstrated the monophyly of section Galoglychia. Within section Galoglychia, we detected several cases of statistically significant cyto-nuclear discordance. Discordances concern both terminal nodes of the phylogenetic trees and one deep node defining relationships between subsections. Because nuclear phylogeny is congruent with morphological taxonomy, discordances were caused by the chloroplast phylogeny. Introgressive hybridization was the most likely explanation for these discordances. We also detected that subsections pollinated by several wasp genera had smaller figs and were pollinated by smaller wasps than subsections pollinated by a single wasp genus. Conclusion As hypothesized, we discovered evidences of past hybridization in Ficus section Galoglychia. Further, introgression was only detected in subsections presenting incongruence between plant and pollinator phylogenies and taxonomy. This supports the hypothesis that host shift is the cause for plant-pollinator incongruence. Moreover, small fig size could facilitate host shifts. Eventually, this study demonstrates that non-coding chloroplast markers are valuable to resolve deep nodes in Ficus phylogeny.

The genealogies of samples of orthologous regions from multiple species can be classified by their shapes. Using a neutral coalescent model of two species, I give exact probabilities of each of four possible genealogical shapes: reciprocal monophyly, two types of paraphyly, and polyphyly. After the divergence that forms two species, each of which has population size N, polyphyly is the most likely genealogical shape for the lineages of the two species. At 651.300N generations after divergence, paraphyly becomes most likely, and reciprocal monophyly becomes most likely at 651.665N generations. For a given species, the time at which 99% of its loci acquire monophyletic genealogies is 655.298N generations, assuming all loci in its sister species are monophyletic. The probability that all lineages of two species are reciprocally monophyletic given that a sample from the two species has a reciprocally monophyletic genealogy increases rapidly with sample size, as does the probability that the most recent common ancestor (MRCA) for a sample is also the MRCA for all lineages from the two species. The results have potential applications for the testing of evolutionary hypotheses.

Abstract Three methods for estimating the average level of gene flow in natural population are discussed and compared. The three methods are FST, rare alleles, and maximum likelihood. All three methods yield estimates of the combination of parameters (the number of migrants [Nm] in a demic model or the neighborhood size [4 D 2] in a continuum model) that determines the relative importance of gene flow and genetic drift. We review the theory underlying these methods and derive new analytic results for the expectation of FST in stepping-stone and continuum models when small sets of samples are taken. We also compare the effectiveness of the different methods using a variety of simulated data. We found that the FST and rare-alleles methods yield comparable estimates under a wide variety of conditions when the population being sampled is demographically stable. They are roughly equally sensitive to selection and to variation in population structure, and they approach their equilibrium values at approximately the same rate. We found that two different maximum-likelihood methods tend to yield biased estimates when relatively small numbers of locations are sampled but more accurate estimates when larger numbers are sampled. Our conclusion is that, although FST and rare-alleles methods are expected to be equally effective in analyzing ideal data, practical problems in estimating the frequencies of rare alleles in electrophoretic studies suggest that FST is likely to be more useful under realistic conditions.

Speciation is an ongoing process. Many recognized species are fully divergent from each other and their ancestors, whereas others are in earlier stages in the diversification process. Such incipient speciation may create patterns when one or a few populations are phenotypically distinct, but lack genomic level coalescence from each other or from their ancestral species. As a result, such progenitor-derivative species pairs are likely to lack reciprocal monophyly or generate paraphyletic ancestral species. Here we examine phylogenetic patterns in the Columnea strigosa (Gesneriaceae) complex to evaluate whether populations that have been named C. kucyniakii are reciprocally monophyletic with C. strigosa, its presumed ancestral species. Molecular phylogenetic results do not support reciprocal monophyly of the two species, implying that incipient speciation is occurring within the C. strigosa complex. We hereby recommend that C. kucyniakii be recognized at the specific rank despite the fact that it creates a paraphyletic C. strigosa. These findings bear importance in taxonomic decisions about paraphyletic taxa and recognizing evolutionary and morphologically distinct lineages.

Eukaryotic cells arose through endosymbiotic uptake of free-living bacteria followed by massive gene transfer from the genome of the endosymbiont to the host nuclear genome. Because this gene transfer took place over a time scale of hundreds of millions of years, direct observation and analysis of primary transfer events has remained difficult. Hence, very little is known about the evolutionary frequency of gene transfer events, the size of transferred genome fragments, the molecular mechanisms of the transfer process, or the environmental conditions favoring its occurrence. We describe here a genetic system based on transgenic chloroplasts carrying a nuclear selectable marker gene that allows the efficient selection of plants with a nuclear genome that carries pieces transferred from the chloroplast genome. We can select such gene transfer events from a surprisingly small population of plant cells, indicating that the escape of genetic material from the chloroplast to the nuclear genome occurs much more frequently than generally believed and thus may contribute significantly to intraspecific and intraorganismic genetic variation.

Chloroplast genomes of Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] and radiata (Monterey) pine [Pinus radiata D. Don], two conifers from the widespread Pinaceae, were mapped and their genomes were compared to other land plants. Douglas-fir and radiata pine lack the large (20-25 kilobases) inverted repeat that characterizes most land plants. To our knowledge, this is only the second recorded loss of this ancient and highly conserved inverted repeat among all lineages of land plants thus far examined. Loss of the repeat largely accounts for the small size of the conifer genome, 120 kilobases, versus 140-160 kilobases in most land plants. Douglas-fir possesses a major inversion of 40-50 kilobases relative to radiata pine and nonconiferous plants. Nucleotide sequence differentiation between Douglas-fir and radiata pine was estimated to be 3.8%. Both conifer genomes possess a number of rearrangements relative to Osmunda, a fern, Ginkgo, a gymnosperm, and Petunia, an angiosperm. Among land plants, structural changes of this degree have occurred primarily within tribes of the legume family (Fabaceae) that have also lost the inverted repeat. These results support the hypothesis that the presence of the large inverted repeat stabilizes the chloroplast genome against major structural rearrangements.

Abstract With the aim of analyzing and interpreting data on DNA polymorphism obtained by DNA sequencing or restriction enzyme technique, a mathematical theory on the expected evolutionary relationship among DNA sequences (nucleons) sampled is developed under the assumption that the evolutionary change of nucleons is determined solely by mutation and random genetic drift. The statistical property of the number of nucleotide differences between randomly chosen nucleons and that of heterozygosity or nucleon diversity is investigated using this theory. These studies indicate that the estimates of the average number of nucleotide differences and nucleon diversity have a large variance, and a large part of this variance is due to stochastic factors. Therefore, increasing sample size does not help reduce the variance significantly The distribution of sample allele (nucleomorph) frequencies is also studied, and it is shown that a small number of samples are sufficient in order to know the distribution pattern.

Phylogeographic inference can be a powerful tool in reconstructing species evolutionary histories; however, although inferred phylogeographic patterns should depend in part on the underlying types and rates of mutations, the effects of different types of mutations have seldom been quantified. In this study we identified two chloroplast minisatellites in the common reed Phragmites australis, and showed that these are more variable than chloroplast microsatellites. We then recreated parsimony networks of the global phylogeography of P. australis based on data that either included or excluded repetitive sequences (minisatellites and microsatellites), thereby illustrating the influence that these repetitive sequences can have on large-scale phylogeographic inference. The resulting networks differed in the numbers of mutational steps, degrees of uncertainty, and total numbers of haplotypes. In addition, the suggested ancestor-descendant relationships among lineages changed substantially depending on whether repetitive sequences were included. We therefore caution against the inclusion of repetitive sequences in large-scale networks because of their high potential for homoplasy. Nevertheless, we advocate the inclusion of repetitive sequences in other analyses: specifically, we show that the ratio of mutations in repetitive vs. non-repetitive regions can provide insight into the relative ages of lineages.

Chloroplast ( trnT ) and nuclear rDNA (ITS) sequence analyses of the Araliaceae provide strong molecular evidence for the monophyly of the genus Hedera. Phylogenetic reconstructions suggest multiple origins and an active polyploidization process not only in the formation of tetraploids (2 n=96), hexaploids (2 n=144), and octoploids (2 n=192), but also of diploids (2 n=48). A high basic chromosome number of x=24, extensive polyphyly in widespread diploids, and terminal placement of Hedera in phylogenies of the Araliaceae reveal that extant diploid taxa may be, in fact, assemblages of ancestral polyploids from plants of n=12. Four major lineages containing four types of chloroplast (chlorotypes I, II, III, and IV), which are defined by different trnT nucleotide substitutions and two large insertions (50- and 30-bp), provide evidence for evolutionary processes and historical biogeography in Hedera. We propose a scenario where an initial colonization in the Mediterranean basin by Asian ancestors (carrying the ancestral Araliaceae chlorotype I) is followed by differentiation into the four chlorotypes of the Mediterranean region, and then recolonization of Asia and northern Europe only by chlorotype III. The Macaronesian taxa ( Hedera azorica, Hedera maderensis ssp. maderensis, and Hedera canariensis) appear to have originated from a single-colonization event to each archipelago with no further contact either with continental or insular species.

In the current study,phylogeographic structure of L-F and rps15-ycf1).Three hundreds and twenty four individuals distributed in 23 natural populations were used in the study covering the QTP and adjacent highlands.The result showed that:(1)The closely related haplotypes in this study tending to occur in same population indicated significantly phylogeographic pattern within the entire distribution range.(2)About 71.4% of the 35 haplotypes detected from all individuals were endemic in certain population;phylogenetic analysis divided all the haplotypes into three highly supported dades,in which H1 was the more ancient and highly distributed haploptype.(3)Mismatch distribution analysis indicated the species had experienced demographic expansion up to some extent since long time,but the expansion was not so rapid.(4)The results of BEAST analysis revealed that cladistics differentiation of S.mongolica occurred about 45 Mya.This research suggested that there are no less than three glacial refugia of S.mongolica in the QTP and adjacent highlands.The distribution of S.mongolica on the QTP and adjacent highlands was due to the combine effects of climatic instability during quaternary glacial-interglacial episodes and the uplifting of QTP.

In the current study,phylogeographic structure of L-F and rps15-ycf1).Three hundreds and twenty four individuals distributed in 23 natural populations were used in the study covering the QTP and adjacent highlands.The result showed that:(1)The closely related haplotypes in this study tending to occur in same population indicated significantly phylogeographic pattern within the entire distribution range.(2)About 71.4% of the 35 haplotypes detected from all individuals were endemic in certain population;phylogenetic analysis divided all the haplotypes into three highly supported dades,in which H1 was the more ancient and highly distributed haploptype.(3)Mismatch distribution analysis indicated the species had experienced demographic expansion up to some extent since long time,but the expansion was not so rapid.(4)The results of BEAST analysis revealed that cladistics differentiation of S.mongolica occurred about 45 Mya.This research suggested that there are no less than three glacial refugia of S.mongolica in the QTP and adjacent highlands.The distribution of S.mongolica on the QTP and adjacent highlands was due to the combine effects of climatic instability during quaternary glacial-interglacial episodes and the uplifting of QTP.

以群落学方法和谱系地理学方法,分析广东省南昆山与大岭山珍稀孑遗植物伯乐树种群之间的关系,揭示它们的遗传结构并比较所处群落的异质性。群落分析发现,南昆山和大岭山伯乐树所在群落无论在区系成分上还是在物种组成上均表现出极大相似性,同时也稍有差异:大岭山伯乐树所在群落的物种多样性不及南昆山;两群落内伯乐树种群龄级结构均不完整,而南昆山伯乐树种群相对表现出更强的生活力。谱系地理分析发现,南昆山和大岭山伯乐树种群的单倍型多样性(hd)分别为0.682和0,核苷酸多样性(π)分别为0.181×10-3和0,两种群间的遗传分化系数GST=0.167,基因交流系数Nm=2.49。研究表明,南昆山和大岭山伯乐树种群为复合种群,南昆山伯乐树种群起源可能更为古老。南昆山伯乐树种群单倍型多样性和核苷酸多样性较之大岭山伯乐树种群为高,更具保护价值。

以群落学方法和谱系地理学方法,分析广东省南昆山与大岭山珍稀孑遗植物伯乐树种群之间的关系,揭示它们的遗传结构并比较所处群落的异质性。群落分析发现,南昆山和大岭山伯乐树所在群落无论在区系成分上还是在物种组成上均表现出极大相似性,同时也稍有差异:大岭山伯乐树所在群落的物种多样性不及南昆山;两群落内伯乐树种群龄级结构均不完整,而南昆山伯乐树种群相对表现出更强的生活力。谱系地理分析发现,南昆山和大岭山伯乐树种群的单倍型多样性(hd)分别为0.682和0,核苷酸多样性(π)分别为0.181×10-3和0,两种群间的遗传分化系数GST=0.167,基因交流系数Nm=2.49。研究表明,南昆山和大岭山伯乐树种群为复合种群,南昆山伯乐树种群起源可能更为古老。南昆山伯乐树种群单倍型多样性和核苷酸多样性较之大岭山伯乐树种群为高,更具保护价值。

绵参(Eriophyton wallichii)是中国青藏高原地区高山流石滩特有植物.为探讨第四纪冰期气候波动对青藏高原高山流石滩植物居群遗传结构以及谱系历史的影响,检测了绵参20个居群187个个体的核糖体内转录间隔区(ITS)的序列变异,共发现19个单倍型,且结果表明大部分居群拥有独自的单倍型.AMOVA分析表明,居群间的变异占了总变异量的89.54％,居群遗传分化指数很高(GST=0.863,NST=0.957),但无明显的谱系结构.我们推测这种独特的谱系地理结构可能主要是由于高山流石滩独特的地形和环境及“孤岛效应”形成的,与先前研究过的青藏高原植物不同,绵参呈现出高度分化的遗传结构,常常呈现出“一个居群,一种单倍型”的模式,这可能是由于第四纪冰期及间冰期尽管绵参的分布范围有扩大和收缩,但始终由于特殊的高山生境和流石滩基质,导致了该物种长期以来居群间很少有基因交流并一直处于隔离状态,造成了现在这样的谱系地理结构.

绵参(Eriophyton wallichii)是中国青藏高原地区高山流石滩特有植物.为探讨第四纪冰期气候波动对青藏高原高山流石滩植物居群遗传结构以及谱系历史的影响,检测了绵参20个居群187个个体的核糖体内转录间隔区(ITS)的序列变异,共发现19个单倍型,且结果表明大部分居群拥有独自的单倍型.AMOVA分析表明,居群间的变异占了总变异量的89.54％,居群遗传分化指数很高(GST=0.863,NST=0.957),但无明显的谱系结构.我们推测这种独特的谱系地理结构可能主要是由于高山流石滩独特的地形和环境及“孤岛效应”形成的,与先前研究过的青藏高原植物不同,绵参呈现出高度分化的遗传结构,常常呈现出“一个居群,一种单倍型”的模式,这可能是由于第四纪冰期及间冰期尽管绵参的分布范围有扩大和收缩,但始终由于特殊的高山生境和流石滩基质,导致了该物种长期以来居群间很少有基因交流并一直处于隔离状态,造成了现在这样的谱系地理结构.

物种自然杂交现象在自然界中普遍存在,并被认为在适宜条件下可以多地独立发生.2013年,在云南元江采集的7种铁线蕨属植物中,其中一种在形态上疑似台湾产梅山铁线蕨,而该种被认为是以假鞭叶铁线蕨为母本、半月形铁线蕨为父本的自然杂交物种.形态学比较显示,该存疑种与铁线蕨属其他种类均有明显差别,用单拷贝核DNA序列CRY2exon 3和叶绿体DNA序列rbc L,atp B,atp A,trn L-F,rps4-trn S对云南元江产梅山铁线蕨存疑种进行研究,结果显示云南元江产梅山铁线蕨存疑种为以苍山铁线蕨为母本和以孟连铁线蕨为父本的自然杂交种,由于形态上与台湾产梅山铁线蕨相似、父母本亲缘关系上与梅山铁线蕨的双亲近缘,因而可称为梅山铁线蕨的隐性杂交种.据此认为,中国可能存在大量的自然杂交种尚未发现,并可以通过核DNA序列和叶绿体DNA序列进行准确鉴定;同时,诸如铁线蕨属这样的蕨类植物,由于存在无性繁殖习性,可以弥补有性生殖隔离带来的适合度降低的缺陷,增加杂交种在自然界中的存活机会,是蕨类植物中自然杂交种普遍存在的重要原因.

物种自然杂交现象在自然界中普遍存在,并被认为在适宜条件下可以多地独立发生.2013年,在云南元江采集的7种铁线蕨属植物中,其中一种在形态上疑似台湾产梅山铁线蕨,而该种被认为是以假鞭叶铁线蕨为母本、半月形铁线蕨为父本的自然杂交物种.形态学比较显示,该存疑种与铁线蕨属其他种类均有明显差别,用单拷贝核DNA序列CRY2exon 3和叶绿体DNA序列rbc L,atp B,atp A,trn L-F,rps4-trn S对云南元江产梅山铁线蕨存疑种进行研究,结果显示云南元江产梅山铁线蕨存疑种为以苍山铁线蕨为母本和以孟连铁线蕨为父本的自然杂交种,由于形态上与台湾产梅山铁线蕨相似、父母本亲缘关系上与梅山铁线蕨的双亲近缘,因而可称为梅山铁线蕨的隐性杂交种.据此认为,中国可能存在大量的自然杂交种尚未发现,并可以通过核DNA序列和叶绿体DNA序列进行准确鉴定;同时,诸如铁线蕨属这样的蕨类植物,由于存在无性繁殖习性,可以弥补有性生殖隔离带来的适合度降低的缺陷,增加杂交种在自然界中的存活机会,是蕨类植物中自然杂交种普遍存在的重要原因.

Abstract The tenet that ecological opportunity drives adaptive diversification has been central to theories of speciation since Darwin, yet no widely accepted definition or mechanistic framework for the concept currently exists. We propose a definition for ecological opportunity that provides an explicit mechanism for its action. In our formulation, ecological opportunity refers to environmental conditions that both permit the persistence of a lineage within a community, as well as generate divergent natural selection within that lineage. Thus, ecological opportunity arises from two fundamental elements: (1) niche availability, the ability of a population with a phenotype previously absent from a community to persist within that community and (2) niche discordance, the diversifying selection generated by the adaptive mismatch between a population's niche-related traits and the newly encountered ecological conditions. Evolutionary response to ecological opportunity is primarily governed by (1) spatiotemporal structure of ecological opportunity, which influences dynamics of selection and development of reproductive isolation and (2) diversification potential, the biological properties of a lineage that determine its capacity to diversify. Diversification under ecological opportunity proceeds as an increase in niche breadth, development of intraspecific ecotypes, speciation, and additional cycles of diversification that may themselves be triggered by speciation. Extensive ecological opportunity may exist in depauperate communities, but it is unclear whether ecological opportunity abates in species-rich communities. Because ecological opportunity should generally increase during times of rapid and multifarious environmental change, human activities may currently be generating elevated ecological opportunity but so far little work has directly addressed this topic. Our framework highlights the need for greater synthesis of community ecology and evolutionary biology, unifying the four major components of the concept of ecological opportunity.

The Qinghai-Tibetan Plateau (QTP) is the highest and one of the most extensive plateaus in the world. Phylogenetic, phylogeographic, and ecological studies support plant diversifications on the QTP through multiple mechanisms such as allopatric speciation via geographic isolation, climatic oscillations and divergences, pollinator-mediated isolation, diploid hybridization and introgression, and allopolyploidy. These mechanisms have driven spectacular radiations and/or species diversifications in various groups of plants such asPedicularisL.,SaussureaDC.,RhododendronL.,PrimulaL.,MeconopsisVig.,RhodiolaL., and many lineages of gymnosperms. Nevertheless, much work is needed toward understanding the evolutionary mechanisms of plant diversifications on the QTP. Well-sampled biogeographic analyses of the QTP plants in the broad framework of the Northern Hemisphere as well as the Southern Hemisphere are still relatively few and should be encouraged in the next decade. This paper reviews recent evidence from phylogenetic and biogeographic studies in plants, in the context of rapid radiations, mechanisms of species diversifications on the QTP, and the biogeographic significance of the QTP in the broader context of both the Northern and Southern Hemisphere biogeography. Integrative multidimensional analyses of phylogeny, morphological innovations, geography, ecology, development, species interactions and diversifications, and geology are needed and should shed insights into the patterns of evolutionary assembly and radiations in this fascinating region.

Comparison of plant mitochondrial (mt), chloroplast (cp) and nuclear (n) DNA sequences shows that the silent substitution rate in mtDNA is less than one-third that in cpDNA, which in turn evolves only half as fast as plant nDNA. The slower rate in mtDNA than in cpDNA is probably due to a lower mutation rate. Silent substitution rates in plant and mammalian mtDNAs differ by one or two orders of magnitude, whereas the rates in nDNAs may be similar. In cpDNA, the rate of substitution both at synonymous sites and in noncoding sequences in the inverted repeat is greatly reduced in comparison to single-copy sequences. The rate of cpDNA evolution appears to have slowed in some dicot lineages following the monocot/dicot split, and the slowdown is more conspicuous at nonsynonymous sites than at synonymous sites.

作为植物细胞器的重要组成部分和光合作用的器官,叶绿体在生物进化的漫长历史中发挥了重要作 用.伴随着生物技术的深入发展,人们发现叶绿体基因组结构和序列的信息在揭示物种起源、进化演变及其不同物种之间的亲缘关系等方面具有重要价值.与此同 时,比核转化具有明显优势的叶绿体转化技术在遗传改良、生物制剂的生产等方面显示出巨大潜力,而叶绿体基因组结构和序列分析则是叶绿体转化的基石.基于叶 绿体的这些重要作用,收集整理了有关的资料,从几个方面归纳了本领域最近的研究进展,希望能使读者对迅速发展的叶绿体基因组研究有更全面的了解,以及对叶 绿体基因组在物种的进化、遗传、系统发育关系等方面的作用有更深刻的认识,同时也希望对叶绿体转化技术的研究和广泛应用产生积极作用.

作为植物细胞器的重要组成部分和光合作用的器官,叶绿体在生物进化的漫长历史中发挥了重要作 用.伴随着生物技术的深入发展,人们发现叶绿体基因组结构和序列的信息在揭示物种起源、进化演变及其不同物种之间的亲缘关系等方面具有重要价值.与此同 时,比核转化具有明显优势的叶绿体转化技术在遗传改良、生物制剂的生产等方面显示出巨大潜力,而叶绿体基因组结构和序列分析则是叶绿体转化的基石.基于叶 绿体的这些重要作用,收集整理了有关的资料,从几个方面归纳了本领域最近的研究进展,希望能使读者对迅速发展的叶绿体基因组研究有更全面的了解,以及对叶 绿体基因组在物种的进化、遗传、系统发育关系等方面的作用有更深刻的认识,同时也希望对叶绿体转化技术的研究和广泛应用产生积极作用.

ABSTRACT Variation in the chloroplast DNA sequence is useful for plant phylogenetic studies. However, the number of variable sequences provided by chloroplast DNA for suggested genes or genomic regions in plant phylogenetic analyses is often inadequate. To identify conserved regions that can be used to design primers and amplify variable sequences for use in plant phylogenetic studies, the complete chloroplast genomic sequences of six plant species (including Oryza sativa, Arabidopsis thaliana, Glycine max, Lotus japonicus, Medicago truncatula, and Phaseolus vulgaris), searched from the taxonomy database of NCBI were investigated. A total of 93 conserved regions, 32 in large single copy and 61 in inverted repeat regions, were identified. A set of five primer pairs were designed according to the conserved sequences located in the psbA~trnK, psbB~psbH, rpl23~trnI, trnR~trnN, and trnY~trnD regions to amplify variable DNA fragments. An additional 18 plant accessions from 14 species were used to validate their utility. Each of the tested species could be distinguished by length polymorphisms of fragments amplified with the five primer pairs. trnR~trnN and rpl23~trnI amplified fragments specific to monocot and legume species, respectively. Three primer pairs located in the psbA~trnK, psbB~psbH, and trnR~trnN regions were applied to amplify variable DNA sequences for phylogenetic analysis using the maximum parsimony method. The consistent result between taxonomy and phylogenetic analysis on the variable sequences amplified with these three primer pairs was revealed. The five newly developed primer pairs are recommended as tools for use in the identification of plant species and in phylogenetic studies.

Chloroplast DNA (cpDNA) markers were developed that provided markers unique to a species or that delimited a large area within a species. These markers were then followed across two hybrid zones: Iris douglasiana/Iris innominata, and Iris chrysophylla/Iris tenax. In each case the cline in haplotype frequency was compared to the cline for a morphologically based hybrid index. In all three transects across the I. douglasiana/I. innominata hybrid zone, the cpDNA cline was displaced 1-2 km relative to the morphologically defined hybrid zone; the displacement was not found in the other hybrid zone. The observed displacement represents introgression of cpDNA from I. douglasiana into I. innominata. It may be that the I. douglasiana/I. innominata hybrid zone has shifted in recent time, leaving the slowly dispersing chloroplast DNA behind. The populations known as Iris thompsonii do not form a phylogenetic species and are best viewed as products of hybridization between I. douglasiana and I. innominata.

Tibetan Plateau (TP) and adjacent regions have the most diverse of alpine plants in the world.The phylogeography of alpine plants in this region could explain the relationship between their demographic history with TP uplift and Quaternary glaciation.Base on the phylogeographic researches of 36 alpine plants in the published literatures,two generalized phylogeographic patterns were concluded:(1)Glacial retreat and subsequent recolonization of the platform from peripheral glacial refugia;(2)Existence of cryptic (or micro-) refugia outside the major glacial refugia,resulted from geographical isolation,followed local expansion during postglacials.The rapid TP uplift promoted intraspecific divergence and allopatric speciation within the late Tertiary,the followed periodic climate fluctuations accelerated the situation.These are the main reasons for shaping the modern phylogeographic patterns.Additionally,Hengduan Mountains as the major glacial refugia,have important implications for Evolutionary Biology and Conservation Biology.At last,several research fields of phylogeography of alpine plant in TP were provided,including species selection,sampling strategy,gene fragment selection and methods.

Tibetan Plateau (TP) and adjacent regions have the most diverse of alpine plants in the world.The phylogeography of alpine plants in this region could explain the relationship between their demographic history with TP uplift and Quaternary glaciation.Base on the phylogeographic researches of 36 alpine plants in the published literatures,two generalized phylogeographic patterns were concluded:(1)Glacial retreat and subsequent recolonization of the platform from peripheral glacial refugia;(2)Existence of cryptic (or micro-) refugia outside the major glacial refugia,resulted from geographical isolation,followed local expansion during postglacials.The rapid TP uplift promoted intraspecific divergence and allopatric speciation within the late Tertiary,the followed periodic climate fluctuations accelerated the situation.These are the main reasons for shaping the modern phylogeographic patterns.Additionally,Hengduan Mountains as the major glacial refugia,have important implications for Evolutionary Biology and Conservation Biology.At last,several research fields of phylogeography of alpine plant in TP were provided,including species selection,sampling strategy,gene fragment selection and methods.

In plants, the maintenance of species integrity despite hybridization has often been explained by the co-adaption of nuclear gene complexes. However, the interaction between plastid and nuclear sub-genomes has been underestimated. Here, we analyzed the genetic structure of aPopulus albaandP. tremulahybrid zone along the Irtysh River system in the Altai region, northwest China, using both nuclear microsatellites and plastid DNA sequences. We found high interspecific differentiation, although the hybridP. canescenswas prevalent. Bayesian inference classified most hybrids into F1, followed by a few back-crosses toP. alba, and fewer F2 hybrids and back-crosses toP. tremula, indicating a few introgressions but preference towardP. alba. When plastid haplotypes in parental species were distinct,P. canescenscarried the haplotypes of both parents, but showed significant linkage between intraspecific haplotype and nuclear genotypes at several microsatellite loci. Selection, rather than migration and assortative mating, might have contributed to such plastid-nuclear disequilibria. By removing later-generated hybrids carrying interspecific combinations of haplotype and nuclear genotypes, plastid-nuclear incompatibility has greatly limited the gene exchange betweenP. albaandP. tremula viabackcrossing with hybrids, demonstrating a significant association between plastid haplotype and the proportion of nuclear admixture.

横断山区作为青藏高原东南部主要的一个冰期避难所,第四纪冰期气候的变化对该地区的植物地理分布和居群遗传结构都产生了重要的影响。为了揭示该地区物种分布的分子系统地理学结构,选取在该地区广泛分布的一种高山灌木-高山绣线菊的叶绿体trnL-trnF序列进行研究。采集了15个居群182个个体进行测序,共发现7个单倍型。总的遗传多样性较高(HT=0.809),但居群内遗传多样性较低(HS=0.236)。分子变异分析(AMOVA)结果表明分布区内高山绣线菊的遗传变异主要存在于居群间(84.48%),且居群间的遗传分化很高(GST=0.708,FST=0.84476,NST=0.863),有着显著的谱系地理学结构(NSTGST,P0.01)和较低的居群间平均基因流(Nm=0.09)。单倍型的系统进化树和进化分支网络分析得到了相似的拓扑结构,7种单倍型都按照地理分布聚为三支:横断山区西部、横断山区东部以及两者的交接地带。本研究推测该物种在横断山区存在多个冰期避难所,而没有表现出大规模的种群集体扩张和迁移的现象。青藏高原隆升、第四纪气候的反复波动以及横断山区特殊的地理环境使得原来连续的居群片段化,并发生范围扩张,从而塑造了高山绣线菊的现代生物地理分布格局。

横断山区作为青藏高原东南部主要的一个冰期避难所,第四纪冰期气候的变化对该地区的植物地理分布和居群遗传结构都产生了重要的影响。为了揭示该地区物种分布的分子系统地理学结构,选取在该地区广泛分布的一种高山灌木-高山绣线菊的叶绿体trnL-trnF序列进行研究。采集了15个居群182个个体进行测序,共发现7个单倍型。总的遗传多样性较高(HT=0.809),但居群内遗传多样性较低(HS=0.236)。分子变异分析(AMOVA)结果表明分布区内高山绣线菊的遗传变异主要存在于居群间(84.48%),且居群间的遗传分化很高(GST=0.708,FST=0.84476,NST=0.863),有着显著的谱系地理学结构(NSTGST,P0.01)和较低的居群间平均基因流(Nm=0.09)。单倍型的系统进化树和进化分支网络分析得到了相似的拓扑结构,7种单倍型都按照地理分布聚为三支:横断山区西部、横断山区东部以及两者的交接地带。本研究推测该物种在横断山区存在多个冰期避难所,而没有表现出大规模的种群集体扩张和迁移的现象。青藏高原隆升、第四纪气候的反复波动以及横断山区特殊的地理环境使得原来连续的居群片段化,并发生范围扩张,从而塑造了高山绣线菊的现代生物地理分布格局。

The topological effect of the Qinghai-Tibet Plateau during the Quaternary climatic oscillation must have important effects on the patterns of spatial distribution and genetic structure of organisms distributed there. To obtain a better understanding of the historical dynamics of present-day species in this region, we examined the population structure of chloroplast trnT-trnF sequence variation in Juniperus przewalskii, an endemic tree species mainly distributed in the northeast plateau. Data were obtained for 392 trees from 20 populations covering the total distributional range of the species. Three cpDNA haplotypes were recorded and were structured into two geographical areas: haplotype A was present on the plateau platform, while all three haplotypes occurred along the plateau edge. Total genetic diversity, HT, was 0.511, with most variation distributed among rather than within populations (GST=0.847). Along the low altitudinal plateau edge, haplotypes A, B and C tended to be fixed or nearly fixed in different populations, indicating that the species may have survived the last glacial period in different refugia within this area, and the bottlenecks due to the repeated reduction and expansion of the population size resulted in the reduction of biodiversity. However, in one population both haplotypes occurred at intermediate frequencies, indicating the possibility of a contact zone or less strong bottleneck effect. On the eastern plateau platform, haplotype A was commonly fixed in the current disjunct populations, suggesting a common postglacial recolonization and a following founder effect. Our results indicate that there might exist in multi-glacial refugia in the plateau edge and both bottleneck and founder effects resulting from the repeated climatic oscillation combined had played important roles in shaping the current haplotype distribution pattern of this tree species.

The topological effect of the Qinghai-Tibet Plateau during the Quaternary climatic oscillation must have important effects on the patterns of spatial distribution and genetic structure of organisms distributed there. To obtain a better understanding of the historical dynamics of present-day species in this region, we examined the population structure of chloroplast trnT-trnF sequence variation in Juniperus przewalskii, an endemic tree species mainly distributed in the northeast plateau. Data were obtained for 392 trees from 20 populations covering the total distributional range of the species. Three cpDNA haplotypes were recorded and were structured into two geographical areas: haplotype A was present on the plateau platform, while all three haplotypes occurred along the plateau edge. Total genetic diversity, HT, was 0.511, with most variation distributed among rather than within populations (GST=0.847). Along the low altitudinal plateau edge, haplotypes A, B and C tended to be fixed or nearly fixed in different populations, indicating that the species may have survived the last glacial period in different refugia within this area, and the bottlenecks due to the repeated reduction and expansion of the population size resulted in the reduction of biodiversity. However, in one population both haplotypes occurred at intermediate frequencies, indicating the possibility of a contact zone or less strong bottleneck effect. On the eastern plateau platform, haplotype A was commonly fixed in the current disjunct populations, suggesting a common postglacial recolonization and a following founder effect. Our results indicate that there might exist in multi-glacial refugia in the plateau edge and both bottleneck and founder effects resulting from the repeated climatic oscillation combined had played important roles in shaping the current haplotype distribution pattern of this tree species.

Michelia yunnanensisFranch., is a traditional ornamental, aromatic, and medicinal shrub that endemic to Yunnan Province in southwest China. Although the species has a large distribution pattern and is abundant in Yunnan Province, the populations are dramatically declining because of overexploitation and habitat destruction. Studies on the genetic variation and demography of endemic species are necessary to develop effective conservation and management strategies. To generate such knowledge, we used 3 pairs of universal cpDNA markers and 10 pairs of microsatellite markers to assess the genetic diversity, genetic structure, and demographic history of 7M. yunnanensispopulations. We calculated a total of 88 alleles for 10 polymorphic loci and 10 haplotypes for a combined 2,089 bp of cpDNA.M. yunnanensispopulations showed high genetic diversity (Ho= 0.551 for nuclear markers andHd= 0.471 for cpDNA markers) and low genetic differentiation (FST= 0.058). Geographical structure was not found amongM. yunnanensispopulations. Genetic distance and geographic distance were not correlated (P> 0.05), which indicated that geographic isolation is not the primary cause of the low genetic differentiation ofM. yunnanensis. Additionally,M. yunnanensispopulations contracted ~20,000 30,000 years ago, and no recent expansion occurred in current populations. Results indicated that the high genetic diversity of the species and within its populations holds promise for effective genetic resource management and sustainable utilization. Thus, we suggest that the conservation and management ofM. yunnanensisshould address exotic overexploitation and habitat destruction.

L-F fragment by PCR-RFLP.H1 and H2 existed in all the populations distributing in mainland of China,but H3 only existed in Huangshan population,and H4 only existed in Lin n population,with H5 and H6 coexisting in Taiwan population.The phylogeographic analysis suggested the H1 and H2 were the ancestor haplotypes,while the other haplotypes were evolved from them.Some 26.39% of the total variation existed among the populations,while 73.61% in the populations suggested by AMOVA.NCA analysis suggested there was slight correlation between the distribution of cpDNA haplotypes and geography,and the genetic structure of T.wallichiana var.mairei resulted from the effect of population fragment or the long-distance colonisation.Variation of cpDNA haplotypes suggested several glacial refugiums of T.wallichiana var.mairei in the east and south of China.

L-F fragment by PCR-RFLP.H1 and H2 existed in all the populations distributing in mainland of China,but H3 only existed in Huangshan population,and H4 only existed in Lin n population,with H5 and H6 coexisting in Taiwan population.The phylogeographic analysis suggested the H1 and H2 were the ancestor haplotypes,while the other haplotypes were evolved from them.Some 26.39% of the total variation existed among the populations,while 73.61% in the populations suggested by AMOVA.NCA analysis suggested there was slight correlation between the distribution of cpDNA haplotypes and geography,and the genetic structure of T.wallichiana var.mairei resulted from the effect of population fragment or the long-distance colonisation.Variation of cpDNA haplotypes suggested several glacial refugiums of T.wallichiana var.mairei in the east and south of China.