272 resultados para ANGIOSPERM PHYLOGENY


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The peritrichs have been recognized as a higher taxon of ciliates since 1968. However, the phylogenetic relationships among them are still unsettled, and their placement within the class Oligohymenophorea has only been supported by the analysis of the small subunit rRNA gene sequence of Opisthonecta henneguyi. DNA was isolated directly from field-sampled species for PCR, and was used to resolve relationships within the genus Epistylis and to confirm the stability of the placement of peritrichs. Small subunit rRNA gene sequences of Epistylis plicatilis, Epistylis urceolata, Epistylis chysemydis, Epistylis hentscheli, Epistylis wenrichi, and Vorticella campanula were sequenced and analyzed using both distance-matrix and maximum-parsimony methods. In phylogenetic trees, the monophyly of both the genus Epistylis and the subclass Peritrichia was strongly supported, while V. campanula clustered with Vorticella microstoma. The topology in which E. plicatilis and E. hentscheli formed a strongly supported sister clade to E. urceolata, E. chrysemydis, and E. wenrichi was consistent with variations in the thickness of the peristomial lip. We concluded that the peristomial area, especially the. peristomial lip, might be the important phylogenetic character within the genus Epistylis.

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SINE (short interspersed element) insertion analysis elucidates contentious aspects in the phylogeny of toothed whales and dolphins (Odontoceti), especially river dolphins. Here, we characterize 25 informative SINEs inserted into unique genomic loci during evolution of odontocetes to construct a cladogram. and determine a total of 2.8 kb per taxon of the flanking sequences of these SINE loci to estimate divergence times among lineages. We demonstrate that: (i) Odontocetes are monophyletic; (ii) Ganges River dolphins, beaked whales, and ocean dolphins diverged (in this order) after sperm whales; (iii) three other river dolphin taxa, namely the Amazon, La Plata, and Yangtze river dolphins, form a monophyletic group with Yangtze River dolphins being the most basal; and (iv) the rapid radiation of extant cetacean lineages occurred some 28-33 million years B.P., in strong accord with the fossil record. The combination of SINE and flanking sequence analysis suggests a topology and set of divergence times for odontocete relationships, offering alternative explanations for several long-standing problems in cetacean evolution.

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1140 bp of cytochrome b gene were amplified and sequenced from 14 species of primitive cyprinid fishes in East Asia. Aligned with other ten cytochrome b gene sequences of cyprinid fish from Europe and North America retrieved from Gene bank, we obtained a matrix of 24 DNA sequences. A cladogram was generated by the method of Maximum likelihood for the primitive cyprinid fishes. The result indicated that subfamily Leuciscinae and Danioninae do not form a monophyletic group. In the subfamily Danioninae, Opsariichthys biden and Zacco platypus are very primitive and form a natural group and located at the root. But the genera in subfamily Danioninae are included in different groups and have not direct relationship. Among them, Aphyocypris chinensis and Yaoshanicus arcus form a monophyletic group. Tanichthys albonubes and Gobiocypris rarus have a close relation to Gobioninae. The genus Danio is far from other genera in Danioninae, In our cladogram, the genera in Leuciscinae were divided into two groups that have no direct relationship. The genera in Leuciscinae distributed in Europe, Sibera and North America, including Leuciscus, Rutilus, Phoxinus, N. crysole, Opsopoeodus emilae, form a monophyletic group. And the Leuciscinae in southern China including Ctenopharyngodon idellus, Mylopharyngodon piceus, Squalibarbus and Ochetobius elongatus have a common origination.

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We surveyed mitochondrial DNA (mtDNA) sequence variation in the subfamily Xenocyprinae from China and used these data to estimate intraspecific, interspecific, and intergeneric phylogeny and assess biogeographic scenarios underlying the geographic structure of lineages. We sequenced 1140 bp of cytochrome b from 30 individuals of Xenocyprinae and one putative outgroup (Myxocypris asiaticus) and also sequenced 297 bp of ND4L, 1380 bp of ND4, 68 bp of tRNA(His), and 69 bp of tRNA(Ser) from 17 individuals of Xenocyprinae and the outgroup (M. asiaticus). We detected high levels of nucleotide variation among populations, species, and genera. The phylogenetic analysis suggested that Distoechodon hupeinensis might be transferred to the genus Xenocypris, the taxonomic status of the genus Plagiognathops might be preserved, and species of Xenocypris and Plagiognathops form a monophyletic group that is sister to the genus Distoechodon and Pseudobrama. The introgressive hybridization might occur among the populations of X. argentea and X. davidi, causing the two species to not be separated by mtDNA patterns according to their species identification, and the process and direction of hybridization are discussed. The spatial distributions of mtDNA lineages among populations of Xenocypris were compatible with the major geographic region, which indicated that the relationship between Hubei + Hunan and Fujian is closer than that between Hubei + Hunan and Sichuan, From a perspective of parasite investigation, our data suggested that the fauna of Hexamita in Xenocyprinae could be used to infer the phylogeny of their hosts. (C) 2001 Academic Press.

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The four species of "river dolphins" are associated with six separate great river systems on three subcontinents and have been grouped for more than a century into a single taxon based on their similar appearance. However, several morphologists recently questioned the monophyly of that group. By using phylogenetic analyses of nucleotide sequences from three mitochondrial and two nuclear genes, we demonstrate with statistical significance that extant river dolphins are not monophyletic and suggest that they are relict species whose adaptation to riverine habitats incidentally insured their survival against major environmental changes in the marine ecosystem or the emergence of Delphinidae.

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Acipenseriformes is an endangered primitive fish group, which occupies a special place in the history of ideas concerning fish evolution, even in vertebrate evolution. However, the classification and evolution of the fishes have been debated. The mitochondrial DNA (mtDNA) ND4L and partial ND4 genes were first sequenced in twelve species of the order Acipenseriformes, including endemic Chinese species. The following points were drawn from DNA sequences analysis: (i) the two species of Huso can be ascribed to Acipenser; (ii) A. dabryanus is the mostly closely related to A. sinensis, and most likely the landlocked form of A. sinensis; (iii) genus Acipenser in trans-Pacific region might have a common origin; (iv) mtDNA ND4L and ND4 genes are the ideal genetic markers for phylogenetic analysis of the order Acipenseriformes.

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用六个线粒体基因片段序列数据,对称猴属的分子系统发育关系进行了分析。8种称猴:台湾猴,北平顶猴,藏酋猴,熊猴,红面猴,J厦河猴,狮尾猴和食蟹猴共40个样本用于本研究。结合从GenBank里下载的雯猴的相应序列,用拂拂作为外群构建了系统发育树。由于六个目的片段均为线粒体基因片段,缺乏重组且作为单一位点遗传,因此我们将六个数据集合并为一个数据集进行分析。我们获得了比前人置信度更高的线粒体系统树。除了红面猴,我们的结果支持Delson(1980)的分组。与前人研究结果一致,红面猴与食蟹猴组成员关系较近(尤其是食蟹猴)。恒河猴(相对于台湾猴),熊猴(相对于藏酋猴)和北平顶猴(相对于狮尾猴)都形成了并系。通过线粒体控制区序列和20个微卫星位点对14个恒河猴地理群体的群体结构进行了研究。微卫星数据和线粒体控制区数据均显示恒河猴有较高的遗传多样度。线粒体数据表明各地区的恒河猴群体间的差异显著大于微卫星数据显示的结果。也就是说,基于微卫星数据所得到的群体间基因流远大于基于线粒体数据所得到的群体间基因流。两种分子标记所得到的结果差异应归因为雌性恒河猴的归家冲动和雄性恒河猴的扩散。线粒体数据显示所有恒河猴群体近期没有发生过快速扩张。微卫星数据显示大部分恒河猴群体近期没有经历过瓶颈效应。恒河猴是分布较广的非人灵长类动物,但它的地理差异研究一直颇有争论且不足以划分其亚种。我们对采自包括印度、越南、缅甸和18个中国地区在内的21个地区共35个恒河猴样品的四个线粒体基因片段(COI,Cofl,COlll和12srRNA)进行了研究。食蟹猴、红面猴、台湾猴和史猴的相应序列作为外群。大约长为100bp的序列用于构建最大似然树和贝叶斯树。与前人结果一致(彭燕章等,1993;蒋学龙等,1995;Groves,2001),西部恒河猴(印度).相当于M.m.mulatta。根据王应祥(2003)的分类,我们的东部恒河猴可分为四组:M.最早分离出来且一个来自缅甸的恒河猴位于该组基部;M.m.brachyurus和M.m.littoralis组成一个大枝,这个大枝与M.m,lasiotus构成姐妹群。

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本工作用线粒体细胞色素b基因全序列探讨了白腹鼠属(Niviventer)分子系统发育关系,利用线粒体控制区全序列研究小家鼠(Mus musculus)在中国的分子系统地理学。 在白腹鼠属的分子系统发育研究中,本研究首次对来自青藏高原东南部和台湾的9种(安氏白腹鼠、川西白腹鼠、社鼠、刺毛鼠、梵鼠、灰腹鼠、褐尾鼠、台湾白腹鼠和台湾社鼠)32个个体的细胞色素b全序列进行了测定和分析。用最大简约法、最大似然法和贝叶斯法三种构树方法构出的进化树一致表明,9种分为3个线粒体DNA系:系A包括社鼠、台湾社鼠、台湾白腹鼠、刺毛鼠、梵鼠和灰腹鼠6种;系B包括安氏白腹鼠和川西白腹鼠2种;系C仅包括褐尾鼠一种。应用DNA分类的分析方法表明台湾社鼠不应是社鼠的亚种,而应是一个独立的种。此外,以来自更新世中晚期(1.2-0.13百万年)的社鼠化石为依据推算该属的分歧时间。结果表明该属的起源于1.64百万年前,系A和系B分化发生在1.46百万年以前,该属的其余物种分化时间为1.29-0.67百万年前。这些分化时间和青藏高原的最后一次隆起、昆黄运动以及第四纪大冰期时间相一致,提示地理隔离和冰期作用可能对该属的形成演化起到重要的作用。 在小家鼠的分子系统地理学研究中,通过测定来自中国12个采集地的184只小家鼠的控制区全序列,并结合了Genbank里的单倍型,首次研究小家鼠在中国的分子系统地理格局、种群历史和生物地理过程。184个样品定义了66个单倍型,单倍型多样性为0.95,核苷酸多样性为0.012。用66个单倍型及Genbank里来自周边国家的62个小家鼠单倍型进行分子系统发育分析,邻接法和贝叶斯法构得的分子系统发育树基本一致。结果表明小家鼠在中国分为两个线粒体DNA系(南方系和北方系)。分子变异等级分析结果表明主要的分子变异发生在两个线粒体DNA系之间(84.03%)。根据研究结果推断长江是两个线粒体DNA系之间基因交流受限的主要地理障碍。单倍型进化网络关系表明两个线粒体DNA是异域扩散。两个系的单倍型歧点分布图呈钟型分布,说明经历了快速的种群扩散。经推算小家鼠在中国北方系的种群扩散时间为16 600(22 000-11 000)年前,而南方系的种群扩散时间早于北方系,扩散时间为20 000(40 000-6 700)年前。依据单倍型和核苷酸的多样性、单倍型进化网络关系以及单倍型歧点分布图综合推断小家鼠在中国的两个系在末冰期时经历了快速的种群扩张导致今天的地理分布格局。根据系统发育树的结果推断,北方系种群的大体扩散方向是从北往南,而南方系是从南往北迁移。小家鼠以长江为界分为南方系和北方系的系统地理模式、扩散方向和种群的扩散时间与人以长江为界分为南方人和北方人、扩散方向、扩散时间(60 000-16 000)基本一致,这可能暗示人类活动对小家鼠的迁移产生的重要影响。

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鰋鮡鱼类共包括9 属43 种(亚种),隶属于鲇形目(Siluriformes)鮡科 (Sisoridae)鰋鮡亚科(Glyptosternae)中鰋鮡族(Tribe Glyptosternini)的鰋鮡 亚族(Subtribe Glyptosternina)。鰋鮡鱼类形态上的共同特征为:无胸吸着器,胸、 腹鳍水平展开,第一根鳍条完全分节或在外缘生出许多软骨细条,被外表皮所裹, 在腹面看到的是许多与分节或软骨细条大致对应的横纹皱褶。鰋鮡鱼类集中分布 于青藏高原周边的水系中,部分属种向西分布到了中亚地区的阿姆河上游,是适 应山区急流环境的一群鱼类。近年来一系列的研究表明,鰋鮡鱼类为一单系类群, 它的起源和演化与青藏高原的隆升有着直接的关系。鰋鮡鱼类的系统发育研究, 可以为青藏高原隆升的年代、幅度和形式提供间接的证据,其系统发育树的拓扑 结构也直接反映了东喜马拉雅地区诸水系的形成与演变。 本研究对鰋鮡鱼类9 属进行了系统整理。1. 发现并描述了异齿鰋属 (Oreoglanis)两新种,分别为分布于景东无量山(澜沧江水系)的景东异齿鰋 (O. jingdongensis)和分布于怒江水系南景河和南滚河的无斑异齿鰋(O. immaculatus)。认为分布于中国的异齿鰋属鱼类均属于尖须异齿鰋种组(O. siamensis species group)。指出区分异齿鰋属两个种组:尖须异齿鰋种组和细尾 异齿鰋种组(O. delacouri species group)的特征是下唇中部是否具有中央缺刻, 尾型的差别(新月型尾或凹型尾)不能用来区分两个种组,给出了异齿鰋属的检 索表,并绘制了异齿鰋属鱼类分布图。2. 对鮡属(Pareuchiloglanis)鱼类进行了 系统整理,通过外部形态度量性状的比较,认为分布于澜沧江水系的,曾经被鉴 定为扁头鮡(P. kamengensis)的标本,应属于大鳍鮡(P. macropterus),扁头鮡 和大鳍鮡之间的最明显的差别在于腹鳍前长的不同。前者的腹鳍前长为体长的 53.2-64.9%,后者的腹鳍前长均不达体长的50%。3. 通过比较金沙江水系分布的 5 种鮡属鱼类,中华鮡(P. sinensis)、前臀鮡(P. anteanalis)、壮体鮡(P. robusta)、 四川鮡(P. sichuanensis)和天全鮡(P. tianquanensis),认为天全鮡和四川鮡之 间分布水系重叠,外部形态亦无差别,天全鮡很可能为四川鮡的同物异名。给出 了鮡属鱼类的系统检索表,并绘制了鮡属鱼类分布图。利用分子系统学的原理和方法对鰋鮡鱼类进行系统发育研究。测定了6 属 15 种鰋鮡鱼类和鮡科中非鰋鮡鱼类4 属7 种共28 个体的线粒体Cyt b 基因部分 片段和全序列(1138 bp),结合从GenBank 下载的相关类群相同的基因序列,以 魾属(Bagarius)的巨魾(B. yarrelli),纹胸鮡属(Glyptothorax)的穴形纹胸鮡 (G. cavia)、亮背纹胸鮡(G. dorsalis)、扎那纹胸鮡(G. zainaensis),福建纹胸 鮡(G. fukiensis fukiensis)、海南纹胸鮡(G. fukiensis hainanensis),黑鮡属(Gagata) 的长丝黑鮡(G. dolichonema)以及褶鮡属(Pseudecheneis)的黄斑褶鮡(P. sulcatus) 和无斑褶鮡(P. immaculatus)作为外类群,采用贝叶斯法(Bayesian)、最简约 法(maximum pasimony, MP)和邻接法(neighbour-joining, NJ)构建系统发育树。 结果显示: 1. 鰋鮡鱼类为一单系类群,并且与褶鮡属互为姐妹群关系; 2. 原鮡属、鰋属和凿齿鮡属是鰋鮡鱼类的三个基部类群; 3. 异齿鰋属构成为一个单系群,大鳍异齿鰋最早从该属的基部分化出来; 4. 石爬鮡属构成一个单系,并与分布于金沙江水系的中华鮡+前臀鮡构成姐妹 群,黄石爬鮡和青石爬鮡的单倍型相互交错;显示两个物种的分类是不合适的, 而是同一水系不同支流种群之间梯度变异的例子,依据本次研究所得出的三个分 支图,结合青石爬鮡自西至东分布于金沙江、雅砻江、大渡河、青衣江、岷江等 的分布格局以及形态特征的分布,显示形态特征的分布变化有以下趋势:自西至 东,腹鳍位置逐渐前移;颌须渐趋缩短;胸鳍趋向发达、伸达腹鳍起点。这些变 化趋势是同一水系不同支流种群之间梯度变异的极好例子; 5. 分布于澜沧江以西(包括澜沧江)水系的鮡属鱼类(扁头鮡、细尾鮡、短鳍 鮡)与分布于怒江和伊洛瓦底江水系的拟鰋属鱼类构成为一支,并且二者共同与 分布于元江上游(红河水系)的大孔鮡构成为单系。其中,短体拟鰋和拟鰋互为 姐妹种;而扁头鮡和短鳍鮡也互为姐妹种;细尾鮡的系统地位则尚不能确定; 6. 无论在MP 树、贝叶斯树还是NJ 树中,石爬鮡属、鮡属、拟鰋属和异齿鰋属 构成一个单系,并且支持率达到了100%。 7. 鰋属位于鰋鮡鱼类的基部,是较早就从鰋鮡鱼类祖先中演化出来的一个类群, 鰋类的口吸盘是一个趋同性状,是在急流环境条件下形成的一种适应性性状,在 鰋类各属中不是同源特征; 8. 鮡属鱼类不是一个单系类群,分布于金沙江流域的鮡属鱼类与同流域分布的石爬鮡属鱼类聚成一支;澜沧江及其以西水系分布的鮡属鱼类与同流域分布的拟 鰋属鱼类聚成一支,之后这两支又同异齿鰋属共同构成为一个大支。 结合鰋鮡鱼类的系统发育分支图对其动物地理学进行研究。鰋鮡鱼类是由类 似纹胸鮡属(Glyptothorax)鱼类的祖先演化而来,鰋属(Exostoma)和凿齿鮡 属(Glaridoglanis)是较早就从从类似原鮡的祖先中演化出来,鮡属、石爬鮡属、 拟鰋属和异齿鰋属这一大支由类似原鮡的祖先演化而来。在青藏高原强烈隆起等 重大地理隔离事件发生之前,类似于现生的原鮡属鱼类已经广布喜马拉雅山脉东 西两侧。在青藏高原强烈隆起等重大地理隔离事件发生后,鰋鮡鱼类在相同的地 理隔离下独自演化为现在的分布格局。

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裂腹鱼类起源鳃亚科鱼类的观点得到了多数学者的支持。但是,鱼巴亚科鱼类是一个非常庞杂的类群,裂腹鱼类究竟起源于鳃亚科的哪一个小类群或裂腹鱼类与哪个小类群更为接近?这样的小类群又怎样随着急剧隆起的青藏高原峥演化发展成为现今的裂腹鱼类?不同的学者从不同的研究领域出发,得出了较为不一致甚至是相互对立的结论。裂腹鱼类(我国有11属,76种和亚种)均为小型鳞片的鱼类,与鳃亚科中一些具有小型鳞片的属种较为相似,即,妒鲤属(凡rCO~)的3个亚种、似鳃属(Lztciocyprinus)的2个种,金线纪属(Sinocyclocheilus)的26种或亚种,本文将这些具细小鳞片的鳃亚科鱼类统称为细鳞鳃鱼类(small-scaledbarbids)。为了探讨细鳞鱼巴类和裂腹鱼类的系统发育关界本文应用分支系统学的方法和原理,选择小裂腹鱼S.(R)parvus、中甸叶须鱼P.c.chunglienensis、松藩裸鲤。Gp.potanini、单纹似鳡L.langsoni、花妒鲤P.pingiregani、抚仙金线鱼巴s.师分别作为裂腹鱼类和细鳞鱼巴类中各分类:阶元的代,作为内a为了确考险状的进化极向并进一步探讨细鳞鱼巴各分类阶元与其他纪亚科和鲤亚科鱼类的系统关系,又选择了鳃亚科中的保山四须纪B.baoshanensis、云南四作为内群,并以螂鱼C.auratus、祀麓鲤C.(C)chilia作为外类群,用来对裂腹鱼类和细鳞鳃类进行性状的极化和系统发育分析。通过对12种24尾标本的外部形态和骨骼特征的全面观察,选择了63个性状进行描述和比较,并构建了供系统发育分析的特征矩阵。应用PAUP程序对特征矩阵进行运算,得出了包括外类群在内最简约的系统发育分支图。主要研究结论如下:(l)鲤亚科与包括裂腹鱼类在内的鳃亚科鱼类之间在很多特征方面存在着比较明显的差异,除了臀鳍最末一根不分枝鳍条坚硬、具锯齿;背鳍分枝多于1。根;第一椎体横突退化等特征外,还有镖的前后比例、前鳄骨的特征、尾舌骨的形状,基枕骨的骨质盘特征等诸多特征存在着显著差异。支持了陈湘舞等(1984)将鲤亚科和纪亚科分别作为独立亚科的观点。卿通过选取不同的外类群,在各种不同的条件下对特征数据矩阵进行运算和分析,除极少数(1个)结果外(当以祀麓鲤为唯一外群时,由34来看,它们都是第三纪末期以来由原始的鳃亚科鱼类演化而来的,适应过共同的寒冷环境(青藏高原的急剧隆升导致的寒冷气候和河川急流和青藏高原的大幅度隆起,引起了地貌环境的急剧改变,云贵高原一带也发生了差异性的升降运动。同时,全球性气温下降,常年性冰盖由北向南大幅度推进的寒冷气候环境)(曹文宣等,1981;王大忠等,2000)。裂腹鱼类和似鳃、妒鲤的起源时间可能稍晚一些。可以设想,第三纪末期的鳃亚科鱼类物种分化不是很多,相互之间的亲缘关系较近,分布于'青藏高原的原始鳃亚科鱼类和分布于云贵高原的原始鳃亚科鱼类分布经历了各自独特的地质、气候等环境条件,演化成为当今的裂腹鱼类和金线纪鱼类。至于似媳和妒鲤,除了经历与鳃亚科鱼类和金线鳃鱼类共同的寒冷水环境外,它们还向着肉食性的方向进行演化。(3)裂腹鱼类的3个代表种都聚在一起,其内部关系和与其他种的关系都较为稳定,为鳃亚科鱼类中较为特化的一支。与陈湘舞等(1984)、曹文宣等(1981)的观点较为一致,即,裂腹鱼类起源于纪亚科鱼类。晗(4)似媳和妒鲤之间的亲缘关系最近,它们很可能起源于最近的共同祖先。尽管二者作为一个单系的支持率并不是很高(大于50%,小于60%),但它们在个别特征(咽喉齿的排列方式、咽骨的形态特征)上表现其他鲤科鱼类所不具有的独特特征(附图1的。(5:)裂腹鱼类与妒鲤+似鳃的系统关系最近,这意味着裂腹鱼类和妒鲤+似鳃有最近的共同祖先或它们之间的关系较它们与金线鳃鱼类的关系更a(e)应用特征分析和系统发育分析所得的结论,综合对鳃系鱼类染色体特征的分析,认为裂腹鱼类和细鳞鳃鱼类在染色体分组组成、哪值等方面的相似特征,极有可能是在进化过程中经历了类似的寒冷水环境。它们之间的核型特征相似,只是说明了它们之间较近的系统发育关系,不支持妒鲤和金线靶是鳃亚科和裂腹鱼亚科之间的中间演化类型的观点。染色体的进化和外部形态特征的进化在纪系鱼类中存在着不平行的现象。(7)通过国内外鱼类学者对鲤科鱼类和纪亚科鱼类各分类单元定义的比较,结合本文的研究结果,认为国外学者的一些分类系统仍然存在一些尚待解决的问题,故建议目前暂不宜将之引入到我国的鱼类分类系统中。

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本文新种部分,记述了分布在横断山地区、台湾以及苏门答腊、菲律宾等地果蝇科的新种55种,异物同名3种。在果蝇属果蝇亚属的伊米果蝇种组建立了一个新的种复组-弯头果蝇种亚组(Drosophila curviceps species subgroup);在quadrilineata种亚组内建立了一个新的种复组-背条果蝇种复组(Drosophila notostriata species-complex)。对横断山地区的果蝇进行研究,计有119个种,主要由东洋种、古北种、特有种和广布种组成;对其渊源关系进行分析。运用分支分类学的观点和方法对伊米果蝇种组中5个种亚组间的系统发育进行研究显示,hypocausta种亚组最原始,位地分支图的最底部,是外群向伊米果蝇种组过渡的一个类群,进化中心在热带的苏门答腊及其附近;其次分化的immigrans种亚组其进化中心仍在热带的苏门答腊及其附近;但该亚组的特种逐渐向北扩散,在中国云南的西双版纳及其附近地区形成次级进化中心。伊米果蝇种组总的进化方向是从热带向温带的逐渐演化。

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青杨组(Section Tacamahaca Spach)杨树是我国重要的乡土经济树种,目前对其分子遗传变异和系统进化的研究还很少,尤其是在青杨组杨树遗传资源极为丰富的川西地区,杨树的分子进化及亲缘关系的研究极为缺乏,非常不利于该树种遗传资源的开发和利用。本研究从川西地区收集了青杨(Populus cathayana)、青海杨(P. prezewalskii)、滇杨(P. yunnanensis)、康定杨(P. kangdingensis)、西南杨(P. schneideri)、小叶杨(P.simonii)和三脉青杨(P. trinervis)这7 个青杨组树种的10 个群体,利用多种分子标记手段对其种间的亲缘关系进行比较,并结合形态和地史资料进行了全面的研究和评价,得到了如下的主要研究结果: 1. SSR 和ISSR 位点变异丰富。通过10 对引物对50 个杨树个体的DNA 样品进行了SSR 分析,所有位点展现了丰富的群体间和种间的多态性,多态位点率达到了100%,每位点的等位基因数变化范围为5 ~ 17,平均为11.9 个;通过11 条ISSR 随机引物对供试的混合DNA 样品进行分析,共检测到130 个标记,其中多态性标记为119 个,多态百分率为91.5%。研究认为,SSR 单个标记能展现高水平信息,而ISSR 单个引物能探测更多数量多态性。通过两个标记的遗传距离、聚类图和PCA 分析,表明:同一种内不同群体间的同源性最高;康定杨和西南杨有较近的亲缘关系;小叶杨和三脉青杨聚合在一起,显示了其相互较近的亲缘关系;滇杨与其它杨树种可能存在着较远的亲缘关系。 2. 采用4 对选择性引物对7 个青杨组杨树种10 个群体进行AFLP 分析,总共扩增出284 个位点,其中200 个位点显示出了多态性,多态位点百分比为70.4%,平均多态带为50 条。TE-AFLP 的分析总共扩增出192 个位点,其中139 个位点显示出了多态性,多态位点百分比为72.4%,平均多态带为34.7 条。比较的结果表明AFLP、TE-AFLP 的遗传信息含量比较接近,略小于ISSR,大约仅为SSR 的1/3;但这两个基于AFLP 的标记系统的信息探察能力也远大于ISSR 和SSR 标记系统。这两个分子标记的聚类结果,显示小叶杨、三脉青杨和滇杨三个种聚为一组,其中小叶杨与三脉青杨的亲缘关系更近;其它几个杨树种聚为一类,西南杨与青杨表现出较近的亲缘关系。 3. 所有7 对cpSSR 引物中,仅有4 个叶绿体位点在种间具有多态性,而在种内群体中并不具有多态性,共检测出13 个条带,组合成了4 种不同的单倍型;对于cpDNA的5 对引物,共检测出了73 条酶切片段,其中52 条是多态带,组合成了9 种不同的单倍型;而5 对mtDNA 通用引物未能检测出多态性的条带,表现出线粒体的保守性。叶绿体的聚类分析认为,小叶杨、三脉青杨和滇杨有较近的母性起源,且依次聚合;其余四种杨树聚为一类,并且康定杨与西南杨表现出最近的亲缘关系,并依次与青杨和青海杨聚合。 4. 根据本文的分子数据,结合形态和生境分布资料分析认为:青杨组杨树种内群体间的遗传变异程度是小于种间的遗传差异,显示了与传统分类一致的结果;三脉青杨和小叶杨有很近的亲缘关系,可能拥有相同的祖先类群;滇杨与小叶杨和三脉青杨之间具有一定的亲缘关系,特别是在其母性祖先的起源上有着一定的同源性;西南杨与青杨和康定杨均保持着较近的亲缘关系,且有可能是这两个种原始祖先杂交后所形成的。 Although western Sichuan is regarded as a natural distribution and variation center forthe Section Tacahamaca of the Populus species in China, little is currently known about themajority of poplar species occurring in this region. In the present study, molecular data wereutilized to determine the genetic relationships among Populus species in Section Tacamahacain western Sichuan including P. cathayana, P. prezewalskii, P. yunnanensis, P. kangdingensis,P. schneideri, P. simonii and P.trinervis. The results are as fellows: 1. The genetic variation at SSR and ISSR loci was abundant. All the 10 SSR loci werepolymorphic, and the number of alleles per locus varied from 5 to 17 with a mean valueequaling 11.9. Based on the 11 ISSR primers, 130 clear and reproducible DNA fragmentswere generated, of which 119 (91.5%) were polymorphic. Our results reveal that single SSRlocus can present more genetic information, while more polymorphic bands can be detectedby single ISSR primer. Moreover, the genetic distance, cluster and PCA analysisdemonstrated that: a close relationship among accessions of the same species and suggestedmonophyly in P. przewalskii and P. cathayana; P. schneideri is genetically highly similar to P.kangdingensis; P. trinervis and P. simonii have a close genetic affinity; P. yunnanensis isdistinct from the other species. 2. Genetic relationships of poplar species in Section Tacamahaca from western Sichuanwere evaluated by means of AFLP and TE-AFLP. For four AFLP primer combinations, atotal of 284 bands were obtained of which 200 (70.4%) were polymorphic with the average of50 polymorphic bands. For four TE-AFLP primer combinations, a total of 192 band wereobtained of which 139 (72.4%) were polymorphic with the average of 34.7 polymorphicbands. Our results indicate that the genetic information of AFLP is similar to that ofTE-AFLP, and little less than that of ISSR, but only about 1/3 of that of SSR. However, theability of information detection of the two AFLP-based markers is much higher than that ofISSR and SSR. In addition, the cluster analysis of AFLP, TE-AFLP and combined data revealthat: P. yunnanensis, P. trinervis and P. simonii clustered together, and P. trinervis and P.simonii showed more closed affinity; the other four poplar species clustered together, P.cathayana and P. schneideri showed more closed origin especially. 3. The cpSSR analysis for seven Populus species belonging to the Section Tacamahaca.Four out of the seven analyzed chloroplast loci were polymorphic, whereas none of the lociwere polymorphic across the accessions within a species. 13 bands and 4 different kinds ofhaplotypes were reduced. Based on 5 pairs of cpDNA primers, 73 fragments (52 polymorphic)and 9 kinds of haplotypes were produced. However, none of the polymorphic was detected bythe 5 mtDNA primer pairs, revealing conservation of mtDNA region. The cluster analysis ofcpDNA revealed that: similar maternal phylogeny among P. yunnanensis, P. trinervis and P.simonii; the other four species clustered together, P. schneideri and P. kangdingensis showedmore closed maternal lineage especially. 4. Our molecular data, morphological characters and nature habitat revealed that: sameto the traditional taxonomy assignment, genetic variation within a same Populus species islower than that among Populus species in Section Tacamahaca; P. yunnanensis may share itschloroplast ancestor with P. trinervis and P. simonii; moreover, sister genetic relationship of P.trinervis and P. simonii indicated their similar origin; P. schneideri clustered with P.kangdingensis and P. cathayana, respectively, and may have derived from an ancienthybridization event involving the ancestors of the two species.

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随着全球气候变暖和温室效应加剧,干旱和荒漠化成为威胁人类生存和发展的主要 灾害,许多被子植物对干旱胁迫的生理、生态和生化响应已逐步得以报道,但很少有开 展干旱胁迫对雌雄异株植物的影响方面的研究。由于这类植物在长期进化过程中已经在 生长、性比、生殖格局、空间分布、资源配置和生物量分配等方面形成了明显的性别差 异,因此,干旱胁迫必将对其雌雄植株产生不同的生理生态影响。本研究以青杨为模式 植物,采用植物生态、生理及生物化学等研究方法,系统研究青杨雌雄植株在常温、增 温以及喷施外源脱落酸的条件下对干旱胁迫的响应,揭示其在生长形态、生物量分配、 光合作用、用水效率和生理生化等方面的性别间差异。主要研究结果如下: 1. 青杨雌雄植株对干旱胁迫的综合响应。 与较好水分条件相比,干旱胁迫显著降低了青杨雌雄植株的光合作用和生长发育, 影响了许多生理生化过程,并导致雌雄植株在生长发育、气体交换、用水效率、膜脂抗 氧化和抗氧化系统酶活性方面表现出显著的性别间差异。在较好水分条件下,雌雄植株 之间在株高、基径、生物量、净光合速率、蒸腾速率、用水效率以及丙二醛、脱落酸和 游离脯氨酸等生化物质含量方面均无显著差异。但在干旱胁迫下,雄株在生长发育、气 体交换、水分利用效率、膜脂过氧化保护和抗氧化系统酶活性方面均显著高于雌株,表 现出比雌株更高的株高、基径、叶面积、总叶片数、总生物量、总色素含量、类胡萝卜 素含量、净光合速率、蒸腾速率、羧化效率、光系统II最大光化学效率、内在水分利用 效率、碳同位素组分、过氧化氢酶和过氧化物酶活性等,而在CO2补偿点、比叶面积、 叶绿素a/b、丙二醛、脱落酸和超氧化物歧化酶活性等指标上显著低于雌株。与雌株相比, 雄株表现出更高的干旱胁迫适应能力,而雌株的生长发育和生理生化过程更易遭受干旱 胁迫的影响。 2. 干旱胁迫下的青杨雌雄植株对增温处理的综合响应 与环境温度相比,增温在干旱胁迫前后均显著促进了雌雄植株的生长发育、气体交 换,降低水分利用效率,影响生化物质含量,并促使青杨雌雄植株之间在干旱胁迫下表 现出显著的差异。在较好水分条件下,增温导致雌株的株高、基径、叶面积、总叶片数、 总生物量和超氧化物歧化酶活性显著高于雄株,而用水效率、丙二醛、脱落酸和游离脯 氨酸、抗坏血酸过氧化物酶和过氧化物酶活性低于雄株。在干旱胁迫下,增温将导致雄 株的株高、基径、叶面积、总生物量、净光合速率、蒸腾速率、气孔导度、总色素含量、 相对含水量、过氧化氢酶和抗坏血酸过氧化物酶活性等显著高于雌株,而光系统II 最大 光化学效率、内在水分利用效率、碳同位素组分、丙二醛、脱落酸、游离脯氨酸和超氧 化物歧化酶活性显著低于雌株。与雄株相比,水分较好条件下的增温有利于促进雌株的 生长发育,并在生理生态特征上优于雄株。而干旱胁迫下的增温则加剧了水分胁迫强度, 致使雌株的生长发育遭受比雄株更多的负面影响。 3. 干旱胁迫下的青杨雌雄植株对喷施外源脱落酸处理的综合响应 与对照相比,在干旱胁迫下喷施外源脱落酸可显著增加青杨雌雄植株的生长发育、 气体交换、降低水分利用效率,影响了生化物质含量,并导致青杨雌雄植株之间在干旱 胁迫下表现出显著的生理生态差异。在干旱胁迫下,喷施外源脱落酸致使雌株的株高、 叶面积、叶干重、细根干重、总生物量、净光合速率、蒸腾速率、气孔导度、光系统II 最大光化学效率、非光化学淬灭系数、相对含水量、总光合色素、类胡萝卜素、脱落酸、 超氧化物歧化酶和过氧化物酶活性的增加量显著高于雄株,而根重比、根冠比、细根/ 总根、比叶面积、内在水分利用效率、碳同位素组分、丙二醛、脯氨酸、过氧化氢酶和 抗坏血酸过氧化物酶活性等指标的减少量上显著低于雄株。与对照相比,干旱胁迫下的 喷施外源脱落酸则一定程度能减缓植株遭受胁迫的压力,促进植株生长和气体交换,减 少了植株体内的过剩自由基数量,并促使雌株的生长发育和光合能力显著提高,增强其 抗干旱胁迫能力。 With development of global warming and greenhouse effect, drought and desertification have been became main natural disasteres in resent years. Studies on ecophysiological responses of most angiosperm species to environmental stress have been reported, but little is known about dioecious plant responses to drought stress. Since significant differences on growth, survival, reproductive patterns, spatial distribution, as well as resource allocation between males and females of dioecious plant have been formed during evolutionary process, sexual different ecophysiological responses should be caused by drought stress. In this experiment, Populus cathayana Rehd. was used as model plant to study the sex-related responses to drought by using the ecological, physiological and biochemical methods under normal atmospheric temperature, elevated temperatures and exogenous abscisic acid (ABA) application treatment respectively, and to expose the sexual differences in growth, biomass allocation, photosynthesis, water use efficiency and some biochemical material contents in the males and females of dioecious plant. The results are follows: 1. A large set of parallel responses of males and females of P. cathayana to drought stress Compared with well-watered treatment, drought significantly decreased growth and photosynthesis of P. cathayana individuals, affected some physiological and biochemical processes, and induced males and females to exhibit obvious sexual differences in growth, gas exchange, water use efficiency, lipid peroxidation protection and antioxidant defenses enzyme system. Under well-watered treatment, there were no significant sexual differences in height growth (HG), basal diameter (BD), dry matter accumulation (DMA), net photosynthesis rate (A), transpiration (E), water use efficiency (WUE), and malondialdehyde (MDA), abscisic acid (ABA) and praline (Pro). However, under drought stress, males were found to exhibit higher HG, BD, leaf area (LA), total leaf number (TLA), DMA, total chlorophyll contents (TC), carotenoids content (Caro), A, E, carboxylation efficiency (CE), the maximum efficiency of PSII (Fv/Fm), intrinsic water use efficiency (WUE ), carbon isotope composition (δ13C), catalase (CAT), peroxidase (POD) and lower CO2 compensation point (Γ), specific leaf area (SLA), chlorophyll a/b ratio (Chla/Chlb), MDA, ABA and superoxide dismutase (SOD) than females. The results suggest that males possess greater drought resistance than do females and females suffer more negative effect on growth and development, physiological and biochemical processes than males under drought stress. 2. A large set of parallel responses of drought-stressed males and females of P. cathayana to elevated temperatures Compared with environmental temperature, elevated temperature treatment significant increased growth and gas exchange, decreased water use efficiency, changed some biochemical material contents of P. cathayana individuals, and induced males and females to exhibit obvious differences under drought stress. Under good water condition, elevated temperature treatment caused females to show significant higher HG, BD, LA, TLN, DMA, SOD activity, and great lower WUE, MDA, ABA, Pro, ascorbate peroxidase (APX) and POD than do males. On contrary, under drought condition, elevated temperature treatment induced males to exhibit higher HG, BD, LA, DMA, A, E, stomatal conductance (gs), relative water content (RWC), CAT, APX activity but lower Fv/Fm, WUE, δ13C, MDA, ABA, Pro, SOD activity than do females. The results suggest that females will benefit from elevating temperature under good water condition by possessing better ecophysiological processes than that of males, but will suffer from greater negative effects than do males when grown under drought stress with elevated temperature treatment. 3. A large set of parallel responses of drought-stressed males and females of P. cathayana to exogenous ABA application Compared with controls, exogenous ABA application under drought greatly increased growth and gas exchange, decreased water use efficiency, changed some biochemical material contents in P. cathayana individuals, and induced males and females to exhibit obvious sexual differences under drought. Under drought stress, exogenous ABA application induced females to exhibit more increases in HG, LA, leaf weight (LW), fine root weight (FRW), DMA, A, E, g, Fv/Fm, non-photochemical quenching coefficient (qN), RWC, TC, Caro, ABA, SOD, POD s activity than males, but to show lower decreases in root/weight ratio (RWR), root mass/foliage area ratio (RF), fine root/total root ratio (FT), SLA, WUE, δ13C, MDA, Pro, CAT, APX than males. The results suggest that exogenous ABA application under drought stress will eliminate negative damages caused by drought stress at a certain extent,promote the growth and gas exchange of plant and decrease the number of superfluous 1O2 in plant cells of males and females of P. cathayana. Furthermore, exogenous ABA application promoted more drought resistance in females than in males by increasing more growth and photosynthetic capacity in females under drought stress.

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对隆肛蛙属的物种构成进行了订正,建立新属肛刺蛙属Yerana gen. nov.;订正后的隆肛蛙属现仅隶2种, 即隆肛蛙F. quadrana和太行隆肛蛙F. taihangnicus。运用形态学分析探讨了隆肛蛙属物种及种群的形态差异和分类关系,通过分子系统学研究探讨了隆肛蛙属物种及种群的分类和系统发育关系,运用动物地理学方法结合系统发育关系探讨了隆肛蛙属种群的地理分布格局成因与历史过程。主要结果和推论如下: 1.隆肛蛙属物种构成的订正及一新属建立 建立新属肛刺蛙属,将隆肛蛙属中的原叶氏隆肛蛙F. yei归隶新属肛刺蛙属并更名为叶氏肛刺蛙Y. yei,,新属建立的主要依据为:(1)雄性肛部隆起,肛孔下方有两个布满黑刺的大的白色球形隆起,具单咽下内声囊, 第一指具婚刺;(2)形态量度分析表明叶氏肛刺蛙与隆肛蛙和太行隆肛蛙的形态差异远大于后两者之间的差异;(3)叶氏肛刺蛙的分布区与隆肛蛙和太行隆肛蛙的分布区距离较远且呈隔离状态;(4)分子系统学研究资料(Jiang et al.,2005)证明叶氏肛刺蛙与隆肛蛙和太行隆肛蛙非单系发生;叶氏肛刺蛙在第二支中位于基部。因此,隆肛蛙属现仅隶2种,即隆肛蛙和太行隆肛蛙。 2.隆肛蛙属种群形态学研究 对隆肛蛙属中隆肛蛙和太行隆肛蛙的15个地理种群565只标本的28项形态性状进行了测量,运用典型判别分析法对其分析的结果表明:(1)太行隆肛蛙与隆肛蛙形态差异明显,支持其为不同的物种;(2)原隆肛蛙河南伏牛山种群和山西中条山种群应为太行隆肛蛙的地理种群;(3)隆肛蛙不同地理种群之间形态差异明显,其中四川安县种群、陕西周至种群和湖北利川种群与模式产地重庆巫山种群的差异可能达到了亚种或亚种以上分化水平。对隆肛蛙属量度分析的15个种群进行定性形态分析表明其分为三种形态型,对应隆肛蛙、过渡型和太行隆肛蛙,其变异特征主要为内跗褶、雄性肛部隆起及疣粒分布、第五趾外侧缘膜等,这与量度分析结果相似。 3.隆肛蛙属种群分子系统学研究 测定隆肛蛙属Feirana的2种19种群的线粒体12S rRNA和16S rRNA基因片段、ND2基因的DNA序列,比对后共计1953bps。(1)遗传多样性与距离分析:结果表明,隆肛蛙属种群具很高的遗传多样性,19个种群样品表现出19种单倍型(遗传多样性指数Hd=1.0); ND2基因的进化信息含量远高于12SrRNA和16SrRNA。隆肛蛙属2种群组内的种群间的遗传距离远小于两种群组间的距离,种群在不同基因上的遗传距离表现的关系与对应的系统树一致。(2)系统发育关系分析:结果表明,不同基因片断基于不同方法构建的隆肛蛙属种群系统发育树结构基本一致,基本表明隆肛蛙属种群为单系发生;它们在系统树中分为两大支,分别对应于隆肛蛙和太行隆肛蛙;支持中条山种群(沁水、历山和济源种群)和伏牛山种群(栾川和内乡种群)为太行隆肛蛙的地理种群,而原隆肛蛙秦岭中东段的部分种群(柞水、宁陕、长安大坝沟种群)也应为太行隆肛蛙的地理种群。(3)亚种分化分析:根据遗传距离分析和系统发育关系分析结果,并考虑形态上的差异情况以及地理分布信息,隆肛蛙所隶种群组可分为2亚种,即隆肛蛙指名亚种F. quadrana quadrana包括四川盆地东缘大巴山东段-巫山-武陵山北麓种群和秦岭中段(周至板房子和长安广货街)种群,他们在系统关系树上聚为一支;安县亚种F. quadrana anxianensis包括四川盆地西缘岷山东麓-龙门山-大巴山和秦岭西段的种群(安县、青川、文县、南江和凤县种群),他们在系统关系树上聚为一支。太行隆肛蛙所隶种群组也可分为2亚种,即太行隆肛蛙指名亚种F. taihangnicus taihangnicus包括中条山的种群(沁水、历山和济源种群)和中东秦岭的部分种群(柞水、长安大坝沟和宁陕种群),他们在系统关系树上聚为一支;太行隆肛蛙伏牛亚种F. taihangnicus funiuensis,为伏牛山地区的种群(栾川和内乡种群),他们在系统关系树上聚为一支。 4.隆肛蛙属种群动物地理学研究 隆肛蛙属19种群的分歧年代分析: 以长江巫山段和黄河三门峡段的形成历史时期为参考点,根据已测隆肛蛙属19种群及其外群包括N. pleski、P. yunnanesis、P. robertingeri、F. limnocharis的1953bps DNA序列构建分子钟,获得各支系的分歧年代。结果表明:①棘蛙族在70Ma左右开始其独立演化历程,这与Roelants et al.(2004)的分析结果~60±15Ma左右开始分化基本一致,后者印证了本文的分子钟。②隆肛蛙属的起始分化年代较早,隆肛蛙和太行隆肛蛙两种群组的最近祖先种群大概在46Ma~50Ma左右;隆肛蛙和太行隆肛蛙种群组内的种群分化年代相对两种群组间晚得多, 隆肛蛙种群组内两亚种分化起始年代约为10Ma左右,而太行隆肛蛙种群组内两亚种分化起始年代约为6Ma。 隆肛蛙属种群分布格局形成过程分析: ①隆肛蛙属的系统关系与地理分布格局密切相关,大部分系统分支分级与地理距离成正比;②隆肛蛙属最近祖先种群的分化中心可能位于秦岭中部地区, 隆肛蛙属的种群分布格局的形成表现为隔离分化与扩散相结合的机制,由隔离分化产生的隆肛蛙祖先种群主要从秦岭中部向西南方向扩散,后隔离分化为两亚种;太行隆肛蛙祖先种群向东北方向扩散也分化为两亚种。 隆肛蛙属种群分布区域地质历史的探讨:本文所建分子钟和种群分化方式印证了该区域的几次主要地质事件,包括岷山-龙门山-西秦岭等地区的快速差异隆起、第四纪冰期等。 The specific composition of the genus Feirana should be revised. A new genus Yerana gen. nov.(Ranidae:Dicroglossinae)was established based on morphological data-set and molecular phylogeny, as a result, only two species F. quadrana and F. taihangnicus are classified into Feirana now. Morphological differences and taxonomy of populations of Feirana were investigated based on morphological and morphometric data; phylogenetic relationships and taxonomy of populations of Feirana were elucidated using molecular data, and then the proceeding of the distribution pattern of populations of Feirana were discussed. The main results and conclusions and proposals were presented as following: 1. Revising of the specific composition of the genus Feirana and establishment of a new genus The new genus Yerana, only containing the type species Y. yei, was established based on the following evidences: (1) In adult male, distinct up-heaved circular vesicle presents around the anal, and under anal there are two white balls on which black spines exist, black horny spines scatter on the upper side of first finger, and internal single subgular vocal sac presents; (2) there is obvious morphometric differences between Yerana and Feirana; (3) Yerana is distributed far from Feirana; (4) evidences of molecular phylogeny(Jiang et al.,2005)suggested that Yerana take a special phylogenetic clade which is different from other genus included in the tribe Paini. As a result, there are only two species in Feirana, i.e., F. quadrana and F. taihangnicus. 2. Morphological research of populations of Feirana Twenty-eight characters of 565 individuals of 15 populations of the genus Feirana were measured, the results of Canonical Discriminant analysis of the morphometric data-set indicated that: (1) there are very prominent differences between the two species F. quadrana and F. taihangnicus. The validity of species F. taihangnicus was approved here; (2) Mt. Funiu population and Mt. Zhongtiao population should belong to the species F. taihangnicus; (3) Obvious differences exist among 12 populations of F. quadrana, the differentiation among Zhouzhi population, Anxian population, Lichuan population, and Wushan population together with the others probably reach sub-specific or specific level. Result of morphological comparison between 15 different populations show that 3 morphological types are recogenized in according with F. quadrana, F. taihangnicus and intergradation, this result conform to the result of morphometric analysis. 3. Molecular phylogenetic study on populaions of Feirana Fragment of 12SrRNA and 16SrRNA genes, and ND2 gene of 19 populations of two species of Feirana were sequenced and aligned, from which 1953 bps were received. (1) analyses of genetic distance and hereditary diversity indicated that: genetic distance between populations in each group were less than distance between two groups of Feirana, 19 haplotypes were recognized from 19 samples of 19 populations, so the hereditary diversity of populations of Feirana was very high (Hd=1.0), phylogenetic information in ND2 gene is more than fragment sequence of 12SrRNA and 16SrRNA genes. (2) Result of molecular phylogeny indicate that the phylogenetic trees constructed using different methods based on different sequence data sets showed the revised genus Feirana is monophyletic since the 19 populations of Feirana were firstly clustered together as one large clade, which was further clustered into two major clades, corresponding to F. quadrana(GroupⅠ) and F. taihangnicus(GroupⅡ), respectively. So populations of Qinshui and Lishan in Mt. Zhongtiao, populations of Luanchuan and Neixiang in Mt. Funiu, and populations of Zhashui, Dabagou of Chang’an and Ningshan in eastern Mt. Qinling should belong to the species F. taihangnicus; (3) Subspecific differentiation. on the basis of genetic distance, phylogenetic trees and geographical distribution, F. quadrana should have two subspecies, i.e., F. quadrana qudadrana, consisting of the populations Guanghuojie of Chang’an and Zhouzhi in Mid-Mt. Qinling, populations in Wushan area and northern Mt. Wuling (Lichuan), and F. qudadrana anxianensis, consisting of the populations in eastern Mt. Ming shan-Mt. Longmen-western Mt. Daba-western Mt. Qinling (Anxian, Qingchuan, Wenxian, Nanjiang and Fengxian); F. taihangnicus should also has two subspecies, i.e., F. taihangnicus taihangnicus, consisting of the populations in Mt. Zhongtiao and eastern Mt. Qinling, and F. taihangnicus funiuensis, consisting of the populations in Mt. Funiu. 4. Zoogeography of populaions of Feirana Analysis for divergent time of 19 populations of Feirana: Using the dates of run-through of Wushan segment of Changjiang River as the time when the population of Lichuan started differentiated from the populations of Wushan and Shennongjia, and the dates of Sanmenxia segment of Yellow River as the time when the populations in Mt. Zhongtiao started differentiated from the population of Dabagou in Chang’an, molecular clock was established using sequences with 1953 bps of 19 populations of Feirana and outgroup including N. pleski, P. yunnanesis, P. robertingeri, F. limnocharis in order to estimate divergent time of all clades. Result of that indicated that: ① the tribe Paini started to evolve independently at about 70Ma when is in consistent with that estimated by Roelants et al.(2004)with result of about ~60±15Ma, they were corroborated by each other, this confirms the validity of this molecular clock; ② divergent time for speciation of Feriana is early, ancestral populations of F. quadrana and F. taihangnicus were found about 46Ma~50Ma; differentiation of populations within species is greatly late to the divergence of the two species, divergent time for F. quadrana is 10Ma and divergent time for F. taihangnicus is 6Ma. Proceeding of distribution pattern of Feirana. Phylogenetic relationships of populations of Feirana matched quite with distribution pattern of them, the relationships among clades showed in phylogenetic trees is direct ratio to geographical distance of them; the estimated date of speciation between two species of Feirana was as early as speciation of Paa yunnanesis and Nanara pleski; middle part of Mt. Qinling is the center of speciation of Feirana, combination of mult-events of dispersal and vicariance are probably the mechanism of speciation of Feirana, F. quadrana colonized the mid-Mt. Qinling and then differentiated into two subspecies in southwest direction, ancestral population of F. taihangnicus colonized the mid-Mt. Qinling and then differentiated into two subspecies in northeast direction. On geological history of the distribution of Feirana. According to molecular clock and speciation model of populations of Feirana, some geological events are confirmed, including special rise of Mt. Minshan- Mt. Longmen-western Mt. Qinling, glacial age.