The uplift of Qinghai-Xizang (Tibet) Plateau and the vicariance speciation of glyptosternoid fishes (Siluriformes : Sisoridae)

Based on the phylogenetic and biogeographical studies of the glyptosternoid fishes in Qinghai-Tibet area, the following hypothesis is proposed: the speciation of this group has a direct relationship with the three uplift intervals of the Qinghai-Tibet Plateau. This process was explained by the theory of vicariance of biogeography. The ancestor of this group was similar to Bagarus and/or Glyptothorax, which still have a wide distribution. At the moment when the Tethys sea closed, the Indian tectonic plate collided with the Eurasian tectonic plate, so the Glyptothorax-like and Bagarus-like ancestors entered Eurasia and gradually became widely distributed. After the Pleistocene, with the enforced colliding, the gradual uplift of the Qinghai-Tibet Plateau brought about the current water environment, and the Glyptosternoids were generated from Glyptothorax-like fish under this environment. The present Glyptosternum, distributed across the Himalayas is the ancestor of Glyptosternoids. In the three uplift intervals of the plateau, the water system of this region was separated gradually and Glyptosternum-like ancestor was isolated in different rivers and evolved into various species. All this resulted in the speciation and formation of the biogeographical pattern of glyptosternoids.

Suckers (Fish, Catostomidae) from the Eocene of China account for the family's current disjunct distributions

Unequivocal Eocene suckers from China are for the first time reported here. This discovery demonstrates that catostomids of the Eocene Epoch (some 55-35 Ma ago) are scattered widely on mainland Asia as well as western North America. The present day disjunct distribution pattern of catostomids, with 68 extant species widespread in North America and the northern part of Middle America and only two in the restricted areas of Asia, is the result of their post-Eocene decline in Asia due to the competitive pressure from cyprinids, their Late Cenozoic radiation in North America, and the vicariant and dispersal events triggered by the changed biogeographic landscape. All of these prove to be a historical product of the geological, biological, and climatic changes throughout the Cenozoic.

OCCURRENCE AND DISTRIBUTION OF SILICA-SCALED CHRYSOPHYTES IN ZHEJIANG, JIANGSU, HUBEI, YUNNAN AND SHANDONG PROVINCES, CHINA

Fifty-one phytoplankton samples were collected from different ecological environments in five provinces located in the central, eastern and southwestern parts of China. 41 taxa of silica-scaled chrysophytes, 8 of them belonging to the Chrysophyceae and 33 to the Synurophyceae, have been identified by means of electron microscopy. Among them, Mallomonas parvula, M. calceolus, M. cyathellata (?), M. costata, M. crassisquama, and Synura echinulata are new to China. The silica-scaled chrysophyte flora and its geographical distribution in this region are analysed and discussed.

澜沧江姬鼠（Apodemus ilex）种群遗传结构及其生物地理分析

澜沧江姬鼠（Apodemus ilex）为小型啮齿类动物，长期以来常被认为是中华姬鼠（A. draco）或长尾姬鼠（A. orestes）的同物异名，但近年基于线粒体Cyt b 的研究证明，该类群与中华姬鼠的遗传距离达到了种间水平，并被认为是一个独立种。本文提取、扩增并测定了来自云南14个地区42个产地156个澜沧江姬鼠的Cyt b 全序列，并将GenBank中的3条澜沧江姬鼠的Cyt b 全序列纳入分析，选取福建武夷山与四川峨眉山的2个中华姬鼠样品的Cyt b 全序列，以及台湾姬鼠、大耳姬鼠、高山姬鼠Cyt b 全序列作为外群，利用贝叶斯法和最大似然法构建系统发育树，并使用Network分析，分子变异分析（AMOVA）分析、错配分析（Mismatch analysis）与Fu`Fs检验等方法，探讨了这些地区澜沧江姬鼠的种群遗传结构，种群动态及生物地理格局。结果表明：云南地区过去被鉴定为中华姬鼠的样品均属于澜沧江姬鼠，澜沧江姬鼠与中华姬鼠间的遗传距离（K2P-ds+v）达到0.091，显著超过姬鼠属内种间差距（0.080），支持澜沧江姬鼠独立种的地位；该种不同地理种群可大致以澜沧江为界分为两大支系，且每个支系又各分为两个亚支系；进一步的AMOVA分析表明遗传变异主要来源于两大支系之间，而错配分析与Fu`Fs检验表明澜沧江姬鼠各地理种群及其两个支系都经历过多次扩张事件，这可能是和冰期与间冰期的轮回交替有关；分子系统树和中介网络图表明地理环境的隔离对于澜沧江姬鼠种群分化存在明显影响，其中澜沧江的阻隔作用最为显著。基于松散分子钟（Relax molecular clock）分析，中华姬鼠和澜沧江姬鼠的分歧时间大约在2.98 Ma（95％HPD: 4.32-1.88），而澜沧江姬鼠则在1.13 Ma（95％HPD: 1.95-0.65）分成两个支系。按照平均分歧时间来看，澜沧江姬鼠两个支系的的分歧事件发生在鄱阳冰期（1.2-0.9 Ma），两个支系内部进一步分化为两支的时间非常接近，分别为：Clade 1中为0.67 Ma （95％HPD: 1.17-0.34）和Clade 2 中为0.64 Ma (95％HPD: 1.17-0.36)，这种隔离种群各自产生分化的时间却非常接近，这提示其分化事件可能是受到古气候如冰期或间冰期的影响所产生。

鰋鮡鱼类(Glyptosternoid fishes)的系统进化和动物地理学研究

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。

九寨沟湖泊湿地植物群落与植物地理研究

九寨沟湖泊湿地在维持九寨沟的生态平衡中起着重要的作用，在旅游产业的发展下，湿地生态系统及生物多样性面临着较大的威胁。尽管九寨沟湿地具有重要的生态价值，但目前对其研究尚比较薄弱。湿地植物群落和植物地理研究可以为湿地资源的可持续利用和监测保护提供科学依据。作者从2004年8月到2007年11月对九寨沟湿地的植物物种组成、地理分布、优势植物群落的结构、生长动态、湿地土壤种子库进行了调查研究。主要结果如下： 1. 九寨沟湿地物种组成、地理分布特点及湿地植物群落特点 九寨沟湿地共有苔藓植物8科13属16种，维管植物为48科107属199种。九寨沟湿地植物的地理成份较为丰富，维管植物在科级水平上有7种地理分布型（变型），在属级水平上有13种地理分布型（变型）， 在种级水平上共有29种地理分布型（变型）。九寨沟湿地植物以温带成份和我国特有成份为主，同时兼有热带、亚热带成份和环极—高山成份。九寨沟湿地植物的分布表现出明显的垂直地带性和水平地带性。湿地植物群落可划分21个群落类型，不同植物群落类型的物种多样性及物种组成存在较大的差异。九寨沟湿地植物的物种多样性和群落多样性以及较高的生产力特征，是维持其湿地生态景观多样性和稳定性的基础。 2. 土壤、水环境、海拔等对湿地植物的分布及生物多样性的影响 九寨沟湿地土壤、水等环境因子存在较大的差异。帕米尔苔草和宽叶香蒲等群落的凋落物较多，土壤有机碳、土壤总磷较高，可能是九寨沟湿地的重要土壤碳库。 九寨沟湿地植物沿水环境梯度的分布规律表现为：沉水植物（轮藻—篦齿眼子菜，水苦荬，杉叶藻）——挺水植物（水木贼，芦苇，宽叶香蒲）——湿生草本（苔草、节节草、披散木贼）——湿生灌木（柳灌丛，小檗灌丛）等。海拔也影响湿地植物的物种组成。 水深对物种多样性有影响，水深与物种丰富度负相关。随着水深的增加，水木贼、芦苇、杉叶藻、宽叶香蒲等群落的物种多样性下降；在长期淹水和季节性淹水的地方，水木贼群落物种多样性存在显著差异。土壤总氮与水木贼群落物种丰富度正相关。 3. 土壤营养元素、水环境对植物生长的影响 水深影响湿地植物生物量的分配。芦苇无性系分株在47 cm水深的环境中单株平均生物量最大；在干滩地中（地面水深0 cm），叶生物量百分比最大，而茎生物量百分比最小，茎的生物量百分比和生长速率随水深的增加而增加；在较干的滩地生境中，开花率、花序的生物量百分比明显大于水较深的生境。 水深与水木贼地上生物量负相关，但水木贼地上生物量在长期淹水和季节性淹水的地方没有显著的差异。在水浅的地方，杉叶藻、水木贼、芦苇等植物群落中，其他伴生物种的生物量占样方总生物量的百分比较大。 土壤有机碳、土壤总氮、土壤总磷等对湿地植物生物量的影响比较大：宽叶香蒲地上生物量与土壤总磷正相关；水木贼地上生物量与土壤总氮正相关；杉叶藻地上生物量与土壤有机碳正相关。 水深、土壤营养成分对湿地植物高度、密度等有影响。水木贼的平均高度在季节性淹水的地方比长期淹水的地方低，平均密度在长期淹水的地方比季节性淹水的地方低；除了5月份，其他观察月份水木贼的密度都与水深负相关，同时与土壤有机碳正相关。另外，芦苇密度与土壤有机碳含量正相关，宽叶香蒲密度与水深负相关，帕米尔苔草高度与土壤有机碳负相关。 4. 优势植物群落的动态变化 在优势植物群落中，优势种的高度、密度、盖度、生物量等在群落中占绝对优势。除五花海，水木贼群落的物种组成、高度、生物量在两年间没有显著的变化。芦苇群落的物种丰富度在近两年有所增加。 湿地植物生长表现为明显的季节动态，生长的峰值大多在7月－8月。优势植物群落的物候与水文周期有关。湿地植物群落的物种组成和密度，可以作为对湿地监测和保护的生物指示。 5. 九寨沟湿地土壤种子库特征及其在湿地生物多样性恢复中的作用 水深和现存植被物种丰富度可以解释湿地土壤种子库的变化。水深可以解释表层物种丰富度45%的变化。现存植被物种丰富度可以分别解释10 cm土层、2－5 cm土层及5－10 cm土层土壤种子库45%、48%和25%的变化。 湿地土壤种子库的密度为0－15945粒m-2, 种子库中共发现23个物种。现存植被优势物种和种子库优势物种不同。各层土壤种子库密度和物种丰富度并不存在显著的差异，但第二层土壤种子库密度最大。海拔、现存植被优势种盖度、土壤总磷、土壤总氮、土壤有机碳对湿地土壤种子库的密度和垂直结构没有影响。土壤种子库物种丰富度小于地上植被物种丰富度。湿地土壤种子库与地上植被的相关性不大。在浅水区域，湿地土壤种子库在湿地植被恢复中有一定作用。但在深水区域，保护现存植被更重要。 The lakeshore wetlands are valuable ecological units of the Jiuzhaigou lakes. Pressure for travel industry development pose a continuing and severe threat to the biodiversity-support function of the wetland system. Despite the ecological importance of wetlands in Jiuzhaigou, they are so far poorly studied. Both general plant communties and biogeographical studies are needed in order to attain basis for sustainable use the wetland resources and adequate protection of these areas. The present study was undertaken to examine aquatic plants distribution and the species compositon, structure and growth dynamics of their communities with variations of environmental factors along altitudes, water depth and soil properities gradients in Jiuzhaigou. Analysis of field survey data collected during August 2004 and November 2007 in lakeshore wetlands in Jiuzhaigou National Nature Reserve, Sichuan, China. The results were as following: (i) Species composition and biogeography in wetland vegetation 8 families, 13 genus, 16 species of moss and 48 families, 107 genus and 199 species of vascular plants in Jiuzhaigou wetlands were found. The floristic compositions were abundunt. Ten geographical distribution types at family level, 13 geographical distributions types at generic level and 29 geographical distribution types at specific level in vascular plants were found. Most species in Jiuzhaigou wetlands are temperate elements and Chinese endemic elements, with a few of tropical and subtropical and some circumarctic elements. And the plant distributions show clear vertical and horizontal patterns. There were 21 major wetland plant community types. Species composition and species richness in different plant communities are different. The species diversity and plant community diversity and their high biomass are the basis for the diversity and stability of wetland landscapes in Jiuzhaigou. (ii) Water depth, soil nutrients and altitudes influence on the species diversity and plant distribution. Total phosphorous and organic cabon in soil were higher in C. pamiernensis and T. latifolia communities, where are important cabon reservoirs in Jiuzhaigou wetlands. Along gradients of water depth, among populations of the dominant plant species present: submerged macrophytes (Chara vulgaris, Potagemonton pectinatus, Veronica anagalis-aquatica，Hippuris vulgaris), emergent macrophytes (Equisetum fluviatile, Phragamites australis, Typha latifolia), helophytes (Carex pamirensis )and shrubs (Salix sp., Berberis sp. ). Altitudes influence on the assemblage of plant communities. Water depth negatively correlated with species richness. Specie richness showed differences between permanently flooded sites and seasonally flooded sites in E. fluvatile communities. And total nitrogen in soil was negatively correlated with species richness in E. fluviatile communities. Altitudes show no significant influence on species richness, but in fact, through our analyses, they do have influence on the assemblage of wetland plants. (iii) Water depth, soil nutrients influence on the plant growth Water depth influences the biomass allocation in Phragmities australis. The average aboveground biomass of a single ramet (4.2 g) was the largest in the habitat with water level 47 cm above the soil surface. At the habitat with water level under soil surface 15 cm (－15 cm), the leaf biomass percentage (of the total ramet biomass) was the largest (46.1%), and the height and percentage of ramose ramets ( with branches on stem )(of the total ramets in a plot) were found obviously different. The deeper in water, the larger the biomass percentage and growth rate of stems were. The flowering rate and biomass of panicles were greater in shallow water than those in deep water. Water depth negatively correlated with aboveground biomass of E. fluviatile. However, above-ground biomass of E. fluviatile showed no significant difference between permanently flooded sites and seasonally flooded sites. But in shallow water, more biomasses of accompanying species were found in dominant plant communities such as H. vulgaris communities, E. fluviatile communities and P. australis communities. Water depth, soil nutrients influence on shoot density and shoot length of wetland plants. The shoot density of E. fluviatile was correlated to water depth in all growth months. Annual average density was significantly lower at permanently flooded sites than at seasonally flooded sites. But the annual average shoot length was significantly lower at seasonally flooded sites than at permanently flooded sites. (iv) Growth dynamics of dominant communities in Jiuzhaigou wetland The shoot length and shoot density, coverage and biomass of domiant species were dominated in plant communities. The species composition increased in P. australis communities in recent two years. The species richness in E. fluviatile communities showed no difference between 2005 and 2007. The above-ground biomass and shoot density in Five－flower Lake from July 2005 to July 2007 were significantly different, while in other sites, the differences were not significant. Shoot height, shoot density and above-ground biomass showed significant seasonal changes in all sites. Growth dynamics correlated with the cycle of water levels in lakes. Most plants growth parameters peaked at July or August. The biomass of T. latifolia peaked in August. But the shoot length of T. latifolia in deeper water peaked in July. The shoot length of E. fluviatile increased significantly from May to August except in seasonally flooded sites in Arrow-bamboo Lake. The species composition of communities and shoot density can be used as bioindicators in Jiuzhaigou wetland. (v) Soil seed bank in Jiuzhaigou wetland and its role in vegetation restoration Seed density in all soil layer samples was negatively correlated to water depth. Water depth can explain 45% variance of species richness in surface layer in sediment. Species richness in extant vegetation can explain 45%, 48%, 25% variance of species richness in total 10 cm and in 2－5 cm and 5－10 cm layer sediment respectively. Mean seed densities in wetlands ranged from 0 to 15945 m–2. A total of 23 species germinated in seed bank. The dominant species in seed bank and extant vegetation showed great difference. The total number of species and seedlings that germinated in different layers was not significantly different. But the second layer had the greatest seed density. In shallow water, seed bank can contribute to vegetation restoration, while in deeper water, protection of extant vegetation may be a better strategy.

角蟾科（Megophryidae; Anura）蝌蚪的形态多样性和生态适应

角蟾科（Megophryidae）是以角蟾属（Megophrys Kuhl and Van Hasselt, 1822）为模式属而建立的，隶于无尾目（Anura），变凹型亚目（Anomocoela）。角蟾科包括2 亚科11 属142 种，分布于东洋界，从巴基斯坦、中国西部向东直到菲律宾和苏达群岛；中国有9 属75 种分布于华中和华南地区。角蟾科被认为是原始的两栖动物之一，其分类学、系统学、生态学、动物地理学的研究均深受中外科学家的瞩目。近年来，通过形态学、古生物学、细胞学、生态学、支序系统学的研究，角蟾科的分类与系统学研究取得了较大进展。与成体形态和分子系统学研究结果相比较，蝌蚪的研究存在更多的问题和挑战，尚需深入研究：（1）角蟾科蝌蚪的形态多样性分析；（2）角蟾科的系统发育关系与蝌蚪的演化，以及口漏斗的起源；（3）角蟾科蝌蚪表型分化与栖息环境和觅食行为的适应演化。针对上述问题，本文对角蟾科9 属30 种蝌蚪的形态特征，包括外部宏观形态和口器外部结构特征、口器内部显微结构、唇齿和角质颌的亚显微结构作了深入细致、多层次的比较研究；通过12s rRNA 和cytochrome b 基因构建最大简约树，采用贝叶斯系统发育进行分析，蝌蚪型的演化采用祖先性状的重建方法分析；得到如下结论：1）初步将角蟾科蝌蚪分为4 种类型；并且建立了2 种新的角蟾科蝌蚪类型。A 型：拟髭蟾型蝌蚪，该型蝌蚪包括拟髭蟾属、髭蟾属、齿蟾属和齿突蟾属的物种；B 型：新类型，掌突蟾型蝌蚪，该型蝌蚪在本文中包括掌突蟾属、小臂蟾属的物种；C 型：新类型，短腿蟾型蝌蚪，一种特化类型，该型蝌蚪在本文中仅包括短腿蟾属的物种；D 型：角蟾型蝌蚪，该型蝌蚪在本文中包括无耳蟾属、小口拟角蟾属和异角蟾属的物种。2）对角蟾科的分类进行了修订：（1）支持角蟾科两个亚科的分类系统；（2）角蟾亚科包括拟角蟾属、异角蟾属、无耳蟾属和短腿蟾属；该亚科形态差异小，系统学关系比较复杂，暂不作族级分类的再划分；（3）拟髭蟾亚科分为2 个族：拟髭蟾族，该族物种具有类型A 的蝌蚪，包括4 个属：拟髭蟾属、髭蟾属、齿蟾属、齿突蟾属；掌突蟾族，该族物种具有类型B 的蝌蚪，包括2 个属：掌突蟾属和小臂蟾属。3）结合分子系统进化关系探讨了4 种蝌蚪类型的演化。（1）角蟾科蝌蚪的最近共同祖先来自于一类具有拟髭蟾型蝌蚪性状的蝌蚪；（2）掌突蟾型蝌蚪和角蟾亚科的蝌蚪是由具有拟髭蟾型蝌蚪性状的祖先蝌蚪分别演化而来；（3）短腿蟾型蝌蚪是角蟾型蝌蚪的一种特化类型；（4）外群蝌蚪具有与拟髭蟾型蝌蚪相似的性状，进一步印证了类拟髭蟾型蝌蚪是角蟾科蝌蚪的最近共同祖先的假说；（5）具有口漏斗的蝌蚪类型是由不具口漏斗的蝌蚪类型演化而来，在角蟾科中口漏斗是一种衍生性状。4）分析了角蟾科四种蝌蚪类型与栖息环境的适应演化。（1）角蟾科蝌蚪的口部和体形的变化反映了该科蝌蚪由缓流向类似静水生境的回水凼的渐变式适应，角蟾科蝌蚪的形态显示了多方面的适应变化；（2）随着蝌蚪类型由A 向D的演化，当水速较大时，拟髭蟾型的蝌蚪营流水攀吸型生活方式；当水速递减时，掌突蟾型蝌蚪营流水附着型生活方式；当水速进一步递减时，具有较小口漏斗的短腿蟾型蝌蚪和具有大漏斗的角蟾型蝌蚪营流水浮泳型生活。角蟾科蝌蚪对于水流递减的适应演化说明蝌蚪的生态学适应是具有进化意义的；（3）蝌蚪口器内部结构的分化揭示了蝌蚪和食性的适应关系，蝌蚪以口部的唇齿与角质颌刮取或吞吸水中的物质，然后，通过口乳突有选择地过滤进入口腔中食物。拟髭蟾亚科蝌蚪的唇齿多而窄，唇齿间距宽，颌鞘粗而稀，反映了其植食性为主的特点；它们的舌前乳突一般为指状，在口腔入口处所占面积小，其机械过滤的作用很多被唇齿和角质颌分担了；而角蟾亚科的蝌蚪，其角质颌弱，其舌前乳突一般为匙状，几乎填满了口腔入口处，因此舌前乳突起了主要的机械过滤作用。The family Megophryidae is the largest and most diverse families inArchaeobatrachia, and most of its species occur in India, Pakistan, and eastward intoChina, Southeast Asia, Borneo and the Philippines to the Sunda Islands. Currently thefamily includes 142 species have been grouped into two subfamilies, Megophryinaeand Leptobrachiinae. The mountains of central and southern China are rich in speciesof Megophryidae, 75 species belong to 9 genera and two subfamilies.The family was supposed to be ideal materials of studies in many fields of biology,such as taxonomy, evolution, systematics, ecology, and biogeography. Recently, therehave a great development in taxonomy and systematics of megophryids throughstudied by morphology, paleontology, cytology, ecology, and cladistics. However,larvae of megophryids were generally unknown, although the tadpoles might be veryimportant for above studies.In this paper, we examined the evolutionary scenario of the tadpoles’ morphologyin the context of a phylogenetic framework. Our objectives are (1) to evaluate thedivergence of larval body shape and oral discs in the family Megophryidae, (2) toexplore the evolutionary trends of the larvae in megophryidae, and test if thefunnel-shaped oral disc is apomorphic, and (3) to explore the relationship of the larvalstructure, diet and microhabitat.We examined larval morphology of 30 megophryid species, the larval body shape,oral discs, the buccopharyngeal cavity, and jaw sheaths and denticles of the Chinesemegophryid frogs were re-examined. We constructed a phylogeny of the species on thebasis of published mitochondrial cytochrome b and 16S rRNA gene segments usingpartitioned Bayesian analyses. Furthermore, hypothetical changes of larval morphologywere inferred using parsimony principle on the phylogeny. The results showed that:1) Four tadpole types in Megophryidae. The larval morphological charactersseries in Chinese megophryids fall into four general categories according to the bodyshape and oral discs: (A) Leptobrachiini type, species from genera Leptobrachium,Oreolalax, Scutiger and, Vibrissaphora share this type of tadpoles. (B) Leptolalax type,species of genus Leptolalax have this type of tadpoles. (C) Brachytarsophrys type,species of the genus Brachytarsophrys have this type of tadpoles. (D) Megophryinitype, species of the genera Atympanophrys, Ophryophryne, and Xenophrys share this type of tadpoles. Of which B and C are two novel types.2）Taxonomic implications. The present study leads us to reconsider the generalclassification of tribes attributed to members of Megophryidae. More specifically,concerning the phylogenetic relationships and the two novel tadpole types describedherein, we propose a provisional taxonomy for the family but suggest that further taxasampling of other megophryids be performed to confirm this taxonomic change. TheMegophryidae is composed of two subfamilies (Leptobrachiinae and Megophryinae).The Leptobrachiinae was recogonized the two tribes: (1) tribe Leptobrachiini sensuDubois, corresponding to the tadpole of type A, including four genera, i.e.,Leptobrachium, Oreolalax, Scutiger and, Vibrissaphora; (2) tribe Leptolalaxini,corresponding to the tadpole of novel type B, including two genera, i.e., Leptolalaxand Leptobrachella. However, the relationships among the genera of Megophryinaewere largely unresolved, they recognized no monophyletic groups above the generalevel. A more thorough sampling will likely foster a better taxonomic solution.3) The larval evolutionary scenario in Megophryidae.Type A is characteristicof normal-mouthed with multiple tooth rows, representing the tadpole type of theMRCA of Chinese megophryids. Type B is characteristic of normal-mouthed withreduced tooth rows, prolonging labium, and integumetary glands. Type C ischaracteristic of no labial teeth and smaller umbeliform oral disc. Type D ischaracteristic of no labial teeth, enlarged umbeliform oral disc, representing the tadpoleof the MRCA of subfamily Megophryinae. A previous hypothesis, referring tofunnel-shaped oral discs as an apomorphy, is supported.4) The larval adaptation to habitats in Megophryidae. Tadpoles generallyadhere to substrates using their mouths, and the microhabitat that the tadpoles occupyreflects the degree of adhesion and oral complexity. The morphological changes inmegophryid tadpoles virtually allow a progressive adaptation to a changing habitatfrom faster water to slower water. Within the tadpoles of Type A to type D, the TOTbecomes smaller and smaller, and the oral disc orientates from anteroventral toumbelliform upturned, and eye position orientates from dorsal to lateral, and the trunkis more and more depressed and tail becomes relatively longer and slender. Within therunning water, the normal-mouthed with multiple tooth rows of Leptobrachiini tadpoles are correlated with lotic-suctorial, benthic feeders with anteroventral oraldisc and the largest body. With the water’s velocity decreasing, the lotic-adherentfeeders of Leptolalax tadpoles have tube-shaped labium with reduced tooth rows andintegumetary glands. And then, the smaller umbeliform in Brachytarsophrys tadpolesand the enlarged umbeliform oral disc in the Megophryini tadpoles are inhabitmicrohabitats of non-flowing backwaters of rivers, indicative of adaptive traits oflotic-neustonic surface feeders. The scheme of megophryid tadpoles andmicrohabitats provided the first clear evidence which congruent with the hypothesis ofAltig and Johnston (1989). The ecological divergence plays a general role in thedivergence and evolution of megophrid larvae. There is a definite correlation amongthe buccopharyngeal cavity, diet and feeding mechanisms, the tadpole graze orswallow the food particles, then through papillae which like a sieve and sort out foodparticles to the oesophagus. The tadpole of Leptobrachiinae possess multiple toothrows, wide intertooth distance as well as thick and sparse jaw sheath, these tadpolesinhabit bottom of the streams and graze on epiphyton or major detritus of organicmatter on the substrates, their prelingual papillae like single finger, the mechanicalpurpose of papillae served share in by tooth and jaw. The tadpoles of Megophryinaeoccur near the water surface of small streams and are the filter feeder, their dietincludes plankton and organic debris floating on the water surface, those tadpolepossess weak jaw, their prelingual papillae like spoon, the mechanical purpose ofpapillae served mostly for sieve.

黄河上游青海沙蜥种组的系统地理学研究

沙蜥属（Phrynocephalus）的卵胎生类群主要分布在我国青藏高原，包括南疆沙蜥（P. forsythii）、西藏沙蜥（P. theobaldi）、红尾沙蜥（P. erythrurus）、贵德沙蜥（P. putjatia）和青海沙蜥（P. vlangalii）。其卵胎生生殖方式适应了高寒生境，与青藏高原隆升有关。纵观前人的研究，上述几种卵胎生沙蜥的分类、系统发育关系以及生物地理都还存在疑问。本文研究了分布在若尔盖湿地的青海沙蜥红原亚种（P. v hongyuanensis）以及分布在黄河上游其它地区青海沙蜥种组的地理分布格局，并探讨了其形成机制。 青海沙蜥在黄河上游主要分布于若尔盖湿地以及青海湖周边地区。若尔盖湿地青海沙蜥红原亚种的生境由于沼泽的形成被切割成不连续的斑块，通过遗传分析可以推测这种特殊生境对它们遗传结构的影响。其次，贵德沙蜥、青海沙蜥的青海湖周边各居群以及若尔盖湿地居群之间的系统地理格局还未见报道。因此本文以居群为单位，将它们作为一个复合体，通过系统地理研究，可以了解其种群遗传结构，据此分析相关的地质历史事件对其分布的影响。主要结果如下： 1. 若尔盖湿地青海沙蜥红原亚种的种群遗传结构： 共研究了三个地理单元（红原（HY）、辖曼（XM）、玛曲（MQ））的7个采集点的72个个体。所有ND4-tRNALeu序列比对得到785 bp的片断，定义了9种单倍型。结果显示总的核苷酸多样性较低，单倍型多样性较高。分子变异分析（AMOVA）显示3个单元间差异显著（P<0.01），遗传变异主要存在于地理单元间，占62.61%。除MQ单元，XM各居群与HY居群混杂在一起，单倍型网络图没有显示出单倍型和地理位置的对应关系。XM单元单倍型的不配对分布（Mismatch distribution）为明显左移的单峰，且Fu’s Fs test为负值，表明XM单元可能经历了近期种群扩张，有足够的时间积累单倍型的多态性，还不足以大幅提高核苷酸多样性，这是其单倍型多样性较高和核苷酸多样性较低的原因。MQ单元遗传多样性低而与其他单元显著分化，推测这与3万年前黄河在若尔盖玛曲之间贯通有关。近期沼泽的形成对XMb居群的隔离时间短，使得其遗传多样性低但还不足以形成大的遗传差异。无论黄河的贯通还是沼泽的形成其隔离形成的时间都不长，其作用改变了单倍型出现的频率，也出现了一些特有单倍型，但共享单倍型还广泛存在，还不足以使得不同居群之间形成较大的遗传距离。 2. 黄河上游青海沙蜥种组的分布格局与地史过程的关系： 黄河上游青海沙蜥种组包括贵德沙蜥、青海沙蜥指名亚种的青海湖周边各居群、青海沙蜥红原亚种若尔盖湿地居群、以及青海湖以西的部分居群（序列由Genbank下载获得），总计22个居群189个样品。所有ND4-tRNALeu序列比对得到703个位点，定义了39种单倍型。以南疆沙蜥为外群构建的贝叶斯树以及MP法构建的无根树，都分为A、B两大组。其中A包括若尔盖湿地居群以及玛多居群（A1）、青海湖以西的居群和兴海居群（A2）、西藏沙蜥；B包括青海湖以南的居群和天祝居群（B1）、青海湖以东北的居群（B2）。单倍型网络图分别对应了系统发育树上的各支。按照系统发育结果分组进行分子变异分析，得到组间变异占88.63％，各组间差异显著（P=0.000）。种群遗传结构分析得到，A1和B2可能经历了近期的种群扩张，前者扩张时间约为0.105－0.189 Ma B.P.（million years before present），后者为0.057－0.102 Ma B.P.，可能与末次间冰期的气候变暖有关。A2和B1对应的两个地理单元都具有较强的种群遗传结构，较为稳定。 青海沙蜥种组A、B两大支之间遗传距离大，分化明显，分化大约发生在4.29－2.38 Ma B.P.，推测青藏运动的A幕运动后复杂的地形变化可能是它们产生分化的原因。B1和B2分化大约发生在1.73－0.96 Ma B.P.，这与湟水流域构造运动发生的时间相符。在早、中更新世时期，B1支内部各居群可能有交流，中更新世末共和盆地出现的抬升以及河流溯源改道等事件可能是引起这支内部多个单倍型丢失的原因。A1、A2支的分化可能与倒数第三次冰期降临之后气候变冷、阿尼玛卿山的大冰帽有关。 The viviparous group of genus Phrynocephalus is mainly distributed in the Qinghai –Tibetan Plateau, including P. forsythii、P. theobaldi、P. erythrurus、P. putjatia and P. vlangalii. These species are adapted well to the cold clime there, and the origin of this group was the result of a vicariance event associated with the uplifting of the Qinghai -Tibetan Plateau. Although many works have been done, there are still several questions about classification、phylogenetic relationships and the biogeography of this group. The phylogeographic pattern of the P. vlangalii complex on the upper reaches of the Yellow River and the P. v. hongyuanensis in Zoige Wetland were studied in this thesis. On the upper reaches of the Yellow River, P. vlangalii complex are distributed in Zoige Wetland and the southeast and northeast region of Kuku-noor Lake. Because of the forming of the wetland in Zoige, the habitats for sand lizards are divided into many discontinuous ones, and it is necessary to analyze genetic structure in these unique habitats. The phylogeographic patter among P. putjatia、populations of P. vlangalii in the southeast region of Kuku-noor Lake and populations of P. vlangalii in Zoige Wetland hasn’t been studied yet, and the complicated geological events of the Plateau may play an important role in the populations’ diversity and species forming there. So these populations were gathered as a complex, and phylogeographic analysis were used to clarify these doubts. According to the two topics above, this thesis has two parts of results as follows: 1. Three geographic units of P. vlangalii hongyuanensis in Zoige Wetland were defined, and they were Xiaman （XM）、Hongyuan （HY） and Maqu （MQ）. 785bp fragments of the mtDNA ND4-tRNAleu were determined from 72 samples and nine haplotypes were identified. As a whole, the nucleotide diversity was low，but the haplotype diversity was high. Analysis of molecular variance （AMOVA） showed that the three units were distinctly different（P<0.01），and 62.61% of the total genetic diversity was attributable to variation among units. There were 3 haplotypes shared among XM and HY，and no geographic clustering was observed except MQ from the TCS network. The results from the mismatch distribution analysis and Fu’s Fs test implied that there might be a recent population expansion in the XM unit, and this may be the reason why XM had a high haplotype diversity but a low nucleotide diversity. We estimate that the MQ and XMb have lower diversities because of some very recent geographic events, such as the formation of the Yellow river’s upriver and the Zoige Wetland. Although they are distinctly different, not enough time has passed for them to have diverged a great genetic distance. 2. 189 samples in 22 populations of P. vlangalii complex were collected, including P. putjatia、populations of P. vlangalii in the southeast and northeast region of Kuku-noor Lake、 populations of P. vlangalii in Zoige Wetland and the data from Genbank. 703bp ND4-tRNALeu sequences identified 39 haplotypes. P. forsythii was selected as outgroup, and both the Bayesian tree and the MP unrooted tree were divided into two groups（A、B）. A included populations in Zoige Wetland and Xinghai（A1）、populations in the west of Kuku-noor Lake（A2）、P. theobaldi, and B included populations in the southeast of Kuku-noor Lake and Tianzhu（B1）、populations in the northeast of Kuku-noor Lake（B2）. The haplotype network agreed with these groups. AMOVA showed that these five groups were distinctly different（P<0.01）, and 88.63% of the total genetic diversity was attributable to variation among groups. There might be recent population expansion in A1 and A2, which corresponded to the dry climate of the last interglacial period. The expansion times were 0.189-0.105 Ma B.P. and 0.102-0.057 Ma B.P., respectively. A2 and B1 had strong genetic structure. The large genetic distance between A and B showed that they had been separated from each other for a long time（about 4.29-2.38 Ma B.P.）, and it corresponded to the A phase of Qingzang Movement. The diversity between B1 and B2 at 1.73-0.96 Ma B.P. may be caused by the geological event in Huangshui valley. In early Pleistocene, populations in B1 may have gene flow because of geographic linkage, and later the uplift of the Plateau and the change of river route there made a few haplotypes lost. A1 and A2 were divided into two parts by A’nyemaqen Mountains at 0.66-0.37 Ma B.P., which maybe corresponded to glaciations at about 0.7 Ma B.P.

Precipitation-use efficiency along a 4500-km grassland transect

National Natural Science Foundation of China [30590381, 40971027]; State Key Technologies RD Program [2006BAC08]; Chinese Academy of Sciences ; National Key Research and Development Program [2010CB833501]

Particle-attached and free-living bacterial communities in a contrasting marine environment: Victoria Harbor, Hong Kong

Diversity of particle-attached and free-living marine bacteria in Victoria Harbor, Hong Kong, and its adjacent coastal and estuarial environments was investigated using DNA fingerprinting and clone library analysis. Denaturing gradient gel electrophoresis (DGGE) analysis of 16S rRNA genes showed that bacterial communities in three stations of Victoria Harbor were similar, but differed from those in adjacent coastal and estuarine stations. Particle-attached and free-living bacterial community composition differed in the Victoria Harbor area. DNA sequencing of 28 bands from DGGE gel showed Alphaproteobacteria was the most abundant group, followed by the Bacteroidetes, and other Proteobacteria. Bacterial species richness (number of DGGE bands) differed among stations and populations (particle-attached and free-living; bottom and surface). BIOENV analysis indicated that the concentrations of suspended solids were the major contributing parameter for the spatial variation of total bacterial community structure. Samples from representative stations were selected for clone library (548 clones) construction and their phylogenetic distributions were similar to those of sequences from DGGE. Approximately 80% of clones were affiliated to Proteobacteria, Bacteroidetes and Cyanobacteria. The possible influences of dynamic pollution and hydrological conditions in the Victoria Harbor area on the particle-attached and free-living bacterial community structures were discussed.