987 resultados para vegetation patterns


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The mechanism of energy balance in an open-channel flow with submerged vegetation was investigated. The energy borrowed from the local flow, energy spending caused by vegetation drag and flow resistance, and energy transition along the water depth were calculated on the basis of the computational results of velocity and Reynolds stress. Further analysis showed that the energy spending in a cross-section was a maximum around the top of the vegetation, and its value decreased progressively until reaching zero at the flume bed or water surface. The energy borrowed from the local flow in the vegetated region could not provide for spending; therefore, surplus borrowed energy in the non-vegetated region was transmitted to the vegetated region. In addition, the total energy transition in the cross-section was zero; therefore, the total energy borrowed from the flow balanced the energy loss in the whole cross-section. At the same time, we found that there were three effects of vegetation on the flow: turbulence restriction due to vegetation, turbulence source due to vegetation and energy transference due to vegetation, where the second effect was the strongest one. Crown Copyright (C) 2010 Published by Elsevier Ltd. All rights reserved.

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青藏高原东缘的亚高山针叶林是长江上游重要的生态屏障,经过近六十年的采伐后,取而代之的是大量人工种植的云杉纯林。目前,这些人工林已经表现出树种单一,结构层次简单等生态问题,其物种多样性及生态效益与同地带天然林相比差距较明显。如何丰富该地区物种多样性,完善人工林生态系统的生态功能是一个十分重要的课题。林下植物是人工林群落的重要组成部分,对维持群落的生物多样性及完善生态系统功能具有明显的作用。因此,研究该地区人工针叶林的林下植被对不同生境的适应性对于理解人工林生态系统物种多样性的形成和维持机制都具有重要的意义。 本文以青藏高原东部亚高山针叶林的主要森林类型----云杉人工林为研究对象,选择林下11种具有不同喜光特性的常见植物,分别设置人工林林冠下及成熟林窗为研究样地,通过对各种植物叶片形态与物质分配特征、叶片解剖学特征、叶片光合生理特性、植物自然分布特征等方面的比较分析,研究林下植物对不同光生境的适应策略及其适应能力,揭示不同物种对人工林生境的适应共性,为西南亚高山地区植被恢复及人工林的经营管理提供科学依据。具体研究结果如下: 在叶片形态和物质分配特征方面:在林窗光生境中,11种林下植物叶片比叶重(LMA)显著高于林下光生境的同种植物。同时,林窗下生长的植物叶片叶片厚度及栅栏细胞长度显著增加,这是影响叶片比叶重变化的直接原因。而多数植物叶重比在两种生境中无明显变化。说明在长期适应自然生境之后,植物可能更多地采取调节叶片组织细胞水平(即叶片功能细胞形态)及叶片器官水平(即单个叶片形态)特征的策略来适应各类生境,而非整株水平上的叶片总比重的增减。 在叶片解剖结构特征方面:多数阔叶物种栅栏组织厚度(PT)、栅栏组织厚度/海绵组织厚度(PT/ST)、栅栏细胞层数及近半数种的气孔密度(SD)在林窗生境中更大或更多,而叶片表皮细胞厚度(UET、LET)气孔长径(SL)及海绵组织厚度(ST)受两种生境影响不大。喜光特性相似的物种在生境适应策略上具有一定的趋同性。 在光合生理特征方面:在林窗生境中多数种植物的最大光合速率(Amax)、暗呼吸速率(Rd)及喜光植物光补偿点(LCP)显著或极显著高于林内生境同种植物。且在同一生境条件下,多数深度耐荫植物比喜光及轻度喜光植物有稍低的Rd和LCP。各植物在林内低光生境中具有更大的内禀光能转化效率,并在中午12:00~14:00之间光强最大的时刻发生了的最深程度的光抑制。多数种能通过调节自身某种光合素含量或色素之间的比例来适应不同的光生境,即通过增加叶绿素含量或降低Chla/b值来适应林内弱光生境,通过提高类胡萝卜素含量或单位叶绿素的类胡萝卜素含量降低强光带来的伤害。绝大多数物种并不采取调节叶片C、N含量的策略来适应不同的光生境。总之,植物部分光合参数(Amax、Rd、LCP)受生境的影响与其自身喜光特性有关,但另一些参数(Fv/Fm日变化、色素含量及比例、叶氮相对含量)受生境影响与其自身喜光特性无明显关联。 在表型可塑性方面:在叶片各表型参数中,器官水平及细胞水平的形态特征参数平均可塑性大于整株水平形态和物质分配特征参数可塑性;叶片光合组织的可塑性大于非光合组织可塑性;反映植物光合能力的参数可塑性大于叶片色素含量参数可塑性。植物叶片形态和物质分配、解剖学特征参数平均可塑性大小与其自身喜光特性基本吻合,即喜光种及轻度耐荫种各参数可塑性最高,深度耐荫种可塑性最小,而这种规律并未在光合生理参数的可塑性大小上体现出来。但是叶片形态和物质分配参数、光合生理参数的平均可塑性水平却大于叶片解剖学参数。 在植物自然分布特征方面:喜光物种云杉幼苗及歪头菜在林内生境中分布密度明显降低,深度耐荫种疏花槭却恰恰相反,更多数物种(7种植物)在两种生境中密度变化趋势不明显。从分布格局来看,7种植物在两种生境中均为聚集分布,但聚集强度为林窗>林内;少数物种桦叶荚迷、直穗小檗、冰川茶藨、黄背勾儿茶在林窗中为聚集型,在林内生境中的分布型发生改变而成为随机型,说明光生境的差异能影响到植物种群的分布特征。但这种影响程度与植物自身的喜光特性无关,同时与各物种叶片表型平均可塑性的大小也无明显关联。 The subalpine coniferous forest area in eastern Qinghai-Tibet Plateau is important ecology-barrier of upriver Yangtze. In past sixty years, those forests had been cut down and replaced with a lot of spruce plantations. At now, there are many ecology problems presenting to us such as singleness species, simple configuration, lower species diversity and ecological benefit than natural forests at the same belt. How to restore the species diversity and enhance the eco-function of the plantations is a very important issue. The understory plants are important part of plantation community, which improved the bio-diversity and eco-function distinctly of forests. So, it is very significance to study the adaptation of understory plants to different environment in plantation, and this study would helping us to understand how plantations to develop and remain their biodiversity. This study was conducted in a 60a spruce plantation in Miyaluo located in western Sichuan, China, and spruce plantation is major types of subalpine coniferous forest in eastern Qinghai-Tibet Plateau. In this paper, the leaf morphological and biomass-distributed characteristics, the anatomical characteristics, the photosynthetic characteristics and the distribution patterns characteristics of eleven different light-requirement understory species grown in two different environments (forest gaps and underneath close canopy) were studied and compared. The purpose of this study was to analyze the adaptation of this forest understory plants, to show up the commonness of these different light-requirement understory species in light acclimation, and to provide some scientific reference to manage and restore the vegetation of subalpine plantation of southwest China. The results were as follows: The leaf morphological and biomass-distributed characteristics: These eleven species in forest gaps had significantly higher dry weight per leaf area (LMA) than those under close canopy. The palisade parenchyma cells of the broad-leaved species in gaps were significantly longer than those grown under the canopy, which been a directed factor for the change of leaf mass per unit area (LMA) in different environment. But the leaf weight ratio (LWR) of most plants species were not evidently changed by the contrasted environments in our study. It was shown the morphological characteristics changing been adopted as a strategy of light acclimation for plants wasn’t on whole plant level (leaf weight ratio) but cellular level (the function cells morphological characteristics) and organic level (the leaf morphological and biomass-distributed characteristics) mostly. The leaf anatomical characteristics: Most broad-leaved plants in gaps increased palisade parenchyma thickness (PT), the palisade parenchyma cell layers and the ratio of palisade to spongy parenchyma (PT/ST). So did as almost about half species in this study in stomatal density (SD). No significant differences in thickness of leaf epidermal cells (UET, LET), stomatal length (SL) and spongy parenchyma (ST) between two environments of most species were observed. The results suggested that species with light-requirement approximately had convergent evolution on adaptation to light condition. The leaf photosynthetic characteristics: The dark respiration rate (Rd) of most plants species, the light compensation point (LCP) of light-demanding plants species in gaps were significantly increased than under close canopy in this study. In a same habitat, most deep-shade-tolerant plants had lower Rd and LCP than those light-demanding plants and slight-shade-tolerant plants. Each species has bigger inherent electron transport rate under close canopy than in gaps, and the greatest photoinhibition happened during 12 to 14 in the daytime. Most species could adapt different light environment by the way of changing their photosynthetic pigments content or the ratio of pigments content. For example, some plants under close canopy increased chlorophyll (Chl) or reduced the values of the ratio Chla/b to adapted the low light condition, some plants in gaps increased carotenoid (Car) or reduced the weight ratio CarChl to avoid been hurt in high light. For most plants, changing the content of C and N in leaf wasn’t a strategy of light acclimation. In conclusion, the variation of some leaf photosynthetic parameters in different light environment such as Fv/Fm, pigments, C and N in leaf related with the light-requirmnet of species, but the others such as Amax, Rd, LCP did not. The leaf plasticity indexes: Among those leaf plasticity indexes, the leaf morphological and biomass-distributed parameters on cellular and organic level were greater than on whole plant level for same species, and the photosynthetic parenchyma parameters were greater than non-photosynthetic parenchyma parameters in same leaf, and photosynthetic capability parameters were greater than photosynthetic pigments content parameters for same species. The average plasticity indexes of leaf morphological and biomass-distributed and anatomical parameters were accordant with plants’ light-requirement approximately: those light-demanding plants and slight-shade-tolerant plants had bigger plasticity indexes than deep-shade-tolerant plants. But this regular wasn’t observed in physiological plasticity indexes for most plants, though the average leaf plasticity indexes of leaf morphological and biomass-distributed, photosynthetic characteristics parameters was greater than the anatomical characteristics parameters. The distribution patterns characteristics: Oppositely to the deep-shade-tolerant specie Acer laxiflorum Pax., the density of light-demanding species Picea asperata Mast. and Vicia unijuga A. Br. in gaps was bigger than under close canopy. Each of the other species has the approximately density in two different environment. The spatial patterns of seven species were aggregated distribution in two environments, but the trend of aggregation of population under close canopy was decrease from in gaps. A few species such as Viburnum betulifoium Batal., Berberis dasystachya Maxim., Ribes glaciale Wall. and Berchemia flavescens Brongn. were aggregated distribution in gaps while random distribution under close canopy. It was shown that the difference between two light environments could affect the distribution pattern of plant population, and the effect didn’t relate with the light-requirement or plasticity indexes of species.

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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.

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海拔梯度造成的环境异质性,如崎岖的地形、复杂的植被结构以及花期延迟等可能会极大地影响到物种的形态和遗传变异格局。理解物种形态和遗传变异的海拔格局对于物种多样性的管理和保护是非常重要的。尽管植物群体遗传学是一个飞速发展的研究领域,然而与海拔相关的形态变异、遗传变异及群体间遗传差异的研究却很少。到目前为止,还不清楚遗传变异与海拔之间是否必然的相关性。 川滇高山栎是一种重要的生态和经济型树种,广泛分布于中国西南的四川、西藏、贵州和云南省的高海拔地区,在保持水土、调节气候方面起着十分重要的作用。尽管主要受阳光限制而仅分布于阳坡,但其海拔梯度范围较大,表明川滇高山栎对不同的环境具有很强的适应性。本文通过叶型及生理响应、微卫星分子标记和扩增性片段长度多态性方法,试图探索川滇高山栎叶沿海拔梯度的形态和生理响应及其沿海拔梯度的遗传变异格局,为川滇高山栎的保护和利用提供进一步的遗传学理论依据和技术指导。 对叶形、含氮量及碳同位素的试验结果表明,平均比叶面积、气孔密度、气孔长度和气孔指数等气孔参数随海拔的升高呈非线性变化。在海拔大于2800 m时,川滇高山栎的比叶面积、气孔长度和气孔指数都随海拔升高而降低,但是在海拔小于2800 m时,这些指标都随海拔的升高而增大。相对而言,单位叶面积的含氮量和碳同位素则表现出相反的变化模式。另外,比叶面积是决定碳同位素沿海拔梯度变化的最重要参数。本研究结果表明,海拔2800 m附近是川滇高山栎生长和发育的最适地带,在这里生长的植物叶片厚度更薄、气孔更大、叶碳同位素值更小。 利用六对微卫星引物对五个不同海拔川滇高山栎群体遗传多样性进行研究,结果表明,群体内表现出较高的遗传多样性,平均每位点等位基因数11.33个,平均期望杂合度达0.820。群体间差异较小,分化仅为6.6%。聚类分析也并没有显示出明显的海拔格局。然而低频率等位基因却与海拔呈显著性正相关(R2=0.97, P < 0.01),表明在高海拔处,川滇高山栎以更多的稀有基因来适应恶劣的环境条件。本试验结果表明由海拔梯度形成的选择性压力对川滇高山栎群体的遗传变异影响并不明显。 为了进一步探讨川滇高山栎群体遗传变异与海拔之间的相互关系,我们还对其进行了扩增性片段长度多态性分析。结果表明:(1)随海拔的升高(从群体WL2到群体WL5),群体内遗传变异降低,而群体间遗传差异增加;(2)低海拔群体WL1表现出最低的遗传变异性(HE = 0.181),同时与其余四个群体间呈现出最大的遗传差异性(平均FST = 0.0596);(3)在除去低海拔群体WL1后,Mantel检测表明群体间遗传距离与海拔距离之间表现出正相关性。另外,研究结果还表明,遗传变异受生境条件(过度的湿热环境)及人为干扰(火烧、砍伐和放牧)的影响,这一点至少在低海拔群体WL1上发生了作用。 通过叶形态、生理及DNA分子水平的研究,结果表明叶形态特征和碳同位素与海拔紧密相关,与海拔之间呈非线性变化,海拔2,800 m附近是川滇高山栎生长和发育的最适地带。海拔梯度在一定程度上会影响到川滇高山栎群体的遗传变异结构,但在这样一个狭窄的地理分布区域里,这种影响并不足以导致群体间较大的遗传分化。同时生境条件及人为干扰也是影响遗传变异的限制性因子,不容忽视。 Altitudinal gradients impose heterogeneous environmental conditions, such as rugged topography, a complex pattern of vegetation and flowering delay, and they likely furthermore markedly affect the morphological and genetic variation pattern of a species. Understanding altitudinal pattern of morphological and genetic variation at a species is important for the management and conservation of species diversity. Although plant population genetics is a fast growing field of research, there are only few recent investigations, which analyzed the genetic differentiation and changes of intra-population variation along altitudinal gradients. At present, it is still unclear whether there are some common patterns of morphological and genetic variation with altitude. Quercus aquifolioides Rehder & E.H. Wilson, which is an important ecological and economical endemic woody plant species, is widely distributed in the Yunnan and Sichuan provinces, Southwest China. Its large range of habitat across different altitudes implies strong adaptation to different environments, although it is mainly restricted to sunny, south facing slopes. It plays a very important role in preventing soil erosion, soil water loss and regulating climate, as well as in retaining ecological stability. In this paper, we tried to understand the altitudinal pattern of morphological and genetic variation along altitudinal gradients through the experiments of leaf morphological and physiological responses, microsatellite analysis and AFLP markers. In leaf morphological and physiological responses experiment, we measured leaf morphology, nitrogen content and carbon isotope composition (as an indicator of water use efficiency) of Q. aquifolioides along an altitudinal gradient. We found that these leaf morphological and physiological responses to altitudinal gradients were non-linear with increasing altitude. Specific leaf area, stomatal length and index increased with increasing altitude below 2,800 m, but decreased with increasing altitude above 2,800 m. In contrast, leaf nitrogen content per unit area and carbon isotope composition showed opposite change patterns. Specific leaf area seemed to be the most important parameter that determined the carbon isotope composition along the altitudinal gradient. Our results suggest that near 2,800 m in altitude could be the optimum zone for growth and development of Q. aquifolioides, and highlight the importance of the influence of altitude in research on plant physiological ecology. Genetic variation and differentiation were investigated among five natural populations of Q. aquifolioides occurring along an altitudinal gradient that varied from 2,000 to 3,600 m above sea level in the Wolong Natural Reserve of China, by analyzing variation at six microsatellite loci. The results showed that the populations were characterized by relatively high intra-population variation with the average number of alleles equaling 11.33 per locus and the average expected heterozygosity (HE) being 0.779. The amount of genetic variation varied only little among populations, which suggests that the influence of altitude factors on microsatellite variation is limited. However, there is a significantly positive correlation between altitude and the number of low-frequency alleles (R2=0.97, P < 0.01), which indicates that Q. aquifolioides from high altitudes has more unique variation, possibly enabling adaptation to severe conditions. F statistics showed the presence of a slight deficiency of heterozygosity (FIS=0.136) and a low level of differentiation among populations (FST=0.066). The result of the cluster analysis demonstrates that the grouping of populations does not correspond to the altitude of the populations. Based on the available data, it is likely that the selective forces related to altitude are not strong enough to significantly differentiate the populations of Q. aquifolioides in terms of microsatellite variation. To further elucidate genetic variation pattern of Q. aquifolioides populations under sub-alpine environments, genetic variation and differentiation were investigated along altitudinal gradients using AFLP markers. The altitudinal populations with an average altitude interval of 400 m, i.e. WL1, WL2, WL3, WL4 and WL5, correspond to the altitudes 2,000, 2,400, 2,800, 3,200 and 3,600 m, respectively. Our results were as follows: (i) decreasing genetic variation (ranging from 0.253 to 0.210) and increasing genetic differentiation with altitude were obtained from the WL2 to the WL5 population; (ii) the WL1 population showed the lowest genetic variation (HE = 0.181) and the highest genetic differentiation (average FST = 0.0596) with the other four populations; (iii) the positive correlation was obtained using Mantel tests between genetic and altitude distances except for the WL1 population. Our results suggest that altitudinal gradients may have influenced the genetic variation pattern of Q. aquifolioides populations to some extent. In addition, habitat environments (unfavorable wet and hot conditions) and human disturbances (burning, grazing and felling) were possible influencing factors, especially to the low-altitude WL1 population. The present study shows that there were close correlations between morphological features and carbon isotope composition in our data. This indicates that a coordinated plant response modified these parameters simultaneously across different altitudes. Around 2,800 m altitude there seems to be an optimum zone for growth and development of Q. aquifolioides, as indicated by thinner leaves, larger stomata and more negative d13C values. All available evidence indicates altitudinal gradients may have influenced the genetic variation pattern of Q. aquifolioides to some extent. Decreasing genetic variation and increasing genetic differentiation with altitude was obtained except for the WL1 population. And the environment of habitats and human disturbances were also contributing factors, which impact genetic variation pattern, especially to the low-altitude WL1 population.

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横断山地区是一个十分自然的植物区系地区,在中国植物区系分区中是作为泛北极植物区中国-喜马拉雅亚区中的一个地区,其种子植物区系具有丰富的科、属、种,地理成分复杂,特有现象和替代现象明显。该地区作为植物区系和生物多样性的研究热点地区,长期以来极受中外植物学家关注。横断山脉东缘是中国-喜马拉雅和中国-日本植物区系的交汇过渡区域,北部的岷江流域以及南部的金沙江流域,孕育了该区丰富的物种资源和植被资源。而岷江干热河谷和金沙江干热河谷的相似性和相关性,更为该区的植物区系和生物多样性南北的对比研究提供了有利的条件。 本研究选择的九顶山西坡和龙肘山分别位于横断山区北部和南部,九顶山属岷江流域而龙肘山属金沙江流域。本研究结合植物区系研究和生物多样性研究,对该区的植物资源进行调查。通过样带调查和样线踏查结合,大量详实的野外样方调查和标本采集,进行传统的区系研究和生物多样性研究。研究该区物种多样性的海拔梯度格局及其潜在的影响影子,并利用新的区系评估质量方法对九顶山西坡的植物区系质量进行定量的研究,以期能更为深刻的理解该区的植物资源,为该区的资源保护和利用提供合理可行的建议。主要研究结论如下: 1)九顶山西坡植物区系的性质和特点 经鉴定和统计,九顶山西坡共有1707 种维管植物,分属617 属和140 科,其中种子植物1616 种,分属572 属117 科。就科的分布区成分构成而言,该区系的热带成分与温带成分相当,热带成分略占优势,表明九顶山西坡的植物区系与热带植物区系有较强的联系。但是,在九顶山西坡属的分布区类型所占的比例上,温带成分远远超过了热带成分,本区的种子植物分布表现出明显的温带性质。且温带分布类型的许多物种组成了九顶山西坡植被的建群种和优势种,是本区系最重要的成分,充分体现了本区系的温带性质。 2)九顶山西坡不同植被带的生物多样性海拔梯度格局 基于对土门-断头崖、茶山-九顶山、雁门沟-光光山三条垂直植被样带的调查,我们发现九顶山西坡的生物多样性沿海拔梯度的变化呈现出一定的规律性,不同样带之间有一定差异。就三条样带的物种组成相似性来看,虽然土门-断头崖样带属于涪江水系,而茶山-九顶山样带和雁门沟-光光山样带属于岷江水系,但不同水系对该区物种组成的影响并不明显。三条样带中,草本层物种丰富度均远远大于灌木层和乔木层,而以乔木层物种丰富度最低;α-多样性指数随着海拔梯度的变化在土门-断头崖样带中呈现单一下降趋势,在茶山-九顶山样带表现为双峰模型,而在雁门沟-光光山样带则表现为不显著波动变化;均匀度指数在土门-断头崖样带呈现出单一下降的趋势,在雁门沟-光光山样带表现为凹形曲线,而在茶山-九顶山样带却无明显的变化规律。β-多样性指数在土门-断头崖样带和茶山-九顶山样带呈现出明显的波动状态,植被类型替代现象明显;而在雁门沟-光光山样带却并未有十分显著的转折点,因其水平植被带受到干扰,同海拔替代现象不显著。 3)九顶山西坡维管植物丰富度的海拔梯度格局 我们考察了九顶山西坡和两条垂直样带(土门-断头崖和雁门沟-光光山样带)的不同分类等级(包括科、属、种)和不同生活型物种(乔木、灌木、禾草、蕨类和其它草本)的丰富度沿着海拔梯度的分布。结果发现,物种的丰富度海拔梯度格局具有不同的模式,单一下降和中间膨胀格局依然是其主流。不同生活型的物种具有不同的丰富度格局,但是对于环境需求相似的类型具有较相似的丰富度格局。不同的丰富度格局可能由多因素导致,包括:气候,海拔跨度,面积,人为干扰等等。 4)九顶山西坡区系质量评估 我们尝试使用传统的区系质量评估方法对九顶山西坡的区系质量进行评估,并尝试使用一种新的区系质量评估体系对该区的区系进行评价。在九顶山西坡随着海拔梯度的上升,平均保守性系数在各条植被带中均呈现出逐渐上升的趋势。区系质量指数随着海拔的升高都表现为双峰模型,在植被交错区区系质量指数相对较高,而在海拔的两极,区系质量指数都很低。大部分地区使用新方法计算所得的加权平均保守性系数和区系质量指数都比传统方法计算的平均保收性系数和区系质量指数要高,说明在九顶山西坡的三条样带中,大部分地区都是那些保守性系数较高的物种占据优势,同时也表明九顶山西坡具有很高质量的区系和自然植被。 5)龙肘山种子植物区系的性质和特点 龙肘山种子植物区系的物种较为丰富,共有154 科,544 属,1156 种。科的优势十分明显,单种属和寡种属数量众多,说明本区系植物成分较为复杂、起源古老、物种多样性指数较高。地理成分复杂,分布类型多样,其中热带成分在总数量上高于温带成分,但是许多温带成分的属是该区植被的重要建群类群和优势类群,表现出明显的亚热带性质。 6)龙肘山生物多样性的现状和特点 在海拔梯度上,龙肘山地区无论是科、属、种的数量,还是不同等级分类单元之间的数量比,均呈现先升后降的趋势,并在中海拔地区达到峰值。物种多样性指数从总体上来说变化幅度不大,略有先升后降的趋势,在中海拔梯度物种多样性最高。乔、灌、草三层的多样性指数表现出乔木层<灌木层<草本层的特征;乔木层均匀度的变化很大,而灌木层和草本层均匀度的变化较小;灌木层均匀度的波动又强于草本层。β-多样性指数呈现单峰模式,中海拔地区最高。就龙肘山东、西坡物种多样性相比较而言,两者虽然在数值上交替上升,但是却体现出了较为一致的趋势,但西坡因受到干热河谷气候的影响,其平均气温要高于东坡,导致了东坡植物群落和物种的分布比西坡要低。在区系成分构成上,低山区的相同海拔段,西坡的热带亚热带成分所占的比例要比东坡高,这是因为西坡的平均气温比东坡稍高,导致了热带、亚热带物种分布更多。而随着海拔的上升,东、西两坡的气候、土壤等条件趋于一致,其植物区系成分的构成格局也趋于一致。 The Hengduan Mountain region is a very natural floristic region; it belongs toChina-Himalaya sub-region of Holarctic region in floristic subarea of China. The flora in this areais rich in family, genus and species; has a very complex composition of geographical elements;especially with high richness of endemic species and obvious substitution phenomenon. Thisregion as a hot-spot area of floristic and biodiversity, has fascinated biologists in the world for along time. The eastern range of Hengduan Mountain is the transition zone of China – Himalayaforest sub-region and China-Japan forest sub-region in floristic. The water systems are quitedifferent, Minjiang River in the north and Jishajiang River in the south grow quit different but alsoabundant plant species and vegetation resources. The similarity and correlativity of Minjiang River dry valleys and Jinshajiang River dry valleys have provided advantageous condition tocontrast flora and biodiversity between north and south. In the present study, the Jiuding Mountainlies in the north of Hengduan Mountain and belongs to Minjiang River, and the LongzhouMountain lies in the south of Hengduan Mountain and belongs to Jinshajiang River. In our study, we combined the methods of floristic research and biodiversity investigation toexplore the resources of plant species and vegetations; sampled with transects along the altitudinalgradients and also with transverse straps with similar elevation; collected the vascular plant specimen with sampling plots of ecology. We explored the plant species richness patterns alongaltitudinal gradients and discussed the underlying factors aroused these patterns; and used a novelmethod to assess the quality of Jiuding Mountain’s flora. All for a deeper comprehension of the plant recourses of this region; and provided feasible and reasonable suggestion for the protectionof resources. The results were as follows: 1 The characteristic of the flora of the west slope of Jiuding Mountain We had collected 1707 species of vascular plants belonging to 617 genera in 140 families inthe west slope of Jiuding Mountain,in which included 1616 seed plant species belonging to 572genera and 117 families. As for the composition of the areal types of the Families of seed plants,tropic components and temperate components are well-balanced, and percentage of tropicscomponents is higher than that of temperate ones for a litter bit. This shows the flora in the westslope of Jiuding Mountain has strong relationship with the tropic flora. But for the composition ofthe areal types of genera, temperate components have far exceeded the tropics ones, indicated thewhole flora with a conspicuous temperate character. Temperate components possess maximumproportion in the west slope of Jiuding Mountain, and many of them belong to constructivespecies and dominant species in the vegetation, are most important components in JiudingMountain’s Flora, also have embodied the temperate character of this area sufficiently. 2 Biodiversity patterns along altitudinal gradients in different vegetation transects in the westslope of Jiuding Mountain Based on the investigation of three vegetation transects (including Tumen-Duantouya transect,Chashan-Jiudingshan transect and Yanmengou-Guangguangshan Transect) in the west slope ofJiuding Mountain, we found the change of biodiversity along the altitude gradients displayedcertain regularity, but have differences among different transects. The three transects belong todifferent water systems; the Tumen-Duantouya transect belongs to Fujiang River, and the othertwo belong to Minjiang River. From the similarity of species compositions of different transects,we found different water system didn’t show obvious impact on the species composition. In all thethree transects, the species richness of herb layer was remarkably higher than shrub and tree layer,and the species richness of tree layer was the lowest one. With the increasing of the altitude, theline of α-diversity was monotonically decreasing curve in Tumen-Duantouya transect, andbimodal curve in Chashan-Jiudingshan transect, but in Yanmengou-Guangguangshan transectshowed a wave-like curve although not very obvious. Species evenness showed monotonicallydecreasing trends in Tumen-Duantouya transect, and very low at mid-altitude in Yanmengou-Guangguangshan transect, but in Chashan-Jiudingshan transect changed irregularly. Changes inβ-diversity corresponded with the transition of vegetation in the Tumen-Duantouya transect andChashan-Jiudingshan transect, and the curve of β-diversity along altitude had obvious turningpoint; but in Yanmengou-Guangguangshan transect had no obvious turning point, and thesubstitution phenomenon was not obvious, transverse vegetation straps distributed interlaced. 3 Richness patterns of vascular plant species along altitude in the west slope of Jiuding Mountain Direct gradient analysis and regression methods were used to describe the species richnesspatterns along the altitudinal for Mt. Jiuding, as well as separately for Tumen-Duantouya Transectand Yanmengou-Guangguangshan Transect. Altitudinal gradient of diversity of units at differenttaxonomic level (including Family, Genus and Species) and at different life form (including tree,shrub, pteridophyte, grass and other herb) were tested to find differences among the richnesspattern. We found altitudinal richness also shows different patterns, and both monotonicallydecreasing pattern and hump-shaped pattern can be founded in vascular species richness. Speciesin different life forms show different altitudinal patterns, but those species with similarrequirements to environmental conditions show similar richness patterns along altitudinalgradients. Different richness patterns can be aroused by different climate, different altitudinal span,area factor, anthropogenic factor and so on. 4 Floristic quality assessments in the west slope of Jiuding Mountain We used both the conventional method broadly adopted in the USA and the new one toassess the floristic quality in the west slope of Jiuding Mountain. The Mean Coefficient ofConservatism (MC) had the trend of increment along the altitudinal gradients. The FloristicQuality Index (FQI) was a bimodal curve with increasing of elevation; FQI got maximum valuesin the transition zones of different vegetations in the middle altitude, and had very low values atthe two end of elevation. In most areas of the west slope of the Jiuding Mountain, the resultscalculated using the new methods were higher than those using the conventional method. Thisindicated the dominant species of the communities had very high coefficients of conservatism inmost areas of Jiuding Mountain, and the communities are relatively kept pristine and the habitats very integrative. 5 The characteristic of the flora of Longzhou Mountain The flora of Longzhou Mountain has very abundant in species composition; there are about1156 species of seed plants belonging to 544 genera in 154 families. In which, twelve families with more than 20 species include totally 232 genera and 532 species, and form the majority of itsflora. The origin of its flora is old, monospecific genera and oligotypic genera amounts to 510 innumber, which constitute 93.75% of total number of genera. The geographical components arevarious in Longzhou Mountain, the majority of flora are temperate and pantropic ones. The tropiccomponents overtopped temperate components on genera quantity, but many temperatecomponents belong to constructive species and dominant species in the vegetation, and the wholeflora shows an obvious subtropical character. 6 Current situation and characteristic of biodiversity in Longzhou Mountain With the increasing of altitude, the number of species, genus, family and the ratios ofdifferent taxonomic levels all displayed a trend of descending after rising first, and peaked atmiddle height area. The change of α-diversity was not very acutely, with the trend of descendingafter rising first in some degree, the middle height area had highest α-diversity. As studying thetree layer, shrub layer and herb layer respectively, the Shannon-Wiener index was in followingorder: tree layer < shrub layer < herb layer; the change of evenness was more complicatedly thanthat of diversity, the tree layer changed acutely, but the shrub layer and herb layer fluctuatedsmoothly. Changes in β-diversity also showed the trend of descending after rising first. TheJaccard index and Cody index all peaked at the middle height forest area. As for the comparison ofplant diversity and evenness between the west and east slope, the numerical values ascendedalternatively, but the trend of changing was similar. The distribution of similar plant communitiesand species in east slope were lower than the west slope for the influence of Jinsha River DryValley. As for the composition of different floristic components, in lower altitude area of westslope, the tropic and sub-tropic plants had higher ratio than east slope’s and even could be equal tothe temperate plants. With the increasing of elevation, the floristic composition become morelikely between the east and west slope and temperate plants dominated the flora.

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从空间、时间、食物3个方面研究了若尔盖湿地3种两栖类的种间竞争,运用生态位理论探讨了3种两栖类利用环境资源的能力以及它们之间的共存模式,研究了3种两栖类年周期食性,并考察了畜牧业对3种两栖类食性及食物竞争格局的影响;此外,还通过实验室研究对2种两栖类幼体的种间竞争策略进行了考察。主要结果如下: 1、两栖类的空间资源利用状况:在3种两栖类成体生态位宽度的比较中,岷山蟾蜍(Bufo minshanicus)成体在牧场性质(0.41)、牛粪数量(0.42)、植被盖度(0.20)、地表温度(0.50)、地表湿度(0.51) 5个维度上的生态位宽度最窄;而倭蛙(Nanorana pleskei)成体在牛粪数量(0.81)、植被高度(0.63)、植被盖度(0.47)、小水体距离(0.68) 4个维度上的生态位宽度最宽。在3种两栖类亚成体生态位宽度的比较中,岷山蟾蜍亚成体在牧场性质(0.66)、牛粪数量(0.58)、植被高度(0.64)、小水体距离(0.51)、地表湿度(0.79) 5个维度的上生态位宽度最宽;倭蛙亚成体在牧场性质(0.39)、牛粪数量(0.30)、地表温度(0.18)、地表湿度(0.33) 4个维度上的生态位宽度最窄。高原林蛙(Rana kukunoris)在地表温度(成体:0.62;亚成体:0.56)、地表湿度(成体:0.84;亚成体:0.60)两个维度上具有较大的生态位宽度值,而在小水体距离维度上(成体:0.27;亚成体:0.14)的生态位宽度值则很小。比较3种无尾两栖类在不同生长阶段(成体、亚成体)的生态位宽度,发现高原林蛙和倭蛙的亚成体对栖息环境的要求更高。3种两栖类空间资源利用的相似程度很高,高原林蛙与倭蛙之间的生态重叠度(0.87)较之它与岷山蟾蜍(0.81)的生态位重叠度更大。 2、两栖类的日活动节律:高原林蛙成体、亚成体、岷山蟾蜍亚成体活动的最低气温为0℃、2℃、8℃;岷山蟾蜍和高原林蛙亚成体出现的数量与气温成极显著的正相关(r=0.797, p<0.001;r=0.794, p<0.001),高原林蛙成体出现的数量与气温有一定相关性(r=0.456, p<0.05);晴天时两栖类的活动性明显高于阴天(p<0.001);多云转晴天气,高原林蛙和岷山蟾蜍亚成体出现两次日活动高峰,分别为中午12:30左右和下午15:30~16:30之间;多云天气,高原林蛙和岷山蟾蜍亚成体出现两次日活动高峰,分别为9:30~10:30之间和15:30~16:30之间。 3、两栖类的食物资源利用状况:春、秋两季,高原林蛙最主要的食物是蜉金龟科(Aphodiidae)昆虫,相对重要性指数(IRI)最高(春季:35.28%,秋季:28.57%),其次为昆虫的幼虫,以及双翅目的毛蚊科(Bibionidae)、蝇科(Muscidae)、丽蝇科(Calliphoridae)昆虫,秋季,蝗虫是高原林蛙食物组成中的重要部分;岷山蟾蜍最主要的食物是蚂蚁(IRI,春季:85.54%,秋季:49.70%),其次为蜉金龟科、象甲科(Curculionidae)、步甲科(Carabidae)、粪金龟科(Geotrupidae) 等鞘翅目昆虫;倭蛙春季的最主要食物也是蜉金龟科昆虫(IRI,春季:13.41%),其次为蚂蚁、毛蚊科昆虫、昆虫的幼虫以及狼蛛科(Lycosidae)。3种两栖类中,倭蛙的食性生态位宽度相对较宽(0.43),而岷山蟾蜍(0.09)和高原林蛙(0.22)的生态位宽度较窄,与春季相比,两栖类在秋季的食谱更宽。以利用食物种类为标准,春季高原林蛙与倭蛙的生态位重叠度(0.40)比它与岷山蟾蜍的生态位重叠度(0.33)更大。 4、畜牧业对两栖类食性及食物竞争格局的影响:以藏牦牛粪为食物或寄居场所的昆虫,如蜉金龟科、粪金龟科、毛蚊科、蝇科、丽蝇科昆虫和某些昆虫幼虫,是3种两栖类食物谱中最主要的组成部分,蜉金龟科昆虫在高原林蛙食谱中的比例更高,高原林蛙可能从畜牧业发展中获得更多的好处,使之在食物竞争方面处于优势地位。与无放牧样地相比,在有放牧样地的中,两栖类食谱中的蜉金龟科昆虫数量更多(有放牧:31.94%;无放牧:21.32%)、出现频率更高(有放牧:76.38%;无放牧:44%)。然而在不同样地上(有放牧/无放牧),两栖类的食物组成无显著性差异(P=0.188),两栖类的数量(P=0.075)、肥满度(P=0.537)均没有显著差别。 5、两栖类幼体的竞争策略:实验室条件下,通过活动性水平,变态时的体重、增长率和完成变态所需时间考察自然条件下常同水塘分布的中华蟾蜍(Bufo gargarizans)和高原林蛙蝌蚪的竞争策略。结果表明:中华蟾蜍蝌蚪在不同食物资源条件下,所选择的生存策略可能不同,即食物资源充足时,增加活动性获取更多食物,食物资源有限时,降低活动性且提前完成变态;与中华蟾蜍蝌蚪相比,在食物资源有限时高原林蛙蝌蚪获取食物能力可能更强。 This paper presented the study of competition of three amphibians (Rana kukunoris, Nanorana pleskei, Bufo minshanicus) based on spatial, temporal and dietary scales in Zoige wetland. We measured coexistence patterns of three amphibians and analyzed their ability of exploiting resource. Effects of grazing on the diet composition and diet competition of amphibians were analyzed by their diet composition during spring and autumn. Furthermore, we examined the competitive ability of larval common frogs (Rana kukunoris)and common toads(Bufo gargarizans) in a laboratory experiment, and analyzed their competitive strategies respectively. The results were as follows: 1 .The status of using spatial resource Niche breadths of B. minshanicus adults on 5 dimensional axes including character of pasture(0.41), number of yaks dung(0.42), vegetation coverage(0.20), temperature (0.50)and humidity(0.51) of ground surface were narrower than adults of R. kukunoris and N. pleskei. Niche breadths of B. minshanicus subadults were broader than R.kukunoris subadults and N.pleskei subadults on 5 dimensional axes including character of pasture (0.66), number of yaks dung (0.58), vegetation height (0.64), distance to small waterbodies (0.51), humidity of ground surface (0.79). Niche breadths of N. pleskei subadults were the narrowest in three anurans subadults on 4 dimensional axes including character of pasture (0.39), number of yaks dung (0.30), temperature (0.18) and humidity (0.33) of ground surface, niche breadths of N. pleskei adults were the broadest in three anurans adults on 4 dimensional axes including number of yaks dung (0.81), vegetation height (0.63) and coverage(0.47), distance to small waterbodies(0.68).Comparatively, niche breadths of R. kukunoris were broader on the two microclimate factors including temperature(adults:0.62;subadults:0.56) and humidity (adults:0.84;subadults:0.60)of ground surface, but was narrow on distance to small waterbodies(adults:0.27;subadults:0.14). Strategies for using habitat resource of adults and subadults of the three species anuran were different. Generally, subadults of R. kukunoris and N. pleskei needs better habitat condition. It was quite similar that three anurans exploited spatial resource, Niche overlap between R. kukunoris and N. pleskei (0.87) was greater than that between R. kukunoris and B.minshanicus(0.81). 2.Daily activity rhythm R. kukunoris audlts were active when air temperatures were as low as 0℃, R. kukunoris subadults were active at 2℃, B.minshanicus subaudlts were active at 8℃. Positive correlation was found between activities of amphibians and air temperature, Subadults of R.kukunoris, (r=0.797, p<0.001), Subadults,of,B.minshanicus, (r=0.794, p<0.001), andbadults,of,R.kukunoris(r=0.456, p<0.05).Amphibians were more active during sunny days than cloudy days. In cloudy turning into sunny, R. kukunoris and B.minshanicus subadults had two active peak: at noon about 12:30 and 15:30~16:30 pm; in cloudy, R. kukunoris and B.minshanicus subadult had two active peak too : 9:30~10:30am,15:30~16:30pm. 3.Diet analysis Aphodiidae was the most commonly consumed food item by R. kukunoris based on index of relative importance (IRI) during spring (35.28%) and autumn (28.57%) in Zogie wetland. Besides Aphodiidae, larval insect, dipterans such as Bibionidae, Muscidae, Calliphoridae also were important food item for R. kukunoris, in autumn, locust was one of important food item for R. kukunoris. The most important food item for B.minshanicus during spring (IRI:85.54%) and autumn (IRI:49.70%) was ants, following, was coleopterans, such as Aphodiidae, dung beetle. Aphodiidae (IRI:13.41%) were the most important consumed food item by N. pleskei during spring too, following, was ants and Bibionidae. Dietary breadth of N. pleskei (0.43) were greater than R. kukunoris (0.22) and B. minshanicus (0.09). As a whole, Dietary breadth of amphibians during aurumn were greater than spring. Based on prey item, dietary overlap between R. kukunoris and N. pleskei (0.40) was greater than that between R. kukunoris and B.minshanicus (0.33) during spring. 4.Effects of grazing on the diet composition and diet competition of amphibians Amphibians are an important part of the pasture ecosystems as prey and predator. In Zogie wetland, major diet of amphibians was closely associated with dung of yaks, for example, Aphodiidae, Bibionidae, Muscidae, dung beetle. Dung of yaks was major diet and habitat of these insects. Proportion of Aphodiidae was higher in diet composition of R. kukunoris than N. pleskei and B.minshanicus, with development of pasturage, R. kukunoris may have a diet competitive advantage over N. pleskei and B.minshanicus. Number of Aphodiidae in diet composition of amphibians was higher in samples with grazing (31.94%) than in those without grazing (21.32%). Occurrence Frequency of Aphodiidae in diet composition of amphibians was higher in samples with grazing (76.38%) than in those without grazing (44%). However, There was not significantly different on diet composition (P=0.188), and number (P=0.075) and the relative fatness (P=0.537) of amphibians between grazing samples and without grazing. 5.Competitive strategies of amphibian larvae I examined the competitive ability of larval toads (Bufo gargarizans) and frogs (Rana kukunoris) which co-occur in the nature pond by activity level, the growth rate and mass at metamorphosis and larval period in a laboratory experiment. The results suggest: In laborary, B.gargarizans adapted himself to different food level by changing activity. At high food level, B. gargarizans increased activity to gain more diet. At low food level, B. gargarizans decreased activity and achieved early metamorphosis. When food resource was limit, R. kukunoris could gain more food than B. gargarizans.

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