Species richness, but not phylogenetic diversity, influences community biomass production and temporal stability in a re-examination of 16 grassland biodiversity studies

* Hundreds of experiments have now manipulated species richness (SR) of various groups of organisms and examined how this aspect of biological diversity influences ecosystem functioning. Ecologists have recently expanded this field to look at whether phylogenetic diversity (PD) among species, often quantified as the sum of branch lengths on a molecular phylogeny leading to all species in a community, also predicts ecological function. Some have hypothesized that phylogenetic divergence should be a superior predictor of ecological function than SR because evolutionary relatedness represents the degree of ecological and functional differentiation among species. But studies to date have provided mixed support for this hypothesis. * Here, we reanalyse data from 16 experiments that have manipulated plant SR in grassland ecosystems and examined the impact on above-ground biomass production over multiple time points. Using a new molecular phylogeny of the plant species used in these experiments, we quantified how the PD of plants impacts average community biomass production as well as the stability of community biomass production through time. * Using four complementary analyses, we show that, after statistically controlling for variation in SR, PD (the sum of branches in a molecular phylogenetic tree connecting all species in a community) is neither related to mean community biomass nor to the temporal stability of biomass. These results run counter to past claims. However, after controlling for SR, PD was positively related to variation in community biomass over time due to an increase in the variances of individual species, but this relationship was not strong enough to influence community stability. * In contrast to the non-significant relationships between PD, biomass and stability, our analyses show that SR per se tends to increase the mean biomass production of plant communities, after controlling for PD. The relationship between SR and temporal variation in community biomass was either positive, non-significant or negative depending on which analysis was used. However, the increases in community biomass with SR, independently of PD, always led to increased stability. These results suggest that PD is no better as a predictor of ecosystem functioning than SR. * Synthesis. Our study on grasslands offers a cautionary tale when trying to relate PD to ecosystem functioning suggesting that there may be ecologically important trait and functional variation among species that is not explained by phylogenetic relatedness. Our results fail to support the hypothesis that the conservation of evolutionarily distinct species would be more effective than the conservation of SR as a way to maintain productive and stable communities under changing environmental conditions.

Foliar fungal pathogens and grassland biodiversity

By attacking plants, herbivorous mammals, insects, and belowground pathogens are known to play an important role in maintaining biodiversity in grasslands. Foliar fungal pathogens are ubiquitous in grassland ecosystems, but little is known about their role as drivers of community composition and diversity. Here we excluded foliar fungal pathogens from perennial grassland by using fungicide to determine the effect of natural levels of disease on an otherwise undisturbed plant community. Importantly, we excluded foliar fungal pathogens along with rabbits, insects, and mollusks in a full factorial design, which allowed a comparison of pathogen effects along with those of better studied plant enemies. This revealed that fungal pathogens substantially reduced aboveground plant biomass and promoted plant diversity and that this especially benefited legumes. The scale of pathogen effects on productivity and biodiversity was similar to that of rabbits and insects, but different plant species responded to the exclusion of the three plant enemies. These results suggest that theories of plant coexistence and management of biodiversity in grasslands should consider foliar fungal pathogens as potentially important drivers of community composition.

The importance of grasslands for animal production and other functions: a review on management and methodological progress in the tropics

The global importance of grasslands is indicated by their extent; they comprise some 26% of total land area and 80% of agriculturally productive land. The majority of grasslands are located in tropical developing countries where they are particularly important to the livelihoods of some one billion poor peoples. Grasslands clearly provide the feed base for grazing livestock and thus numerous high-quality foods, but such livestock also provide products such as fertilizer, transport, traction, fibre and leather. In addition, grasslands provide important services and roles including as water catchments, biodiversity reserves, for cultural and recreational needs, and potentially a carbon sink to alleviate greenhouse gas emissions. Inevitably, such functions may conflict with management for production of livestock products. Much of the increasing global demand for meat and milk, particularly from developing countries, will have to be supplied from grassland ecosystems, and this will provide difficult challenges. Increased production of meat and milk generally requires increased intake of metabolizable energy, and thus increased voluntary intake and/or digestibility of diets selected by grazing animals. These will require more widespread and effective application of improved management. Strategies to improve productivity include fertilizer application, grazing management, greater use of crop by-products, legumes and supplements and manipulation of stocking rate and herbage allowance. However, it is often difficult to predict the efficiency and cost-effectiveness of such strategies, particularly in tropical developing country production systems. Evaluation and on-going adjustment of grazing systems require appropriate and reliable assessment criteria, but these are often lacking. A number of emerging technologies may contribute to timely low-cost acquisition of quantitative information to better understand the soil-pasture-animal interactions and animal management in grassland systems. Development of remote imaging of vegetation, global positioning technology, improved diet markers, near IR spectroscopy and modelling provide improved tools for knowledge-based decisions on the productivity constraints of grazing animals. Individual electronic identification of animals offers opportunities for precision management on an individual animal basis for improved productivity. Improved outcomes in the form of livestock products, services and/or other outcomes from grasslands should be possible, but clearly a diversity of solutions are needed for the vast range of environments and social circumstances of global grasslands.

Defoliation and patchy nutrient return drive grazing effects on plant and soil properties in a dairy cow pasture

Large herbivores can influence plant and soil properties in grassland ecosystems, but especially for belowground biota and processes, the mechanisms that explain these effects are not fully understood. Here, we examine the capability of three grazing mechanisms-plant defoliation, dung and urine return, and physical presence of animals (causing trampling and excreta return in patches)-to explain grazing effects in Phleum pratense-Festuca pratensis dairy cow pasture in Finland. Comparison of control plots and plots grazed by cows showed that grazing maintained original plant-community structure, decreased shoot mass and root N and P concentrations, increased shoot N and P concentrations, and had an inconsistent effect on root mass. Among soil fauna, grazing increased the abundance of fungivorous nematodes and Aporrectodea earthworms and decreased the abundance of detritivorous enchytraeids and Lumbricus earthworms. Grazing also increased soil density and pH but did not affect average soil inorganic-N concentration. To reveal the mechanisms behind these effects, we analyzed results from mowed plots and plots that were both mowed and treated with a dung and urine mixture. This comparison revealed that grazing effects on plant attributes were almost entirely explained by defoliation, with only one partly explained by excreta return. Among belowground attributes, however, the mechanisms were more mixed, with effects explained by defoliation, patchy excreta return, and cow trampling. Average soil inorganic-N concentration was not affected by grazing because it was simultaneously decreased by defoliation and increased by cow presence. Presence of cows created great spatial heterogeneity in soil N availability and abundance of fungivorous nematodes. A greenhouse trial revealed a grazing-induced soil feedback on plant growth, which was explained by patchiness in N availability rather than changes in soil biota. Our results show that grazing effects on plant attributes can be satisfactorily predicted using the effects of defoliation, whereas those on soil fauna and soil N availability need understanding of other mechanisms as well. The results indicate that defoliation-induced changes in plant ecophysiology and the great spatial variation in N availability created by grazers are the two key mechanisms through which large herbivores can control grassland ecosystems.

草地生态系统碳循环及其对全球变化的响应_锡林河流域羊草草原生态系统循环研究

本研究从三个方面研究了草地生态系统的碳循环及其对全球变化的响应： 1)系统整理了世界范围内各种类型草原群落碳循环主要过程和影响因素的有关资料，在此基础上，对草地生态系统巾的碳素贮量和主要流量进行了全球估计;在CENTURY模型碳循环业模型的基础上，对草地生态系统碳循环的分室模型进行了说明;阐述了土壤呼吸作用在草地碳循环中的作用，综述了草地土壤呼吸的测定方法、影响草地呼吸的主要环境因素，对世界范围内草地土壤呼吸量和土壤中的有机碳周转速度进行了初步估计;讨论了全球气候变化、 C02浓度增加和土地利用变化对草地生态系统的可能影响。 2)对内蒙古锡林河流域一个固定样地内的羊草草原群落的碳循环特点进行了初步研究;采用实测数据和CENTURY模型模拟相结合，对该群落中碳素的输入与输出量，碳素贮量和土壤中有机碳的周转口寸间进行了首次估计，得出了“未利用状态下羊草草原群落基本处于碳素平衡状态”的结论。 3)对世界范围内草地利用现状的资料分析表明，全世界70%的草地出现了不同程度的退化， 过度放牧是导致草地退化的主要原因，在退化草地总面积中，约有35%是由于过度放牧造成的。过度放牧是锡林河流域羊草草原退化的主要入为因素，40年来，过度放牧共导致了羊草草地O - 20cm土层中23%的土壤有机碳损失。结合CENTURY模型的预测结果，土地利用变化对羊草草原土壤有机碳贮量的影响将比气候变化和C02浓度增加的影响更加重大。

土地利用方式对内蒙古农牧交错区草地生态系统的影响

陆地生态系统是人类赖以生存与持续发展的基础。由于其巨大的碳储存能力陆地生态系统在全球碳循环过程中发挥着重要作用，尤其是在延缓因大气中CO2浓度升高所引起的气候变化方面作用更为突出。作为世界上最广布的植被类型之一，草地生态系统是陆地生态系统的重要组成部分，其面积占到全球陆地面积的五分之一。中国有着世界第二大的草地生态系统，面积为3.55×108 hm2，约占世界草地面积的6％～8％，其碳储量为世界总碳储量的9％～16％。以内蒙古高原为主体的北方草地，面积约占国土总面积的40％左右，是一种重要的可再生的农牧业资源;作为一种重要的绿色生态屏障，北方草地在调节气候，涵养水源，防风固沙，净化空气和美化环境等方面起着重要作用。由于人口的增长和经济发展的需求，许多土地利用方式呈现在北方农牧交错区。地处内蒙古高原南麓的多伦县属于典型的农牧交错区，良好的气候条件和复杂的人文环境决定了多种土地利用方式在这一地区的并存，为开展不 同土地利用方式对草地生态系统影响的研究提供了天然的研究场所。 本研究选择了多伦县境内的6种土地利用方式作为研究对象，即围封禁牧样地、刈割禁牧样地、自由放牧样地、人工（紫花苜蓿）草地、农田（青贮玉米）和弃耕地，主要研究在这几种不同的土地利用方式影响下，土壤碳氮储量、草地生态系统功能（生产力等）和群落结构的变化。通过对土壤碳氮储量变化的研究发现： (1) 不同土地利用方式因其地上净初级生产力的不同而影响了土壤的碳氮储量，但是高的地上初级生产力并不一定能提高土壤的碳氮储量，如青贮玉米地。这主要是因为土壤碳氮储量与根系和凋落的输入相关。 (2) 不同土地利用方式对植被的浅层（0-20 cm）根系影响较大，继而影响了土壤的碳氮储量。这是因为大约80％的根系碳氮和50％的土壤碳氮储量都集中分布在这一层次。而土壤潜在库容的进一步扩大则要依靠深层根系生物量的增加，以提高深层土壤的碳氮含量，如人工（苜蓿）草地。 (3) 在合理利用方式或者科学管理措施下，草地土壤有机碳储量处于8 kg C m-2到10 kg C m-2之间，比过度退化草地的有机碳储量提高了50％。 (4) 经过短期恢复，土壤的库容量会很快增加，表现出较强的储存功能。因此，通过合理的土地利用方式和管理措施，中国北方农牧交错区可以转变成一个巨大的碳汇。 通过对不同利用方式下草地生态系统的功能、群落结构和物种多样性恢复的研究，得到如下实验结果： (1) 与自由放牧相比，短期（5年）禁牧（围封和刈割）提高了草地的地上净初级生产力和物种数目。围封禁牧和刈割禁牧草地的地上净初级生产力分别比自由放牧草地高出107.3％和50％;而物种数目则分别比自由放牧草地增加了14.8％和25.9％。 (2) 围封和刈割禁牧，尤其是后者，主要增加了非禾本科草本植物在群落中所占的比例;而在自由放牧草地，灌木和半灌木种群在群落中占有绝对优势。 (3) 物种多样性和生产力间的关系因外部干扰程度的不同而变化。在轻度干扰（围封禁牧）和中度干扰（刈割禁牧）下，生产力和物种多样性间呈指数正相关关系;而在重度干扰（自由放牧）下，群落地上净初级生产力随着物种多样性的增加呈指数递减的趋势。 通过对不同土地利用（或干扰）方式下，草地生态系统小尺度上的空间属性进行分析后，得到以下研究结果： (1) 群落和物种的地上生物量以及土壤养分均表现出显著的小尺度空间结构特征，并且在不同利用方式影响下，这种空间属性表现出较大的差异。 (2) 禁牧（围封和刈割）草地的土壤有机碳和氮含量比较高。土壤有机碳的变程较小，变化范围为0.83～1.40 m，其在三种类型土地利用的变异系数都维持在18.0％左右;土壤全氮的空间自相关尺度为0.46～14.69m，变异系数为12.10％～30.86％。 (3) 群落地上生物量以禁牧（围封和刈割）草地较高。在围封禁牧和刈割禁牧草地内的变异系数分别为26.48％和19.94％，在自由放牧草地的变异系数为67.47％;其空间自相关尺度为0.74～3.9m。在不同的土地利用方式下，物种、功能群以及多样性指数都表现出显著的空间异质性，并且其空间属性变化各异。 (4) 土壤养分和植被的空间分布相互影响，互为因果关系。这种小尺度的空间格局差异是群落演替的驱动力，而不同的土地利用方式正是通过改变群落内小尺度的空间属性来影响了群落的演替方向。 在不同土地利用方式影响下，草地生态系统不仅在宏观功能，如生态系统生产力、群落结构和土壤的碳氮储量等方面表现出较大的差异，而且在微观结构上，如土壤养分和物种的空间属性等方面也发生了明显了变化，并且宏观和微观上的变化相互促进相互影响，它们的相互作用在很大程度上决定了群落的演替方向和草地生态系统全球变化过程中的具体作用。

内蒙古草原中生物土壤结皮生态学研究

1. 生物土壤结皮是干旱半干旱生态系统中的重要组成部分，它可通过增加土壤肥力和稳定性、影响水分再分配和植物萌发、成活、生长和繁殖而对生态系统结构和功能产生重要影响。为阐明生物结皮在内蒙古草地中的作用，本报告对两个草地生态系统进行了为期三年的调查，对生物结皮的氮素输入、对放牧的响应及其与植物的关系进行了综合研究。 2. 利用乙炔还原法，本研究对内蒙古退化草地中的生物土壤结皮中的蓝藻、地衣和地耳的氮素输入进行了为期30个月的连续测定。研究发现：1) 生物结皮固氮活性主要集中于5-10月，呈单峰型曲线，表明生物土壤结皮的固氮作用主要受温度和降雨影响;2) 生物结皮年固氮量为12.99-129.9 kg N•ha-1，98%的氮素固定于6-9月份;3) 按固氮量排序，蓝藻 (61%) > 地衣 (33%) > 地耳 (6%)，表明物种组成和丰度对生物结皮的氮素输入具重要影响。生物结皮的固氮量和季节变化表明生物结皮可以是影响退化草地中植物生长和促进退化草地恢复的重要影响因子。 3. 本研究选择三个放牧处理（长期放牧、短期围封和近期放牧）对生物结皮固氮活性进行了为期3年的研究。结果表明，与短期围封相比，长期放牧造成生物土壤结皮固氮活性下降了99.5%。固氮活性在放牧时间不足11个月时即可下降至最低水平，因此，放牧持续时间短于4个月的轮牧可能有利于生物土壤结皮的固氮。 4. 本研究选择6个放牧梯度（对照：0.00 羊/公顷, 极轻度放牧：1.33羊/公顷，轻度放牧：2.67羊/公顷，中度放牧：4.00羊/公顷，重度放牧：5.33羊/公顷，极重度放牧：6.67羊/公顷)，研究放牧强度对于生物结皮丰度、物种组成和固氮输入的影响。不同放牧强度对生物结皮丰度、物种组成和固氮输入具有重要影响，表明长期放牧可抑制生物土壤结皮在氮素输入和土壤固定方面的作用。极轻度放牧对生物土壤结皮影响不大;轻度放牧造成氮素输入降低了50%;重度和极重度放牧造成氮素输入降低了90%，并可使移动性较强的物种成为生物土壤结皮的优势组分，从而可抑制其土壤固定作用。因此，极轻度和轻度放牧是有利于生物土壤结皮固氮和固定土壤的草地利用方式。 5. 在处于恢复早期的一个退化草地中，我们对生物结皮和植物之间的关系进行了为期2年的研究。结果表明，生物结皮的丰度和物种组成与植物地上生物量和盖度高度相关。生物结皮丰度和氮素输入随植物生物量和盖度下降。结果还表明生物结皮是退化草地的主要氮素输入来源，尤其是在草地恢复初期。植物组织δ15N 低于土壤，这种差异随植物生产力增高而减小，表明生物结皮所固定氮素首先被植物利用，而后返回土壤。生物结皮的固氮输入变化可能是这种变化模式的主要原因，在分解作用和氮素损失中的同位素分馏，以及菌根真菌对于氮素的转运可能也是这种变化模式的原因。结果还显示生物土壤结皮与植物之间可能存在负反馈关系。这种自我调节的反馈过程可能是影响退化草地生态系统生产力和氮素循环的重要调节机制。

施肥和降水增加对半干旱草地生产力和温室气体排放的影响

人类活动能显著地改变生态系统的属性和功能。这些变化不仅发生在容易观察的局部层面上，如土壤沙化、植被退化等;还能更深刻地影响到微生物控制的温室气体释放，进而影响大尺度上气候的变化。本论文根据2 年田间试验结果报告了人类干扰（草地退耕和施肥）和降水增加30%对中国北部农牧交错区（内蒙古多伦县）半干旱草地生态系统生产力和温室气体排放的影响。 本研究发现氮是影响该区草地和弃耕地生产力的最重要因子，即使低量的氮肥应用(5 g N y-1)也能使该区草地地上净初级生产力（ANPP）增加36-46%。然而氮对草地生产力影响主要集中在地上部分，对地下净初级生产力（BNPP） 影响比较微弱。单独使用磷肥对ANPP 和BNPP 均无显著影响，氮磷混施对ANPP 有显著正交互作用。增加的降水对草地和弃耕地ANPP 和BNPP 有弱的正效应，增加30%降水使草地对照ANPP 增加9-25%，BNPP 增加12%-17%，弃耕地对照ANPP 增加7%-37%，BNPP 增加36%-37% 。草地和弃耕地对照ANPP 并无显著差别，由于高于弃耕地1-3 倍的BNPP，总体上草地比弃耕地有更高的固碳能力。这个结果也说明草地能更有效地把植物地上部分固定的碳转移到地下，这与草地长期发育积累的丰富地下根系和高根冠比物种组成有关。 水肥处理及土地利用方式也深刻改变了温室气体排放。温带草地和弃耕地CO2 排放（系统呼吸：Re）存在显著季节变化，温度、水分和地上生物量是控制这些变化的主要因子。在气候温暖的6-9 月份，生物量和土壤水分是决定系统呼吸季节变化主要因子，而与温度无明显相关。随着氮肥使用，草地和弃耕地呼吸速率加大，其温度敏感系数Q10 也明显升高，这主要与氮肥使用后增加的地上生物量有关。由于水分促进作用和生物量的弱增长，增加的降水也加大了草地和弃耕地CO2 排放。低的地下生物量和有机碳氮并没有导致弃耕地低的CO2 排放，显示了农垦后土壤结构破坏导致的有机碳分解增加不会在农业弃耕后短期内（5-6 年）停止。 除了对生物量的不敏感性外，中国北部半干旱生态系统N2O 排放表现出与CO2 排放类似的变化规律：水分和温度是N2O 季节变化的主导因子;氮肥使用和降水增加都增加了N2O 排放;N2O 排放在草地对照和弃耕地对照之间没有显著差别。更少的根系对无机氮的竞争导致了弃耕地比草地有更高的N2O 释放因子（单位氮添加所引起N2O 释放）。低的土壤水分（<70% WFPS ）和无机氮都是限制反硝化发生的重要条件，因此硝化可能是草地和弃耕地N2O 的最主要来源。半干旱草地生态系统N2O 排放与CO2 排放之间存在着显著线性关系。 中国北部半干旱草地和弃耕地都起着CH4 汇的生态功能。水分是控制CH4 吸收季节变异的主导变量，土壤空隙水含量（WFPS）与CH4 指数递减方程能解释其变异的64%-83% 。增加的降水导致了土壤水分增加，从而引起甲烷氧化菌所需基质（CH4 和O2）扩散受阻，减小了土壤CH4 吸收功能。施肥并没有抑制CH4 吸收，相反由于施肥后植物速生对土壤水的抽干作用，施肥增加了植物速生期时的CH4 吸收。CH4 吸收对农垦引起的土壤微生物生境结构破坏非常敏感，在农业措施停止5-6 年后的弃耕地，CH4 吸收仍比草地低24%。 本研究初步分析了影响中国北部农牧交错区草地生态系统生产力及温室气体排放的自然环境因子和人为因子，给出了它们之间的定性、定量关系，为科学地管理中国北部草地，充分发挥其生态和经济效益功能提供了有效信息。

The distribution and economic losses of alien species invasion to China

Invasive alien species have become one of the most serious environmental issues in the world. Data of taxon, origin, pathway, and environmental impacts of invasive alien microorganisms, invertebrates, amphibians and reptiles, fish, birds, mammals, herbs, trees, and, marine organisms in terrestrial, aquatic, and marine ecosystems of China were analyzed during 2001 and 2003, based on literature retrieval and field survey. There were 283 invasive alien species in China, and the number of species of invasive alien microorganisms, aquatic plants, terrestrial plants, aquatic invertebrates, terrestrial invertebrates, amphibians and reptiles, fish, and mammals were 19, 18, 170, 25, 33, 3, 10, and 5, respectively. The proportion of invasive alien species originated from America, Europe, Asia, Africa, and Oceania were 55.1, 21.7, 9.9, 8.1, and 0.6%, respectively. Methods for estimation of direct economic losses to agriculture, forestry, stockbreeding, fishery, road and water transportation, storage, water conservancy, environment and public facilities, and human health were established. Methods for estimation of indirect economic losses caused by invasive alien species to service functions of forest ecosystems, agricultural ecosystems, grassland ecosystems, and wetland ecosystems were also established. The total economic losses caused by invasive alien species to China were to the time of USD 14.45 billion, with direct and indirect economic losses accounting for 16.59% and 83.41% of total economic losses, respectively.

草地生态系统土壤呼吸及其影响因素研究进展

土壤呼吸在全球碳收支中占有重要的地位,笔者对草地生态系统土壤呼吸在陆地生态系统碳平衡中的作用、土壤呼吸的分类及其影响因素等方面进行了综述。结果表明,草地生态系统土壤呼吸在不同时间空间各组分所占比例不同,生物、非生物及人为活动等因素对草地土壤呼吸影响各异,主要从土壤温度、气候变暖、土壤湿度、降水、干旱化、土壤C/N等非生物因素,叶面积指数、植物光合作用、植被凋落物等生物因素以及人类干扰活动等方面具体阐述这些因素变化对土壤呼吸产生的影响,并对草地土壤呼吸的Q10值及各影响因素间的交互作用进行归纳总结。提出草地生态系统土壤呼吸研究存在的问题和今后重点发展方向,并对未来草地生态系统土壤呼吸的研究工作做了进一步的展望。

Considerable effects of diversity indices and spatial scales on conclusions relating to ecological diversity

The relationships between ecological diversity and ecosystem functions such as stability and productivity have long been debated and have no final conclusion until now. It is ignored that the debate should be firstly based on the same diversity index, which should be theoretically complete, and on same observation scale. For the issue on the scale of ecotope observation, ecosystems should be distinguished according to intensity of human disturbance. For the issue on the scale of species observation, either number diversity or biomass diversity should be identified. This paper takes grassland ecosystems located within the Bayin Xile grassland of Xilin Gol League of Inner Mongolia Autonomous Region as an example to analyze effects of different diversity indices and spatial scales on the conclusions of ecological diversity and its relationships with ecosystem functions. The analysis results both on the scale of ecotope observation and on the scale of species observation show that different diversity indices might give different conclusions and spatial resolution has a great effect on the relative conclusions. (c) 2005 Elsevier B.V. All rights reserved.

Relationship between species diversity and ecotope diversity

The relationship between species diversity and ecotope diversity has long been debated. But these debates seem meaningless because most of them were based on different definitions. In this paper, diversity has two components: richness based on the total number and evenness based on the relative abundance. Species diversity is distinguished into individual-counting diversity and biomass-based diversity. Ecotope diversity is divided into individual ecotope-counting diversity and ecotope-area based diversity. Under this definition, we make a comprehensive investigation into Dongzhi tableland of Loess Plateau by cooperating with local technicians. We find that individual-counting diversity is significantly correlated with biomass-based diversity in grassland ecosystems; individual ecotope-counting diversity and ecotope-area based diversity have a significant correlation. Therefore, it is unnecessary to divide species diversity into individual-counting diversity and biomass-based diversity in grassland ecosystems and to distinguish ecotope diversity into individual ecotope-counting and ecotope-area based diversity for the issues that have no special requirement for accuracy. But the analyses of the investigation data demonstrate that species diversity has no significant correlation with ecotope diversity.

Depression of net ecosystem CO2 exchange in semi-arid Leymus chinensis steppe and alpine shrub

Uptake and release of carbon in grassland ecosystems is very critical to the global carbon balance and carbon storage. In this study, the dynamics of net ecosystem CO2 exchange (FNEE) of two grassland ecosystems were observed continuously using the eddy covariance technique during the growing season of 2003. One is the alpine shrub on the Tibet Plateau, and the other is the sem-arid Leymus chinensis steppe in Inner Mongolia of China. It was found that the FNEE of both ecosystems was significantly depressed under high solar radiation. Comprehensive analysis indicates that the depression of FNEE in the L. chinensis steppe was the results of decreased plant photosynthesis and increased ecosystem respiration (R-eco) under high temperature. Soil water stress in addition to the high atmospheric demand under the strong radiation was the primary factor limiting the stomatal conductance. In contrast, the depression of FNEE in the alpine shrub was closely related to the effects of temperature on both photosynthesis and ecosystem respiration, coupled with the reduction of plant photosynthesis due to partial stomatal closure under high temperature at mid-day. The R,c of the alpine shrub was sensitive to soil temperature during high turbulence (u* > 0.2 m s(-1)) but its FNEE decreased markedly when the temperature was higher than the optimal value of about 12 degrees C. Such low optimal temperature contrasted the optimal value (about 20 degrees C) for the steppe, and was likely due to the acclimation of most alpine plants to the long-term low temperature on the Tibet Plateau. We inferred that water stress was the primary factor causing depression of the FNEE in the semi-arid steppe ecosystem, while relative high temperature under strong solar radiation was the main reason for the decrease of FNEE in the alpine shrub. This study implies that different grassland ecosystems may respond differently to climate change in the future. (c) 2006 Elsevier B.V All rights reserved.

Stability of alpine meadow ecosystem on the Qinghai-Tibetan Plateau

The meadow ecosystem on the Qinghai-Tibetan Plateau is considered to be sensitive to climate change. An understanding of the alpine meadow ecosystem is therefore important for predicting the response of ecosystems to climate change. In this study, we use the coefficients of variation (Cv) and stability (E) obtained from the Haibei Alpine Meadow Ecosystem Research Station to characterize the ecosystem stability. The results suggest that the net primary production of the alpine meadow ecosystem was more stable (Cv = 13.18%) than annual precipitation (Cv = 16.55%) and annual mean air temperature (Cv= 28.82%). The net primary production was insensitive to either the precipitation (E = 0.0782) or air temperature (E = 0.1113). In summary, the alpine meadow ecosystem on the Qinghai-Tibetan Plateau is much stable. Comparison of alpine meadow ecosystem stability with other five natural grassland ecosystems in Israel and southern African indicates that the alpine meadow ecosystem on the Qinghai-Tibetan Plateau is the most stable ecosystem. The alpine meadow ecosystem with relatively simple structure has high stability, which indicates that community stability is not only correlated with biodiversity and community complicity but also with environmental stability. An average oscillation cycles of 3-4 years existed in annual precipitation, annual mean air temperature, net primary production and the population size of consumers at the Haibei natural ecosystem. The high stability of the alpine meadow ecosystem may be resulting also from the adaptation of the ecosystem to the alpine environment.

Energy exchange between the atmosphere and a meadow ecosystem on the Qinghai-Tibetan Plateau

To reveal the potential contribution of grassland ecosystems to climate change, we examined the energy exchange over an alpine Kobresia meadow on the northeastern Qinghai-Tibetan Plateau. The annual pattern of energy exchange showed a clear distinction between periods of frozen soil with the daily mean soil temperature at 5 cm (T-s5 &LE; 0 &DEG; C) and non-frozen soil (T-s5 > 0 &DEG; C). More than 80% of net radiation was converted to sensible heat (H) during the frozen soil period, but H varied considerably with the change in vegetation during the non-frozen soil period. Three different sub-periods were further distinguished for the later period: (1) the pre-growth period with Bowen ratio (β) > 1 was characterized by a high β of 3.0 in average and the rapid increase of net radiation associated with the increases of H, latent heat (LE) and soil heat; (2) during the Growth period when β &LE; 1, the LE was high but H fluxes was low with β changing between 0.3 and 0.4; (3) the post-growth period with average β of 3.6 when H increased again and reached a second maximum around early October. The seasonal pattern suggests that the phenology of the vegetation and the soil water content were the major factors affecting the energy partitioning in the alpine meadow ecosystem. © 2005 Elsevier B.V. All rights reserved.