草原灌丛化对生态系统多功能性的影响

草原灌丛化现象在干旱半干旱区广泛发生,影响了生态系统的结构、过程和功能。生态系统具有同时提供多种功能的能力,即生态系统多功能性。灌丛化是否会引起草原生态系统多功能性的减少,其内在的作用机制又是什么？这些问题仍有待明晰。理解草原灌丛化对生态系统多功能性的影响,对于促进草原地区"草-畜-人"平衡和实现区域可持续发展至关重要。从响应规律、影响路径和控制因素三个方面总结评述了草原灌丛化对生态系统多功能性影响的研究进展,主要包括:(1)阐明了单一生态系统功能和多种生态系统功能对草原灌丛化的响应特征;(2)从生物路径、非生物路径以及气候变化和人类活动的影响方面探讨了灌丛化对生态系统多功能性的影响路径;(3)从灌丛化物种、灌丛化阶段和草原类型三个方面明晰了草原灌丛化对生态系统多功能性影响的控制因素。在此基础上,针对灌丛化对生态系统多功能性的影响机制,对生产-生态功能权衡的影响等方面对未来研究进行了展望,并面向可持续发展目标探讨了灌丛化生态系统的可持续管理路径。研究可为我国灌丛化草原的恢复和管理提供支撑。     

Detecting shrub encroachment in seminatural grasslands using UAS LiDAR

1. Shrub encroachment in seminatural grasslands threatens local biodiversity unless management is applied to reduce shrub density. Dense vegetation of Cytisus scoparius homogenizes the landscape negatively affecting local plant diversity. Detecting structural change (e.g., biomass) is essential for assessing negative impacts of encroachment. Hence, exploring new monitoring tools to achieve this task is important for effectively capturing change and evaluating management activities.
2. This study combines traditional field‐based measurements with novel Light Detection and Ranging (LiDAR) observations from an Unmanned Aircraft System (UAS). We investigate the accuracy of mapping C. scoparius in three dimensions (3D) and of structural change metrics (i.e., biomass) derived from ultrahigh‐density point cloud data (>1,000 pts/m²). Presence–absence of 12 shrub or tree genera was recorded across a 6.7 ha seminatural grassland area in Denmark. Furthermore, 10 individuals of C. scoparius were harvested for biomass measurements. With a UAS LiDAR system, we collected ultrahigh‐density spatial data across the area in October 2017 (leaf‐on) and April 2018 (leaf‐off). We utilized a 3D point‐based classification to distinguish shrub genera based on their structural appearance (i.e., density, light penetration, and surface roughness).
3. From the identified C. scoparius individuals, we related different volume metrics (mean, max, and range) to measured biomass and quantified spatial variation in biomass change from 2017 to 2018. We obtained overall classification accuracies above 86% from point clouds of both seasons. Maximum volume explained 77.4% of the variation in biomass.
4. The spatial patterns revealed landscape‐scale variation in biomass change between autumn 2017 and spring 2018, with a notable decrease in some areas. Further studies are needed to disentangle the causes of the observed decrease, for example, recent winter grazing and/or frost events.
5. Synthesis and applications: We present a workflow for processing ultrahigh‐density spatial data obtained from a UAS LiDAR system to detect change in C. scoparius. We demonstrate that UAS LiDAR is a promising tool to map and monitor grassland shrub dynamics at the landscape scale with the accuracy needed for effective nature management. It is a new tool for standardized and nonbiased evaluation of management activities initiated to prevent shrub encroachment.

     

人工灌丛化对荒漠草原生态系统碳储量的影响

宁夏盐池县从20世纪70年代开始在荒漠草原上人工种植柠条灌木用以防风固沙和生态恢复,这一人为措施极大地改变了区域生态系统的植被结构和碳循环,而定量评估人工灌丛化对荒漠草原生态系统碳储量的影响,不仅能够揭示人类活动的碳循环反馈机制,而且可为地方政府生态治理提供理论指导。结合Biome-BGC模型和Logistics生长模型模拟了1958—2017年间荒漠草原人工灌丛化前后的碳储量变化,定量分析了人工灌丛化对生态系统碳储量和组分的影响。结果表明:(1)结合Biome-BGC模型和Logistics生长模型可以较准确地模拟出荒漠草原人工灌丛化过程中生态系统碳储量的变化。(2)人工灌丛化会快速改变荒漠草原的碳储量累积特征,柠条灌木种植后的快速生长阶段极大增强了生态系统的总碳储量,导致生态系统的碳储量特征由草地型向灌木型转变。(3)人工灌丛化改变了生态系统各类型碳储量的组分结构,其对地上植被和枯落物碳储量的影响非常明显,灌丛化后生态系统的植被和枯落物碳分别增加了6倍和1.76倍;因植被碳向土壤碳转化过程较慢,故人工灌丛化对地下土壤碳储量的影响在短期内较为微弱。以上结果显示,荒漠草原人工灌丛化能显...     

Using consecutive prescribed fires to reduce shrub encroachment in grassland by increasing shrub mortality

Woody plant encroachment into open grasslands occurs worldwide and causes multiple ecological and management impacts. Prescribed fire could be used to conserve grassland habitat but often has limited efficacy because many woody plants resprout after fire and rapidly reestablish abundance. If fire‐induced mortality could be increased, prescribed fire would be a more effective management tool. In California's central coast, shrub encroachment, especially of Baccharis pilularis (coyote brush), is converting coastal prairie into shrub‐dominated communities, with a consequent loss of native herbaceous species and open grassland habitat. B. pilularis has not been successfully controlled with single prescribed fire events because the shrub resprouts and reestablishes cover within a few years. We investigated whether two consecutive annual burns would control B. pilularis by killing resprouting shrubs, without reducing native herbaceous species or encouraging invasive plants. As expected, resprouting did occur; however, 2 years after the second burn, B. pilularis cover on burned plots was only 41% of the cover on unburned plots. Mortality of B. pilularis more than doubled following the second burn, likely maintaining a reduction in B. pilularis cover for longer than a single burn would have. Three native coastal prairie perennial grasses did not appear to be adversely affected by the two burns, nor did the burns result in increased cover of invasive species. Managers wanting to restore coastal prairie following B. pilularis encroachment should consider two consecutive annual burns, especially if moderate fire intensity is achievable.     

Carabid beetles as indicators for shrub encroachment in dry grasslands

Semi-natural grasslands are key habitats for biodiversity conservation in Central Europe. Shrub encroachment is one of the most threatening drivers of grassland degradation and affects soil properties, microclimate, and vegetation with possible impacts on higher trophic levels. We aimed to analyse the impact of shrub encroachment with broom (Cytisus scoparius) on carabid beetle diversity, species composition, and functional traits. In a field study on dry grasslands on the island of Hiddensee (Germany) we studied 15 sites along a gradient of increasing broom encroachment and classified them into three dry grassland types with low, medium, and high shrub cover. Our results provide evidence that shrub encroachment initially has positive effects on species richness and activity densities of dry grassland carabids. Carabid species composition differed among differently shrub-covered dry grassland types, and sites with low and high shrub cover were each characterised by unique carabid assemblages. The species composition of sites with a medium shrub biomass had a transitional character and contained species which are typical for open dry grassland, but also shared species with sites with a high shrub cover. Among functional trait parameters investigated, especially the body size of carabid beetles was related to environmental parameters associated with shrub encroachment. Body size was positively correlated to shrub biomass and soil humidity, but negatively to temperature. Eurytopy values of carabids were related to high litter cover, i.e. habitat generalist (eurytopic) species mainly occurred in densely shrub-encroached sites. In order to preserve unique carabid assemblages of open dry grasslands with stenotopic and smaller species, it is most important to prevent a shrub encroachment higher than about 60% cover. For management we suggest extensive grazing (by cattle, sheep or horses) to prevent shrub encroachment on dry grasslands. In areas with high shrub cover additionally the use of goats or mechanical removal of shrubs might be necessary.     

Shrub encroachment effect on the evapotranspiration and its component—A numerical simulation study of a shrub encroachment grassland in Nei Mongol,China

Aims Shrub encroachment is a common global change phenomenon occurring in arid and semi-arid regions. Due to the difficulty of partitioning evapotranspiration into shrub plants, grass plants and soil in the field, there are few studies focusing on shrub encroachment effect on the evapotranspiration and its component in China. This study aims to illustrate shrub encroachment effect on evapotranspiration by the numerical modeling method. Methods A two-source model was applied and calibrated with the measured evapotranspiration (ET) by the Bowen ratio system to simulate evapotranspiration and its component in a shrub encroachment grassland in Nei Mongol, China. Based on the calibrated model and previous shrub encroachment investigation, we set three scenarios of shrub encroachment characterized by relative shrub coverage of 5%, 15% and 30%, respectively, and quantified their effects caused by shrub encroachment through localized and calibrated two-source model.Important findings The two-source model can well reconstruct the evapotranspiration characteristics of a shrub encroachment grassland. Sensitivity analysis of the model shows that errors for the input variables and parameters have small influence on the result of partitioning evapotranspiration. The result shows that shrub encroachment has relatively small influence on the total amount of ET, but it has clear influence on the proportion of the components of evapotranspiration (E/ET). With shrub coverage increasing from 5% to 15% and then 30%, the evapotranspiration decreased from 182.97 to 180.38 and 176.72 W·m^-2, decreasing amplitude values of 0.34% and 0.44%, respectively. On average, E/ET rises from 52.9% to 53.9% and 55.5%, increasing amplitude values to 2.04% and 3.25%. Data analysis indicates that shrub encroachment results in smaller soil moisture changes, but clear changes of ecosystem structure (decreasing ecosystem leaf area index while increasing vegetation height) which lead to the decrease of transpiration fraction through decreasing canopy conductance. The research highlights that, with the shrub encroachment, more water will be consumed as soil evaporation which is often regarded as invalid part of evapotranspiration and thus resulting in the decrease of water use efficiency.     

Mixtures with grass litter may hasten shrub litter decomposition after shrub encroachment into mountain grasslands

Aims

Shrub encroachment in mesic grasslands alters the identity and quality of litters entering the system. As litter from shrubs and grasses can differ in their quality, this can lead to differences in litter decomposition by the direct effect of quality, but also to litter interaction during decomposition. The objective of this study was to examine the occurrence of non-additive effects of litter mixtures on the decomposition rates of legume shrub litter (poor in P) or conifer shrub litter (poor in N) and grass litter.

Methods

In addition to single litter type litterbags for the three species, we mixed litters of each pair of possible combinations to determine the influence of each species on mass loss. Litterbags were placed in the field and collected after 1, 6, 8, 12 and 24 months. In each collection, litter of each species remaining in mixed bags was separated, dry weighed and analyzed for C, N and P.

Results

With respect to shrub litter decomposing alone, mass loss of shrub litter when mixed with grass showed a 9–10 % increase in decomposition rate for conifer and a 3 % increase for legume litter. These litter mixture effects varied with time and they were detected after a decomposition period of 1 year in legume litter and of 2 years in conifer litter.

Conclusions

Grass litter hastened conifer and legume litter decomposition in leaf litter mixtures, at least during the first stages of the process. The potential consequences of this result to alter litter accumulation patterns and thus carbon sequestration rates after shrub encroachment into grasslands will depend on whether the observed trends are maintained in the advanced decomposition stages.     

内蒙古典型草原灌丛化对生物量和生物多样性的影响

通过样方调查,研究了内蒙古典型草原不同退化程度草地中小叶锦鸡儿 (Caragana microphylla Lam.)灌丛斑块空间分布格局、草地生物量及生物多样性特征。结果表明,从轻度到中度、重度退化草地,草本植物生物量呈减少趋势,小叶锦鸡儿生物量呈增加趋势,总生物量呈先减少后增加趋势;灌丛斑块生物多样性呈增加趋势,草地斑块生物多样性呈先减少后增加趋势,其中轻度退化草地中灌丛斑块生物多样性低于草地斑块,中度和重度退化草地中灌丛斑块生物多样性高于草地斑块。本研究认为,内蒙古典型草原灌丛化过程中,生态系统可能存在草本植物占优势或小叶锦鸡儿占优势这样两种稳定状态,这两种状态能维持较高生物量和生物多样性,而在由草本植物占优势向小叶锦鸡儿占优势转化的过渡阶段,系统不稳定,仅能维持较低生物量和生物多样性。     

Soil fungal composition changes with shrub encroachment in the northern Chihuahuan Desert

Woody species encroachment of grasslands globally causes many socioecological impacts, including loss of grazing pastures and decreased biodiversity. Soil microbial communities may partially regulate the pace of shrub encroachment, as plant-microbial interactions can strongly influence plant success. We measured fungal composition and activity under dominant plant species across a grassland to shrubland transition to determine if shrubs cultivate soil microbial communities as they invade. Specifically, soil microbial communities, abiotic soil properties, and extracellular enzyme activities were quantified for soils under four common Chihuahuan Desert plant species (three grasses, one shrub) in central New Mexico, U.S.A. Extracellular enzyme activity levels were fairly consistent under different plant species across the grassland to shrubland transition. Activity levels of two enzymes (alkaline phosphatase and beta-N-acetyl-glucosaminidase) were lower in the ecotone, presumably because soil organic matter content was also lower in ecotone soils. Community composition of soil fungi mirrored patterns in the plant community, with distinct plant and fungal communities in the shrubland and grassland, while grassland-shrubland ecotone soils hosted a mix of taxa from both habitats. We show that shrubs cultivate a distinct microbial community on the leading edge of the invasion, which may be necessary for shrub colonization, establishment, and persistence.     

Impacts of shrub encroachment on ecosystem structure and functioning: towards a global synthesis

Encroachment of woody plants into grasslands has generated considerable interest among ecologists. Syntheses of encroachment effects on ecosystem processes have been limited in extent and confined largely to pastoral land uses or particular geographical regions. We used univariate analyses, meta-analysis and structural equation modelling to test the propositions that (1) shrub encroachment does not necessarily lead to declines in ecosystem functions and (2) shrub traits influence the functional outcome of encroachment. Analyses of 43 ecosystem attributes from 244 case studies worldwide showed that some attributes consistently increased with encroachment (e.g. soil C, N), and others declined (e.g. grass cover, pH), but most exhibited variable responses. Traits of shrubs were associated with significant, though weak, structural and functional outcomes of encroachment. Our review revealed that encroachment had mixed effects on ecosystem structure and functioning at global scales, and that shrub traits influence the functional outcome of encroachment. Thus, a simple designation of encroachment as a process leading to functionally, structurally or contextually degraded ecosystems is not supported by a critical analysis of existing literature. Our results highlight that the commonly established link between shrub encroachment and degradation is not universal.     

The impact of shrub encroachment on savanna bird diversity from local to regional scale

Aim  Evidence is accumulating of a general increase in woody cover of many savanna regions of the world. Little is known about the consequences of this widespread and fundamental ecosystem structural shift on biodiversity.
Location  South Africa.
Methods  We assessed the potential response of bird species to shrub encroachment in a South African savanna by censusing bird species in five habitats along a gradient of increasing shrub cover, from grassland/open woodland to shrubland dominated by various shrub species. We also explored historical bird species population trends across southern Africa during the second half of the 20th century to determine if any quantifiable shifts had occurred that support an ongoing impact of shrub encroachment at the regional scale.
Results  At the local scale, species richness peaked at intermediate levels of shrub cover. Bird species composition showed high turnover along the gradient, suggesting that widespread shrub encroachment is likely to lead to the loss of certain species with a concomitant decline in bird species richness at the landscape scale. Finally, savanna bird species responded to changes in vegetation structure rather than vegetation species composition: bird assemblages were very similar in shrublands dominated by Acacia mellifera and those dominated by Tarchonanthus camphoratus .
Main conclusions  Shrub encroachment might have a bigger impact on bird diversity in grassland than in open woodland, regardless of the shrub species. Species recorded in our study area were associated with historical population changes at the scale of southern Africa suggesting that shrub encroachment could be one of the main drivers of bird population dynamics in southern African savannas. If current trends continue, the persistence of several southern African bird species associated with open savanna might be jeopardized regionally.     

Effects of shrub encroachment on the anuran community in periodically flooded grasslands of the largest Neotropical wetland

In many parts of the world, replacement of natural grasslands by woody plants has resulted in a decrease of pasture areas and in habitat loss for a variety of animal species, including amphibians. Wetlands are especially susceptible to invasive plants, both native and exotic, but the effects of such invasions on animal assemblages remain poorly understood. Here, we present information on the impact of selected environmental variables, especially coverage by the native shrub Combretum laxum Jacq., on the structure of an anuran assemblage in the Pantanal, a huge flood‐pulsed South American wetland. Anurans were surveyed during the rainy season in 17 plots, which differed in extent of C. laxum coverage, leaf litter volume, soil moisture and distance to permanently wet areas. Effects of these environmental variables on the species number, relative abundance and composition of the anuran assemblage were evaluated using multivariate statistical analyses. We captured 1203 anurans, of 21 species from four families. Both the number of species and the relative abundance of anurans were lower in plots with greater C. laxum coverage, which also influenced anuran species composition. Number of species was highest in plots located closest to permanently wet areas, which provide protection from desiccation and other resources during the Pantanal dry season, and so could be considered source areas of anurans. While many anuran species were negatively affected by the homogenization of the landscape caused by shrub encroachment, some seemed to be favoured in such circumstances. For these, dense shrub encroachment into natural grasslands may provide safer migratory routes to permanently wet habitats. Thus, at the mesoscale, a mosaic of areas with different levels of coverage by C. laxum (shrub islands) may aid anuran assemblages in the Pantanal wetlands, facilitating the maintenance of higher beta and gamma diversity.     

Spatial patterns of shrub encroachment in neighbouring grassland communities in the Pyrenees: floristic composition heterogeneity drives shrub proliferation rates

We studied the role of floristic composition and associational resistance in shrub dynamics by comparing spatial patterns of shrub cover after prescribed burning in neighbouring grassland communities with different palatability. The study focused on the shrub Cytisus balansae ssp. europaeus (G. López and Jarvis) Muñoz Garmendia. Seven two-dimensional transects (20 × 0.5 m) were established to monitor shrub cover for at least 10 years after prescribed burning. Shrub cover and spatial patterns were assessed in each transect. Floristic similarity between transects and Cytisus associations with different species were estimated. Over an entire transect, shrub cover and shrub scale of pattern and patch size were lowest in the unpalatable Festuca eskia grasslands and highest in F. paniculata grasslands. At short distances, we found negative associations between Cytisus and most of the grasses, except for F. nigrescens and Agrostis capillaris, which showed positive associations with Cytisus. Thus, the effects of associational resistance on shrub encroachment were not as marked as expected, F. eskia grasslands showing the lowest shrub encroachment rates after fire. By contrast, Cytisus was positively associated with the most palatable grasses in the site, namely F. nigrescens and A. capillaris. We conclude that differences in floristic composition drive shrub encroachment rates in these spatially heterogeneous communities.     

Seed dispersal by cattle may cause shrub encroachment of Grewia flava on southern Kalahari rangelands

Abstract. Shrub encroachment, i.e. the increase in woody plant cover, is a major concern for livestock farming in southern Kalahari savannas. We developed a grid‐based computer model simulating the population dynamics of Grewia flava, a common, fleshy‐fruited encroaching shrub. In the absence of large herbivores, seeds of Grewia are largely deposited in the sub‐canopy of Acacia erioloba. Cattle negate this dispersal limitation by browsing on the foliage of Grewia and dispersing seeds into the grassland matrix. In this study we first show that model predictions of Grewia cover dynamics are realistic by comparing model output with shrub cover estimates obtained from a time series of aerial photographs. Subsequently, we apply a realistic range of intensity of cattle‐induced seed dispersal combined with potential precipitation and fire scenarios. Based on the simulation results we suggest that cattle may facilitate shrub encroachment of Grewia. The results show that the severity of shrub encroachment is governed by the intensity of seed dispersal. For a high seed dispersal intensity without fire (equivalent to a high stocking rate) the model predicts 56% shrub cover and 85% cell cover after 100 yr. With fire both recruitment and shrub cover are reduced, which may, under moderate intensities, prevent shrub encroachment. Climate change scenarios with two‐fold higher frequencies of drought and wet years intensified shrub encroachment rates, although long‐term mean of precipitation remained constant. As a management recommendation we suggest that shrub encroachment on rangelands may be counteracted by frequent fires and controlling cattle movements to areas with a high proportion of fruiting Grewia shrubs.     

祁连山东缘高寒草甸灌丛化对植被群落与土壤理化特性的影响

为明晰祁连山东缘高寒草甸灌丛化对其植被群落与土壤理化特性的影响,选取甘肃省天祝藏族自治县高寒草甸灌丛化区域,调查其植被,采集土壤测定理化性质,分析植被与土壤的互馈关系。结果表明:高寒草甸植被的均匀度指数、高度、盖度、地上生物量、土壤含水率随灌丛化过程增加(P<0.05),而多样性指数和容重均降低;全磷含量在各土层以灌丛区最高,但C ∶ P和N ∶ P以草甸区最高;土壤N ∶ P<14表明土壤养分主要受限于氮;土壤脲酶和过氧化氢酶活性在草甸区各土层显著高于灌丛区(P<0.05);冗余分析显示前两个排序轴的累计解释量为54.88%,土壤过氧化氢酶与植被特征拟合的结果最好(Pr=0.010),表明高寒草甸灌丛化过程中土壤过氧化氢酶受植被特征的制约最大。可见,高寒草甸灌丛化显著影响了植被群落结构与土壤理化特性,在祁连山生态保护和管理中应该予以重视。     

Cancer heterogeneity—a multifaceted view

Cancers of various organs have been categorized into distinct subtypes after increasingly sophisticated taxonomies. Additionally, within a seemingly homogeneous subclass, individual cancers contain diverse tumour cell populations that vary in important cancer‐specific traits such as clonogenicity and invasive potential. Differences that exist between and within a given tumour type have hampered significantly both the proper selection of patients that might benefit from therapy, as well as the development of new targeted agents. In this review, we discuss the differences associated with organ‐specific cancer subtypes and the factors that contribute to intra‐tumour heterogeneity. It is of utmost importance to understand the biological causes that distinguish tumours as well as distinct tumour cell populations within malignancies, as these will ultimately point the way to more rational anti‐cancer treatments.EMBO reports advance online publication 12 July 2013; doi:10.1038/embor.2013.92