小叶锦鸡儿灌丛化对典型草原群落结构与生态系统功能的影响

草原灌丛化是全球干旱半干旱地区面临的重要生态问题。灌丛化对草原生态系统结构与功能的影响较为复杂, 有待于在更广泛的区域开展研究。该研究在内蒙古锡林郭勒典型草原选择轻度、中度和重度灌丛化草地, 通过群落调查, 结合植物功能性状和土壤理化性质观测, 研究了小叶锦鸡儿(Caragana microphylla)灌丛化对草原群落结构(物种多样性、功能多样性和功能群组成)和生态系统功能(初级生产力、植被和土壤养分库)的影响。结果表明: 1)不同程度灌丛化草地的物种丰富度、功能性状多样性和群落加权性状平均值差异显著, 其中, 中度灌丛化草地的物种多样性和功能多样性较高, 表明一定程度的灌丛化有利于生物多样性维持。2)重度灌丛化草地的地上净初级生产力(ANPP)显著高于轻度和中度灌丛化草地, 其原因主要是随着灌丛化程度加剧, 群落内一/二年生草本植物显著增加, 而多年生禾草和多年生杂类草显著减少。三个灌丛化草地的植被叶片和土壤碳、氮库差异均不显著。3)灌丛化对草原生态系统功能包括ANPP、植被和土壤养分库均没有直接的影响, 而是通过影响功能群组成、土壤理化性质和功能多样性, 间接地影响生态系统功能; 灌丛化导致功能群发生替代和土壤旱碱化是最重要的生物和非生物因素。     

On the Importance of Shrub Encroachment by Sprouters, Climate, Species Richness and Anthropic Factors for Ecosystem Multifunctionality in Semi-arid Mediterranean Ecosystems

One of the most important changes taking place in drylands worldwide is the increase of the cover and dominance of shrubs in areas formerly devoid of them (shrub encroachment). A large body of research has evaluated the causes and consequences of shrub encroachment for both ecosystem structure and functioning. However, there are virtually no studies evaluating how shrub encroachment affects the ability of ecosystems to maintain multiple functions and services simultaneously (multifunctionality). We aimed to do so by gathering data from ten ecosystem functions linked to the maintenance of primary production and nutrient cycling and storage (organic C, activity of β-glucosidase, pentoses, hexoses, total N, total available N, amino acids, proteins, available inorganic P, and phosphatase activity), and summarizing them in a multifunctionality index (M). We assessed how climate, species richness, anthropic factors (distance to the nearest town, sandy and asphalted road, and human population in the nearest town at several historical periods) and encroachment by sprouting shrubs impacted both the functions in isolation and M along a regional (ca. 350 km) gradient in Mediterranean grasslands and shrublands dominated by a non-sprouting shrub. Values of M were higher in those grasslands and shrublands containing sprouting shrubs (43 and 62%, respectively). A similar response was found when analyzing the different functions in isolation, as encroachment by sprouting shrubs increased functions by 2–80% compared to unencroached areas. Encroachment was the main driver of changes in M along the regional gradient evaluated, followed by anthropic factors and species richness. Climate had little effects on M in comparison to the other factors studied. Similar responses were observed when evaluating the functions in isolation. Overall, our results showed that M was higher at sites with higher sprouting shrub cover, longer distance to roads and higher perennial plant species richness. Our study is the first documenting that ecosystem multifunctionality in shrublands is enhanced by encroaching shrubs differing in size and leaf attributes. Our findings reinforce the idea that encroachment effects on ecosystem functioning cannot be generalized, and that are largely dependent on the traits of the encroaching shrub relative to those of the species being replaced.     

Shrub and precipitation interactions shape functional diversity of nematode communities on the Qinghai–Tibet Plateau

Land use and climate change alter biodiversity patterns and ecosystem functioning worldwide. Land abandonment with consequent shrub encroachment and changes in precipitation gradients are known factors in global change. Yet, the consequences of interactions between these factors on the functional diversity of belowground communities remain insufficiently explored. Here, we investigated the dominant shrub effects on the functional diversity of soil nematode communities along a precipitation gradient on the Qinghai–Tibet Plateau. We collected three functional traits (life-history C–P value, body mass, and diet) and calculated the functional alpha and beta diversity of nematode communities using kernel density n-dimensional hypervolumes. We found that shrubs did not significantly alter the functional richness and dispersion, but significantly decreased the functional beta diversity of nematode communities in a pattern of functional homogenization. Shrubs benefited nematodes with longer life-history, larger body mass, and higher trophic levels. Moreover, the shrub effects on the functional diversity of nematodes depended strongly on precipitation. Increasing precipitation reversed the effects shrubs have on the functional richness and dispersion from negative to positive but amplified the negative effects shrubs have on functional beta diversity of nematodes. Benefactor shrubs had stronger effects on the functional alpha and beta diversity of nematodes than allelopathic shrubs along a precipitation gradient. A piecewise structural equation model showed that shrubs and its interactions with precipitation indirectly increased the functional richness and dispersion through plant biomass and soil total nitrogen, whereas it directly decreased the functional beta diversity. Our study reveals the expected changes in soil nematode functional diversity following shrub encroachment and precipitation, advancing our understanding of global climate change on nematode communities on the Qinghai–Tibet Plateau.     

Shrub encroachment can reverse desertification in semi-arid Mediterranean grasslands

The worldwide phenomenon of shrub encroachment in grass-dominated dryland ecosystems is commonly associated with desertification. Studies of the purported desertification effects associated with shrub encroachment are often restricted to relatively few study areas, and document a narrow range of possible impacts upon biota and ecosystem processes. We conducted a study in degraded Mediterranean grasslands dominated by Stipa tenacissima to simultaneously evaluate the effects of shrub encroachment on the structure and composition of multiple biotic community components, and on various indicators of ecosystem function. Shrub encroachment enhanced vascular plant richness, biomass of fungi, actinomycetes and other bacteria, and was linked with greater soil fertility and N mineralization rates. While shrub encroachment may be a widespread phenomenon in drylands, an interpretation that this is an expression of desertification is not universal. Our results suggest that shrub establishment may be an important step in the reversal of desertification processes in the Mediterranean region.     

The essential role of biodiversity in the key axes of ecosystem function

Biodiversity is essential for maintaining the terrestrial ecosystem multifunctionality (EMF). Recent studies have revealed that the variations in terrestrial ecosystem functions are captured by three key axes: the maximum productivity, water use efficiency, and carbon use efficiency of the ecosystem. However, the role of biodiversity in supporting these three key axes has not yet been explored. In this study, we combined the (i) data collected from more than 840 vegetation plots across a large climatic gradient in China using standard protocols, (ii) data on plant traits and phylogenetic information for more than 2,500 plant species, and (iii) soil nutrient data measured in each plot. These data were used to systematically assess the contribution of environmental factors, species richness, functional and phylogenetic diversity, and community-weighted mean (CWM) and ecosystem traits (i.e., traits intensity normalized per unit land area) to EMF via hierarchical partitioning and Bayesian structural equation modeling. Multiple biodiversity attributes accounted for 70% of the influence of all the variables on EMF, and ecosystems with high functional diversity had high resource use efficiency. Our study is the first to systematically explore the role of different biodiversity attributes, including species richness, phylogenetic and functional diversity, and CWM and ecosystem traits, in the key axes of ecosystem functions. Our findings underscore that biodiversity conservation is critical for sustaining EMF and ultimately ensuring human well-being.     

Disturbances by desert rodents are more strongly associated with spatial changes in soil texture than woody encroachment

Background and Aims

Soil texture is an important determinant of ecosystem structure and productivity in drylands, and may influence animal foraging and, indirectly, plant community composition.

Methods

We measured the density and composition of surface disturbances (foraging pits) of small, soil-foraging desert vertebrates in shrubland and grasslands, both with coarse- and fine-textured soils. We predicted that the density and functional significance of disturbances would be related more to differences in texture than shrub encroachment.

Results

Soil texture had a stronger influence on animal foraging sites than shrub encroachment. There were more disturbances, greater richness and abundance of trapped seed, and greater richness of germinating plants on coarse- than fine-textured soils. Pits in coarse soils trapped 50 % more litter than those in finer soils. Apart from slightly more soil removal and greater litter capture in shrubland pits, there were no effects of encroachment.

Conclusions

Although the process of woody encroachment has been shown to have marked effects on some ecosystem properties, it is likely to have a more subordinate effect on surface disturbances and therefore their effects on desert plant communities than soil texture. Our results highlight the importance of animal activity in shaping desert plant communities, and potentially, in maintaining or reinforcing shrub dominant processes.     

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.     

草灌植被转变对草地生态系统及其水碳过程的影响研究进展

气候变暖和过度放牧的共同作用使全球草地出现明显的灌丛化现象,灌木去除是草地灌丛化控制的重要方式,识别这些草灌植被转变对生态系统、生态水文、土壤侵蚀和侵蚀碳流失的影响对草地可持续管理具有重要意义。综述了草地灌木入侵及其控制对植物群落和土壤功能(如土壤有机碳)的影响,以及这些草灌植被变化对生态水文、土壤侵蚀和土壤侵蚀碳流失等水碳耦合过程的影响机制。针对目前草地灌木入侵和去除对植物群落、植被格局、水土过程和功能影响研究的薄弱环节,对未来相关研究提出以下建议：(1)需深化草灌植被转变对碳、氮等生物地球化学循环的影响机制研究,(2)需重视核磁共振光谱、生物标志物、同位素等新技术和植被格局的指数与连通性等新方法在草灌植被转变的水、碳等生态效应研究中的应用,(3)需加强草灌植被格局和生态水文、土壤侵蚀与土壤侵蚀碳等水碳过程的多要素、多过程和多尺度的综合研究。本文旨在为灌丛化草地科学有效的生态恢复与多目标的土地利用管理提供理论支撑。     

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...     

Interactive effects of species richness and species traits on functional diversity and redundancy

The importance of species diversity for ecosystem function has emerged as a key question for conservation biology. Recently, there has been a shift from examining the role of species richness in isolation towards understanding how species interact to effect ecosystem function. Here, we briefly review theoretical predictions regarding species contributions to functional diversity and redundancy and further use simulated data to test combined effects of species richness, number of functional traits, and species differences within these traits on unique species contributions to functional diversity and redundancy, as well as on the overall functional diversity and redundancy within species assemblages. Our results highlighted that species richness and species functional attributes interact in their effects on functional diversity. Moreover, our simulations suggested that functional differences among species have limited effects on the proportion of redundancy of species contributions as well as on the overall redundancy within species assemblages, but that redundancy rather was determined by number of traits and species richness. Our simulations finally indicated scale dependence in the relative effects of species richness and functional attributes, which suggest that the relative influence of these factors may affect individual contributions differently compared to the overall ecosystem function of species assemblages. We suggest that studies on the relationship between biological diversity and ecosystem function will benefit from focusing on multiple processes and ecological interactions, and that the relative functional attributes of species will have pivotal roles for the ecosystem function of a given species assembly.     

Shrub encroachment effect on the evapotranspiration and its component-A numerical simulation study of a shrub encroachment grassland in Nei Mongol,China北大核心CSCD

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.     

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.     

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。     

Progressively excluding mammals of different body size affects community and trait structure of ground beetles

Mammalian grazing induces changes in vegetation properties in grasslands, which can affect a wide variety of other animals including many arthropods. However, the impacts may depend on the type and body size of these mammals. Furthermore, how mammals influence functional trait syndromes of arthropod communities is not well known. We progressively excluded large (e.g. red deer, chamois), medium (e.g. alpine marmot, mountain hare), and small (e.g. mice) mammals using size‐selective fences in two vegetation types (short‐ and tall‐grass vegetation) of subalpine grasslands. We then assessed how these exclusions affected the community composition and functional traits of ground beetles (Coleoptera, Carabidae), and which vegetation characteristic mediated the observed effects. Total carabid biomass, the activity densities of carabids with specific traits (i.e. small eyes, short wings), the richness of small‐eyed species and the richness of herbivorous species were significantly higher when certain mammals were excluded compared to when all mammals had access, regardless of vegetation type. Excluding large and medium mammals increased the activity density of herbivorous carabid species, but only in short‐grass vegetation. Similarly, excluding large mammals (ungulates) altered carabid species composition in the short‐, but not in the tall‐grass vegetation. All these responses were related to aboveground plant biomass, but not to plant Shannon diversity or vegetation structural heterogeneity. Our results indicate that changes in aboveground plant biomass are key drivers of mammalian grazers’ influence on carabids, suggesting that bottom–up forces are important in subalpine grassland systems. The exclusion of ungulates provoked the strongest carabid response. Our results, however, also highlight the ecological significance of smaller herbivorous mammals. Our study furthermore shows that mammalian grazing not only altered carabid community composition, but also caused community‐wide functional trait shifts, which could potentially have a wider impact on species interactions and ecosystem functioning.     

荒漠草原人工灌丛化对蒸散发及其组分的影响——以盐池县为例

我国西北防沙治沙工程中大量的种植中间锦鸡儿(Caragana intermedia)会导致荒漠草原发生灌丛化现象,研究人工灌丛化对荒漠草原蒸散发的影响,不仅能够揭示半干旱区人为活动对生态系统水循环的影响机理,还可以指导区域生态治理实践。以宁夏盐池县荒漠草原为例,基于植被的生理生态参数和荒漠草原水热条件,采用生物地球化学模型(Biome Bio-Geochemical Cycles,Biome-BGC)和地球呼吸系统模拟模型(Breathing Earth System Simulator,BESS)结合的方法,模拟荒漠草原生态系统人工灌丛引入前后蒸散发及其组分的变化,定量研究荒漠草原人工灌丛化对区域生态水文循环中蒸散发的影响。结果表明,人工灌丛的引入使植被结构及特征发生了变化,叶面积指数(Leaf Area Index,LAI)年最大值由0.20增加为0.67,改变了植被年内与年际变化特征。荒漠草原人工灌丛化后,生态系统年均蒸散发由251.74 mm增加到了281.42 mm;人工灌丛化对生长季的蒸散发增强明显,8月蒸散发峰值时,日均蒸散发由1.27 mm/d增加到1.56 mm/d。...     

Testing the Link between Functional Diversity and Ecosystem Functioning in a Minnesota Grassland Experiment

The functional diversity of a community can influence ecosystem functioning and reflects assembly processes. The large number of disparate metrics used to quantify functional diversity reflects the range of attributes underlying this concept, generally summarized as functional richness, functional evenness, and functional divergence. However, in practice, we know very little about which attributes drive which ecosystem functions, due to a lack of field-based tests. Here we test the association between eight leading functional diversity metrics (Rao’s Q, FD, FDis, FEve, FDiv, convex hull volume, and species and functional group richness) that emphasize different attributes of functional diversity, plus 11 extensions of these existing metrics that incorporate heterogeneous species abundances and trait variation. We assess the relationships among these metrics and compare their performances for predicting three key ecosystem functions (above- and belowground biomass and light capture) within a long-term grassland biodiversity experiment. Many metrics were highly correlated, although unique information was captured in FEve, FDiv, and dendrogram-based measures (FD) that were adjusted by abundance. FD adjusted by abundance outperformed all other metrics in predicting both above- and belowground biomass, although several others also performed well (e.g. Rao’s Q, FDis, FDiv). More generally, trait-based richness metrics and hybrid metrics incorporating multiple diversity attributes outperformed evenness metrics and single-attribute metrics, results that were not changed when combinations of metrics were explored. For light capture, species richness alone was the best predictor, suggesting that traits for canopy architecture would be necessary to improve predictions. Our study provides a comprehensive test linking different attributes of functional diversity with ecosystem function for a grassland system.     

Changes in arthropod diversity along a land use driven gradient of shrub cover in savanna rangelands: identification of suitable indicators

Shrub encroachment linked to heavy grazing has dramatically changed savanna landscapes, and is a major form of rangeland degradation. Our understanding of how shrub encroachment affects arthropod communities is poor, however. Here, we investigate the effects of shrub encroachment on abundance and diversity of ground-dwelling (wingless) arthropods at varying levels of shrub cover in the southern Kalahari. We also ascertain if invertebrate assemblage composition changes with habitat structure and identify which aspects of habitat structure (e.g., grass cover, herbaceous plant cover, shrub density) correlate most strongly with these changes. Ant, scorpion and dung beetle abundance increased with shrub cover, whereas grasshoppers and solifuges declined. Spider and beetle abundance exhibited hump-shaped relationships with shrub cover. RTU richness within orders either mirrored abundances, or exhibited no trend. Shrub density was the habitat component most correlated with similarities between invertebrate assemblages. Ground-dwelling arthropods showed clear shifts in species assemblage composition at a similarity level of 65% according to shrub density. Changes in indicator species showed that within the Tenebrionidae (darkling beetles), certain species respond positively to shrub thickening, replacing other species within the Family. Small-bodied, wingless Scarabaeidae (dung beetles) tended to increase with increased shrub density and three species emerged as significant indicators of more thickened habitats, although this might be a response to greater dung availability, rather than habitat structure itself. We conclude that because ground-dwelling invertebrates showed such clear responses in species assemblage composition, they present excellent candidates for use as indicator species in further studies into bush encroachment.     

Shrub expansion stimulates soil C and N storage along a coastal soil chronosequence

Expansion of woody vegetation in grasslands is a worldwide phenomenon with implications for C and N cycling at local, regional and global scales. Although woody encroachment is often accompanied by increased annual net primary production (ANPP) and increased inputs of litter, mesic ecosystems may become sources for C after woody encroachment because stimulation of soil CO₂ efflux releases stored soil carbon. Our objective was to determine if young, sandy soils on a barrier island became a sink for C after encroachment of the nitrogen‐fixing shrub Morella cerifera, or if associated stimulation of soil CO₂ efflux mitigated increased litterfall. We monitored variations in litterfall in shrub thickets across a chronosequence of shrub expansion and compared those data to previous measurements of ANPP in adjacent grasslands. In the final year, we quantified standing litter C and N pools in shrub thickets and soil organic matter (SOM), soil organic carbon (SOC), soil total nitrogen (TN) and soil CO₂ efflux in shrub thickets and adjacent grasslands. Heavy litterfall resulted in a dense litter layer storing an average of 809 g C m^?2 and 36 g N m^?2. Although soil CO₂ efflux was stimulated by shrub encroachment in younger soils, soil CO₂ efflux did not vary between shrub thickets and grasslands in the oldest soils and increases in CO₂ efflux in shrub thickets did not offset contributions of increased litterfall to SOC. SOC was 3.6–9.8 times higher beneath shrub thickets than in grassland soils and soil TN was 2.5–7.7 times higher under shrub thickets. Accumulation rates of soil and litter C were highest in the youngest thicket at 101 g m^?2 yr^?1 and declined with increasing thicket age. Expansion of shrubs on barrier islands, which have low levels of soil carbon and high potential for ANPP, has the potential to significantly increase ecosystem C sequestration.     

Direct and indirect facilitation affect community productivity through changes in functional diversity in an alpine system

Background and AimsFacilitation is an important ecological process for plant community structure and functional composition. Although direct facilitation has accrued most of the evidence so far, indirect facilitation is ubiquitous in nature and it has an enormous potential to explain community structuring. In this study, we assess the effect of direct and indirect facilitation on community productivity via taxonomic and functional diversity.MethodsIn an alpine community on the Tibetan Plateau, we manipulated the presence of the shrub Dasiphora fruticosa and graminoids in a fenced meadow and a grazed meadow to quantify the effects of direct and indirect facilitation. We measured four plant traits: height, lateral spread, specific leaf area (SLA) and leaf dry matter content (LDMC) of forbs; calculated two metrics of functional diversity [range of trait and community-weighted mean (CWM) of trait]; and assessed the responses of functional diversity to shrub facilitation. We used structural equation modelling to explore how shrubs directly and indirectly drove community productivity via taxonomic diversity and functional diversity.Key ResultsWe found stronger effects from herbivore-mediated indirect facilitation than direct facilitation on productivity and taxonomic diversity, regardless of the presence of graminoids. For functional diversity, the range and CWM of height and SLA, rather than lateral spread and LDMC, generally increased due to direct and indirect facilitation. Moreover, we found that the range of traits played a primary role over taxonomic diversity and CWM of traits in terms of shrub effects on community productivity.ConclusionsOur study reveals that the mechanism of shrub direct and indirect facilitation of community productivity in this alpine community is expanding the realized niche (i.e. expanding range of traits). Our findings indicate that facilitators might increase trait dispersion in the local community, which could alleviate the effect of environmental filters on trait values in harsh environments, thereby contributing to ecosystem functioning.     

Multiple trade‐offs regulate the effects of woody plant removal on biodiversity and ecosystem functions in global rangelands

Woody plant encroachment is a major land management issue. Woody removal often aims to restore the original grassy ecosystem, but few studies have assessed the role of woody removal on ecosystem functions and biodiversity at global scales. We collected data from 140 global studies and evaluated how different woody plant removal methods affected biodiversity (plant and animal diversity) and ecosystem functions (plant production, hydrological function, soil carbon) across global rangelands. Our results indicate that the impact of removal is strongly context dependent, varying with the specific response variable, removal method, and traits of the target species. Over all treatments, woody plant removal increased grass biomass and total groundstorey diversity. Physical and chemical removal methods increased grass biomass and total groundstorey biomass (i.e., non‐woody plants, including grass biomass), but burning reduced animal diversity. The impact of different treatment methods declined with time since removal, particularly for total groundstorey biomass. Removing pyramid‐shaped woody plants increased total groundstorey biomass and hydrological function but reduced total groundstorey diversity. Environmental context (e.g., aridity and soil texture) indirectly controlled the effect of removal on biomass and biodiversity by influencing plant traits such as plant shape, allelopathic, or roots types. Our study demonstrates that a one‐size‐fits‐all approach to woody plant removal is not appropriate, and that consideration of woody plant identity, removal method, and environmental context is critical for optimizing removal outcomes. Applying this knowledge is fundamental for maintaining diverse and functional rangeland ecosystems as we move toward a drier and more variable climate.