The effect of grazing on the spatial heterogeneity of vegetation

Grazing can alter the spatial heterogeneity of vegetation, influencing ecosystem processes and biodiversity. Our objective was to identify why grazing causes increases in the spatial heterogeneity of vegetation in some cases, but decreases in others. The immediate effect of grazing on heterogeneity depends on the interaction between the spatial pattern of grazing and the pre-existing spatial pattern of vegetation. Depending on the scale of observation and on the factors that determine animal distribution, grazing patterns may be stronger or weaker than vegetation patterns, or may mirror the spatial structure of vegetation. For each possible interaction between these patterns, we make a prediction about resulting changes in the spatial heterogeneity of vegetation. Case studies from the literature support our predictions, although ecosystems characterized by strong plant-soil interactions present important exceptions. While the processes by which grazing causes increases in heterogeneity are clear, how grazing leads to decreases in heterogeneity is less so. To explore how grazing can consistently dampen the fine-scale spatial patterns of competing plant species, we built a cell-based simulation model that features two competing plant species, different grazing patterns, and different sources of vegetation pattern. Only the simulations that included neighborhood interactions as a source of vegetation pattern produced results consistent with the predictions we derived from the literature review.     

Herbivores as architects of savannas: inducing and modifying spatial vegetation patterning

In this paper, we address the question whether and through which mechanisms herbivores can induce spatial patterning in savanna vegetation, and how the role of herbivory as a determinant of vegetation patterning changes with herbivore density and the pre-existing pattern of vegetation. We thereto developed a spatially explicit simulation model, including growth of grasses and trees, vertical zonation of browseable biomass, and spatially explicit foraging by grazers and browsers. We show that herbivores can induce vegetation patterning when two key assumptions are fulfilled. First, herbivores have to increase the attractiveness of a site while foraging so that they will revisit this site, e.g. through an increased availability or quality of forage. Second, foraging should be spatially explicit, e.g. when foraging at a site influences vegetation at larger spatial scales or when vegetation at larger spatial scales influences the selection and utilisation of a site. The interaction between these two assumptions proved to be crucial for herbivores to produce spatial vegetation patterns, but then only at low to intermediate herbivore densities. High herbivore densities result in homogenisation of vegetation. Furthermore, our model shows that the pre-existing spatial pattern in vegetation influences the process of vegetation patterning through herbivory. However, this influence decreases when the heterogeneity and dominant scale of the initial vegetation decreases. Hence, the level of adherence of the herbivores to forage in pre-existing patches increases when these pre-existing patches increase in size and when the level of vegetation heterogeneity increases. The findings presented in this paper, and critical experimentation of their ecological validity, will increase our understanding of vegetation patterning in savanna ecosystems, and the role of plant–herbivore interactions therein.     

Formation of banded vegetation patterns resulted from interactions between sediment deposition and vegetation growth

This research investigates the formation of banded vegetation patterns on hillslopes affected by interactions between sediment deposition and vegetation growth. The following two perspectives in the formation of these patterns are taken into consideration: (a) increased sediment deposition from plant interception, and (b) reduced plant biomass caused by sediment accumulation. A spatial model is proposed to describe how the interactions between sediment deposition and vegetation growth promote self-organization of banded vegetation patterns. Based on theoretical and numerical analyses of the proposed spatial model, vegetation bands can result from a Turing instability mechanism. The banded vegetation patterns obtained in this research resemble patterns reported in the literature. Moreover, measured by sediment dynamics, the variation of hillslope landform can be described. The model predicts how treads on hillslopes evolve with the banded patterns. Thus, we provide a quantitative interpretation for coevolution of vegetation patterns and landforms under effects of sediment redistribution.     

Spatial heterogeneity and irreversible vegetation change in semiarid grazing systems

Recent theoretical studies have shown that spatial redistribution of surface water may explain the occurrence of patterns of alternating vegetated and degraded patches in semiarid grasslands. These results implied, however, that spatial redistribution processes cannot explain the collapse of production on coarser scales observed in these systems. We present a spatially explicit vegetation model to investigate possible mechanisms explaining irreversible vegetation collapse on coarse spatial scales. The model results indicate that the dynamics of vegetation on coarse scales are determined by the interaction of two spatial feedback processes. Loss of plant cover in a certain area results in increased availability of water in remaining vegetated patches through run-on of surface water, promoting within-patch plant production. Hence, spatial redistribution of surface water creates negative feedback between reduced plant cover and increased plant growth in remaining vegetation. Reduced plant cover, however, results in focusing of herbivore grazing in the remaining vegetation. Hence, redistribution of herbivores creates positive feedback between reduced plant cover and increased losses due to grazing in remaining vegetated patches, leading to collapse of the entire vegetation. This may explain irreversible vegetation shifts in semiarid grasslands on coarse spatial scales.     

Indirect effects of cattle grazing on shrub spatial pattern in a mediterranean scrub community

Identifying the mechanisms and interactions that influence the spatial structure of vegetation is important for both scientific and practical purposes. Grazing is one of the most fundamental interactions in ecology but so far its effect on vegetation spatial pattern received little attention. In this study we propose a conceptual model that can be used to predict the effect of grazing on shrub spatial pattern in water-limited ecosystems where shrubs grow within a matrix of annual vegetation. According to the model, grazing may increase or decrease clumping in shrub distribution, depending on (1) the relative palatability of shrubs vs. annual plants to the herbivores, and (2) the manner (negative or positive) by which adult shrubs and annual plants affect the establishment of shrub seedlings. We tested our model in a Mediterranean scrub ecosystem by analyzing the development of shrub spatial pattern over a period of 40 years in plots characterized by contrasting intensities of cattle grazing. As predicted by the model, all plots showed a clumped pattern of shrub distribution in the absence of cattle grazing while intense cattle grazing reduced the clumpiness of the vegetation and generated a more random pattern of shrub distribution. Interestingly, plots representing the two grazing regimes did not differ significantly in their shrub cover, suggesting that shrub spatial pattern may be more sensitive to grazing than overall shrub cover.     

Large‐scale prerain vegetation green‐up across Africa

Information on the response of vegetation to different environmental drivers, including rainfall, forms a critical input to ecosystem models. Currently, such models are run based on parameters that, in some cases, are either assumed or lack supporting evidence (e.g., that vegetation growth across Africa is rainfall‐driven). A limited number of studies have reported that the onset of rain across Africa does not fully explain the onset of vegetation growth, for example, drawing on the observation of prerain flush effects in some parts of Africa. The spatial extent of this prerain green‐up effect, however, remains unknown, leaving a large gap in our understanding that may bias ecosystem modelling. This paper provides the most comprehensive spatial assessment to‐date of the magnitude and frequency of the different patterns of phenology response to rainfall across Africa and for different vegetation types. To define the relations between phenology and rainfall, we investigated the spatial variation in the difference, in number of days, between the start of rainy season (SRS) and start of vegetation growing season (SOS); and between the end of rainy season (ERS) and end of vegetation growing season (EOS). We reveal a much more extensive spread of prerain green‐up over Africa than previously reported, with prerain green‐up being the norm rather than the exception. We also show the relative sparsity of postrain green‐up, confined largely to the Sudano‐Sahel region. While the prerain green‐up phenomenon is well documented, its large spatial extent was not anticipated. Our results, thus, contrast with the widely held view that rainfall drives the onset and end of the vegetation growing season across Africa. Our findings point to a much more nuanced role of rainfall in Africa's vegetation growth cycle than previously thought, specifically as one of a set of several drivers, with important implications for ecosystem modelling.     

Slowing down in spatially patterned ecosystems at the brink of collapse

Predicting the risk of critical transitions, such as the collapse of a population, is important in order to direct management efforts. In any system that is close to a critical transition, recovery upon small perturbations becomes slow, a phenomenon known as critical slowing down. It has been suggested that such slowing down may be detected indirectly through an increase in spatial and temporal correlation and variance. Here, we tested this idea in arid ecosystems, where vegetation may collapse to desert as a result of increasing water limitation. We used three models that describe desertification but differ in the spatial vegetation patterns they produce. In all models, recovery rate upon perturbation decreased before vegetation collapsed. However, in one of the models, slowing down failed to translate into rising variance and correlation. This is caused by the regular self-organized vegetation patterns produced by this model. This finding implies an important limitation of variance and correlation as indicators of critical transitions. However, changes in such self-organized patterns themselves are a reliable indicator of an upcoming transition. Our results illustrate that while critical slowing down may be a universal phenomenon at critical transitions, its detection through indirect indicators may have limitations in particular systems.     

A model for small‐scale seed bank and standing vegetation connection along time

Since inter‐annual climatic variability influences composition and structure of seed bank and extant vegetation, it is expected that it also affects the relationship between both compartments at small scales along time. We hypothesize that seed bank and aboveground vegetation are closely and sequentially linked at small spatial and time scales. We conducted a field spatially explicit investigation to explore the spatial and temporal relationship between both compartments. Abundance, composition and spatial structure at different small‐scales of seed bank and aboveground vegetation were analysed through 100 permanent plots during two consecutive (dry and wet) periods of growth. Following a conceptual transition path model, we analysed changes in correlation values in composition and spatial aggregation between both compartments along time including seasonal variations of seed bank. Shape of spatial structures were evaluated using partial Mantel correlograms. Annuals and perennials guilds were studied separately. During the wet year, annuals increased their aerial spatial aggregation and cover, whereas the opposite happened for perennials. Density and spatial aggregation of the seed bank increased for both guilds in the two seasons following the rain period, especially in annuals. The clumped structure of the transient seed bank is a consequence of the addition of spatial structure of extant vegetation and the persistent seed bank. Expression of the persistent seed bank is much lower during the dry period for the annual guild. In spite of the strong yearly variability, the community maintained a highly structured spatiotemporal pattern. The mechanisms promoting this stability differed for annual and perennial guilds. Temporal persistence in perennials relied mainly in established plants longevity, whereas annual guild persistence depended solely on seed bank. This tight structure was coherent with the existence of successional dynamics in the community, although persistent seed bank could moderate the pace. Longer term studies of seed bank–standing vegetation dynamics are required to fully understand this process.     

黄土丘陵区延河流域潜在植被分布预测与制图

潜在植被的分布预测与制图对植被恢复规划具有重要的指导价值.利用广义相加模型（generalized additive model,GAM）,结合GIS空间分析技术和环境梯度分层采样技术,为延河流域24个地带性物种建立了分布模型,并在考虑群落内部物种种间关系及其分布概率的基础上,对物种分布进行运算,模拟预测了延河流域37种植物群落的分布状况和延河流域的潜在植被分布.结果表明: 研究区植被分布预测值与实际调查值间的差异不显著,预测的植被空间分布较好地反映了延河流域潜在的植被分布状况,表明该模型具有较好的预测能力,对于区域植被恢复的目标设定和恢复规划具有重要意义.     

Brain expansion in early hominins predicts carnivore extinctions in East Africa

While the anthropogenic impact on ecosystems today is evident, it remains unclear if the detrimental effect of hominins on co‐occurring biodiversity is a recent phenomenon or has also been the pattern for earlier hominin species. We test this using the East African carnivore fossil record. We analyse the diversity of carnivores over the last four million years and investigate whether any decline is related to an increase in hominin cognitive capacity, vegetation changes or climatic changes. We find that extinction rates in large carnivores correlate with increased hominin brain size and with vegetation changes, but not with precipitation or temperature changes. While temporal analyses cannot distinguish between the effects of vegetation changes and hominins, we show through spatial analyses of contemporary carnivores in Africa that only hominin causation is plausible. Our results suggest that substantial anthropogenic influence on biodiversity started millions of years earlier than currently assumed.     

Application of geographically weighted regression in estimating the effect of climate and site conditions on vegetation distribution in Haihe Catchment,China

Climate and topography are the two key factors influencing vegetation pattern, distribution, and plant growth. Traditionally, studies on the relationship between vegetation and climate rely largely on field data from limited samples. Now, digital elevation model (DEM) and remote sensing data readily provide huge amounts of spatial data on site-specific conditions like elevation, aspect, and climate, while recent development of geographically weighted regression (GWR) analysis facilitates efficient spatial evaluation of interactions among vegetation and site conditions. Using Haihe Catchment as a case study, GWR is applied in establishing spatial relations among leaf area index (LAI; a critical vegetation index from Moderate Resolution Imaging Spectroradiometer (MODIS)) and interpolated climate variables and site conditions including elevation, aspect, and Topographic Wetness Index (TWI). This study suggests that the GWR solution to spatial effect of climate and site conditions on vegetation is much better than ordinary least squares (OLS). In most of the study area, effects of elevation, aspect change from south to north, and precipitation on LAI are positive, while temperature, TWI, and potential evapotranspiration have a negative influence. Spatially, models perform better in places with large spatial variations in LAI—primarily driven by strong spatial variations in temperature and precipitation. On the contrary, the effect of topographic and climatic factors on vegetation is weak in regions with small spatial variations in LAI. This study shows that overall water availability is a determining factor for spatial variations in vegetation.     

The effect of spatial scale on evenness

Abstract. The effect of spatial scale on species evenness has not previously been investigated. As the area of each sample of vegetation (i.e. the spatial grain) increases, evenness could in theory increase, decrease, or stay the same, though the simplest model predicts an increase. We use biomass data from four dune slack sites and two semi-arid grasslands, sampled to allow calculation of evenness at a range of spatial grains. In all six sites, evenness decreases as grain size increases, almost monotonically. It is hypothesized that such a pattern is a result of a general feature of plant species abundance distributions and of vegetation response to environmental microheterogeneity.     

Complexity and coexistence in a simple spatial model for arid savanna ecosystems

Tree–grass coexistence is broadly observed in tropical savannas. Recent studies indicate that, in arid savannas, such coexistence is stable and related to water availability. The role of different factors (from niche separation to demographic structure) has been explored. Nevertheless, spatial mechanisms of water–vegetation interactions have been rarely taken into account, despite their well-known importance for vegetation distribution. Here, we introduce a spatial model including tree and grass biomass dynamics, together with soil and surface water dynamics. We consider two water–vegetation feedbacks. Grasses increase water infiltration into the soil, while tree shadow limits evaporation, and both mechanisms increase soil water availability, leading to positive feedbacks. The infiltration feedback can also lead to spatial pattern formation. Despite the fact that trees and grasses compete for the same resource, namely water, we observe stable coexistence as a possible model outcome. The system displays a complex behavior, with multiple stable states and possible catastrophic shifts between states, e.g., patterned grassland, bare soil and forest. In our model, coexistence is always linked with multi-stability and spatial pattern formation, driven by grass infiltration feedback. Given such complex model solutions, we expect that, under real conditions, heterogeneities and disturbances, acting on the multi-stable states, may further foster coexistence.     

遥感反演植被理化参数的光谱和空间尺度效应

植被理化参数是生态系统中碳和养分等物质循环与能量交换的重要指标,利用遥感技术反演是获取区域及全球植被理化参数的重要手段,但光谱和空间尺度效应的存在,限制了源自不同遥感传感器植被理化参数产品的统一应用。阐述了遥感反演植被理化参数光谱尺度效应的概念及其产生原因,主要从光谱波段位置和波段宽度两方面对国内外相关研究进行了介绍和评述。同时,从遥感反演植被理化参数的空间尺度效应产生原因、空间异质性描述方法和空间尺度转换方法等方面对其国内外研究现状进行了归纳和评述。最后,总结了遥感反演植被理化参数光谱和空间尺度效应研究的不足之处和发展趋势,并指出光谱和空间耦合效应的研究将是一大趋势,而在生态学等领域形成的尺度效应研究的理论和方法也值得借鉴参考。     

Short range co-operativity competing with long range inhibition explains vegetation patterns

We model the non-local dynamics of vegetation communities and interpret the formation of vegetation patterns as a spatial instability of intrinsic origin: the wavelength of the patterns predicted within the framework of this approach is determined by the parameters governing the dynamics rather than by boundary conditions and/or geometrical constraints. The spatial periodicity results from an interplay between short-range co-operative interactions and long-range self-inhibitory interactions inside the vegetation community. The influence of environmental anisotropies on pattern symmetry and orientation is discussed. As a case study, the approach is applied to a system of vegetation bands situated in the north-west of Burkina Faso. The parameters describing the co-operative and inhibitory interactions at the origin of the patterns are evaluated.     

内蒙古荒漠草原植被盖度的空间异质性动态分析

利用半方差函数分析法对内蒙古荒漠草原生长盛期(6—8月)的植被盖度时空变异特征的研究表明,荒漠草原生长盛期的植被盖度半方差函数形态符合指数模型,但函数曲线的形态和各参数在不同月份变化较大。其中,6月的植被盖度变程最大,达到100 m;7月植被盖度的半方差函数形态具有巢式等级结构;8月植被盖度的变程最小,仅为15 m,但空间变异程度最高。3个月的结构比介于72%—85%,具有较强的空间自相关。各向异性分析表明,6月植被盖度在135°方向的半方差函数值明显低于其它3个方向(0°、45°、90°),具有各向异性特征;而7月和8月植被盖度的各向异性比接近于1,表现为各向同性。研究结果表明,荒漠草原植被盖度空间异质性的时间动态不容忽视,在野外采样或制图时,要根据时间合理控制采样范围。     

Patterns of spatial autocorrelation of assemblages of birds, floristics, physiognomy, and primary productivity in the central Great Basin, USA

We fitted spatial autocorrelation functions to distance-based data for assemblages of birds and for three attributes of birds' habitats at 140 locations, separated by up to 65 km, in the Great Basin (Nevada, USA). The three habitat characteristics were taxonomic composition of the vegetation, physical structure of the vegetation, and a measure of primary productivity, the normalized difference vegetation index, estimated from satellite imagery. We found that a spherical model was the best fit to data for avifaunal composition, vegetation composition, and primary productivity, but the distance at which spatial correlation effectively was zero differed substantially among data sets ( c . 30 km for birds, 20 km for vegetation composition, and 60 km for primary productivity). A power-law function was the best fit to data for vegetation structure, indicating that the structure of vegetation differed by similar amounts irrespective of distance between locations (up to the maximum distance measured). Our results suggested that the spatial structure of bird assemblages is more similar to vegetation composition than to either vegetation structure or primary productivity, but is autocorrelated over larger distances. We believe that the greater mobility of birds compared with plants may be responsible for this difference.     

Spatial extent of bird species response to landscape changes: colonisation/extinction dynamics at the community-level in two contrasting habitats

Animal community dynamics in changing landscapes are primarily driven by changes in vegetation structure and ultimately by how species respond to these changes and at which spatial scale. We consider two major components of local community dynamics, species colonisation and extinction. We hypothesise that (1) the optimal spatial extent needed to accurately predict them will differ between these two processes; (2) it will also likely differ from species to species as a result of life history traits differences related to differences in habitat selection and (3) that a species' primary habitat will determine the spatial extent at which it perceives change in vegetation structure. We used data collected over 25 yr in a changing Mediterranean landscape to study bird species local colonisation and extinction patterns in two groups of species typical from two habitats: open farmland and woodland. Vegetation changes were measured at spatial extents ranging from 0.2 to 79 ha. Local species colonisation and extinction estimates were computed using a method accounting for heterogeneity in detection probability among species. We built linear models between local species colonisation/extinction estimates and vegetation changes and examined variations in model quality with respect to the spatial extent at which vegetation changes had been measured. Models for open habitat species showed that colonisation processes operated at the landscape scale (79 ha), while extinction was more tightly linked to local habitat requirements (0.2 ha). Models for woodland species presented a low and constant model quality whatever the spatial extent considered. Our results suggest that the dynamics of the woodland species considered responded to a combination of vegetation changes at several scales and, in particular, to changes in the vertical structure of the vegetation. We highlight the need to explicitly consider spatial extent in studies of habitat selection and of habitat and population dynamics to improve our understanding of the biological consequences of land use changes and guide more effective conservation efforts.     

Scale matters: fire–vegetation feedbacks are needed to explain tropical tree cover at the local scale

At a broad (regional to global) spatial scale, tropical vegetation is controlled by climate; at the local scale, it is believed to be determined by interactions between disturbance, vegetation and local conditions (soil and topography) through feedback processes. It has recently been suggested that strong fire–vegetation feedback processes may not be needed to explain tree‐cover patterns in tropical ecosystems and that climate–fire determinism is an alternative possibility. This conclusion was based on the fact that it is possible to reproduce observed patterns in tropical regions (e.g. a trimodal frequency distribution of tree cover) using a simple model that does not explicitly incorporate fire–vegetation feedback processes. We argue that these two mechanisms (feedbacks versus fire–climate control) operate at different spatial and temporal scales; it is not possible to evaluate the role of a process acting at fine scales (e.g. fire–vegetation feedbacks) using a model designed to reproduce regional‐scale pattern (scale mismatch). While the distributions of forest and savannas are partially determined by climate, many studies are providing evidence that the most parsimonious explanation for their environmental overlaps is the existence of feedback processes. Climate is unlikely to be an alternative to feedback processes; rather, climate and fire–vegetation feedbacks are complementary processes at different spatial and temporal scales.     

Vegetation Pattern Formation Due to Interactions Between Water Availability and Toxicity in Plant–Soil Feedback

Development of a comprehensive theory of the formation of vegetation patterns is still in progress. A prevailing view is to treat water availability as the main causal factor for the emergence of vegetation patterns. While successful in capturing the occurrence of multiple vegetation patterns in arid and semiarid regions, this hypothesis fails to explain the presence of vegetation patterns in humid environments. We explore the rich structure of a toxicity-mediated model of the vegetation pattern formation. This model consists of three PDEs accounting for a dynamic balance between biomass, water, and toxic compounds. Different (ecologically feasible) regions of the model’s parameter space give rise to stable spatial vegetation patterns in Turing and non-Turing regimes. Strong negative feedback gives rise to dynamic spatial patterns that continuously move in space while retaining their stable topology.