Crust Composition and Disturbance Drive Infiltration Through Biological Soil Crusts in Semiarid Ecosystems

Soil crusts influence many soil parameters that affect how water moves into and through the soil, and therefore, critically influence water availability, erosion processes, nutrient fluxes, and vegetation distribution patterns in semiarid ecosystems. Soil crusts are quite sensitive to disturbance, and their alteration can lead to modification of the local hydrological regime, thus affecting general functioning of the ecosystem. The aim of this study was to analyze the influence of different types of soil crusts, physical, and biological in different developmental stages, as well as the impact of their disturbance, on infiltration. This was assessed by means of rainfall simulations conducted in two semiarid ecosystems in southeast Spain characterized by different lithologies, topographies, and soil crust distributions. Two consecutive rainfall simulation experiments (50 mm h⁻¹ rainfall intensity), the first on dry soil and the second on wet soil, were carried out in microplots (0.25 m²) containing the most representative soil crust types at each site, each crust type subjected to three disturbance treatments: (a) undisturbed, (b) trampling, and (c) removal. Infiltration in the crusts was higher on coarse- than on fine-textured soils and almost two times greater on dry than on wet soil. Biological soil crusts (BSC) showed higher infiltration rates than physical soil crusts (PSC). Within BSC, infiltration increased as cyanobacterial biomass increased and was the highest in moss crusts. However, late-successional crustose and squamulose lichen crusts showed very low infiltration rates. Trampling reduced infiltration rates, especially when soil was wet, whereas crust removal enhanced infiltration. But this increase in infiltration after removing the crust decreased over time as the soil sealed again due to raindrop impact, making runoff rates in the scraped microplots approach those registered in the respective undisturbed crust types. Our results demonstrate that water redistribution in semiarid ecosystems strongly depends on the type of crusts that occupy the interplant spaces and the characteristics of the soils which they overly, as well as the antecedent moisture conditions of the soil. Disturbance of these crust patches results in increased runoff and erosion, which has important consequences on general ecosystem functioning.     

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.     

Spatial heterogeneity and plant species richness at different spatial scales under rabbit grazing

Herbivores influence spatial heterogeneity in soil resources and vegetation in ecosystems. Despite increasing recognition that spatial heterogeneity can drive species richness at different spatial scales, few studies have quantified the effect of grazing on spatial heterogeneity and species richness simultaneously. Here we document both these variables in a rabbit-grazed grassland. We measured mean values and spatial patterns of grazing intensity, rabbit droppings, plant height, plant biomass, soil water content, ammonia and nitrate in sites grazed by rabbits and in matched, ungrazed exclosures in a grassland in southern England. Plant species richness was recorded at spatial scales ranging between 0.0001 and 150 m(2). Grazing reduced plant height and plant biomass but increased levels of ammonia and nitrate in the soil. Spatial statistics revealed that rabbit-grazed sites consisted of a mixture of heavily grazed patches with low vegetation and nutrient-rich soils (lawns) surrounded by patches of high vegetation with nutrient-poor soils (tussocks). The mean patch size (range) in the grazed controls was 2.1 +/- 0.3 m for vegetation height, 3.8 +/- 1.8 m for soil water content and 2.8 +/- 0.9 m for ammonia. This is in line with the patch sizes of grazing (2.4 +/- 0.5 m) and dropping deposition (3.7 +/- 0.6 m) by rabbits. In contrast, patchiness in the ungrazed exclosures had a larger patch size and was not present for all variables. Rabbit grazing increased plant species richness at all spatial scales. Species richness was negatively correlated with plant height, but positively correlated to the coefficient of variation of plant height at all plot sizes. Species richness in large plots (<25 m(2)) was also correlated to patch size. This study indicates that the abundance of strong competitors and the nutrient availability in the soil, as well as the heterogeneity and spatial pattern of these factors may influence species richness, but the importance of these factors can differ across spatial scales.     

Interactive Effects of Three Ecosystem Engineers on Infiltration in a Semi-Arid Mediterranean Grassland

The redistribution of water in semi-arid environments is critical for the maintenance and survival of vegetation patches. We used a systems approach to examine the interactive effects of three engineers—Stipa tenacissima, biological soil crusts, and the European rabbit (Oryctolagus cuniculus)—on infiltration processes in a model gypseous semi-arid Mediterranean grassland. We measured the early (sorptivity) and later (steady-state infiltration) stages of infiltration at two supply potentials using disk permeameters, which allowed us to determine the relative effects of different engineers and soil micropores on water flow through large macropores. We detected few effects under tension when flow was restricted to matrix pores, but under ponding, sorptivity and steady-state infiltration adjacent to Stipa tussocks were 2–3 times higher than in intact or rabbit-disturbed biological soil crusts. Structural Equation Modeling (SEM) showed that both Stipa and biological soil crust cover exerted substantial and equal positive effects on infiltration under ponding, whereas indirectly, rabbit disturbance negatively affected infiltration by reducing crust cover. Under tension, when macropores were prevented from conducting water, Stipa had a direct negative effect and biological soil crust cover was relatively unimportant. More complex SEM models demonstrated that (1) Stipa primarily influenced biological soil crusts by reducing their richness, (2) rabbits exerted a small negative effect on crust richness, and (3) lichens were negatively, and mosses positively, correlated with a derived “infiltration” axis. Our results highlight the importance of biological soil crusts as key players in the maintenance of infiltration processes in Stipa grasslands, and demonstrate the modulating role played by rabbits through their surface disturbances.     

Numerical investigation of spatial pattern in a vegetation model with feedback function

The vegetative cover in semi-arid lands typically occurs as patches of individual species more or less separated from one another by bare ground. Klausmeier [1999. Regular and irregular patterns in semiarid vegetation. Science 284 (5421), 1826-1828] reported that the vegetation striped patterns can grow lying along the contours of gentle slopes. He has proposed a model of vegetation stripes based on competition for water. In this paper, our main aim is to study the positive feedback effects between the water and biomass on the vegetation spatial pattern formation within a nonsaturated soil, which arises from the suction of water by the roots and processes of water resource redistribution. According to the dispersion relation formula, we discuss the changes of the wavelength, wave speed, as well as the conditions of the spatial pattern formation. Our numerical results show that trees are more sensitive than grasses to the positive feedback function to format the spatial heterogenous pattern, and the stronger positive feedback increases the parameters region where vegetation bands occur, which indicates that the positive feedback raises the possibility of shift from green to desert states in semi-arid areas for the long term. Our numerical results also show that the positive feedback can increase the migration velocity of the vegetation stripes.     

The Importance of Coprophagous Macrodetritivores for the Maintenance of Vegetation Heterogeneity in an African Savannah

Grazing ecosystems are often characterized by dynamic vegetation structure mosaics of short grazing lawns and tall grass vegetation that are important for the biodiversity and functioning of these ecosystems. Grazing-induced trampling, causing soil compaction and reduced water infiltration, has been shown to be an important mechanism for lawn grass formation. However, insights in reverse bioturbation mechanisms were mostly lacking, especially how tall vegetation persists under continuous grazing by herbivores. In this study, we explore if defecation by large herbivores in combination with different groups of coprophagous macrodetritivores can locally convert compacted grazing lawn patches back to tall bunch grasslands with a more loose soil. Across a rainfall gradient in an African savannah, we separated the potential roles in this process between dung beetles versus earthworms and termites. We placed different mesh sizes under dung piles and studied the consequences for soil, vegetation, and hydrological properties. We found that soil water infiltration rate, soil organic matter content, electrical conductivity, bunch grass cover, and bunch grass biomass were significantly promoted by dung addition, irrespective of position along the rainfall gradient. In addition, the presence of tunneling dung beetles significantly increased water infiltration rate and biomass of bunch grasses, pointing at a new mechanism whereby macrodetritivores affect the structure and diversity of plant communities. We conclude that coprophagous macrodetritivores interact with large herbivores in contributing to the maintenance of structural heterogeneity in the vegetation of grazing ecosystems, with a special role played by soil-tunneling dung beetles.     

Grazing changes topography-controlled topsoil properties and their interaction on different spatial scales in a semi-arid grassland of Inner Mongolia, P.R. China

Semiarid steppe ecosystems account for large terrestrial areas and are considered as large carbon (C) sinks. However, fundamental information on topsoil sensitivity to grazing is lacking across different spatial scales including the effects of topography. Our interdisciplinary approach considering soil chemical, physical, and vegetation properties included investigations on pit scale (square-metre scale), plot scale (hectare scale), and the scale of a landscape section (several hectares). Five different sites, representing a grazing intensity gradient, ranging from a long-term grazing exclosure to a heavily grazed site were used. On the pit scale, data about aggregate size distribution, quantity of different soil organic carbon (SOC) pools, SOC mineralisation, hydraulic conductivity and shear strength was available for topsoil samples from representative soil profiles. Spatial variability of topographical parameters, topsoil texture, bulk density, SOC, water repellency, and vegetation cover was analysed on the basis of regular, orthogonal grids in differently grazed treatments by using two different grid sizes on the plot scale and landscape section. On the pit scale, intensive grazing clearly decreased soil aggregation and the amount of fresh, litter-like particulate organic matter (POM). The weak aggregation in combination with animal trampling led to an enhanced mineralisation of SOC, higher topsoil bulk densities, lower infiltration rates, and subsequently to a higher risk of soil erosion. On the plot scale, the effects of soil structure disruption due to grazing are enhanced by the degradation of vegetation patches and resulted in a texture-controlled wettability of the soil surface. In contrast, topsoils of grazing exclosures were characterised by advantageous mechanical topsoil characteristics and SOC-controlled wettability due to higher POM contents. A combined geostatistical and General Linear Model approach identified topography as the fundamental factor creating the spatial distribution of texture fractions and related soil parameters on the scale of a landscape section. Grazing strongly interfered with the topography-controlled particle relocation processes in the landscape and showed strongest effects on the aboveground biomass production and biomass-related soil properties like SOC stocks. We conclude that interdisciplinary multi-scale analyses are essential (i) to differentiate between topography- and grazing-controlled spatial patterns of topsoil and vegetation properties, and (ii) to identify the main grazing-sensitive processes on small scales that are interacting with the spatial distribution and relocation processes on larger scales.     

Plant life on gypsum: a review of its multiple facets

The adaptation of plants to particular soil types has long intrigued biologists. Gypsum soils occupy large areas in many regions of the world and host a striking biological diversity, but their vegetation has been much less studied than that developing over serpentine or saline soils. Herein, we review all aspects of plant life on gypsum ecosystems, discuss the main processes driving their structure and functioning, and highlight the main conservation threats that they face. Plant communities in gypsum habitats typically show distinctive bands at very small spatial scales, which are mainly determined by topography. Plants living on gypsum soils can be classified into three categories: (i) wide gypsophiles are specialists that can penetrate the physical soil crust during early life stages and have physiological adjustments to cope with the chemical limitations imposed by gypsum soils; (ii) narrow gypsophiles are refugee plants which successfully deal with the physical soil crust and can tolerate these chemical limitations but do not show specific adaptations for this type of soils; and (iii) gypsovags are non‐specialist gypsum plants that can only thrive in gypsum soils when the physical crust is absent or reduced. Their ability to survive in gypsum soils may also be mediated by below‐ground interactions with soil microorganisms. Gypsophiles and gypsovags show efficient germination at low temperatures, seed and fruit heteromorphism within and among populations, and variation in seed dormancy among plants and populations. In gypsum ecosystems, spatio‐temporal changes in the composition and structure of above‐ground vegetation are closely related to those of the soil seed bank. Biological soil crusts (BSCs) dominated by cyanobacteria, lichens and mosses are conspicuous in gypsum environments worldwide, and are important drivers of ecosystem processes such as carbon and nitrogen cycling, water infiltration and run‐off and soil stability. These organisms are also important determinants of the structure of annual plant communities living on gypsum soils. The short‐distance seed dispersal of gypsophiles is responsible for the high number of very narrow endemisms typically found in gypsum outcrops, and suggests that these species are evolutionarily old taxa due to the time they need to colonize isolated gypsum outcrops by chance. Climate change and habitat fragmentation negatively affect both plants and BSCs in gypsum habitats, and are among the major threats to these ecosystems. Gypsum habitats and specialists offer the chance to advance our knowledge on restrictive soils, and are ideal models not only to test important evolutionary questions such as tolerance to low Ca/Mg proportions in soils, but also to improve the theoretical framework of community ecology and ecosystem functioning.     

Recovery of some surface soil properties of ecological interest after sheep grazing in a semi-arid woodland

Solodic soils within a semi-arid poplar box (Eucalyptus populnea) woodland at Wycanna, in southern Queensland, have been subjected to impacts from sheep and cattle grazing for at least 100 years. The micromorphology of the surface of two soils, a gradational texture profile (Gn 3.13, Paleustalf) and a duplex profile (Db 1.23, Paleustalf) showed that compaction had occurred. Recovery of the soil surfaces following removal of animal grazing was measured in terms of porosity, presence of illuvial layers, surface soil strength, some chemical properties and water infiltration rates. The grazing impact was greatest on the Gn 3.13 soil and visual signs of the grazing impact were still evident 16 years following sheep removal. Reduced soil organic matter and increased surface soil hardness as a result of grazing, rather than surface crusting or changes in water infiltration rates are suggested as the mechanisms controlling the observed increases in woody plant populations in these semi-arid woodlands.     

Changes in semi-arid plant species associations along a livestock grazing gradient

In semi-arid ecosystems, vegetation is heterogeneously distributed, with plant species often associating in patches. These associations between species are not constant, but depend on the particular response of each species to environmental factors. Here, we investigated how plant species associations change in response to livestock grazing in a semi-arid ecosystem, Cabo de Gata-Níjar Natural Park in South East Spain. We established linear point-intercept transects at four sites with different grazing intensity, and recorded all species at each point. We investigated plant associations by comparing the number of times that each pair of species occurred at the same spatial point (co-occurrences), with the expected number of times based on species abundances. We also assessed associations for each shrub and grass species by considering all their pairs of associations and for the whole plant community by considering all pairs of associations on each site. At all sites, the plant community had a negative pattern of association, with fewer co-occurrences than expected. Negative association in the plant community increased at maximum grazing intensity. Most species associated as expected, particularly grass species, and positive associations were most important at intermediate grazing intensities. No species changed its type of association along the grazing gradient. We conclude that in the present plant community, grazing-resistant species compete among themselves and segregate in space. Some shrub species act as refuges for grazing-sensitive species that benefit from being spatially associated with shrub species, particularly at intermediate grazing intensities where positive associations were highest. At high grazing intensity, these shrubs can no longer persist and positive associations decrease due to the disappearance of refuges. Spatial associations between plant species and their response to grazing help identify the factors that organize plant communities, and may contribute to improving management of semi-arid ecosystems.     

Composition and diversity of herbaceous patches in woody vegetation: The effects of grazing,soil seed bank,patch spatial properties and scale

Herbaceous plants contribute much to plant diversity in Mediterranean-type ecosystems though mostly occupying relatively small patches within the dense woody vegetation. While studying species diversity in the herbaceous patches, we hypothesized that grazing, soil seed bank, and spatial properties of the patch affect plant diversity and composition at different spatial scales. The study site was in an LTER site located in the Mediterranean region in north Israel. We determined herbaceous species composition in: (1) randomly sampled quadrats in herbaceous patches in grazed and un-grazed plots; (2) soil seed bank samples taken from the same patches and germinated under optimal greenhouse conditions; (3) quadrats in the same patches sown with a homogenous mixture of local soil samples. Using GIS methods, we determined small-scale spatial characteristics of the herbaceous patches. Alpha and beta diversities were calculated at the patch and plot scales using Shannon's entropy H. Grazing increased alpha diversity of local untreated seed bank samples but decreased alpha diversity of the artificial homogenous soil seed bank mixture at both patch and plot scales. Positive relation between alpha diversity and patch area was detected only under grazing. Grazing increased beta diversity in all three treatments at the patch scale. Grazing decreased the similarity in species composition between above-ground vegetation and soil seed bank. The results indicate that moderate cattle-grazing affects species diversity in the herbaceous patches within the dense maquis. This effect is scale-dependent, and interacts with the effects of soil seed bank and patch spatial-properties: without grazing soil seed bank plays a more important role than patch spatial properties, but under grazing the size and the accessibility of the patch are more important in the determination of herbaceous species composition.     

荒漠结皮层藓类植物死亡对表层土壤水分蒸发和入渗的影响

水分是荒漠植物生长最主要的限制因子,藓类结皮作为荒漠土壤表层重要覆被物,对土壤水分蒸发入渗具有重要影响。研究表明,在全球气候变化背景下,不确定的降水格局变化导致结皮层藓类植物出现集群死亡现象,但这一过程对荒漠地表土壤水分蒸发与入渗过程的影响及其机理尚不清楚。以古尔班通古特沙漠齿肋赤藓结皮为研究对象,利用便携式渗透计和蒸发仪,研究了结皮层藓类植物死亡对土壤水分蒸发与入渗的影响。结果表明,与裸沙相比,藓类结皮的存在显著抑制了水分入渗,而藓类植物死亡的结皮层抑制作用最大,其初渗速率、稳渗速率和累积入渗量分别是活藓类结皮的39.89%、85.91%及64.48%,仅为裸沙的5.96%、13.13%及20.42%。在水分蒸发初期,裸沙的水分蒸发速率明显高于活藓类结皮和藓类植物死亡的结皮层,但藓类植物死亡的结皮层维持相对稳定的蒸发速率的时间长于裸沙和活藓类结皮,这也导致最终累计蒸发量以藓类植物死亡的结皮层最高、裸沙最低。可见,荒漠生物土壤结皮中藓类植物死亡会明显减少土壤水分入渗、增大水分蒸发,进一步影响荒漠表层土壤水分格局,从而影响生物土壤结皮与维管植物的水分利用关系。     

Spatial patterns of species distributions in grazed subalpine grasslands

Spatial patterns of species diversity have important influences on the functioning of ecosystems, and the effect of livestock grazing on spatial heterogeneity can differ depending on the scale of the analysis. This study examined the effects of grazing on the spatial patterns of species distributions and whether the effects of grazing on the spatial distributions of a species vary with the scale of the analysis. Data were collected at three locations in the subalpine grasslands of Ordesa-Monte Perdido National Park and Aísa Valley, Central Pyrenees, Spain, which differed in mean stocking rates. Aspect explained about one-third of the environmental variation in species distributions. In flat areas, spatial variation in species composition varied with grazing intensity at two scales. At a coarse scale (among vegetation patches), grazing promoted patchiness, and among-transect variation in species diversity and grazing intensity were positively correlated. At a fine scale (within vegetation patches), the disruption of the self-organizing processes of the species spatial distributions resulted in a reduction in the long-range spatial autocorrelations of some of the characteristic species and in the homogenization of species spatial distributions. The presence of encroaching Echinospartum horridum had a significant influence on the effect of grazing on south-facing grassland slopes.     

Diversity and Abundance of Biological Soil Crust Taxa in Relation to Fine and Coarse-Scale Disturbances in a Grassy Eucalypt Woodland in Eastern Australia

Over the last 200 years the box woodlands of eastern Australia have been considerably altered by European farming practices. These changes have been accompanied by a reduction in the size and number of patches of vegetation as well as the quality of the understorey vegetation and underlying soil surface. We measured diversity and abundance of soil crust taxa in relation to habitat complexity, remnant area and width, diversity of vascular plants as well as the number, size and separation of patches of vegetation and grass butts (coarse-scale patchiness), and an index of surface stability derived from measures of seven soil surface features of small microsites (patches of bare/crusted, litter- or grass-covered soil; micro-scale) on both coarse- and fine-textured soils at 35 sites in south-eastern Australia. Fifty taxa were recorded from the 35 sites, and there were more taxa from sites with fine-textured soils (12.7) compared with coarse-textured soils (4.4). The soil crust community was dominated by a few relatively common species, with many species occurring at only a few sites. Half the number of species accounting for <1% of total abundance. Bare and crusted microsites supported more species and greater cover compared with grassy microsites. Crust diversity declined with increasing coarse-level disturbances (i.e. declines in habitat complexity, remnant area and width, and diversity of vascular plants) but the results were not consistent between soil types. No measures of fine-scale disturbance were related significantly to any of the crust diversity or abundance measures, and there was no evidence of a recent grazing effect on crust composition. The fact that few sites had many species (and visa versa) suggests to us that many sites are probably required to conserve soil crust taxa in these highly fragmented landscapes     

Spatial and temporal variation of soil moisture in dependence of multiple environmental parameters in semi-arid grasslands

Grazing of grasslands changes soil physical and chemical properties as well as vegetation characteristics, such as vegetation cover, species composition and biomass production. In consequence, nutrient allocation and water storage in the top soil are affected. Land use and management changes alter these processes. Knowledge on the impacts of grazing management on nutrient and water fluxes is necessary because of the global importance of grasslands for carbon sequestration. Soil water in semi-arid areas is a limiting factor for matter fluxes and the intrinsic interaction between soil, vegetation and atmosphere. It is therefore desirable to understand the effects of grazing management and stocking rate on the spatial and temporal distribution of soil moisture. In the present study, we address the question how spatio-temporal soil moisture distribution on grazed and ungrazed grassland sites is affected by soil and vegetation properties. The study took place in the Xilin river catchment in Inner Mongolia (PR China). It is a semi-arid steppe environment, which is characterized by still moderate grazing compared to other regions in central Inner Mongolia. However, stocking rates have locally increased and resulted in a degradation of soils and vegetation also in the upper Xilin River basin. We used a multivariate geostatistical approach to reveal spatial dependencies between soil moisture distribution and soil or vegetation parameters. Overall, 7 soil and vegetation parameters (bulk density, sand, silt and clay content, mean weight diameter, mean carbon content of the soil, vegetation cover) and 57 soil moisture data sets were recorded on 100 gridded points on four sites subject to different grazing intensities. Increasing stocking rates accelerated the influence of soil and vegetation parameters on soil moisture. However, the correlation was rather weak, except for a site with high stocking rate where higher correlations were found. Low nugget ratios indicate spatial dependency between soil or plant parameters and soil moisture on a long-term ungrazed site. However, the effect was not found for a second ungrazed site that had been excluded from grazing for a shorter period. Furthermore the most important soil and vegetation parameters for predicting soil moisture distribution varied between different grazing intensities. Therefore, predicting soil moisture by using secondary variables requires a careful selection of the soil or vegetation parameters.     

Seed bank spatial structure in semi-arid environments: beyond the patch-bare area dichotomy

The prevalence of patchy structures in vegetation is a common feature in semi-arid ecosystems. Although the effect of patches on seed density is widely known, we still lack information on how patch features affect seed bank density and composition. Our aim was to answer two basic questions: (1) How do seed bank density and composition vary within and outside patch aboveground physical limits? and (2) Do patch characteristics affect soil seed bank density and composition? We sampled 50 shrub patches in a semi-arid gypsum system in Central Spain, measuring patch size, composition and structure, and seed bank at three locations per shrub (centre, edge and outside). We calculated the effect of interior patch location, patch composition and structure on seed density and composition. Patches acted both as seed sources, increasing seed density in neighbouring areas and as seed sinks by trapping seeds from bare areas. Patch structure (erect perennial cover) had the greatest effect on seed bank density, whereas patch size and microslope had the greatest influence on bare area density. Patch structure, composition and interior location explained the variation in seed bank composition. Patch effect extends to the surrounding bare matrix creating a seed bank gradient in density and composition. This effect is modulated by patch structure and composition and affects seed bank composition. Our results suggest that the spatial structure of gypsum community seed banks may act as a mechanism for a spatial storage effect contributing to the maintenance of high levels of diversity in semi-arid environments     

Grazing and productivity alter individual grass size dynamics in semi-arid woodlands

The spatial arrangement of perennial vegetation is critical for ecosystem function in drylands. While much is known about how vegetation patches respond to grazing and abiotic conditions, the size dynamics of individual plants is mostly limited to theoretical studies. We measured the size distribution (mean, variance, skewness) and density of individual grasses, and grass species composition at 451 sites spanning a range of grazing intensities across three broad vegetation communities in semi-arid eastern Australia. We assessed the relative role of grazing by livestock (cattle and sheep), native (kangaroos) and introduced (rabbits) free ranging herbivores, and several environmental measures (productivity, diversity, composition and groundstorey plant cover) on the size distribution and density of individual grasses. We found mean grass size and density were more sensitive to shifts in grazing intensity and environmental conditions than size variance or the frequency of the smallest individuals (skewness), and shifts were mostly driven by site productivity and cattle and kangaroo grazing. Sheep grazing only reduced mean grass size, and rabbit grazing had no consistent effects. Importantly, we found that site productivity and species composition altered the impacts of grazing on grass density and size distribution. For example, increasing cattle grazing led to larger grasses in low productivity sites. It also led to larger, denser, more variable-sized grasses among grass species from sites with finer soil texture. Increasing kangaroo grazing led to smaller, denser individuals among grass species from sites with coarse soil texture. At high diversity sites kangaroo grazing led to denser, more homogenised grass sizes with a lower frequency of small individuals. Understanding the in situ response of individual plant sizes gives us insights into the processes driving shifts in perennial vegetation patchiness, improving our ability to predict how the spatial arrangement of ecosystems might change under global change scenarios.     

Spatial patterns of surface soil properties and vegetation in a Mediterranean semi-arid steppe

In arid and semi-arid areas with sparse vegetation cover, the spatial pattern of surface soil properties affects water and nutrient flows, and is a question of considerable interest for understanding degradation processes and establishing adequate management measures. In this study, we investigate the spatial distribution of vegetation and surface soil properties (biological crusts, physical crusts, mosses, rock fragments, earthworm casts, fine root accumulation and below-ground stones) in a semi-arid Stipa tenacissima L. steppe in SE Spain. We applied the combination of spatial analysis by distance indices (SADIE) and geostatistics to assess the spatial pattern of soil properties and vegetation, and correlation analyses to explore how these patterns were related. SADIE analysis detected significant clumped patterns in the spatial distribution of vegetation, mosses, fine root accumulation and below-ground stone content. Contoured SADIE index of clustering maps suggested the presence of patchiness in the distribution of earthworm casts, fine roots, below-ground stone content, mosses and biological crusts. Correlation analyses suggested that spatial pattern of some soil properties such as biological crusts, moss cover, surface rock fragments, physical crusts and fine roots were significantly related with above-ground plant distribution. We discuss the spatial arrangement of surface soil properties and suggest mechanistic explanations for the observed spatial patterns and relationships.     

Influence of grazing management on vegetation,soil structure and nutrient distribution and the infiltration of applied rainfall in a semi-arid chenopod shrubland

The experiment utilized a fenceline contrast in vegetation and soil condition that was clearly visible on Landsat imagery. Measurements of vegetation cover, soil structure and chemistry, and infiltration were made. The greatest vegetation change was at the soil surface where the loss of litter and lichen crust cover under heavy grazing accompanied the loss of perennial shrubs. Although grazing caused changes in soil structure and chemistry to less than 10 cm in depth, these changes are quite significant for plant growth. Consistent differences in the infiltration of applied rainfall at two intensities were measured between the grazed and ungrazed sites. At both intensities of application the absence of a lichen crust increased infiltration three-fold on the heavily grazed site compared with the ungrazed site. The implications of these observations on the long-term functioning of this landscape are discussed.     

Grazing‐induced changes in plant–soil feedback alter plant biomass allocation

Large vertebrate herbivores, as well as plant–soil feedback interactions are important drivers of plant performance, plant community composition and vegetation dynamics in terrestrial ecosystems. However, it is poorly understood whether and how large vertebrate herbivores and plant–soil feedback effects interact. Here, we study the response of grassland plant species to grazing‐induced legacy effects in the soil and we explore whether these plant responses can help us to understand long‐term vegetation dynamics in the field. In a greenhouse experiment we tested the response of four grassland plant species, Agrostis capillaris, Festuca rubra, Holcus lanatus and Rumex acetosa, to field‐conditioned soils from grazed and ungrazed grassland. We relate these responses to long‐term vegetation data from a grassland exclosure experiment in the field. In the greenhouse experiment, we found that total biomass production and biomass allocation to roots was higher in soils from grazed than from ungrazed plots. There were only few relationships between plant production in the greenhouse and the abundance of conspecifics in the field. Spatiotemporal patterns in plant community composition were more stable in grazed than ungrazed grassland plots, but were not related to plant–soil feedbacks effects and biomass allocation patterns. We conclude that grazing‐induced soil legacy effects mainly influenced plant biomass allocation patterns, but could not explain altered vegetation dynamics in grazed grasslands. Consequently, the direct effects of grazing on plant community composition (e.g. through modifying light competition or differences in grazing tolerance) appear to overrule indirect effects through changes in plant–soil feedback.