Effects of grazing on patch structure in a semi‐arid two‐phase vegetation mosaic

Abstract. Question: What are the grazing effects in the spatial organization and the internal structure of high and low cover patches from a two‐phase vegetation mosaic? Location: Patagonian steppe, Argentina. Methods: We mapped vegetation under three different grazing conditions: ungrazed, lightly grazed and heavily grazed. We analysed the spatial patterns of the dominant life forms. Also, in each patch type, we determined density, species composition, richness, diversity, size structure and dead biomass of grasses under different grazing conditions. Results: The vegetation was spatially organized in a two‐phase mosaic. High cover patches resulted from the association of grasses and shrubs and low cover patches were represented by scattered tussock grasses on bare ground. This spatial organization was not affected by grazing, but heavy grazing changed the grass species involved in high cover patches and reduced the density and cover of grasses in both patch types. Species richness and diversity in high cover patches decreased under grazing conditions, whereas in low cover patches it remained unchanged. Also, the decrease of palatable grasses was steeper in high cover patches than in low cover patches under grazing conditions. Conclusions: We suggest that although grazing promotes or inhibits particular species, it does not modify the mosaic structure of Patagonian steppe. The fact that the mosaic remained unchanged after 100 years of grazing suggests that grazing does not compromize population processes involved in maintaining patch structure, including seed dispersal, establishment or biotic interactions among life forms.     

Effects of Grazing Regimes on Plant Traits and Soil Nutrients in an Alpine Steppe,Northern Tibetan Plateau

Understanding the impact of grazing intensity on grassland production and soil fertility is of fundamental importance for grassland conservation and management. We thus compared three types of alpine steppe management by studying vegetation traits and soil properties in response to three levels of grazing pressure: permanent grazing (M1), seasonal grazing (M2), and grazing exclusion (M3) in the alpine steppe in Xainza County, Tibetan Plateau. The results showed that community biomass allocation did not support the isometric hypothesis under different grassland management types. Plants in M1 had less aboveground biomass but more belowground biomass in the top soil layer than those in M2 and M3, which was largely due to that root/shoot ratios of dominant plants in M1 were far greater than those in M2 and M3. The interramet distance and the tiller size of the dominant clonal plants were greater in M3 than in M1 and M2, while the resprouting from rhizome buds did not differ significantly among the three greezing regimes. Both soil bulk density and soil available nitrogen in M3 were greater than in M1 at the 15–30 cm soil depth (P = 0.05). Soil organic carbon and soil total nitrogen were greater in M3 than in M1 and M2 (P = 0.05). We conclude that the isometric hypothesis is not supported in this study and fencing is a helpful grassland management in terms of plant growth and soil nutrient retention in alpine steppe. The extreme cold, scarce precipitation and short growing period may be the causation of the unique plant and soil responses to different management regimes.     

Direct and indirect effects of grazing constrain shrub encroachment in semi‐arid Patagonian steppes

Question: What are the long‐term effects of grazing exclusion on the population structure and dynamics of, and interactions among, three dominant shrub species? Location: Grass‐shrub Patagonian steppe, Chubut, Argentina. Methods: Permanent plots were established in grazed paddocks and paddocks excluded from grazing in representative Patagonian rangelands. Shrub abundance, population size‐structure, short‐term (two 3‐yr periods) and long‐term (matrix models) population dynamics, and neighborhood interactions of three native and codominant shrub species (Mulinum spinosum, Senecio filaginoides and Adesmia volckmanni) were measured and analysed using different statistical approaches. Results: The total density of shrubs was 74% higher in paddocks excluded from grazing, owing mainly to increases in Mulinum (80%) and Senecio (68%) species. However, differences in size structure between ungrazed and grazed paddocks were only detected in Mulinum. Demographic rates differed between shrub species, time‐periods and grazing conditions. In particular, recruitment in the short term (especially in wet years) and population growth rate in the long term (λ) were higher in paddocks excluded from grazing only in Mulinum populations. Senecio populations showed a marginal increase in recruitment and mortality independent of the grazing condition in the wet and dry period. Grazing exclusion modified the balance of neighborhood interactions among the three shrub species. In grazing‐exclusion paddocks, there was a balance between positive and negative interspecific interactions, while in grazed paddocks there were more negative intraspecific and interspecific interactions, resulting in a net negative balance of neighborhood interactions. Conclusions: Our understanding of woody encroachment in arid rangelands can be informed through evaluation of direct and indirect effects of grazing exclusion on the abundance and demography of dominant woody species. In Patagonian arid steppes, the occurrence of woody encroachment in rangelands excluded from grazing can be explained by altered responses in plant‐animal and plant‐plant interactions among shrub species.     

内蒙古高原典型草原区河漫滩湿地植物群落退化表征

草原区河流河漫滩草甸是生物多样性表现最充分和生物生产力最高的地段, 但由于过度放牧利用, 绝大部分草甸处于退化状态。该文以锡林河流域中游的河漫滩草甸为研究对象, 比较分析了围封保育湿地与放牧退化湿地的群落组成、地上生物量, 以及共有植物种的植株高度、节间长、叶长、叶宽, 土壤含水量、容重, 群落地下根量及根的分布, 土壤微生物生物量碳、氮的变化。结果表明: 1)放牧使得湿地植物群落优势种发生变化, 原有湿生植物逐渐向旱生化转变, 同时地上及地下生物量明显降低。2)退化湿地的植物呈现显著小型化现象。3)放牧退化湿地的土壤含水量较围封保育湿地低, 其垂直分布及地下根的垂直分布也发生变化。在低河漫滩, 土壤水分随土层的增加而增加, 根量也趋于深层化。但在高河漫滩湿地, 土壤含水量接近典型草原, 根未出现深层化分布趋势。4)放牧践踏引起土壤容重和土壤紧实度增加。5)放牧使得低河漫滩湿地土壤微生物生物量增加, 而在过渡区及高河漫滩湿地, 放牧使得土壤微生物生物量碳、氮含量显著降低。     

Temporal asynchrony in fine‐root biomass may contribute to shrub and grass coexistence in mixed patches

We described seasonal changes in fine‐root biomass of a grass and a shrub dominant species in a plant community characteristic of the arid Patagonian Monte and then we inferred to want extent the observed differences could contribute to the species coexistence. We selected representative plant patches of the natural vegetation arrangement consisting of one isolated plant of the dominant shrub Larrea divaricata (Ld), grass patches formed by one or more bunches of the dominant grass Nassella tenuis (Nt), and mixed patches consisting of one individual of L. divaricata with bunches of N. tenuis under its canopy (LdNt). We assessed the biomass and temporal changes in fine roots of each species in the upper soil (50 cm depth) of each patch type at three‐month intervals during 2 years. The temporal series of fine‐root biomass were compared among patch types and in relation to above‐ground phenology, as well as climate variables (precipitation, arid index and air temperature). Seasonal changes in fine‐root biomass showed similar cycles in the three plant patches with a maximum in spring. The maximum increase in root biomass in Ld and Nt patches occurred during the onset of reproductive growth in winter and spring, respectively. Fine‐root changes in LdNt patches mimicked that in Ld patches. Precipitation inputs were significantly positively and temperature negatively related to fine‐root changes in Nt patches. Fine‐root changes in Ld and LdNt patches were related to the aridity index (positively) and temperature (negatively). We concluded that the observed asynchronies in the date of the largest increases in root biomass and its climate control between the studied grass and shrub species could contribute to the coexistence of plants of both life forms when they overlap their root systems growing in mixed patches. Mechanisms underlying the root patterns observed should be further explored.     

Differential Controls of Water Input on Litter Decomposition and Nitrogen Dynamics in the Patagonian Steppe

Studies of the effects of precipitation on litter decomposition and nitrogen mineralization in arid and semiarid environments have demonstrated contradictory results. We conducted a manipulative experiment with rainout shelters in the semiarid Patagonian steppe, aimed at assessing the direct effects of water availability on litter decomposition and net nitrogen mineralization while isolating the indirect effects. We created four levels of precipitation input: control and three levels (30, 55 and 80%) of precipitation interception and we examined litter decomposition and nutrient release of a dominant grass species, Stipa speciosa, inorganic soil nitrogen, and in situ net nitrogen mineralization over two consecutive years. Litter decomposition rates (k, year⁻¹) varied significantly (P < 0.001) among precipitation interception treatments and were positively correlated with incoming annual precipitation (APPT, mm/year) (k = 0.0007 × APPT + 0.137). In contrast, net N mineralization was not correlated with incoming precipitation. Soil NO₃⁻ significantly decreased with increasing precipitation input, whereas soil NH₄⁺ concentration did not differ among precipitation interception treatments. Controls of water input on litter decomposition appear to be different from controls on N mineralization in the semiarid Patagonian steppe. We suggest that although water availability affects both the mineralization of C and N, it differentially affects the movement and fate of the inorganic products. A consequence of the accumulation of inorganic N during dry episodes is that periods of maximum water and soil nutrient availability may occur at different times. This asynchrony in the availability of N and water in the soil may explain the observed lags in the response of primary production to increases in water availability.     

Production and turnover rates of shallow fine roots in rangelands of the Patagonian Monte, Argentina

Selective sheep grazing in arid rangelands induces a decrease in total cover and grass cover and an increase in the dominance of shrubs. Both life forms differ in aboveground and belowground traits. We hypothesized that grazing disturbance leads to the replacement of grass by shrub fine roots in the upper soil, and this is reflected in changes in the seasonal dynamics of shallow fine roots at the community level. In two sites representative of non-grazed and grazed vegetation states in the Patagonian Monte, we assessed the canopy structure, and the fine root biomass, N concentration, production, and turnover during two consecutive years. The non-grazed site exhibited higher total, grass, and shrub cover than the grazed site. The grazed site had larger or equal fine root biomass than the non-grazed site except for late spring of the second year. This could be associated with the ability of shrubs to develop dimorphic-root systems occupying the soil freed by grasses at the grazed site, and with the larger contribution of grass than shrub fine roots in relation to an extraordinary precipitation event at the non-grazed site. This was consistent with the N concentration in fine roots. Fine root production was positively correlated to temperature at the grazed site and with precipitation at the non-grazed site. Fine root turnover did not differ between sites. Our results indicate that grazing leads to a shifting in the seasonality and main climatic controls of fine root production, while fine root turnover is mostly affected by changes in soil water conditions.     

The role of plant–soil feedbacks and land-use legacies in restoration of a temperate steppe in northern China

Plant–soil feedbacks affect plant performance and plant community dynamics; however, little is known about their role in ecological restoration. Here, we studied plant–soil feedbacks in restoration of steppe vegetation after agricultural disturbance in northern China. First, we analyzed abiotic and biotic soil properties under mono-dominant plant patches in an old-field restoration site and in a ‘target’ steppe site. Second, we tested plant–soil feedbacks by growing plant species from these two sites on soils from con- and heterospecific origin. Soil properties generally did not differ between the old-field site and steppe site, but there were significant differences among mono-dominant plant patches within the sites. While soil species origin (i.e., the plant species beneath which the soil was collected) affected biomass of individual plant species in the feedback experiment, species-level plant–soil feedbacks were ‘neutral’. Soil site origin (old-field, steppe) significantly affected biomass of old-field and steppe species. For example, old-field species had higher biomass in old-field soils than in steppe soils, indicating a positive land-use legacy. However, soil site origin effects depended on the plant species beneath which the soils were collected. The predictive value of abiotic and biotic soil properties in explaining plant biomass differed between and within groups of old-field and steppe species. We conclude that the occurrence of positive land-use legacies for old-field species may retard successional replacement of old-field species by steppe species. However, high levels of idiosyncrasy in responses of old-field and steppe plant species to con- and heterospecific soils indicate interspecific variation in the extent to which soil legacies and plant–soil feedbacks control successional species replacements in Chinese steppe ecosystems.     

Using ants to manage sustainable grazing: Dynamics of ant faunas along sheep grazing gradients conform to four global patterns

Ants are considered an important faunal group for the functioning of arid rangelands, they have a long history of use for environmental monitoring, and exhibit four global patterns in grazing lands: (i) soil and vegetation type are primary determinants of ant community composition, and have a far greater effect on ant community composition than grazing; (ii) grazing induces species compositional change, but does not necessarily affect species richness or abundance; (iii) a species response to grazing is not necessarily consistent across habitats; and (iv) approximately one‐quarter to one‐half of species that are common enough for statistical analysis have significant responses to grazing. Here we report the patterns of arid zone ant faunas as they exist after several decades of sheep grazing in southern Australia, and examine the extent to which they conform to the four global patterns. We measured ant faunas along grazing gradients (varying distance to water) in seven paddocks containing two soil and two vegetation types on five pastoral properties. Total site abundance and richness of ants did not differ significantly with distance from water, but the abundance of 10 (34%) of the 29 most common species did differ; three were increasers, three were decreasers, and four had mixed responses dependent on soil/vegetation type. Rare species showed no trend with grazing intensity. The ant fauna of the more structurally complex vegetation types appeared to be the most vulnerable to grazing effects. Multivariate analysis showed soil type was the primary factor influencing ant faunal composition, followed by vegetation structure; however, grazing treatment effects were present. This study fully supports the recently identified global patterns of ant responses to grazing. It also shows that sampling regional ant faunas using widely dispersed traps can detect ant faunal patterns comparable to studies that use smaller‐scale grids of traps.     

Spatial patterns of the germinable soil seed bank of coexisting perennial-grass species in grazed shrublands of the Patagonian Monte

We compared the size and spatial pattern of the germinable soil seed bank (GSB) of the three dominant perennial-grass species in the arid Patagonian Monte of Argentina. These species differ in plant functional traits, seed morphology, and in their preference by grazers. During three consecutive years, we sampled the soil seed bank at 1 m intervals and the intercepted diameter of perennial-grass species and shrub patches along 50-m linear transects at three sites with different grazing pressure. Soil samples were incubated at field capacity during 12 weeks and emerged seedlings counted. The largest-seeded and most xerophytic Stipa speciosa did not form a GSB. The other two grass species with more mesophytic traits and smaller seeds than S. speciosa formed different GSB. Poa ligularis formed a smaller and more heterogeneous GSB than Stipa tenuis. Grazing had not effect on GSB size of P. ligularis but increased the proportion of seeds under dense shrub canopies. In contrast, grazing reduced the size and also increased the proportion of the GSB of S. tenuis under shrub canopies. We conclude that the size and the spatial pattern of the GSB of the dominant perennial-grass species in the arid Patagonian Monte were related not only to seed and plant traits but also to the spatial distribution of grass plants and their exposure to grazers.     

Effects of differential grazing on decomposition rate and nitrogen availability in a productive mountain grassland

Background and aims

Grazing may influence nutrient cycling in several ways. In productive mountain grasslands of central Argentina cattle grazing maintain a mosaic of different vegetation patches: lawns, grazed intensively and dominated by high quality palatable plants, and open and closed tussock grasslands dominated by less palatable species. We investigated if differences in the resources deposited on soil (litter and faeces) were associated with litter decomposition rates and soil nitrogen (N) availability across these vegetation patches.

Methods

We compared the three vegetation patches in terms of litter and faeces quality and decomposability, annual litterfall and faeces deposition rate. We determined decomposition rates of litter and faces in situ and decomposability of the same substrates in a common garden using “litter bags”. We determined soil N availability (with resin bags) in the vegetation patches. Also, we performed a common plant substrates decomposition experiment to assess the effect of soil environment on decomposition process. This technique provides important insights about the soil environmental controls of decomposition (i.e. the sum of soil physicochemical and biological properties, and microclimate), excluding the substrate quality.

Results

The litter quality and faeces deposition rate were higher in grazing lawns, but the total amounts of carbon (C) and nitrogen (N) deposited on soil were higher in tussock grasslands, due to higher litterfall in these patches. The in situ decomposition rates of litter and faeces, and of the two common plant substrates were not clearly related to either grazing pressure, litterfall or litter quality (C, N, P, lignin, cellulose or hemicellulose content). In situ litter decomposition rate and soil ammonium availability were correlated with the decomposition rates of both common plant substrates. This may suggest that difference in local soil environment among patch types is a stronger driver of decomposition rate than quality or quantity of the resource that enter the soil.

Conclusions

Our results show that, although high grazing pressure improves litter quality and increases faeces input, the reduction in biomass caused by herbivores greatly reduces C and N input for the litter decomposition pathway. We did not find an accelerated decomposition rate in grazing lawns as proposed by general models. Our results point to soil environment as a potential important control that could mask the effect of litter quality on field decomposition rates at local scale.     

Do soil enzymes respond to long-term grazing in an arid ecosystem?

Background and aims

Our objective was to assess the effects of long-term continuous grazing on soil enzyme activities in relation to shifts in plant litter attributes and soil resources in an arid ecosystem, considering both spatial and temporal variations.

Methods

We randomly extracted soil samples with the respective litter cover at 5 modal size plant-covered patches (PCP) and the nearest inter-canopy areas (IC) at Patagonian Monte sites with low, medium and high grazing intensity in winter and summer from 2007 to 2009. We analyzed enzyme activities (dehydrogenase, ß-glucosidase, protease, alkaline and acid phosphatase), microbial biomass-C, organic-C, total soil-N, and moisture in soil and mass and quality in plant litter. We assessed faeces density and plant cover in the field.

Results and conclusions

Grazing led to reduced grass cover, decreasing plant litter mass with increasing soluble phenolics, and reduced phosphatases, ß-glucosidase and microbial biomass-C at PCP. A localized nutrient input from animal excreta seems to promote microbial biomass-C, alkaline phosphatase and dehydrogenase activities but only at IC from the site with high grazing intensity. Plant heterogeneous distribution, plant litter quantity and quality, nutrient inputs from grazers and seasonal variation in soil moisture, also affecting soil resources and microbial biomass, modulate soil enzyme responses to long-term grazing in the arid Patagonian Monte.     

半干旱草地长期封育进程中针茅植物根系格局变化特征

以云雾山不同封育年限草地针茅植物根系和土壤为研究对象,对其根系特征、土壤特性及两者关系进行研究,以探讨分析封育对针茅根系格局的影响。结果表明:(1)针茅植物根系生物量、根长密度、根表面积和根体积在封育初期轻微下降,之后缓慢上升,并在封育30 a草地得到显著增加。(2)随封育年限增加,各根系指标在3种针茅物种间的组成格局具有类似变化规律,具体表现为:长芒草在放牧草地所占比例最高,之后逐渐降低,并在封育30 a草地消失;大针茅所占比例呈先升后降变化规律,并在封育22 a草地达到最大值;甘青针茅仅出现于封育30 a草地,且占据优势地位。(3)大针茅和甘青针茅0—0.6 mm径级根系比例高于大针茅,使其根系直径显著低于大针茅,比根长和比根面积显著高于大针茅;此外,长芒草根组织密度显著高于长芒草和甘青针茅。(4)长期封育在显著提高土壤水分、养分含量和土壤氮磷比的同时显著降低土壤碳氮比,但对微生物生物量碳、氮无明显影响。(5)针茅根系特征与土壤指标的关联性分析显示针茅根系受土壤氮资源的显著影响。     

Plant and soil nitrogen dynamics in mediterranean grasslands: a comparison of annual and perennial grasses

Summary The predominance of annual species in the rangelands of southwestern Spain is not due only to climatic factors but is also strongly influenced by grazing management. Manipulating the grazing system in an experimental plot gave a vegetation structure with patches of annual grasses (mainly Vulpia ssp. and Bromus hordeaceus) and patches of perennial grasses (mainly Phalaris aquatica). This vegetation change allowed us to test the hypothesis that life-cycle differences between annual and perennial grasses affect soil nitrogen availability and plant uptake. Nitrogen availability, measured by in situ incubation, and nitrogen uptake were measured through the growing period (October to June). Amounts of in situ mineralized nitrogen over the whole growth phase were more important for soils supporting perennials (37 ppm) than for soils supporting annuals (27 ppm). The difference between the mineral nitrogen produced in situ and the mineral nitrogen accumulated during the same time in the soil allowed an estimation of the maximum mineral nitrogen quantity which can be taken up by the vegetation during each incubation period. The quantities accumulated over the year were 47 and 38 ppm (or 103 and 83 kg/ha) for soils supporting perennials and annuals respectively. For the same period, amounts of nitrogen immobilized in biomass production were 90 and 70 kg/ha for perennials and annuals respectively. During the autumn, a large proportion of mineral nitrogen was leached from soils supporting annual plants which had only just commenced germination. By contrast, the ability to use mineral nitrogen as soon as autumn rains occurred gave a competitive advantage to the perennial species, but only if they were protected from grazing during this period. The higher mineralization and use of this nitrogen reserve by perennials indicates that they made more efficient use of nitrogen resources than annuals, and validate the initial hypothesis.     

Short-term regrowth responses of four steppe grassland species to grazing intensity, water and nitrogen in Inner Mongolia

Inner Mongolia steppe grasslands are widely used for livestock farming and the regrowth ability of grassland species is therefore strongly influenced not only by water and nutrient availability but also quite heavily by grazing. However, little is known on how grazing, water and nitrogen interactively affect the dominant C3 species (Leymus chinensis, Stipa grandis, Agropyron cristatum) and the C4 species (Cleistogenes squarrosa). Therefore in the 2007 and 2008 growing seasons, a field experiment was carried out to test the hypothesis that under different grazing intensities the dominant species show different short-term regrowth response to simultaneous variation in the availability of water and nitrogen. Single factor and interaction effects of the addition of water (rainfed vs. simulated wet-year) and nitrogen (0 or 25 kg N ha^?1) were analysed along a gradient of four grazing intensities (ungrazed, lightly, moderately and heavily grazed) after one month of grazing exclusion. Water and nitrogen addition affected short-term regrowth of all species in a similar way whereas species responded differently to grazing. Simulated wet-year water availability consistently resulted in higher standing biomass, relative growth rate and cellulase digestible organic matter yield. Supplementary nitrogen promoted standing biomass and crude protein concentration. The nutritive value of all species’ standing biomass showed a similar increase with more intensive grazing. However, heavy grazing led to a clear shift in the relative biomass of the species, i.e. mainly a promotion of the C4 grass, C. squarrosa. In contrast to our hypothesis, there were no differences among species in their response to water or nitrogen addition, whereas, heavy grazing induced the expected species-specific response. Our results suggest that heavy grazing rather than nitrogen or water determine short-term shifts in species composition of the investigated steppe ecosystem. Furthermore, differences in the species-specific growth response to grazing may increase the proportion of the C4 grass C. squarrosa in steppe communities, whereas higher availability of nitrogen and water may lead to higher forage biomass and nutritive value of all investigated species but in short-term cannot compensate for the grazing induced changes in species composition.     

Nutrient foraging traits in 10 co-occurring plant species of contrasting life forms

1 Responses to spatial heterogeneity of soil nutrients were tested in 10 plant species that differ in life form and successional status, but which co-occur in the South Carolina coastal plain. The morphological responses of the root system were tested by assessing scale (represented by root mass and root length densities), precision (preferential proliferation of roots in nutrient-rich patches compared with less fertile patches) and discrimination (ability to detect and proliferate within the richest patches when patches vary in nutrient concentration). We also investigated sensitivity (growth benefits gained as spatial heterogeneity of nutrients increases, measured as total biomass).
2 Ten individuals of each species were grown in pots under four treatments that had differing nutrient distribution but the same overall nutrient addition. Plants were harvested when roots reached pot edge.
3 We observed high variation between species in scale, precision and sensitivity. No significant discrimination responses were observed, although greatest root mass density occurred at intermediate fertility levels for all species.
4 We rejected the hypothesis that scale and precision are negatively correlated. Indeed, in herbaceous species alone, scale and precision were positively correlated.
5 Sensitivity was not closely related to precision, indicating that proliferation of roots in fertile patches does not always yield growth benefits in heterogeneous soils. Further, some sensitive species had very low precision, suggesting that a positive growth response in heterogeneous environments may be related to plasticity in physiology or root life span, rather than morphology.
6 Plant life form was not correlated with precision or sensitivity. However, scale of response was greater in herbs than in woody plants, possibly because the two life forms develop root systems at different rates.     

Multivariate control of root biomass in a semi-arid grassland on the Loess Plateau,China

Aim

Root biomass has long been under-represented in biodiversity–ecosystem functioning studies, despite its dominance in biomass in many arid and semi-arid ecosystems. We aimed to explore the multivariate control over root biomass by plant diversity, together with other biotic and abiotic factors and to evaluate the relative importance of these factors.

Methods

Above- and below-ground traits of 13 communities and soil properties were measured in semi-arid grasslands on the Loess Plateau, China. Structural equation modeling (SEM) was used to evaluate the relative importance of the community and soil characteristics, emphasizing the direct and indirect effects of plant diversity on root biomass.

Results

Significant indirect effects of plant species richness on root biomass were found, although no direct correlation was detected between them. In the indirect pathways, plant species richness showed a positive effect on soil total nitrogen, but a significant negative influence on soil total carbon. Soil total nitrogen and plant diversity had the largest and smallest total effect respectively on root biomass in the model.

Conclusions

Plant species richness was not the strongest determinant of root biomass but had a significant indirect effect, mediated through soil total carbon and nitrogen. This study suggests that greater plant species richness, through a positive influence on soil total nitrogen, may indirectly promote root carbon stock.     

Islands of Fertility: A Byproduct of Grazing?

Empirical work in arid shrubland systems has documented a distinct spatial pattern of soil nutrient distribution, with higher concentrations of nutrients under shrub canopies compared to bare ground interspaces between shrubs. This “fertile island” pattern is considered characteristic of arid ecosystems. However, recent work at a desertified shrubland site reported no island of fertility pattern for nitrogen inside a long-term grazing exclosure despite a strong fertile island pattern outside the exclosure, leading to a hypothesis that the pattern may be associated with grazing. In this paper we examine this hypothesis in two ways. First, we collected soil nitrogen data inside and outside two long-term livestock exclosures. Second, we examined published work on the fertile island pattern for nitrogen, phosphorus and carbon, and its association with livestock grazing. At the field sites, there was a strong fertile island pattern for soil nitrogen in the grazed area outside each exclosure but no fertile island pattern in the ungrazed area inside the fence. From the literature, we identified 49 studies, which represent 99 cases that examine differences in soil nutrients under and between shrubs (that is, tests of the fertile island pattern). In 92 cases, fertile islands were present for at least one of the nutrients. All cases where livestock grazing was present exhibited fertile island patterns for nitrogen and most did so for carbon and phosphorus. In contrast, fertile islands were less consistent where livestock grazing was absent; all nine cases that had been ungrazed for more than 30 years did not exhibit fertile islands for nitrogen. Our empirical data and literature review suggest that the fertile island pattern may be an artifact of grazing rather than an inherent property of arid shrublands. Further, we suggest that the grazing history of a site needs to be explicitly considered when documenting spatial patterns of soil nutrients in arid systems. This has significant implications for our understanding of soil nutrient dynamics in arid systems, the management of rangelands, and the potential for reversal of desertification.     

Sheep Grazing Decreases Organic Carbon and Nitrogen Pools in the Patagonian Steppe: Combination of Direct and Indirect Effects

We explored the net effects of grazing on soil C and N pools in a Patagonian shrub–grass steppe (temperate South America). Net effects result from the combination of direct impacts of grazing on biogeochemical characteristics of microsites with indirect effects on relative cover of vegetated and unvegetated microsites. Within five independent areas, we sampled surface soils in sites subjected to three grazing intensities: (1) ungrazed sites inside grazing exclosures, (2) moderately grazed sites adjacent to them, and (3) intensely grazed sites within the same paddock. Grazing significantly reduced soil C and N pools, although this pattern was clearest in intensely grazed sites. This net effect was due to the combination of a direct reduction of soil N content in bare soil patches, and indirect effects mediated by the increase of the cover of bare soil microsites, with lower C and N content than either grass or shrub microsites. This increase in bare soil cover was accompanied by a reduction in cover of preferred grass species and standing dead material. Finally, stable isotope signatures varied significantly among grazed and ungrazed sites, with δ¹⁵N and δ¹³C significantly depleted in intensely grazed sites, suggesting reduced mineralization with increased grazing intensity. In the Patagonian steppe, grazing appears to exert a negative effect on soil C and N cycles; sound management practices must incorporate the importance of species shifts within life form, and the critical role of standing dead material in maintaining soil C and N stocks and biogeochemical processes. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author Contributions  RAG designed study, performed research, analyzed data, wrote the paper; ATA designed study, wrote the paper; CGGM designed study, performed research, analyzed data; MGP performed research; OES designed study; RBJ designed study, contributed new methods.     

What causes changes in plant litter quality and quantity as consequence of grazing in the Patagonian Monte: Plant cover reduction or changes in species composition?

In Patagonian Monte, as in other arid ecosystems, grazing has triggered changes in vegetation and soil such as plant cover reduction, changes in species composition and soil nutrient losses. Several mechanisms were proposed interconnecting these changes, but evidence supporting them is very scarce. On the basis of published data concerning plant cover by species along grazing gradients and leaf litter production of dominant species, we estimated the effects of grazing on a – quality (N, soluble phenolics and lignin concentrations) and b – quantity (leaf litterfall (LLF) and inputs of nitrogen, soluble phenolics and lignin to the soil) of leaf litter in the Patagonian Monte, discriminating the effect of plant cover reduction from that of species composition. We also evaluated the relationship between senesced leaves traits and the response of species to grazing (i.e. their relative change in plant cover). Grazing causes a reduction in LLF and in the inputs of nitrogen, soluble phenolics and lignin to the soil. In the case of LLF, this reduction was not only a result of the decrease in plant cover but also due to changes in species composition. In contrast, our results showed that the reduction in nitrogen, soluble phenolics and lignin inputs to the soil by LLF is only a consequence of plant cover reduction. Additionally, litter quality was affected through increasing concentration of N and secondary compounds (soluble phenolics and lignin). N and soluble phenolics concentration on senesced leaves were positively related to the response of species to grazing, suggesting that other factors instead of N are relevant to sheep foraging decisions.