Sheep Wool δ13C Reveals No Effect of Grazing on the C3/C4 Ratio of Vegetation in the Inner Mongolia–Mongolia Border Region Grasslands

We tested whether the abundance of C₄ vegetation in grasslands of the Mongolian plateau is influenced by grazing conditions. The analysis exploited the politically originated contrast that exists between Mongolia (low stocking rate, transhumant system) and the district of Inner Mongolia, China (high stocking rate, sedentary system). We estimated the proportion of C₄ carbon (P_C4) in grazed vegetation from the relative carbon isotope ratio (δ¹³C) of sheep wool sampled from 298 annual shearings originating from 1996 to 2007. Annual stocking rates varying over time and between the districts of both countries were taken from regional statistics. The P_C4 pattern within the 0.7 million km² sampling area was geostatistically analyzed and related to stocking rates and temperature gradients. For similar climatic conditions, P_C4 was the same in both countries. Further, a unique relationship was found between P_C4 and July temperature on both sides of the border, which explained 71% of the pattern. Stocking rate and grazing system had no significant influences on present-day C₃/C₄ abundance ratio. This finding suggests that recent changes in the C₃/C₄ ratio of these grasslands are mainly a consequence of regional warming, not overgrazing.     

Nitrogen-15 signals of leaf-litter-soil continuum as a possible indicator of ecosystem nitrogen saturation by forest succession and N loads

Understanding forest carbon cycling responses to atmospheric N deposition is critical to evaluating ecosystem N dynamics. The natural abundance of ¹⁵N (??¹⁵N) has been suggested as an efficient and non-invasive tool to monitor N pools and fluxes. In this study, three successional forests in southern China were treated with four levels of N addition. In each treatment, we measured rates of soil N mineralization, nitrification, N₂O emission and inorganic N leaching as well as N concentration and ?? ¹⁵N of leaves, litters and soils. We found that foliar N concentration and ??¹⁵N were higher in the mature broadleaf forest than in the successional pine or mixed forests. Three-year continuous N addition did not change foliar N concentration, but significantly increased foliar ?? ¹⁵N (p < 0.05). Also, N addition decreased the ?? ¹⁵N of top soil in the N-poor pine and mixed forests and significantly increased that of organic and mineral soils in N-rich broadleaf forests (p < 0.05). In addition, the soil N₂O emission flux and inorganic N leaching rate increased with increasing N addition and were positively correlated with the ¹⁵N enrichment factor (?? _p/s) of forest ecosystems. Our study indicates that ?? ¹⁵N of leaf, litter and soil integrates various information on plant species, forest stand age, exogenous N input and soil N transformation and loss, which can be used to monitor N availability and N dynamics in forest ecosystems caused by increasing N deposition in the future.     

Do grazer hair and faeces reflect the carbon isotope composition of semi-arid C3/C4 grassland?

The carbon isotope composition (δ¹³C) of C3/C4 mixed grassland is reflected in the δ¹³C of diet, hair or faeces of grazers, if ¹³C discrimination (¹³Δ) between grassland vegetation and these tissues is known and constant. However, these relationships could be modified by selective grazing or differential digestibility of the C3 and C4 components, potentially creating a bias between grassland and grazer tissue δ¹³C. Importantly, these factors have never been studied in detail. We investigated the relation between δ¹³C of C3/C4 grassland vegetation and that of faeces and hair of sheep in a 3-year (2005–2007) experiment in the Inner Mongolian semi-arid steppe. The experiment employed six stocking rates (0.375–2.25 sheep ha^?1 year^?1; four replications), which allowed for a large variation in species composition, digestibility, and diet selection. Faecal-nitrogen content, a proxy for digestibility, decreased from 1.9% to 1.5% during the grazing period due to aging of the herbage. At the same time, the C3/C4 ratio decreased due to the later growth initiation of C4 species. ¹³Δ between diet and faeces (¹³Δ_DF; 0.6‰) and between diet and hair (¹³Δ_DH; ?3.9‰) were not influenced by stocking rate, period in the season or C3/C4 ratio. Moreover, faeces–hair discrimination (¹³Δ_FH; ?4.3‰), which reflects differences between digestibility of the C3 and C4 components, did not vary along the different gradients. The δ¹³C of grassland vegetation can be estimated from the δ¹³C of sheep faeces and hair, provided that ¹³Δ was accounted for. This is useful for landscape- or regional-scale investigations or reconstruction of C3/C4 vegetation distribution from faeces and hair, which provide different temporal and spatial integration of grassland isotope signals.     

Feedback of grazing on gross rates of N mineralization and inorganic N partitioning in steppe soils of Inner Mongolia

Plant-microbe interactions are crucial regulators of belowground nitrogen cycling in terrestrial ecosystems. However, such interactions have mostly been excluded from experimental setups for the investigation of gross inorganic N fluxes and N partitioning to plants and microorganisms. Ungulate grazing is likely to feed back on soil N fluxes, and hence it is of special importance to simultaneously investigate grazing effects on both plant and microbial N fluxes in intact plant-soil systems, where plant-microbe interactions persist during the experimental incubation. Based on the homogenous ¹⁵NH ₄ ⁺ labelling of intact plant-soil monoliths we investigated how various stocking rates (0, 2.35, 4.8 and 7.85 sheep ha^?1 grazing season^?1) in steppe of Inner Mongolia feedback on gross rates of N mineralization and short-term inorganic N partitioning between plant, microbial and soil N pools. Our results showed that the effect of grazing on gross N mineralization was non-uniform. At low stocking rate gross N mineralization tended to decrease but increased with higher grazing pressure. Hence, there was no significant correlation between stocking rate and gross N mineralization across the investigated grazing intensities. Grazing decreased ¹⁵N recovery both in plant and microbial N pools but strongly promoted NO ₃ ^? accumulation in the soil and thus negatively affected potential ecosystem N retention. This appeared to be closely related to the grazing-induced decline in easily degradable soil C availability at increasing stocking rate.     

Optimizing Carbon Storage Within a Spatially Heterogeneous Upland Grassland Through Sheep Grazing Management

Livestock grazing is known to influence carbon (C) storage in vegetation and soil. Yet, for grazing management to be used to optimize C storage, large scale investigations that take into account the typically heterogeneous distribution of grazers and C across the landscape are required. In a landscape-scale grazing experiment in the Scottish uplands, we quantified C stored in swards dominated by the widespread tussock-forming grass species Molinia caerulea. The impact of three sheep stocking treatments (‘commercial’ 2.7 ewes ha^?1 y^?1, ‘low’ 0.9 ewes ha^?1 y^?1 and no livestock) on plant C stocks was determined at three spatial scales; tussock, sward and landscape, and these data were used to predict long-term changes in soil organic carbon (SOC). We found that tussocks were particularly dense C stores (that is, high C mass per unit area) and that grazing reduced their abundance and thus influenced C stocks held in M. caerulea swards across the landscape; C stocks were 3.83, 5.01 and 6.85 Mg C ha^?1 under commercial sheep grazing, low sheep grazing and no grazing, respectively. Measured vegetation C in the three grazing treatments provided annual C inputs to RothC, an organic matter turnover model, to predict changes in SOC over 100 years. RothC predicted SOC to decline under commercial sheep stocking and increase under low sheep grazing and no grazing. Our findings suggest that no sheep and low-intensity sheep grazing are better upland management practices for enhancing plant and soil C sequestration than commercial sheep grazing. This is evaluated in the context of other upland management objectives.     

Long-Term Increase in Aboveground Carbon Stocks Following Exclusion of Grazers and Forest Establishment in an Alpine Ecosystem

Ecosystem stores of carbon are a key component in the global carbon cycle. Many studies have examined the impact of climate change on ecosystem carbon storage, but few have investigated the impact of land-use change and herbivory. However, land-use change is a major aspect of environmental change, and livestock grazing is the most extensive land use globally. In this study, we combine a grazing exclosure experiment and a natural experiment to test the impact of grazer exclusion on vegetation dynamics and ecosystem carbon stores in the short term (12-year exclosures), and the long term (islands inaccessible to livestock), in a heavily grazed mountain region in Norway. Following long-term absence of sheep, birch forest was present. The grazing-resistant grass Nardus stricta, dominated under long-term grazing, whilst the selected grass Deschampsia flexuosa and herb species dominated the vegetation layer in the long-term absence of sheep. The established birch forest led to vegetation carbon stocks being higher on the islands (0.56 kg C m^?2 on the islands compared to 0.18 kg C m^?2 where grazed) and no difference in soil carbon stocks. In the short-term exclusion of sheep, there were minor differences in carbon stocks reflecting the longer term changes. These results show that aboveground carbon stocks are higher in the long-term absence of sheep than in the continual presence of high sheep densities, associated with a vegetation state change between tundra and forest. The reduction of herbivore populations can facilitate forest establishment and increase aboveground carbon stocks, however, the sequestration rate is low.     

Annual methane uptake by temperate semiarid steppes as regulated by stocking rates,aboveground plant biomass and topsoil air permeability

Overgrazing‐induced degradation of temperate semiarid steppes may affect the soil sink for atmospheric methane (CH₄). However, previous studies have primarily focused on the growing season and on single grazing patterns. Thus, the response of annual CH₄ uptake by steppes compared with various grazing practices is uncertain. In this study, we investigated the effects of grazing on the annual CH₄ uptake by two typical Eurasian semiarid steppes (the Stipa grandis steppe and the Leymus chinensis steppe) located in Inner Mongolia, China. The CH₄ fluxes were measured year‐round using static chambers and gas chromatography at 12 field sites that differed primarily in grazing intensities. Our results indicated that steppe soils were CH₄ sinks throughout the year. The annual CH₄ uptake correlated with stocking rates, whereas the seasonality of CH₄ uptake was primarily dominated by temperature. The annual CH₄ uptake at all sites averaged 3.7±0.7 kg C ha^?1 yr^?1 (range: 2.3–4.5), where approximately 35% (range: 23–40%) occurred during the nongrowing season. Light‐to‐moderate grazing (stocking rate≤1 sheep ha^?1 yr^?1) did not significantly change the annual CH₄ uptake compared with ungrazed steppes, but heavy grazing reduced annual CH₄ uptake significantly (by 24–31%, P<0.05). These findings imply that easing the pressure of heavily grazed steppes (e.g. moving to light or moderate stocking rates) would help restore steppe soil sinks for atmospheric CH₄. The empirical equations based on the significant relationships between annual CH₄ uptake and stocking rates, aboveground plant biomass and topsoil air permeability (P<0.01) could provide simple approaches for the estimation of regional CH₄ uptake by temperate semiarid steppes.     

Nitrogen transfer between herbivores and their forage species

Herbivores may increase the productivity of forage plants; however, this depends on the return of nutrients from faeces to the forage plants. The aim of this study was to test if nitrogen (N) from faeces is available to forage plants and whether the return of nutrients differs between plant species using ¹⁵N natural abundance in faeces and plant tissue. To investigate the effect of grazing on N transfer, we carried out a grazing experiment in wet and mesic tundra on high Arctic Spitsbergen using barnacle geese (Branta leucopsis) as the model herbivore. N inputs (from faeces) increased with grazing pressure at both the wet and mesic sites, with the greatest N input from faeces at the wet site. The δ¹⁵N ratio in plant tissue from grazed plots was enriched in mosses and the dwarf shrub species, reflecting the δ¹⁵N signature of faeces-derived N, but no such pattern was observed in the dominant grasses. This study demonstrates that the δ¹⁵N signature of faeces and forage species is a useful tool to explore how grazing impacts on N acquisition. Our findings suggest that plant species which acquire their N close to the soil surface (e.g. mosses) access more of the N from faeces than species with deeper root systems (e.g. grasses) suggesting a transfer of N from the preferred forage species to the mosses and dwarf shrubs, which are less preferred by the geese. In conclusion, the moss layer appears to disrupt the nitrogen return from herbivores to their forage species.     

Feeding ecology and trophic level of the banded guitarfish, Zapteryx exasperata, inferred from stable isotopes and stomach contents analysis

The feeding ecology and trophic level of the banded guitarfish, Zapteryx exasperata, from the Gulf of California, were assessed using stable isotope analysis (SIA) of carbon and nitrogen in muscle samples, and stomach contents analysis (SCA). Males and females showed similar diet preferences with a high diet overlap (C???=?0.99) and similar values of ??¹³C and ??¹⁵N (ANOVA, F?=?0.0 p?=?0.94 for ??¹³C; F?=?0.1 p?=?0.78 for ??¹⁵N). The main prey groups found were demersal fishes dominated by the daisy midshipman, Porichthys margaritatus, (>90%IRI), however, pelagic fishes were also found (Engraulis mordax 2%IRI). An ontogenetic diet shift was found, indicated by the importance of crustaceans in the juvenile diet and the significant isotopic differences between juveniles and adults (??¹³C, ANOVA, F?=?13.3 p?=?0.0004 and ??¹⁵N, ANOVA, F?=?4.7 p?=?0.03). An overall trend of increase in ??¹⁵N values with increasing body length (F?=?8.15 p?=?0.005) was observed. The mean trophic level estimated in this work by the two methods was ~4.1, indicating that this species is a top predator in the Gulf of California. Zapteryx exasperata is a specialist feeder (Bi?=?0.11, H???=?2.32), consuming mainly benthic fishes in coastal areas of the Gulf of California. The importance of this species in the benthic ecosystem equilibrium as a top predator is an important point to keep in mind when developing future management plans of the batoid fishery in the Gulf of California.     

不同起始状态对草原群落恢复演替的影响

内蒙古典型草原,由于过度放牧利用,绝大部分草原处于退化状态.为了使退化草原得到较好的恢复,以锡林郭勒盟白音锡勒牧场典型草原为研究对象,比较分析了在不同起始状态下的草原群落,经过6a的自然恢复,其各自的群落组成,地上生物量及共有种的植株高度、节间长、叶长、叶宽,土壤紧实度和容重.结果表明:1)不同放牧率的植物群落,经过6a的禁牧恢复,群落类型发生了变化且群落趋于一致.2)当放牧率SR≤5.33羊/hm2(SR4)时,演替起始状态对草原群落地上生物量的恢复没有影响;当放牧率SR＞5.33羊/hm2时,演替起始状态对草原群落地上生物量的恢复产生影响,其结果是导致当前生物量降低,不利于草原的恢复.3)不同放牧率植物群落的植物个体特征趋于一致,“个体小型化”现象消失.同时,也说明群落恢复演替的起点不同,正常化的时间没有太大的变化.4)不同放牧率植物群落的土壤紧实度和容重经过6a的禁牧恢复,没有得到完全恢复,但均达到一致的水平.     

Incorporating sheep into dryland grain production systems: II. Impact on changes in biomass and weed density

Weed control in fallow management to conserve soil moisture and nutrients is the largest variable cost to dryland grain production. Our objective was to compare burning, grazing, tilling, trampling and clipping wheat stubble fields on changes in total aboveground biomass and weed density. Treatments were evaluated in three experiments using a randomized complete block design for each experiment with four replications at each site. Contrasts statements were used to make pre-planned comparisons. For experiment 1, treatments were fall tilled, fall grazed, spring grazed, fall and spring combined (Fall/Spr) grazed, and an untreated control. For grazing treatments, five mature ewes were confined with electric fence to 111 m² plot for 24 h for fall and spring resulting in a stocking rate of 452 sheep day/ha. For Fall/Spr the stocking rate was 904 sheep day/ha. For experiment 2, treatments were fall grazed, fall burned, fall tilled, and an untreated control. In experiment 3, treatments were fall trampling by sheep, spring trampling by sheep, fall and spring combined (Fall/Spr) trampling by sheep, stubble hand clipped to a height of 4.5 cm, and an untreated control. Trampling treatments were applied at the same stocking rates as grazing treatments but sheep were muzzled to prevent intake. Data were collected in the fall, prior to treatment imposition, and spring, after treatments had been removed. Post treatment biomass and weed density were greater (P < 0.05) in either control or tilled plots when compared to grazed plots. Post treatment biomass and weed density were greater (P < 0.01) for control than burned plots. Post treatment biomass, weed density, and percent change in these variables, did not differ (P > 0.08) between burned and tilled, and burned and grazed treatments. These results indicate the potential for using grazing sheep as a component in fallow management to reduce biomass and control weeds.     

Stable N isotope composition of nitrate reflects N transformations during the passage of water through a montane rain forest in Ecuador

Knowledge of the fate of deposited N in the possibly N-limited, highly biodiverse north Andean forests is important because of the possible effects of N inputs on plant performance and species composition. We analyzed concentrations and fluxes of NO₃ ^??CN, NH₄ ⁺?CN and dissolved organic N (DON) in rainfall, throughfall, litter leachate, mineral soil solutions (0.15?C0.30 m depths) and stream water in a montane forest in Ecuador during four consecutive quarters and used the natural ¹⁵N abundance in NO₃ ^? during the passage of rain water through the ecosystem and bulk ??¹⁵N values in soil to detect N transformations. Depletion of ¹⁵N in NO₃ ^? and increased NO₃ ^??CN fluxes during the passage through the canopy and the organic layer indicated nitrification in these compartments. During leaching from the organic layer to mineral soil and stream, NO₃ ^? concentrations progressively decreased and were enriched in ¹⁵N but did not reach the ??¹⁵N values of solid phase organic matter (??¹⁵N = 5.6?C6.7??). This suggested a combination of nitrification and denitrification in mineral soil. In the wettest quarter, the ??¹⁵N value of NO₃ ^? in litter leachate was smaller (??¹⁵N = ?1.58??) than in the other quarters (??¹⁵N = ?9.38 ± SE 0.46??) probably because of reduced mineralization and associated fractionation against ¹⁵N. Nitrogen isotope fractionation of NO₃ ^? between litter leachate and stream water was smaller in the wettest period than in the other periods probably because of a higher rate of denitrification and continuous dilution by isotopically lighter NO₃ ^??CN from throughfall and nitrification in the organic layer during the wettest period. The stable N isotope composition of NO₃ ^? gave valuable indications of N transformations during the passage of water through the forest ecosystem from rainfall to the stream.     

Thyme travels: 15N isoscapes of Thymus vulgaris L. invasion in lightly grazed pastoral communities

Alterations to ecosystem nitrogen (N) cycling by introduced plant species may increase the invasibility of habitat providing a positive feedback for the introduced species to become invasive. Spatial patterns of foliar and soil δ¹⁵N ratios reflect variation in rates and process of N‐cycling across invaded landscapes and provide insight into N‐source uptake and utilization strategies of invasive plant species. To evaluate invasion‐associated changes in soil and foliar δ¹⁵N at different scales: regional (among different sites), local (between north‐ and south‐facing aspect at the same site), and microsite (within populations in the same community), we measured foliar and soil δ¹⁵ N, animal faeces cover (as a proxy for grazing intensity) and N₂‐fixing species cover from inside to outside Thymus vulgaris L. (thyme)‐invaded lightly grazed pastoral communities in Central Otago, southern South Island, New Zealand. Mean thyme foliar δ¹⁵N were near‐zero across the invaded landscape, and did not change across the advancing edge of invasion or with aspect. There was no evidence that associations with N₂‐fixing species provide a potential N source. Soil δ¹⁵N was lower inside of thyme compared to at the edge or outside of thyme and was varied between aspects at some sites. Animal faeces cover as a proxy for grazing intensity explained only 23% of this observed variation of soil δ¹⁵N. Thyme invasion may result in lowered soil δ¹⁵N reflecting alterations to N dynamics. Associated invasion‐related impacts of animal grazing may also impact soil δ¹⁵N. Further studies are required to distinguish the underlying mechanism responsible for the observed patterns of foliar and soil δ¹⁵N values across thyme‐invaded Central Otago landscapes.     

Insensitivity of Soil Microbial Activity to Temporal Variation in Soil N in Subarctic Tundra: Evidence from Responses to Large Migratory Grazers

Large migratory grazers commonly influence soil processes in tundra ecosystems. However, the extent to which grazing effects are limited to intensive grazing periods associated with migration has not previously been investigated. We analyzed seasonal patterns in soil nitrogen (N), microbial respiration and extracellular enzyme activities (EEAs) in a lightly grazed tundra and a heavily grazed tundra that has been subjected to intensive grazing during reindeer (Rangifer tarandus L.) migration for the past 50 years. We hypothesized that due to the fertilizing effect of the reindeer, microbial respiration and EEAs related to microbial C acquisition should be higher in heavily grazed areas compared to lightly grazed areas and that the effects of grazing should be strongest during reindeer migration. Reindeer migration caused a dramatic peak in soil N availability, but in contrast to our predictions, the effect of grazing was more or less constant over the growing season and the seasonal patterns of microbial activities and microbial N were strikingly uniform between the lightly and heavily grazed areas. Microbial respiration and the EEAs of β-glucosidase, acid-phosphatase, and leucine-aminopeptidase were higher, whereas that of N-acetylglucosamidase was lower in the heavily grazed area. Experimental fertilization had no effect on EEAs related to C acquisition at either level of grazing intensity. Our findings suggest that soil microbial activities were independent of grazing-induced temporal variation in soil N availability. Instead, the effect of grazing on soil microbial activities appeared to be mediated by substrate availability for soil microorganisms. Following a shift in the dominant vegetation in response to grazing from dwarf shrubs to graminoids, the effect of grazing on soil processes is no longer sensitive to temporal grazing patterns; rather, grazers exert a consistent positive effect on the soil microbial potential for soil C decomposition.     

Nitrogen cycling in grazed pastures at elevated CO2: N returns by ruminants

In pastures grazed by large herbivores, nutrients cycle both through litter and animal excreta. We compared nitrogen (N) returns from sheep grazing a temperate pasture exposed to ambient or elevated CO₂ (475 μmol mol^?1) in a FACE (Free Air CO₂ Enrichment) experiment established in the spring of 1997. In the spring of 2000 and 2001, we measured the chemical composition of the diet, sheep faeces and of individual plant species before grazing to characterize feed intake and to compare the intake of N to the N produced in faeces. In both years under elevated CO₂, leaves of the individual species exhibited lower N concentrations and higher water‐soluble carbohydrate (WSC) concentrations. There was a significantly greater proportion of legume in the diet at elevated CO₂ but, together with the changes in chemical composition of individual species, this resulted in diets that had similar N but higher WSC and digestibility for both ambient and elevated CO₂. We found that a greater proportion of dietary N was partitioned to urine at elevated CO₂, probably because of the higher proportion of legume N in the diet, with possible differences in protein quality. A potentially significant consequence of this change in partitioning is greater N loss through volatilization at higher CO₂ levels.     

Control of nematode parasites by grazing management—II. Decontamination of sheep and cattle pastures by varying periods of grazing with the alternate host

Commencing in December 1970, paddocks of a uniform series of sheep pastures were grazed for 6, 12 or 24 weeks by either yearling steers is or yearling ewes. Cattle pastures were treated similarly. All ewes and steers were pre-dosed with anthelmintic.At the conclusion of alternate grazing the effectiveness of the grazing treatments was evaluated by grazing each paddock for 1 month with either ten worm-free lambs (sheep pastures) or three wormfree calves (cattle pastures). The test animals were then slaughtered for total worm counts. The grazing of sheep pastures with cattle for 6, 12 or 24 weeks from December onwards resulted in reductions in numbers of Haemonchus contortus and Trichostrongylus colubriformis in test lambs. In comparison with continuous grazing by sheep, Nematodirus spp was only reduced after 24 weeks grazing by cattle. Cattle pastures grazed by sheep for 6 weeks showed no reduction in numbers of Ostertagia ostertagi or Cooperia oncophora in test calves. However after 12 weeks with sheep, numbers of O. ostertagi though not C. oncophora were reduced and after 24 weeks of alternate grazing both these species were reduced.Calves following the 6 week sheep treatments acquired both Haemonchus contortus and Trichostrongylus colubriformis and the calves from the 12 week sheep treatment paddock also carried H. contortus. For sheep the only evidence of cross-transmission was the occurrence of small numbers of Cooperia oncophora in test lambs from the 24 week cattle grazing treatment.The results provide evidence that sequential stocking with cattle and sheep in conjunction with anthelmintic treatment is an effective management strategy for preparing parasitologically safer pastures, but further information is required to determine the optimum timing of sequential stocking in farming situations.     

Effect of sheep grazing on seed circulation on the Loess Plateau

In grazing ecosystems, mature seeds fall directly to the soil to form the soil seed bank (SSB), or are ingested by grazing livestock to become part of the dung seed bank (DSB; i.e., seed circulation). Both the SSB and DSB form the basis for the natural regeneration of vegetation. However, little is known about the relationships between the SSB, DSB, and aboveground vegetation (AGV) community under different stocking rates (SRs). This study investigated the relationships between the SSB, seeds in Tan sheep (Ovis aries) dung, and AGV at different SRs (0, 2.7, 5.3, and 8.7 sheep ha^–1) in a semiarid region of the Loess Plateau in China. We found that Tan sheep grazing increased the species richness heterogeneity of grassland vegetation, and negatively influenced the density of AGV. Under natural conditions, 17 species from soil‐borne seeds and 10 species from Tan sheep dung germinated. There was low species similarity between the soil and DSBs and AGV. Sheep SR and the seed banks (soil and dung) were negatively correlated with AGV. Seeds are cycled from herbage to livestock to soil during cold season grazing; the seasonal nature of this seed dispersal is an adaptation to harsh, semiarid environments. Increased seed bank diversity under sheep grazing facilitates grassland regeneration on the Loess Plateau, similarly to other semiarid regions globally.     

Dynamics of vegetation and soil carbon and nitrogen accumulation over 26 years under controlled grazing in a desert shrubland

For decades, arid desert ecosystems in northwest China, covering one-fourth the country’s land surface, have experienced a rapid decline in plant species diversity, productivity and soil carbon stock owing to degradation by overgrazing. In this study, plant community composition, diversity and productivity, as well as soil carbon (C) and nitrogen (N) stocks, were monitored over 26 years from 1981 to 2006 in a severely degraded Haloxylon ammodendron-dominated shrubland where livestock densities were reduced from 4–5 to 1–2 dry sheep equivalent ha^-1. The objective was to assess long-term grazing effects on vegetation and soil C and N accumulation dynamics. Results showed that the reduction of grazing pressure significantly increased vegetation cover, plant diversity and productivity, resulting primarily from an increase in livestock-preferred species. Controlled grazing also led to marked increases in soil C and N stocks in the top 30 cm of soil. This increase was strongly associated with increased plant species richness, vegetation cover and biomass production. Averaged over 26 years, soil C and N accumulated at rates of 89.9 g?C and 8.4 g?N m^-2?year^-1, respectively, but rates of C and N accumulation varied greatly at different time periods. The greatest species regeneration occurred in the first 8 years, but the largest C and N accumulation took place during years 9–18, with a time-lag in response to changes in vegetation. Our results provide insights into the long-term recovery patterns of different ecosystem components from the influence of prolonged overgrazing disturbance that cannot be inferred from a short-term study. The findings are important for assessing the resilience of these livestock-disturbed desert ecosystems and developing a more effective strategy for the management of this important biome from a long-term perspective.     

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.     

Coarser wool is not a necessary consequence of sheep aging: allometric relationship between fibre diameter and fleece-free liveweight of Saxon Merino sheep

The mean fibre diameter (MFD) of wool is the primary determinant of price, processing performance and textile quality. This study determines the primary influences on MFD as Saxon Merino sheep age, by allometrically relating MFD to fleece-free liveweight (FFLwt). In total, 79 sheep were grazed in combinations of three stocking rates and two grazing systems (GS: sheep only; mixed with Angora goats) and studied over 3 years. Measurements were made over 14 consecutive periods (Segments), including segments of FFLwt gain or FFLwt loss. Using shearing and liveweight records and dye-bands on wool, the FFLwt and average daily gain (ADG) of each sheep were determined for each segment. The mean and range in key measurements were as follows: FFLwt, 40.1 (23.1 to 64.1) kg; MFD, 18.8 (12.7 to 25.8) μm. A random coefficient restricted maximum likelihood (REML) regression mixed model was developed to relate the logarithm of MFD to the logarithm of FFLwt and other effects. The model can be written in the form of β(A) is a random animal effect, γ(Segment.Plot) a random effect associated with Segment.plot combinations, and κ a constant that depends on GS, random animal effects, random Segment.plot combination effects, Segment and ADG. Thus, MFD was allometrically related to the cube root of FFLwt over seasons and years for sheep, but to the square root of FFLwt for sheep grazed with goats. The result for sheep grazed alone accords with a primary response being that the allocation of nutrients towards the cross-sectional growth of wool follicles is proportional to the changes in the skin surface area arising from changes in the size of the sheep. The proportionality constant varied systematically with ADG, and in sheep only grazing, was about 5 when sheep lost 100 g/day and about 6 when sheep gained 100 g/day. The proportionality constant did not systematically change with chronological age. The variation in the allometric coefficient between individual sheep indicates that some sheep were more sensitive to changes in FFLwt than other sheep. Key practical implications include the following: (a) the reporting of systematic increases in MFD with age is likely to be a consequence of allowing sheep to increase in size during shearing intervals as they age; (b) comparisons of MFD between sheep are more likely to have a biological basis when standardised to a common FFLwt and not just to a common age; (c) wool quality (MFD, staple strength) are most likely to be optimised in management systems that maintain constant FFLwt of adult sheep within and between years.