Livestock turnover and dynamic livestock carrying capacity are crucial factors for alpine grassland management: The Qinghai-Tibetan plateau as a case study.

Alpine grasslands are distributed widely on high-elevated ranges and plateaus from the wet tropics to polar regions, accounting for approximately 3% of the world's land area. The Qinghai-Tibetan Plateau (QTP) is the highest and largest plateau in the world, and approximately 60% of the plateau consists of alpine grassland, which is used mainly for grazing animals. Livestock structure was determined in Guinan (GN), Yushu (YS) and Maqu counties (MQ) on the QTP by interviewing 235 local pastoralists. Based on data collected from GN, the livestock carrying capacity was calculated using herbage dry matter biomass intake (LCCm) by the livestock, and the metabolizable energy yield (LCCe) and digestible crude protein (LCCp) available in pasture. The pasture area per household differed among the regions of the QTP, which was the main reason for the difference in livestock stocking rate. The householders raised the appropriate proportion of breeding females and young yaks and sheep in GN and MQ, but not in YS, to maintain a constant turnover. Most pasture in YS was used at the community level, especially in summer. The calculated carrying capacities based on metabolizable energy yield (LCCe) of the pasture and dry matter biomass (LCCm) were similar in most months except for August, when the value of LCCe was higher than LCCm. Based on the digestible protein of the pasture, the calculated livestock carrying capacity overestimated the actual carrying capacity during the herbage growing season from May to September. Appropriate practices should be taken in different regions of QTP, such as providing supplementary feed, especially protein, during the forage non-growing season. Livestock carrying capacity should be adjusted dynamically, and calculated by a number of parameters. The stocking rate should be controlled to optimize livestock production and curb or minimize grassland degradation to generate a sustainable system. This study examined the grasslands and LCC on the QTP, but the results could be applied to grasslands worldwide.

Variation of root traits and its influences on soil organic carbon stability in response to altered precipitation in an alpine meadow.

Soil organic carbon (SOC) dynamics are strongly controlled by plant roots. Yet, how variation of root traits under precipitation change influences SOC stability remains unclear. As part of a 5-year field experiment manipulating precipitation including 90 % (0.1P), 50 % (0.5P), 30 % (0.7P) decrease, and 50 % increase (1.5P), this study was designed to assess the effects of changing precipitation on root traits and production dynamics by minirhizotron and examine how such influences regulate SOC stability in an alpine meadow on the Qinghai-Tibetan Plateau. We found that root length density (RLD), specific root length (SRL), root branching intensity (RBI), and root residue carbon input (RC input) exhibited no significant response, whereas root turnover (RT), root carbon (C), nitrogen (N) concentrations and C/N ratio were altered by precipitation change with nonlinear trends. Absorptive root RT positively correlated to manipulated precipitation within the interannual precipitation range in topsoil, but it showed no significant change under extreme drought treatment. Alpine meadows can maintain the SOC content and density under varied precipitation. However, it showed significant variation in aggregate stability and organic carbon (OC) distribution in aggregates in topsoil, which were mainly due to the strong direct effects of soil moisture and partly related to RLD and RC input of transport roots. Although subsurface soil aggregate stability and OC associated with aggregates were not modified, our results indicated a risk of SOC stability variation in subsurface soil if absorptive root RT and SRL changed. These findings provide vital information to predict responses of SOC dynamics of alpine meadow to future climate change.

Precipitation affects soil nitrogen fixation by regulating active diazotrophs and nitrate nitrogen in an alpine grassland of Qinghai-Tibetan Plateau.

Soil asymbiotic nitrogen (N) fixation provides a critical N source to support plant growth in alpine grasslands, and precipitation change is expected to lead to shifts in soil asymbiotic N fixation. However, large gaps remain in understanding the response of soil asymbiotic N fixation to precipitation gradients. Here we simulated five precipitation gradients (10 % (0.1P), 50 % (0.5P), 70 % (0.7P), 100 % (1.0P) and 150 % (1.5P) of the natural precipitation) in an alpine grassland of Qinghai-Tibetan Plateau and examined the soil nitrogenase activity and N fixation rate for each gradient. Quantitative PCR and high-throughput sequencing were used to measure the abundance and community composition of the soil nifH DNA (total diazotrophs) and nifH RNA reverse transcription (active diazotrophs) gene. Our results showed that the soil diazotrophic abundance, diversity and nifH gene expression rate peaked under the 0.5P. Soil nitrogenase activity and N fixation rate varied in the range 0.032-0.073 nmol·C2H4·g-1·h-1 and 0.008-0.022 nmol·N2·g-1·h-1 respectively, being highest under the 0.5P. The 50 % precipitation reduction enhanced the gene expression rates of Azospirillum and Halorhodospira which were likely responsible for the high N fixation potential. The 0.5P treatment also possessed a larger and more complex active diazotrophic network than the other treatments, which facilitated the resistance of diazotrophic community to environmental stress and thus maintained a high N fixation potential. The active diazotrophic abundance had the largest positive effect on soil N fixation, while nitrate nitrogen had the largest negative effect. Together, our study suggested that appropriate precipitation reduction can enhance soil N fixation through promoting the abundance of the soil active diazotrophs and decreasing soil nitrate nitrogen, and soil active diazotrophs and nitrate nitrogen should be considered in predicting soil N inputs in the alpine grassland of Qinghai-Tibetan Plateau under precipitation change.

Astragalus root extract improved average daily gain, immunity, antioxidant status and ruminal microbiota of early weaned yak calves.

BACKGROUND: Early weaning in yak calves is being attempted to improve yak reproduction rate. However, this has to be done with caution because of the high mortality rate of calves due to the lack of nutrients and the harsh environmental conditions. Twenty-four weaned male yak calves were used in a 60 day feeding trial in which astragalus root extract (ARE) was supplemented. They were assigned randomly to one of four dietary treatments (n = six per treatment) that differed in ARE level: 0 g kg-1 (control), ARE0 ; 20 g kg-1 , ARE20 ; 50 g kg-1 , ARE50; and 80 g kg-1 dry matter intake (DMI), ARE80 . RESULTS: Final bodyweight and average daily gain (ADG) were significantly higher and the DMI/ADG ratio was significantly lower in calves with ARE supplementation than control (ARE0 ) calves. Ruminal concentrations of acetate and propionate and serum concentration of superoxide dismutase in ARE80 calves were higher than in the other groups and serum concentration of insulin was higher in ARE80 calves than in ARE20 calves. Serum immunoglobulin G (IgG) and interleukin-2 (IL-2) concentrations in ARE-fed calves were higher than in controls. Serum tumor necrosis factor (TNF-α) concentration was higher in ARE50 and ARE80 groups than ARE0 calves and serum interleukin-6 (IL-6) concentration was higher in ARE80 than in ARE0 calves. Serum immunoglobulin A (IgA), IgG and immunoglobulin M (IgM) concentrations increased with age in ARE-fed calves. ARE supplementation increased the abundance of fiber degrading bacteria. CONCLUSION: ARE at a dosage of 5% to 8% DMI can be supplemented to early weaned yak calves to improve growth performance, antioxidant capacity and immunity. © 2020 Society of Chemical Industry.

Rangeland sharing by cattle and bees: moderate grazing does not impair bee communities and resource availability.

Rangelands are a dominant anthropogenic land use and a main driver of natural habitat loss worldwide. Land sharing, the integration of agricultural production and biodiversity conservation, may provide a platform for managing rangelands to fulfill multiple ecosystem services. However, livestock grazing can greatly affect biodiversity and little is known about its effects on providers of focal ecosystem services, such as pollinators. We investigated the effect of cattle grazing on bee communities and their foraging and nesting resources in Mediterranean rangelands. Specifically, we explored the effect of moderate cattle grazing on flowering plant abundance, species richness and composition, the diversity of nesting substrates, and consequently, the possible effects on wild bee and honey bee foraging activity, species diversity, and community composition. We conducted field research in the Mediterranean rangelands of Israel during the main bee activity season, in the spring of 2012 and 2013, comparing paired cattle-grazed and ungrazed areas. The availability of floral and nesting resources for bees was unaffected or positively affected by grazing. Similarly, wild bee abundance, species richness, and composition were not affected by grazing, but were instead shaped by spatiotemporal factors. Nor was honey bee activity level impaired by grazing. The foraging preferences of bees, as well as flower species composition and peak bloom differed between grazed and ungrazed areas. Therefore, in our studied rangelands, grazing had its main effect on the foraging choices of honey bees and wild bees, rather than on their abundance and diversity. Moreover, our results indicate the potentially important role of ungrazed patches in increasing nectar and pollen diversity and availability in rangelands for both honey bees and wild bees in the spring. Hence, maintaining a mosaic of moderately grazed and ungrazed patches is expected to provide the greatest benefits for wild bee conservation and honey bee activity in Mediterranean rangelands. Our findings support the notion of rangeland sharing by cattle and bees in Mediterranean ecosystems under moderate grazing intensities, mimicking the coexistence of honey bees, wild bees, and cattle in Mediterranean ecosystems on an evolutionary timescale.

Quantifying Competitive Exclusion and Competitive Release in Ecological Communities: A Conceptual Framework and a Case Study.

A fundamental notion in community ecology is that local species diversity reflects some balance between the contrasting forces of competitive exclusion and competitive release. Quantifying this balance is not trivial, and requires data on the magnitude of both processes in the same system, as well as appropriate methodology to integrate and interpret such data. Here we present a novel framework for empirical studies of the balance between competitive exclusion and competitive release and demonstrate its applicability using data from a Mediterranean annual grassland where grazing is a major mechanism of competitive release. Empirical data on the balance between competitive exclusion and competitive release are crucial for understanding observed patterns of variation in local species diversity and the proposed approach provides a simple framework for the collection, interpretation, and synthesis of such data.

Climate change scenarios of herbaceous production along an aridity gradient: vulnerability increases with aridity.

Climate change is expected to reduce annual precipitation by 20% and increase its standard deviation by 20% in the eastern Mediterranean. We have examined how these changes may affect herbaceous aboveground net primary production (ANPP) and its inter-annual coefficient of variation (CV) in natural rangelands along a desert-Mediterranean precipitation gradient, at five sites representing arid, semi-arid, and Mediterranean-type ecosystems, respectively, all showing positive linear relationships between herbaceous ANPP and annual precipitation. Scenarios of reduced annual precipitation and increased inter-annual precipitation variability were defined by manipulating mean annual precipitation (MAP) and its standard deviation. We simulated precipitation and calculated ANPP using current ANPP-precipitation relationships. Our model predicts that reduced precipitation will strongly reduce ANPP in arid and semi-arid sites. Moreover, the effect of reduced precipitation on the CV of ANPP along the entire gradient may be modified by changes in inter-annual variability in MAP. Reduced precipitation combined with increased precipitation variability was the scenario most relevant to the wet end of the gradient, due to the increased likelihood for both dry and rainy years. In contrast, the scenario most relevant to the arid end of the gradient combined reduced precipitation with decreased precipitation variability, due to the strong effect on mean ANPP. All scenarios increased variability of ANPP along the entire gradient. However, the higher sensitivity of vegetation at arid and semi-arid sites (i.e., lower forage production) to future changes in the precipitation regime emphasizes the need to adapt grazing management in these ecosystems to secure their long-term viability as sustainable rangelands.

Testing the limits of resistance: a 19-year study of Mediterranean grassland response to grazing regimes.

A synthesis of a long-term (19 years) study assessing the effects of cattle grazing on the structure and composition of a Mediterranean grassland in north-eastern Israel is presented, with new insights into the response of the vegetation to grazing management and rainfall. We hypothesized that the plant community studied would be resistant to high grazing intensities and rainfall variability considering the combined long history of land-use and unpredictable climatic conditions where this community evolved. Treatments included manipulations of stocking densities (moderate, heavy, and very heavy) and of grazing regimes (continuous vs. seasonal), in a factorial design. The effect of interannual rainfall variation on the expression of grazing impacts on the plant community was minor. The main effects of grazing on relative cover of plant functional groups were related to early vs. late seasonal grazing. Species diversity and equitability were remarkably stable across all grazing treatments. A reduction in tall grass cover at higher stocking densities was correlated with increased cover of less palatable groups such as annual and perennial thistles, as well as shorter and prostrate groups such as short annual grasses. This long-term study shows that interannual fluctuations in plant functional group composition could be partly accounted for by grazing pressure and timing, but not by the measured rainfall variables. Grazing affected the dominance of tall annual grasses. However, the persistence of tall grasses and more palatable species over time, despite large differences in grazing pressure and timing, supports the idea that Mediterranean grasslands are highly resistant to prolonged grazing. Indeed, even under the most extreme grazing conditions applied, there were no signs of deterioration or collapse of the ecosystem. This high resistance to grazing intensity and interannual fluctuation in climatic conditions should favor the persistence of the plant community under forecasted increasing unpredictability due to climate change.

Quantifying drylands' drought resistance and recovery: the importance of drought intensity, dominant life history and grazing regime.

Projected global change will increase the level of land-use and environmental stressors such as drought and grazing, particularly in drylands. Still, combined effects of drought and grazing on plant production are poorly understood, thus hampering adequate projections and development of mitigation strategies. We used a large, cross-continental database consisting of 174 long-term datasets from >30 dryland regions to quantify ecosystem responses to drought and grazing with the ultimate goal to increase functional understanding in these responses. Two key aspects of ecosystem stability, resistance to and recovery after a drought, were evaluated based on standardized and normalized aboveground net primary production (ANPP) data. Drought intensity was quantified using the standardized precipitation index. We tested effects of drought intensity, grazing regime (grazed, ungrazed), biome (grassland, shrubland, savanna) or dominant life history (annual, perennial) of the herbaceous layer to assess the relative importance of these factors for ecosystem stability, and to identify predictable relationships between drought intensity and ecosystem resistance and recovery. We found that both components of ecosystem stability were better explained by dominant herbaceous life history than by biome. Increasing drought intensity (quasi-) linearly reduced ecosystem resistance. Even though annual and perennial systems showed the same response rate to increasing drought intensity, they differed in their general magnitude of resistance, with annual systems being ca. 27% less resistant. In contrast, systems with an herbaceous layer dominated by annuals had substantially higher postdrought recovery, particularly when grazed. Combined effects of drought and grazing were not merely additive but modulated by dominant life history of the herbaceous layer. To the best of our knowledge, our study established the first predictive, cross-continental model between drought intensity and drought-related relative losses in ANPP, and suggests that systems with an herbaceous layer dominated by annuals are more prone to ecosystem degradation under future global change regimes.

Competitive exclusion, beta diversity, and deterministic vs. stochastic drivers of community assembly.

Species diversity has two components - number of species and spatial turnover in species composition (beta-diversity). Using a field experiment focusing on a system of Mediterranean grasslands, we show that interspecific competition may influence the two components in the same direction or in opposite directions, depending on whether competitive exclusions are deterministic or stochastic. Deterministic exclusions reduce both patch-scale richness and beta-diversity, thereby homogenising the community. Stochastic extinctions reduce richness at the patch scale, but increase the differences in species composition among patches. These results indicate that studies of competitive effects on beta diversity may help to distinguish between deterministic and stochastic components of competitive exclusion. Such distinction is crucial for understanding the causal relationship between competition and species diversity, one of the oldest and most fundamental questions in ecology.

Inference of the activity timeline of cattle foraging on a Mediterranean woodland using GPS and pedometry.

The advent of the Global Positioning System (GPS) has transformed our ability to track livestock on rangelands. However, GPS data use would be greatly enhanced if we could also infer the activity timeline of an animal. We tested how well animal activity could be inferred from data provided by Lotek GPS collars, alone or in conjunction with IceRobotics IceTag pedometers. The collars provide motion and head position data, as well as location. The pedometers count steps, measure activity levels, and differentiate between standing and lying positions. We gathered synchronized data at 5-min resolution, from GPS collars, pedometers, and human observers, for free-grazing cattle (n = 9) at the Hatal Research Station in northern Israel. Equations for inferring activity during 5-min intervals (n = 1,475), classified as Graze, Rest (or Lie and Stand separately), and Travel were derived by discriminant and partition (classification tree) analysis of data from each device separately and from both together. When activity was classified as Graze, Rest and Travel, the lowest overall misclassification rate (10%) was obtained when data from both devices together were subjected to partition analysis; separate misclassification rates were 8, 12, and 3% for Graze, Rest and Travel, respectively. When Rest was subdivided into Lie and Stand, the lowest overall misclassification rate (10%) was again obtained when data from both devices together were subjected to partition analysis; misclassification rates were 6, 1, 26, and 17% for Graze, Lie, Stand, and Travel, respectively. The primary problem was confusion between Rest (or Stand) and Graze. Overall, the combination of Lotek GPS collars with IceRobotics IceTag pedometers was found superior to either device alone in inferring animal activity.