The influence of soil depth on plant species response to grazing within a semi-arid savanna

Grassland patches within a semi-arid savanna were evaluated over 45-years for (1) local temporal dynamics of basal area for five dominant grass species within long-term heavily grazed and ungrazed treatments, (2) the influence of soil depth (resource availability) on vegetation dynamics, and (3) the applicability of community-level grazing response groups over fine-scale patterns of soil heterogeneity. Temporal patterns in species composition and basal area were dependent upon soil depth. In the heavy grazed treatment, Hilaria belangeri dominated deep soils while Erioneuron pilosum and Bouteloua trifida were restricted to shallow soils. In the ungrazed treatment, removal of grazing resulted in successional changes that were significantly different across soil depths. After 45 years without grazing, Eriochloa sericea was most abundant on deep soils while Bouteloua curtipendula was more abundant on intermediate and shallow soils. Community-level functional groups that are based on grazing were not appropriate when multiple pattern-driving variables were considered across multiple scales indicating that functional groups should only be applied to certain processes at specific scales. Within the ungrazed treatments, variable soil depths have resulted in a shifting mosaic in time and space where early- and late-successional species co-exist continuously but spatially separated within the community. In the heavily grazed treatment, species are somewhat spatially arranged by soil depths, but much of the inherent heterogeneity is eliminated and species composition is dominated by the three grazing-resistant short-grasses. Broad scale successional changes may appear linear and predictable while at finer scales, the same changes may be described as non-linear and dependent upon soil depth resulting in thresholds that are partially explained by weather patterns, seed bank limitations and competitive inhibitions.     

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.     

Grassland composition affects season shifts in seed preference by Pogonomyrmex barbatus (Hymenoptera: Myrmicinae) in the Edwards Plateau, Texas

The effects of season and community composition as generated by livestock herbivory of differing intensity on seed species preference by Pogonomyrmex barbatus (F. Smith) were studied in a semiarid savanna on the Edward's Plateau, TX. Seasonal differences in nutrient requirements of the colony could lead to differential preferences for seeds harvested in spring and fall. Field cafeteria studies were conducted to test the hypothesis that late successional species, with their high nutrient content, would be chosen regardless of grazing intensity or season. Commercial seeds of known nutrient content were used to test the hypothesis that high protein levels would be chosen in spring and high carbohydrate levels in the fall. Naturally occurring seeds were differentially harvested and some were preferred regardless of relative availability. Total seed harvest in cafeteria experiments was higher in spring than in fall. Commercial seeds were harvested equally among treatments within a season; thus, nutrient selection was indistinguishable. Preference for native species was significantly different in both seasons but was influenced by a significant interaction with grazing treatments. Bouteloua curtipendula, a late successional mid-grass, was harvested significantly more in the spring than the fall and at higher rates in the heavily grazed treatment, rejecting the hypothesis that they would be chosen regardless of treatment or season. Seed preference for late successional grasses within heavily grazed communities may slow succession after grazing. During disturbance recovery, late successional species may be reduced by forager preference and rates of spring harvest.     

The impact of herbivores on nitrogen mineralization rate: consequences for salt-marsh succession

Soil net N-mineralization rate was measured along a successional gradient in salt-marsh sites that were grazed by vertebrate herbivores, and in 5-year-old exclosures from which the animals were excluded. Mineralization rate was significantly higher at ungrazed than at grazed sites. In the absence of grazing, mineralization rate increased over the course of succession, whereas it remained relatively low when sites were grazed. The largest differences in mineralization rate between grazed and ungrazed sites were found at late successional stages where grazing pressure was lowest. The amount of plant litter was significantly lower at grazed sites. In addition, the amount of litter and potential litter (non-woody, live shoots) was linearly related to net N-mineralization rate. This implies that herbivores reduced mineralization rate by preventing litter accumulation. Bulk density was higher at grazed salt-marsh sites than at ungrazed sites. This factor may also have contributed to the differences in net N-mineralization rate between grazed and ungrazed sites. Received: 30 November 1997 / Accepted: 27 August 1998     

Does selective defoliation mediate competitive interactions in a semiarid savanna? A demographic evaluation

Abstract. Three patterns of target-neighbor plant defoliation were imposed on a late-seral, perennial, C₄-grass, Bouteloua curtipendula, in three long-term grazing regimes to determine the influence of selective defoliation on competitive interactions and species replacement in a semiarid savanna on the Edwards Plateau, Texas, USA. Short-term (3-yr) target plant defoliation did not significantly affect either tiller or plant responses in any of the three grazing regimes. Neighbor plant defoliation, either alone or in combination with target plants, produced a significant defoliation interaction with time for tiller number and basal area per plant, but not for tiller recruitment or mortality. The minimal effect of selective defoliation on the intensity of competitive interactions in this semiarid community indicates that selective grazing has a less definitive role in mediating herbivore-induced species replacement than it does in mesic grasslands and savannas. This interpretation is discussed within the context of long-term (45-yr) change in herbaceous vegetation associated with grazing in this community. Cumulative tiller recruitment in the intensively grazed regime was only 44% of that in the ungrazed regime because of greater plant mortality and fewer surviving plants that recruited tillers. Target plant mortality (50%) only occurred in the intensively grazed regime and the proportion of target plants that initiated tillers decreased by 70, 48 and 32% in the ungrazed, moderately and intensively grazed regimes, respectively, during the final two years of the investigation. The decrease in cumulative tiller recruitment in all grazing regimes was probably mediated by a drought-induced increase in median tiller age the second year of the study. However, tiller per tiller recruitment rate among plants that recruited at least one tiller remained relatively constant among grazing regimes and years. Intensive, long-term grazing has modified the population structure of this late-seral perennial grass to the extent that population responses to both herbivory and periodic drought have been altered in comparison with those of ungrazed and moderately grazed populations. Ecological consequences of a herbivore-induced transition in population structure may be to minimize the effect of selective herbivory on competitive interactions and to function as an avoidance mechanism to reduce the probability of localized population extinction in response to intensive long-term herbivory.     

Simulated dynamics of succession in a North American subtropical Prosopis savanna

Abstract. A transition matrix model was used to explore the dynamics, rate and potential extent of changes in landscape vegetation patterns on a southern Texas Prosopis savanna. Transitions between seven vegetation classes were determined for the periods 1941–1960 and 1960–1983 on aerial photographs of three sites. During these periods, the sites were heavily grazed by cattle and were fire-free. Vegetation states assessed in grids of 20 m x 20 m cells superimposed on photographs ranged from grass-dominated to woody plant-dominated. The 1941–1960 period (denoted DRY) was characterized by prolonged drought, whereas annual rainfall during the 1960–1983 period (denoted WET) was typically normal to above-normal. The 1941 landscape consisted of herbaceous zones (6% of cells), woodland (50% of cells) and savanna parkland (44% of cells with grass/woody plant mixtures). The woodland state was the most stable, with probabilities of no change being 0.970 and 0.873 in WET and DRY periods, respectively. The herbaceous state was least stable, with corresponding values of 0.074 and 0.353. Past and future landscape structure was modelled by randomly selecting DRY or WET transitions at 20 year time steps. The model was run under a series of rainfall scenarios where the probability of selecting the WET transition matrix (P[WET]) ranged from 0 (DRY always chosen) to 1 (WET always chosen). Historical records indicate P[WET] has approximated 0.3 to 0.4 in the region. The rate of succession to states of greater woody cover increased as P[WET] increased. Forward simulations based on P[WET] > 0.2 suggest the present landscape is unstable and will develop into a closed-canopy woodland within the next 180 years, assuming the processes operating between 1941 and 1983 continue (e.g. grazing by cattle and lack of fire). Reverse simulations concur with historical observations and projections derived from woody plant growth rates in other studies and suggest that 200 to 300 yr BP these landscapes contained a substantially greater proportion of cells dominated by grassland or grassland with scattered woody plants (43 to 74%) than was present in 1983 (19%). Based upon elapsed time between predicted past and future steady states, succession from open savanna to closed-canopy woodland may occur in ca. 400 to 500 yr for P(WET) ≥ 0.33. Arresting or reversing the projected trend may require changes in climate and/or changes in livestock grazing and land management practices. The approaches employed in this study illustrate how time series maps, aerial photographs and satellite imagery can be analyzed and used to interpret, project and reconstruct local and regional changes in ecosystem structure. Difficulties and limitations associated with the use of Markov chains to model succession are identified and discussed.     

Scaling effects of grazing in a semi-arid grassland

Abstract. Previous studies have demonstrated relationships between spatial scale and spatial pattern and developed general hypotheses of scaling effects. Few studies, however, have examined the interactive relationship between scale and pattern-driving processes such as grazing. The goal of this study is to evaluate scale-dependent patterns across three spatial scales for three grazing intensities over 45 yr and to identify some mechanisms that may be associated with scale related differences. Correlation analysis and analysis of the coefficients of variation indicate that the relationships between units are dependent upon spatial scale and treatment. Across all grazing treatments, the relationship between units of the same scale becomes stronger as the spatial scale is increased. However, the rate of increase in the correlation coefficient is different for each treatment. The coefficient of variation responded inversely across scales with the greatest variation between small-scale units and little difference between the intermediate- and large scales. In addition to different relationships between units at each scale, differences in heterogeneity within treatments over time is illustrated by the relationship between small-scale units within each treatment and their associated larger scale units. The strongest relationship occurred in the heavily grazed treatments where correlation coefficients of small-scale units with intermediate- and large-scale units were ca. 0.60, indicating similar dynamics across scales. For the moderately grazed and ungrazed treatments this relationship varied from 0.40 to 0.47. Results from this study suggest that grazing alters scaling effects. Variability between small-scale units was greatest in the ungrazed treatment which had greater heterogeneity and less predictability than grazed treatments because of the influence of grazing on plant morphology, demography and composition. At the intermediate scale, relationships between units were fairly similar with the least variation occurring in the moderately grazed treatment. Alternatively, variation between large-scale units was greatest in the moderately grazed treatment because of the relationship between rest cycles, weather patterns, and patch grazing. Therefore, grazing can have a positive, a negative, or no influence on heterogeneity between units depending upon the scale of observation. Evaluation of long-term dynamics across these treatments at the same small spatial scale results in different variances within each treatment which may violate assumptions of some statistical and experimental designs. Therefore, evaluations of temporal dynamics should consider scale relative to the relationship between plant size, density and longevity (relative scale).     

Influence of Grazing on Soil Seed Banks Determines the Restoration Potential of Aboveground Vegetation in a Semi‐arid Savanna of Ethiopia

Species composition, number of emerging seedlings, species diversity and functional group of the soil seed banks, and the influence of grazing on the similarity between the soil seed banks and aboveground vegetation, were studied in 2008 and 2009 in a semi‐arid savanna of Ethiopia. We tested whether the availability of persistent seeds in the soil could drive the transition from a degraded system under heavy grazing to healthy vegetation with ample perennial grasses. A total of 77 species emerged from the soil seed bank samples: 21 annual grasses, 12 perennial grasses, 4 herbaceous legumes, 39 forbs, and 1 woody species. Perennial grass species dominated the lightly grazed sites, whereas the heavily grazed sites were dominated by annual forbs. Heavy grazing reduced the number of seeds that can germinate in the seed bank. Species richness in the seed bank was, however, not affected by grazing. With increasing soil depth, the seed density and its species richness declined. There was a higher similarity in species composition between the soil seed bank and aboveground vegetation at the lightly grazed sites compared with the heavily grazed sites. The mean similarity between the seed banks and aboveground vegetation was relatively low, indicating the effect of heavy grazing. Moreover, seeds of perennial grasses were less abundant in the soil seed banks under heavy grazing. We concluded that restoration of grass and woody species from the soil seed banks in the heavily grazed areas could not be successful in semi‐arid savannas of Ethiopia.     

Herbaceous vegetation change in variable rangeland environments: The relative contribution of grazing and climatic variability

Abstract. A 44‐yr record of herbaceous vegetation change was analysed for three contrasting grazing regimes within a semi‐arid savanna to evaluate the relative contribution of confined livestock grazing and climatic variability as agents of vegetation change. Grazing intensity had a significant, directional effect on the relative composition of short‐ and mid‐grass response groups; their composition was significantly correlated with time since the grazing regimes were established. Interannual precipitation was not significantly correlated with response group composition. However, interannual precipitation was significantly correlated with total plant basal area while time since imposition of grazing regimes was not, but both interannual precipitation and time since the grazing regimes were established were significantly correlated with total plant density. Vegetation change was reversible even though the herbaceous community had been maintained in an altered state for ca. 60 yr by intensive livestock grazing. However, ca. 25 yr were required for the mid‐grass response group to recover following the elimination of grazing and recovery occurred intermittently. The increase in mid‐grass composition was associated with a significant decrease in total plant density and an increase in mean individual plant basal area. Therefore, we failed to reject the hypotheses based on the proportional change in relative response group composition with grazing intensity and the distinct effects of grazing and climatic variability on response group composition, total basal area and plant density. Long‐term vegetation change indicates that grazing intensity established the long‐term directional change in response group composition, but that episodic climate events defined the short‐term rate and trajectory of this change and determines the upper limit on total basal area. The occurrence of both directional and non‐directional vegetation responses were largely a function of (1) the unique responses of the various community attributes monitored and (2) the distinct temporal responses of these community attributes to grazing and climatic variation. This interpretation supports previous conclusions that individual ecosystems may exist in equilibrial and non‐equilibrial states at various temporal and spatial scales.     

Composition of grazed and cleared temperate grassy woodlands in eastern Australia: patterns in space and inferences in time

Abstract. A regional vegetation survey of the temperate grassy woodlands (temperate savanna) in Australia was designed to assess the effects of clearing and grazing on the composition of vegetation remnants and the adjacent pasture matrix. Vegetation was sampled across a range of habitats using 77 0.1024‐ha quadrats; the relative abundance of species was recorded. Classification analysis clustered the sites into three main groups that corresponded to intensity of grazing/clearing followed by groups based on underlying lithology (basalt, metasediment, granites). Using Canonical Correspondence Analysis, exogenous disturbance and environmental variables were related to the relative abundance of species; grazing intensity had the highest eigenvalue (0.27) followed by tree canopy cover (0.25), lithology (0.18), altitude (0.17) and slope (0.10). Based on two‐dimensional ordination scores, six species response groups were defined relating to intensity of pastoralism and nutrient status of the landscape. Abundance and dominance of native shrubs, sub‐shrubs, twiners and geophytes were strongly associated with areas of less‐intense pastoralism on low‐nutrient soils. The strongest effects on species richness were grazing followed by canopy cover. Continuously grazed sites had lower native species richness across all growth forms except native grasses. There was no indication that intermediate grazing intensities enhanced forb richness as a result of competitive release. Species richness for all native plants was lowest where trees were absent especially under grazed conditions. Canopy cover in ungrazed sites appeared to promote the co‐existence of shrubs with the herbaceous layer. Predicted declines in forb richness in treeless, ungrazed, sites were not detected. The lack of a disturbance‐mediated enhancement of the herbaceous layer was attributed to habitat heterogeneity at 0.1 ha sampling scale.     

A hierarchical model for analysing the stability of vegetation patterns created by grazing in temperate pastures

Questions: Does vegetation structure display any stability over the grazing season and in two successive years, and is there any correlation between the stability of these spatial patterns and local sward composition? Location: An upland grassland in the French Massif Central. Method: The mosaic of short and tall vegetation stands considered as grazed and ungrazed patches respectively is modeled as the realization of a Boolean process. This method does not require any arbitrarily set sward‐height thresholds to discriminate between grazed and ungrazed areas, or the use of additional variables such as defoliation indexes. The model was validated by comparing empirical and simulated sward‐height distributions and semi‐variograms. Results: The model discriminated between grazed and ungrazed patches at both a fine (1 m²) and a larger (500 m²) scale. Selective grazing on legumes and forbs and avoidance of reproductive grass could partly explain the stability of fine‐scale grazing patterns in lightly grazed plots. In these plots, the model revealed an inter‐annual stability of large‐scale grazing patterns at the time peak biomass occurred. At the end of the grazing season, lightly grazed plots showed fluctuating patch boundaries while heavily grazed plots showed a certain degree of patch stability. Conclusion: The model presented here reveals that selective grazing at the bite scale could lead to the creation of relatively stable patches within the pasture. Locally maintaining short cover heights would result in divergent within‐plot vegetation dynamics, and thus favor the functional diversity of vegetation.     

Changes in the vegetation of Quercus pubescens woodland after cessation of coppicing and grazing

Abstract. The cessation of coppicing and grazing in Quercus woodlands, with its subsequent changes in the structure, composition and functioning of vegetation communities, is becoming more frequent throughout the Mediterranean Basin. In southern France, we have studied successional changes in Quercus pubescens woodlands by visiting previously studied sites 18 yr later. Changes in vertical structure, species richness, floristic composition, life form and dispersal type were analysed and compared between woodlands that had previously been grazed or ungrazed. Both successions showed a decline in vegetation cover in the 0–25 cm height class and in the height class immediately under the canopy layer, due to oak litter accumulation and tree ageing. In post‐grazing succession, the abandonment of grazing and associated burning has allowed the vegetation cover to increase in the 0.25‐2 m height class. In both successions, grassland species decreased in frequency and forest species increased, a trend which was stronger in undisturbed succession. Species richness decreased with time in the undisturbed succession, but remained stable in the post‐grazing succession mainly because of the slow decline of plants linked to grazing. In undisturbed succession, therophytes and hemicryptophytes decreased. In contrast, therophytes and hemicryptophytes remained stable in post‐grazing succession. In both successions, endozoochorous species (notably Ruscus aculeatus) increased. Plants dispersed by non‐animal vectors decreased in undisturbed succession, but stability was observed in most of the dispersal types in post‐grazing succession. These results showed that a time‐lag existed between undisturbed succession and post‐grazing succession, the latter remaining at a younger stage of successional development due to more recent impact of grazing. However, both successions have converged suggesting that most of the traces of grazing on vegetation will disappear within a few years. The vegetation of these coppices, regardless of the previous grazing regime, will become increasingly similar to the vegetation of undisturbed woodlands. However, their floristic composition will probably never be identical to that of undisturbed woodlands, mainly because of the rarity of these undisturbed woodlands and of the short‐distance dispersal of many forest plant species.     

Intensity of livestock grazing in relation to habitat use by brown hares (Lepus europaeus)

There is very limited information concerning livestock (sheep and goats) and brown hare Lepus europaeus interaction when both coexist. The effect of the intensity of livestock grazing on seasonal habitat use by hares, in a typical Mediterranean rangeland, was evaluated using the pellet-count method. Lightly grazed pastures were less preferred by hares compared with moderately grazed ones, whereas ungrazed pastures were used less intensively than grazed ones. Because livestock grazing reduces the quantity of standing biomass proportionally to its grazing intensity, forage resource was not the driving force for pasture selection. The increased use of moderately grazed pastures by hares in relation to lightly and ungrazed ones, where vegetation was more abundant, could be attributed to their reduced herbage height and density. This behaviour is probably a tactic that hares follow for predator avoidance, because they are more likely to detect visually approaching predators when feeding in a biotope with a limited herbaceous layer. The conclusion of this research is that livestock and brown hare coexistence may be compatible and beneficial rather than competitive when stocking rates do not exceed grazing capacity, leading to the conclusion that proper livestock grazing and hare population management can be feasible in practice.     

Effects of grazing,topography, and precipitation on the structure of a semiarid grassland

Structural aspects of the shortgrass steppe plant community, functional groups, and species populations were examined in response to long-term heavy grazing and exclosure from grazing, contiguous wet or dry years, and an environmental gradient of topography. Of the three factors, relatively greater differences in community similarity were observed between catena positions, particularly on the ungrazed treatments. Grazing was intermediate between catena position and short-term weather in shaping plant community structure. Grazed treatments and ridgetops had a less variable species composition through fluctuations in weather.An increase with grazing of the dominant, heavily grazed species was observed. Basal cover and density of total species was also greater on grazed sites. The more uniform grazing lawn structure of the grazed plant communities had an influence on segregation of plant populations along topographical gradients. Segregation was less on grazed catenas, but diversity and the abundance of introduced and opportunistic-colonizer species was also less.Although the shortgrass steppe community was relatively invariant, less abundant species were dynamic and interactions occurred with respect to grazing, weather, and catena position. The effects of grazing may be mitigated by favorable growing seasons but magnified in unfavorable years in populations that are adapted to favorable sites. Grazing can be considered a disturbance at the level of the individual but it may or may not be a disturbance at the level of the population, and it is not a disturbance at the level of the community in this particular grassland.     

Relationships between the effects of insect herbivory and sheep grazing on seasonal changes in an early successional plant community

Summary The effects of spring grazing by sheep and of natural levels of insect herbivory were studied in 1985 on a limestone field abandoned from arable land for four years. A split-plot design was adopted in which paddocks, arranged in Latin squares, were either left ungrazed or heavily grazed by sheep for ten days in April. Within each paddock plots were either sprayed regularly with Malathion-60 or untreated.Natural levels of insect herbivory, compared to the reduced levels in insecticide-treated plots, had effects of similar magnitude to those from the short burst of spring grazing. Many attributes of the grazed/insecticide-treated sward were either increased or decreased by a factor of two within a season. Both types of herbivore caused changes in the direction of plant succession as well as in its rate. Effects on early successional species were large and similar when caused by either type of herbivore. Effects on later successional species were often smaller, but also showed differences in the action of the two herbivore types, as did effects on sward height, species richness and total cover. The effects of sheep and insect herbivory were not always additive or in the same direction.The results suggest that manipulations of both mammal and insect herbivores may be powerful tools for directing changes in plant community composition.     

Grazing as a mediator for maintenance of offspring diversity: Sexual and clonal recruitment in alpine grassland communities

To understand the effects of grazing on grassland plants sexual and clonal recruitment, we conducted a demographic field investigation of species recruitment along a grazing gradient in the Tibetan alpine grassland. Grazing intensity had significant effects on quantity and diversity of sexual and clonal recruitment. Sexual recruitment increased significantly, but clonal offspring production decreased significantly with increased grazing intensity. Grazing intensity had different, significant effects on offspring recruitment of the various functional groups in the community, grasses (GG), sedges (SG), legumes (LG) and forbs (FG). Higher grazing intensity reduced offspring recruitment of GG and SG; it increased offspring recruitment of LG and FG. Seedlings were significantly more abundant in lightly grazed, moderately grazed and heavily grazed meadows than in non-grazed grasslands. Offspring diversity from sexual recruitment was significantly higher than that from clonal recruitment in grazed than in non-grazed grasslands. Our studies indicate that moderate grazing had positive effects on seedling recruitment and offspring diversity, but heavy gazing may alter community succession by affecting recruitment patterns among the four plant functional groups.     

Managing open habitats by wild ungulate browsing and grazing: A case‐study in North‐Eastern Germany

Question: Can wild ungulates efficiently maintain and restore open habitats? Location: Brandenburg, NE Germany. Methods: The effect of wild ungulate grazing and browsing was studied in three successional stages: (1) Corynephorus canescens‐dominated grassland; (2) ruderal tall forb vegetation dominated by Tanacetum vulgare; and (3) Pinus sylvestris‐pioneer forest. The study was conducted over 3 yr. In each successional stage, six paired 4 m²‐monitoring plots of permanently grazed versus ungrazed plots were arranged in three random blocks. Removal of grazing was introduced de novo for the study. In each plot, percentage cover of each plant and lichen species and total cover of woody plants was recorded. Results: Wild ungulates considerably affected successional pathways and species composition in open habitats but this influence became evident in alteration of abundances of only a few species. Grazing effects differed considerably between successional stages: species richness was higher in grazed versus ungrazed ruderal and pioneer forest plots, but not in the Corynephorus sites. Herbivory affected woody plant cover only in the Pioneer forest sites. Although the study period was too short to observe drastic changes in species richness and woody plant cover, notable changes in species composition were still detected in all successional stages. Conclusion: Wild ungulate browsing is a useful tool to inhibit encroachment of woody vegetation and to conserve a species‐rich, open landscape.     

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

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

Grassland dynamics under sheep grazing in an Australian Mediterranean type climate

Grassland dynamics in a degraded disclimax grassland dominated by Danthonia caespitosa Gaudich. are examined using both demographic and multivariate approaches in an experiment designed to determine the effect of grazing intensity and exclosure on pasture dynamics. The experiment ran for 20 years from 1949 to 1968, using permanent quadrats at 3 grazing intensities and within exclosures. Demographic studies of some perennial grass species demonstrated markedly different responses to grazing; Danthonia caespitosa was unaffected by grazing but responsive to seasonal rainfall differences. Enteropogon acicularis survived only on protected sites. Numerical classification of total species set (121 species) for six observation periods demonstrated that community types were sensitive to differences in winter rainfall, and time since the start of experiment. Principal component analysis of permanent quadrat observations for individual years demonstrates quadrat trajectories which confirm this and indicate progressive divergence of the successional trends of the grazed and ungrazed quadrats. Repeated analysis on grazed quadrats only, shows that three components of pasture dynamics can be recognized; these are trend (succession?) and seasonal differences, each of which account for about 20% of the variance, and differences due to soil heterogeneity in the experimental paddock (8% of variance accounted for). No effect of grazing intensity was detected. Multivariate techniques can provide a clear partitioning of types of dynamic behaviour present in grassland communities. It is concluded that partitioning of environmental heterogeneity prior to demographic studies would increase their sensitivity.

     

Effects of Grazing Intensity and Environmental Factors on Species Composition and Diversity in Typical Steppe of Inner Mongolia,China

In the present study, we aim to analyze the effect of grazing, precipitation and temperature on plant species dynamics in the typical steppe of Inner Mongolia, P.R. China. By uncoupling biotic and abiotic factors, we provide essential information on the main drivers determining species composition and species diversity. Effects of grazing by sheep were studied in a controlled experiment along a gradient of seven grazing intensities (from ungrazed to very heavily grazed) during six consecutive years (2005–2010). The results show that plant species composition and diversity varied among years but were little affected by grazing intensity, since the experimental years were much dryer than the long term average, the abiotic constraints may have overridden any grazing effect. Among-year differences were predominantly determined by the abiotic factors of precipitation and temperature. Most of the variation in species dynamics and coexistence between C3 and C4 species was explained by seasonal weather conditions, i.e. precipitation and temperature regime during the early-season (March-June) were most important in determining vegetation dynamics. The dominant C3 species Stipa grandis was highly competitive in March-June, when the temperature levels were low and rainfall level was high. In contrast, the most common C4 species Cleistogenes squarrosa benefited from high early-season temperature levels and low early-season rainfall. However, biomass of Stipa grandis was positively correlated with temperature in March, when effective mean temperature ranges from 0 to 5°C and thus promotes vernalization and vegetative sprouting. Our results suggest that, over a six-year term, it is temporal variability in precipitation and temperature rather than grazing that determines vegetation dynamics and species co-existence of grazed steppe ecosystems. Furthermore, our data support that the variability in the biomass of dominant species, rather than diversity, determine ecosystem functioning. The present study provides fundamental knowledge on the complex interaction of grazing – vegetation – climate.