Early changes in species composition of upland sown grassland under extensive grazing management

Abstract. Due to economic pressures and policy changes Lolium perenne‐Trifolium repens sown swards in upland UK sheep systems are likely to become less intensively managed. We present results from the first 5 yr of a long‐term experiment studying vegetation change under more extensive grazing management at three sites. One treatment was representative of current, intensive management and 5 were unfertilized with different intensities of seasonal grazing. The species composition of unfertilized, ungrazed swards changed dramatically within 2 yr and the sown species had virtually disappeared by year 5. Ranunculus repens, Poa trivialis, Agrostis gigantea, Juncus spp. and Carex spp. became dominant at the wettest site. Grasses were dominant at the other sites. In contrast, the sown species were retained in the unfertilized, grazed treatments; there were small shifts in abundance of the species present initially and few additions or losses of species. Some colonizing species were present in the seed bank whereas others with a transient seed bank appeared to have invaded from neighbouring vegetation. Implications of these results for compensation schemes to reduce animal output and increase biodiversity are discussed.     

The vertical distribution of below-ground biomass in grassland communities in relation to grazing regime and habitat characteristics

Abstract. 40 sites, representing different pasture types in Northwest Spain, were sampled in respect of their floristic composition, distribution of above and below-ground biomass and environmental and physical variables. Five plant community types were identified by classification techniques of plant species composition. These communities were then characterized in terms of the percentage of ground covered by herbaceous and shrub vegetation, stones, rocks and gaps as well as their topographic location and characteristics of the shallow soil (pH, organic matter, nitrogen and calcium content). Bio-mass was assessed in terms of above-ground structures, surface crowns and three below-ground layers to a depth of 10 cm. Three types of grazing regime were distinguished: Concentrated Intense Grazing in early spring (CIG), Extended Intense Grazing throughout the spring (EIG), and Non-Intense Grazing (NIG). Grazing regime showed the highest association with plant community type and three broad categories were identified: xeric stressed pastures, which nevertheless received CIG, mesic pastures with EIG, and three kinds of NIG mesic pastures. The xeric communities had the highest proportion of aboveground biomass, as a consequence of their greater proportion of woody perennials. These xeric communities displayed a more gradual reduction in below-ground biomass with depth than mesic pastures, a likely consequence of the low water content in the upper soil layers. The mesic communities had a high concentration of below-ground biomass in the upper layers when they were intensely grazed. However, when grazing was low (i.e. NIG situations), these communities had greater variability in biomass profiles than any of the other pasture types. Possible causes of the patterns in biomass distribution of the intensely grazed pastures are 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).     

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.     

Impact of grazing on vegetation dynamics in former ricefields

Abstract. In the Rhône delta, Juncus gerardi and Scirpus maritimus are often the dominant species in abandoned rice fields which are artificially flooded in early spring to improve forage production. Under these conditions they occur either in mixed communities, or form monospecific stands. Monitoring the vegetation dynamics in quadrats located in six abandoned rice fields artificially flooded from November to April confirmed the important role of grazing. In ungrazed plots, communities dominated by Scirpus maritimus mixed with Juncus gerardi developed fast. After 42 months of management Scirpus maritimus had established in nearly all quadrats and continued to expand, whereas Juncus gerardi had started to decline. In grazed plots Juncus gerardi alone dominated and continued to increase in cover up to the 42nd month. Scirpus maritimus established at low densities mainly in quadrats where Juncus gerardi was initially absent. Introduction of seeds of Scirpus maritimus in communities of Juncus gerardi under controlled conditions demonstrated the existence of the phenomenon of preemption. The increase in cover of Juncus gerardi suggests that the preemption of Juncus gerardi over Scirpus maritimus plays a more pronounced role in the field in the presence of grazing.     

Long-term vegetation dynamics mediated by herbivores,weather and fire in a Juniperus-Quercus savanna

Abstract. Long-term (45-yr) basal area dynamics of dominant graminoid species were analyzed across three grazing intensity treatments (heavily grazed, moderately grazed and ungrazed) at the Texas A&M University Agricultural Research Station on the Edwards Plateau, Texas. Grazing intensity was identified as the primary influence on long-term variations in species composition. Periodic weather events, including a severe drought (1951–1956), had little direct influence on composition dynamics. However, the drought interacted with grazing intensity in the heavily grazed treatment to exacerbate directional changes caused by grazing intensity. Species response to grazing was individualistic and noisy. Three response groups were identified. Taller, more productive mid-grasses were most abundant under moderate or no grazing. Short grasses were most abundant under heavy grazing. Intermediate species were most abundant under moderate grazing and opportunistic to weather patterns. Graminoid diversity increased with the removal or reduction of grazing intensity. The moderately and ungrazed treatments appeared most resistant to short-term weather fluctuations, while the heavily grazed treatment demonstrated significant resilience when grazing intensity was reduced after over 110 yr of overgrazing. Identification of a ‘climax’ state is difficult. Significant directional change, which took nearly 20 yr, appears to continue in the ungrazed treatment after 45 yr of succession. The observed, relatively linear patterns of perennial grass composition within the herbaceous patches of this savanna were generally explained by traditional Clementsian succession. However, when dynamics of the herbaceous community are combined with the woody component of this savanna, the frequency and intensity of fire becomes more important. Across the landscape, successional changes follow several pathways. When vegetation change is influenced by several factors, a multi-scale model is necessary to demonstrate interactions and feedbacks and accurately describe successional patterns. Absence of fires, with or without grazing, leads ultimately to a Juniperus/Quercus woodland with grazing intensity primarily influencing the fuel load and hence fire intensity.     

Dry grassland plant diversity conservation using low-intensity sheep and goat grazing management: case study in Prague (Czech Republic)

After abandonment, dry grassland (Festuco-Brometea) areas decline due to gradual overgrowing by woody species and the expansion of perennial tall grass species. Dry grassland vegetation was formed by extensive livestock grazing, thus grazing is considered one of the most natural methods for managing this type of vegetation. Six years after introducing low-intensity sheep and goat grazing in seven nature reserves in Prague (Czech Republic), the following impact of this management on dry grassland vegetation was observed: The cover of expansive woody species, particularly Ligustrum vulgare, and to a smaller extent Cornus sanguinea and Prunus spinosa declined. In addition, a significant, long-term declining trend of the expansive species Arrhenatherum elatius was also observed. Also the cover of Pimpinella saxifraga and Allium senescens declined significantly with regard to statistical evaluation. On the contrary, the cover of Achillea millefolium, Centaurea stoebe, Securigera varia, Elytrigia repens, Erysimum crepidifolium, Falcaria vulgaris, Fallopia convolvulus and Verbascum lychnitis increased. The cover of species characteristic of dry grasslands (Festuco-Brometea) increased significantly. No changes were observed in the number and cover of the Red List species. In addition, the presence of nitrophilous and ruderal species increased. Species diversity also significantly increased. From our findings we can conclude that managing dry grasslands with low-intensity grazing can help to keep dry grassland vegetation in good condition and conserve its plant diversity. Nomenclature: Kubát et al. (2002) for taxa and Moravec et al. (1995) for syntaxa.     

Impact of different sheep grazing intensities on salt marsh vegetation in northern Germany

Abstract. Mainland salt marshes in Schleswig-Holstein (northern Germany) have been grazed intensively by sheep for several decades. In 1988 experimental sites were established in the lower and middle salt marsh of Sönke-Nissen-Koog and subjected to different grazing intensities. From 1989 to 1993 the impact of sheep grazing on the composition and structure of the salt marsh vegetation was studied through the yearly analysis of permanent plots, vegetation mapping and measurements of the vegetation height. The intensively grazed site (10 sheep/ha) is covered by a short monotonous Puccinellia maritima sward with Salicornia europaea and Suaeda maritima. Halimione portulacoides and Aster tripolium, especially flowering plants, are rare. On the sites with 1.5 and 3 sheep/ha Puccinellia maritima remained dominant. The population density of Salicornia europaea decreases after reduction of the grazing intensity, whereas Suaeda maritima finds optimal growing conditions. Stands of Halimione portulacoides and flowering Aster tripolium plants are rare near the sea dike but their cover and size increases further away from the dike. In the plot with 1.5 sheep/ha the height of the vegetation increases along the gradient from the dike towards the tidal flats, due to local differences in actual grazing intensity. When grazing is stopped, Puccinellia maritima is successively replaced by Festuca rubra, Halimione portulacoides and Aster tripolium. High variability of vegetation height indicates structural diversity. Patches of higher and lower vegetation correspond with the distribution pattern of different plant species. In terms of nature conservation cessation of grazing is recommended.     

The effect of horse, cattle and sheep grazing on the diversity and abundance of land snails in nutrient-poor calcareous grasslands

Livestock grazing is a common management practise in semi-natural grasslands in Central Europe. Different types of livestock (horses, cattle, sheep) and grazing intensity are known to affect the richness and composition of plant species. However, knowledge of grazing-dependent effects on invertebrates is limited. We examined the influence of horse, cattle and sheep grazing on the richness, abundance and composition of land snail species in 21 calcareous nutrient-poor grassland areas in the northwestern Jura Mountains, Switzerland. Grazing by different livestock species did not affect the species richness, abundance and species composition of land snails. Furthermore, the number of open-land species and the ratio of large- to small-sized snail species or individuals did not differ among the three pasture types. However, independent of livestock species, grazing intensity negatively influenced the snail fauna. Snail species richness, abundance and number of Red list species decreased with increasing grazing intensity. Grazing intensity also affected the occurrence of individual snail species (Truncatellina cylindrica, Cecilioides acicula, Candidula unifasciata and Trichia plebeia). To preserve the snail fauna in nutrient-poor grasslands, pastures can be stocked with horses, cattle or sheep. However, both maximum stocking rate (number of livestock units per hectare) and grazing duration (number of grazing days per year) must be carefully defined for the proper management of the pastures.     

Effects of reindeer grazing on understorey vegetation in dry Pinus sylvestris forests

Abstract. Data on floristic composition and environmental variables were collected in floristically homogeneous oligotrophic pine (Pinus sylvestris) forests with heath-like under- storey vegetation in eastern Fennoscandia, and ordinated by non-linear multidimensional scaling (NMDS) in order to study the effect of lichen grazing by reindeer on the understorey vegetation. The study sites included areas with varying grazing pressure, as well as 50-yr old grazing exclosures. Sites rich in respectively bryophytes and lichens were placed at opposite ends of the ordination axes, and heavily grazed sites were placed in between them. Reindeer grazing increased the abundance of bryophytes, especially Dicranum spp. and Pleurozium schreberi. Grazing changed the vegetation to the extent that it resembled more mesotrophic sites, but this did not show any relationship with tree volume or other site productivity indicators. This was observed both in the ordination and, in a more compelling way, when exclosures with adjacent grazed areas were compared. No such signs were evident at ungrazed sites, where especially Cladina spp. spatially replace Cladonia spp. and tiny bryophytes like Barbilophozia spp., Polytrichum spp. and Pohlia nutans during succession. Cladina stellaris had almost disappeared from the most intensively grazed sites. The soil at ungrazed sites was characterized by high Al and Fe concentrations and bryophyte-rich sites by high Mn concentrations. Shannon's diversity index, depth of humus layer and proportion of bare ground also increased in sites getting richer in bryophytes.     

A vegetation analysis of the pastoral landscapes of upland Wales,UK

Abstract. The upland moorlands of Wales are situated on the oceanic fringe of western Europe, and have experienced a long history of pastoral management. Recent vegetation data are analysed to assess the relative contribution of abiotic and anthropogenic factors to variation in habitat composition among the major upland ranges of this region. From a numerical analysis of plant community cover data, recorded from 65 sites covering 260 000 ha, a six‐cluster site classification emerged with striking biogeographical coherence. Direct gradient analysis and variance partitioning revealed strong correlation between vegetation composition and spatially‐structured climatic gradients, in particular temperature, rainfall and oceanicity; differences in bedrock geology appear to have a lesser role. The analysis also indicates a close correlation between habitat variation and anthropogenic parameters, especially grazing intensity, burning frequency, and sulphur and nitrogen deposition levels. At this regional scale, anthropogenic impacts appear to have accentuated, rather than obscured, vegetation patterns which are primarily determined by climate and other abiotic variables. The findings have considerable relevance for conservation planning and also for predictive studies on the consequences of climatic change for the biota of the uplands of southern Britain.     

Restoration of high species density in calcareous grassland: the role of seed rain and soil seed bank

Abstract. A small-scale restoration experiment concerning a species-rich calcareous grassland was conducted over a 7-yr period. 20 yr of abandonment (1971-1990) resulted in both a strong reduction of the number of species and an absolute dominance of the shrub Crataegus monogyna and the grass Brachypodium pinnatum - which contributed 98% of the above-ground herb phytomass. The plot (3 m x 6 m) was cleared in September 1990 and thereafter yearly mown in autumn. Small soil seed bank samples were taken immediately after clearing and again in the autumn of 1996. The inner part of the plot showed a reduced soil seed bank compared to the plot underneath the vegetation outside the plot in 1990. Six years later an increased number of seeds in the soil in this inside area could be recorded mainly in the upper soil layer, although the amount was only 40% compared to the outside or control area. Ca. 20% of the species present in the restored vegetation were recorded in seed traps during the main seed dispersal period in 1997. Restoration of high species richness in the plot was a fast process during the first few years: from seven species in 1990 towards 39 three years later. This was significantly faster than the restoration process of the calcareous grassland at the same slope 20 yr earlier. This was probably due to the increased size of the community species pool at the site as a result of appropriate management during the last decades. Relative species number increase was highest at the largest scale during this experiment, viz. at the total plot area. The success of restoration of species-rich, dry grassland vegetation largely depends on the interaction between several conditions, e.g. seed availability, either originating from the seed rain or soil seed bank, favourable germination conditions and a successful development to a mature, seed producing individual. Germination and establishment strongly depend on suitable vegetation structure as a result of sustainable management.     

Effects of litter accumulation on riparian vegetation: Importance of particle size

Abstract. The floristic effects of river‐borne litter that accumulates in riparian zones may vary in space and time depending on variations in mass and particle size of the deposited litter. To analyse the effects of litter mass and size we applied differentsized litter (natural uncut pieces and powder) to riparian vegetation at different quantities. Vegetation responses were analysed after one season at the community level (total biomass or richness for all species) and species traits (biomass or richness for groups of species). At the community level uncut litter, but not powder, reduced species richness and both uncut and ground litter reduced above‐ground biomass. At the species trait level uncut litter had a stronger effect than powder on species richness and biomass. The only positive effect of litter addition was that powder increased graminoid species richness. The topsoil conditions indicated that the major impact of deposited, river‐borne litter was that it acted as a physical barrier directly preventing established plants from penetrating the litter layer and reducing light and soil temperature.     

Formalized reproduction of an expert‐based phytosociological classification: A case study of subalpine tall‐forb vegetation

Abstract. Delimitation of vegetation units in phytosociology is traditionally based on expert knowledge. Applications of expert‐based classifications are often inconsistent because criteria for assigning relevés to vegetation units are seldom given explicitly. Still, there is, e.g. in nature conservation, an increasing need for a consistent application of vegetation classification using computer expert systems for unit identification. We propose a procedure for formalized reproduction of an expert‐based vegetation classification, which is applicable to large phytosociological data sets. This procedure combines Bruelheide's Cocktail method with a similarity‐based assignment of relevés to constancy columns of a vegetation table. As a test of this method we attempt to reproduce the expert‐based phytosociological classification of subalpine tall‐forb vegetation of the Czech Republic which has been made by combination of expert judgement and stepwise numerical classification of 718 relevés by TWINSPAN. Applying the Cocktail method to a geographically stratified data set of 21794 relevés of all Czech vegetation types, we defined groups of species with the statistical tendency of joint occurrences in vegetation. Combinations of 12 of these species groups by logical operators AND, OR and AND NOT yielded formal definitions of 14 of 16 associations which had been accepted in the expert‐based classification. Application of these formal definitions to the original data set of 718 relevés resulted in an assignment of 376 relevés to the associations. This assignment agreed well with the original expert‐based classification. Relevés that remained un‐assigned because they had not met the requirements of any of the formal definitions, were subsequently assigned to the associations by calculating similarity to relevé groups that had already been assigned to the associations. A new index, based on frequency and fidelity, was proposed for calculating similarity. The agreement with the expert‐based classification achieved by the formal definitions was still improved after applying the similarity‐based assignment. Results indicate that the expert‐based classification can be successfully formalized and converted into a computer expert system.     

Establishment of Picea abies seedlings in a central European mountain grassland: an experimental study

Abstract. Secondary meadows in the Kru?né hory mountains, central Europe, are not colonized by woody species, although they have been abandoned for half a century and are surrounded by Picea abies (Norway spruce) forests. The causes of inhibition of establishment of Norway spruce seedlings in the meadows were tested experimentally. The experiment was started in a masting year to ensure sufficient seed input. Four treatments (vegetation cut; vegetation cut and litter removed; all above-ground biomass and topsoil removed; control) were combined in a factorial design. The effect of browsing, mainly by deer, was assessed by fencing half of the experimental plots. Seedling establishment differed among treatments (all pair-wise differences were significant) but was independent of fencing. In contrast, seedling survival was influenced both by the treatment and fencing. At the end of the second year, only the plots with all biomass and organic topsoil removed supported viable populations of spruce seedlings. Dense herb cover and a thick layer of slowly decaying litter are considered the main factors inhibiting the establishment of woody species.     

放牧、刈割及摘顶对亚热带人工草地牧草种群的影响

通过比较放牧、刈割及摘顶3种干扰发现,适度放牧、刈割提高了亚热带高山人工草地鸭茅、黑麦草种群的适应性,部分原因在于2种干扰均对牧草进行了摘顶．摘顶解除了牧草的顶端优势及生殖生长。放牧、刈割对牧草产生的生态影响在某种程度上可通过摘顶而实现。从而提高种群密度、热值及能量积累．单因子的人工摘顶试验表明,牧草顶端优势的维持降低了牧草种群密度,抑制了能量积累。从抑制了种间竞争．持续摘顶解除了牧草的顶端优势。促进了牧草密度及能量的增长,从而提高了牧草种群的竞争力．     

Patch colonisation in a savanna grassland

Patches (1 m diameter) were cleared in a heavily grazed and a lightly grazed savanna in South Africa; half of them were covered once with grass litter. The greatest colonisation over three years occurred on heavily grazed patches with litter, the least on open, lightly grazed patches. Annual rainfall affected colonisation rate. Basal cover was lower on patches than in the surrounding vegetation. Patch composition was weakly affected by treatment, and was partly related to the composition of adjacent vegetation, but the proportional representation of species on patches differed from the surrounding vegetation. The stoloniferous Digitaria eriantha and the obligate seed reproducers Aristida bipartita and Heteropogon contortus were major components of patch communities but were uncommon in the surrounding vegetation and in the seed bank. Setaria incrassata and Themeda triandra were the predominant components of the surrounding vegetation and of the seed bank, but Setaria established very poorly and Themeda established well only on heavily grazed patches with litter. The pattern of seedling establishment was the same as that of colonisation. Most seedlings emerged atthe beginning of the wet season, with ca. 50% mortality soon thereafter. The colonising species can be partly predicted from the availability of propagules (vegetative or seed), emergence and establishment success, and subsequent growth.     

Vegetation change following exclusion of grazing animals in depleted grassland,Central Otago,New Zealand

Abstract. Models of semiarid vegetation dynamics were evaluated to explain changes in the grassland of interior South Island, New Zealand. Annual records were taken for six years of plant species height frequency and percentage ground cover in five plots established in 1986. One subplot at each site was fenced to exclude sheep, one to exclude rabbits and sheep, and one remained unfenced as a control. Records from 1986–1992 were analysed by ordination. The overall pattern of vegetation change shows considerable year-to-year variation. At some sites, variation in vegetation composition between years was as great as, or greater than, that between grazed and ungrazed subplots. Such variation is particularly evident in grazed vegetation, perhaps because it is under greater stress than ungrazed vegetation. At one site changes in vegetation total cover and species composition could be statistically related to rainfall during the first half of the growing season. The only general trends following cessation of grazing were for perennials to increase in frequency, and for year-to-year changes to become smaller with time. Total vegetation cover values seldom changed as a result of cessation of grazing, but tended to follow year-to-year changes in species frequency. The results do not in general support switch/state-and-transition models of semi-arid vegetation dynamics. Vegetation change follows changes in grazing and climate with little lag. This most closely conforms with the Pulse-phase dynamic model.