Different responses of plants and herbivore insects to a gradient of vegetation height: an indicator of the vertebrate grazing intensity and successional age

Empirical studies have suggested that species richness of grassland insects usually decreases under grazing management. By contrast, grazing has been shown to increase the species density and richness of vascular plants, especially on productive soils. In order to test the suggested differences in response to management between plants and insects, we simultaneously studied species richness of vascular plants and their insect herbivores, butterflies and moths, in 68 semi-natural grasslands with varying grazing intensity and frequency in SW Finland. Species richness of plants and insects was for the first time related to a quantitative measure of disturbance intensity and successional age, mean vegetation height, by using generalized additive models (GAM). The effects of soil nutrients on vegetation height were accounted for by using phosphorus concentration as a productivity surrogate.
The results showed that species richness of butterflies and moths peaked in taller vegetation compared with vascular plants, corresponding to a lower disturbance intensity and increasing time since abandonment. These patterns are discussed in the light of two hypotheses, the "structural diversity hypothesis" and "dynamic equilibrium model" of Huston, both suggesting a weaker disturbance tolerance of insects compared with plants. Butterflies and moths which are specialists in their larval host-plant use (monophagous and oligophagous species) preferred lower vegetation (higher disturbances) compared with generalists (polyphagous species), as predicted by Huston's model. This difference indicates a stronger relationship with plant species richness for specialist than for generalist butterflies and moths. Our results support the application of regionally and temporally varying grazing intensities in grassland conservation management.     

Does resource availability,resource heterogeneity or species turnover mediate changes in plant species richness in grazed grasslands?

Grazing by large ungulates often increases plant species richness in grasslands of moderate to high productivity. In a mesic North American grassland with and without the presence of bison (Bos bison), a native ungulate grazer, three non-exclusive hypotheses for increased plant species richness in grazed grasslands were evaluated: (1) bison grazing enhances levels of resource (light and N) availability, enabling species that depend on higher resource availability to co-occur; (2) spatial heterogeneity in resource availability is enhanced by bison, enabling coexistence of a greater number of plant species; (3) increased species turnover (i.e. increased species colonization and establishment) in grazed grassland is associated with enhanced plant species richness. We measured availability and spatial heterogeneity in light, water and N, and calculated species turnover from long-term data in grazed and ungrazed sites in a North American tallgrass prairie. Both regression and path analyses were performed to evaluate the potential of the three hypothesized mechanisms to explain observed patterns of plant species richness under field conditions. Experimental grazing by bison increased plant species richness by 25% over an 8-year period. Neither heterogeneity nor absolute levels of soil water or available N were related to patterns of species richness in grazed and ungrazed sites. However, high spatial heterogeneity in light and higher rates of species turnover were both strongly related to increases in plant species richness in grazed areas. This suggests that creation of a mosaic of patches with high and low biomass (the primary determinant of light availability in mesic grasslands) and promotion of a dynamic species pool are the most important mechanisms by which grazers affect species richness in high productivity grasslands.     

Vegetation and soil seed bank in a 23-year grazing exclusion chronosequence in a Mediterranean dry grassland

Long-term grazing shaped plant diversity in dry Mediterranean grasslands. Abandonment of grazing affects plant diversity especially in the northern Mediterranean. Considerable efforts are, therefore, under way for grassland conservation and restoration. Yet, we do not know at which temporal scales impacts of grazing abandonment appear and in particular how soil seed banks evolve after longer grazing abandonment. Here, we provide detailed data from one of the very few long-term experiments available. These experiments provide data for up to 23 years (1982–2005) of grazing exclusion built in 1982, 1989, 2000 and 2001. Grazing exclusion decreased species richness, modified vegetation structure and changed soil parameters. Decline in species richness appears in communities that experienced 16 and 23 years of grazing exclusion. Only four to nine plant species of this Mediterranean grassland built persistent soil seed banks appearing after grazing exclusion, compared to 40–50 species in the established vegetation of grazed plots. Hence, similarity between vegetation and soil seed bank decreased with time of grazing exclusion. Even 23 years after abandonment, no woody plants colonised the experiments. We conclude that vegetation will recover fast from grazing abandonment in the short-term. Nevertheless, longer abandonment will impact diversity due to reduced soil seed banks.     

Influence of experimental soil disturbances on the diversity of plants in agricultural grasslands

Aims and Methods Disturbance is supposed to play an important role for biodiversity and ecosystem stability as described by the intermediate disturbance hypothesis (IDH), which predicts highest species richness at intermediate levels of disturbances. In this study, we tested the effects of artificial soil disturbances on diversity of annual and perennial vascular plants and bryophytes in a field experiment in 86 agricultural grasslands differing in land use in two regions of Germany. On each grassland, we implemented four treatments: three treatments differing in application time of soil disturbances and one control. One year after experimental disturbance, we recorded vegetation and measured biomass productivity and bare ground. We analysed the disturbance response taking effects of region and land-use-accompanied disturbance regimes into account.Important findings Region and land-use type strongly determined plant species richness. Experimental disturbances had small positive effects on the species richness of annuals, but none on perennials or bryophytes. Bare ground was positively related to species richness of bryophytes. However, exceeding the creation of 12% bare ground further disturbance had a detrimental effect on bryophyte species richness, which corresponds to the IDH. As biomass productivity was unaffected by disturbance our results indicate that the disturbance effect on species richness of annuals was not due to decreased overall productivity, but rather due to short-term lowered inter- and intraspecific competition at the newly created microsites. Generally, our results highlight the importance of soil disturbances for species richness of annual plants and bryophytes in agricultural grasslands. However, most grasslands were disturbed naturally or by land-use practices and our additional experimental soil disturbances only had a small short-term effect. Overall, total plant diversity in grasslands seemed to be more limited by the availability of propagules rather than by suitable microsites for germination. Thus, nature conservation efforts to increase grassland diversity should focus on overcoming propagule limitation, for instance by additional sowing of seeds, while the creation of additional open patches by disturbance might only be appropriate where natural disturbances are scarce.     

Effects of grazing disturbance on plant diversity,community structure and direction of succession in an alpine meadow on Tibet Plateau,China

To elucidate the effects of grazing intensity and grazing time on plant diversity and community structure, as well as the successional differentiation in an alpine meadow, a controlled grazing trial, with six grazing intensities on an alpine meadow was conducted in the eastern Qilian Mountain region for four years. Using species accumulation curves, RDA ordination and variance decomposition, we analyzed the changes in proportion of dominant species, richness, abundance, as well as the life forms of plant communities under grazing disturbance. Both the grazing intensity and grazing time had a significant effect on these dominant species, richness, abundance, as well as the life forms in the plant community (P < 0.01). More detailed results showed that: (1) The richness and abundance of plant species were highest in the light grazing plot, and these increased as time passed. In the heavy grazing plot, the abundance of plant species decreased as time passed, but the richness of these species did not change significantly. (2) The abundance of Gramineae and Umbelliferae were negatively and significantly correlated with the duration of grazing treatments, whereas Plantaginaceae and Geraniaceae were positively and significantly correlated with the grazing intensity. Over time, the abundance of bunch-type plants decreased and other life forms of plants have increased. With the increase in grazing intensity, the plants' abundance with the rosette type did not change, but other life forms of plants decreased. The results of variance decomposition indicated that grazing disturbance has had greater effects on life forms and plant populations, followed by the changes in the dominant species and their abundance, with lesser effects on the richness of the species. Grazing intensity had a greater effect than the duration of the grazing treatment had. The results of PCA showed that the climax community in both the lowest and the highest grazing intensity plots had changed over time. In the sample plots with light grazing intensity, the plant community changed to an Elymus nutans + Poa crymophila community, but later changed to a Melilotoides ruthenicus + Kobresia humilis community under heavy grazing.     

Effects of grazing disturbance on plant diversity,community structure and direction of succession in an alpine meadow on Tibet Plateau,China

To elucidate the effects of grazing intensity and grazing time on plant diversity and community structure, as well as the successional differentiation in an alpine meadow, a controlled grazing trial, with six grazing intensities on an alpine meadow was conducted in the eastern Qilian Mountain region for four years. Using species accumulation curves, RDA ordination and variance decomposition, we analyzed the changes in proportion of dominant species, richness, abundance, as well as the life forms of plant communities under grazing disturbance. Both the grazing intensity and grazing time had a significant effect on these dominant species, richness, abundance, as well as the life forms in the plant community (P < 0.01). More detailed results showed that: (1) The richness and abundance of plant species were highest in the light grazing plot, and these increased as time passed. In the heavy grazing plot, the abundance of plant species decreased as time passed, but the richness of these species did not change significantly. (2) The abundance of Gramineae and Umbelliferae were negatively and significantly correlated with the duration of grazing treatments, whereas Plantaginaceae and Geraniaceae were positively and significantly correlated with the grazing intensity. Over time, the abundance of bunch-type plants decreased and other life forms of plants have increased. With the increase in grazing intensity, the plants' abundance with the rosette type did not change, but other life forms of plants decreased. The results of variance decomposition indicated that grazing disturbance has had greater effects on life forms and plant populations, followed by the changes in the dominant species and their abundance, with lesser effects on the richness of the species. Grazing intensity had a greater effect than the duration of the grazing treatment had. The results of PCA showed that the climax community in both the lowest and the highest grazing intensity plots had changed over time. In the sample plots with light grazing intensity, the plant community changed to an Elymus nutans + Poa crymophila community, but later changed to a Melilotoides ruthenicus + Kobresia humilis community under heavy grazing.     

Influence of grazing and fire frequency on small-scale plant community structure and resource variability in native tallgrass prairie

In grasslands worldwide, grazing by ungulates and periodic fires are important forces affecting resource availability and plant community structure. It is not clear, however, whether changes in community structure are the direct effects of the disturbance (i.e. fire and grazing) or are mediated indirectly through changes in resource abundance and availability. In North American tallgrass prairies, fire and grazing often have disparate effects on plant resources and plant diversity, yet, little is known about the individual and interactive effects of fire and grazing on resource variability and how that variability relates to heterogeneity in plant community structure, particularly at small scales. We conducted a field study to determine the interactive effects of different long-term fire regimes (annual vs four-year fire frequency) and grazing by native ungulates ( Bos bison ) on small-scale plant community structure and resource variability (N and light) in native tallgrass prairie. Grazing enhanced light and nitrogen availability, but did not affect small-scale resource variability. In addition, grazing reduced the dominance of C₄ grasses which enhanced species richness, diversity and community heterogeneity. In contrast, annual fire increased community dominance and reduced species richness and diversity, particularly in the absence of grazing, but had no effect on community heterogeneity, resource availability and resource variability. Variability in the abundance of resources showed no relationship with community heterogeneity at the scale measured in this study, however we found a relationship between community dominance and heterogeneity. Therefore, we conclude that grazing generated small-scale community heterogeneity in this mesic grassland by directly affecting plant community dominance, rather than indirectly through changes in resource variability.     

Perennial grassland dynamics on fertile plains: Is coexistence mediated by disturbance?

Abstract The response of grasslands to disturbance varies with the nature of the disturbance and the productivity of the landscape. In highly productive grasslands, competitive exclusion often results in decreased species richness and grazing may allow more species to coexist. Once widespread, grasslands dominated by Dichanthium sericeum (Queensland bluegrass) and Astrebla spp. (Mitchell grass) occur on fertile plains but have been reduced in extent by cultivation. We tested the effects of exclusion of livestock grazing on these grasslands by comparing the floristic composition of sites in a nature reserve with an adjacent stock reserve. In addition, sites that had been cultivated within the nature reserve were compared with those where grazing but no cultivation had occurred. To partition the effects of temporal variation from spatial variation we sampled sites in three different years (1998, 2002 and 2004). Some 194 taxa were recorded at the nature reserve and surrounding stock routes. Sampling time, the occurrence of past cultivation and livestock grazing all influenced species composition. Species richness varied greatly between sampling periods relating to highly variable rainfall and water availability on heavy clay soils. Native species richness was significantly lower at previously cultivated sites (13–22 years after cultivation), but was not significantly influenced by grazing exclusion. After 8 years it appears that reintroducing disturbance in the form of livestock grazing is not necessary to maintain plant species richness in the reserve. The highly variable climate (e.g. droughts) probably plays an important role in the coexistence of species by negating competitive exclusion and allowing interstitial species to persist.     

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.     

Butterfly and grasshopper diversity patterns in humid Mediterranean grasslands: the roles of disturbance and environmental factors

The present paper studies butterfly, grasshopper and vascular plant communities in ten seasonally flooded grasslands with different anthropogenic disturbance regimes (NW Greece). Disturbance intensity was assessed on the basis of disturbance frequency and type (grazing, mowing, trampling, constructions). The distribution patterns of butterflies are regulated by humidity and elevation (Redundancy Analysis). Elevation, flower-heads abundance, low disturbance intensity and plant species richness predict grasshopper species richness well, while the latter together with humidity predict plant species richness (Generalized Linear Models). Chorthippus lacustris, a critically endangered endemic grasshopper species, is positively associated with humid microhabitats with high flower-heads abundance. An indicator value procedure reveals four butterfly species as being typical species for habitats with a pronounced character of hedgerows and tree lines. Conservation management of grassland butterflies should focus on the maintenance of the humid character of the humid grasslands as well as on the maintenance of hedgerows and tree lines. The reduction of human-induced disturbance towards occasional grazing and mowing seems to benefit both butterfly and grasshopper communities. Finally, we suggest the use of grasshoppers as surrogates for vascular plants and vice versa, given their congruent species richness patterns.     

From plant neighbourhood to landscape scales: how grazing modifies native and exotic plant species richness in grassland

The interactive effect of grazing and soil resources on plant species richness and coexistence has been predicted to vary across spatial scales. When resources are not limiting, grazing should reduce competitive effects and increase colonisation and richness at fine scales. However, at broad scales richness is predicted to decline due to loss of grazing intolerant species. We examined these hypotheses in grasslands of southern Australia that varied in resources and ungulate grazing intensity since farming commenced 170 years ago. Fine-scale species richness was slightly greater in more intensively grazed upper slope sites with high nutrients but low water supply compared to those that were moderately grazed, largely due to a greater abundance of exotic species. At broader scales, exotic species richness declined with increasing grazing intensity whether nutrients or water supply were low or high. Native species richness declined at all scales in response to increasing grazing intensity and greater resource supply. Grazing also reduced fine-scale heterogeneity in native species richness and although exotics were also characterised by greater heterogeneity at fine scales, grazing effects varied across scales. In these grasslands patterns of plant species richness did not match predictions at all scales and this is likely to be due to differing responses of native and exotic species and their relative abundance in the regional species pool. Over the past 170 years intolerant native species have been eliminated from areas that are continually and heavily grazed, whereas transient, light grazing increases richness of both exotics and natives. The results support the observation that the processes and scales at which they operate differ between coevolved ungulate—grassland systems and those in transition due to recent invasion of herbivores and associated plant species.     

草地植物生产力主要影响因素研究综述

草地是全球分布面积最大的陆地生态系统,植物初级生产力是反映草地功能的重要指标。从植物种多样性、资源有效性、放牧、退化草地恢复和气候变化等方面较系统综述了影响草地植物生产力的关键要素和驱动力。大量研究表明,植物多样性与生产力的关系尚未有一致的结论,依据试验地点、起始状态甚至度量指标不同而不同;特别是资源有效性调节着生产力水平并对植物多样性和生产力关系产生显著影响;放牧改变了植物群落特征和养分有效性进而影响生产力的形成过程,也改变了资源有效性-植物多样性-生产力之间的关系;对于退化生态系统,在退化草地恢复过程中植物与土壤资源有效性的互作效应对植物生产力的变化起着关键作用;而在未来气候变化特别是增温对植物生产力的影响因地点和生态系统的不同而异,但多数研究结果显示增温提高了草地植物生产力。与国外其它草地分布区相比,国内的相关研究不仅在数量上明显不足,更重要是欠缺机理上的深入研究。在放牧和未来气候变化背景下如何维持和提高草地生产力,如何加速退化草地生态系统的恢复,进而实现生态安全建设和经济社会协调发展,是我国当前急需解决的理论和实践问题。     

Intraspecific variation overrides origin effects in impacts of litter-derived secondary compounds on larval amphibians

Vertebrate herbivores can be key determinants of grassland plant species richness, although the magnitude of their effects can largely depend on ecosystem and herbivore characteristics. It has been demonstrated that the combined effect of primary productivity and body size is critical when assessing the impact of herbivores on plant richness of perennial-dominated grasslands; however, the interaction of site productivity and herbivore size as determinants of plant richness in annual-dominated pastures remains unknown. We experimentally partitioned primary productivity and herbivore body size (sheep and wild rabbits) to study the effect of herbivores on the plant species richness of a Mediterranean semiarid annual plant community in central Spain over six years. We also analyzed the effect of grazing and productivity on the evenness and species composition of the plant community, and green cover, litter, and plant height. We found that plant richness was higher where the large herbivore was present at high-productivity sites but barely changed at low productivity. The small herbivore did not affect species richness at either productivity site despite its large effects on species composition. We propose that adaptations to resource scarcity and herbivory prevented plant richness changes at low-productivity sites, whereas litter accumulation in the absence of herbivores decreased plant richness at high productivity. Our results are consistent with predictions arising from a long history of grazing and highlight the importance of both large and small herbivores to the maintenance of plant diversity of Mediterranean annual-dominated pastures.     

Grazing, Environmental Heterogeneity, and Alien Plant Invasions in Temperate Pampa Grasslands

Temperate humid grasslands are known to be particularly vulnerable to invasion by alien plant species when grazed by domestic livestock. The Flooding Pampa grasslands in eastern Argentina represent a well-documented case of a regional flora that has been extensively modified by anthropogenic disturbances and massive invasions over recent centuries. Here, we synthesise evidence from region-wide vegetation surveys and long-term exclosure experiments in the Flooding Pampa to examine the response of exotic and native plant richness to environmental heterogeneity, and to evaluate grazing effects on species composition and diversity at landscape and local community scales. Total plant richness showed a unimodal distribution along a composite stress/fertility gradient ranging several plant community types. On average, more exotic species occurred in intermediate fertility habitats that also contained the highest richness of resident native plants. Exotic plant richness was thus positively correlated with native species richness across a broad range of flood-prone grasslands. The notion that native plant diversity decreases invasibility was supported only for a limited range of species-rich communities in habitats where soil salinity stress and flooding were unimportant. We found that grazing promoted exotic plant invasions and generally enhanced community richness, whereas it reduced the compositional and functional heterogeneity of vegetation at the landscape scale. Hence, grazing effects on plant heterogeneity were scale-dependent. In addition, our results show that environmental fluctuations and physical disturbances such as large floods in the pampas may constrain, rather than encourage, exotic species in grazed grasslands.     

Effects of grazing exclusion on plant species richness and phytomass accumulation vary across a regional productivity gradient

Question: Does long‐term grazing exclusion affect plant species diversity? And does this effect vary with long‐term phytomass accumulation across a regional productivity gradient? Location: Lowland grassy ecosystems across the state of Victoria, southeast Australia. Methods: Floristic surveys and phytomass sampling were conducted across a broad‐scale productivity gradient in grazing exclusion plots and adjacent grazed areas. Differences in species richness, evenness and life‐form evenness between grazed and ungrazed areas were analysed. The environmental drivers of long‐term phytomass accumulation were assessed using multiple linear regression analysis. Results: Species richness declined in the absence of grazing only at the high productivity sites (i.e. when phytomass accumulation was >500 g m^?2). Species evenness and life‐form evenness also showed a negative relationship with increasing phytomass accumulation. Phytomass accumulation was positively associated with both soil nitrogen and rainfall, and negatively associated with tree cover. Conclusions: Competitive dominance is a key factor regulating plant diversity in productive grassy ecosystems, but canopy disturbance is not likely to be necessary to maintain diversity in less productive systems. The results support the predictions of models of the effects of grazing on plant diversity, such as the dynamic equilibrium model, whereby the effects of herbivory are context‐dependent and vary according to gradients of rainfall, soil fertility and tree cover.     

Functional dependence underlies a positive plant-grasshopper richness relationship

A central focus of ecology is identifying the factors that shape spatial patterns of species diversity and this is particularly relevant in an era of global change. Positive relationships between plant and consumer diversity are common, but could be driven by direct responses of each trophic level to underlying environmental gradients, or indirectly where changes in environmental conditions propagate through food webs. Here we use structural equation modeling to examine the relative importance of soil resource availability and disturbance (fire) in mediating relationships between plant and grasshopper richness in insular grasslands. We found a positive relationship between plant and consumer richness that became stronger after accounting for disturbance, despite unique responses of plants and consumers to the two environmental gradients. Plant richness responded to an underlying gradient in soil resource availability. Time since the last fire had a direct positive effect on grasshopper richness but had no effect on plant richness. This work supports that plant and consumer richness are functionally linked, rather than having similar responses to environmental gradients. By disentangling the direct and indirect processes underlying a positive relationship between plant and consumer diversity in a natural system that spans multiple environmental gradients, we demonstrate the importance of investigating biodiversity through explicit multivariate models.     

Determinants of bryophyte species composition and diversity on the Great Alvar of Öland,Sweden

Factors driving the species richness and distribution of bryophytes are poorly studied and not well understood, particularly in grasslands. We analysed the occurrence of bryophyte species and variation in species richness across 674 plots (0.5?m?×?0.5?m) in alvar vegetation (grassland on limestone pavement with thin or no soil) on Öland (Sweden) in relation to substrate characteristics and chemistry, inundation frequency, grazing pressure and geographical variables. We found 148 taxa, including 11 nationally red-listed ones. Species richness per plot was significantly associated with substrate type, positively associated with pH and grazing intensity, but negatively associated with soil depth. However, richness of species typical of, or restricted to, alvar habitats responded differently to richness of species more common in other habitats. Typical alvar species were favoured by high pH, shallow soil and low phosphate availability, while generalists preferred relatively low pH, higher phosphate availability and organic or mull soil types. Distance from the alvar margin had only weak effects. Concerning the effects on individual species and community composition, inundation frequency and pH were found to have the largest effects, although other factors (substrate type, soil depth, bare soil, bare stone, phosphate availability and grazing pressure) were more important for some individual species, stressing the importance of microsite variability and variability in management for regional species richness. From a conservation perspective, it is concluded that grazing is generally positive whilst factors increasing phosphate availability may disadvantage the typical alvar species, and proximity to the alvar margin is not a major problem.     

Temporary grazing exclusion promotes rapid recovery of species richness and productivity in a long‐term overgrazed Campos grassland

Moderate grazing intensity is considered the basic requirement to enhance ecosystem function in grasslands. Yet, deterioration by overgrazing is common in many biomes, including Campos grasslands in South America. Understanding how grazing management can lead to recovery of ecosystem function is essential to design and implement effective strategies for sustainable use of this resource. In a long‐term field experiment carried out in Southern Brazil, we studied the effects of temporal grazing exclusions (spring or fall) at moderate and severe livestock grazing intensities (maintained by adjusting contrasting forage allowances) on the species richness, botanical composition, forage mass, sward height, and photosynthetic active radiation intercepted. The experiment was arranged in a completely randomized design with three replications of grazing exclusions, applied simultaneously at moderate and severe grazing intensities. Moderate grazing intensity showed a bimodal structure of shorter and taller canopies, and high species richness. Severe grazing created a shorter and homogeneous sward structure characterized by less standing biomass and species loss. In response to grazing exclusions, sward height, standing biomass, and light interception recovered almost to the levels of moderate grazing. Further, within 2 years grass species richness increased and botanical composition changed toward grasses with erect habit prevailing in moderate grazing intensity. Our study confirms that (1) moderate grazing intensities allow the coexistence of high number of species and (2) spring grazing exclusions of long‐term overgrazed grasslands can lead to a quick start to recover the grass species richness, primary productivity, and species composition like that prevailing in well‐managed grasslands.     

Moderate grazing promotes the root biomass in Kobresia meadow on the northern Qinghai–Tibet Plateau

Grazing is an important modulator of both plant productivity and biodiversity in grassland community, yet how to determine a suitable grazing intensity in alpine grassland is still controversy. Here, we explore the effects of different grazing intensities on plant biomass and species composition, both at community level and functional group level, and examines the productivity–species richness relationship under four grazing patterns: no grazing (CK), light grazing (LG), moderate grazing, (MG) and heavy grazing (HG), attempt to determine a suitable grazing intensity in alpine grassland. The results were as follows. The total aboveground biomass (AGB) reduced with increasing grazing intensity, and the response of plant functional groups was different. AGB of both sedges and legumes increased from MG to HG, while the AGB of forbs reduced sharply and the grass AGB remained steady. There was a significant positive relationship between productivity and species richness both at community level and functional group level. In contrast, the belowground biomass (BGB) showed a unimodal relationship from CK to HG, peaking in MG (8,297.72 ± 621.29 g/m²). Interestingly, the grassland community tends to allocate more root biomass to the upper soil layer under increasing grazing intensities. Our results suggesting that moderate levels of disturbance may be the optimal grassland management strategy for alpine meadow in terms of root production.     

Co-limitation of plant primary productivity by nitrogen and phosphorus in a species-rich wooded meadow on calcareous soils

High small-scale species richness of calcareous grasslands is generally thought to result from evening of species competitive potentials by limited N availability, because of relatively low herb N/P ratios in these communities. However, P mobility is low in alkaline soils as well. We studied soil chemistry and productivity of herb and moss layers in a very diverse calcareous meadow (up to 76 vascular plant species per m²) to test the hypotheses of a co-limitation of herb productivity by both soil N and P availabilities and moss productivity primarily by P availability. The effect of nutrient supply on productivity was investigated using both a natural productivity gradient as well as fertilization experiments. We observed strong positive correlations of soil P availability and total soil N with the above-ground productivity of herb layer. A long-term fertilization experiment demonstrated that P alone and N and P together increased productivity of vascular species, and that the productivity continuously declined after cessation of fertilization with the effect of previous fertilization occasionally visible even 14 years after treatment termination. A short-term fertilization experiment further demonstrated that N and P when supplied alone increase productivity of vascular plants, suggesting that both elements were limiting. Furthermore, there was a significant interaction between N and P on productivity, indicating that simultaneous N and P supply increased productivity more than separate nutrient additions. Moss productivity was negatively associated with vascular plant productivity. In particular, N addition decreased moss productivity, but moss productivity did not decline in P addition treatments. P requirements of mosses were larger than those of vascular plants. Our data indicate co-limitation of herb productivity by both soil N and P in this highly diverse grassland, while limitation of moss productivity mainly by P. We suggest that N and P co-limitations are common in calcareous diverse grasslands, and may partly explain the extreme small-scale species diversity in these communities.