Direct and indirect effects of land‐use intensification on ant communities in temperate grasslands

Land‐use intensification is a major driver of local species extinction and homogenization. Temperate grasslands, managed at low intensities over centuries harbored a high species diversity, which is increasingly threatened by the management intensification over the last decades. This includes key taxa like ants. However, the underlying mechanisms leading to a decrease in ant abundance and species richness as well as changes in functional community composition are not well understood. We sampled ants on 110 grassland plots in three regions in Germany. The sampled grasslands are used as meadows or pastures, being mown, grazed or fertilized at different intensities. We analyzed the effect of the different aspects of land use on ant species richness, functional trait spaces, and community composition by using a multimodel inference approach and structural equation models. Overall, we found 31 ant species belonging to 8 genera, mostly open habitat specialists. Ant species richness, functional trait space of communities, and abundance of nests decreased with increasing land‐use intensity. The land‐use practice most harmful to ants was mowing, followed by heavy grazing by cattle. Fertilization did not strongly affect ant species richness. Grazing by sheep increased the ant species richness. The effect of mowing differed between species and was strongly negative for Formica species while Myrmica and common Lasius species were less affected. Rare species occurred mainly in plots managed at low intensity. Our results show that mowing less often or later in the season would retain a higher ant species richness—similarly to most other grassland taxa. The transformation from (sheep) pastures to intensively managed meadows and especially mowing directly affects ants via the destruction of nests and indirectly via loss of grassland heterogeneity (reduced plant species richness) and increased soil moisture by shading of fast‐growing plant species.     

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.     

An attempt to restore a central European species‐rich mountain grassland through grazing

Abstract. This paper describes the effects of re‐establishing seasonal cattle grazing by 0.7 animal.ha^‐1 on vegetation in a long‐term abandoned, and partly degraded, semi‐natural mountain pasture in the ?umava National Park, Czech Republic. There was very uneven grazing intensity inside the locality, and grazing preference changed during the season: cattle grazed most of the time in productive but species‐poor Deschampsia cespitosa swards, but changed to a species‐rich Violion caninae stand in the middle of the summer. A species‐rich Carex rostrata community was only grazed at the end of the season. Species‐poor swards dominated by Nardus stricta and Carex brizoides were mainly used as resting areas. Both grazing and excluding from grazing had a negative effect on species diversity of the Deschampsia cespitosa swards. The soil seed bank contained only few species that are characteristic of mountain grassland communities, and seed dispersal of the target species by cattle dung was also found to be very limited. Thus both grazing and exclusion from grazing are probably of limited value for the restoration of species‐rich grasslands from species‐poor Deschampsia cespitosa swards in this case.     

After the hotspots are gone: Land use history and grassland plant species diversity in a strongly transformed agricultural landscape

Question: We asked how landscape configuration and present management influence plant species richness and abundance of habitat specialists in grasslands in a ‘modern’(much exploited and transformed) agricultural Swedish landscape. Location: Selaön, south‐eastern Sweden (59°24’ N, 17°10’ E). Methods: Present and past (150 and 50 years ago) landscape pattern was analysed in a 25 km² area. Species richness was investigated in 63 different grassland patches; grazed and abandoned semi‐natural grasslands, and grazed ex‐arable fields. Influence of landscape variables; area, past and present grassland connectivity, present management on total species richness, density and abundance of 25 grassland specialists was analysed. Results: Semi‐natural grasslands (permanent unfertilised pastures or meadows formed by traditional agricultural methods) had declined from 60% 150 years ago to 5% today. There was a significant decline in species richness and density in abandoned semi‐natural grasslands. Total species richness was influenced by present management, size and connectivity to present and past grassland pattern. Landscape variables did not influence species density in grazed semi‐natural grassland suggesting that maintained grazing management makes grassland patches independent of landscape context. The abundance of 16 grassland specialists was mainly influenced by management and to some extent also by landscape variables. Conclusion: Although species richness pattern reflect management and to some extent landscape variables, the response of individual species may be idiosyncratic. The historical signal from past landscapes is weak on present‐day species richness in highly transformed, agricultural landscapes. Generalizations of historical legacies on species diversity in grasslands should consider also highly transformed landscapes and not only landscapes with a high amount of diversity hotspots left.     

Patch dynamics in grazed subtropical native pastures in south‐east Queensland

Abstract Patch formation is common in grazed grasslands but the mechanisms involved in the formation and maintenance of patches are not clear. To increase our knowledge on this subject we examined possible reasons for patch formation and the influence of management on changes between patch states in three experiments in native pasture communities in the Crows Nest district, south‐east Queensland. In these communities, small‐scale patches (tall grassland (dominated by large and medium tussock grasses), short swards (dominated by short tussock grasses and sedges), and lawns (dominated by stoloniferous and/or rhizomatous grasses)) are readily apparent. We hypothesized that the formation of short sward and lawn patches in areas of tall grassland was due to combinations of grazing and soil fertility effects. This was tested in Experiment 1 by applying a factorial combination of defoliation, nutrient application and transplants of short tussock and stoloniferous species to a uniform area of tall grassland. Total species density declined during the experiment, was lower with high nutrient applications, but was not affected by defoliation. There were significant changes in abundance of species that provided support for our hypotheses. With light defoliation and low nutrients, the tall grassland remained dominated by large tussock grasses and contained considerable amounts of forbs. With heavy defoliation, the pastures were dominated by medium tussock grasses and there were significant decreases in forbs and increases in sedges (mainly with low nutrients) and stoloniferous grasses (mainly with high nutrients). Total germinable seed densities and those of most species groups were significantly lower in the heavy defoliation than the light defoliation plots. Total soil seed numbers were not affected by nutrient application but there were fewer seeds of the erect forbs and more sedge seeds in plots with high nutrients. The use of resting from grazing and fire to manage transitions between patches was tested. In Experiment 2 , changes in species density and abundance were measured for 5 years in the three patch types with and without grazing. Experiment 3 examined the effects of fire, grazing and resting on short sward patches over 4 years. In Experiment 2 , total species density was lower in lawn than short sward or tall grassland patches, and there were more species of erect forbs than other plant groups in all patch types. The lawn patches were originally dominated by Cynodon spp. This dominance continued with grazing but in ungrazed patches the abundance of Cynodon spp. declined and that of forbs increased. In the short sward patches, dominance of short tussock grasses continued with grazing but in ungrazed plots their abundance declined while that of large tussock grasses increased. The tall grassland patches remained dominated by large and medium tussock species. In Experiment 3 , fire had no effect on species abundance. On the grazed plots the short tussock grasses remained dominant but where the plots were rested from grazing the small tussock grasses declined and the large tussock grasses increased in abundance. The slow and relatively small changes in these experiments over 4 or 5 years showed how stable the composition of these pastures is, and that rapid changes between patch types are unlikely.     

Impact of grazing on germination trait selection in an alpine grassland on the Tibet Plateau

放牧对青藏高原高寒草地种子萌发性状选择的影响 以前的研究表明放牧能够引起草地生物与非生物环境的显著变化,但这种变化影响草地群落对种子萌发特征的选择机制尚不清楚。因此,我们旨在回答：放牧是否对草地群落中萌发特征的组成和多样性产生显著影响。我们在实验室检测了研究草地群落内主要物种种子的萌发特性,并比较了这些植物在放牧和非放牧草地上的表现。在此基础上,比较了放牧草地和非放牧草地的各萌发性状的群落加权平均值和萌发性状多样性,从而了解放牧草地和非放牧草地是否存在不同的萌发性状结构。研究结果表明,在物种水平上,放牧和非放牧草地各物种的多度变化与物种的萌发性状无显著关系。但在群落水平上,与非放牧草地相比,放牧草地的物种普遍具有较高的种子萌发率;放牧草地种子萌发对变温的正响应显著大于非放牧草地,而且放牧草地种子萌发温度生态位宽度小于非放牧草地。与非放牧草地相比,放牧草地种子萌发性状多样性增加,萌发性状均匀度降低。放牧可以改变微生境,从而通过环境过滤改变草地群落对萌发性状的选择,导致草地群落的萌发性状于放牧前后在群落水平上存 在显著差异。     

内蒙古锡林郭勒草地的草种组分与放牧压的关系研究

通过了采样调查,对内蒙古锡林河流域的放牧草地群落的组成成分,现存生物量,物种多样性等与放牧强度的关系进行了研究。结果表明,距离定居点越远放牧强度越弱,出现了以羊草(Leymus chinensis)和大针茅(Stipa grandis)为优势种的群落。距离定居点越近放牧强度越强,出现了以糙隐子草(Cleistogenes squarrosa)和寸苔草(Carex korshinskyi)为优势种的群溶。单位样方内(1m^2)地上部的现存生物量在强放牧压下的127．3g和较弱放牧压下的303g之间变化。物种多样性的变化则独立于放牧强度差异,在3．13至3．64之间波动。为了判断放牧草地的牧草的生长状态,采用了生长状态质量指数(Stand quality index：SQI)来评价了实验样地群落的生长状态。在定居点的内部,SQI为278,而在离定居点4公里远的样地,指数则变为667。禁牧13年的原退化样地的SQI值从420上升至624。     

Fuzzy set ordination and classification of Serengeti short grasslands,Tanzania

Abstract. Species abundance data from 32 representative stands are used to evaluate the vegetation in the heavily grazed Serengeti short grasslands. The abundant species are: Kyllinga nervosa, Sporobolus ioclados, S. kentro-phyllus and Eustachys paspaloides, all occurring in elevated areas, and Hypoestes forskalei, which is found in the drainage areas. Numerical classification and fuzzy set ordination revealed four main grassland communities associated with a topographic gradient. The communities are considered distinct since their niches, as computed in the community and environmental fuzzy system space are not overlapping. Leaching, erosion and animal disturbance effects correlated with the topographic gradient are found to determine species composition and overall community structure in the grasslands.     

Resilience of a high‐conservation‐value,semi‐arid grassland on fertile clay soils to burning,mowing and ploughing

In grassland reserves, managed disturbance is often necessary to maintain plant species diversity. We carried out experiments to determine the impact of fire, kangaroo grazing, mowing and disc ploughing on grassland species richness and composition in a nature reserve in semi‐arid eastern Australia. Vegetation response was influenced by winter–spring drought after establishment of the experiments, but moderate rainfall followed in late summer–autumn. Species composition varied greatly between sampling times, and the variability due to rainfall differences between seasons and years was greater than the effects of fire, kangaroo grazing, mowing or disc ploughing. In the fire experiment, species richness and composition recovered more rapidly after spring than autumn burning. Species richness and composition were similar to control sites within 12 months of burning and mowing, suggesting that removal of the dominant grass canopy is unnecessary to enhance plant diversity. Two fires (separated by 3 years) and post‐fire kangaroo grazing had only minor influence on species richness and composition. Even disc ploughing caused only a small reduction in native richness. The minor impact of ploughing was explained by the small areas that were ploughed, the once‐off nature of the treatment, and the high degree of natural movement and cracking in these shrink‐swell soils. Recovery of the composition and richness of these grasslands was rapid because of the high proportion of perennial species that resprout vegetatively after fire and mowing. There appears to be little conservation benefit from fire, mowing or ploughing ungrazed areas, as we could identify no native plant species dependent on frequent disturbance for persistence in this grassland community. However, the ability of the Astrebla‐ and Dichanthium‐dominated grasslands to recover quickly after disturbance, given favourable seasonal conditions, suggests that they are well adapted to natural disturbances (e.g. droughts, fire, flooding and native grazing).     

Use of vegetation classification and plant indicators to assess grazing abandonment in Estonian coastal wetlands

Question: What are the effects of grazing abandonment on the vegetation composition of Estonian coastal wetlands? Location: Vormsi Island and Silma Nature Reserve in western Estonia, Europe. Methods: Local knowledge and field reconnaissance were used to identify current and historical management levels of wetland sites within the west Estonian study area. Nine study sites, with varying management histories, were selected comprising an area of 287 ha. A total of 198 quadrats were taken from 43 distinct vegetation patches in five of the sites. TWINSPAN analysis was used to identify community type, and a phytosociological key was constructed for character taxa. This vegetation classification was then applied within a GIS‐based context to classify all the study sites, using a ground survey technique and 1:2000 scale air photos. Results: We identified 11 different brackish coastal wetland community types. Indicator species were defined with community characteristics for the seven main vegetation types readily recognisable in the field. Coastal wet grasslands were most extensive in grazed sites, or sites that had been more intensively grazed, while abandoned sites were largely composed of Phragmites australis stands, tall grassland, and scrub. Site variations based on vegetation composition were significantly correlated with past grazing intensity. Plant community types showed significant edaphic differences, with particularly low soil moisture and high conductivity and pH for open pioneer patches compared to other vegetation types. Conclusion: Abandonment of traditionally grazed coastal grasslands threatens their characteristic biodiversity. This study found that grazing abandonment reduced the extent of coastal wetland grasslands of particular conservation value. Nevertheless, plant species of conservation interest were found across the sequence of community types described. The study shows that grazing is an important factor influencing coastal wetland plant communities but suggests that vegetation distribution is affected by environmental variables, such as topography.     

Restoration of semi-natural grasslands,a success for phytophagous beetles (Curculionidae)

Semi-natural pastures have rich plant and animal communities of high conservation value which depend on extensive management. As the area of such land decreases, abandoned semi-natural grasslands have been restored to re-establish biodiversity. Restoration schemes, which include thinning of woody plants and reintroduction of grazing, are mainly designed according to the responses of well-studied groups (such as vascular plants and birds). Weevils (Curculionidae) are a very diverse phytophagous beetle family. Here, we evaluated the restoration success of pastures for weevils (Curculionidae), by comparing their species diversity in abandoned, restored, and continuously grazed semi-natural pastures on 24 sites in central Sweden. Weevils were sampled by sweep-netting. We recorded 3019 weevil individuals belonging to 104 species. There was no statistically significant difference in species numbers between the pasture management treatments. However, weevil species composition of abandoned pastures differed from those in restored and continuously managed pastures, but there was no significant difference in community composition between restored and continuously grazed pastures. The abandoned sites tended to be dominated by polyphagous species, whereas the grazed sites contained more monophagous and oligophagous species. The number of weevil species was positively related to understory vegetation height and connectivity to other semi-natural grasslands and negatively related to the cover of trees and shrubs in the pastures. We conclude that restoration of abandoned semi-natural pastures is a good approach to restore weevil communities. To maintain a species rich weevil community, pastures should be managed to be relatively open, but still have patches of tall field-layer vegetation. Restoration and conservation measures should primarily be targeted on regions and landscapes where a high proportion of semi-natural grassland still remains.     

Recreating Grasslands in Swedish Rural Landscapes – Effects of Seed Sowing and Management History

Recent loss of plant species richness in Swedish semi-natural grasslands has led to an increase in grassland recreation and restoration. To increase the establishment of declining species favoured by grazing and to re-establish original species richness, seed sowing has been discussed as a conservation tool. In this study, I examined to what extent seed sowing in former arable fields increases species richness and generates a species composition typical of semi-natural grasslands. Six grassland species favoured by grazing (target species) and six generalist species favoured by ceased grazing, were studied in a seed-addition experiment. Four different seed densities were used on four different grassland categories, two grazed former arable fields, one continuously grazed grassland and one abandoned grassland. Target and generalist species emerged in all grassland categories, but seedling emergence was higher in the grazed than in the abandoned grassland. Target species had higher emergence in the two grasslands with the longest grazing continuity. Seedling emergence and frequency of established plants of each target species were positively associated. The largest fraction of seeds germinated at an intermediate sowing density, 20–50 seeds/dm², suggesting that aggregation of seeds positively affects emergence up to a certain threshold. In conclusion, artificial seed sowing may induce the recreation of typical grassland communities on former arable fields, which may be an important contribution to increase the total grassland area and species richness in the landscape.     

Cattle grazing in CRP grasslands during the nesting season: effects on avian abundance and diversity

The Conservation Reserve Program (CRP) is a primary tool for restoring grassland in the United States, in part as wildlife habitat, which has benefited declining grassland bird populations. Among potential mid-contract management practices used to maintain early-successional CRP grasslands, cattle grazing had been prohibited and is currently disincentivized during the primary nesting season for birds (much of the growing season), despite the important role that large herbivores historically played in structuring grassland ecosystems. Conservative grazing of CRP grasslands could increase spatial heterogeneity in vegetation structure and plant diversity, potentially supporting higher densities of some grassland bird species and higher bird diversity. Our objective was to determine the effect of experimental cattle grazing on species-specific relative abundance and occupancy, species diversity, and community dissimilarity of grassland birds on CRP grasslands across the longitudinal extent of Kansas, USA (a 63.5-cm precipitation gradient) during the 2017–2019 avian breeding seasons. Fifty-three of 108 fields were grazed by cattle during the growing seasons of 2017 and 2018 and all fields were rested from grazing in 2019. For all analyses, we examined separate model sets for semiarid western versus more mesic eastern Kansas. Using data from line transect surveys, we modeled relative abundances of 5 songbird species: grasshopper sparrow (Ammodramus savannarum), dickcissel (Spiza americana), eastern meadowlark (Sturnella magna), western meadowlark (Sturnella neglecta), and brown-headed cowbird (Molothrus ater). Grazing had delayed yet positive effects on abundances of grasshopper sparrow in western Kansas, and eastern meadowlark in eastern Kansas, but negative effects on dickcissel abundance in western Kansas and especially on burned fields in eastern Kansas. Somewhat counterintuitively, brown-headed cowbirds in western Kansas were more abundant on ungrazed versus grazed fields in the years after grazing began. In addition, we modeled multi-season occupancy of 3 gamebird species (ring-necked pheasant [Phasianus colcicus], northern bobwhite [Colinus virginianus], mourning dove [Zenaida macroura]) and Henslow's sparrow (Centronyx henslowii); grazing did not affect occupancy of these species. In eastern Kansas, species diversity was highest in grazed, unburned fields. In western Kansas, bird communities in grazed and ungrazed fields were dissimilar, as determined from multivariate analysis. Though regionally variable, conservative stocking of cattle on CRP grasslands during the nesting season as a mid-contract management tool might increase bird species diversity by restructuring habitat that accommodates a greater variety of species and decreasing abundances of species associated with taller, denser stands of vegetation.     

Spontaneous steppe restoration on abandoned cropland in Kazakhstan: grazing affects successional pathways

After the collapse of the Soviet Union in 1991, 12 million ha of cropland were abandoned in the steppe zone of Kazakhstan. At the same time livestock numbers crashed, leaving large areas of steppe without any grazing. We aimed to investigate, to which degree spontaneous succession on former cropland leads to the restoration of native steppe vegetation, and how this process is influenced by changing grazing patterns. We recorded biomass and vegetation characteristics as well as local soil and landscape variables in 151 quadrats of 100 m², covering 89 plots on former cropland abandoned in the early 1990s and 62 reference plots of near-natural steppe grassland that was never ploughed. About half of the plots of each category were located in a remote region where grazing has been absent for ca. 20 years, whereas the other half was located in a region with moderate livestock grazing. While there were no differences in the diversity, structure and plant life form composition of currently grazed and un-grazed near-natural steppe grasslands, corresponding successional plots on abandoned arable land exhibited significant differences. Grazed plots on former fields showed higher species richness and a higher cover of dwarf shrubs (mostly Artemisia spec.), ruderals and perennial herbs. At the same time, immigration of typical steppe species was much more successful. Contrary, in the absence of any grazing we found species-poor swards dominated by Stipa lessingiana and Leymus ramosus exhibiting an increasing frequency of wildfires due to litter accumulation. After 15–20 years, secondary steppe grasslands still differed substantially from their near-natural references. Our results suggest that grazing is mandatory to fully restore the original near-natural steppe vegetation and the underlying processes of pyric herbivory.     

Biodiversity of semi‐arid Mediterranean grasslands: Impact of grazing and afforestation

Question: What is the impact of grazing and/or afforestation on grassland diversity, species composition and cover parameters? Location: Semi‐arid Mediterranean grasslands of Jordan. Methods: Vegetation, litter, bare soil and rock cover were compared among four management types – free grazing and protected from grazing with three levels of tree cover. Species composition, plant cover, species richness and evenness were used to evaluate differences in vegetation among management types. Species composition differences among management types were also investigated. Results: Semi‐arid Mediterranean grasslands harbour appreciable levels of plant biodiversity. Grazing did not affect plant diversity, indicating the high resilience against and adaptation to grazing; however,grazing affected species composition and cover parameters. Afforestation seems to protect soil through higher litter cover but its impact on plant biodiversity was negative and markedly affected species composition. Conclusions: Neither protection from grazing or massive afforestation alone are sufficient for conserving biodiversity in this system. A management model is suggested where the landscape should be maintained as a mosaic of four management types: complete protection from grazing, grazing rotation, planting sparse trees in eroded areas and revegetating degraded areas using native, herbaceous and grazing tolerant species.     

The conservation benefit of mowing vs grazing for management of species‐rich grasslands: a multi‐site,multi‐year field experiment

Species‐rich semi‐natural grasslands in Europe are becoming more fragmented and many species that depend on this habitat type are rare and threatened today. Management methods like mowing and grazing are needed to preserve remaining grasslands. Because management is costly it is important to use the most cost‐effective as well as the most beneficial management method, but few studies have compared mowing and grazing. We investigated the effect of mowing and grazing on grassland vegetation using data from 11 long‐term field trials situated in southern Sweden. We calculated the change in the odds of finding species belonging to three different groups of indicators at the start of the treatment and after 8 and 14 years. The used indicator groups were indicators of good management, excess nitrogen and poor management. The results revealed an increase in the odds of finding indicators of good management in mowed plots and an increase in finding indicators of excess nitrogen in grazed plots. The odds of finding indicators of poor management remained unchanged. Results from sub‐analysis of the grazing intensity showed a more negative effect from grazing with low grazing intensity than normal/high grazing intensity. Therefore, mowing is the best long‐term management method for semi‐natural grasslands in Sweden and grazing using a low grazing intensity should be avoided.     

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.     

Changes in vegetation states in grazed and ungrazed Mackenzie Basin grasslands, New Zealand, 1990-2000

Changes in vegetation from 1990 to 2000 were examined at 10 high country localities, representing four grassland types: fescue tussock (Festuca novae-zelandiae), snow tussock (Chionochloa rigida), red tussock (C. rubra), and silver tussock (Poa cita). At each locality, three treatments were established: ambient sheep+rabbit grazing, rabbit grazing only, and no grazing. The mutivariate methods of classification and ordination were used on individual-quadrat cover data to define vegetation states and to examine transitions between them over time. Vegetation states in quadrats already dominated by Hieracium pilosella(> 50% cover) in 1990 showed little change in species composition regardless of grassland type and grazing treatment. In fescue tussock grassland, H. pilosellaincreased regardless of grazing treatment in states with low initial H. pilosellacover (< 5%), while the cover of Carex colensoi, Aira caryophyllea and Rumex acetosella decreased. In the single silver tussock locality, Poa citadecreased markedly in the ungrazed treatment as adventive species such as Dactylis glomerataand Echium vulgare increased. However, Poa citaalso decreased, probably due to drought, in the grazed treatment. Snow tussock and red tussock grassland states were more stable than those in short tussock grasslands, but there was also a general trend towards increasing H. pilosellacover in intertussock vegetation regardless of treatment. However, at one snow tussock locality, transitions from H. piloselladominated to C. rigida-dominated states occurred in ungrazed quadrats, while the reverse occurred in grazed vegetation. Implications for the management of tussock grasslands for conservation are discussed.     

Disproportionate effects of non‐colonial small herbivores on structure and diversity of grassland dominated by large herbivores

The response of semiarid grasslands to small, non‐colonial herbivores has received little attention, focusing primarily on the effects of granivore assemblages on annual plant communities. We studied the long‐term effects of both small and large herbivores on vegetation structure and species diversity of shortgrass steppe, a perennial semiarid grassland considered marginal habitat for small mammalian herbivores. We hypothesized that 1) large generalist herbivores would affect more abundant species and proportions of litter‐bare ground‐vegetation cover through non‐selective herbivory, 2) small herbivores would affect less common species through selective but limited consumption, and 3) herbivore effects on plant richness would increase with increasing aboveground net primary production (ANPP). Plant community composition was assessed over a 14‐year period in pastures grazed at moderate intensities by cattle and in exclosures for large (cattle) and large‐plus‐small herbivores (additional exclusion of rabbits and rodents). Exclusion of large herbivores affected litter and bare ground and basal cover of abundant, common and uncommon species. Additional exclusion of small herbivores did not affect uncommon components of the plant community, but had indirect effects on abundant species, decreased the cover of the dominant grass Bouteloua gracilis and total vegetation, and increased litter and species diversity. There was no relationship between ANPP and the intensity of effects of either herbivore body size on richness. Exclusion of herbivores of both body sizes had complementary and additive effects which promoted changes in vegetation composition and physiognomy that were linked to increased abundance of tall and decreased abundance of short species. Our findings show that small mammalian herbivores had disproportionately large effects on plant communities relative to their small consumption of biomass. Even in small‐seeded perennial grasslands with a long history of intensive grazing by large herbivores, non‐colonial small mammalian herbivores should be recognized as an important driver of grassland structure and diversity.     

Can re-establishment of cattle grazing restore bryophyte diversity in abandoned mesic semi-natural grasslands?

Changes of agricultural practices have led to decline of semi-natural habitats sustained by traditional animal husbandry in many European regions. The abandonment of semi-natural pastures leads to increase of vascular plant biomass and subsequent decline of weak competitors such as bryophytes. Re-establishing traditional animal husbandry may potentially restore biodiversity but the success of such measures remains insufficiently known. In this study, we asked if re-establishing cattle grazing on previously abandoned grasslands will restore their bryophyte communities. The effect of cattle grazing on bryophyte communities of mesic semi-natural grasslands was studied in south-western Finland in a comparison of (i) continuously grazed pastures, (ii) previously abandoned pastures where grazing was re-established during 1990s, and (iii) abandoned pastures, where grazing had ceased during late 1960s to early 1980s. The average cover, species richness, species density and species diversity of bryophytes were significantly higher in the continuously grazed than in the abandoned grasslands. Ordination analyses revealed clear differences also in community structure between the management classes. Re-established grasslands were ecologically heterogeneous and situated in between the continuously grazed and abandoned grasslands in all characteristics, indicating variable effect of the restoration measure. Seventeen bryophyte species were recognized as significant indicators of the three grassland classes, four of which could be used as indicators of valuable grassland habitats. Although there was variation in the consequences of re-introduction of grazing, the results give evidence of positive effect of grazing on regaining bryophyte diversity of abandoned grasslands.