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The succession of species-rich wetland vegetation in dune valleys into species-poor dwarf shrub vegetation was followed by means of permanent vegetation plots, in which the cover of vascular plant, moss and lichen species were recorded over a period of up to 33 years. Low density cattle grazing is an effective substitute for rabbits in stalling this succession, thus preserving the local plant species diversity. The influence of direct rabbit grazing pressure on the vegetation was studied in exclosures. The differences were significant, but not long-lasting after the exclosures had been opened again to the rabbits. Cladina lichen-rich Empetrum-heathland in the dunes is a stage in a probably cyclic succession that might be triggered by grazing. The total vascular plant, moss and lichen diversity of the dune ecosystem can be maintained by a combination of extensive cattle grazing and a regular but limited re-creation of pioneer situations. [ABSTRACT FROM AUTHOR]

The relative importance of predation risk and food quality on spatial grazing pressure and activity patterns was tested in a central-place foraging herbivore: the European rabbit. Rabbits grazed less with increasing distance from their burrows into adjacent grassland, thereby creating a gradient of increasing vegetation height and plant biomass and decreasing plant nutrient concentration. When nitrogen concentration was experimentally increased by 150% through fertilizing and mowing, rabbits visited these plots four times more frequently than the untreated control plots. Addition of predator scent (mink pellets) did not result in different patch use by rabbits. The combined addition of fertilizer and mink pellets had the same effect as addition of fertilizer alone. However, the mink pellets changed the temporal activity pattern of rabbits as measured with infrared detectors. Rabbits were predominantly nocturnal but shifted their activities to the day when mink pellets were added, resulting in equal activities during night and day. We conclude that rabbits are sensitive to perceived predation risk, but that this does not influence their spatial grazing pressure. A selection for the highest food quality on the other hand can explain the observed natural rabbit grazing gradient. Food quality was highest close to the burrows, therefore rabbits selecting for high quality food should forage most intensely close to the burrows and only move further away for higher quality items or when the vegetation close to their burrows is depleted. Through intensive grazing close to the burrows rabbits facilitated for themselves either through stimulating fresh protein rich re-growth or the return of nutrients through faeces or both. This is in contrast with central-place foraging theory where intense feeding close to the burrow is assumed to lead to reduced food resources. [ABSTRACT FROM AUTHOR]

Herbivores are reported to slow down as well as enhance nutrient cycling in grasslands. These conflicting results may be explained by differences in herbivore type. In this study we focus on herbivore body size as a factor that causes differences in herbivore effects on N cycling. We used an exclosure set-up in a floodplain grassland grazed by cattle, rabbits and common voles, where we subsequently excluded cattle and rabbits. Exclusion of cattle lead to an increase in vole numbers and a 1.5-fold increase in net annual N mineralization at similar herbivore densities (corrected to metabolic weight). Timing and height of the mineralization peak in spring was the same in all treatments, but mineralization in the vole-grazed treatment showed a peak in autumn, when mineralization had already declined under cattle grazing. This mineralization peak in autumn coincides with a peak in vole density and high levels of N input through vole faeces at a fine-scale distribution, whereas under cattle grazing only a few patches receive all N and most experience net nutrient removal. The other parameters that we measured, which include potential N mineralization rates measured under standardized laboratory conditions and soil parameters, plant biomass and plant nutrient content measured in the field, were the same for all three grazing treatments and could therefore not cause the observed difference. When cows were excluded, more litter accumulated in the vegetation. The formation of this litter layer may have added to the higher mineralization rates under vole grazing, through enhanced nutrient return through litter or through modification of microclimate. We conclude that different-sized herbivores have different effects on N cycling within the same habitat. Exclusion of large herbivores resulted in increased N annual mineralization under small herbivore grazing. [ABSTRACT FROM AUTHOR]

In this study we analysed the effects of large herbivores on smallrodent communities in different habitats using large herbivore exclosures. Westudied the effects of three year grazing introduction by red deer( Cervus elaphus L.) in previously ungrazed pine and oakwoodland and the exclusion of grazing by red deer, roe deer( Capreoluscapreolus L.) and mouflon ( Ovis ammon musiminL.) in formerly, heavily grazed pine woodland and heathland. At eight exclosuresites within each habitat type, small rodents were captured with live trapsusing trapping grids. At each trapping grid, seed plots of beechnuts( Fagus sylvatica L.) and acorns ( Quercusrobur L.) were placed to measure seed predation by rodents.Exclusion of grazing by large herbivores in formerly, heavily grazedhabitats had a significant effect on small rodent communities. Insideexclosureshigher densities of mainly wood mice ( Apodemus sylvaticusL.) and field voles ( Microtus agrestis L.) were captured.Introduction of grazing by red deer appeared to have no significant negativeeffects on small rodent communities. The seed predation intensity of beechnutsand acorns by small rodents was significantly higher in ungrazed situations,particularly in habitats that were excluded from grazing. The differencesbetween grazing introduction and exclusion effects on small rodent communitiescan be explained by differences in vegetation structure development. Therecovery of heavily browsed understory vegetation after large herbivore grazingexclusion proceeded faster than the understory degradation due to grazingintroduction. Small rodents depend on structural rich vegetations mainly forshelter. We conclude that large herbivores can have significant effects onvegetation dynamics not only via direct plant consumption but also throughindirect effects by reducing the habitat quality of small rodent habitats. [ABSTRACT FROM AUTHOR]

We analyzed the dynamics of dominant plant species in a grazed grassland over 17 years, and investigated whether local shifts in these dominant species, leading to vegetation mosaics, could be attributed to interactions between plants and soil-borne pathogens. We found that Festuca rubra and Carex arenaria locally alternated in abundance, with different sites close together behaving out of phase, resulting in a shifting mosaic. The net effect of killing all soil biota on the growth of these two species was investigated in a greenhouse experiment using gamma radiation, controlling for possible effects of sterilization on soil chemistry. Both plant species showed a strong net positive response to soil sterilization, indicating that pathogens (e.g., nematodes, pathogenic fungi) outweighed the effect of mutualists (e.g., mycorrhizae). This positive growth response towards soil sterilization appeared not be due to effects of sterilization on soil chemistry. Growth of Carex was strongly reduced by soil-borne pathogens (86% reduction relative to its growth on sterilized soil) on soil from a site where this species decreased during the last decade (and Festuca increased), while it was reduced much less (50%) on soil from a nearby site where it increased in abundance during the last decade. Similarly, Festuca was reduced more (67%) on soil from the site where it decreased (and Carex increased) than on soil from the site where it increased (55%, the site where Carex decreased). Plant-feeding nematodes showed high small-scale variation in densities, and we related this variation to the observed growth reductions in both plant species. Carex growth on unsterilized soil was significantly more reduced at higher densities of plant-feeding nematodes, while the growth reduction in Festuca was independent of plant-feeding nematode densities. At high plant-feeding nematode densities, growth of Carex was reduced more than Festuca, while at low nematode densities the opposite was found. Each plant species thus seems to be affected by different (groups of) soil-borne pathogens. The resulting interaction web of plants and soil-borne pathogens is discussed. We hypothesize that soil disturbances by digging ants and rabbits may explain the small-scale variation in nematode densities, by locally providing “fresh” sand. We conclude that soil-borne pathogens may contribute to plant diversity and spatial mosaics of plants in grasslands. [ABSTRACT FROM AUTHOR]

A study was conducted to analyse the effect of habitat productivity on woody species colonisation. Three soil types were distinguished: a relatively poor sandy soil type (1), a somewhat richer sandy type (2) and a relatively rich sandy loamy type (3). Chronosequences were established on these three soil types of 38 (type 1), 20 (type 2) and 54 years (type 3) after abandonment. In total 117 vegetation relevées were used to analyse life form change and species responses during old field succession via regression models. On the rich soil type the colonisation rate of woody species was slower than on the poor soil type. This can be explained by higher abundances of perennial species during the first 20 years after abandonment on the rich soil type in contrast to the poor soil type. Perennial species may delay the woody species colonisation. First they close the bare ground which inhibits germination and next they compete with woody seedlings for light, water and nutrients. The effect of habitat productivity on woody species colonisation can only be determined appropriately by taking life history traits into account. Early successional 'pioneer' woody species dispersed by wind have less difficulties colonising old fields than late successional 'forest' species; they colonise old fields prior to the development of a dense perennial sward. Forest species depend on animals to be dispersed which are attracted by vegetation structure. In ± 30 years on the poor soil type and in ± 45 years on the rich soil type woody species become dominant relative to other life forms. Forest species like Quercus robur L. invaded relatively early (<5 years) in contrast to other studies which probably coincides with the distance to seed sources (forest edges). [ABSTRACT FROM AUTHOR]

Vegetation and soil development during succession in coastal dune slacks on Terschelling island, the Netherlands, was investigated, by comparing neighbouring ecosystems on similar substrates that had been developing for 1, 5, 35 and 76 years since the vegetation and organic soil layer had been removed. In this successional sequence, soil organic matter accumulated rapidly due to the production of litter and dead roots. N mineralization was extremely low, increasing from 0.2 g m-2 yr-1 after 5 years to 0.8 g m-2 yr-1 after 76 years. It was accompanied by a decline in the pH (KCl) in the upper 10 cm of the soil from 6.8 to 4.4. Most of the above-ground biomass accumulated in the shrub species Oxycoccus macrocarpos and Salix repens. The 5- year-old plots harboured many plant species (18 species per 0.25 m2), but plant species diversity was much lower in the older plots. It is concluded that most changes in species composition and the decline in diversity occurred because early successional plant species were gradually outshaded by the thick litter layer and the accumulated shrub biomass. [ABSTRACT FROM AUTHOR]

An attempt was made to relate variation in life-history characteristics within a population of Plantago major ssp. pleiosperma to small-scale environmental variability. At a beach plain, embanked in 1966, a mosaic environment was distinguished with spatial variability in vegetation structure as well as in nutrient availability and water content of the soil. Differences between three subsites in comtemporary selection were demonstrated, e.g. in shoot morphology and allocation to reproductive tissue. The effects of nutrient supply and waterlogging on morphology and life history were studied on lines from the three subsites in a greenhouse. For most of the traits high levels of phenotypic plasticity were observed, covering almost entirely the observed phenotypic variability at the beach plain. In all treatments lines from the shrubs had, however, a higher leaf-area ratio as well as delayed flowering when compared to lines from more open subsites. In addition, in a reciprocal transplant experiment it was demonstrated that lines from the shrubs had larger shoots with e.g. broader leaves in the shady environment of the shrubs than other lines. From the experiments no indications were obtained that lines from any subsite were especially adapted to specific levels of nutrient supply or water content of the soil. With respect to these environmental factors P. major ssp. pleisoperma might occur and reproduce at all subsites by means of phenotypic plasticity, e.g. in plant form. However, it is suggested that spatial variability in vegetation structure caused a population subdivision in allocation patterns, leaf form and life history at the beach plain, over distances of about 15-25 m. This differentation occurred during primary succession over a period of twenty years. [ABSTRACT FROM AUTHOR]

Herbivores alter plant biodiversity (species richness) in many of the world's ecosystems, but the magnitude and the direction of herbivore effects on biodiversity vary widely within and among ecosystems. One current theory predicts that herbivores enhance plant biodiversity at high productivity but have the opposite effect at low productivity. Yet, empirical support for the importance of site productivity as a mediator of these herbivore impacts is equivocal. Here, we synthesize data from 252 large-herbivore exclusion studies, spanning a 20-fold range in site productivity, to test an alternative hypothesis-that herbivore-induced changes in the competitive environment determine the response of plant biodiversity to herbivory irrespective of productivity. Under this hypothesis, when herbivores reduce the abundance (biomass, cover) of dominant species (for example, because the dominant plant is palatable), additional resources become available to support new species, thereby increasing biodiversity. By contrast, if herbivores promote high dominance by increasing the abundance of herbivory-resistant, unpalatable species, then resource availability for other species decreases reducing biodiversity. We show that herbivore-induced change in dominance, independent of site productivity or precipitation (a proxy for productivity), is the best predictor of herbivore effects on biodiversity in grassland and savannah sites. Given that most herbaceous ecosystems are dominated by one or a few species, altering the competitive environment via herbivores or by other means may be an effective strategy for conserving biodiversity in grasslands and savannahs globally.