Search

Questions In African savannas, Macrotermes termites contribute to small-scale heterogeneity by constructing large mounds. Operating as islands of high nutrient and water availability and low fire frequency, these mounds support distinct, diverse communities of trees that have been shown to be highly attractive to browsers. However, the distinct traits of tree species on termite mounds have hardly been studied, even though this may help to understand processes determining (1) their characteristic community structure and (2) attractiveness for browsers. Here, we compare functional trait and browser preference values between tree species on and off termite mounds. Location Hluhluwe-i Mfolozi Park, Kwazulu- Natal, South Africa. Methods We recorded tree community compositions for 16 large Macrotermes natalensis mounds and 16 control plots of 100 m2 each in a paired design. For each observed tree species we measured 22 traits, related to water and nutrient use, fire tolerance, light competition and anti-herbivore defence, and compared average trait values between mound and control communities. Furthermore, we investigated the feeding preferences of ungulate browsers for the most common tree species and how this was linked to their associated traits. Results Termite mounds supported tree communities that were distinct from the surrounding savanna vegetation. Mounds hosted more evergreen and less leguminous tree species than control communities, and the dominant species were less mechanically defended, less nutritious, had larger leaves and lower wood density than the species dominating control plots. Browsers preferred leguminous tree species with high leaf N and P content, which were relatively rare on termite mounds. Conclusions Overall, we conclude that termite mounds in this savanna form small refuges for tree species that seem less adapted to fire (more evergreens), have low nutrient availability (less nitrogen fixers) and suffer from water stress (larger leaf sizes) than typical savanna trees. Surprisingly, despite their reputation as browsing hotspots, the tree species dominating mounds are less nutritious and less preferred by browsers than tree species of the surrounding savanna, which may be explained by the relatively nutrient-rich nature of this savanna or intraspecific trait differences. [ABSTRACT FROM AUTHOR]

Herbivores may influence the nitrogen (N) recycling rates and consequently increase or decrease the productivity of grasslands. Plant N concentration emerged as a critical parameter to explain herbivore effects from several conceptual models, which predict that herbivores decrease soil N availability when plant N concentration is low whereas they increase it when plant N concentration is high ( Hobbs 1996 , Ritchie et al. 1998 , Pastor et al. 2006 ). However, a broader cross-site comparison among published studies to test these predictions is hampered by the different methodologies used to measure soil N availability or a proxy thereof, and a lack of measurements of plant N concentration. Therefore it remains unclear whether these model predictions are generally valid across a range of grasslands. We tested whether there is a relationship between plant N concentration and herbivore impact on soil N availability (measured with resin bags) with a study of replicate 6–8 year old exclosures (with an unfenced control) of vertebrate herbivores (>1 kg) established at each of seven grassland sites in North America and Europe. Contrary to model predictions, we found a negative relationship between the effect of herbivores on resin bag soil N availability and plant N concentration. Our study confirms the importance of plant N concentration as a predictor of herbivore effect on soil N availability across grasslands, but contradicts the models. A possible explanation may be that the results represent a transient situation as the exclosures were relatively young whereas the models may refer to an equilibrium state. Simultaneous measurements of both plant N concentration and herbivore effect on soil N availability from more grassland sites, preferably with contrasting plant N concentrations and including exclosures of different ages, should resolve the contrast between model predictions and our field measurements. [ABSTRACT FROM AUTHOR]

Light-demanding trees and thorny shrubs in temperate plant communities may reflect adaptations to now-extinct large grazers, such as aurochs and tarpans, rendering these adaptations ecological anachronisms. We explored the ecological functions of plant traits of Quercus robur and Prunus spinosa in areas grazed by cattle and horses, the domesticated descendants of aurochs and tarpans. Specifically, we tested the hypothesis that grazing induces a shifting mosaic of grassland, shrub thickets and woodlands through the key process of associational resistance: the protection of palatable young trees by thorny shrubs. An exclosure experiment with transplanted Q. robur seedlings revealed that Q. robur grew best in grassland exclosures and on the edge of thorny shrub thickets, which may be viewed as an optimal balance between sufficient protection from large herbivores and sufficient light availability. A cross-site comparison of four floodplain woodlands in north-western Europe showed that Q. robur can regenerate in the presence of large herbivores through spatial association with P. spinosa. However, we found that expansion of P. spinosa shrubs and Q. robur coincided with periods of low rabbit abundance and not with livestock density. From this, it appears that the process of associational resistance does not work with rabbits. With extensive grazing by large (domesticated) grazers in temperate floodplains, a shifting mosaic of grassland, shrubs and trees may develop that has high conservation value. Palatable, light-demanding Q. robur seedlings can successfully regenerate in spiny P. spinosa shrubs through associational resistance. This process does not offer protection from abundant small herbivores, such as rabbits, that can inhibit the recruitment of shrubs and trees in this mosaic vegetation. In floodplain meadows frequent flooding may be an efficient way to reduce rabbit populations, with dry conditions in summer and wet in winter. When floodplain meadows are combined with adjacent higher grounds, large herbivores can escape the floods through migration. [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]

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]

Grasslands are often characterised by small-scale mosaics in plant community composition that contribute to their diversity. Although above- and belowground biota can both cause such mosaics, few studies have addressed their interacting effects. We studied multi-trophic interactions between aboveground vertebrate grazers, subterranean ants, plant-pathogenic soil biota (especially nematodes) and the vegetation in a temperate grassland. We found that when rabbits and cattle locally omit vegetation patches, yellow ants (Lasius flavus ) respond to the taller vegetation by digging up more sand from deeper soil layers (hence making taller nest mounds), probably to maintain sufficiently high soil temperatures. We found that this ant digging affects other soil biota, as the mounds contain fewer plant-parasitic and fungivore nematodes. Also, the mounds have lower moisture content and soil bulk densities, and higher pH and available nutrient content than the directly surrounding soil. The clonal sedge Carex arenaria grows vigorously on the mounds, producing more shoots and shorter rhizome internode lengths than in surrounding vegetation. Other plant species, such as the grass Festuca rubra , dominate the surrounding vegetation. A greenhouse bioassay experiment revealed that harmful soil organisms (as plant-parasitic nematodes and pathogenic fungi) outweighed the effect of beneficial organisms (e.g., mycorrhizae) in this system. Rhizome biomass and shoot production of C. arenaria were indeed inhibited less by biota in soil from ant mounds than by biota in soil from the surrounding vegetation. However, the total biomass production of C. arenaria was inhibited as strongly in both soil types. F. rubra was inhibited more strongly by biota in the surrounding soil. We suggest that various direct and indirect interactions between above- and belowground biota can contribute to community mosaics and hence diversity in grasslands. [ABSTRACT FROM AUTHOR]

Evaluates an experiment where fertilizer application to mown grassland was stopped in an attempt to restore former species-rich vegetation. Crop and species composition of different fields after cessation of fertilizer application; Dynamics of species replacement under a Gaussian response model; Discusses of results with regard to diversity-productivity relationships, nutrient cycling and adaptation of species to nutrient-poor conditions.

The results of a restoration experiment carried out on a permanent grassland on peaty, heavy clay in the Netherlands are described. The experiment started in 1985, 7 years after fertilizer application had ceased, and was designed to provide insight into ecologically significant processes accompanying restoration. An analysis was made of the effect of management regime and of raising the water table on nutrient availability, dry matter production, tissue nutrient concentration, dynamics of species numbers and plant species replacement. Three management practices were compared: cutting and removal (RR), cutting and mulching (MM), sod removal in 1985, and thereafter cutting and removal of the hay (RS). Data are presented on changes during a 5-year period. No trend was discernible in soil pH, total C, N and P in the RR treatment; extractable P and K decreased sharply in the field with the raised groundwater level. Nine years after fertilizer application ceased, dry matter production had fallen from 10–11 to 6–7 t ha&sup-1; years&up-1;. In the subsequent 5 years of the experiment it declined to 5–5 t ha&sup-1; year&sup-1; when all cut biomass was removed, and to about 4 t ha&sup-1; year&sup-1; after sod removal. Mulching caused an increase to 11 t ha&sup-1; year&sup-1;. No effect was seen of the raised water level. The dry matter yield of the first June cut in the RR treatment decreased. The tissue K concentration also decreased, but no increase of the tissue P concentration was detected. It was concluded that the availability of K and to some extent of P was more important than N availability in explaining the decrease in dry matter production. The tissue nutrient concentrations were not influenced by the water table. Sod removal to a depth of 5 cm resulted in the lowest productivity and the lowest tissue concentrations of P, while tissue concentrations of N and K were not affected. Raising the water level resulted in a more rapid establishment of species indicative of wet conditions, some of which invaded from nearby ditches. The: trends of dominant species are described with a set of response models. The species were ranked from disappearing to colonizing species. The relationship between rank order of replacement and indicator values of species was investigated. Raising: the water table resulted in species indicative of wet conditions becoming dominant, independently of vegetation management. The removal of nutrients resulted in the appearance of species with a lower maximal height, indicative of lower P and K availability. [ABSTRACT FROM AUTHOR]

Analyzes the germination of 91 herbaceous species in response to a temperature gradient and to alternating temperature and stratification. Principal components analysis of the species; Changes in canopy structure; Relation of changes in species composition of species present at different stages to changes in germination attributes.