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The coexistence of different species of large herbivores (ungulates) in grasslands and savannas has fascinated ecologists for decades. However, changes in climate, land‐use and trophic structure of ecosystems increasingly jeopardise the persistence of such diverse assemblages. Body size has been used successfully to explain ungulate niche differentiation with regard to food requirements and predation sensitivity. But this single trait axis insufficiently captures interspecific differences in water requirements and thermoregulatory capacity and thus sensitivity to climate change. Here, we develop a two‐dimensional trait space of body size and minimum dung moisture content that characterises the combined food and water requirements of large herbivores. From this, we predict that increased spatial homogeneity in water availability in drylands reduces the number of ungulate species that will coexist. But we also predict that extreme droughts will cause the larger, water‐dependent grazers as wildebeest, zebra and buffalo–dominant species in savanna ecosystems – to be replaced by smaller, less water‐dependent species. Subsequently, we explore how other constraints such as predation risk and thermoregulation are connected to this two‐dimensional framework. Our novel framework integrates multiple simultaneous stressors for herbivores and yields an extensive set of testable hypotheses about the expected changes in large herbivore community composition following climate change. [ABSTRACT FROM AUTHOR]

Coastal food webs can be supported by local benthic or pelagic primary producers and by the import of organic matter. Distinguishing between these energy sources is essential for our understanding of ecosystem functioning. However, the relative contribution of these components to the food web at the landscape scale is often unclear, as many studies lack good taxonomic and spatial resolution across large areas. Here, using stable carbon isotopes, we report on the primary carbon sources for consumers and their spatial variability across one of the world's largest intertidal ecosystems (Dutch Wadden Sea; 1460 km2 intertidal surface area), at an exceptionally high taxonomic (178 species) and spatial resolution (9,165 samples from 839 locations). The absence of overlap in δ13C values between consumers and terrestrial organic matter suggests that benthic and pelagic producers dominate carbon input into this food web. In combination with the consistent enrichment of benthic primary producers (δ13C −16.3‰) relative to pelagic primary producers (δ13C −18.8) across the landscape, this allowed the use of a two-food-source isotope-mixing model. This spatially resolved modelling revealed that benthic primary producers (microphytobenthos) are the most important energy source for the majority of consumers at higher trophic levels (worms, molluscs, crustaceans, fish, and birds), and thus to the whole food web. In addition, we found large spatial heterogeneity in the δ13C values of benthic primary producers (δ13C −19.2 to −11.5‰) and primary consumers (δ13C −25.5 to −9.9‰), emphasizing the need for spatially explicit sampling of benthic and pelagic primary producers in coastal ecosystems. Our findings have important implications for our understanding of the functioning of ecological networks and for the management of coastal ecosystems. [ABSTRACT FROM AUTHOR]

ABSTRACT Both arthropods and large grazing herbivores are important components and drivers of biodiversity in grassland ecosystems, but a synthesis of how arthropod diversity is affected by large herbivores has been largely missing. To fill this gap, we conducted a literature search, which yielded 141 studies on this topic of which 24 simultaneously investigated plant and arthropod diversity. Using the data from these 24 studies, we compared the responses of plant and arthropod diversity to an increase in grazing intensity. This quantitative assessment showed no overall significant effect of increasing grazing intensity on plant diversity, while arthropod diversity was generally negatively affected. To understand these negative effects, we explored the mechanisms by which large herbivores affect arthropod communities: direct effects, changes in vegetation structure, changes in plant community composition, changes in soil conditions, and cascading effects within the arthropod interaction web. We identify three main factors determining the effects of large herbivores on arthropod diversity: ( i) unintentional predation and increased disturbance, ( ii) decreases in total resource abundance for arthropods (biomass) and ( iii) changes in plant diversity, vegetation structure and abiotic conditions. In general, heterogeneity in vegetation structure and abiotic conditions increases at intermediate grazing intensity, but declines at both low and high grazing intensity. We conclude that large herbivores can only increase arthropod diversity if they cause an increase in (a)biotic heterogeneity, and then only if this increase is large enough to compensate for the loss of total resource abundance and the increased mortality rate. This is expected to occur only at low herbivore densities or with spatio-temporal variation in herbivore densities. As we demonstrate that arthropod diversity is often more negatively affected by grazing than plant diversity, we strongly recommend considering the specific requirements of arthropods when applying grazing management and to include arthropods in monitoring schemes. Conservation strategies aiming at maximizing heterogeneity, including regulation of herbivore densities (through human interventions or top-down control), maintenance of different types of management in close proximity and rotational grazing regimes, are the most promising options to conserve arthropod diversity. [ABSTRACT FROM AUTHOR]

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]

Trait-based community assembly theory suggests that trait variation among co-occurring species is shaped by two main processes: abiotic filtering, important in stressful environments and promoting similarity, and competition, more important in productive environments and promoting dissimilarity. Previous studies have indeed found trait similarity to decline along productivity gradients. However, these studies have always been done on single trophic levels. Here, we investigated how interactions between trophic levels affect trait similarity patterns along environmental gradients. We propose three hypotheses for the main drivers of trait similarity patterns of plants and herbivores along environmental gradients: (1) environmental control of both, (2) bottom-up control of herbivore trait variation, and (3) top-down control of grass trait variation. To test this, we collected data on the community composition and trait variation of grasses (41 species) and grasshoppers (53 species) in 50 plots in a South African savanna. Structural equation models were used to investigate how the range and spacing of within-community functional trait values of both grasses and their insect herbivores (grasshoppers; Acrididae) respond to (1) rainfall and fire frequency gradients and (2) the trait similarity patterns of the other trophic level. The analyses revealed that traits of co-occurring grasses became more similar toward lower rainfall and higher fire frequency (environmental control), while showing little evidence for top-down control. Grasshopper trait range patterns, on the other hand, were mostly directly driven by vegetation structure and grass trait range patterns (bottom-up control), while environmental factors had mostly indirect effects via plant traits. Our study shows the potential to expand trait-based community assembly theory to include trophic interactions. [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]

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]

Presents a study which looks at the interactions between soil changes, vegetation and herbivory, since different stages of salt marsh. Detailed information on the methodology used to conduct the study; Discussion based on the results of the study.

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.

Studies the species dynamics and nutrient accumulation during early primary succession in coastal sand dunes on the Dutch island of Schiermonnikoog. Determinants of variance in species composition; Influence of nitrogen in limiting above aground biomass production; Comparison of nitrogen content in the soil organic layer in the plains versus sand dunes.

The sandy seabed of shallow coastal shelf seas displays morphological patterns of various dimensions. The seabed also harbors a rich ecosystem. Increasing pressure from human‐induced disturbances necessitates further study on drivers of benthic community distributions over morphological patterns. Moreover, a greater understanding of the sand ripple distribution over tidal sand waves may improve morphological model predictions. Here we analyzed the biotic abundance and ripple morphology in sand wave troughs and crests using video transects. We found that both the epibenthos and endobenthos are significantly more abundant in sand wave troughs, where ripples are less abundant and more irregularly shaped. Finally, we show that camera systems are relatively quick and effective tools to study biotic spatial patterns in relation to seabed morphology. Plain Language Summary: Coastal seabeds are important areas for human activities and constructions and are also inhabited by many organisms, collectively referred to as benthos. The interaction of tides, wind, sand and biology leads to the growth of large, rhythmic seabed patterns. They can grow up to several meters high, hundreds of meters long, and shift a few meters per year. Computer models can predict this behavior, but they lack understanding of biological processes and variations in sand ripple patterns. To increase the understanding of these effects, we present findings from two video transects, for which we lowered a video camera to the seabed and recorded both organisms and sand ripples for ~200 m. We show that, relative to crests, troughs have more benthic organisms, and sand ripples are often absent or less regularly shaped. Finally, we show that a camera is a quick and effective tool to study benthic spatial patterns. Key Points: Video transects over tidal sand waves yielded detailed spatial information of both benthic organisms and small‐scale seabed morphologyBoth epibenthos and endobenthos are more abundant in sand wave troughs compared to the crestsSand ripples are less abundant and more often irregularly shaped in the troughs, suggesting variations in seabed roughness over sand waves [ABSTRACT FROM AUTHOR]