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Ontogenetic niche shifts have helped to understand population dynamics. Here we show that ontogenetic niche shifts also offer an explanation, complementary to traditional concepts, as to why certain species show seasonal migration. We describe how demographic processes (survival, reproduction and migration) and associated ecological requirements of species may change with ontogenetic stage (juvenile, adult) and across the migratory range (breeding, non-breeding). We apply this concept to widely different species (dark-bellied brent geese (Branta b. bernicla), humpback whales (Megaptera novaeangliae) and migratory Pacific salmon (Oncorhynchus gorbuscha) to check the generality of this hypothesis. Consistent with the idea that ontogenetic niche shifts are an important driver of seasonal migration, we find that growth and survival of juvenile life stages profit most from ecological conditions that are specific to breeding areas. We suggest that matrix population modelling techniques are promising to detect the importance of the ontogenetic niche shifts in maintaining migratory strategies. As a proof of concept, we applied a first analysis to resident, partial migratory and fully migratory populations of barnacle geese (Branta leucopsis). We argue that recognition of the costs and benefits of migration, and how these vary with life stages, is important to understand and conserve migration under global environmental change.

After an historical absence, over the last decades Eurasian Spoonbills Platalea leucorodia leucorodia have returned to breed on the barrier islands of the Wadden Sea. The area offers an abundance of predator-free nesting habitat, low degrees of disturbance, and an extensive intertidal feeding area with increasing stocks of brown shrimp Crangon crangon , the assumed main prey of P. leucorodia leucorodia . Nevertheless, newly established and expanding colonies of spoonbills have surprisingly quickly reached plateau levels. Here we tested the often stated assertion that spoonbills mainly rely on brown shrimp as food, by quantifying the diet of chicks on the basis of regurgitates and by analysis of blood isotopes using stable isotope Bayesian mixing models. Both methods showed that, rather than brown shrimp being the staple food of spoonbill chicks, small flatfish (especially plaice Pleuronectes platessa ) and gobies ( Pomatoschistus spp.) were their main prey. Unlike shrimp, small flatfish have been reported to be rather scarce in the Wadden Sea in recent years, which may explain the rapid saturation of colony size due to food-related density-dependent recruitment declines of growing colonies. By way of their diet and colony growth characteristics, spoonbills may thus indicate the availability of small fish in the Wadden Sea. We predict that the recovery to former densities of young flatfish and other juvenile/small fish in the Wadden Sea will be tracked by changing diets (more fish) and an increase in the size of Eurasian Spoonbill colonies across the Wadden Sea.

Savanna grasslands are characterized by high spatial heterogeneity in vegetation structure, aboveground biomass and nutritional quality, with high quality short-grass grazing lawns forming mosaics with patches of tall bunch grasses of lower quality. This heterogeneity can arise because of local differences in consumption, because of differences in productivity, or because both processes enforce each other (more production and consumption). However, the relative importance of both processes in maintaining mosaics of lawn and bunch grassland types has not been measured. Also their interplay been not been assessed across landscape gradients. In a South African savanna, we, therefore, measured the seasonal changes in primary production, nutritional quality and herbivore consumption (amount and percentage) of grazing lawns and adjacent bunch grass patches across a rainfall gradient. We found both higher amounts of primary production and, to a smaller extent, consumption for bunch grass patches. In addition, for bunch grasses primary production increased towards higher rainfall while foliar nitrogen concentrations decreased. Foliar nitrogen concentrations of lawn grasses decreased much less with increasing rainfall. Consequently, large herbivores targeted the biomass produced on grazing lawns with on average 75 % of the produced biomass consumed. We conclude that heterogeneity in vegetation structure in this savanna ecosystem is better explained by small-scale differences in productivity between lawn and bunch grass vegetation types than by local differences in consumption rates. Nevertheless, the high nutritional quality of grazing lawns is highly attractive and, therefore, important for the maintenance of the heterogeneity in species composition (i.e. grazing lawn maintenance).

African savannahs are among the few places on earth where diverse communities of mega- and meso-sized ungulate grazers dominate ecosystem functioning. Less conspicuous, but even more diverse, are the communities of herbivorous insects such as grasshoppers, which share the same food. Various studies investigated the community assembly of these groups separately, but it is poorly known how ungulate communities shape grasshopper communities. Here, we investigated how ungulate species of different body size alter grasshopper communities in a South African savannah. White rhino is the most abundant vertebrate herbivore in our study site. Other common mesoherbivores include buffalo, zebra and impala. We hypothesized that white rhinos would have greater impact than mesoherbivores on grasshopper communities. Using 10-year-old exclosures, at eight sites we compared the effects of ungulates on grasshopper communities in three nested treatments: (i) unfenced plots ('control plots') with all vertebrate herbivores present, (ii) plots with a low cable fence, excluding white rhino ('megaherbivore exclosures'), and (iii) plots with tall fences, excluding all herbivores larger than rodents ('complete ungulate exclosures'). In each plot, we collected data of vegetation structure, grass and grasshopper community composition. Complete ungulate exclosures contained 30% taller vegetation than megaherbivore exclosures and they were dominated by different grass and grasshopper species. Grasshoppers in complete ungulate exclosures were on average 3.5 mm longer than grasshoppers in megaherbivore exclosures, possibly due to changes in plant communities or vegetation structure. We conclude that surprisingly, in this megaherbivore hotspot, mesoherbivores, instead of megaherbivores, most strongly affect grasshopper communities.

In community ecology, broad-scale spatial replication can accommodate contingencies in patterns within species groups, but contingencies in processes across species groups remain problematic. Here, based on a focused review of grazing and browsing by large mammals, we use one trophic guild as a "control" for the other to identify generalities that are not contingent upon specific consumer-resource interactions. An example of such a generality is the Jarman-Bell principle, which explains how allometries of metabolism and digestion influence dietary tolerance and thereby enable resource partitioning within both guilds at multiple scales. By comparing the grazing succession with browsing stratification we show how competition from smaller herbivores, rather than facilitation from larger ones, is the underlying process structuring ungulate assemblages when shared resources become limiting. Also, grazing lawns and browsing hedges are functionally similar. In each case, plants expressing tolerance traits can withstand chronic grazing or browsing in sites where the nutritive value of the local food resource is enhanced in positive feedback to the actions of its consumers. The debate over whether ungulates accelerate or decelerate nutrient cycling can be resolved by comparing grazing and browsing effects in the same ecosystem type. Evidence from African savannas points to the rate of nutrient cycling being controlled by the mix of tolerance and resistance traits in plants; not the relative dominance of grazing or browsing by local herbivores. We recommend this across-guild comparative approach as a novel solution with widespread utility for resolving contingencies in community processes.

Studies addressing the role of large herbivores on nitrogen cycling in grasslands have suggested that the direction of effects depends on soil fertility. Via selection for high quality plant species and input of dung and urine, large herbivores have been shown to speed up nitrogen cycling in fertile grassland soils while slowing down nitrogen cycling in unfertile soils. However, recent studies show that large herbivores can reduce nitrogen mineralization in some temperate fertile soils, but not in others. To explain this, we hypothesize that large herbivores can reduce nitrogen mineralization in loamy or clay soils through soil compaction, but not in sandy soils. Especially under wet conditions, strong compaction in clay soils can lead to periods of soil anoxia, which reduces decomposition of soil organic matter and, hence, N mineralization. In this study, we use a long-term (37-year) field experiment on a salt marsh to investigate the hypothesis that the effect of large herbivores on nitrogen mineralization depends on soil texture. Our results confirm that the presence of large herbivores decreased nitrogen mineralization rate in a clay soil, but not in a sandy soil. By comparing a hand-mown treatment with a herbivore-grazed treatment, we show that these differences can be attributed to herbivore-induced changes in soil physical properties rather than to above-ground biomass removal. On clay soil, we find that large herbivores increase the soil water-filled porosity, induce more negative soil redox potentials, reduce soil macrofauna abundance, and reduce decomposition activity. On sandy soil, we observe no changes in these variables in response to grazing. We conclude that effects of large herbivores on nitrogen mineralization cannot be understood without taking soil texture, soil moisture, and feedbacks through soil macrofauna into account.

Aboveground and belowground organisms influence plant community composition by local interactions, and their scale of impact may vary from millimeters belowground to kilometers aboveground. However, it still poorly understood how large grazers that select their forage on large spatial scales interact with small-scale aboveground-belowground interactions on plant community heterogeneity. Here, we investigate how cattle (Bos taurus) modify the effects of interactions between yellow meadow ants (Lasius flavus) and European brown hares (Lepus europaeus) on the formation of small-scale heterogeneity in vegetation composition. In the absence of cattle, hares selectively foraged on ant mounds, while under combined grazing by hares and cattle, vertebrate grazing pressure was similar on and off mounds. Ant mounds that were grazed by only hares had a different plant community composition compared to their surroundings: the cover of the grazing-intolerant grass Elytrigia atherica was reduced on ant mounds, whereas the relative cover of the more grazing-tolerant and palatable grass Festuca rubra was enhanced. Combined grazing by hares and cattle, resulted in homogenization of plant community composition on and off ant mounds, with high overall cover of F. rubra. We conclude that hares can respond to local ant-soil-vegetation interactions, because they are small, selective herbivores that make their foraging decisions on a local scale. This results in small-scale plant patches on mounds of yellow meadow ants. In the presence of cattle, which are less selective aboveground herbivores, local plant community patterns triggered by small-scale aboveground-belowground interactions can disappear. Therefore, cattle modify the consequences of aboveground-belowground interactions for small-scale plant community composition.

A key aspect of savannah vegetation heterogeneity is mosaics formed by two functional grassland types, bunch grasslands, and grazing lawns. We investigated the role of termites, important ecosystem engineers, in creating high-nutrient patches in the form of grazing lawns. Some of the ways termites can contribute to grazing lawn development is through erosion of soil from aboveground mounds to the surrounding soil surface. This may alter the nutrient status of the surrounding soils. We hypothesize that the importance of this erosion varies with termite genera, depending on feeding strategy and mound type. To test this, we simulated erosion by applying mound soil from three termite genera ( Macrotermes , Odontotermes , and Trinervitermes ) in both a field experiment and a greenhouse experiment. In the greenhouse experiment, we found soils with the highest macro nutrient levels (formed by Trinervitermes ) promoted the quality and biomass of both a lawn ( Digitaria longiflora ) and a bunch ( Sporobolus pyramidalis ) grass species. In the field we found that soils with the highest micro nutrient levels (formed by Macrotermes ) showed the largest increase in cover of grazing lawn species. By linking the different nutrient availability of the mounds to the development of different grassland states, we conclude that the presence of termite mounds influences grassland mosaics, but that the type of mound plays a crucial role in determining the nature of the effects.

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.

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.

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.

Recent discussions on determinants of competitive success during succession require the study of the combined effect of light and nutrient availability on growth and allocation. These effects can be used to predict the outcome of competition at changing resource availabilities. This work is part of a study on the successional sequence in permanent grassland starting after fertilizer application is stopped, but with continued mowing, in order to restore former species-rich communities. This yields a successional sequence which proceeds from grasslands with a high nutrient availability and a closed canopy, to grasslands with a low nutrient availability and an open canopy. If allocation is related to competitive ability, species from the productive stages would be expected to allocate more biomass and nitrogen to leaves, which could make them better competitors for light, while species from the unproductive stages would allocate more biomass to roots, which could make them better nutrient competitors. This study reports on growth, specific leaf area (SLA), vertical display of leaves, and allocation of biomass and nitrogen of six grassland species from this successional sequence at 16 combinations of light and nutrient supply. Species from the poorer successional stages reached a lower final dry weight than species from the richer stages, over all treatment combinations. The experimental design made it possible to test for unique effects of the resource ratio effect of light and nutrients on allocation characteristics. This resource-ratio effect was defined as the ratio light intensity/(light intensity + nutrient supply rate), using standardized levels for the treatments. The within-species variation (plasticity) in both allocation of dry matter and nitrogen was linearly related to this resource-ratio effect. Some interspecific differences in this relationship were found which could be related to the position of the species along the successional gradient. However, the range of plasticity in allocation pattern expressed within each species was much larger than the differences between species. It was concluded that allocation differences between these grassland species are relatively unimportant, given the large amount of plasticity in these traits. Interspecific differences in SLA and vertical stature seemed to be more important in explaining the position of species along the successional gradient.

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.