Plant communities are often dispersal-limited and zoochory can be an efficient mechanism for plants to colonize new patches of potentially suitable habitat. We predicted that seed dispersal by ungulates acts as an ecological filter -which differentially affects individuals according to their characteristics and shapes species assemblages -and that the filter varies according to the dispersal mechanism (endozoochory, fur-epizoochory and hoof-epizoochory). We conducted two-step individual participant data meta-analyses of 52 studies on plant dispersal by ungulates in fragmented landscapes, comparing eight plant traits and two habitat indicators between dispersed and non-dispersed plants. We found that ungulates dispersed at least 44% of the available plant species. Moreover, some plant traits and habitat indicators increased the likelihood for plant of being dispersed. Persistent or nitrophilous plant species from open habitats or bearing dry or elongated diaspores were more likely to be dispersed by ungulates, whatever the dispersal mechanism. In addition, endozoochory was more likely for diaspores bearing elongated appendages whereas epizoochory was more likely for diaspores released relatively high in vegetation. Hoof-epizoochory was more likely for light diaspores without hooked appendages. Furepizoochory was more likely for diaspores with appendages, particularly elongated or hooked ones. We thus observed a gradient of filtering effect among the three dispersal mechanisms. Endozoochory had an effect of rather weak intensity (impacting six plant characteristics with variations between ungulate-dispersed and non-dispersed plant species mostly below 25%), whereas hoof-epizoochory had a stronger effect (eight characteristics included five ones with above 75% variation), and fur-epizoochory an even stronger one (nine characteristics included six ones with above 75% variation). Our results demonstrate that seed dispersal by ungulates is an ecological filter whose intensity varies according to the dispersal mechanism considered. Ungulates can thus play a key role in plant community dynamics and have implications for plant spatial distribution patterns at multiple scales.Plant communities are often dispersal-limited and zoochory can be an efficient mechanism for plants to colonize new patches of potentially suitable habitat. Our analysis is the first synthesis of ungulate seed dispersal that compares characteristics from both non-dispersed and dispersed diaspores, distinguishing the three zoochory mechanisms ungulates are involved in: endozoochory, hoof-epizoochory and fur-epizoochory. We confirmed that seed dispersal by ungulates is an ecological filter whose intensity increases from endozoochory, then hoof-epizoochory to finally fur-epizoochory. By filtering seed traits through dispersal, ungulates can thus play a key role in plant community dynamics and have implications for plant spatial distribution patterns at multiple scales. Synthesis

Aim In several ecosystems, the diversity of functional species traits has been shown to have a stronger effect on ecosystem functioning than taxonomic diversity alone. However, few studies have explored this idea at a large geographical scale. In a multisite experiment, we unravelled the relationship between ecosystem function and functional completeness of species assemblages using dung beetles as a model group, focusing on dung removal and secondary seed dispersal. Location Seventeen grassland locations across the Western Palaearctic. Methods We used a randomized block design with different exclosure types to control the dung and seed removing activities of individual functional groups of the local dung beetle assemblage. We classified dung beetle species according to resource specialization and into functional groups based on dung processing behaviour (dwellers, tunnellers, rollers) and body size (small, large). Additionally, we assessed the role of other soil macro‐invertebrates. By sampling the dung beetle community and measuring the remaining dung and seeds after the experiment, the impact of each functional group was estimated. Results Dung beetle assemblages differed along a north–south and east–west gradient. Dwellers dominated northernmost sites, whereas at lower latitudes we observed more tunnellers and rollers indicating a functional shift. Resource specialists were more abundant in southern and eastern areas. Overall, functional group diversity enhanced dung removal. More dung (+46.9%) and seeds (+32.1%) were removed in the southern sites and tunnellers and rollers were more effective. At the northernmost sites, where tunnellers were scarce or absent, other soil macro‐invertebrates removed the majority of dung. Main conclusions The conservation of functionally complete dung beetle assemblages is crucial to maintain the ecosystem functions provided by dung beetles. Given the latitudinal variation in functional group diversity, it is reasonable to expect compositional changes due to climate change. These changes could lead to increased dung removal and a higher secondary seed dispersal rate in northern regions.

Question: Is it possible to restore dry calcareous inland sand ecosystems with their characteristic plant community structure within a 4-yr period by means of combined abiotic-biotic techniques (topsoil replacement, inoculation with raked/mown plant material from target areas)? Location: Upper Rhine valley, Germany. Methods: Two 4-year experiments were carried out on former arable land, each in the proximity of a reference area bearing a similar complex of threatened sandy grasslands (experiment 1: fine-scale; experiment 2: landscape scale). In both experiments we used nutrient-poor deep-sand substrate (abiotic approach), raked/mown inoculation material from target areas and grazing as management tool (biotic and management approach). The vegetation of the restoration and donor areas was sampled once a year and analysed by non-metric multidimensional scaling (NMDS) ordination and target-species ratios. Mixed linear models were calculated to determine effects of grazing (experiment 1) and year (both experiments). Results: NMDS revealed a continuous development of the restored sites towards the corresponding donor sites. Similarly, target-species ratios of the restored sites tended towards the ratios of the donor sites. To date, grazing effects have mainly been structural: reduction of a carpet-forming pleurocarpous moss species and of litter. In addition, cover of target species in relation to total plant cover was significantly enhanced by grazing in the last two study years. Conclusions: The combination of nutrient-poor substrate, inoculation with raked/mown plant material and grazing proved to be a very effective restoration method for dry base-rich sand ecosystems. After 4 yr the restored plant communities serve as well-developed parts of a habitat network. Nomenclature: Wisskirchen & Haeupler (1998) for vascular plant species, Koperski et al. (2000) for bryophytes and Scholz (2000) for lichens.Abbreviations: NMDS 5 non-metric multidimensional scaling; TSR qual 5 qualitative target-species ratio; TSR quant 5 quantitative target-species ratio.Eichberg, C.

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