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

Food-plant and foraging-site selection by semidomesticated female reindeer (Rangifer tarandus tarandus L.) was studied to shed light on the searching and foraging behaviour of this herbivore. The aims of the study were (i) to determine the role of food biomass and (or) plant nitrogen content in feeding-site selection and (ii) to analyse the extent to which movement patterns of reindeer are related to availability of food resources using several models of searching behaviour (fractal analysis, correlated random walks, and Lévy flights). The study was conducted in summer 1999 in a mountainous area of northern Sweden. Reindeer selected different plant communities during this period and changed search pattern in late summer. We found that reindeer selected feeding sites with higher green biomass of Betula spp. and Salix spp. However, there was no sharp threshold for foraging as suggested by some models. Contrary to qualitative predictions of optimal-foraging theory, we found no selection of feeding sites on the basis of the nitrogen content of food. The changed search pattern in late summer and the discrepancy between reindeer foraging paths and a correlated random walk model suggests that reindeer were responding to their environment by changing their searching behaviour.

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Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant-plant interactions and through soil disturbance. In forest ecosystems, researchers' attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation-wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light-demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non-forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape-level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.

Habitat fragmentation contributes to the decline of plant species by decreasing gene flow among populations. Restoring connectivity among habitat patches is therefore a major issue for plant conservation. However, deciding where to focus restoration efforts requires identifying suitable dispersers for each target plant species. We collected data from the literature on wild and domesticated ungulates, known to be effective seed dispersers, and on the plants they dispersed in Europe via epi‐ and/or endozoochory. We performed a systematic literature review to identify plant and animal traits relevant for seed dispersal. We first modeled the relationships between epi‐ or endozoochory and a priori selected plant traits (diaspore releasing height, length, shape and morphology, and habitat openness). The differences we underlined between the two dispersal mechanisms justified splitting our analyses accordingly. Then, for each dispersal mechanism, we asked whether basic plant traits could be used to predict specific traits of ungulates as endozoochorous or epizoochorous seed dispersers. We modeled the relationships between a priori selected ungulate traits for epizoochory (habitat openness, shoulder height, hair curliness, and hair length) and for endozoochory (habitat openness, body mass, feeding type and digestive system) and plant traits. Plant habitat openness and diaspore morphology were the predictors that most often explained differences among ungulates for epizoochory, whereas plant habitat openness and diaspore releasing height most often explained differences for endozoochory. Our trait‐based predictive models can help improve our ability to propose more precise management decisions for the conservation of plant populations worldwide by taking into account ungulate dispersers.

Semi‐natural grasslands are rich in biodiversity and thus important habitats for conservation, yet they are experiencing rapid declines due to agricultural intensification and abandonment. Promoting a more diverse mammalian herbivore community, including large and megaherbivores, may result in positive cascade effects for biodiversity and ecosystem functioning. Therefore, reintroducing an ecologically functional substitute of an extinct large herbivore could mitigate current biodiversity declines and foster semi‐natural grassland conservation. To test this hypothesis, we set up a 3‐year rewilding experiment where 12 feral horses were introduced in three 10‐hectare enclosure replicates (four horses per enclosure). We used community‐weighted mean plant functional traits to elucidate plant community changes induced by grazing through time. We also investigated the effects of this experimental treatment on insect pollinated plants and on pollinator habitat use. The grassland community exerted a mixed tolerance/avoidance response to grazing. This resulted in plant functional compositional changes which favoured prostrate plant species with higher specific leaf area, characteristic of ruderal communities. Plant species richness was higher in grazed compared to ungrazed areas. Butterfly and bumblebee habitat use, as well as feeding and resting activities were also higher in grazed areas. Moreover, the number of pollinators increased with plant species richness. Synthesis and applications. This study demonstrates that, to enhance the diversity of a given herbivore community with ecological replacements of extinct wild horses can have significant effects on the functional composition of grasslands. It can also mitigate plant species declines, in particular bee‐dependent plants, and boost pollinator habitat use. Novel management alternatives are urgently needed to reverse the negative effect of land abandonment in European agricultural landscapes. Thus, rewilding interventions with large mammalian herbivores may offset current biodiversity declines by maintaining important functional links between plants and pollinators in grassland ecosystems.

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SummaryMost range shift predictions focus on the dispersal phase of the colonization process. Because moving populations experience increasingly dissimilar nonclimatic environmental conditions as they track climate warming, it is also critical to test how individuals originating from contrasting thermal environments can establish in nonlocal sites.We assess the intraspecific variation in growth responses to nonlocal soils by planting a widespread grass of deciduous forests (Milium effusum) into an experimental common garden using combinations of seeds and soil sampled in 22 sites across its distributional range, and reflecting movement scenarios of up to 1600 km. Furthermore, to determine temperature and forest-structural effects, the plants and soils were experimentally warmed and shaded.We found significantly positive effects of the difference between the temperature of the sites of seed and soil collection on growth and seedling emergence rates. Migrant plants might thus encounter increasingly favourable soil conditions while tracking the isotherms towards currently 'colder' soils. These effects persisted under experimental warming. Rising temperatures and light availability generally enhanced plant performance.Our results suggest that abiotic and biotic soil characteristics can shape climate changedriven plant movements by affecting growth of nonlocal migrants, a mechanism which should be integrated into predictions of future range shifts.