In intensively used landscapes, remnant grassland fragments are often restricted to places unsuitable for agricultural cultivation. Such refuges are the ancient burial mounds called “kurgans,” which are typical landscape elements of the Eurasian steppe and forest steppe zone. Due to their hill‐like shape, loose soil structure and undisturbed status kurgans provide proper habitats for burrowing mammals. Accordingly, grassland vegetation on kurgans is often exposed to bioturbation, which can influence the habitat structure and plant species pool. In our study, we explored the effect of fox burrows and landscape context on the habitat properties and vegetation composition of small landscape elements, using kurgans as model habitats. We surveyed the vegetation of fox burrows and that of the surrounding grassland on five kurgans situated in cleared landscapes surrounded by arable lands and five kurgans in complex landscapes surrounded by grazed grasslands. We recorded the percentage cover of vascular plants, the amount of litter, and soil moisture content in twelve 0.5 m × 0.5 m plots per kurgan, in a total of 120 plots. We found that foxes considerably transformed habitat conditions and created microhabitats by changing the soil nutrient availability and reducing total vegetation cover and litter. Several grassland specialist species, mostly grasses (Agropyron cristatum, Elymus hispidus, and Stipa capillata) established in the newly created microhabitats, although the cover of noxious species was also considerable. We found that landscape context influenced the sort of species which could establish on kurgans by affecting the available species pool and soil moisture. Our results revealed that foxes act as ecosystem engineers on kurgans by transforming abiotic and biotic conditions by burrowing. Their engineering activity maintains disturbance‐dependent components of dry grasslands and increases local environmental heterogeneity.

Sowing of grass seed mixtures is a feasible and cost-effective method for landscape-scale grassland restoration. However, sowing only grasses usually leads to species-poor and dense grass sward, where the establishment of target forbs is hampered both by microsite and propagule limitation. To overcome these limitations and increase the diversity of species-poor sown grasslands we developed a novel method by creating 'establishment gaps'. We used tillage to open gaps of 1 m 2 , 4 m 2 and 16 m 2 size in the dense grass sward of six species-poor sown grasslands in the Great Hungarian Plain. We sowed high-diversity seed mixtures of 35 native species into all gaps. We analyzed vegetation development during the first five years after setting up the trial. We also studied the colonization dynamics of the sown species along four 20-m transects around each gap, resulting in a total of 1440 plots of 1 m 2 size that were studied. Our results indicated that most of the sown species were able to establish permanently in the establishment gaps. The total cover and the cover of perennial sown species increased independently of gap size. Meanwhile the cover of short-lived sown species 2 decreased during the five years. There was only a moderate level of weed abundance in the gaps, and weed cover decreased over the years. The sown target species started to colonize the species-poor grasslands surrounding the establishment gaps within five years. The highest number of species and individuals dispersed from the 4 m 2 -sized gaps, as they had a more stable development than small gaps and were exposed to lower grazing pressure than large ones. Implications for practice Establishment gaps are widely applicable tools to increase the diversity of speciespoor grasslands. Gaps of 4 m 2 represent a more feasible solution compared to larger openings also for the farmers, because there is only a moderate level of weed encroachment and smaller soil disturbance occurs during their creation.  We recommend sowing high-diversity seed mixtures containing both short-lived species that can establish in the first year and perennial species, which guarantee a high cover of target species later on.  Gaps sown with high-diversity seed mixture are highly resistant to unfavorable climatic conditions: increasing grass abundance in dry years does not hamper the recovery of target grassland species in the following years.

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Questions Plant invasions are considered among the biggest threats to biodiversity worldwide. In a full‐factorial greenhouse experiment we analysed the effect of soil burial depth and litter cover on the germination of invasive plants. We hypothesised that: (a) burial depth and litter cover affect the germination of the studied species; (b) the effects of burial and litter cover interact with each other, and (c) the effects are species‐specific, but dependent on seed size. Methods We tested the germination and seedling growth of 11 herbaceous invasive species in a full‐factorial experiment using four levels of seed burial depths and litter cover. We analysed the effect of burial, litter cover, and their interactions on germination, seedling length and biomass across species and at the species level. Results Soil burial depth and litter cover had a significant effect on the germination of the studied species, but there were considerable differences between species. We observed a general trend of species with bigger seeds being not or less seriously affected by soil burial and litter cover than smaller‐seeded species. Correlations between seed weight and effect sizes mostly confirmed this general trend, but not in the case of soil burial. Conclusions Our findings confirmed that seed size is a major driver of species’ response to litter cover and to the combined effects of litter cover and soil burial, but there is no general trend regarding the response to soil burial depth. Despite its very small seeds, the germination of Cynodon dactylon was not affected by soil burial. The germination of Ambrosia artemisiifolia was hampered by both soil burial and litter cover despite its relatively large seeds. Thus, specific information on species’ response to burial depth and litter accumulation is crucial when planning management or restoration in areas threatened by plant invasions.