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Establishment of calcareous grassland on ex-arable fields by introducing target species is one of the most frequently used methods to restore the species assemblages of this highly endangered habitat type. The present study evaluates the long-term success of calcareous grassland restoration on former arable land in the vicinity of one of the oldest nature reserves in Bavaria, the “Garchinger Heide”. The restoration experiment combined different measures like topsoil removal, transfer of freshly cut seed-containing hay and additional sowing to the following variants in a 21-year experiment: (1) No topsoil removal, no hay transfer (control), (2) no topsoil removal with immediate hay transfer, (3) topsoil removal with immediate hay transfer and (4) topsoil removal with hay transfer 10 years after the start of restoration. Eleven Red List species which had not been transferred successfully were additionally sown after 9 to 19 years. Due to a limited availability of seeds, sowing of these species was mainly restricted to areas with topsoil removal, where better establishment was expected due to low vegetation cover. Five rare species with abundant seed production were also sown to plots without topsoil removal and hay transfer. The nature reserve served both as the donor area of the target species and as the reference to evaluate restoration success. Regarding aboveground biomass and total vegetation cover, greatest similarity to the donor site was observed on plots without topsoil removal. In contrast, the highest numbers of target species occurred on plots with topsoil removal, hay transfer and additional sowing. Similarity in species composition between restoration sites and the reference area increased over time, but species composition of restored sites did not fully reflect the reference after 21 years. One reason for the remaining dissimilarity was probably that topsoil removal favored stress tolerant species which were less common on the mature and more fine-grained soils of the nature reserve. Plots without topsoil removal still differed from the reference by their high vegetation cover and a significantly higher proportion of mesophytic grassland species. The study also showed that 19 Red List species were successfully established on the former arable fields, eight of them presumably by sowing. Nevertheless, various other rare species have not been observed yet. Results on functional traits characterizing environmental adaptation and reproduction also underlined the differences between restoration plots and the reference site. Our study presents a ʽdynamic restoration approachʼ where managers evaluated the original factorial treatments after a decade and modified them by additional treatments where development was sub-optimal. Such additional treatments may have confounded the experimental design, but from a management perspective proved to be a promising option to establish species rich grassland of high conservation value with a reasonable expenditure of time.
During recent decades, many studies have shown that the successful restoration of species-rich grasslands is often seed-limited because of depleted seed banks and limited seed dispersal in modern fragmented landscapes. In Europe, commercial seed mixtures, which are widely used for restoration measures, mostly consist of species and varieties of non-local provenance. The regional biodiversity of a given landscape, however, can be preserved only when seeds or plants of local provenance are used in restoration projects. Furthermore, the transfer of suitable target species of local provenance can strongly enhance restoration success.
We review and evaluate the success of currently used near-natural methods for the introduction of target plant species (e.g. seeding of site-specific seed mixtures, transfer of fresh seed-containing hay, vacuum harvesting, transfer of turves or seed-containing soil) on restoration sites, ranging from dry and mesic meadows to floodplain grasslands and fens. Own data combined with literature findings show species establishment rates during the initial phase as well as the persistence of target species during long-term vegetation development on restoration sites.
In conclusion, our review indicates that seed limitation can be overcome successfully by most of the reviewed measures for species introduction. The establishment of species-rich grasslands is most successful when seeds, seed-containing plant material or soil are spread on bare soil of ex-arable fields after tilling or topsoil removal, or on raw soils, e.g. in mined areas. In species-poor grasslands without soil disturbance and on older ex-arable fields with dense weed vegetation, final transfer rates were the lowest. For future restoration projects, suitable measures have to be chosen carefully from case to case as they differ considerably in costs and logistic effort. Long-term prospects for restored grassland are especially good when management can be incorporated in agricultural systems.