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Non-native Berberis aquifolium is an invasive species in Belgian coastal dunes. With its strong clonal growth through suckers, this evergreen shrub outcompetes native species and affects dune succession. To prevent further secondary spread and mitigate its impact, there was an urgent need for knowledge on the effectiveness of control measures, both at the plant and habitat level. Here, we report on a first control experiment. Individual B. aquifolium clones were subjected to one of four treatments (manual uprooting, foliar herbicide application, stem cutting followed by herbicide or salt application), with regrowth being measured up to one year after treatment. We analyzed the relationship between kill rate, treatment, dune area, plant volume and number of plant stems using a generalized linear model. Berberis aquifolium plants proved most susceptible to foliar herbicide application (5% glyphosate solution), resulting in 88% (64%–97%) of the clones dying after treatment. The predicted kill rate decreased with an increasing number of stems under all treatments. We discuss the limitations of our experiment and the potential for actual field application of the different treatments. We present some guidelines for future control that may become further refined as experience builds up and we provide some recommendations for tackling invasive alien species in Atlantic dune ecosystems.

Coastal dunes in Northwestern Europe were subject to substantial scrub expansion during the past decades, a process often considered as a major problem for conservation management. This paper explores this process in De Westhoek nature reserve (Belgium), focussing on the three major constituents of the early scrub: Salix repens, Hippophae rhamnoides and Ligustrum vulgare. We study changes in dominance pattern between 1953 and 2010 based on vegetation maps and the demography of these species based on growth ring counting. We relate this data to the edaphic factors soil moisture and soil age. Early scrub development occurred in three phases: (1) rapid, mainly secondary scrub expansion after release of grazing pressure, (2) a main shift in relative abundance of the three main species, and (3) further succession into tall scrub and woodland along with local scrub decay. Edaphic factors play a major role in vegetation dynamics, but their relative importance diminishes along with landscape ageing. Simultaneously, the outcome of competitive interaction between scrub species, in which population demography plays a key role, gains importance. Primary and secondary scrub development seem

In coastal dunes, landscape changes are a rule, rather than an exception. This paper gives an overview of changes in landscape and vegetation with a focus on the past century. The history of dunes is characterised by phases of sand drift, alternated with geomorphological stability. The historical development of dune woodland during these stable phases has been documented for sites all over Europe. Vegetation reconstructions of historical open dune habitats however is very difficult due to limited preservation of fossil remains. People have drastically altered coastal dune landscapes through centuries of exploitation and more recently development of the coast. Historical land use has generally pushed vegetation back into a semi-natural state. During roughly the past century a tendency of increasing fixation and succession is observed on coastal dunes throughout northwest Europe. Six causes of change are discussed. 1) Changes in land use, mainly abandonment of agricultural practices, have led to the development of late successional stages such as scrub and woodland. 2) Crashing rabbit populations due to myxomatosis in the 1950s caused vigorous grass growth and probably stimulated scrub development. 3) A general tendency of landscape fixation is observed due to both natural and anthropogenic factors. 4) Eutrophication, mainly due to atmospheric nitrogen deposition is clearly linked to grass encroachment on acidic but also on some calcareous dunes. 5) The impact of climate change on vegetation is still unclear but probably lengthening of growing season and maybe enhanced CO₂ concentrations have led to an acceleration of succession. 6) A general anthropogenisation of the landscape occurs with rapid spread of non-native species as an important consequence. The reconstruction of a natural reference landscape is considered largely unattainable because of irreversible changes and the long tradition of human impact, in many cases since the development of the dunes. Two contradictory elements need reconciliation. First, the general acceleration of succession and scrub and woodland development in particular is partly caused by a decreased anthropogenic interference in the landscape and deserves more appreciation. Second, most biodiversity values are largely linked to open, early succession dune habitats and are threatened by the same tendency. Apart from internal nature management, in which grazing plays an important part, re-mobilisation of stable, senescent dunes is an important challenge for dune management.

Extensive grazing often has a strong influence on the structure and composition of herbaceous plant communities with increasing population sizes for some species and decreasing presence in others. Herbivores affect plant communities directly by selective grazing of plant species, and indirectly by either epizoochory or endozoochory. Helianthemum nummularium is considered an increasing species because its distribution increased after the introduction of large, free-ranging grazers in at least two coastal dune grassland areas in Belgium. However, its seeds lack any obvious adaptations for epizoochory, and direct observations of plant/seed consumption are scarce. Through field and lab experiments, we assessed the dispersal ability of H. nummularium via endozoochory and epizoochory. In a differentiated grazer exclusion experiment, evidence was found that plants are grazed by large domestic ungulates and small wild herbivores although these incidences were rare. Direct endozoochory evidence remained scarce. No seeds were found germinating in field-collected dung, and only few seedlings emerged following a seed feeding experiment. However, once deposited, we found higher growth rates when seeds were mixed with dung and decreased establishment success when seeds were sown in combination with competitively superior species. Epizoochory was plausible because both fur and hooves of cattle and horses were potentially capable of contributing to the transport of H. nummularium seeds. We conclude that herbivores play a role in seed dispersal, while their selective grazing behaviour most probably creates an appropriate environment for Helianthemum establishment and maintenance.

Non-native Berberisaquifolium is an invasive species in Belgian coastal dunes. With its strong clonal growth through suckers, this evergreen shrub outcompetes native species and affects dune succession. To prevent further secondary spread and mitigate its impact, there was an urgent need for knowledge on the effectiveness of control measures, both at the plant and habitat level. Here, we report on a first control experiment. Individual B.aquifolium clones were subjected to one of four treatments (manual uprooting, foliar herbicide application, stem cutting followed by herbicide or salt application), with regrowth being measured up to one year after treatment. We analyzed the relationship between kill rate, treatment, dune area, plant volume and number of plant stems using a generalized linear model. Berberisaquifolium plants proved most susceptible to foliar herbicide application (5% glyphosate solution), resulting in 88% (64%–97%) of the clones dying after treatment. The predicted kill rate decreased with an increasing number of stems under all treatments. We discuss the limitations of our experiment and the potential for actual field application of the different treatments. We present some guidelines for future control that may become further refined as experience builds up and we provide some recommendations for tackling invasive alien species in Atlantic dune ecosystems.

Grey dunes are a priority habitat type of the European Union Habitats Directive and demand special attention for conservation and management. Knowledge of the ecology of coastal grey dunes can contribute to this policy. Dune grassland succession is initiated by fixation and driven by the complex of soil formation (humus accumulation) and vegetation development. Leaching and mobilization of CaCO3, which are important in nutrient dynamics, complicate the picture. At present, grass- and scrub encroachment greatly overrules these fine scaled soil processes and causes substantial loss of regional biodiversity. Belgium has an international responsibility in grey dune conservation because of the limited range of its characteristic vegetation, flora and fauna. As biomass removal seems essential in grassland preservation, grazing is an important management tool. Evaluation of management measures focuses on biodiversity measurements on the levels of landscape, community and species.

Nature-based solutions to mitigate the impact of future climate change depend on restoring biological diversity and natural processes. Coastal foredunes represent the most important natural flood barriers along coastlines worldwide, but their area has been squeezed dramatically because of a continuing urbanisation of coastlines, especially in Europe. Dune development is steered by the development of vegetation in interaction with sand fluxes from the beach. Marram grass (Calamagrostis arenaria, formerly Ammophila arenaria) is the main dune building species along most European coasts, but also in other continents where the species was introduced. Engineering of coastal dunes, for instance by building dunes in front of dikes, needs to be based on a solid understanding of the species' interactions with the environment. Only quantitative approaches enable the further development of mechanistic models and coastal management strategies that encapsulate these biomorphogenic interactions. We here provide a quantitative review of the main biotic and physical interactions that affect marram grass performance, their interactions with sand fluxes and how they eventually shape dune development. Our review highlights that the species spatial organisation is central to dune development. We further demonstrate this importance by means of remote sensing and a mechanistic model and provide an outlook for further research on the use of coastal dunes as a nature-based solution for coastal protection. Competing Interest Statement The authors have declared no competing interest. Footnotes * https://github.com/dbonte/EndureModel.git