1. Increasing the number and height of the new foredunes, seaward of the manmade dune, based on the initial nourishment, calcareous marine sand, as well as
subsequent accumulation of less calcareous marine sand.
2. Increasing the distance of the man-made dune, as well as that of the new dunes,
from the sea, causing a decline in the relative influence of the sea and
increasing stabilization.
3. Successive immigration of species from local and regional species pools.
4. Change in the dominating dispersal agent—the seawater—during the first years
of dune building and the wind during the later stages.
5. Change from a high proportion of coastal plant species to a high proportion of
inland plant species.
6. Replacement of beach annuals and dune-building rhizome geophytes by
hemicryptophytes and inland annuals.
7. Increasing habitat differentiation in time and space.
The increasing habitat differentiation of the initially man-made dune system
was indicated by the increasing number of TWINSPAN groups with increasing
dune age and distance from the coastline. This differentiation matches well with
dune succession found elsewhere in younger parts of natural dune systems. After
24 years of development there is no indication so far—neither from soil processes
like leaching and acidification (Vestergaard 2006) nor from the presence of species
typical of stabilized, acid dunes such as Corynephorus canescens—of continued
succession toward plant communities found in the inner, older parts of natural
Baltic and NW European dune systems (e.g., Olsson 1974; Hundt 1985). It can be
expected, however, that the landward-most part of the dune system will in due
course be invaded by Rosa rugosa scrub, as observed in other parts of the seaside
park. Such a development will probably reduce the species richness of the dunes
(Isermann 2008).
Acknowledgments Most of the field work was carried out in cooperation with the late Professor
Kjeld Hansen. I wish to thank Hans Henrik Bruun, Marisa Martinez, Patrick Hesp, and Juan B.
Gallego-Fernandez for constructive comments on the manuscript.
References
Andersen UV (1993) Dispersal strategies of Danish seashore plants. Ecography 16:289–298
Andersen UV (1995) The influence of human trampling on the vegetation on artificial dune and
coastal grassland in Denmark. In: Salman AHPM, Berends H, Bonazountas M (eds) Coastal
management and habitat conservation, vol 1. EUCC, Leiden, pp 427–438
Böcher TW (1954) Studies in European calcareous fixed dune communities. Plant Ecol 5–6:562–570
Box EO (1996) Plant functional types and climate at the global scale. J Veg Sci 9:309–320
Bradshaw AD (1993) Introduction: understanding the fundamentals of succession. In: Miles J,
Walton DWH (eds) Primary succession on land. Blackwell, Oxford, pp 1–3
Carter RWG (1988) Coastal environments. Academic Press, London
4 Natural Plant Diversity Development on a Man-Made Dune System
63
subsequent accumulation of less calcareous marine sand.
2. Increasing the distance of the man-made dune, as well as that of the new dunes,
from the sea, causing a decline in the relative influence of the sea and
increasing stabilization.
3. Successive immigration of species from local and regional species pools.
4. Change in the dominating dispersal agent—the seawater—during the first years
of dune building and the wind during the later stages.
5. Change from a high proportion of coastal plant species to a high proportion of
inland plant species.
6. Replacement of beach annuals and dune-building rhizome geophytes by
hemicryptophytes and inland annuals.
7. Increasing habitat differentiation in time and space.
The increasing habitat differentiation of the initially man-made dune system
was indicated by the increasing number of TWINSPAN groups with increasing
dune age and distance from the coastline. This differentiation matches well with
dune succession found elsewhere in younger parts of natural dune systems. After
24 years of development there is no indication so far—neither from soil processes
like leaching and acidification (Vestergaard 2006) nor from the presence of species
typical of stabilized, acid dunes such as Corynephorus canescens—of continued
succession toward plant communities found in the inner, older parts of natural
Baltic and NW European dune systems (e.g., Olsson 1974; Hundt 1985). It can be
expected, however, that the landward-most part of the dune system will in due
course be invaded by Rosa rugosa scrub, as observed in other parts of the seaside
park. Such a development will probably reduce the species richness of the dunes
(Isermann 2008).
Acknowledgments Most of the field work was carried out in cooperation with the late Professor
Kjeld Hansen. I wish to thank Hans Henrik Bruun, Marisa Martinez, Patrick Hesp, and Juan B.
Gallego-Fernandez for constructive comments on the manuscript.
References
Andersen UV (1993) Dispersal strategies of Danish seashore plants. Ecography 16:289–298
Andersen UV (1995) The influence of human trampling on the vegetation on artificial dune and
coastal grassland in Denmark. In: Salman AHPM, Berends H, Bonazountas M (eds) Coastal
management and habitat conservation, vol 1. EUCC, Leiden, pp 427–438
Böcher TW (1954) Studies in European calcareous fixed dune communities. Plant Ecol 5–6:562–570
Box EO (1996) Plant functional types and climate at the global scale. J Veg Sci 9:309–320
Bradshaw AD (1993) Introduction: understanding the fundamentals of succession. In: Miles J,
Walton DWH (eds) Primary succession on land. Blackwell, Oxford, pp 1–3
Carter RWG (1988) Coastal environments. Academic Press, London
4 Natural Plant Diversity Development on a Man-Made Dune System
63