An ecological comparison between ancient and other forest plant species of Europe, and the implications for forest conservation

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Abstract

An analysis is presented of the ecological characteristics of ancient forest plant species in deciduous forests of Europe. Twenty-two literature sources were used to generate a list of 132 ancient forest plant species, described from at least eight countries in Europe. The affinity for ancient forests of these species differs considerably from country to country, but they have a definite ecological profile. There is a significant difference in the response of the ancient forest plant species compared with other forest plant species for a variety of ecological characteristics, based on Ellenberg indicators, plant strategies and phytosociological associations. Ancient forest plant species tend to be more shade-tolerant than the other forest plant species; dry and wet sites are avoided. They are typical of forest sites with an intermediate pH and nitrogen availability. Geophytes and hemicryptophytes are more frequent amongst ancient forest plant species. The stress-tolerant plant strategy type is significantly more abundant under the ancient forest species than expected when compared with other forest plant species and vice versa for the competitive plant strategy. This distinct ecological profile suggests that ancient forest plant species may be considered as a guild. The poor ability of these species to colonize new forest sites may be attributed to a complex of interacting variables: limited dispersal abilities (many have a short-distance dispersal strategy), low diaspore production and recruitment problems (e.g. low competitive ability). The regional variation in ancient forest plant species suggests that regional lists are more appropriate for assessing the nature conservation value of forests than one global European list. Due to their distinct ecological profile and low colonizing abilities, ancient forest plant species may be considered as important biodiversity indicators for forests.

Introduction

In a fragmented, dynamic landscape, a species must be able to successfully colonize new habitats at least as fast as it is lost from existing habitats. Those species which are unable to do so, are threatened by extinction from the landscape. Such is considered to be the case for ancient forest plant species. While some plants can colonize new forests quickly, (e.g. Geum urbanum and Urtica dioica), other forest species may take centuries (Peterken, 1981, Peterken and Game, 1984). Such slow-colonizing species are termed ancient forest species, because their presence suggests a long continuous history for the habitat patch, and also because they therefoe may be indicative of more original forest conditions (Peterken, 1974, Rackham, 1980, Whitney and Foster, 1988). They are considered important in terms of nature conservation, because lists of ancient forest species combine both qualitative (forest quality) and quantitative (diversity) conservation criteria (Peterken, 1974, Peterken, 1977, Peterken, 1996, Honnay et al., 1999a). However, what are the ecological characteristics of this group of plants, other than a tendency to be associated with old forests?

Previously there has been no attempt at an ecological comparison between ancient forest plants species and other species at the European scale. Such an overview could inform the debate on forest biodiversity indicators and also help identify the conditions needed for the long-term persistence of these plants within European landscapes. The objective of this paper is to create a species list of ancient forest plants from the literature, and secondly to compare ecological attributes for species within this list with those for other plant species with forests. We then discuss the causes of failure of ancient forest plant species to colonize new forests and the consequences for nature conservation.

Section snippets

Identification of European ancient forest plants

A list of ancient forest plant species was developed from 22 publications, all from deciduous forests of northwestern and central Europe (Belgium, the Czech Republic, Denmark, Germany, Great Britain, the Netherlands, Poland, and Sweden, Table 1). The quality and nature of these publications vary considerably, from detailed, large scale studies (Hermy, 1985, Peterken and Game, 1984, Rackham, 1980, Wulf, 1994) to scattered and more anecdotical data on forest species (e.g. Roisin and Thill, 1952,

Lists of ancient forest species and total forest flora

A review of the literature produced a list of 132 forest plant species considered to have a clear affinity for ancient forests (Table 2), out of a list total of 627 forest species including the two species with non-specific behaviour. Note that five ancient forest species indicative of grasslands and heathlands according to Ellenberg et al. (1991) were not omitted (Table 2), because they are associated with forests in western Europe.

The total list of all forest species can be divided into five

The list of ancient forest plant species and regional variation

While our list of ancient plant species is informative, it must be interpreted with caution for several reasons. The absence of an ancient forest species from a certain list may mean that the species was simply unrecorded (particularly with anecdotal sources, see methods section), that it was not present in the studied forests, or that it was present but was not considered to be an ancient forest species. No exact conclusion can be reached from the available literature without further data.

Acknowledgements

This work is part of the Landeconet project, an EC funded research project on landscape ecology in changing agricultural landscapes (contract EV5VCT940528). Thanks to Kris Verheyen, an anonymous referee and Dr. Jean-Luc Dupouey for comments on the manuscript.

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