Formerly coppiced old growth stands act as refugia of threatened biodiversity in a managed steppic oak forest
Introduction
Vegetation is a dynamic system, and change in vegetation is a universal phenomenon (Miles, 2012). Besides external conditions, the inevitability of death of plant individuals and the capability of plants to reproduce and disperse are constant intrinsic drivers of this change. However, the rate of change may substantially differ between sites and habitats. Persistence is as universal as change itself, and different mechanisms have been proposed for what is sometimes called biological inertia of plant communities (Gorham, 1957, Von Holle et al., 2003). Already Watt (1947) pointed out that species longevity is an important factor in this respect and that we can expect a lower degree of correspondence between site conditions and community structure in species assemblages with long-lived dominants; forests are an obvious case. As a result, present vegetation is a mixture of plant communities which differ not only due to varied site conditions, but also due to unique histories (Roleček et al., 2015).
Historical and modern forest management forms are usually accompanied by specific disturbance regimes and associated dynamics of plant and animal populations (Zavala and Burkey, 1997, Kopecký et al., 2013, Kirby et al., 2017). In temperate forests of Western and Central Europe, a major change in use has taken place since the 18th century, when historical forms such as coppicing, grazing and litter raking started to be abandoned (von Carlowitz, 1713, Evans, 1992, Müllerová et al., 2015). These forests have often been replaced by plantations of fast-growing trees managed by means of clear-cutting. Even if tree species composition was preserved, planting of young individuals of seed origin has become a favoured form of tree regeneration to the detriment of coppicing, which has been stigmatised as inferior forest management (e.g. Kostler, 1953, Normand, 1960). This shift towards large-scale plantations of trees, in temperate Europe frequently oaks (mainly Quercus robur and Q. petraea), may be surprizing as it in many cases involved expensive and labour-intensive measures such as stump removal, soil tillage and repeated manual weeding of the clear-cuts (Fujimori, 2001; Řepka, 2009). The destructive effects on plant and animal communities were hardly taken into consideration.
In the past decades, biodiversity decline has become a major concern, and restoration or emulation of historical forest management forms has been proposed as a nature-friendly alternative for forest management (Müllerová et al., 2015, Kirby et al., 2017). It is a particularly tempting option for abandoned coppices which have not been transformed to plantations and have preserved some ancient features, such as their characteristic biota (Vild et al., 2013, Roleček et al., 2017). Unfortunately, this option has not been yet implemented into management plans of many protected lowland forests in the Czech Republic, and the efforts to restore historical forest management meets considerable resistance not only from forestry institutions, but sometimes also from nature conservation authorities (Řepka, 2009, Miklín and Čížek, 2014). We suppose that besides economic reasons this resistance also stems from a lack of experience with alternative management forms, absence of consistent policy in this area and inability to communicate effectively with the stakeholders (Dušek, 2014, Miklín and Čížek, 2014).
The aim of the present study is to show the consequences for plant biodiversity of the transformation of an old coppice, historically linked to ancient wood pasture, to oak plantation. The study was performed in Dúbrava forest – the most valuable complex of lowland steppic oak forests in the Czech Republic and a Site of Community Importance. Besides simple effects on species richness, we focused on differences in ecological spectra of the species assemblages, which may indicate major environmental drivers of the observed dynamics.
Section snippets
Study site
Dúbrava forest (Fig. 1) is a large complex (about 34 km2) of steppic oak forests on sand located between the municipalities of Hodonín, Mutěnice, Dubňany, Ratíškovice and Rohatec (south-eastern Czech Republic; 48°53′N, 17°06′E). It is situated on a low plateau above the Morava river valley at an altitude of 160–205 m a.s.l., in a wine-growing region with a relatively warm and moderately dry climate (mean July temperature 19.8 °C, mean January temperature −1.5 °C, mean annual rainfall 585 mm;
Results
The 135 analysed oak stands significantly differed in species richness and other biodiversity-related characteristics depending on their age and management history. Species richness (Fig. 2) was the highest in clear-cuts, the lowest in young growth stands after area-wide site preparation and intermediate in old growth stands. When standardised to equal area, oak stands showing signs of past coppicing were significantly richer than other old growth stands. Species composition, ecological spectra
General pattern of species richness and composition
The objective of the present study was to show the consequences for plant diversity of the transformation of old coppices in a steppic oak forest to intensive oak plantations. We paid particular attention to changes in species composition and shifts in species’ ecological spectra indicating potential environmental drivers of the observed dynamics. Our results show that clear-cutting and establishment of oak plantations has had profound effects on plant species richness and composition: the
CRediT authorship contribution statement
Jan Roleček: Methodology, Data curation, Formal analysis, Visualization, Writing - original draft, Writing - review & editing. Radomír Řepka: Conceptualization, Methodology, Investigation, Data curation, Writing - review & editing.
Declaration of Competing Interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
Acknowledgements
The first author was supported by long-term research development project RVO 67985939. We are grateful to F. Koliba and P. Szabó for the information on the management history of oak stands in the Dúbrava forest and J. W. Jongepier for correcting our English. O. Hájek, J. Sladký and J. Sladký jun. are acknowledged for drawing the map.
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