Spatial and temporal patterns in the climate-growth relationships of Fagus sylvatica across Western Europe, and the effects on competition in mixed species forest

Increases in temperature, altered precipitation patterns, and the occurrence and severity of extreme climatic events have been important characteristics of the climate change observed to date. This has had many and diverse impacts upon the living world, with one recent observation being a global reduction in the net primary production of all terrestrial vegetation. Increases in temperature and the frequency of extreme events are predicted to continue throughout the 21st century, and can be expected to have far reaching effects on global terrestrial ecosystems. Increases in temperature and drought occurrence could fundamentally impact upon the growth rates, species composition and biogeography of forests in many regions of the world, with many studies indicating that this process is already underway. European beech, Fagus sylvatica, is one of Europe’s most widespread and significant broadleaved tree species, forming an important and frequently dominant component of around 17 million hectares of forest. However, the species is also considered to be drought sensitive. Thus, much research interest has focused on eliciting the details of its physiological response to increased water stress, whilst dendroecological studies have attempted to identify sites and regions where reductions in growth might be found. A significant knowledge gap exists regarding a multi-regional, range-wide view of growth trends, growth variability, climate sensitivity, and drought response for the species. Predicting the potential effects of climate change on competition and species composition in mixed species forests remains an important challenge. In order to address this knowledge gap, a multi-regional tree-ring network was constructed comprising of 46 sites in a latitudinal transect across the species’ Western European range. This consisted of 2719 tree cores taken from 1398 individual trees, which were used to construct tree-ring chronologies for each site in the network. As a first step in a multi-regional assessment for F. sylvatica, a combination of the tree-ring chronologies and environmental data derived from a large scale gridded climate dataset were used in a multivariate analysis. Sites in the latitudinal transect were partitioned into geographically meaningful regions for further analysis. The resulting regions were then studied using climate-growth analysis, pointer year analysis of drought years, analysis of growth trends and growth variability, in order to examine regional variation in the response of the species to climate. Furthermore, a combination of long-term monitoring data from one specific site was combined with tree-ring sampling of multiple cohorts of F. sylvatica and one co-dominant competitor, Quercus petraea, to study the effects of an extreme drought event in 1976 on mortality and subsequent recovery. Key results of the multi-regional analysis are that large scale growth reductions are not evident in even the most southerly and driest portions of the species’ range. Radial growth is increasing, both in the north and in the core of the species’ range, with southern range edge forests maintaining stable growth. However, the variability of growth from year to year is increasing for all regions, indicative of growing stress. Crucially, the southern range edge, which previous studies had identified as an ‘at risk’ region, was shown to be more robust than expected. Climate sensitivity and drought impacts were low for this region. Instead, forests in the core of the species range, both in continental Europe and in the south of the UK, were identified as having the highest climate sensitivity, highest drought impacts, and experiencing periodic reductions in growth as a result. Northern range edge forests showed little sign of being affected by drought, instead having low climate sensitivity and strongly increasing growth trends. Extreme drought was found to affect species differently: the dominant species (F. sylvatica) failed to recover pre-drought levels of growth, whilst a transient effect of competitive release occurred for the co-dominant species (Q. petraea). There was also a long term effect on the relative abundance of the two species within the woodland, due to differences in the levels of drought induced mortality experienced by the species. This shows that in the case of extreme climatic events where thresholds in the ability of species to tolerate water stress are breached, the effects of drought can be rapid and long lasting. Drought impacts can cascade beyond that experienced by the most drought sensitive species, due to changes in competitive interactions between species in mixed species forests. The implications of this work suggest opportunities, risks and strengths for F. sylvatica. In the northern portion of the species’ range, predicted increases in productivity are confirmed by recent growth trends, indicating a good outlook for the species. At the southern range edge, F. sylvatica forests exist either in locations where precipitation is high or locations where local environmental conditions buffer them from an inhospitable regional climate. These factors result in southern range edge forests which are highly resilient to the effects of increasing climate stress. It is instead in the core of the species’ range where the most sensitive forests are found. The effects of extreme drought on a range core forest demonstrated here provide a cautionary note: where drought stress tolerance thresholds are breached, rapid and long lasting effects on growth and mortality can occur, even in regions where drought has not previously been considered to pose a strong risk to the species.