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Fast response of Scots pine to improved water availability reflected in tree-ring width and δ13C
(2010)
Drought-induced forest decline, like the Scots pine mortality in inner-Alpine valleys, will gain in importance as the frequency and severity of drought events are expected to increase. To understand how chronic drought affects tree growth and tree-ring δ13C values, we studied mature Scots pine in an irrigation experiment in an inner-Alpine valley. Tree growth and isotope analyses were carried out at the annual and seasonal scale. At the seasonal scale, maximum δ13C values were measured after the hottest and driest period of the year, and were associated with decreasing growth rates. Inter-annual δ13C values in early- and latewood showed a strong correlation with annual climatic conditions and an immediate decrease as a response to irrigation. This indicates a tight coupling between wood formation and the freshly produced assimilates for trees exposed to chronic drought. This rapid appearance of the isotopic signal is a strong indication for an immediate and direct transfer of newly synthesized assimilates for biomass production. The fast appearance and the distinct isotopic signal suggest a low availability of old stored carbohydrates. If this was a sign for C-storage depletion, an increasing mortality could be expected when stressors increase the need for carbohydrate for defence, repair or regeneration.
In Valais, Switzerland, Scots pines (Pinus sylvestris L.) are declining, mainly following drought. To assess the impact of drought on tree growth and survival, an irrigation experiment was initiated in 2003 in a mature pine forest, approximately doubling the annual precipitation. Tree crown transparency (lack of foliage) and leaf area index (LAI) were annually assessed. Seven irrigated and six control trees were felled in 2006, and needles, stem discs and branches were taken for growth analysis. Irrigation in 2004 and 2005, both with below-average precipitation, increased needle size, area and mass, stem growth and, with a 1-year delay, shoot length. This led to a relative decrease in tree crown transparency (−14%) and to an increase in stand LAI (+20%). Irrigation increased needle length by 70%, shoot length by 100% and ring width by 120%, regardless of crown transparency. Crown transparency correlated positively with mean needle size, shoot length and ring width and negatively with specific leaf area. Trees with high crown transparency (low growth, short needles) experienced similar increases in needle mass and growth with irrigation than trees with low transparency (high growth, long needles), indicating that seemingly declining trees were able to ‘recover’ when water supply became sufficient. A simple drought index before and during the irrigation explained most of the variation found in the parameters for both irrigated and control trees.
An increasing number of studies have reported on forest declines and vegetation shifts triggered by drought. In the Swiss Rhone valley (Valais), one of the driest inner-Alpine regions, the species composition in low elevation forests is changing: The sub-boreal Scots pine (Pinus sylvestris L.) dominating the dry forests is showing high mortality rates. Concurrently the sub-Mediterranean pubescent oak (Quercus pubescens Willd.) has locally increased in abundance. However, it remains unclear whether this local change in species composition is part of a larger-scale vegetation shift. To study variability in mortality and regeneration in these dry forests we analysed data from the Swiss national forest inventory (NFI) on a regular grid between 1983 and 2003, and combined it with annual mortality data from a monitoring site. Pine mortality was found to be highest at low elevation (below 1000 m a.s.l.). Annual variation in pine mortality was correlated with a drought index computed for the summer months prior to observed tree death. A generalized linear mixed-effects model indicated for the NFI data increased pine mortality on dryer sites with high stand competition, particularly for small-diameter trees. Pine regeneration was low in comparison to its occurrence in the overstorey, whereas oak regeneration was comparably abundant. Although both species regenerated well at dry sites, pine regeneration was favoured at cooler sites at higher altitude and oak regeneration was more frequent at warmer sites, indicating a higher adaptation potential of oaks under future warming. Our results thus suggest that an extended shift in species composition is actually occurring in the pine forests in the Valais. The main driving factors are found to be climatic variability, particularly drought, and variability in stand structure and topography. Thus, pine forests at low elevations are developing into oak forests with unknown consequences for these ecosystems and their goods and services.