Incidences of oak decline have occurred repeatedly during the past three centuries as well as in the most recent decades. On the basis of historical records and dendrochronological measurements, oak decline in Central Europe has been attributed to the single or combined effects of climatic extremes (winter frost, summer drought), defoliating insects, and pathogenic fungi. Starting from a literature review, we discuss the possible roles of various abiotic (air pollution, nitrogen eutrophication, soil chemical stress, climatic extremes, site conditions) and biotic factors (insect defoliation, borer attack, infection by pathogenic fungi, microorganisms) that have been related to oak decline. On the basis of investigations on Quercus petraea and Quercus robur at three different levels (from experiments with young trees to monitoring on a supraregional scale), we suggest a conceptual model of the interaction of abiotic and biotic factors responsible for the onset of oak decline. This model should be valid for Central European oak stands at more acidic sites (soil pH (H 2 O) £ 4.2; on soils with higher pH, pathogenic Phytophthora species may contribute to oak decline). The combination of severe insect defoliation in at least two consecutive years with climatic extremes is the most significant complex of factors in the incidence of oak decline. Combined with defoliation, summer drought or winter ⁄ spring frost or both have to occur within the same year or in consecutive years to trigger major outbreaks of decline. Important additional stress factors are the following: (1) hydromorphic site conditions which, particularly in the case of Q. robur, render the trees more susceptible to drought stress as a result of an impairment of root growth in the subsoil; and (2), possibly, excess nitrogen which, in combination with drought stress, results in distinct decreases in the foliar concentrations of allelochemicals in Q. robur, thereby probably making the trees more susceptible to insect defoliation. Air pollution, soil chemical stress (including excess manganese), and nitrogen-induced nutritional imbalance do not seem to be important causal factors in the complex of oak decline. On the basis of the model, the appearance of the most recent oak decline in North-western Germany can be adequately explained.
The production and composition of leaf litter, soil acidity, exchangeable nutrients, and the amount and distribution of soil organic matter were analyzed in a broad‐leaved mixed forest on loess over limestone in Central Germany. The study aimed at determining the current variability of surface‐soil acidification and nutrient status, and at identifying and evaluating the main factors that contributed to the variability of these soil properties along a gradient of decreasing predominance of European beech (Fagus sylvatica L.) and increasing tree‐species diversity. Analyses were carried out in (1) mature monospecific stands with a predominance of beech (DL 1), (2) mature stands dominated by three deciduous‐tree species (DL 2: beech, ash [Fraxinus excelsior L.], lime [Tilia cordata Mill. and/or T. platyphyllos Scop.]), and (3) mature stands dominated by five deciduous‐tree species (DL 3: beech, ash, lime, hornbeam [Carpinus betulus L.], maple [Acer pseudoplatanus L. and/or A. platanoides L.]). The production of leaf litter was similar in all stands (3.2 to 3.9 Mg dry matter ha–1 y–1) but the total quantity of Ca and Mg deposited on the soil surface by leaf litter increased with increasing tree‐species diversity and decreasing abundance of beech (47 to 88 kg Ca ha–1 y–1; 3.8 to 7.9 kg Mg ha–1 y–1). The soil pH(H2O) and base saturation (BS) measured at three soil depths down to 30 cm (0–10 cm, 10–20 cm, 20–30 cm) were lower in stands dominated by beech (pH = 4.2 to 4.4, BS = 15% to 20%) than in mixed stands (pH = 5.1 to 6.5, BS = 80% to 100%). The quantities of exchangeable Al and Mn increased with decreasing pH and were highest beneath beech. Total stocks of exchangeable Ca (0–30 cm) were 12 to 15 times larger in mixed stands (6660 to 9650 kg ha–1) than in beech stands (620 kg ha–1). Similar results were found for stocks of exchangeable Mg that were 4 to 13 times larger in mixed stands (270 to 864 kg ha–1) than in beech stands (66 kg ha–1). Subsoil clay content and differences in litter composition were identified as important factors that contributed to the observed variability of soil acidification and stocks of exchangeable Ca and Mg. Organic‐C accumulation in the humus layer was highest in beech stands (0.81 kg m–2) and lowest in stands with the highest level of tree‐species diversity and the lowest abundance of beech (0.27 kg m–2). The results suggest that redistribution of nutrients via leaf litter has a high potential to increase BS in these loess‐derived surface soils that are underlain by limestone. Species‐related differences of the intensity of soil–tree cation cycling can thus influence the rate of soil acidification and the stocks and distribution of nutrients.
The hypothesis that water relations and growth of phreatophytic Tamarix ramosissima Ledeb. and Populus euphratica Oliv. on dunes of varying height in an extremely arid Chinese desert depend on vertical distance to a permanent water table was tested. Shoot diameter growth of P. euphratica was inversely correlated with groundwater depth (GD) of 7 to 23 m (adj. R 2 = = = = 0.69, P = = = = 0.025); growth of T. ramosissima varied independent of GD between 5 and 24 m ( P = = = = 0.385). Pre-dawn (pd) and midday (md) water potentials were lower in T. ramosissima (minimum pd − − − − 1.25 MPa, md − − − − 3.6 MPa at 24 m GD) than in P. euphratica (minimum pd − − − − 0.9 MPa, md − − − − 3.05 MPa at 23 m GD) and did not indicate physiologically significant drought stress for either species. Midday water potentials of P. euphratica closely corresponded to GD throughout the growing season, but those of T. ramosissima did not. In both species, stomatal conductance was significantly correlated with leaf water potential ( P. euphratica : adj. R 2 = = = = 0.84, P < < < < 0.0001; T. ramosissima : adj. R 2 = = = = 0.64, P = = = = 0.011) and with leaf-specific hydraulic conductance ( P. euphratica : adj. R 2 = = = = 0.79, P = = = = 0.001; T. ramosissima : adj. R 2 = = = = 0.56, P = = = = 0.019); the three variables decreased with increasing GD in P. euphratica . Stomatal conductance of P. euphratica was more strongly reduced ( > > > > 50% between − − − − 2 and − − − − 3 MPa) in response to decreasing leaf water potential than that of T. ramosissima (30% between − − − − 2 and − − − − 3 MPa). Tolerance of lower leaf water potentials due to higher concentrations of leaf osmotically active substances partially explains why leaf conductance, and probably leaf carbon gain and growth, of T. ramosissima was less severely affected by GD. Additionally, the complex below-ground structure of large clonal T. ramosissima shrub systems probably introduces variability into the assumed relationship of xylem path length with GD.
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