Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks.
We present evidence of a recent drying in the eastern Mediterranean, based on weather and tree-ring data for Samos, an island of the eastern Aegean Sea. Rainfall declined rapidly after the late 1970s following trends for the entire Mediterranean and was associated with reduced tree-ring width in Pinus brutia. The most recent decline led to the lowest annual radial stem increment after the last 100 years (as far as records reach). As moisture availability decreased best correlations of tree growth with rainfall were obtained for progressively longer integration periods (1-2 years in moister periods, 5-6 years during the severe dryness of 20th century's last decades), suggesting increasing dependency in deep soil water. Such long-term integration periods of tree-growth responses to precipitation have not been reported before. They may reflect a tree-rooting pattern adapted to cope with even several successive dry years. In late summer 2000, moisture reserves became exhausted, however, and a substantial fraction of low altitude pines died, including some 80-year-old trees, which underlines the exceptional extent this trend had reached. Our findings provide empirical support for Intergovernmental Panel on Climate Change projections derived from global circulation models that the Mediterranean, its eastern basin in particular, should become drier as temperature rises, as was the case in the recent past.
Evidence is presented of how Pinus halepensis Miller from dry habitats at <300 m elevation of four Greek island regions have responded to climatic conditions of the last two centuries. We compared historical periods of low growth due to low precipitation with the recent period of significant precipitation decline. In all cases trees' growth patterns across the twentieth century were consistent with trends in annual (rather than seasonal) precipitation, with lowest values in both precipitation and radial growth during the last two decades of the twentieth century, the worst conditions for tree growth in more than 200 years. The data are compared with trends across different vegetation belts of the northern Mediterranean basin. Drought related tree mortality in Greece in 2000 and 2007 coincided with the most severe fire outbreaks on record. IPCC WG I (2007) climate scenarios for the Mediterranean suggest a further decline in precipitation, particularly in the eastern regions. Should this occur, growth reduction in trees, tree mortality and damage from forest fires are likely to become more severe.
The Eastern Mediterranean region is among the regions which were predicted to become drier under IPCC climate scenarios. Here, we document a gradual reduction of rainfall and tree growth and the loss of rural springs during the last decades of the twentieth century. Years with severe drought are associated with very low tree growth (dendrochronology) and dry falling of springs as evidenced by interviews with local stakeholders. The paper discusses the consequences of accelerating drought on natural vegetation and agriculture and points at the interaction with fire dynamics and economy, both likely to enhance the drought effect.
We investigated forest responses to global warming by observing: (1) planted Pinus halepensis forests, (2) an aridity gradient-with annual precipitation (P) ranging from ~300 to ~700 mm, and (3) periods of wet and dry climate that included the driest period during at least the last 110 years. We examined: (1) how the length of climatic integration periods to which trees are most responsive varies in space and time, (2) the extent to which competition modulates growth decline during drought (2011) and subsequent recovery (2012) years. The temporal scale of rainfall that was most influential on growth shortened in progressing southward, and in the drier than in the wetter period. Long-term underground water storage, as reflected in the relationship of growth to multiple-year rainfall, remained significant up to the point where P ≈ 500 mm. Under drier conditions (P < 500 mm) in both space and time, influential rainfall scales shortened, probably reflecting a diminishing role of water storage. These drier locations are the first from which the species would be likely to retreat if global warming intensified. Competition appeared to set an upper limit to growth, while growth variation among individual trees increased as competition-intensity decreased. That upper limit increased in 2012 compared with 2011. The observed insensitivity of slow-growing trees to competition implies that mortality risk may be density independent, when even any potential for higher soil moisture availability in open stands is lost to evapotranspiration before it can benefit tree growth.
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