Old-growth forests remove carbon dioxide from the atmosphere at rates that vary with climate and nitrogen deposition. The sequestered carbon dioxide is stored in live woody tissues and slowly decomposing organic matter in litter and soil. Old-growth forests therefore serve as a global carbon dioxide sink, but they are not protected by international treaties, because it is generally thought that ageing forests cease to accumulate carbon. Here we report a search of literature and databases for forest carbon-flux estimates. We find that in forests between 15 and 800 years of age, net ecosystem productivity (the net carbon balance of the forest including soils) is usually positive. Our results demonstrate that old-growth forests can continue to accumulate carbon, contrary to the long-standing view that they are carbon neutral. Over 30 per cent of the global forest area is unmanaged primary forest, and this area contains the remaining old-growth forests. Half of the primary forests (6 x 10(8) hectares) are located in the boreal and temperate regions of the Northern Hemisphere. On the basis of our analysis, these forests alone sequester about 1.3 +/- 0.5 gigatonnes of carbon per year. Thus, our findings suggest that 15 per cent of the global forest area, which is currently not considered when offsetting increasing atmospheric carbon dioxide concentrations, provides at least 10 per cent of the global net ecosystem productivity. Old-growth forests accumulate carbon for centuries and contain large quantities of it. We expect, however, that much of this carbon, even soil carbon, will move back to the atmosphere if these forests are disturbed.

Overviewing the European carbon (C), greenhouse gas (GHG), and non-GHG fluxes, gross primary productivity (GPP) is about 9.3 Pg yr-1, and fossil fuel imports are 1.6 Pg yr-1. GPP is about 1.25% of solar radiation, containing about 360 x 1018 J energy - five times the energy content of annual fossil fuel use. Net primary production (NPP) is 50%, terrestrial net biome productivity, NBP, 3%, and the net GHG balance, NGB, 0.3% of GPP. Human harvest uses 20% of NPP or 10% of GPP, or alternatively 1 parts per thousand of solar radiation after accounting for the inherent cost of agriculture and forestry, for production of pesticides and fertilizer, the return of organic fertilizer, and for the C equivalent cost of GHG emissions. C equivalents are defined on a global warming potential with a 100-year time horizon. The equivalent of about 2.4% of the mineral fertilizer input is emitted as N(2)O. Agricultural emissions to the atmosphere are about 40% of total methane, 60% of total NO-N, 70% of total N(2)O-N, and 95% of total NH(3)-N emissions of Europe. European soils are a net C sink (114 Tg yr-1), but considering the emissions of GHGs, soils are a source of about 26 Tg CO(2) C-equivalent yr-1. Forest, grassland and sediment C sinks are offset by GHG emissions from croplands, peatlands and inland waters. Non-GHGs (NH(3), NOx) interact significantly with the GHG and the C cycle through ammonium nitrate aerosols and dry deposition. Wet deposition of nitrogen (N) supports about 50% of forest timber growth. Land use change is regionally important. The absolute flux values total about 50 Tg C yr-1. Nevertheless, for the European trace-gas balance, land-use intensity is more important than land-use change. This study shows that emissions of GHGs and non-GHGs significantly distort the C cycle and eliminate apparent C sinks

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