• allometry;
  • competition;
  • disturbance;
  • ecosystem services;
  • metabolic scaling theory;
  • Nothofagus;
  • PPA modelling;
  • productivity;
  • self-thinning;
  • southern beech;
  • WBE theory


1. Forests are an important, yet poorly understood, component of the global carbon cycle. We develop a general integrative framework for modelling the influences of stand age, environmental conditions, climate change and disturbance on woody biomass production and carbon sequestration. We use this framework to explore drivers of carbon cycling in New Zealand mountain beech forests, using a 30-year sequence of data from 246 permanent inventory plots.

2. A series of disturbance events (wind, snow storms, earthquakes and beetle outbreaks) had major effects on carbon fluxes: by killing large trees, they removed significant quantities of carbon from the woody biomass pool, and by creating canopy gaps, they reduced the crown area index (CAI) of stands (i.e. canopy area per unit ground area) and woody biomass production. A patch-dynamics model, which we parameterized using permanent plot data, predicts that episodic disturbance events can create long-term (c. 100-year) oscillations in carbon stocks at the regional scale.

3. Productivity declined with stand age, as shown in many other studies, but the effect was hard to detect because of canopy disturbance. Individual trees can increase productivity by adjusting the positioning, nutrient content and angle of leaves within canopies. We show that such optimization is most effective when trees are large and suggest it reduces the impact of water and nutrient limitation in old stands.

4. We found no evidence that forests were responding to changing climatic conditions, although strong altitudinal trends in biomass production indicate that global warming could alter carbon fluxes in future.

5.Synthesis.Our study emphasizes the critical role of disturbance in driving forest carbon fluxes. Losses of biomass arising from tree death (particularly in older stands) exceeded gains arising from growth for most of the 30-year study, moving 0.3 Mg C ha−1 year−1 from biomass to detritus and atmospheric pools. Large-scale disturbance events are prevalent in many forests world-wide, and these events are likely to be a driving factor in determining forest carbon sequestration patterns over the next century.