Long-term decline of the Amazon carbon sink.

Atmospheric carbon dioxide records indicate that the land surface has acted as a strong global carbon sink over recent decades, with a substantial fraction of this sink probably located in the tropics, particularly in the Amazon. Nevertheless, it is unclear how the terrestrial carbon sink will evolve as climate and atmospheric composition continue to change. Here we analyse the historical evolution of the biomass dynamics of the Amazon rainforest over three decades using a distributed network of 321 plots. While this analysis confirms that Amazon forests have acted as a long-term net biomass sink, we find a long-term decreasing trend of carbon accumulation. Rates of net increase in above-ground biomass declined by one-third during the past decade compared to the 1990s. This is a consequence of growth rate increases levelling off recently, while biomass mortality persistently increased throughout, leading to a shortening of carbon residence times. Potential drivers for the mortality increase include greater climate variability, and feedbacks of faster growth on mortality, resulting in shortened tree longevity. The observed decline of the Amazon sink diverges markedly from the recent increase in terrestrial carbon uptake at the global scale, and is contrary to expectations based on models.

Leaf traits and herbivory rates of tropical tree species differing in successional status.

We evaluated leaf characteristics and herbivory intensities for saplings of fifteen tropical tree species differing in their successional position. Eight leaf traits were selected, related to the costs of leaf display (specific leaf area [SLA], water content), photosynthesis (N and P concentration per unit mass), and herbivory defence (lignin concentration, C:N ratio). We hypothesised that species traits are shaped by variation in abiotic and biotic (herbivory) selection pressures along the successional gradient. All leaf traits varied with the successional position of the species. The SLA, water content and nutrient concentration decreased, and lignin concentration increased with the successional position. Herbivory damage (defined as the percentage of damage found at one moment in time) varied from 0.9-8.5% among the species, but was not related to their successional position. Herbivory damage appeared to be a poor estimator of the herbivory rate experienced by species, due to the confounding effect of leaf lifespan. Herbivory rate (defined as percentage leaf area removal per unit time) declined with the successional position of the species. Herbivory rate was only positively correlated to water content, and negatively correlated to lignin concentration, suggesting that herbivores select leaves based upon their digestibility rather than upon their nutritive value. Surprisingly, most species traits change linearly with succession, while resource availability (light, nutrients) declines exponentially with succession.

Photosynthetic rates in relation to leaf phosphorus content in pioneer versus climax tropical rainforest trees.

In Guyana dense rainforest occurs on intensely weathered acid soils, low in soil phosphorus. To investigate whether low P availability limits photosynthesis of trees growing on these soils more than N does, leaf P and N content, and their relationship with the photosynthetic capacity (A sat, µmol CO2 m-2 s-1) were studied for nine pioneer and climax tree species in a range of light climates. Light environment was described using hemispherical photographs. For both pioneer and climax species, leaf P content (r 2=0.71 and 0.23, respectively) is a more important determinant of A sat than leaf N content (r 2=0.54 and 0.12, respectively). Pioneer species have a higher leaf P and N content than climax species. At similar P or N content, pioneers have a higher A sat than climax species. The saplings studied had a relatively high A sat, considering their low P concentration (15-30 µmol P g-1). All species studied had a constant leaf P and N concentration and photosynthetic capacity across light climates, because specific leaf mass (g m-2) increased similarly with light availability. This acclimation to a change in light environment makes a possible limitation of A sat by P or N independent of light environment.

Phenotypic plasticity in response to nitrate supply of an inherently fast-growing species from a fertile habitat and an inherently slow-growing species from an infertile habitat.

The aim of the present study was to investigate possible differences in plasticity between a potentially fast-growing and a potentially slow-growing grass species. To this end, Holcus lanatus (L.) and Deschampsia flexuosa (L.) Trin., associated with fertile and infertile habitats, respectively, were grown in sand at eight nitrate concentrations. When plants obtained a fresh weight of approximately 5 g, biomass allocation, specific leaf area, the rate of net photosynthesis, the organic nitrogen concentration of various plant parts and the root weight at different soil depths were determined. There were linear relationships between the morphological and physiological features studied and the In-transformed nitrate concentration supplied, except for the specific leaf area and root nitrogen concentration of H. lanatus, which did not respond to the nitrate concentration. The root biomass of H. lanatus was invariably distributed over the soil layers than that of D. flexuosa. However, D. flexuosa allocated more root biomass to lower soil depths with decreasing nitrate concentration, in contrast to H. lanatus, which did not respond. The relative response to nitrate supply, i.e. the value of a character at a certain nitrate level relative to the value of that character at the highest nitrate supply, was used as a measure for plasticity. For a number of parameters (leaf area ratio, root weight ratio, root nitrogen concentration, vertical root biomass distribution and rate of net photosynthesis per unit leaf weight) the potentially slow-growing D. flexuosa exhibited a higher phenotypic plasticity than the potentially fast-growing H. lanatus. These findings are in disagreement with current literature. Possible explanations for this discrepancy are discussed in terms of differences in experimental approach as well as fundamental differences in specific traits between fast- and slow-growing grasses.

The relation between above- and belowground biomass allocation patterns and competitive ability.

In a 2-year experiment, the evergreen shrubsErica tetralix andCalluna vulgaris (dominant on nutrient-poor heathland soils) and the perennial deciduous grassMolinia caerulea (dominant on nutrient-rich heathland soils) were grown in replacement series in a factorial combination of four competition types (no competition, only aboveground competition, only belowground competition, full competition) and two levels of nutrient supply (no nutrients and 10 g N+2 g P+10 g K m-2 yr-1). Both in the unfertilized and in the fertilized treatmentsMolinia allocated about twice as much biomass to its root system than didErica andCalluna. In all three species the relative amount of biomass allocated to the roots was lower at high than at low nutrient supply. The relative decrease was larger forMolinia than forErica andCalluna. In the fertilized monocultures biomass of all three species exceeded that in the unfertilized series.Molinia showed the greatest biomass increase. In the unfertilized series no effects of interspecific competition on the biomass of each species were observed in either of the competition treatments. In the fertilized mixtures where only belowground competition was possibleMolinia increased its biomass at the expense of bothErica andCalluna. When only aboveground competition was possible no effects of interspecific competition on the biomass of the competing species were observed. However, in contrast with the evergreens,Molinia responded by positioning its leaf layers relatively higher in the canopy. The effects of full competition were similar to those of only belowground competition, so in the fertilized series belowground competition determined the outcome of competition. The high competitive ability ofMolinia at high nutrient supply can be attributed to the combination of (1) a high potential productivity, (2) a high percentage biomass allocation to the roots, (3) an extensive root system exploiting a large soil volume, and (4) plasticity in the spatial arrangement of leaf layers over its tall canopy. In the species under study the allocation patterns entailed no apparent trade-off between the abilities to compete for above- and belowground resources. This study suggests that this trade-off can be overcome by: (1) plasticity in the spatial arrangement of leaf layers and roots, and (2) compensatory phenotypic and species-specific differences in specific leaf area and specific root length.