title: Trees increase their P:N ratio with size.
abstract: OBJECTIVE
  Phosphorus (P) tends to become limiting in aging terrestrial ecosystems, and its resorption efficiency is higher than for other elements such as nitrogen (N). We thus hypothesized that trees should store more P than those other elements such as N when tree size increases and that this process should be enhanced in slow-growing late successional trees.
  
  
  METHODS
  Catalan forests.
  
  
  METHODS
  We have used data from the Catalan Forest Inventory that contains field data of the P and N contents of total aboveground, foliar and woody biomasses of the diverse Mediterranean, temperate and alpine forests of Catalonia (1018 sites). We used correlation and general lineal models (GLM) to analyze the allometric relationships between nutrient contents of different aboveground biomass fractions (foliar, branches and stems) and total aboveground biomass.
  
  
  RESULTS
  Aboveground forest P content increases proportionally more than aboveground forest N content with increasing aboveground biomass. Two mechanisms underlie this. First, woody biomass increases proportionally more than foliar biomass having woody biomass higher P:N ratio than foliar biomass. Second, wood P:N ratio increases with tree size. These results are consistent with the generally higher foliar resorption of P than of N. Slow-growing species accumulate more P in total aboveground with size than fast-growing species mainly as a result of their large capacity to store P in wood.
  
  
  CONCLUSIONS
  Trees may have thus developed long-term adaptive mechanisms to store P in biomass, mainly in wood, thereby slowing the loss of P from the ecosystems, reducing its availability for competitors, and implying an increase in the P:N ratio in forest biomass with aging. This trend to accumulate more P than N with size is more accentuated in slow-growing, large, long-living species of late successional stages. This way they partly counterbalance the gradual decrease of P in the soil.
authors.affiliation: CSIC, Global Ecology Unit CREAF-CEAB-CSIC-UAB, Cerdanyola del Vallés, 08193 Barcelona, Catalonia, Spain. ; CREAF, Cerdanyola del Vallés, 08193 Barcelona, Catalonia, Spain.
authors.forename: J
authors.initials: J
authors.lastname: Sardans
authors.affiliation: CSIC, Global Ecology Unit CREAF-CEAB-CSIC-UAB, Cerdanyola del Vallés, 08193 Barcelona, Catalonia, Spain. ; CREAF, Cerdanyola del Vallés, 08193 Barcelona, Catalonia, Spain.
authors.forename: J
authors.initials: J
authors.lastname: Peñuelas
keywords: Early-succesional species; N:P; forest; late-succesional species; leaf:wood ratio; nitrogen; phosphorus; soil; stoichiometry