Wood density is related to aboveground biomass and productivity along a successional gradient in upper Andean tropical forests

Wood density (WD) is a key functional trait related to ecological strategies and ecosystem carbon dynamics. Despite its importance, there is a considerable lack of information on WD in tropical Andean forests, particularly regarding its relationship with forest succession and ecosystem carbon cycling. Here, we quantified WD in 86 upper Andean tree and shrub species in central Colombia, with the aim of determining how WD changes with forest succession and how it is related to productivity. We hypothesized that WD will increase with succession because early successional forests will be colonized by acquisitive species, which typically have low WD, while the shaded understory of older forests should favor higher WD. We measured WD in 481 individuals from 27 shrub and 59 tree species, and quantified aboveground biomass (AGB), canopy height, net primary production (NPP) and species composition and abundance in 14, 400-m(2), permanent plots. Mean WD was 0.513 ± 0.114 (g/cm(3)), with a range between 0.068 and 0.718 (g/cm(3)). Shrubs had, on average, higher WD (0.552 ± 0.095 g/cm(3)) than trees (0.488 ± 0.104 g/cm(3)). Community weighted mean WD (CWMwd) decreased with succession (measured as mean canopy height, AGB, and basal area); CWMwd also decreased with aboveground NPP and stem growth. In contrast, the percentage of NPP attributed to litter and the percent of shrubs in plots increased with CWMwd. Thus, our hypothesis was not supported because early successional forests had higher CWMwd than late successional forests. This was related to a high proportion of shrubs (with high WD) early in succession, which could be a consequence of: 1) a low seed availability of trees due to intense land use in the landscape and/or 2) harsh abiotic conditions early in succession that filter out trees. Forest with high CWMwd had a high %NPP attributed to litter because they were dominated by shrubs, which gain little biomass in their trunks. Our findings highlight the links between WD, succession and carbon cycling (biomass and productivity) in this biodiversity hotspot. Thus, WD is an important trait that can be used to understand upper Andean forest recovery and improve forest restoration and management practices.