Including tree spatial extension in the evaluation of neighborhood competition effects in Bornean rain forest

Classical tree neighborhood models use size variables acting at point distances. In a new approach here, trees were spatially extended as a function of their crown sizes, represented impressionistically as points within crown areas. Extension was accom- panied by plasticity in the form of crown removal or relocation under the overlap of taller trees. Root systems were supposedly extended in a similar manner. For the 38 most abundant species in the focal size class (10- < 100 cm stem girth) in two 4-ha plots at Danum (Sabah), for periods P 1 (1986-1996) and P 2 (1996-2007), stem growth rate and tree survival were individually regressed against stem size, and neighbor- hood conspecific (CON) and heterospecific (HET) basal areas within incremented steps in radius. Model parameters were critically assessed, and statistical robustness in the modeling was set by randomization testing. Classical and extended models dif- fered importantly in their outcomes. Crown extension weakened the relationship of CON effect on growth versus plot species' abundance, showing that models without plasticity overestimated negative density dependence. A significant negative trend of difference in CON effects on growth (P 2 −P 1 ) versus CON or HET effect on survival in P 1 was strongest with crown extension. Model outcomes did not then support an explanation of CON and HET effects being due to (asymmetric) competition for light alone. An alternative hypothesis is that changes in CON effects on small trees, largely incurred by a drought phase (relaxing light limitation) in P 2 , and following the more shaded (suppressing) conditions in P 1 , were likely due to species-specific (symmetric) root competition and mycorrhizal processes. The very high variation in neighborhood composition and abundances led to a strong "neighborhood stochasticity" and hence to largely idiosyncratic species' responses. A need to much better understand the roles of rooting structure and processes at the individual tree level was highlighted.