When all leaning trees were considered together, regardless of social status, the mean proportion of heartwood (HW) was significantly greater on the side of the tree above the direction of lean, compared to both perpendicular and below the direction of lean. No significant differences were found in HW proportion between suppressed trees (black bars), co-dominant trees (white bars) and dominant trees (hatched bars). Where letters in superscript differ, mean data are significantly different with regard to the direction of stem lean.