Forest growth along a rainfall gradient in Hawaii: Acacia koa stand structure, productivity, foliar nutrients, and water- and nutrient-use efficiencies

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Abstract
We tested whether variation in growth of native koa (Acacia koa) forest along a rainfall gradient was attributable to differences in leaf area index (LAI) or to differences in physiological performance per unit of leaf area. Koa stands were studied on western Kauai prior to Hurricane Iniki, and ranged from 500 to 1130 m elevation and from 850 to 1800 mm annual precipitation. Koa stands along the gradient had basal area ranging from 8 to 42 m2/ha, LAI ranging from 1.4 to 5.4, and wood increment ranging from 0.7 to 7.1 tonnes/ha/year. N, P, and K contents by weight of sun leaves (phyllodes) were negatively correlated with specific leaf mass (SLM, g m-2) across sites; on a leaf area basis, N increased whereas P and K decreased with SLM. LAI, aboveground woody biomass increment, and production per unit leaf area (E) increased as phyllode δ13C became more negative. The δ13C data suggested that intrinsic water-use efficiency (ratio of assimilation to conductance) increased as water availability decreased. In five of the six sites, phyllode P contents increased as LAI increased, but biomass increment and E were not correlated with phyllode nutrient contents, suggesting that productivity was limited more by water than by nutrient availability. Because vapor pressure deficits increased with decreasing elevation, actual water-use efficiency (ratio of assimilation to transpiration) was lower at drier, low-elevation sites. There was a trade-off between intrinsic water-use efficiency and production per unit of canopy N or P across the gradient. In summary, koa responds to water limitation both by reducing stand LAI and by adjusting gas exchange, which results in increased intrinsic water-use efficiency but decreased E.