Carbon uptake, growth and resource-use efficiency in one invasive and six native Hawaiian dry forest tree species.

Photosynthetic gas exchange, nitrogen- and water-use efficiency, leaf water potential and seasonal patterns of leaf production were studied in seven, dominant dry-forest species from the island of Lana'i, Hawaii, including the rapidly colonizing, non-native Schinus terebinthifolius (Raddi). We evaluated whether unique physiological characteristics of the invasive species explain its capacity to rapidly invade dry forests throughout the Hawaiian Islands. Apparent anomalies in stable carbon isotope data (delta13C) relative to other results led us to study effects of environmental conditions and physiological performance during leaf expansion on delta13C. Species that expanded all their foliage at the beginning of the wet season had more negative leaf delta13C values during the dry season than species with continuous leaf expansion. Among species, S. terebinthifolius had a strong seasonal pattern of leaf production and the most negative delta13C (-29 per thousand). With respect to almost every trait measured, S. terebinthifolius fell at an end of the range of values for the native species. Rapid growth of S. terebinthifolius in this ecosystem may be partially explained by its high maximum CO2 assimilation rates (15 micromol m-2 s-1), low leaf mass per area, high photosynthetic nitrogen-use efficiency per unit leaf mass or area and large decrease in stomatal conductance during the dry season. Relative to the native species, the invasive species exhibited striking phenotypic plasticity, including high rates of stem growth and water and CO2 uptake during the wet season, and maintenance of leaves and high leaf water potentials, as a result of reduced water loss, during the dry season, enabling it to utilize available resources effectively.

Temporal and spatial partitioning of water resources among eight woody species in a Hawaiian dry forest.

Lowland dry forests are unique in Hawaii for their high diversity of tree species compared with wet forests. We characterized spatial and temporal partitioning of soil water resources among seven indigenous and one invasive dry forest species to determine whether the degree of partitioning was consistent with the relatively high species richness in these forests. Patterns of water utilization were inferred from stable hydrogen isotope ratios (δD) of soil and xylem water, zones of soil water depletion, plant water status, leaf phenology, and spatial patterns of species distribution. Soil water δD values ranged from -20‰ near the surface to -48‰ at 130 cm depth. Metrosideros polymorpha, an evergreen species, and Reynoldsia sandwicensis, a drought-deciduous species, had xylem sap δD values of about -52‰, and appeared to obtain their water largely from deeper soil layers. The remaining six species had xylem δD values ranging from -33 to -42‰, and apparently obtained water from shallower soil layers. Xylem water δD values were negatively correlated with minimum annual leaf water potential and positively correlated with leaf solute content, an integrated measure of leaf water deficit. Seasonal patterns of leaf production ranged from dry season deciduous at one extreme to evergreen with near constant leaf expansion rates at the other. Species tapping water more actively from deeper soil layers tended to exhibit larger seasonality of leaf production than species relying on shallower soil water sources. Individuals of Myoporum sandwicense were more spatially isolated than would be expected by chance. Even though this species apparently extracted water primarily from shallow soil layers, as indicated by its xylem δD values, its nearly constant growth rates across all seasons may have been the result of a larger volume of soil water available per individual. The two dominant species, Diospyros sandwicensis and Nestegis sandwicensis, exhibited low leaf water potentials during the dry season and apparently drew water mostly from the upper portion of the soil profile, which may have allowed them to exploit light precipitation events more effectively than the more deeply rooted species. Character displacement in spatial and temporal patterns of soil water uptake was consistent with the relatively high diversity of woody species in Hawaiian dry forests.