Acacia Koa Research Articles

Genetic differentiation and diversity of Acacia koa populations in the Hawaiian Islands

AbstractAcacia koa A. Gray (koa) is a leguminous tree endemic to the Hawaiian Islands and can be divided into morphologically distinguishable groups of A. koaia Hillebrand, A. koa and populations that are intermediate between these extremes. The objectives of this investigation were to distinguish among divergent groups of koa at molecular levels, and to determine genetic diversity within and among the groups. Phylogenetic analyses using the ITS/5.8S rDNA and trnK intron sequences did not separate the representative koa types into distinct clusters. An unweighted pair group method with arithmetic mean cluster analysis and principal coordinate analysis, based on allele profiles of 12 microsatellite loci for 215 individual koa samples, separated the population into three distinct clusters consistent with their morphology, A. koaia, A. koa and intermediate forms. There was an average of 8.8 alleles per polymorphic locus (AP) among all koa and koaia individuals. The intermediate populations had the highest genetic diversity (H′ = 1.599), AP (7.9) and total number of unique alleles (21), whereas A. koaia and A. koa showed similar levels of genetic diversity (H′ = 0.965 and 0.943, respectively). No correlation was observed between geographic distance and genetic distance as determined by a Mantel test (r = 0.027, P = 0.91). The data presented here support previous recommendations that morphological variation within koa should be recognized at the subspecific level rather than as distinct species.

Differentiation of Acacia koa forest stands across an elevation gradient in Hawai‘i using fine-resolution remotely sensed imagery

Koa (Acacia koa) forests are found across broad environmental gradients in the Hawaiian Islands. Previous studies have identified important environmental factors controlling stand structure and productivity at the plot level, but these have not been applied at the landscape level because of small-scale spatial variability. The goal of this study is to compare the differentiation of koa forest types across an elevation/temperature gradient ranging from 1200 to 2050 m asl (17–13°C mean annual temperature (MAT)) through the analysis of field measurements of forest structure and fine-resolution remotely sensed imagery. Several vegetation indices (VIs) (atmospherically resistant vegetation index (ARVI), enhanced vegetation index (EVI), normalized difference vegetation index (NDVI), soil-adjusted vegetation index (SAVI), modified soil-adjusted vegetation index (MSAVI), simple ratio (SR) and modified simple ratio (MSR)) are calculated from IKONOS satellite imagery of these stands and analysed using supervised classification techniques. This procedure allows a clear differentiation of koa stands from areas dominated by grasses, shrubs and bare lava. Across the elevation gradient, VIs allow differentiation of three koa forest stand classes at upper, intermediate and lower elevations. In agreement with the image classification, analysis of variance (ANOVA) of tree height and leaf phosphorus (P) suggests that there are also three significantly different groups of koa stands at those elevations. A landscape-scale map of land cover and koa stand classes demonstrates both the general trend with elevation and the small-scale heterogeneity that exists across the elevation gradient. Application of these classification techniques with fine spatial resolution imagery can improve the characterization of different koa stand types across the islands of Hawai‘i, which should aid both the conservation and utilization of this ecologically important species.

Early field performance of Acacia koa seedlings grown under subirrigation and overhead irrigation

Anthony S Davis, Jeremiah R Pinto and Douglass F Jacobs

Assessment of Acacia koa forest health across environmental gradients in Hawai'i using fine resolution remote sensing and GIS.

Koa (Acacia koa) forests are found across broad environmental gradients in the Hawai‘ian Islands. Previous studies have identified koa forest health problems and dieback at the plot level, but landscape level patterns remain unstudied. The availability of high-resolution satellite images from the new GeoEye1 satellite offers the opportunity to conduct landscape-level assessments of forest health. The goal of this study was to develop integrated remote sensing and geographic information systems (GIS) methodologies to characterize the health of koa forests and model the spatial distribution and variability of koa forest dieback patterns across an elevation range of 600–1,000 m asl in the island of Kaua‘i, which correspond to gradients of temperature and rainfall ranging from 17–20 °C mean annual temperature and 750–1,500 mm mean annual precipitation. GeoEye1 satellite imagery of koa stands was analyzed using supervised classification techniques based on the analysis of 0.5-m pixel multispectral bands. There was clear differentiation of native koa forest from areas dominated by introduced tree species and differentiation of healthy koa stands from those exhibiting dieback symptoms. The area ratio of healthy koa to koa dieback corresponded linearly to changes in temperature across the environmental gradient, with koa dieback at higher relative abundance in warmer areas. A landscape-scale map of healthy koa forest and dieback distribution demonstrated both the general trend with elevation and the small-scale heterogeneity that exists within particular elevations. The application of these classification techniques with fine spatial resolution imagery can improve the accuracy of koa forest inventory and mapping across the islands of Hawai‘i. Such findings should also improve ecological restoration, conservation and silviculture of this important native tree species.

NURSERY RESPONSE OF ACACIA KOA SEEDLINGS TO CONTAINER SIZE, IRRIGATION METHOD, AND FERTILIZATION RATE

Planting koa (Acacia koa A. Gray) in Hawai‘i, USA aids in restoration of disturbed sites essential to conservation of endemic species. Survival and growth of planted seedlings under vegetative competition typically increases with initial plant size. Increasing container size and fertilizer rate may produce larger seedlings, but high fertilization can lead to surface and groundwater pollution and relatively low fertilizer use efficiency. Subirrigation systems may help mitigate this problem. Our study objective was to evaluate koa seedling growth with overhead or subirrigation over a range of container volumes (50 to 656 mL) and fertilizer rates (0 to 9.6 kg·m−3). Increasing container volume from 50 to 656 mL yielded koa seedlings with 200% more height and stem diameter growth. Subirrigation resulted in less nutrient leaching losses and yielded seedlings of similar vigor as overhead irrigated seedlings. Subirrigation helps optimize fertilizer delivery, which may improve fertilizer use efficiency and reduce environmental contamination.

Turning over a new ‘leaf’: multiple functional significances of leaves versus phyllodes in Hawaiian Acacia koa

Hawaiian endemic tree Acacia koa is a model for heteroblasty with bipinnately compound leaves and phyllodes. Previous studies suggested three hypotheses for their functional differentiation: an advantage of leaves for early growth or shade tolerance, and an advantage of phyllodes for drought tolerance. We tested the ability of these hypotheses to explain differences between leaf types for potted plants in 104 physiological and morphological traits, including gas exchange, structure and composition, hydraulic conductance, and responses to varying light, intercellular CO(2) , vapour pressure deficit (VPD) and drought. Leaf types were similar in numerous traits including stomatal pore area per leaf area, leaf area-based gas exchange rates and cuticular conductance. Each hypothesis was directly supported by key differences in function. Leaves had higher mass-based gas exchange rates, while the water storage tissue in phyllodes contributed to greater capacitance per area; phyllodes also showed stronger stomatal closure at high VPD, and higher maximum hydraulic conductance per area, with stronger decline during desiccation and recovery with rehydration. While no single hypothesis completely explained the differences between leaf types, together the three hypotheses explained 91% of differences. These findings indicate that the heteroblasty confers multiple benefits, realized across different developmental stages and environmental contexts.

Comparison of Dissolved Organic Carbon Bioavailability from Native and Invasive Vegetation along a Hawaiian River

Riparian litter fall is an important source of organic matter to rivers and accounts for a large fraction of their dissolved organic carbon (DOC) load. DOC is metabolically important in rivers, and therefore changes in riparian vegetation species composition should affect riverine DOC bioavailability. Worldwide, invasive vegetation composes a large percentage of riparian vegetation. In Hawai'i, riparian vegetation changes from native to invasive species with decreasing elevation. To assess how changes in riparian vegetation affect riverine DOC dynamics, we compared DOC bioavailability from native (Acacia koa and Metrosideros polymorpha) and invasive (Falcataria moluccana and Psidium cattleianum) riparian trees to freshwater and estuarine bacteria from the Wailuku River on Hawai'i Island through dark bioassays. DOC bioavailabilities in riverine and estuarine waters were similar among all riparian vegetation types. In contrast, vegetation-derived DOC was more bioavailable (52% ± 4%) than the riverine and estuarine DOC (14% ± 3%). Combining DOC bioavailability and leaf litter input data from our native and invaded riparian sites suggests that a shift in leaf litter inputs from native to invasive species may increase the amount of bioavailable DOC entering Hawaiian rivers and streams. This DOC input has the potential to impact the metabolism and food webs of downstream ecosystems.

Top-Down Analysis of Forest Structure and Biogeochemistry Across Hawaiian Landscapes

Technical and analytical improvements in aircraft-based remote sensing allow synoptic measurements of structural and chemical properties of vegetation across whole landscapes. We used the Carnegie Airborne Observatory, which includes waveform light detection and ranging (LiDAR) and high-fidelity imaging spectroscopy, to evaluate the landscapes surrounding four well-studied sites on a substrate age gradient across the Hawaiian Islands. The airborne measurements yielded variations in ground topography, canopy height, and canopy nitrogen (N) concentration more accurately than they could have been obtained by any reasonable intensity of ground-based sampling. We detected spatial variation in ecosystem properties associated with the properties of different species, including differences in canopy N concentrations associated with the native species Metrosideros polymorpha and Acacia koa, and differences brought about by invasions of the biological N fixer Morella faya. Structural and chemical differences associated with exotic tree plantations and with dominance of forest patches by the native mat-forming fern Dicranopteris linearis also could be analyzed straightforwardly. This approach provides a powerful tool for ecologists seeking to expand from plot-based measurements to landscape-level analyses.

Physiological and anatomical responses of Acacia koa (Gray) seedlings to varying light and drought conditions

Abstract True leaves of Acacia koa (Gray) seedlings play a vital role in the early stages of seedling establishment, particularly in the Hawaiian archipelago where regeneration occurs under very heterogeneous light and moisture conditions. Given the importance of understanding seedling regeneration in the highly disturbed forests of Hawaii, we studied the growth, biomass allocation, physiological, and anatomical characteristics of A. koa seedlings with true leaves in response to three levels of light (low, intermediate, and high sunlight) and two levels of moisture regimes (100% field capacity, 20% field capacity) in a greenhouse experiment. With increasing light intensity, seedlings exhibited greater growth, biomass accumulation, intrinsic water-use efficiency, more foliar C, and higher stomatal density. Their leaflets were thicker, containing more palisade and spongy parenchyma. Low light intensity ameliorated drought-related effects of carbon assimilation on A. koa seedlings, while seedling growth of drought-stressed seedlings was reduced under intermediate and high sunlight. Our results suggest that true leaves of A. koa seedlings are adaptive when light is the only limiting environmental variable. The low adaptive capacity of true leaves to drought stress under high light intensity negatively affected seedling performance, which possibly explains environmental conditions under which phyllodes are likely to emerge.

Poor Stem Form as a Potential Limitation to Private Investment in Koa Plantation Forestry in Hawaii

Providing economic incentives to landholders is an effective way of promoting sustainable forest management, conservation and restoration. In Hawaii, the main native hardwood species with commercial value is Acacia koa (koa), but lack of successful examples of koa plantation forestry hinders private investment. Financial models, which have been offered to encourage investment, assume that a particular minimum volume of merchantable wood will be harvested at rotation age. No studies have been done to determine the appropriateness of the assumed volumes for koa plantations. Three 18- to 28-year-old koa plantations were studied to determine butt-log size, crown class and crop tree potential. Estimates of present and future merchantable sawtimber volume were made for each plantation. Results indicate that most existing plantation koa trees fork so close to the ground that they will produce little to no merchantable wood. Projected sawtimber yields from crop trees in these plantations at age 45 years are 10, 15 and 90 m 3 /ha, which are below the 135 m 3 /ha yield used in the most recent financial analysis of returns on investment in koa forestry. Relaxing the crop tree criteria gives higher volume yields, but two plantations still fall short of the assumed target yield by 43 and 52%.

Tree invasion in managed tropical forests facilitates endemic species

AbstractAim To determine whether different abundances of introduced species of Cinchona (Rubiaceae) affect species composition and facilitate species richness in managed tropical forests, to test whether any facilitative effects on understorey species depend on forest type, and to investigate whether facilitative effects can be attributed to the ‘substitutive facilitation model’.Location Makawao Forest Reserve on Maui, Hawai’i, USA.Methods Cinchona species (Cinchona pubescens and Cinchona calisaya) were mapped within various forest types. In three forest types (ageing Eucalyptus and Pinus plantations, and near‐natural Acacia koa forests), we analysed environmental parameters (e.g. canopy cover, litter cover, pH value and soil depth) and the species composition of Cinchona‐invaded and non‐invaded plots; data were compared based on Cinchona cover and forest types. Habitat modelling for several endemic species and tree ferns was carried out to test whether Cinchona cover is an important variable for the probability of occurrence of these endemics.Results Cinchona species have naturalized mainly in Eucalyptus and Pinus plantations and Acacia koa forests and here add an additional shrub layer. In contrast to other studies, we revealed facilitative effects of Cinchona on native species within all forest types. Species richness is about 20% higher in invaded plots than in non‐invaded plots, and these show a nearly 50% higher proportion of endemic species, including tree ferns. The proportion of endemics even increases with increasing Cinchona cover. For several endemics, Cinchona is found to be an important variable for the probability of occurrence, and the removal of Cinchona cover as an explanatory variable lowers the model fit. In addition to Cinchona, variables delineating vegetation structure and light availability have a strong effect on the model fit.Main conclusions In the structurally simplified Hawaiian forests studied, Cinchona facilitated endemic species in accordance with the ‘substitutive facilitation model’. This contrasts with the results of an earlier study in the naturally treeless Galápagos highlands, which revealed a sharp decrease in the abundance of endemics under Cinchona canopy. These results illustrate that, through the same structural change (addition of a vegetation layer), an invasive species may exert divergent effects across different ecosystem types. The facilitation of endemic understorey species by invasive tree species in managed forests leads to a dilemma in conservation but also to new perspectives for ecosystem restoration.

A Recent Outbreak of the Hawaiian Koa Moth,Scotorythra paludicola(Lepidoptera: Geometridae), and a Review of Outbreaks between 1892 and 2003

The koa moth, Scotorythra paludicola, is an endemic Hawaiian moth that undergoes sporadic outbreaks in koa forests in Hawai‘i, causing vast defoliations of its host plant, Acacia koa. We studied one such outbreak that occurred on East Maui in 2003, in which approximately 16 km2 of forest were defoliated. We collected adult moths and larvae, and recorded size-class distribution of larvae in defoliated regions. Larvae at a given site tended to be of a similar size class, suggesting that outbreaks were synchronous, and mean development time from first instar to adulthood was 42 days under laboratory conditions. Mortality of field-collected, laboratory-reared larvae due to disease was high (80%), making it impossible to quantify meaningful parasitism rates, but three nonnative hymenopteran primary parasitoids were reared (the braconids Meteorus laphygmae and Cotesia marginiventris, and the ichneumonid Hyposoter exiguae). One ichneumonid hyperparasitoid, Gelis sp., was also reared. No native parasitoids were reared. We found no relationship between occurrence of five koa moth outbreaks on East Maui between 1920 and 2006 and annual or monthly precipitation or temperature during that period.

Effects of fertilisation on phosphorus pools in the volcanic soil of a managed tropical forest

Acacia koa forests benefit from phosphorus fertilisation, but it is unknown if fertilisation is a short or long term effect on P availability. Past research suggests that P cycling in soils with high P sorption capacity, such as Andisols, was through organic pathways. We studied leaf P and soil P fractions in a tropical forest Andisol for 3 years after fertilisation with triple super phosphate. Leaf P concentration and labile P remained high after fertilisation. Fertilisation had increased all the inorganic P fractions over the length of the study, while organic P fractions had not. The results suggested that the organic P fractions had a reduced role as a source of labile P after fertilisation. The size and dynamics of the sodium hydroxide- and hydrochloric acid-extractable P pools would suggest that either pool could be major sources of labile P. Because of the high level of poorly crystalline minerals in Andisols (allophone and imogolite), it would be expected that applied P would quickly lead to strong P sorption onto mineral surfaces and thus a rapid decline in P availability. We propose that the high organic matter present in these soils had masked some of the Al and Fe sorption sites, which allowed the sorption and desorption when large amounts of addition P were applied.

Inoculating Acacia koa with Bradyrhizobium and Applying Fertilizer in the Nursery: Effects on Nodule Formation and Seedling Growth

Restoration of Acacia koa A. Gray (koa) forests on degraded sites in Hawaii is important for conservation of rare, endemic plants and animals and is often accomplished by planting nursery-grown seedlings. To be successful after outplanting, koa seedlings must access sufficient nutrients from the soil and outcompete other vegetation. Forming symbiotic associations with nitrogen-fixing microorganisms may help alleviate inherent nutrient deficiencies found on volcanic sites, but interactions between inoculation and fertilization during production of koa seedlings are not well studied. Under operational nursery conditions, we fertilized koa seedlings at six controlled-release fertilizer (CRF) rates (ranging from 0 to 11.9 kg·m−3 Osmocote Plus 15N–9P–12K), which were also either inoculated with a locally collected source of nitrogen-fixing Bradyrhizobium spp. or not. Fertilized seedlings were significantly larger than the control; on average, fertilized seedlings were 300% taller and had 180%, 700%, and 350% more root-collar diameter, shoot biomass, and root biomass, respectively. Nitrogen (N) concentration was 23% to 119% greater in fertilized seedlings than nonfertilized seedlings. Inoculation with Bradyrhizobium had no significant affect on seedling characteristics except for root N concentration, which was 8% higher for inoculated seedlings. Fertilizer rate and inoculation interacted to significantly affect nodule number and nodule biomass/seedling. With inoculation, we observed an increase in nodule number (16.5 to 25.5) and biomass (5.8 mg to 11.7 mg) with fertilizer rate up to 4.7 kg·m−3 but a decline to nearly zero at the highest rates. Without inoculation, nodule formation was scarce (3.2 nodules/seedling or less) and only observed when CRF was 2.3 kg·m−3 or less. Our results suggest that high rates of fertilization and successful nodulation are not mutually exclusive nursery practices. Concurrent use of fertilization and inoculation in the nursery may provide opportunity to produce larger seedlings that have high potential to reduce the time needed to achieve canopy closure and thereby help achieve restoration objectives.

Altering Light Availability to Restore Invaded Forest: The Predictive Role of Plant Traits

Many studies have demonstrated that reduced light availability, which can be manipulated at local scales by planting or seeding canopy species, can curtail the growth of invasive species and promote the growth of native species. Species differences in functional traits, such as light use and stress tolerance, may be used to determine how native and invasive species will respond to these resource manipulations. We altered light availability in a mesic Hawaiian forest understory and found that low light levels reduced the biomass and growth of two invasive grasses (Pennisetum clandestinum and Ehrharta stipoides) relative to two native shrubs (Pipturus albidus and Coprosma rhynchocarpa) and two native canopy species (Metrosideros polymorpha and Acacia koa). Native species generally displayed traits associated with shade tolerance (high quantum yield, chlorophyll content, and leaf mass per area), whereas the two invasive grasses displayed traits associated with shade intolerance (high photosynthetic rate and growth rate). Several key traits pertaining to light acquisition and shade tolerance (quantum yield, chlorophyll content, and leaf mass per area) predicted seedling survival in low-light treatments. Our data suggest that differences in light use among native and invasive species can help to determine the utility of resource manipulation as a restoration tool and, more specifically, to predict which native species will be optimal for restoration efforts that manipulate light availability.

Development and characterization of microsatellite markers for analysis of population differentiation in the tree legumeAcacia koa(Fabaceae: Mimosoideae) in the Hawaiian Islands

The aim of this research was to develop and use microsatellite markers to characterize the high-value timber tree Acacia koa (koa), which is endemic to the Hawaiian Islands. Genomic DNA fragments of 300-1000 bp were cloned and sequenced following enrichment for microsatellite motifs by PCR using 7 oligonucleotide repeat primers in separate reactions. Among 96 sequences analyzed, 63 contained unique microsatellite motifs flanked by variable sequences. A dual PCR method involving a primer walking step was used to develop 15 primer pairs. Another 16 primer pairs were developed directly from the variable sequences on both sides of the microsatellite motifs. These 31 primer pairs were tested on 172 koa plants representing 11 populations collected from 4 of the major Hawaiian Islands. Nine of the primers that identified polymorphic microsatellite loci and 3 that detected unique alleles exclusively in some populations were used for genetic diversity studies of koa. Cluster analysis and multidimensional scaling of the allelic phenotype data revealed that koa from Kauai formed a distinct group separate from koa of the neighboring islands of Oahu, Maui, and Hawaii. The oldest of the four islands, Kauai, also had the most diverse populations of koa.

Wood Production and Native Vegetation Conservation in Hawaii's Uplands

Abstract Stand basal area, a stochastic growth model (KOA), and GIS were used to simulate and spatially integrate potential timber production of the native koa (Acacia koa) in uplands on the island of Hawaii. Vegetation patterns were examined to estimate potential impacts on the ecosystem from increased forest activities and to select suitable areas for wood production and ecological preservation. The KOA model predicted a production of 82,200 m3 of merchantable koa wood at age 30 from nonmanaged, naturally regenerated stands. The spatial analysis revealed four corridors with high wood production. Mean vegetation patch size was 69 ha, with a greater frequency of small patches than of large patches. An index of patch shape and complexity was similar for patches from different categories, but it was inversely correlated to patch size, indicating different processes affecting landscape structure. A shape diversity index was greater for savannah patches than for closed canopy units, implying higher complexity in more disturbed areas. Patches within the same category were highly aggregated, and the core of the area with lower degree of disturbance matched low-productivity sites for koa.

Responses of native and invasive plant species to selective logging in an Acacia koa‐Metrosideros polymorpha forest in Hawai'i

AbstractQuestions: Is the introduced timber species Fraxinus uhdei invasive in Hawai'i? Has logging disturbance facilitated the spread of Fraxinus and other alien species?Location: Windward Mauna Kea, island of Hawai'i.Methods: We surveyed 29 plots which were established before selective logging of the native tree Acacia koa in 1971 to determine if Fraxinus spread beyond the borders of an existing plantation and if other alien species increased. We created gaps in the canopy of the Fraxinus plantation and measured seed rain and regeneration, and we sampled foliar and soil nutrients inside and around the plantation.Results: Basal area of Fraxinus increased from 0.7 m2.ha‐1in 1971 to 10.8 m2.ha‐1 in 2000. Fraxinus was not found in plots that were located more than 500 m from those where it occurred in 1971 except along a road. Basal area of Acacia koa decreased after logging but subsequently recovered. Occurrence of the alien vine Passiflora tarminiana and alien grass Ehrharta stipoides decreased. Seedling regeneration of Fraxinus was prolific in gaps but did not occur under the canopy. Basal area of Fraxinus did not correlate with soil nutrient concentrations.Conclusions: Fraxinus was able to regenerate following logging more rapidly than native tree species. Basal area growth of Fraxinus was great enough to offset a decline in native trees and cause an increase in forest productivity. If the Fraxinus plantation is harvested, managers should plan ways of favoring regeneration of the native Acacia which is more valuable both for timber and for conservation.

Assessing the conservation value of a human-dominated island landscape: Plant diversity in Hawaii

The conversion of native habitats to pasture and other working lands, unbuilt lands modified by humans for production, is one of the greatest threats to biodiversity. While some human-dominated landscapes on continents support relatively high native biodiversity, this capacity is little studied in oceanic island systems characterized by high endemism and vulnerability to invasion. Using Hawaii as a case study, we assessed the conservation value of working landscapes on an oceanic island by surveying native and non-native plant diversity in mature native forest and in the three dominant land covers/uses to which it has been converted: native, Acacia koa timber plantations, wooded pasture, and open pasture. As expected, native plant diversity (richness and abundance) was significantly higher and non-native abundance significantly lower in mature native forests than any other site type. A. koa plantations and wooded pasture supported four and three times greater, respectively, species richness of native understory plants than open pasture. Also, A. koa plantations and wooded pasture supported similar species communities with about 75% species in common. Conservation and restoration of mature native forest in Hawaii is essential for the protection of native, rare species and limiting the spread of non-native species. A. koa plantations and wooded pasture, however, may help harmonize production and conservation by supporting livelihoods, more biodiversity than open pasture, and some connectivity between native forest remnants important for sustaining landscape-level conservation value into the future.

Sudden and sustained response of Acacia koa crop trees to crown release in stagnant stands

We examined the growth response of Acacia koa Gray (koa) in dense (10 000 trees/ha), ~25-year-old, second-growth stands on Hawai‘i to crown release (thinning) and grass control. Koa diameter increment increased within 2 months of thinning across the range of residual densities (200–900 trees/ha). After 3 years, diameter increment in the most intense thinning treatment was 180% greater than in the control treatment (1.4 vs. 0.5 cm/year); in the least intense thinning treatment diameter increment was 100% greater than the control (1.0 vs. 0.5 cm/year). Koa’s growth response was independent of all measures of tree size (diameter at breast height, height, and leaf area) across the range of tree sizes sampled. A grass control treatment to determine if reduced competition for soil water would improve koa growth showed no improvement in growth response. Koa phyllode δ13C values, which represent an integrated measure of tree water status, showed no evidence of tree water stress (mean δ13C = –28.3‰) and were not significantly different between the grass removal and control treatments. These results demonstrate that koa is capable of responding quickly and positively to crown release despite years of suppressed growth and suggest that landowners interested in managing koa forests may have considerable flexibility in the timing and application of thinning treatments to koa stands.