Restoration of root hemiparasite trees, such as Hawaiʻi's endemic Santalum species (ʻiliahi), may benefit from underplanting in stands of suitable hosts like the nitrogen‐fixing native tree, Acacia koa (koa). At a pasture site on Hawaiʻi Island previously reforested with koa, we underplanted seedlings of the island‐endemic sandalwood species, Santalum paniculatum , to examine the tradeoff between access to an established root network (distance to the nearest koa tree) under variable overstory shading (8.8–90.1% canopy openness range) during regeneration establishment. We hypothesized that there is an optimal parasite–host spacing and canopy openness that balance parasitic resource transfer with light availability. ʻIliahi seedling survival was 96% with no survival treatment differences. ‘Iliahi seedling growth was positively related to canopy openness but negatively related to the distance to the nearest koa tree, and the slope of these relationships increased over time. Leaf photosynthetic light compensation points, light saturation points, and stomatal density mostly followed similar trends as growth. These results demonstrate that ‘iliahi can be successfully underplanted in an established koa stand, which benefits ‘iliahi plantings and contributes to diversifying initial restoration and reforestation plantings. There appears to be a significant tradeoff in planting distance between benefits from and competition with the host; however, the improvement in growth with increased canopy openness appeared to be much greater than the effect of planting distance. Underplanting into an established host stand with sufficient canopy openness can help restore functionally compatible and abundant ‘iliahi regeneration into forests.

Root hemiparasitic trees can photosynthesize yet also acquire resources from host plants, which may benefit the long-term survival and growth of the hemiparasite. Experimental evaluation of planting distance between tree hemiparasites and their hosts can lead to biological insights and contribute to practitioner decision making. On an abandoned pasture site in a historically tropical dry forest in Hawai'i, we studied the effects of two host species and four planting distances on the survival and growth of 'iliahi (Hawaiian sandalwood, Santalum pan-iculatum), an endemic root hemiparasitic tree. Treatments included a control with no host or one of two native host species: 'a'ali'i (Dodonaea viscosa), a fast-growing shrub, or koa (Acacia koa), a fast-growing nitrogen-fixing tree. 'Iliahi and host seedlings were planted at a distance of <0.2, 0.5, 1.0, or 2.0 m from each other. After three years, survival of 'iliahi seedlings was greatest for 'iliahi paired with koa at 1.0 m (88 %) compared to the control with no host (53 %) and when paired with koa at 2.0 m (55 %). Height and ground line diameter of 'iliahi seedlings were greater when paired with koa at closer distances than with 'a'ali'i at any distance or the control with no host. Specifically, 'iliahi growth was greatest when paired with koa at <0.2 m distance. Foliar nitrogen concentration was greater for 'iliahi paired with koa at <0.2, 0.5, and 1.0 m distances, whereas the other foliar nutrient concentrations were typically greater for the 'iliahi control with no host. For sites with few or no pre-established hosts, such as abandoned pastures, 'iliahi growth can be greatly improved by co-planting in proximity to koa. As with other Santalum spp., early and abundant parasitic root connections with a host, especially a nitrogen-fixing host, are likely important for 'iliahi establishment and early growth.

Herbivory is a common forest regeneration challenge across ecosystems. While fencing effectively reduces animal browse damage, it can be impractical. Tree shelters are an alternative forest restoration tool to protect seedlings from herbivory that may also provide a favorable microclimate. Yet, regeneration responses to tree shelters often vary among tree species, environmental conditions, and shelter specifications. To restore the once dominant Quercus virginiana (live oak) and its associated conservation values to subtropical U.S. maritime forests, control of animal browsing is critical. We evaluated the effects of tube and mesh tree shelters to exclude animal browse, combined with the use of controlled-release fertilizer to promote Q. virginiana seedling growth. After two growing seasons, mean seedling survival was 83% for protected seedlings, either from tube or mesh shelters, compared to 68% with non-sheltered seedlings. Seedlings in solid-wall tube shelters had significantly less browse incidence compared to both seedlings in mesh shelters and non-sheltered seedlings. Seedlings in tube shelters had greater height and diameter, followed by mesh shelters, and lastly, the no shelter treatment. Fertilizer resulted in higher browsing incidence and greater seedling height and diameter after the first growing season only, with no shelter treatment interactions. Our findings illustrate the efficacy of tree shelters to improve early regeneration success of Q. virginiana and may have application to the restoration of other forest ecosystems.

Panax quinquefolius, American ginseng, is one of the most valuable non-timber forest products providing provisioning and cultural ecosystem services in eastern temperate forests. Although ginseng has a broad distribution range, populations are declining due to several factors including overharvesting and habitat degradation. We designed experiments to study the effects of soil and aspect on ginseng to better identify potential reintroduction sites. We hypothesized soil and aspect would have a significant effect on ginseng performance. Performance included growth (leaf area, height, number of prongs) and reproduction (inflorescence presence). We predicted ginseng would have greatest performance in calcium-rich loam soil, on north-facing slopes. After four growing seasons, we found only soil had a significant effect on growth. This was a consistent result over four growing seasons. On average, ginseng grown in limed loam soil were 55-mm taller with 20,000 mm(2) more leaf area and more prongs than ginseng grown in soils lower in calcium. Aspect did not affect any measured variables. Given these results, we suggest identifying soils with higher levels of calcium regardless of aspect as potential ginseng habitat. This experimental study contributes to a more complete understanding of the ecology of ginseng. Further, these results can be incorporated into developing strategies for restoration and management of this valuable species.

Quercus spp. (oaks) are generally intermediate in shade tolerance, yet there is large variation within the genus in shade tolerance and plasticity in response to varying resource availability. Ecophysiological knowledge specific to semi-evergreen Quercus spp. from subtropical maritime forests is lacking relative to temperate deciduous oaks. We studied the influence of light availability and plant competition on leaf physiology and performance of semi-evergreen Quercus virginiana on a barrier island along the US southern Atlantic coast. Seedlings were underplanted in pine (Pinus taeda) plantation stands with varying overstory density (clear-cut, heavy thin, light thin, and non-thinned; creating a gradient of understory light availability) and vegetation (no competition removal or herbaceous competition removal) treatments. After 2 years, seedling survival was higher with increasing light availability (clear-cut = heavy thin > light thin > non-thinned). Seedling growth (i.e., diameter, height, and crown width) increased similarly with increasing thinning intensity, while vegetation control was mainly beneficial to seedling growth in clear-cuts. These responses were partially explained by foliar nitrogen and leaf trait measurements, which followed the same pattern. Q. virginiana seedlings demonstrated high plasticity in their ability to acclimate to varying resource availability, as indicated by light response curves, specific leaf area, stomatal density, stomatal pore index, and maximum theoretical stomatal conductance. Light compensation and saturation points illustrated seedling capacity to increase net CO2 assimilation with increased light availability. Leaves on trees in the high light environment had the highest net CO2 assimilation, stomatal density, stomatal pore index, maximum theoretical stomatal conductance, and lowest specific leaf area. Although we demonstrated the relative shade tolerance of Q. virginiana in lower light environments (i.e., heavy and light thin plots), this semi-evergreen species shows high plasticity in capacity to respond to varying resource availability, similar to other Quercus spp. from mesic and Mediterranean environments.

Herbivory and competition during the regeneration phase influence forest successional dynamics. We demonstrated the importance of using the Target Plant Concept to identify and overcome site limiting factors for subtropical maritime forest restoration associated with deer browsing and competition. Quercus virginiana Mill. (live oak) bareroot seedlings were planted into clearcuts along the US Southern Atlantic coast with different treatment combinations of herbivory control (fenced or non-fenced) against white-tailed deer (Odocoileus virginianus Zimm.) browsing and competing vegetation removal (none, one-year, or two-years). After three growing seasons, mean seedling survival was 61% with no significant treatment differences. Control of browse and vegetation interacted to facilitate growth of live oak; seedlings were significantly larger for all response parameters (diameter, height, crown width) when fenced and treated with vegetation control. Removal of vegetation improved seedling performance only in fenced plots, however, indicating a shift in pressure from herbivory to competition as the most limiting site factor when deer were excluded. After the second growing season, foliar nitrogen was greater in fenced plots than non-fenced plots and greater in two-year vegetation control subplots than non-vegetation control subplots. This result, however, was absent after the third growing season. Three years after clearcutting, there was no evidence of Q. virginiana natural regeneration in non-fenced plots. Even with artificial regeneration in non-fenced plots, Q. virginiana growth was slow, indicating that herbivory was a key limiting factor. Our findings illustrate the importance of accounting for site limiting factors and may aid in developing management prescriptions to promote semi-evergreen oak regeneration in ecosystems with high pressure from herbivory and competing vegetation.