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Geology topics

Stephanie G. Yelenik

Publications and source records attributed to Stephanie G. Yelenik.

36 records · Page 2Linked to original sources
USGS2020

Successful restoration of Metrosideros polymorpha (ʻōhiʻa) is possible in forest sites with active Rapid ‘Ōhiʻa Death infections

Rapid ʻŌhiʻa Death (ROD), caused by the fungal pathogen Ceratocystis , is killing large numbers of ʻōhiʻa trees ( Metrosideros polymorpha ) in Hawaiʻi. ʻŌhiʻa are a dominant tree in Hawaiian forests, have a range that goes from arid to wet forest climates, and are important for endangered species habitat and ecosystem function. To test whether actively planting ʻōhiʻa seedlings is a viable restoration strategy in areas with high ROD mortality, we planted ʻōhiʻa in a ROD‐affected forest and crossed this with weeding and fencing treatments to compare ROD mortality to other stressors. We also tested for viable Ceratocystis spores in soils around planting areas. We found that seedlings were more likely to die in unweeded and unfenced treatments than controls. Although viable Ceratocystis spores were found in soil, none of the 41 dead seedlings tested positive for Ceratocystis . This indicates that competition from exotic plants and exotic feral ungulate damage are more likely to kill seedlings than ROD within the first year after planting.

Hawaii↗
USGS2020

Architecture of remnant trees influences native woody plant recruitment in abandoned Hawaiian pastures

Abandoned tropical pastures offer opportunities for passive and active restoration of native forest communities. Tree architecture of remnant canopy trees may be one important factor that can facilitate native plant recruitment in abandoned pastures but has largely been overlooked. Here, we evaluated patterns of native woody plant recruitment under remnant trees in abandoned pastures on Hawai’i Island and how these might be related to both tree architectural features and landscape variables. We measured native woody stems (excluding sprouts of the tree itself) in a 5 m radius around the base of each tree and modeled total basal area of native stems as a function of tree architectural characteristics. Recruitment was positively correlated with tree height as well as horizontal woody area below 1 m (tree structure that occurred below 1 m and was < 45° angle from the ground) around the base of trees. Tall trees likely attract more avian seed dispersers due to their higher visibility on the landscape and increased crown volume. Horizontal woody area likely provides establishment microsites that are above the pasture grass layer, similar to how dead or decaying logs act as nurse substrates. Unlike previous studies, we found little evidence that landscape variables such as distance to the intact forest or nearest canopy neighbor influenced understory recruitment. Tree architectural characteristics can be important predictors of native plant recruitment in abandoned tropical pastures and should be considered in addition to local and landscape-level variables.

Hawaii↗
USGS2020

Restoration at the landscape scale as a means of mitigation and adaptation to climate change

Purpose of Review Although landscape-scale restoration efforts are gaining traction worldwide, their success is generally unknown. We review landscape-scale restorations to gain insight to whether focal ecological outcomes have been achieved, in the face of changing environmental conditions. Recent Findings Only 9% of the 477 articles that resulted from our search were studies of landscape-scale restorations. The majority (73%) of the landscape restorations from our study have occurred since the 1990s, indicating that this type of restoration has gained in popularity in the last 30 years. Furthermore, 67% of these restoration studies occurred in a single country: China. Many scientific studies have addressed the ability of a species to shift ranges with climate change, yet few of the landscape-scale restoration studies used for our study addressed this question. Instead, 87% of the studies focused on ecosystem function, rather than community-level processes, as a result of restoration. Summary There is a clear need for more research to be undertaken on the ecological outcomes of landscape-scale restorations to understand whether they enable species and communities to shift their ranges or adapt to climate change. Conservation practitioners could utilize our decision matrix as a tool to guide restoration of individual sites within a landscape context, as well as current and future climatic conditions, to guide ecological outcomes of interest. Optimal biodiversity maintenance requires habitat conservation in concert with restoration activities at the landscape scale, and the latter, likely increasingly so in a world of changing climate.

Current Landscape Ecology Reports↗
USGS2020

Local to landscape-level controls of water fluxes through Hawaiian forests: Effects of invasive animals and plants on soil infiltration capacity across substrate and moisture gradients

Given the potential effect of invasive plants and animals to water fluxes through forests, the invasive-driven degradation of native ecosystems is a topic of great concern for many downstream land and water managers. The infiltration rate determines the partitioning between runoff and infiltration into soil in Hawaiian forests and beyond. Thus, to explore the ecohydrological effects of plant and animal invasion in mesic and wet forests in Hawaii, we measured soil infiltration capacity in multiple fenced (i.e., ungulate-free)/unfenced and native/invaded forest sites along moisture and substrate age gradients across the islands of Hawai‘i and Kaua‘i. We also characterized forest composition and structure and soil characteristics at these sites to assess the direct and vegetation-mediated impacts of invasive species on infiltration capacity. Infiltration capacity is highly variable across forested sites and the wider landscape. Much of this variability is determined by a complex set of soil, vegetation, and disturbance factors that affect infiltration capacity at the immediate surrounding of measurement plots. Consequently, the effect of any given factor can be masked by variability in other factors. However, by controlling for variability in soil and vegetation conditions at a local plot level, we found that the presence of invasive species in forests has complex and sometimes non-intuitive effects on infiltration. Our final models showed that invasive ungulates negatively affect soil infiltration capacity consistently across the wide moisture and substrate age gradients considered. Additionally, because several soil characteristics known to be affected by ungulates were associated with local infiltration rates (e.g., soil organic matter, bare soil cover, soil depth), the long-term secondary effects of high ungulate densities in Hawaiian forests may be higher than effects observed in this study. These results provide clear evidence for land managers that ungulate control efforts likely improve ecohydrologic function to mesic and wet forest systems critical to protecting downstream and nearshore resources and maintaining groundwater recharge. Compared to ungulate effects, the effect of invasive plants on water infiltration capacity in Hawaiian forests appeared much more complex. In general, elements of forest structure including increased canopy, understory and floor cover, greater presence of large roots, and lower grass and bare soil covers were positively associated with water infiltration. Whether native or not, a plant species’ potential to alter infiltration rates in Hawaiian forests was likely to depend on its physiognomy and how it affects forest community structure. For instance, while the cover of native dominant tree ‘ōhi‘a, Metrosideros polymorpha, was found to be positively associated with infiltration capacity (perhaps as an indicator of overall forest integrity), invasive Himalayan ginger, Hedychium gardnerianum, was also positively correlated with infiltration capacity, possibly due to preferential flow channels created by the presence of large root mats. Few studies have conducted comprehensive integrated ecological and hydrological sampling in forests of high conservation value. While we show there are large benefits to understanding how conservation efforts may help shape water fluxes, we also found that the commonly used study design for infiltration studies used here and elsewhere (i.e., adjacent paired sites) could be modified to provide more accurate effects of invasion in future studies for ecosystems in Hawaii and beyond.

Hawaii↗
USGS2019

Environmental gradients influence differences in leaf functional traits between native and non-native plants

Determining the characteristics of non-native plants that can successfully establish and spread is central to pressing questions in invasion ecology. Evidence suggests that some non-native species establish and spread in new environments because they possess characteristics (functional traits) that allow them to either successfully compete with native residents or fill previously unfilled niches. However, the relative importance of out-competing native species vs. filling empty niche space as potential mechanisms of invasion may depend on environmental characteristics. Here, we measured plant functional traits, proxies indicative of competitive and establishment strategies, to determine if these traits vary among native and invasive species and if their prevalence is dependent on environmental conditions. Using a natural environmental gradient in Hawai’i Volcanoes National Park, we evaluated how functional traits differ between native and non-native plant communities and if these differences change along an environmental gradient from hot, dry to cool, wet conditions. Functional trait differences suggested that both competition and open niche space may be important for invasion. Non-native communities tended to have traits associated with faster growth strategies such as higher specific leaf area and lower leaf thickness. However, native and non-native community traits became more dissimilar along the gradient, suggesting that non-native species may be occupying previously unfilled niche space at the hot, dry end of the gradient. We also found that most of the variation in functional trait values amongst plots was due to species turnover rather than intraspecific variation. These results highlight the role of environmental context when considering invasion mechanisms.

Hawaii↗
USGS2019

Bryophyte abundance, composition and importance to woody plant recruitment in natural and restoration forests

Restoration of tropical forests can lead to enhanced ecosystem services and increases in native biodiversity. Bryophytes may be an integral part of the forest restoration process and can serve a critical role in forest functioning. However, the recovery of bryophytes and their ability to facilitate woody plant establishment during restoration remains poorly studied, especially in the tropics. We investigate how bryophyte abundance and community composition, as well as woody plant seedling associations with bryophyte mats and other ground cover types change from under the canopy of intact forest to under trees in restoration corridor plantings in Hawaii. Restoration corridors consisted of corridors of koa (Acacia koa) trees that were planted roughly 30 years ago. Some corridors were planted around remnant ʻōhiʻa trees (Metrosideros polymorpha) that can be several hundred years old. We sampled under ʻōhiʻa in intact forest and both koa and ʻōhiʻa trees in restoration corridors. In restoration corridors, bryophyte abundance was low relative to intact forest and species diversity was a subset of that found in intact forest despite restoration corridors being several decades old. Seedlings strongly associated with bryophytes across all habitats suggesting that bryophytes may significantly enhance forest seedling establishment when present in restoration corridors. Other ground cover types like woody litter and nurse logs also had a positive association with forest seedlings but were rare in restoration corridors. Grass remained a dominant ground cover type in restoration corridors under koa and remnant ʻōhiʻa trees and only a single seedling was ever found growing in this grass. Enhancing bryophyte growth and recovery within restoration plantings through the reduction of grass cover could facilitate native plant establishment.

Hawaii↗
USGS2018

Invasive rat control is an efficient, yet insufficient, method for recovery of the critically endangered Hawaiian plant hau kuahiwi (Hibiscadelphus giffardianus)

Biological invasions of rodents and other species have been especially problematic on tropical islands. Invasive Rattus rattus consumption of Hibiscadelphus giffardianus (Malvaceae; common Hawaiian name hau kuahiwi) fruit and seeds has been hypothesized to be the most-limiting factor inhibiting the critically endangered tree, but this has not been experimentally tested, and little is known about other factors affecting seed dispersal, germination, and seedling establishment. Thus, we do not know if rat removal is sufficient to increase hau kuahiwi recruitment. This study aims to evaluate the effect of rat population control on the ability of hau kuahiwi to retain fruit and establish seedlings. We compared hau kuahiwi fruiting and seedling recruitment in a stand treated to reduce rat abundance and a neighbouring control stand. Fruit retention increased following treatment but seedling establishment did not. Although rat control improves the ability of hau kuahiwi to retain fruit, other, presently unknown inhibitors to seed dispersal, germination, and/or seedling development remain. Seed and seedling predation by other species, competition from numerous invasive plant species, unsuitable climate, and/or other factors may be primary inhibitors in the absence of rats, but we emphasize that progressive isolation of these factors at individual hau kuahiwi life stages may be necessary to identify the remaining threats to the conservation of this critically endangered plant.

PLoS ONE↗
USGS2018

Long-term impacts of exotic grazer removal on native shrub recovery, Santa Cruz Island, California

A combination of overgrazing and exotic species introduction has led to the degradation of habitats worldwide. It is often unclear whether removal of exotic ungulates will lead to the natural reestablishment of native plant communities without further management inputs. I describe here my return to sites on Santa Cruz Island, California, 12 years after initial sampling in order to gain a long-term view on native shrub reestablishment into exotic grasslands after exotic grazer removal. Santa Cruz Island was grazed by feral sheep and cattle for over a century; these exotic grazers were removed in the late 1980s and feral pigs were removed in 2005–2006. I resampled 5 sites on south-facing slopes in the Central Valley of the island to quantify native shrub cover, density, and size. Previous data suggested that one species, Eriogonum arborescens, would be able to naturally recruit in exotic grass–dominated areas. Native shrubs have shown a modest increase in cover over time, although more striking was a sharp increase in the amount of dead shrub cover and density. Recruitment events during high rainfall years probably led to the slight increase in Eriogonum cover between sampling periods. Recent drought periods, however, have probably increased mortality and otherwise slowed shrub reestablishment in these arid sites.

California↗
USGS2017

Ecosystem vs. community recovery 25 years after grass invasions and fire in a subtropical woodland

Despite a large body of research documenting invasive plant impacts, few studies have followed individual invaded sites over decades to observe how they change, and none have contrasted how compositional impacts from invasion compare to ecosystem-process impacts over a multi-decadal time-scale. Using direct measurements of plant density and composition and of ecosystems processes, we evaluate how ecosystem structure, above-ground net primary production (ANPP), and above-ground and soil nutrient pools compare over 25 years since fire and C4 grass invasions disrupted seasonally dry Hawaiian woodlands. We compare structure and function between primary woodland that has never burned and is largely native species-dominated, with sites that had been the same woodland type but burned in alien-grass-fuelled fires in the 1970s and 1980s. The sites have not experienced fires since 1987. We report here that woody plant composition and structure continue to be dramatically changed by the initial invasions and fires that occurred 25 years ago and invaders continue to dominate in burned sites. This is reflected in continued low plant carbon pools in burned compared to unburned sites. Yet ANPP and N storage, which were dramatically lower in the initial decade after invasive-grass fuelled fires, have increased and are now indistinguishable from values measured in intact woodlands. Soil carbon pools were resilient to both invasion and fire initially and over time. Above-ground net primary production has recovered because of invasion of burned sites by a non-native N-fixing tree rather than because of recovery of native species. This invasive N-fixing tree is unlikely to return C storage of the invaded sites to those of unburned woodland because of its tissue and growth characteristics and its interactions with invasive grasses. It does not facilitate native species but rather promotes a persistent invasive grass/N-fixer savanna. Synthesis. We conclude that fire, an unusual disturbance in this system, has perpetuated the dominance of these sites by invasive species and that despite the dramatic recovery of above-ground net primary production and N pools, the ecosystem continues to be in a distinctly different state than the pre-fire, pre-Melinis community. Thus, despite the absence of further disturbance (fire), there is no evidence that succession towards the original ecosystem is occurring. The fact that N pools and above-ground net primary production recover because of a new invader (Morella faya), highlights the unpredictability of ecosystem trajectories in the face of altered regional species pools.

Hawaii↗
USGS2017

The influence of soil resources and plant traits on invasion and restoration in a subtropical woodland

It has been shown in some cases that nitrogen (N) addition to soil will increase abundance of plant invaders because many invaders have traits that promote rapid growth in response to high resource supply. Similarly, it has been suggested, and sometimes shown, that decreasing soil N via carbon (C) additions can facilitate native species recovery. Yet all species are unlikely to respond to resource supply in the same way. We asked how soil nutrients and competition affect native and exotic woody species in a restoration experiment where we added N or C, and crossed soil manipulation with the manipulation of dominant exotic grass abundance in a Hawaiian subtropical woodland. We related changes in survival and growth of outplanted individuals to native/exotic status and plant traits. As a group, N-fixers showed reduced survival compared to non-fixers in response to added N, with Morella faya (exotic) and Acacia koa (native) having dramatic negative responses. Among non-fixers, species with greater foliar %N had more positive survival responses to increasing soil N. Specific leaf area was not predictive of responses to nutrients or competition. In general, responses to carbon addition were weak, although reducing competition from existing exotic grasses was beneficial for all outplanted species, with N-fixers showing the most positive response. We conclude that commonly used restoration strategies to clear exotic species or lower soil resources with C addition will most greatly benefit N-fixing species, which themselves may be unwanted invaders. Thus statements about the influence of increased soil N on invasions should be carefully dissected by considering the traits (such as N-fixation status) of the regional species pool.

Hawai'i↗
USGS2017

Rapid colonization of a Hawaiian restoration forest by a diverse avian community

Deforestation of tropical forests has led to widespread loss and extirpation of forest bird species around the world, including the Hawaiian Islands which have experienced a dramatic loss of forests over the last 200–800 years. Given the important role birds play in forest ecosystem functions via seed dispersal and pollination, a bird community's response to forest restoration is an important measure of the success of such conservation actions. We evaluated the bird response to reforestation at an important bird sanctuary, Hakalau Forest National Wildlife Refuge, Hawai′i Island, using 26 years of bird count data. We show that most species from within the diverse avian community increased significantly, but species colonized the restoration forest at different rates. Distance from intact forest and time since restoration were both important predictors of colonization rate, interacting such that for most species it took more time to colonize areas farther from the intact forest. In addition, both forest cover and understory diversity helped to explain bird densities, but the effect varied among species, suggesting that different habitat requirements may help drive variation in colonization rates. This article provides the first detailed evaluation of how a diverse community of birds has responded to one of the largest, ongoing reforestation projects in Hawai′i.

Hawai′i↗
USGS2017

Interactions among invasive plants: Lessons from Hawai‘i

Most ecosystems have multiple-plant invaders rather than single-plant invaders, yet ecological studies and management actions focus largely on single invader species. There is a need for general principles regarding invader interactions across varying environmental conditions, so that secondary invasions can be anticipated and managers can allocate resources toward pretreatment or postremoval actions. By reviewing removal experiments conducted in three Hawaiian ecosystems (a dry tropical forest, a seasonally dry mesic forest, and a lowland wet forest), we evaluate the roles environmental harshness, priority effects, productivity potential, and species interactions have in influencing secondary invasions, defined here as invasions that are influenced either positively (facilitation) or negatively (inhibition/priority effects) by existing invaders. We generate a conceptual model with a surprise index to describe whether long-term plant invader composition and dominance is predictable or stochastic after a system perturbation such as a removal experiment. Under extremely low resource availability, the surprise index is low, whereas under intermediate-level resource environments, invader dominance is more stochastic and the surprise index is high. At high resource levels, the surprise index is intermediate: Invaders are likely abundant in the environment but their response to a perturbation is more predictable than at intermediate resource levels. We suggest further testing across environmental gradients to determine key variables that dictate the predictability of postremoval invader composition.

Hawai'i↗
USGS2017

Methods for measuring bird-mediated seed rain: Insights from a Hawaiian mesic forest

Amount and diversity of bird-dispersed seed rain play important roles in determining forest composition, yet neither is easy to quantify. The complex ecological processes that influence seed movement make the best approach highly context specific. Although recent advances in seed rain theory emphasize quantifying source-specific seed shadows, many ecological questions can be addressed u sing a less mechanistic approach that requires fewer assumptions. Using seed rain rates from 0.38 m 2 hoop traps sampled twice monthly over the course of a year, we show that number of traps required to identify changes in seed rain varies across seed species and forest type. Detecting a 50% increase in amount of seed rain required from 65 to >300 traps, while detecting a 200% increase generally required ≤⃒50 traps. Trap size and ecological context dictate the number of seeds found in each trap, but the coefficient of variation (CV) across traps in a given ecological context can help inform future studies about number of traps needed to detect change. To better understand factors influencing variation around estimates of seed rain, we simulated both clustered and evenly distributed patterns of fecal deposition using three different levels of seed aggregation (number of seeds in each fecal deposit). When patterns of fecal deposition were clustered, rather than evenly dispersed across the study area, they required >1.5 times the number of traps to identify a 100% increase in seed rain. Similarly, we found that low seed aggregation required >1.5 times the number of traps to detect a 100% change than when aggregation was medium or high. At low aggregations, fewer seed rain traps contained seeds (low, 33 ± 5%; medium, 23 ± 4%; high, 24 ± 5%), resulting in more variation across traps than medium and high aggregations. We also illustrate the importance of training observers to discern between morphologically similar seeds from different species and provide resources to help identify bird-dispersed seeds commonly found within midelevation mesic Hawaiian forests.

Hawaii↗
USGS2017

Influence of restored koa in supporting bird communities

Deforestation of Hawaiian forests has adversely impacted native wildlife, including forest birds, bats and arthropods. Restoration activities have included reforestation with the native koa (Acacia koa), a dominant canopy tree species that is easy to propagate, has high survivorship, and has fast growth rates. We review recent research describing the ecological benefits of koa restoration on wildlife colonization/use, plant dispersal, and native plant recruitment. In general, planting monotypic koa stands can provide forest habitats for species that need them but does not automatically lead to natural regeneration of a diverse forest species assemblage and may require additional restoration activities such as outplanting of other native plants and alien grass control to achieve more natural forest systems. Although early signs of forest and wildlife recovery have been encouraging, the goals of restoration for wildlife conservation versus commercial grade harvesting require different restoration methods.

Hawai'i↗
USGS2016

Linking dominant Hawaiian tree species to understory development in recovering pastures via impacts on soils and litter

Large areas of tropical forest have been cleared and planted with exotic grass species for use as cattle pasture. These often remain persistent grasslands after grazer removal, which is problematic for restoring native forest communities. It is often hoped that remnant and/or planted trees can jump-start forest succession; however, there is little mechanistic information on how different canopy species affect community trajectories. To investigate this, I surveyed understory communities, exotic grass biomass, standing litter pools, and soil properties under two dominant canopy trees— Metrosideros polymorpha (‘ōhi‘a) and Acacia koa (koa)—in recovering Hawaiian forests. I then used structural equation models (SEMs) to elucidate direct and indirect effects of trees on native understory. Native understory communities developed under ‘ōhi‘a, which had larger standing litter pools, lower soil nitrogen, and lower exotic grass biomass than koa. This pattern was variable, potentially due to historical site differences and/or distance to intact forest. Koa, in contrast, showed little understory development. Instead, data suggest that increased soil nitrogen under koa leads to high grass biomass that stalls native recruitment. SEMs suggested that indirect effects of trees via litter and soils were as or more important than direct effects for determining native cover. It is suggested that diverse plantings which incorporate species that have high carbon to nitrogen ratios may help ameliorate the negative indirect effects of koa on natural understory regeneration.

Hawai‘i↗
USGS2014

Evaluating nurse plants for restoring native woody species to degraded subtropical woodlands

Harsh habitats dominated by invasive species are difficult to restore. Invasive grasses in arid environments slow succession toward more desired composition, yet grass removal exacerbates high light and temperature, making the use of &ldquo;nurse plants&rdquo; an appealing strategy. In this study of degraded subtropical woodlands dominated by alien grasses in Hawai'i, we evaluated whether individuals of two native ( Dodonaea viscosa , Leptocophylla tameiameia ) and one non-native ( Morella faya ) woody species (1) act as natural nodes of recruitment for native woody species and (2) can be used to enhance survivorship of outplanted native woody species. To address these questions, we quantified the presence and persistence of seedlings naturally recruiting beneath adult nurse shrubs and compared survival and growth of experimentally outplanted seedlings of seven native woody species under the nurse species compared to intact and cleared alien-grass plots. We found that the two native nurse shrubs recruit their own offspring, but do not act as establishment nodes for other species. Morella faya recruited even fewer seedlings than native shrubs. Thus, outplanting will be necessary to increase abundance and diversity of native woody species. Outplant survival was the highest under shrubs compared to away from them with few differences between nurse species. The worst habitat for native seedling survival and growth was within the unmanaged invasive grass matrix. Although the two native nurse species did not differentially affect outplant survival, D. viscosa is the most widespread and easily propagated and is thus more likely to be useful as an initial nurse species. The outplanted species showed variable responses to nurse habitats that we attribute to resource requirements resulting from their typical successional stage and nitrogen fixation capability.

Hawaii↗
USGS2014

Dynamics and ecological consequences of the 2013−2014 koa moth outbreak at Hakalau Forest National Wildlife Refuge.

A massive outbreak of the koa moth (Geometridea: Scotorythra paludicola) defoliated more than a third of the koa (Acacia koa) forest on Hawai&lsquo;i Island during 2013&minus;2014. This was the largest koa moth outbreak ever recorded and the first on the island since 1953. The outbreak spread to sites distributed widely around the island between 800&minus;2,000 m elevation and in wet rainforest to dry woodland habitats. We monitored the outbreak at two windward forest sites (Laupāhoehoe and Saddle Road Kīpuka) and one leeward forest site (Kona), and we studied the dynamics of the outbreak and its impacts on the forest ecosystem at Hakalau Forest National Wildlife Refuge, our higher elevation windward site. Study sites at Hakalau included two stands of koa that were planted (reforestation stands) in former cattle pastureland about 20 years earlier and two stands of koa that were dominated by &lsquo;ōhi&lsquo;a (Metrosideros polymorpha) and that were naturally recovering from cattle grazing (forest stands). We observed one outbreak at Hakalau, multiple outbreaks at the two other windward sites, but no outbreak at the leeward site. Caterpillars at Hakalau reached peak estimated abundances of more than 250,000 per tree and 18,000,000 per hectare, and they removed between 64&minus;93% of the koa canopy in managed forest stands. Defoliation was more extensive in naturally recovering forest, where &lsquo;ōhi&lsquo;a dominated and koa was less abundant, compared to the planted stands, where koa density was high. Koa trees were still growing new foliage six months after being defoliated, and leaves were produced in greater proportion to phyllodes, especially by small koa (&le; 8 cm dbh) and by larger trees in forest stands, where light levels may have remained relatively low after defoliation due to the high cover of &lsquo;ōhi&lsquo;a. Small branches of many trees apparently died, and canopy regrowth was absent or low in 9% of koa trees and seedlings, which indicates the likely level of mortality. Between 2,000&minus;5,000 kg/ha of frass fell during the defoliation event, resulting in the deposition of up to 200 kg/ha of highly labile nitrogen on the forest floor in less than two months. The deposition of nitrogen was detected as pulses in resin-available nitrogen in the top 5&minus;10 cm of soil at two of three sites. These sites showed elevated soil nitrogen for about seven months. Nitrogen content of understory plant foliage, which is indicative of nitrogen uptake, suggested weak and variable effects of nitrogen deposition in the soil. Foliar nitrogen increased slightly in alien pasture grasses four months after the deposition of frass, although distinctive increases were not detected in native woody species. Birds responded to the abundance of caterpillars by increasing their activity in koa during the buildup of caterpillars and decreasing their use of koa after defoliation. During the outbreak, caterpillars increased in the diets of the two generalist insectivores we examined, and nearly all species gained weight. Bats responded to the abundance of moths by compression of active foraging into the first three hours of darkness each night after presumably having reached a digestive bottleneck. Reduced foraging activity by bats also resulted in lower indices of detectability based upon acoustic monitoring when compared to non-outbreak years. Parasitoid wasps tracked caterpillar abundance, but the low rate at which they attacked caterpillars suggests that they had little influence on the population. The predatory yellowjacket (Vespula pensylvanica) did not respond to the outbreak. Although a single, protracted outbreak occurred at Hakalau, multiple outbreaks and defoliations occurred at lower elevations. Our results provide a broad foundation for evaluating the dynamics and impacts of future Scotorythra outbreaks.

Hawaii↗