Hakalau Forest National Wildlife Refuge Research Articles

Hawaiian Endemic Honeycreepers (Drepanidinae) are Nectar Robbers of the Invasive Banana Poka (Passiflora tarminiana, Passifloraceae)

The human transport and subsequent naturalization of species outside their natural ranges has led to novel interactions between introduced and native species throughout the world. Understanding how introduced species impact pollination networks is useful for both invasive species management and native species conservation and restoration. Banana poka (Passiflora tarminiana), a hummingbird pollinated liana native to South America, has naturalized in higher elevation forests on the islands of Kauaʻi, Maui and Hawaiʻi in the Hawaiian archipelago, habitats in which endemic honeycreepers still occur. To develop an understanding of the interaction between banana poka and honeycreepers, we undertook a floral visitation study at Hakalau Forest National Wildlife Refuge on the island of Hawaiʻi where three nectivorous honeycreepers and banana poka co-occur. Two honeycreeper species, ʻiʻiwi (Drepanis coccinea) and Hawaiʻi ʻamakihi (Chlorodrepanis virens), nectar robbed all of the banana poka flowers that they visited, ostensibly due to the length of the corolla tubes (60–90 mm long) which physically inhibits both honeycreeper species from accessing nectar via the mouth of the corolla. In addition, the standing crop and sugar composition of banana poka floral nectar were assessed. Flowers produced large standing crops (375 ± 132 μL) of nectar containing 29.1 ± 1% (w/v) of sugar that was sucrose-dominant (mean: 95.6 ± 0.5% sucrose in each sample). Our observations suggest that the floral nectar of banana poka may form a substantial component of the diet of both honeycreeper species at the study site. Further research is needed to understand how infestations of banana poka affect bird pollination networks at this and other sites in Hawaiʻi.

Hypotheses and lessons from a native moth outbreak in a low‐diversity, tropical rainforest

AbstractOutbreaks of defoliating insects in low‐diversity tropical forests occur infrequently but provide valuable insights about outbreak ecology in temperate environments and in general. We investigated an extensive outbreak of the endemic koa moth (Scotorythra paludicola), which defoliated endemic koa trees (Acacia koa) over a third of their range on Hawai‘i Island during 2013 and 2014. At Hakalau Forest National Wildlife Refuge, we observed the dynamics of the outbreak and its effects on host trees, nutrient cycling, and insectivorous consumers in reforestation stands of densely planted koa and in natural forest stands of mixed koa and ‘ōhi‘a (Metrosideros polymorpha). Contrary to predictions of the resource concentration hypothesis, caterpillar biomass and defoliation severity were greater in the natural forest sites, where koa density was relatively low. Caterpillars preferentially consumed the most palatable koa foliage type (phyllodes), and koa initially refoliated with the least palatable foliage type (true leaves). Lightly defoliated small trees refoliated more quickly than did heavily defoliated ones but the opposite was true for large trees, which also produced a greater proportion of phyllodes. Mortality was greatest for heavily defoliated small koa. Caterpillar frass caused larger increases in soil nitrogen (N) than phosphorus (P) availability, with the greatest N increases in fine‐textured soils. Foliar N increased in alien grasses under koa canopies compared to grasses away from koa and to native woody understory species. Bird activity was influenced by ‘ōhi‘a flower abundance and the severity of koa defoliation; birds switched to outbreaking caterpillar prey, and they gained weight during the outbreak. Bat foraging times decreased during the outbreak, apparently because they became satiated quickly each night. Parasitoid wasps increased with caterpillar abundance but had little influence on outbreak dynamics. Reducing alien grass cover and increasing tree diversity would likely reduce the impacts of insect outbreaks and similar perturbations to native forests.

Fostering real-time climate adaptation: Analyzing past, current, and forecast temperature to understand the dynamic risk to Hawaiian honeycreepers from avian malaria

Various vector control options are increasingly being considered to safeguard forest birds in their natural habitats from avian malaria transmission. However, vector control options require localized deployment that is not logistically, ethically, ecologically, nor economically viable everywhere and all the time. Based on thermal tolerances of the sporogonic stages of avian malaria (Plasmodium relictum) parasite and its vector, the southern house mosquito (Culex quinquefasciatus), we examined the long-term weather trends for three high value, forest bird refuges (Alakai Wilderness Preserve on Kaua’i, Hanawi Natural Area Reserve on Maui, and Hakalau Forest National Wildlife Refuge on Hawai’i Island) to understand the temporal and site-specific differences of temperature-driven suitability for localized avian malaria transmission.On average, Alakai had mean ambient temperatures suitable for both the vector’s immature stage development and parasite sporogonic development most of the time (85.3%), indicating that observed variability in vector abundance or disease transmission may be driven by other factors. At higher elevation sites like Hakalau and Hanawi, current mean ambient temperatures suitable only for vector development prevail (91.7% and 96.6%, respectively), while mean ambient temperatures for both vector and parasite sporogonic development seldom occur (4.4% and 0% respectively). Our results not only show differences in the temperature suitability for transmission across elevation, but also different levels of vulnerability to avian malaria transmission with any additional projected increase in temperature. For instance, under a conservative warming scenario of 1.0 °C, the joint temperature suitability of parasite and vector development increases at higher elevation sites such as Hakalau (+35.8%) and Hanawi (+15.4%). While mean ambient temperatures suitable for both vector and parasite development already occur most of the time at Alakai, the occurrence also increases (+8.4%) as well under this conservative warming scenario.By linking current site-specific weather data to real-time weather forecasts, we developed a real-time avian malaria warning system to assist managers in identifying conditions when vector control is most needed at these three selected study sites. This online tool determines when conditions are likely to be suitable for local development of P. relictum and C. quinquefasciatus at Alakai, Hanawi, and Hakalau. This tool illustrates how managers can incorporate climate and current weather patterns into decision making without having to consider the uncertainties of long-term climatic and ecological projections.

Forest regeneration following ungulate removal in a montane Hawaiian wet forest

Most Hawaiian forests lack resiliency following disturbance due to the presence of non‐native and invasive plant and animal species. The montane wet forest within Hakalau Forest National Wildlife Refuge on Hawai'i island has a long history of ungulate disturbance but portions of the refuge were fenced and most ungulates excluded by the early 1990s. We examined patterns of regeneration within two 100 ha study sites in this forest following the removal of ungulates and in the absence of invasive woody tree species to determine, in part, if passive restoration techniques can be successful under these conditions. We characterized growth, mortality, and basal area (BA) changes for approximately 7,100 marked individuals of all native tree species present in two surveys over a 17–18‐year period within two hundred 30 m diameter forest plots. Considerable recruitment within plots of new trees of all species significantly changed size class distributions and erased deficits in small‐sized trees observed during the first survey, particularly for the codominant canopy tree, koa (Acacia koa). Overall, growth of established dominant 'ōhi'a trees (Metrosideros polymorpha) and recruitment of mid‐canopy trees contributed to increases in BA while high levels of mortality for largeA.koatrees contributed to decreased BA. This resulted in a slight increase in BA between the two surveys (+1.9%). This study demonstrates that fencing and ungulate removal may have rescued theA.koapopulation by facilitating the first real pulse in recruitment in over a century, and that passive restoration can be a successful management strategy in this forest.

Evaluating community-level response to management actions across a diverse Hawaiian forest bird community.

Although species-specific approaches are necessary to understand the dynamics of individual species composing a community, they do not offer a framework for making optimal management decisions at the community level. Here, we present a simple framework for comparing the response of entire communities to multiple management scenarios. Our approach uses a weighted average of standardized species-specific responses to produce a single integrative measure of the community response and employs mixed-effect linear models to quantify the increase in the community response due to each management action, or combination of actions. We demonstrate our approach with a simulation study assessing the potential benefits of multiple management actions on the avian community of Hakalau Forest National Wildlife Refuge, Hawai'i, which is composed of eight native species with conservation status varying from endangered to least concern. Management actions considered included a reduction of avian malaria transmission risk, reducing rat predation, and increasing forest habitat. We used three different prioritization (weighting) schemes to assess whether the response to management actions differed along a conservation gradient (from endangered to least concern). We also investigated whether future changes in disease distribution as a result of climate change will alter the relationship between management practices and community response. Our community-level analysis produced three important insights, highlighting the need to consider the response from multiple species to changing threats and management actions. First, increasing the amount of habitat always had the greatest positive impact on the avian community, regardless of the weighting scheme. Second, the community response to management was different under current vs. future conditions, with increased benefit in the future when disease risks were higher. Third, the response to management varied along the conservation gradient. Reducing malaria transmission risk in 2100 had the greatest benefit to endangered species, while increasing forest habitat had the greatest benefit to "least concern" species. However, reducing rat predation appeared to benefit a subset of species based on ecological factors unrelated to rarity, such as nest accessibility. Our approach is widely applicable using experimental, observational, or simulation-based data, allowing managers to consider the response of all species while weighting their priority levels.

Seasonality and Prevalence of Pollen Collected from Hawaiian Nectarivorous Birds1

Hawaiian nectarivorous forest birds play a vital ecological role as pollinators in Hawaiian ecosystems. However, little is known about what nectar resources are utilized by Hawai‘i's nectarivorous birds, how seasonality influences nectar availability, and how nectar preference differs by bird species. We sampled pollen from the heads of ‘i‘iwi (Drepanis coccinea), ‘apapane (Himatione sanguinea), Hawai‘i ‘amakihi (Chlorodrepanis virens), and the non-native Japanese white-eye (Zosterops japonicas) captured at Hakalau Forest National Wildlife Refuge on Hawai‘i Island. ‘Ōhi‘a (Metrosideros polymorpha) was the most prevalent pollen species, observed throughout the sampling period while other pollen species were more seasonal in occurrence. Consistent with the peak flowering phenology of the plant species, pollen from koa (Acacia koa), māmane (Sophora chrysophylla), and gorse (Ulex europaeus) plant speciesweremore commonly sampled frombirds in the winter months, while ‘ōhelo (Vaccinium reticulatum), ‘ākala (Rubus hawaiensis), and blackberry (Rubus argutus)were more prevalent during the spring months.We also found an association between bird species and pollen resources, with ‘i‘iwi and Hawai‘i ‘amakihi having a higher diversity of pollen than ‘apapane and Japanese white-eye, which primarily had just ‘ōhi‘a. These results demonstrate that ‘ōhi‘a is likely the most important nectar resource for Hawai‘i's nectar feeding birds, but seasonally abundant nectar may be important for some species.

Predicting effects of large‐scale reforestation on native and exotic birds

AbstractAimEcological restoration is critical for recovering biodiversity and ecosystem services, yet designing interventions to achieve particular outcomes remains fraught with challenges. In the extensive regions where non‐native species are firmly established, it is unlikely that historical conditions can be fully reinstated. To what degree, and how rapidly, can human‐dominated areas be shifted via restoration into regimes that benefit target species, communities or processes?LocationWe explore this question in a >20‐year‐old reforestation effort underway at Hakalau Forest National Wildlife Refuge in montane Hawaii. This large‐scale planting of Acacia koa trees is designed to secure populations of globally threatened bird species by transitioning the site rapidly from pasture to native forest.MethodsWe surveyed all forest birds in multiple corridors of young planted trees, remnant corridors of mature trees along gulches and at sites within mature forest. Using a Bayesian hierarchical approach, we identified which factors (distance from forest, habitat type and surrounding tree cover) had the most important influence on native and exotic bird abundance in the reforestation area.ResultsWe found that 90% of native and exotic bird species responded quickly, occupying corridors of native trees approximately a decade after planting. However, native and exotic forest birds responded to markedly different characteristics of the reforested area. Native bird abundance was strongly predicted by proximity to mature forest and remnant corridors; conversely, exotic bird abundance was best predicted by overall tree cover throughout the area reforested.Main conclusionsOur results demonstrate that large‐scale tree planting in corridors adjacent to mature forest can catalyse rapid recovery (both increased abundance and expanded distribution) of forest birds and that it is possible to design reforestation to benefit native species in novel ecosystems.

No evidence of critical slowing down in two endangered Hawaiian honeycreepers.

There is debate about the current population trends and predicted short-term fates of the endangered forest birds, Hawai`i Creeper (Loxops mana) and Hawai`i `Ākepa (L. coccineus). Using long-term population size estimates, some studies report forest bird populations as stable or increasing, while other studies report signs of population decline or impending extinction associated with introduced Japanese White-eye (Zosterops japonicus) increase. Reliable predictors of impending population collapse, well before the collapse begins, have been reported in simulations and microcosm experiments. In these studies, statistical indicators of critical slowing down, a phenomenon characterized by longer recovery rates after population size perturbation, are reported to be early warning signals of an impending regime shift observable prior to the tipping point. While the conservation applications of these metrics are commonly discussed, early warning signal detection methods are rarely applied to population size data from natural populations, so their efficacy and utility in species management remain unclear. We evaluated two time series of state-space abundance estimates (1987–2012) from Hakalau Forest National Wildlife Refuge, Hawai`i to test for evidence of early warning signals of impending population collapse for the Hawai`i Creeper and Hawai`i `Ākepa. We looked for signals throughout the time series, and prior to 2000, when white-eye abundance began increasing. We found no evidence for either species of increasing variance, autocorrelation, or skewness, which are commonly reported early warning signals. We calculated linear rather than ordinary skewness because the latter is biased, particularly for small sample sizes. Furthermore, we identified break-points in trends over time for both endangered species, indicating shifts in slopes away from strongly increasing trends, but they were only weakly supported by Bayesian change-point analyses (i.e., no step-wise changes in abundance). The break-point and change-point test results, in addition to the early warning signal analyses, support that the two populations do not appear to show signs of critical slowing down or decline.

Scaling Up Restoration Efforts in the Pacific Islands: A Call for Clear Management Objectives, Targeted Research to Minimize Uncertainty, and Innovative Solutions to a Wicked Problem

Today, restoration of resilient native ecosystems that minimize extinction risk and maximize ecosystem services has never been more important. Success stories, such as that of Hakalau Forest National Wildlife Refuge and papers in this special feature, demonstrate the value in large-scale efforts. However, the number of decision makers involved and siloed agencies responsible for land and nearshore coastal management, incomplete or contradictory knowledge regarding community and ecosystem dynamics, the large costs associated with complete restoration, and the interconnectedness of restoration decisions with social, cultural, policy, and economic dimensions, clearly classify the process of attempting large-scale restoration of degraded systems as a wicked problem. We wish to outline three focal areas that specifically address each of the challenges associated with wicked problems and that may improve the likelihood of achieving restoration goals. First, managers and decision makers must identify clear objectives that will guide decisions regarding restoration actions. Second, research funded through conservation initiatives should be aimed at reducing uncertainty in ways that increase the probability of choosing a set of management actions likely to have a desirable outcome. Finally, we need innovative solutions that borrow from industries that have already discovered economy of scale, as well as partnerships among those in the fields of social science, economics, policy, and the natural sciences. The collection of articles in this special feature illustrate how objective-driven studies, high-value research in areas of uncertainty, and collaborations among economists, cultural practitioners, and scientists can move us toward the identification of optimal solutions. Ultimately, we wish to restore systems in ways that reduce conservation reliance and result in resilient, self-sustaining native ecosystems.

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

AbstractDeforestation 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.

Altitudinal migration and the future of an iconic Hawaiian honeycreeper in response to climate change and management

AbstractAltitudinal movement by tropical birds to track seasonally variable resources can move them from protected areas to areas of increased vulnerability. In Hawaiʻi, historical reports suggest that many Hawaiian honeycreepers such as the ‘I‘iwi (Drepanis coccinea) once undertook seasonal migrations, but the existence of such movements today is unclear. Because Hawaiian honeycreepers are highly susceptible to avian malaria, currently minimal in high‐elevation forests, understanding the degree to which honeycreepers visit lower elevation forests may be critical to predict the current impact of malaria on population dynamics and how susceptible bird populations may respond to climate change and mitigation scenarios. Using radio telemetry data, we demonstrate for the first time that a large fraction of breeding adult and juvenile ‘I‘iwi originating from an upper‐elevation (1,920 m) population at Hakalau Forest National Wildlife Refuge exhibit post‐breeding movements well below the upper elevational limit for mosquitoes. Bloom data suggest seasonal variation in floral resources is the primary driver of seasonal movement for ‘I‘iwi. To understand the demographic implications of such movement, we developed a spatial individual‐based model calibrated using previously published and original data. ʻI‘iwi dynamics were simulated backward in time, to estimate population levels in the absence of avian malaria, and forward in time, to assess the impact of climate warming as well as two potential mitigation actions. Even in disease‐free ‘refuge’ populations, we found that breeding densities failed to reach the estimated carrying capacity, suggesting the existence of a seasonal “migration load” as a result of travel to disease‐prevalent areas. We predict that ‘I‘iwi may be on the verge of extinction in 2100, with the total number of pairs reaching only ~ 0.2–12.3% of the estimated pre‐malaria density, based on an optimistic climate change scenario. The probability of extinction of ‘I‘iwi populations, as measured by population estimates for 2100, is strongly related to their estimated migration propensity. Long‐term conservation strategies likely will require a multi‐pronged response including a reduction of malaria threats, habitat restoration and continued landscape‐level access to seasonally variable nectar resources.

Comparative demographics of a Hawaiian forest bird community

Estimates of demographic parameters such as survival and reproductive success are critical for guiding management efforts focused on species of conservation concern. Unfortunately, reliable demographic parameters are difficult to obtain for any species, but especially for rare or endangered species. Here we derived estimates of adult survival and recruitment in a community of Hawaiian forest birds, including eight native species (of which three are endangered) and two introduced species at Hakalau Forest National Wildlife Refuge, Hawaiʻi. Integrated population models (IPM) were used to link mark–recapture data (1994–1999) with long‐term population surveys (1987–2008). To our knowledge, this is the first time that IPM have been used to characterize demographic parameters of a whole avian community, and provides important insights into the life history strategies of the community. The demographic data were used to test two hypotheses: 1) arthropod specialists, such as the ‘Akiapōlā‘au Hemignathus munroi, are ‘slower’ species characterized by a greater relative contribution of adult survival to population growth, i.e. lower fecundity and increased adult survival; and 2) a species’ susceptibility to environmental change, as reflected by its conservation status, can be predicted by its life history traits. We found that all species were characterized by a similar population growth rate around one, independently of conservation status, origin (native vs non‐native), feeding guild, or life history strategy (as measured by ‘slowness’), which suggested that the community had reached an equilibrium. However, such stable dynamics were achieved differently across feeding guilds, as demonstrated by a significant increase of adult survival and a significant decrease of recruitment along a gradient of increased insectivory, in support of hypothesis 1. Supporting our second hypothesis, we found that slower species were more vulnerable species at the global scale than faster ones. The possible causes and conservation implications of these patterns are discussed.

Evaluating abundance and trends in a Hawaiian avian community using state-space analysis

SummaryEstimating population abundances and patterns of change over time are important in both ecology and conservation. Trend assessment typically entails fitting a regression to a time series of abundances to estimate population trajectory. However, changes in abundance estimates from year-to-year across time are due to both true variation in population size (process variation) and variation due to imperfect sampling and model fit. State-space models are a relatively new method that can be used to partition the error components and quantify trends based only on process variation. We compare a state-space modelling approach with a more traditional linear regression approach to assess trends in uncorrected raw counts and detection-corrected abundance estimates of forest birds at Hakalau Forest National Wildlife Refuge, Hawai‘i. Most species demonstrated similar trends using either method. In general, evidence for trends using state-space models was less strong than for linear regression, as measured by estimates of precision. However, while the state-space models may sacrifice precision, the expectation is that these estimates provide a better representation of the real world biological processes of interest because they are partitioning process variation (environmental and demographic variation) and observation variation (sampling and model variation). The state-space approach also provides annual estimates of abundance which can be used by managers to set conservation strategies, and can be linked to factors that vary by year, such as climate, to better understand processes that drive population trends.

Diffuse competition can be reversed: a case history with birds in Hawaii

Diffuse competition, based on niche overlap of an invader with most native species in a community, is considered a mechanism preventing its successful establishment in the community. However, native species must themselves have extensive niche overlap, implying that diffuse competition can be reversed if some external force increases numbers of the introduced species while native community numbers remain unchanged. Reverse diffuse competition, defined as an invader that outcompetes most of the native community, can then lower numbers of native species. We illustrate these principles by documenting diffuse competition between the introduced Japanese White‐eye (Zosterops japonicus) and eight species of Hawaiian forest birds at Hakalau Forest National Wildlife Refuge followed by reverse diffuse competition. Competition was based on extensive overlap of multiple foraging substrates with native species. The external force which reversed diffuse competition was a restoration area contiguous with much of the old‐growth forest where diffuse competition previously occurred. Propagule pressure from that area increased white‐eye numbers in the old‐growth forest, without initial change in numbers of native birds, resulting in greater than sixfold increase in adult white‐eyes on our study site. This increase resulted in stunted skeletal growth of young birds, and diverse changes in molt of both young and adults of native birds and white‐eyes. While native birds declined from stunted growth, white‐eyes increased adult survival. Changes in condition of native birds in time and space matched associated changes in white‐eye density. Normalizing natural selection of lower mass and shorter bills resulted in loss of almost half of native birds in a 3373 ha area of the refuge, adjacent to the restoration area, and 10% of native birds in a lower 1998 ha area. Competition with female native birds was initially stronger than with males as indicated by more male‐biased adult sex ratios in five species. Structure of the forest, small size of arthropods available, greater use of the understory, and lower minimum air temperature resulted in successful white‐eye competition with the entire native community. Reverse diffuse competition is a process in community ecology and a way an additional species can be packed into a community.

Hawaiian forest bird trends: Using log-linear models to assess long-term trends is supported by model diagnostics and assumptions (reply to Freed and Cann 2013)

ABSTRACT Freed and Cann (2013) criticized our use of linear models to assess trends in the status of Hawaiian forest birds through time (Camp et al. 2009a, 2009b, 2010) by questioning our sampling scheme, whether we met model assumptions, and whether we ignored short-term changes in the population time series. In the present paper, we address these concerns and reiterate that our results do not support the position of Freed and Cann (2013) that the forest birds in the Hakalau Forest National Wildlife Refuge (NWR) are declining, or that the federally listed endangered birds are showing signs of imminent collapse. On the contrary, our data indicate that the 21-year long-term trends for native birds in Hakalau Forest NWR are stable to increasing, especially in areas that have received active management.

Females lead population collapse of the endangered Hawaii creeper.

Population collapses result from drastic environmental changes, but the sexes may differ in vulnerability. Collapse of the endangered Hawaii creeper (Oreomystis mana) at Hakalau Forest National Wildlife Refuge resulted from food limitation associated with increased numbers of an introduced bird (Japanese white-eye, Zosterops japonicus), which competes with the creeper for food. Both creeper sexes had stunted bill growth and the greatest change in molt of native species in the community. With a surge in numbers of white-eyes, a recent cohort of adult females had very low survival after breeding, while adult males from the same cohort, and older females and males, continued to have high survival. Lower female survival resulted in a significantly more male-biased adult sex ratio. Recent low female survival was based on a great cost of reproduction, indicated by molt-breeding overlap that was previously avoided, and lower fat during the lengthy fledgling period. The difference in female survival between cohorts was associated with stunted bills from being reared in and then breeding in an increasingly poor food environment. Trend analysis of survey data indicate that the bird is declining throughout the refuge, with males being 72–80% of adults left six years after the white-eye increased. Competition over time was consistent with that previously documented over space on the Island of Hawaii. Adaptive management to recover the bird in this protected area needs to focus on improving both adult female survival and the adult sex ratio.

Passive restoration augments active restoration in deforested landscapes: The role of root suckering adjacent to planted stands of Acacia koa

Active forest restoration in Hawaii’s Hakalau Forest National Wildlife Refuge has produced a network of Acacia koa tree corridors and islands in deforested grasslands. Passive restoration by root suckering has potential to expand tree cover and close gaps between planted stands. This study documents rates of encroachment into grassland, clonal stand structure, and tree architecture. Data were collected from random replicate strip transects that started inside 23-year-old koa plantations and ended either in open grassland or in adjacent planted stands. For the former, sucker densities increased from near zero inside planted stands to a maximum of 5–38 stemsm−2 5–14m away from the edge of the plantation canopy, and then decreased to zero–a typical pattern for trees invading grassland. No suckers occurred more than 28m from the canopy edge on east-facing slopes, or more than 18m on south-facing slopes. Rates of expansion into grassland ranged from 0.8 to 1.5myr−1; suckers had already filled gaps between closely spaced plantation stands located on north-facing slopes. Continued suckering should result in the eventual re-establishment of tree cover on deforested areas between planted tree islands and corridors, and without additional active restoration.

More Misleading Trend Analysis of Hawaiian Forest Birds

Abstract. Accurate analysis of trends in densities estimated from survey data depends on selection of a model that fits the data. When such analysis uses data recorded at different times in the same area but from different transects, then the target species' density and spatial heterogeneity become an issue. It is possible, even in the absence of environmental change, that one set of transects may underestimate or overestimate the density entirely on the basis of location. The resulting risk is that the density estimated earlier may falsely indicate a decline or an increase when compared to the densities estimated later. We investigate this problem for trend analysis of two species of endangered Hawaiian birds at Hakalau Forest National Wildlife Refuge, island of Hawaii. Assertions based on Bayesian regression, and accepted by the U.S. Fish and Wildlife Service, purportedly indicate increases in the density of the endangered Hawaii Akepa (Loxops coccineus coccineus) from 1977 to 2007 and of the endangered...

Parent tree effects on reestablishment of Acacia koa in abandoned pasture and the influence of initial density on stand development

Increasingly private landholders in Hawaii are considering native forest restoration for their lands, and some public agencies have already started such work. Initial efforts have focused on reestablishing Acacia koa to recover alien-grass-dominated sites. This study was done in Hakalau Forest National Wildlife Refuge, Island of Hawaii, to determine the efficacy of disk plowing to stimulate natural regeneration of koa from buried seeds. Sites with four different koa parent tree configurations were treated-single live overhead koa canopy, multiple live canopies, downed snags, and no parent koa tree. Tree growth and survival were assessed periodically over 21 years. Average initial stand den- sities ranged from 100 to 1,500 trees ha -1 of scarified land, although some open areas had as few as 20 trees ha -1 . The distributions of seedlings with increasing distance from plot center were variable within and between parent tree configurations. Initial seedling density was significantly greater for the multiple-live-parent than for the no-parent configuration. Densities for the single-live and dead configurations differed from the no-parent config- uration only when densities were based on the entire scarified area of each plot. Stand densities declined 10-67 % during the next 20 years. Survival was a negative, non-linear function of initial stand density. Initial stand density exerted strong control over stem diameter and crown size at age 21-years, but had little effect on the proportion of trees with single-stems. The relationships between stand basal area and density at 21 years con- formed to the existing koa stocking guidelines. While moderate to high densities of natural regeneration can be expected from scarifying around live and dead koa trees, single trees or low density stands are likely in open areas.

Increase of an introduced bird competitor in old-growth forest associated with restoration

Many successful invasions involve long initial periods in which the invader exists at low densities followed by sudden population increases. The reasons for such time-lags remain poorly understood. Here we document a sudden increase in density of the introduced Japanese white-eye (Zosterops japonicus) in a restoration area contiguous with old-growth forest at Hakalau Forest National Wildlife Refuge on the Island of Hawaii. The refuge, with very high density of native birds, existed in a pocket of low white-eye density that persisted for at least 20 years since the late 1970s. The refuge began an extensive native trees restoration project in 1989 within a 1314 ha abandoned pasture above old-growth forest. This area was soon colonized by white-eyes and their population grew exponentially once the trees had grown tall enough to develop a canopy. This increase was in turn followed by significantly more white-eyes in the open and closed forests adjacent to the restoration area. Competition between white-eyes and native species was documented on study sites within these forests. Density data indicate that competition was more widespread, with loss of tens of thousands of native birds in the 5371 ha area surveyed. Our results are consistent with the view that ecological barriers may delay the population increase of invaders and that human-derived activities may help invaders cross these barriers by creating new ecological opportunities. Control of white-eye numbers may be essential for recovery of native species.