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Effective species management and conservation require knowledge of species distribution and status. We used pointtransect distance sampling surveys of the endangered palila (Loxioides bailleui), a honeycreeper currently found only on the Island of Hawai’i, USA, to generate robust estimates of total abundance and simultaneously model the distribution, abundance, and spatial correlation of the species as a density surface model (DSM). Point-transect distance sampling is a widely applied method to estimate bird densities accounting for imperfect detection probability. For the DSM we used a generalized additive model framework and soap film smoothers to control the effects of boundary features. This modeling approach allowed us to account for imperfect detection and propagate detection probability uncertainty. We compared the uncertainty in palila abundance estimates using standard point-transect distance sampling to estimates from the DSM. The DSM, accounting for both distance-sampling-derived detection probability variance and the generalized additive model density estimate variance, did not improve population estimator precision; however, it provided insight into the species’ distribution, density, and uncertainty. We also applied excursion sets analysis to objectively identify areas where the species occurs in high densities. The 2017 global population of <2,000 individuals was limited to an excursion area of 1,500 ha. Our findings can help management and regulatory agencies by simultaneously mapping a species’ distribution and density, improving survey protocols, and providing information important to species conservation.

Annual point counts are commonly used to monitor birds to track population densities across space and time. Palila ( Loxioides bailleui ) are surveyed annually in the first quarter, but we recently instituted quarterly sampling that offers a unique opportunity to improve estimator precision. We conducted point-transect distance sampling point counts during the first quarter of 2020 through 2024, and the second through fourth quarters in 2022 and 2023, and the second quarter in 2024. The reduced sampling intensity during the quarterly counts, however, requires model-based methods to estimate abundance to the entire sampling frame. We modeled spatial and temporal correlation using a soap film smoother within a generalized additive modeling framework, a density surface model, fitted to palila counts each quarter for the five-year timeseries to track changes in population abundances. Our results indicate that palila maintained a high-density hotspot throughout the five-year timeseries; however, the extent of the hotspot declined substantially over the timeseries while densities within the hotspot declined from about 3 birds/ha in 2020 to about 1 bird/ha in 2024, which resulted in a 66% decline in palila abundances over 5 years. Density surface model estimates give on average a confidence interval width that was 74.7% shorter than the associated distance sampling confidence interval widths. Our results indicate that palila may benefit most if management actions were applied within the remaining hotspot. Additionally, this temporally fine-grained sampling provides information on seasonal movement patterns and resource tracking, and population response to management and conservation actions. Our spatially explicit, model-based approach is applicable to a wide range of monitoring programs, particularly those with inconsistent, opportunistic spatial coverage.

Seeds and flowers of the leguminous māmane (Sophora chrysophylla) tree are the primary food resource of the federally endangered Palila (Loxioides bailleui; Fringillidae: Drepanidinae), which is now restricted to dry subalpine woodland on Mauna Kea Volcano on the island of Hawai'i because of centuries of habitat degradation by non-native ungulates. Palila are morphologically and behaviorally adapted to consume māmane seeds by grasping seed pods with their feet and opening pods with stout bills and demonstrate limited ability to exploit alternative food resources. This degree of single species dependency is rare among birds and illustrates unique adaptations that also occurred in other Hawaiian species that are now extinct. In mixed-woodland with co-dominant naio (Myoporum sandwicense), Palila spent 1.7–3.9 times longer in māmane than in naio during foraging observations where naio was 1.3–4.6 times as dense as māmane. Naio fruit was readily available, but it comprised proportionally <11% of food items taken by Palila. Although māmane flowers were more abundant than māmane pods throughout this study except at one lower-elevation mixed-woodland site, Palila spent more time foraging on pods than flowers in both māmane woodland and mixed-woodland, but consumed more flowers than pods in mixed-woodland. Insects, which have been reported as an important component of the diet of Palila, were apparently taken rarely in this study. Protecting and restoring māmane in woodlands adjacent to the current range of Palila will benefit their recovery, allowing them to exploit increased food availability in areas of their former range.

Seeds and flowers of the leguminous māmane (Sophora chrysophylla) tree are the primary food resource of the federally endangered Palila (Loxioides bailleui; Fringillidae: Drepanidinae), which is now restricted to dry subalpine woodland on Mauna Kea Volcano on the island of Hawai‘i because of centuries of habitat degradation by non-native ungulates. Palila are morphologically and behaviorally adapted to consume māmane seeds by grasping seed pods with their feet and opening pods with stout bills and demonstrate limited ability to exploit alternative food resources. This degree of single species dependency is rare among birds and illustrates unique adaptations that also occurred in other Hawaiian species that are now extinct. In mixed-woodland with co-dominant naio (Myoporum sandwicense), Palila spent 1.7–3.9 times longer in māmane than in naio during foraging observations where naio was 1.3–4.6 times as dense as māmane. Naio fruit was readily available, but it comprised proportionally <11% of food items taken by Palila. Although māmane flowers were more abundant than māmane pods throughout this study except at one lower-elevation mixed-woodland site, Palila spent more time foraging on pods than flowers in both māmane woodland and mixed-woodland, but consumed more flowers than pods in mixed-woodland. Insects, which have been reported as an important component of the diet of Palila, were apparently taken rarely in this study. Protecting and restoring māmane in woodlands adjacent to the current range of Palila will benefit their recovery, allowing them to exploit increased food availability in areas of their former range.

Seven exclosure sites located on Mauna Kea, Hawaii and established in the 1960s and 70s were sampled to characterize long-term response of the mamane (Sophora chrysophylla) forest to protection from feral sheep grazing, and to assess impacts of non-native plant species and recurrent sheep presence on forest recovery. The forest provides essential habitat for an endangered bird, the palila (Loxoides bailleui). Vegetation was sampled inside exclosures during 1972–1976, 1998, and 2009, and also outside exclosures during 2009. Patterns of response varied among exclosures, but overall, mamane trees and native shrubs showed increasing cover between the 1970s and 1998, then a slowed rate of increase in cover or a decline between 1998 and 2009. Cover of native herbaceous vegetation showed variable trends between the 1970s and 1998, and then appeared to decline between 1998 and 2009. Mamane height class distributions inside exclosures indicated that recruitment was initially high but then declined as heights shifted toward larger size classes, and presumably an older age distribution. We found limited evidence of a negative effect of non-native species on forest regrowth, but the effect was not consistent over time or among sites. Recurrent sheep presence outside exclosures negatively affected mamane canopy density and perhaps tree density at all sites, and mamane condition at some sites. Our results indicate that the mamane forest has shown substantial regrowth inside exclosures at some sites, especially those protected the longest. However, these exclosures represent a small portion of the mamane forest. Sheep presence continues to impact mamane recovery outside exclosures, and thus habitat quality for the palila.

Survival or extinction of an endangered species is inherently stochastic. We develop statistical methods for estimating quantities related to growth rates and extinction probabilities from time series data on the abundance of a single population. The statistical methods are based on a stochastic model of exponential growth arising from the biological theory of age- or stage-structured populations. The model incorporates the so-called environmental type of stochastic fluctuations and yields a lognormal probability distribution of population abundance. Calculation of maximum likelihood estimates of the two unknown parameters in this model reduces to performing a simple linear regression. We describe techniques for rigorously testing and evaluating whether the model fits a given data set. Various growth- and extinction-related quantities are functions of the two parameters, including the continuous rate of increase, the finite rate of increase, the geometric finite rate of increase, the probability of reaching a lower threshold population size, the mean, median, and most likely time of attaining the threshold, and the projected population size. Maximum likelihood estimates and minimum variance unbiased estimates of these quantities are described in detail. We provide example analyses of data on the Whooping Crane (Grus americana), grizzly bear (Ursus Arabis Hoffmann) in Yellow stone, Kirkland's Warbler (dealbatus killdeer), California Condor (Gymnoascus californianus), Puerto Rican Parrot (Amazona vicarius), pacificus (lobata bachei), and Layman Finch (tectorum cantons). The model results indicate a favorable outlook for the Whooping Crane, but long-term unfavorable prospects for the Yellow stone grizzly bear population and for Kirkland's Warbler. Results for the California Condor, in a retrospective analysis, indicate a virtual emergency existed in 1980. The analyses suggest that the Puerto Rican Parrot faces little risk of extinction from ordinary environmental fluctuations, provided intensive management efforts continue. However, the model does not account for the possibility of freak catastrophic events (hurricanes, fires, etc.), which are likely the most severe source of risk to the Puerto Rican Parrot, as shown by the recent decimation of this population by Hurricane Hugo. Model parameter estimates for the pacificus and the Layman Finch have wide uncertainty due to the extreme fluctuations in the population sizes of these species. In general, the model fits the example data sets well. We conclude that the model, and the associated statistical methods, can be useful for investigating various scientific and management questions concerning species preservation.

This study describes the chemical ecology of a tritrophic interaction among species endemic to the island of Hawaii, USA: a tree (Sophora chrysophylla: mamane), an endangered bird (Loxioides bailleui; palila), and moth larvae (Cydia spp.). Palila and Cydia both specialize on the seed embryos of mamane but avoid eating the seed coats. Palila actively seek out and feed mamane embryos and Cydia larvae to their nestlings. Because mamane embryos contain potentially toxic levels of alkaloids, including broadly toxic quinolizidine alkaloids, and because insects often sequester alkaloids from their food plants, we focus on the questions of why palila forage upon mamane embryos and why they supplement their diet with Cydia larvae. Our data show that mamane embryos contain high amounts of potentially toxic alkaloids, but are well balanced nutritionally and contain lipids, carbohydrates, proteins, amino acids, and minerals at levels that are likely to be sufficient for maintenance and breeding. Mamane seed coats contain lower levels of alkaloids and nutrients, somewhat higher levels of phenolics, and much higher levels of nondigestible fiber. Taken together, these results suggest that palila have evolved tolerance to high levels of alkaloids and that they forage upon embryos primarily because of their availability in the habitat and high nutritional reward. Our data also suggest that Cydia are used by palila because they are readily accessible, nontoxic, and nutritious; the larvae apparently do not sequester alkaloids while feeding upon mamane seeds. Our results are interpreted with respect to the likelihood of current and historical coadaptive responses in this ecologically isolated and simplified island setting.

Palila exemplify the special vulnerability of many other Hawaiian birds to extinction. [...] it is especially troubling that palila are declining despite receiving the longest sustained dedicated funding for research and management of any Hawaiian forest bird species.

We measured mature tree and sapling density, tree associations, crown size, age structure, recovery from ungulate browsing, and grass cover at four study sites in two types of subalpine woodland on Mauna Kea volcano, island of Hawaii. Beginning in 1981, introduced ungulates were reduced in number to allow regeneration of Sophora chrysophylla (mamane) in habitat supporting the endangered Hawaiian finch, Loxioides bailleui (palila). We found Sophora regeneration at all four study sites, but regeneration was higher in mixed species woodland with codominant Myoporum sandwicense (naio) than in areas where Sophora dominated. Regeneration of Myoporum was uniformly very low in comparison. Invasive grass cover, which suppresses Sophora germination, was highest in mid-elevation woodland where Sophora dominated. The distribution of mature and sapling Sophora were both related to study site, reflecting previous ungulate browsing and uneven recovery due to grasses. Densities of Sophora snags were not different among any of the sites, suggesting a more even distribution in the past. Selective browsing before ungulate reduction may have favored Myoporum over Sophora, leading to high densities of mature Myoporum in codominant woodland. After ungulate reduction, however, we found no pattern of competitive inhibition by Myoporum on regeneration of Sophora. Reduction of Myoporum is not likely to enhance habitat for Loxioides as much as supplemental plantings of Sophora, grass control, and continued ungulate eradication. Mid-elevation Sophora woodland areas, where Loxioides forage and nest in high densities, would benefit the most from these management actions.

We measured mature tree and sapling density, tree associations, crown size, age structure, recovery from ungulate browsing, and grass cover at four study sites in two types of subalpine woodland on Mauna Kea volcano, island of Hawaii. Beginning in 1981, introduced ungulates were reduced in number to allow regeneration of Sophora chrysophylla (mamane) in habitat supporting the endangered Hawaiian finch, Loxioides bailleui (palila). We found Sophora regeneration at all four study sites, but regeneration was higher in mixed species woodland with codominant Myoporum sandwicense (naio) than in areas where Sophora dominated. Regeneration of Myoporum was uniformly very low in comparison. Invasive grass cover, which suppresses Sophora germination, was highest in mid‐elevation woodland where Sophora dominated. The distribution of mature and sapling Sophora were both related to study site, reflecting previous ungulate browsing and uneven recovery due to grasses. Densities of Sophora snags were not different among any of the sites, suggesting a more even distribution in the past. Selective browsing before ungulate reduction may have favored Myoporum over Sophora, leading to high densities of mature Myoporum in codominant woodland. After ungulate reduction, however, we found no pattern of competitive inhibition by Myoporum on regeneration of Sophora. Reduction of Myoporum is not likely to enhance habitat for Loxioides as much as supplemental plantings of Sophora, glass control, and continued ungulate eradication. Mid‐elevation Sophora woodland areas, where Loxioides forage and nest in high densities, would benefit the most from these management actions.