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David L. Leonard

Publications and source records attributed to David L. Leonard.

8 recordsLinked to original sources

Evaluating the long-term management of introduced ungulates to protect the palila, an endangered bird, and its criticial habitat in subalpine forest of Mauna Kea, Hawai'i

Under the multiple-use paradigm, conflicts may arise when protection of an endangered species must compete with other management objectives. To resolve such a conflict in the Critical Habitat of the endangered Hawaiian honeycreeper, palila (Loxioides bailleui), federal courts ordered the eradication of introduced ungulates responsible for damaging the māmane (Sophora chrysophylla) forest on which palila depend. During 1980–2011, a total of 18,130 sheep (Ovis aries and O. gmelini musimon) and 310 goats (Capra hircus) were removed from Palila Critical Habitat (PCH) primarily by public hunters (54%) and secondarily by aerial shooting. Nevertheless, our analysis indicates that ungulates have increased over time. Palila numbers have declined sharply since 2003 due to long-term habitat degradation by ungulates and drought. Although culling ungulate populations has allowed some habitat improvement, their complete removal is necessary for palila to recover, especially given the potential for continued drought. Introduced predators are being controlled to reduce palila mortality, māmane and other native trees are being planted to restore some areas, and fencing is being constructed to prevent ungulate immigration. Funds are recently available for more effective eradication efforts, which are urgently needed to eliminate browsing damage in PCH and protect the palila from extinction.

Hawaii

Collapsing avian community on a Hawaiian island

The viability of many species has been jeopardized by numerous negative factors over the centuries, but climate change is predicted to accelerate and increase the pressure of many of these threats, leading to extinctions. The Hawaiian honeycreepers, famous for their spectacular adaptive radiation, are predicted to experience negative responses to climate change, given their susceptibility to introduced disease, the strong linkage of disease distribution to climatic conditions, and their current distribution. We document the rapid collapse of the native avifauna on the island of Kaua‘i that corresponds to changes in climate and disease prevalence. Although multiple factors may be pressuring the community, we suggest that a tipping point has been crossed in which temperatures in forest habitats at high elevations have reached a threshold that facilitates the development of avian malaria and its vector throughout these species’ ranges. Continued incursion of invasive weeds and non-native avian competitors may be facilitated by climate change and could also contribute to declines. If current rates of decline continue, we predict multiple extinctions in the coming decades. Kaua‘i represents an early warning for the forest bird communities on the Maui and Hawai‘i islands, as well as other species around the world that are trapped within a climatic space that is rapidly disappearing.

Science Advances

An assessment of arthropod prey resources at Nakula Natural Area Reserve, a potential site of reintroduction for Kiwikiu (Pseudonestor xanthophrys) and Maui `Alauahio (Parareomyza montana)

Hawaiian forest birds have declined dramatically since humans arrived in the archipelago. Birds from all foraging guilds have been affected but insectivorous species are currently at greatest risk of extinction. On the island of Maui, populations and ranges of the insectivorous kiwikiu (Maui parrotbill; Pseudonestor xanthophrys) and Maui ‘alauahio (Maui creeper; Paroreomyza montana) have declined significantly from historic levels primarily due to habitat loss, predation,disease, and food web disruption, leading to federal listings of endangered species and species of concern, respectively. Recovery plans for these birds include reestablishment of populations in parts of their former range. Nakula Natural Area Reserve on the leeward side of HaleakalāVolcano has been targeted for release of wild-caught or captive-bred individuals. The mesic, montane koa-‘ōhi‘a (Acacia koa-Metrosideros polymorpha) forest at Nakula has been heavily impacted through grazing by feral ungulates, but recent management actions to exclude these animals are promoting forest recovery. The objective of this study was to assess the arthropod prey base at Nakula in preparation for reintroductions of kiwikiu and Maui ‘alauahio. To accomplish that goal, we compared arthropod abundances at Nakula to those at Hanawi Natural Area Reserve and Waikamoi Preserve, areas where kiwikiu and Maui ‘alauahio are currently found. We also identified diets of kiwikiu and Maui ‘alauahio from fecal samples to better understand and evaluate the prey base at Nakula. Assessment methods included clipping branch tips to sample arthropods within the foliage of koa and ‘ōhi‘a, using traps to quantify arthropods on koa and ‘ōhi‘a bark surfaces, counting exit holes to quantify abundances of beetles (Coleoptera) within dead branches of koa, and measuring the density of arthropods within the stems of ‘ākala (Rubus hawaiiensis). The diet of kiwikiu was dominated by caterpillars (Lepidoptera larvae), which comprised 90% of all prey items for 50 adult birds and 98% of all prey for two nestlings. Caterpillars were also the most important prey for Maui ‘alauahio (43% for 104 adult birds) although spiders (Araneae, 16%), beetles (12%) and true bugs, planthoppers and psyllids (Hemiptera; 12%) were also important. Caterpillars were generally the most abundant type of arthropod in the foliage of koa and ‘ōhi‘a, although spiders, beetles and hemipterans were also common. Total arthropod biomass and caterpillar biomass at Nakula was as great, or greater, than that observed at Hanawi and Waikamoi per unit of foliage of both koa and ‘ōhi‘a. Spiders generally dominated the bark fauna on both koa and ‘ōhi‘a at all sites although isopods (Isopoda), millipedes (Myriapoda: Millipeda) and lacewings (Neuroptera) were also abundant at Waikamoi and Hanawi. Total arthropod biomass on bark, as well as the biomass of several individual taxa, was significantly lower at Nakula than the other sites. Our measurement of the density of beetle exit holes in dead koa branches found no difference between Nakula and Waikamoi. Finally, no difference existed in the abundance of arthropods (primarily caterpillars and moth pupae) within ‘ākala stems among sites. With the exception of bark surfaces, our results suggest that the arthropod prey base for birds on primary foraging substrates at Nakula is similar to that found at two sites within the current range of kiwikiu and Maui ‘alauahio. However, our results should be viewed with caution because they are limited to the scale of individual branch, tree, or ‘ākala stem. To complete the assessment, our results should be scaled up to the landscape level by determining the density of each substrate within each site. Key arthropod prey of kiwikiu and Maui ‘alauahio are available at Nakula and, as habitat restoration continues, food abundance should increase to the point at which populations of these birds can be supported.

Hawaii

Richness, diversity, and similarity of arthropod prey consumed by a community of Hawaiian forest birds.

We evaluated the diet richness, diversity, and similarity of a community of seven endemic and two introduced passerine birds by analyzing the composition of arthropod prey in fecal samples collected during 1994–1998 at Hakalau Forest National Wildlife Refuge, Hawai‘i Island. Most prey fragments were identified to order, but we also distinguished among morpho-species of Lepidoptera based on the shape of larval (caterpillar) mandibles for higher resolution of this important prey type. Diets were compared among feeding specialists, generalists, and “intermediate” species and among introduced and three endangered Hawaiian honeycreeper (Fringillidae) species. Lepidoptera (moths), especially the larval (caterpillar) stage, comprised the greatest proportion of prey in samples of all bird species except for the introduced Japanese white-eye ( Zosterops japonicus ; JAWE). Araneae (spiders) was the most abundant order in JAWE samples and the second most abundant order for most other species. The two specialist honeycreepers ranked lowest in the richness and diversity of arthropod orders, but only the ‘akiapōlā‘au ( Hemignathus munroi , AKIP) was significantly lower than the three generalist or intermediate honeycreeper species. The diversity of arthropod orders was significantly lower for the three endangered honeycreeper species compared to the two introduced species. No significant differences were observed among the five honeycreepers with respect to the arthropod orders they consumed. The use of arthropod orders taken by endangered honeycreepers and introduced species was significantly different in all paired comparisons except for JAWE and ‘ākepa ( Loxops coccineus ; AKEP). In terms of richness and diversity of caterpillar morpho-species in the diet, only the specialist, AKEP, was significantly lower than all three generalist and intermediate species. Both AKEP and AKIP consumed a significantly different diet of caterpillar morpho-species compared to at least one honeycreeper generalist or intermediate species. Among the endangered honeycreepers and introduced species, the richness and diversity of caterpillar morpho-species was significantly lower only for AKEP compared to both introduced species. Significant differences were not observed between endangered and introduced species in the distribution of caterpillar morpho-species in the diet. Only three morpho-species were heavily exploited, with one being consumed by all bird species. The heavy exploitation of very few morpho-species by specialists underscored their greater vulnerability to changes in forest food webs and threats to key arthropod prey. When evaluated together with data on overlap in foraging behavior, our results could be useful in evaluating competition between bird species at Hakalau. Nevertheless, invasive parasitoid wasps may impact key caterpillar prey more substantially than do introduced birds, highlighting the need for additional research to understand the ecology of caterpillar species and their interactions with both invertebrate and vertebrate consumers. The severe decline of specialist bird species historically and recently is a reminder of the importance of maintaining food web resilience, potentially through vigorous habitat restoration, to withstand the continuing and perhaps increasing threats from a diverse array of invasive species and climate change.

Hawaii

Survival estimates of wild and captive-bred released Puaiohi, an endangered Hawaiian thrush

Estimating and monitoring adult and juvenile survival are vital to understanding population status, informing recovery planning for endangered species, and quantifying the success of management. We used mark&ndash;recapture models to estimate apparent annual survival of the Puaiohi ( Myadestes palmeri ), an endangered thrush endemic to the Hawaiian island of Kauai, from 2005 to 2011. Our sample included 87 wild birds and 123 captive-bred birds that were released at various ages. Survival was higher for wild adult males (0.71 &plusmn; 0.09) than for wild adult females (0.46 &plusmn; 0.12). Survival of wild juveniles (0.23 &plusmn; 0.06) was lower than that of wild adults of both sexes, indicating that recruitment may limit population growth. Captive-bred birds released when <1 yr old had survival (0.26 &plusmn; 0.21) comparable with that of wild juveniles, but captive-bred birds released at 1&ndash;3 yr old had very low survival (0.05 &plusmn; 0.06). Only 8 of 123 (7%) captive birds were seen again after release. Two wild birds resighted five years after marking are the oldest known individuals, being at least six years of age. Malarial infection did not affect survival of wild Puaiohi, unlike many Hawaiian forest birds. The difference between adult male and adult female survival is consistent with rat ( Rattus spp.) predation of females on the nest as a major source of mortality. As such, attempting to reduce nest predation by controlling rats may be the best available management option. Releasing captive-bred birds has had little effect on the wild population in recent years.

The Condor

Palila restoration research, 1996−2012

The Palila Restoration Project was initiated in 1996 by the U.S. Geological Survey to assist government agencies mitigate the effects of realigning Saddle Road (Highway 200) through Palila Critical Habitat (U.S. Fish and Wildlife Service 1998, Federal Highway Administration 1999). Ecological research on the palila (Loxioides bailleui), an endangered Hawaiian forest bird, carried out by the U.S. Geological Survey (formerly organized as the Research Division of U.S. Fish and Wildlife Service) since 1987 and research conducted by the Palila Restoration Project provided the scientific bases for developing a recovery strategy (U.S. Fish and Wildlife Service 2006) and its adaptive implementation. The main objectives of the Palila Restoration Project were to develop techniques for reintroducing the palila to a portion of its former range, investigate the biological threats to the palila and its habitat, and synthesize the existing body of ecological knowledge concerning the palila. Five broad study themes formed the research framework: 1. Population reintroduction and restoration 2. Demography and breeding ecology 3. Habitat use and food ecology 4. Vegetation ecology 5. Predator ecology and management An element that was not included in the research program of the project was the ecology and management of introduced ungulates, which has historically constituted the single greatest threat to Palila Critical Habitat (Banko et al. 2009). The absence of ungulate studies should not be interpreted to mean that we believe ungulates no longer damage palila habitat. Other research has already established that removing alien browsers and grazers from Mauna Kea is essential for the recovery of the subalpine forest on which palila now depend (Scowcroft and Giffin 1983; Scowcroft and Sakai 1983; Scowcroft and Conrad 1988, 1992; Hess et al. 1999). Moreover, the Federal District Court of Hawai&lsquo;i has ordered the state of Hawai&lsquo;i to remove browsing ungulates from Palila Critical Habitat (Banko et al. 2009, Hess and Banko 2011). This final report summarizes results of Palila Restoration Project research from December 1996 to December 2012. Even though some results contained in this report have been published in scientific journals and other technical reports (Appendix I), they are included here to provide a comprehensive chronicle of all project activities.

Hawaii

Response of palila and other subalpine Hawaiian forest bird species to prolonged drought and habitat degradation by feral ungulates

Extinction has claimed half of all historically-known Hawaiian passerines, and today many extant species are increasingly threatened due to the combined effects of invasive species and climate change. Habitat disturbance has affected populations of feeding specialists most profoundly, and our results indicate that specialists continue to be most vulnerable, although even some abundant, introduced, generalist species also may be affected. Surveys of passerines during 1998–2011 in subalpine woodland habitat on Mauna Kea Volcano, Island of Hawai′i, revealed that the abundance of the critically endangered palila ( Loxioides bailleui ), a seed specialist, declined by 79% after 2003. The ′akiapōlā′au ( Hemignathus munroi ), an endangered specialist insectivore, was not detected in the survey area after 1998. The Hawai′i ′amakihi ( Hemignathus virens virens ), a generalist feeder and the most abundant species on Mauna Kea, was the only native species to maintain a stable population. The Japanese white-eye ( Zosterops japonicus ), a well-entrenched generalist and one of the three most common introduced species, declined. Drought prevailed in 74% of months during 2000–2011, and dry conditions contributed to the recent decline of the palila by reducing the annual māmane ( Sophora chrysophylla ) seed pod crop, which influences palila breeding and survival. Sustained browsing by introduced ungulates also lowered habitat carrying capacity, and their elimination should reduce the effects of drought and promote forest restoration. Our results illustrate how the feeding ecology of a species can influence its response to interacting environmental perturbations, and they underscore the value of long-term monitoring to detect population trends of sensitive species.

Hawai'i

2011 Kiwikiu (Maui Parrotbill) and Maui 'Alauahio abundance estimates and the effect of sampling effort on power to detect a trend

The Kiwikiu ( Pseudonestor xanthophrys ), also called the Maui Parrotbill, is an endangered, forest bird found only in high elevation, wet forest of the eastern portion of Maui Island. Recent surveys, conducted at five year intervals, have revealed wide variation in abundance estimates (Camp et al. 2009). Effective management and conservation requires accurate estimates of abundance, which is difficult for rare species such as the Kiwikiu because low density leads to few observations, resulting in low sample size and high uncertainty in abundance estimates. In addition to being rare, they occur in remote, difficult to access terrain, which makes them difficult to detect and further reduces the accuracy of counts. The Maui `Alauahio ( Paroreomyza montana ), sometimes called the Maui Creeper, historically occupied the entire island of Maui (Gorresen et al. 2009). It has since been extirpated from much of its original habitat and now occurs in forested areas of East Maui where its habitat overlaps with that of the Kiwikiu. Though they share the same habitat, the `Alauahio is much more abundant—by more than two orders of magnitude—and occurs over a wider range than the Kiwikiu. Both species appear to have no statistically significant population trend from 1980–2001, but abundance estimates vary widely from survey to survey and have wide uncertainties (Camp et al. 2009). Ideally survey design should result in estimates precise enough to be able to detect significant declines in abundance that may trigger management intervention. We wished to improve the accuracy of Kiwikiu abundance estimates and the ability to detect significant trends in abundance. To that end, in 2011, repeated point count surveys were conducted across the Kiwikiu range, excluding Haleakalā National Park (Figure 1). The increased sampling effort increases sample size and improves the precision of estimates, and repeat samples also allowed us to partition within-year and between-year variation in surveys, increasing the statistical power to detect trends.

Hawaii