Geology ReportsSearch

Geology topics

Carter T. Atkinson

Publications and source records attributed to Carter T. Atkinson.

At least 37 records · Page 2Linked to original sources

Genetic variation and structure in remnant population of critically endangered Melicope zahlbruckneri

The distribution and amount of genetic variation within and between populations of plant species are important for their adaptability to future habitat changes and also critical for their restoration and overall management. This study was initiated to assess the genetic status of the remnant population of Melicope zahlbruckneri–a critically endangered species in Hawaii, and determine the extent of genetic variation and diversity in order to propose valuable conservation approaches. Estimated genetic structure of individuals based on molecular marker allele frequencies identified genetic groups with low overall differentiation but identified the most genetically diverse individuals within the population. Analysis of Amplified Fragment Length Polymorphic (AFLP) marker loci in the population based on Bayesian model and multivariate statistics classified the population into four subgroups. We inferred a mixed species population structure based on Bayesian clustering and frequency of unique alleles. The percentage of Polymorphic Fragment (PPF) ranged from 18.8 to 64.6% for all marker loci with an average of 54.9% within the population. Inclusion of all surviving M. zahlbruckneri trees in future restorative planting at new sites are suggested, and approaches for longer term maintenance of genetic variability are discussed. To our knowledge, this study represents the first report of molecular genetic analysis of the remaining population of M. zahlbruckneri and also illustrates the importance of genetic variability for conservation of a small endangered population.

Journal of Biodiversity & Endangered Species

Effects of Climate and land use on diversity, prevalence, and seasonal transmission of avian hematozoa in American Samoa

The indigenous forest birds of American Samoa are increasingly threatened by changing patterns of rainfall and temperature that are associated with climate change as well as environmental stressors associated with agricultural and urban development, invasive species, and new introductions of avian diseases and disease vectors. Long term changes in their distribution, diversity, and population sizes could have significant impacts on the ecological integrity of the islands because of their critical role as pollinators and seed dispersers. We documented diversity of vector borne parasites on Tutuila and Ta‘u Islands over a 10-year period to expand earlier observations of Plasmodium, Trypanosoma , and filarial parasites, to provide better parasite identifications, and to create a better baseline for detecting new parasite introductions. We also identified potential mosquito vectors of avian Plasmodium and Trypanosoma , determined whether land clearing and habitat alterations associated with subsistence farming within the National Park of American Samoa can influence parasite prevalence, and determined whether parasite prevalence is correlated with seasonal changes in rainfall, temperature and wind speed. Three taxonomically distinct lineages of Plasmodium were identified from mosquito vectors and forest birds based on partial sequence data from parasite mitochondrial genes. All three have been described from passerine and galliform birds in Australasia. Two lineages, SCEDEN01 and ORW1, had elongate gametocytes and large schizonts that were consistent with species of Plasmodium in the subgenus Giavannolaia, but were taxonomically distinct from known morphological species of Plasmodium based on a Bayesian phylogenetic analysis of a 478 bp region of the parasite cytochrome b gene. Both are candidates for description as new species. The third lineage (GALLUS02) was detected only in mosquito vectors on Tutuila and was similar in cytochrome b sequence to P. juxtanucleare , a pathogenic species of Plasmodium from chickens and other galliform birds from Australasia, Africa, and South America. Plasmodium relictum , the malarial parasite that has had such a devastating impact on Hawaiian forest birds, was not detected. We observed large, striated trypanosomes in avian hosts from both Tutuila and Ta‘u Islands that fell within the same taxonomic clade as T. corvi and T. culicavium based on 18S ribosomal DNA sequence. We also observed sheathed microfilariae with pointed tails that had some morphological similarities to microfilaria from species of Pelecitus, Struthiofilaria and Eulimdana , but identification will require recovery and examination of adult filarial worms from the connective tissue or body cavities of infected birds. We also observed one or more species of haemococcidians (Isospora, synonym = Atoxoplasma ) within circulating lymphocytes from multiple avian host species. Overall prevalence of Plasmodium was higher on Ta‘u (22%, 75/341) than Tutuila (9.2%, 27/294), with most infections occurring in Polynesian starlings, Samoan starlings, Wattled honeyeaters, and Cardinal honeyeaters. Prevalence was relatively constant from year to year and between seasons at individual study sites, but varied among study sites, with highest rates of infection in areas with agricultural activity at Faleasao (37.4%, 73/195, Ta‘u Island) and Amalau Valley (9.7%, 21/216, Tutuila Island). Prevalence in more remote areas of the National Park of American Samoa was lower, ranging from 1.4% (2/146) at Laufuti and Luatele on Ta‘u to 7.7% (6/78) at Olo Ridge on Tutuila. Similar trends were evident for infections with Trypanosoma and filarial worms. Overall prevalence was not influenced significantly by warmer, wet (summer) or cooler, dry (winter) season. We detected Plasmodium infections in Culex sitiens and C. quinquefasciatus through either salivary gland and midgut dissections or PCR amplification of parasite cytochrome b genes in pooled or individual samples of mosquitoes that were collected on Tutuila. Pooled or individual Aedes oceanicus, A. polynesiensis, A. tutuilae, A. upolensis, A. nocturnus, Aedes (Finlaya) (mixed pools of A. samoanus, A. oceanicus, A. tutuilae), Aedes (Stegomyia) (mixed pools of A. aegypti, A. upolensis, A. polynesiensis ), and C. annulirostris were negative for Plasmodium , but we detected infections with Trypanosoma through midgut and salivary gland dissections in a single C. sitiens from Amalau Valley, Tutuila and three A. oceanicus from Faleasao, Ta‘u. Two of the A. oceanicus from Faleasao amplified successfully with Trypanosoma primers, but sequences were distinctly different from those obtained from avian hosts. We found a strong association between land use and prevalence of mosquito-transmitted parasites on Ta‘u Island with odds of being infected more than 20 times greater in agricultural plots than more remote native forest. This relationship was evident on Tutuila Island but not statistically significant because of the close proximity of study sites and observed movement of birds between native forest and agricultural land. Our data support previous studies that have suggested that Plasmodium and other vector-borne parasites are part of the indigenous parasite fauna in American Samoa. Transmission dynamics appear to be affected by environmental changes associated with land use practices.

HI

Genetic diversity of Wolbachia endosymbionts in Culex quinquefasciatus from Hawai`i, Midway Atoll, and Samoa

Incompatible insect techniques are potential methods for controlling Culex quinquefasciatus and avian disease transmission in Hawai‘i without the use of pesticides or genetically modified organisms. The approach is based on naturally occurring sperm-egg incompatibilities within the Culex pipiens complex that are controlled by different strains of the bacterial endosymbiont Wolbachia pipientis (wPip). Incompatibilities can be unidirectional (crosses between males infected with strain A and females infected with strain B are fertile, while reciprocal crosses are not) or bidirectional (reciprocal crosses between sexes with different wPip strains are infertile). The technique depends on release of sufficient numbers of male mosquitoes infected with an incompatible wPip strain to suppress mosquito populations and reduce transmission of introduced avian malaria ( Plasmodium relictum ) and Avipoxvirus in native forest bird habitats. Both diseases are difficult to manage using more traditional methods based on removal and treatment of larval habitats and coordination of multiple approaches may be needed to control this vector. We characterized the diversity of Wolbachia strains in C. quinquefasciatus from Hawai‘i, Kaua‘i, Midway Atoll, and American Samoa with a variety of genetic markers to identify compatibility groups and their distribution within and between islands. We confirmed the presence of wPip with multilocus sequence typing, tested for local genetic variability using 16 WO prophage genes, and identified similarities to strains from other parts of the world with a transposable element (tr1). We also tested for genetic differences in ankyrin motifs (ank2 and pk1) which have been used to classify wPip strains into five worldwide groups (wPip1–wPip5) that vary in compatibility with each other based on experimental crosses. We found a mixture of both widely distributed and site specific genotypes based on presence or absence of WO prophage and transposable element markers on Hawai‘i Island (Volcano, Pu‘u Wa‘awa‘a, Laupāhoehoe, Kaumana, Kahuku, Nīnole, and Maulua Gulch), Kaua‘i Island (Kawaikōī, Mōhihi, Kalāheo, Lāwa‘i and Hanapepe) and Midway Atoll. Genotypes from American Samoa were unique and formed their own clade. Based on analysis of ankyrin motifs, wPip strains from Hawai‘i, Kaua‘i, and Midway Atoll were most similar to wPip5 strains of Australasian origin. By contrast, Wolbachia strains from Culex quinquefasciatus collected in American Samoa were most similar to wPip3 strains of American origin. We detected a single Culex mosquito from Pu‘u Wa‘awa‘a on Hawai‘i Island that was infected with a unique wPip3 genotype. This discovery, plus a rarefaction analysis of genotypes from Kaua‘i and Hawai‘i Islands suggests that limited sampling may have underestimated diversity of wPip in our study. Mosquitoes infected with wPip5 and wPip3 are bidirectionally compatible with each other based on prior studies, which would support their ability to coexist within the same population on Hawai‘i Island. Available evidence from prior studies suggests that genotype wPip4 from Africa, the Middle East, Europe, and Asia is bidirectionally incompatible with genotype wPip5 and varies in compatibility with genotype wPip3 depending on geographic origin. Since wPip5 appears to be the most common compatibility group in Hawai‘i based on limited sampling, logical next steps are to 1) expand the current survey to include additional islands and localities, 2) infect a laboratory colony of Hawaiian Culex with wPip4 through tetracycline treatment of Hawaiian mosquitoes and backcross with Culex from Europe, North Africa, and the Middle East that are naturally infected with wPip4, 3) conduct cage trials to confirm bidirectional incompatibilities between Hawaiian Culex infected with wPip4 and wPip5, and 4) conduct field trials to evaluate whether release of incompatible males can be applied at small scales to suppress local populations.

American Samoa, Hawaii

Changes in the prevalence of avian disease and mosquito vectors at Hakalau Forest National Wildlife Refuge: a 14-year perspective and assessment of future risk

Throughout the main Hawaiian Islands, introduced mosquito-borne disease has had, and continues to have, a profound impact on the distributions and abundance of native Hawaiian forest birds. Populations of remaining native forest birds are largely restricted to high elevation forests where mean temperatures are marginal for vector and parasite development and limited availability of larval mosquito habitat constrains mosquito populations and disease transmission. Hakalau Forest National Wildlife Refuge (HFNWR) was established for the preservation of endemic avifauna in 1985. Since its creation, native bird communities there have remained intact and most species populations are stable or increasing. However, avian malaria had been detected at HFNWR in the past and, in light of documented climate change, new concerns have been raised regarding the long-term fate of the refuge’s forest birds. To examine the possible changes in avian malaria transmission at HFNWR we sampled forest birds for blood parasites, trapped adult mosquitoes and surveyed larval mosquito habitat at three sites during 2012 and compared our results with similar data collected between 1998 and 1999. We tested blood samples by polymerase chain reaction (PCR), immunoblotting, and microscopy to determine prevalence of acute and chronic infection and used attractive gravid traps to sample the vector mosquito Culex quinquefasciatus. Our study documented spatial trends and temporal changes in the prevalence of avian malaria, mosquito presence, larval mosquito habitat and feral pig activity at HFNWR. We found evidence of local transmission in high elevation forests, a general pattern of increasing prevalence at lower elevations and along a South to North gradient and a two-fold decrease in the prevalence of avian malaria in the intervening 14 years. Despite considerable effort, we were unable to detect larval C. quinquefasciatus and captured only one adult indicating that the vector of avian malaria has a very limited presence at HFNWR. We did, however, document the establishment of another invasive mosquito, Aedes japonicus japonicus , and its occurrence in tree fern cavities and rock pools as larval habitat in the lower forests of HFNWR. We suggest that interspecific competition by A. j. japonicus and predation by a suite of native predators may provide biotic resistance to the establishment of permanent C. quinquefasciatus populations. While current predictions of climate change in the Hawaiian Islands include a gradual warming and enhanced transmission by mid-century, the current cooling trend recorded at high elevation HFNWR illustrates the importance of monitoring to document fine scale temporal and site specific changes in prevalence. Long term changes in precipitation may have a more profound effect on local transmission of malaria than temperature and we may have already seen some potential impacts of an extended drought at HFNWR with a decrease in feral pig activity and pig associated larval mosquito habitat and increases in stream-associated larval mosquito habitat.

Hawaii

Will a warmer and wetter future cause extinction of native Hawaiian forest birds?

Isolation of the Hawaiian archipelago produced a highly endemic and unique avifauna. Avian malaria ( Plasmodium relictum ), an introduced mosquito-borne pathogen, is a primary cause of extinctions and declines of these endemic honeycreepers. Our research assesses how global climate change will affect future malaria risk and native bird populations. We used an epidemiological model to evaluate future bird-mosquito-malaria dynamics in response to alternative climate projections from the Coupled Model Intercomparison Project (CMIP). Climate changes during the second half of the century accelerate malaria transmission and cause a dramatic decline in bird abundance. Different temperature and precipitation patterns produce divergent trajectories where native birds persist with low malaria infection under a warmer and dryer projection (RCP4.5), but suffer high malaria infection and severe reductions under hot and dry (RCP8.5) or warm and wet (A1B) futures. We conclude that future global climate change will cause significant decreases in the abundance and diversity of remaining Hawaiian bird communities. Because these effects appear unlikely before mid-century, natural resource managers have time to implement conservation strategies to protect this unique avifauna from further decimation. Similar climatic drivers for avian and human malaria suggest that mitigation strategies for Hawai'i have broad application to human health.

Hawai'i

Avian malaria in Hawaiian forest birds: Infection and population impacts across species and elevations

Wildlife diseases can present significant threats to ecological systems and biological diversity, as well as domestic animal and human health. However, determining the dynamics of wildlife diseases and understanding the impact on host populations is a significant challenge. In Hawai‘i, there is ample circumstantial evidence that introduced avian malaria ( Plasmodium relictum ) has played an important role in the decline and extinction of many native forest birds. However, few studies have attempted to estimate disease transmission and mortality, survival, and individual species impacts in this distinctive ecosystem. We combined multi-state capture-recapture (longitudinal) models with cumulative age-prevalence (cross-sectional) models to evaluate these patterns in Apapane, Hawai‘i Amakihi, and Iiwi in low-, mid-, and high-elevation forests on the island of Hawai‘i based on four longitudinal studies of 3–7 years in length. We found species-specific patterns of malaria prevalence, transmission, and mortality rates that varied among elevations, likely in response to ecological factors that drive mosquito abundance. Malaria infection was highest at low elevations, moderate at mid elevations, and limited in high-elevation forests. Infection rates were highest for Iiwi and Apapane, likely contributing to the absence of these species in low-elevation forests. Adult malaria fatality rates were highest for Iiwi, intermediate for Amakihi at mid and high elevations, and lower for Apapane; low-elevation Amakihi had the lowest malaria fatality, providing strong evidence of malaria tolerance in this low-elevation population. Our study indicates that hatch-year birds may have greater malaria infection and/or fatality rates than adults. Our study also found that mosquitoes prefer feeding on Amakihi rather than Apapane, but Apapane are likely a more important reservoir for malaria transmission to mosquitoes. Our approach, based on host abundance and infection rates, may be an effective alternative to mosquito blood meal analysis for determining vector-host contacts when mosquito densities are low and collection of blood-fed mosquitoes is impractical. Our study supports the hypothesis that avian malaria has been a primary factor influencing the elevational distribution and abundance of these three species, and likely limits other native Hawaiian species that are susceptible to malaria.

Hawai'i

Avian disease and mosquito vectors in the Kahuku unit of Hawai`i Volcanoes National Park and Ka`u Forest Reserve

While avian disease has been well-studied in windward forests of Hawai&lsquo;i Island, there have been few studies in leeward Ka&lsquo;u. We surveyed four altitudinal sites ranging from 1,200 to 2,200 m asl in the Kahuku Unit of Hawai&lsquo;i Volcanoes National Park (Kahuku) and three altitudinal sites ranging from 1,200 to 1,500 m asl in the Ka&lsquo;u Forest Reserve (Ka&lsquo;u) for the prevalence of avian disease and presence of mosquitoes. We collected blood samples from native and non-native forest birds and screened for avian malaria (Plasmodium relictum) using PCR diagnostics. We examined birds for signs of avian pox (Avipoxvirus sp.), knemidokoptic mange (Knemidokoptes jamaicensis) and feather ectoparasites. We also trapped adult mosquitoes (Culex quinquefasciatus and Aedes japonicus japonicus) and surveyed for available larval habitat. Between September, 2012 and October, 2014, we completed 3,219 hours of mist-netting in Kahuku capturing 515 forest birds and 3,103 hours of mist-netting in Ka&lsquo;u capturing 270 forest birds. We screened 750 blood samples for avian malaria. Prevalence of avian malaria in all species was higher in Ka&lsquo;u than Kahuku when all sites were combined for each tract. Prevalence of avian malaria in resident Hawai&lsquo;i &lsquo;amakihi (Chlorodrepanis virens) was greatest at the lowest elevation sites in Kahuku (26%; 1,201 m asl) and Ka&lsquo;u (42%; 1,178 m asl) and in general, prevalence decreased with increasing elevation and geographically from east to west. Significantly higher prevalence was seen in Ka&lsquo;u at comparable low and mid elevation sites but not at comparable high elevation sites. The overall presumptive pox prevalence was 1.7% (13/785) for both tracts, and it was higher in native birds than non-native birds, but it was not significant. Presumptive knemidokoptic mange was detected at two sites in lower elevation Kahuku, with prevalence ranging from 2‒4%. The overall prevalence of ectoparasites (Analges and Proctophyllodes spp.) was 6.7% (53/785). The site with the highest prevalence was Lower Glover in Kahuku (7.2%; 10/138) and Maka&lsquo;alia in Ka&lsquo;u. In general, mosquito larval habitat was more prevalent at lower elevation sites than higher elevation sites within the Kahuku&mdash;Ka&lsquo;u landscape, and more prevalent in Ka&lsquo;u than Kahuku. We observed significantly more available larval mosquito habitat in total belt transect plots in Ka&lsquo;u than Kahuku for both hapu&lsquo;u cavities (&Chi;2 = 47.06, df = 1, p < 0.01) and other habitat types combined (i.e., ground pools, rock holes, tree holes) (&Chi;2 = 104.35, df = 1, p < 0.01). Mosquitoes were most abundant at low elevation Kahuku, but were captured at all sites up to 1,532 m asl in Kahuku. The malarial infection rate of live mosquitoes was 21% (39/186) at Kahuku and 25% (2/8) at Ka&lsquo;u. There were 19 times more larval habitats available in Ka&lsquo;u than Kahuku on survey transects, yet we captured 53 times more C. quinquefasciatus mosquitoes in Kahuku. We captured very few adult A. j. japonicus across the landscape (Ntotal = 6) and no Aedes albopictus were detected in this study. Larval surveys along ranch roads and infrastructure revealed that ground pools along rutted, overgrown ranch roads were the likely source of Kahuku mosquitoes. We did not find mosquito larvae associated with ranching infrastructure. Unlike the low elevation forests on windward Hawai&lsquo;i Island, avian malaria prevalence, mosquito abundance, and the density of available larval habitat in Kahuku and Ka&lsquo;u were relatively low. Although altitudinal variations in climate appear to be the primary factors limiting the distribution of avian disease, habitat type, avian movements, human activity, and feral pig (Sus scrofa) management all may play important roles in determining the prevalence of avian malaria across the Kahuku&mdash;Ka&lsquo;u landscape.

Hawaii

Global phylogeography of the avian malaria pathogen Plasmodium relictum based on MSP1 allelic diversity

Knowing the genetic variation that occurs in pathogen populations and how it is distributed across geographical areas is essential to understand parasite epidemiology, local patterns of virulence, and evolution of host-resistance. In addition, it is important to identify populations of pathogens that are evolutionarily independent and thus &lsquo;free&rsquo; to adapt to hosts and environments. Here, we investigated genetic variation in the globally distributed, highly invasive avian malaria parasite Plasmodium relictum , which has several distinctive mitochondrial haplotyps (cyt b lineages, SGS1, GRW11 and GRW4). The phylogeography of P. relictum was accessed using the highly variable nuclear gene merozoite surface protein 1 (MSP1), a gene linked to the invasion biology of the parasite. We show that the lineage GRW4 is evolutionarily independent of GRW11 and SGS1 whereas GRW11 and SGS1 share MSP1 alleles and thus suggesting the presence of two distinct species (GRW4 versus SGS1 and GRW11). Further, there were significant differences in the global distribution of MSP1 alleles with differences between GRW4 alleles in the New and the Old World. For SGS1, a lineage formerly believed to have both tropical and temperate transmission, there were clear differences in MSP1 alleles transmitted in tropical Africa compared to the temperate regions of Europe and Asia. Further, we highlight the occurrence of multiple MSP1 alleles in GRW4 isolates from the Hawaiian Islands, where the parasite has contributed to declines and extinctions of endemic forest birds since it was introduced. This study stresses the importance of multiple independent loci for understanding patterns of transmission and evolutionary independence across avian malaria parasites.

Ecography

Changing climate and the altitudinal range of avian malaria in the Hawaiian Islands: an ongoing conservation crisis on the island of Kaua'i

Transmission of avian malaria in the Hawaiian Islands varies across altitudinal gradients and is greatest at elevations below 1500 m where both temperature and moisture are favorable for the sole mosquito vector, Culex quinquefasciatus, and extrinsic sporogonic development of the parasite, Plasmodium relictum. Potential consequences of global warming on this system have been recognized for over a decade with concerns that increases in mean temperatures could lead to expansion of malaria into habitats where cool temperatures currently limit transmission to highly susceptible endemic forest birds. Recent declines in two endangered species on the island of Kaua'i, the &lsquo;Akikiki (Oreomystis bairdi) and &lsquo;Akeke'e (Loxops caeruleirostris), and retreat of more common native honeycreepers to the last remaining high elevation habitat on the Alaka'i Plateau suggest that predicted changes in disease transmission may be occurring. We compared prevalence of malarial infections in forest birds that were sampled at three locations on the Plateau during 1994&ndash;1997 and again during 2007&ndash;2013, and also evaluated changes in the occurrence of mosquito larvae in available aquatic habitats during the same time periods. Prevalence of infection increased significantly at the lower (1100 m, 10.3% to 28.2%), middle (1250 m, 8.4% to 12.2%), and upper ends of the Plateau (1350 m, 2.0% to 19.3%). A concurrent increase in detections of Culex larvae in aquatic habitats associated with stream margins indicates that populations of the vector are also increasing. These increases are at least in part due to local transmission because overall prevalence in Kaua'i &lsquo;Elepaio (Chasiempis sclateri), a sedentary native species, has increased from 17.2% to 27.0%. Increasing mean air temperatures, declining precipitation, and changes in streamflow that have taken place over the past 20 years are creating environmental conditions throughout major portions of the Alaka'i Plateau that support increased transmission of avian malaria.

Hawai'i

Experimental evidence for evolved tolerance to avian malaria in a wild population of low elevation Hawai`i `Amakihi ( Hemignathus virens )

Introduced vector-borne diseases, particularly avian malaria ( Plasmodium relictum ) and avian pox virus ( Avipoxvirus spp.), continue to play significant roles in the decline and extinction of native forest birds in the Hawaiian Islands. Hawaiian honeycreepers are particularly susceptible to avian malaria and have survived into this century largely because of persistence of high elevation refugia on Kaua‘i, Maui, and Hawai‘i Islands, where transmission is limited by cool temperatures. The long term stability of these refugia is increasingly threatened by warming trends associated with global climate change. Since cost effective and practical methods of vector control in many of these remote, rugged areas are lacking, adaptation through processes of natural selection may be the best long-term hope for recovery of many of these species. We document emergence of tolerance rather than resistance to avian malaria in a recent, rapidly expanding low elevation population of Hawai‘i ‘Amakihi ( Hemignathus virens ) on the island of Hawai‘i. Experimentally infected low elevation birds had lower mortality, lower reticulocyte counts during recovery from acute infection, lower weight loss, and no declines in food consumption relative to experimentally infected high elevation Hawai‘i ‘Amakihi in spite of similar intensities of infection. Emergence of this population provides an exceptional opportunity for determining physiological mechanisms and genetic markers associated with malaria tolerance that can be used to evaluate whether other, more threatened species have the capacity to adapt to this disease.

Hawai'i

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

Distribution and prevalence of knemidokoptic mange in Hawai`i `Amakihi on the island of Hawaii

Knemidokoptic mange was first observed on two Hawai&lsquo;i &lsquo;Amakihi (Hemignathus virens) mist netted in Manuka Natural Area Reserve (NAR) on the Island of Hawai&lsquo;i in June 2007. Microscopic examination of skin scrapings from lesions of the infested individuals revealed the scaley-leg mite, Knemidokoptes jamaicensis. Continued surveillance at Manuka NAR (2007-2009) documented a 24% (15/63) prevalence of mange among Hawai&lsquo;i &lsquo;Amakihi distributed from coastal habitat to 1,500 m above sea level (asl). From 2012-2014, we conducted an island-wide survey of wild passerine birds from several leeward sites (Manuka NAR, Kahuku Unit of Hawai&lsquo;i Volcanoes National Park (HAVO), Pu&lsquo;u Wa&lsquo;awa&lsquo;a Forest Bird Sanctuary, and Kipahoehoe NAR) and windward sites (Hakalau Forest National Wildlife Refuge, &lsquo;Ᾱinahou Ranch of HAVO, Malama Ki Forest Reserve, and Keauohana Forest Reserve) to determine the current distribution and host range of knemidokoptic mange. We also determined the prevalence of malaria in Hawai&lsquo;i &lsquo;Amakihi populations where mange was present and treated a subset of infested Hawai&lsquo;i &lsquo;Amakihi mange with a single, topical dose of moxidectin. We mist netted and examined a total of 1,734 passerines, including 738 Hawai&lsquo;i &lsquo;Amakihi. Mange was present in Hawai&lsquo;i &lsquo;Amakihi at Manuka NAR (595 and 305 m asl), Kahuku Ranch Unit of HAVO (Glover site: 1,201 m asl and Kipuka Akala site: 1,532 m asl), Malama Ki Forest Reserve and Keauohana Forest Reserve (293 m asl). No other passerine birds (n = 995) were infected. Mange prevalence ranged from a high of 69% (40/58) in Keauohana Forest Reserve to a low of 2% (1/65) in the Kahuku Ranch Unit of HAVO (Kipuka Akala). At Manuka NAR prevalence had decreased from 26% in 2010 to 10% (7/81) in 2012&ndash;2014. We found no significant relationship between the prevalence of mange and the prevalence of avian malaria in mesic habitats at Manuka NAR (P = 0.59 (FET, n = 81)), but there was a significant association between the prevalence of mange and the prevalence of malaria in lowland wet forests in Puna Forest Reserves (P < 0.01 (FET, n = 72)). This apparent association may be a reflection of the high prevalence of malaria (>80%) in these areas. There was no difference in the frequency of recapture of birds that were infested versus un-infested at first capture at our long-term sites (Manuka NAR and Puna sites) (&chi;2(1, n = 227) = 1.51, P = 0.22, but when all sites with mange present were pooled, there was a significant difference in the frequency of recaptures between infested and un-infested birds (&chi;2(1, n = 424) = 7.13, P = 0.01). There was a significant association between parasitemia level (per 10,000 RBCs) and the ranked stage of mange present in infested individuals. We treated 24 Hawai&lsquo;i &lsquo;Amakihi with moxidectin and upon recapture (n = 2), found a reduction in both the size and stage of mange lesions, such that a single dose, topical treatment of moxidectin appears to be an effective treatment for knemidokoptic mange in wild populations. Our results suggest that knemidokoptic mange is currently limited to Hawai&lsquo;i &lsquo;Amakihi and prevalent in low elevation sites on both the windward and leeward sides of the island.

Hawaii

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

When worlds collide: challenges and opportunities for conservation of biodiversity in the Hawaiian Islands

This chapter identifies four key challenges and opportunities for long-term conservation of biodiversity in the Hawaii's Islands. Following are the challenges that need to be resolved for remaining species of native forest birds to survive into the next century: invasive species, landscape processes, social factors, and climate change. These challenges are also relevant to other threatened terrestrial taxonomic groups (i.e., plants and invertebrates) in the Hawaiian Islands. Such threats are familiar to conservation biologists the world over, but rarely do they act as synergistically as they do in the Hawaiian Islands. The chapter reviews conservation successes and failures in Hawaii, and provides an example of the possible future course of conservation in other island communities.

Hawai'i

Population genetic structure of rare and endangered plants using molecular markers

This study was initiated to assess the levels of genetic diversity and differentiation in the remaining populations of Phyllostegia stachyoides and Melicope zahlbruckneri in Hawai`i Volcanoes National Park and determine the extent of gene flow to identify genetically distinct individuals or groups for conservation purposes. Thirty-six Amplified Fragment Length Polymorphic (AFLP) primer combinations generated a total of 3,242 polymorphic deoxyribonucleic acid (DNA) fragments in the P. stachyoides population with a percentage of polymorphic bands (PPB) ranging from 39.3 to 65.7% and 2,780 for the M. zahlbruckneri population with a PPB of 18.8 to 64.6%. Population differentiation (Fst) of AFLP loci between subpopulations of P. stachyoides was low (0.043) across populations. Analysis of molecular variance of P. stachyoides showed that 4% of the observed genetic differentiation occurred between populations in different k ī puka and 96% when individuals were pooled from all k ī puka. Moderate genetic diversity was detected within the M. zahlbruckneri population. Bayesian and multivariate analyses both classified the P. stachyoides and M. zahlbruckneri populations into genetic groups with considerable sub-structuring detected in the P. stachyoides population. The proportion of genetic differentiation among populations explained by geographical distance was estimated by Mantel tests. No spatial correlation was found between genetic and geographic distances in both populations. Finally, a moderate but significant gene flow that could be attributed to insect or bird-mediated dispersal of pollen across the different k ī puka was observed. The results of this study highlight the utility of a multi-allelic DNA-based marker in screening a large number of polymorphic loci in small and closely related endangered populations and revealed the presence of genetically unique groups of individuals in both M. zahlbruckneri and P. stachyoides populations. Based on these findings, approaches that can assist conservation efforts of these species are proposed.

Hawai`i

Immunological markers for tolerance to avian malaria in Hawai`i `Amakihi: new tools for restoring native Hawaiian forest birds?

We evaluated three assays for non-specific or innate immune capacity to see if measurements were independent of malarial infection and capable of distinguishing malaria-tolerant, low-elevation Hawaiʽi ʽAmakihi from malaria-susceptible, high-elevation ʽAmakihi. ʽAmakihi were captured at Malama Ki Forest Reserve (20 m), Hakalau Forest National Wildlife Refuge (1800 m), and Upper Waiakea Forest Reserve (1700 m), bled for collection of plasma and packed erythrocytes for malarial diagnostics, and either transported to Kīlauea Field Station Aviary and held in captivity for 48 hours for inoculation of wing webs with phytohemagglutinin A (PHA) or released immediately in the field after collection of a blood sample. All birds were tested by polymerase chain reaction (PCR) and microscopy to determine infection status. We found no significant association between malarial infection status and degree of wing web swelling after inoculation with PHA (T = -0.174, df = 13, P = 0.864) and no association between origin of birds from low- and high-elevation populations and degree of wing web swelling (T = 0.113, df = 52, P = 0.911). Infected ʽAmakihi from low elevation had significantly higher small molecule plasma antioxidant capacity than uninfected individuals from the same population (T = -2.675, df = 21, P = 0.014), so we limited comparisons to uninfected birds. Uninfected ʽAmakihi from low elevations did not differ in small molecule plasma antioxidant capacity from uninfected ʽAmakihi from high elevation (T = -0.260, df = 46, P = 0.796). Compared to high-elevation birds, low-elevation ʽAmakihi had significantly higher titers of natural antibodies (NAb) as measured by complement-mediated lysis of rabbit erythrocytes (Mann-Whitney U = 27, X 2 = 32.332, df = 1, P < 0.0001). This innate immunological difference may be related to ability to survive malarial infection and may prove to be important for understanding possible mechanisms for the evolution of disease tolerance in Hawaiʽi’s native bird species.

Hawai'i

Avian disease assessment in seabirds and non-native passerines birds at Midway Atoll NWR

Midway Atoll in the Northwestern Hawaiian Islands supports the largest breeding colony of Laysan albatross ( Phoebastria immutabilis ) in the world and is a proposed site for the translocation of endangered Northwestern Hawaiian Island passerine birds such as the Nihoa finch ( Telespiza ultima ), Nihoa millerbird ( Acrocephalus familiaris kingi ), or Laysan finch ( Telespiza cantans ). On the main Hawaiian Islands, introduced mosquito-borne avian malaria ( Plasmodium relictum ) and avian pox ( Avipoxvirus ) have contributed to the extinction and decline of native Hawaiian avifauna. The mosquito vector ( Culex quinquefasciatus ) is present on Sand Island, Midway Atoll, where epizootics of Avipoxvirus have been reported among nestling Laysan albatross, black-footed albatross ( Phoebastria nigripes ), and red-tailed tropicbirds ( Phaethon rubricauda ) since 1963. Two introduced passerines, the common canary ( Serinus canaria ) and the common myna ( Acridotheres tristis ), are also present on Sand Island and may serve as reservoirs of mosquito-borne pathogens. Assessing disease prevalence and transmission potential at Midway Atoll National Wildlife Refuge (NWR) is a critical first step to translocation of Nihoa endemic passerines. In May 2010 and April 2012 we surveyed Midway Atoll NWR for mosquitoes and evidence of mosquito-borne disease. Although we did not observe active pox infections on albatross nestlings in May 2010, active infections were prevalent on albatross nestlings in April 2012. Presumptive diagnosis of Avipoxvirus was confirmed by PCR amplification of the Avipoxvirus 4b core protein gene from lesions collected from 10 albatross nestlings. Products were sequenced and compared to 4b core protein sequences from 28 Avipoxvirus isolates from the Hawaiian Islands and other parts of the world. Sequences from all Midway isolates were identical and formed a clade with other Avipoxvirus isolates from seabirds that was distinct from other Avipoxvirus isolates from the Hawaiian Islands. Tissue from three presumptive avian pox lesions from common canaries tested negative for Avipoxvirus . Blood samples from 124 canaries and 61 mynas tested negative for Plasmodium by one or more diagnostic tests based on microscopy, serology, or PCR diagnostics. Prevalence of Avipoxvirus infection was highest among albatross nestlings (94.6%) in the vicinity of the septic tanks where adult C. quinquefasciatus reached their highest densities, and data from all sites suggest a positive correlation between mosquito abundance and Avipoxvirus prevalence. Adult C. quinquefasciatus were also locally abundant around fishless, constructed wetlands. Since 1996, infrastructure removal and source reduction efforts by the refuge have greatly reduced the availability of underground and container habitats for larval mosquitoes on Sand Island. However, the creation of artificial wetlands and a central septic system on Sand Island has resulted in new, highly productive larval mosquito habitat for C. quinquefasciatus . Despite the presence of endemic Avipoxvirus in albatross nestlings and the introduction of mosquito vectors and two susceptible passerine species in the last century, we found no evidence of the avian malaria Plasmodium relictum or a passerine-infecting Avipoxvirus on Midway Atoll NWR that would interfere with the successful translocation of endemic Northwestern Hawaiian Island passerines. Without eradication of mosquitoes from Midway Atoll, however, periodic epizootics of Avipoxvirus among nestling seabirds will likely continue, and the introduction of malaria and passerine strains of Avipoxvirus from migratory birds will remain a long-term threat to passerine restoration programs.

Hawai'i

Modeling future conservation of Hawaiian Honeycreepers by mosquito management and translocation of disease-tolerant Amakihi

Avian malaria is an important cause of the decline of endemic Hawaiian honeycreepers. Because of the complexity of this disease system we used a computer model of avian malaria in forest birds to evaluate how two proposed conservation strategies: 1) reduction of habitat for mosquito larvae and 2) establishment of a low-elevation, malaria-tolerant honeycreeper (Hawaii Amakihi) to mid-elevation forests would affect native Hawaiian honeycreeper populations. We evaluated these approaches in mid-elevation forests, where malaria transmission is seasonal and control strategies are more likely to work. Our model suggests the potential benefit of larval habitat reduction depends on the level of malaria transmission, abundance of larval cavities, and the ability to substantially reduce these cavities. Permanent reduction in larval habitat of >80% may be needed to control abundance of infectious mosquitoes and benefit bird populations. Establishment of malaria-tolerant Amakihi in mid-elevation forests increases Amakihi abundance, creates a larger disease reservoir, and increases the abundance of infectious mosquitoes which may negatively impact other honeycreepers. For mid-elevation sites where bird populations are severely affected by avian malaria, malaria-tolerant Amakihi had little impact on other honeycreepers. Both management strategies may benefit native Hawaiian honeycreepers, but benefits depend on specific forest characteristics, the amount of reduction in larval habitat that can be achieved, and how malaria transmission is affected by temperature.

Hawai'i