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.     

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.     

Pathogenicity of avian malaria in experimentally-infected Hawaii Amakihi

The introduction of avian malaria (Plasmodium relictum) and mosquitoes (Culex quinquefasciatus) to the Hawaiian Islands (USA) is believed to have played a major role in the decline and extinction of native Hawaiian honeycreepers (Drepanidinae). This introduced disease is thought to be one of the primary factors limiting recovery of honeycreepers at elevations below 1,200 m where native forest habitats are still relatively intact. One of the few remaining species of honeycreepers with a wide elevational distribution is the Hawaii Amakihi (Hernignathus virens). We measured morbidity and mortality in experimentally-infected Hawaii Amakihi that were captured in a high elevation, xeric habitat that is above the current range of the mosquito vector. Mortality among amakihi exposed to a single infective mosquito bite was 65% (13/20). All infected birds had significant declines in food consumption and a corresponding loss in body weight over the 60 day course of the experiment. Gross and microscopic lesions in birds that succumbed to malaria included enlargement and discoloration of the spleen and liver and parasitemias as high as 50% of circulating erythrocytes. Mortality in experimentally-infected amakihi was similar to that observed in Apapane (Himnatione sanguinea) and lower than that observed in Iiwi (Vestiaria coccinea) infected under similar conditions with the same parasite isolate. We conclude that the current elevational and geographic distribution of Hawaiian honeycreepers is determined by relative susceptibility to avian malaria.     

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. 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.     

A PCR test for avian malaria in Hawaiian birds

The decline of native Hawaiian forest birds since European contact is attributed to factors ranging from habitat destruction to interactions with introduced species. Remaining populations of Hawaiian honeycreepers (Fringillidae: Drepanidinae) are most abundant and diverse in high elevation refuges above the normal range of disease-carrying mosquitoes. Challenge experiments suggest that honeycreepers are highly susceptible to avian malaria (Plasmodium sp.) but resistance exists in some species. In order to detect low levels of malarial infection and quantify prevalence of Plasmodium in high elevation natural populations of Hawaiian birds, a polymerase chain reaction (PCR) based diagnostic test was developed that identifies rRNA genes of Plasmodium in avian blood samples. Quantitative competitive PCR (QC-PCR) experiments indicate that the detection limit of our test is an order of magnitude greater than that reported for human malaria DNA blot tests. Compared with standard histological methods, the PCR test detected a higher prevalence of diseased birds at mid-elevations. Malaria was detected in three species of native birds living in a high elevation wildlife refuge on the island of Hawaii and in four species from Maui. Our results show that avian malaria is more widespread in Hawaiian forests than previously thought, a finding that has important conservation implications for these threatened species.     

Assessing the potential of translocating vulnerable forest birds by searching for novel and enduring climatic ranges

Hawaiian forest birds are imperiled, with fewer than half the original >40 species remaining extant. Recent studies document ongoing rapid population decline and project complete climate‐based range losses for the critically endangered Kaua'i endemics ‘akeke’e (Loxops caeruleirostris) and ‘akikiki (Oreomystis bairdi) by end‐of‐century due to projected warming. Climate change facilitates the upward expansion of avian malaria into native high elevation forests where disease was historically absent. While intensified conservation efforts attempt to safeguard these species and their habitats, the magnitude of potential loss and the urgency of this situation require all conservation options to be seriously considered. One option for Kaua’i endemics is translocation to islands with higher elevation habitats. We explored the feasibility of interisland translocation by projecting baseline and future climate‐based ranges of ‘akeke’e and ‘akikiki across the Hawaiian archipelago. For islands where compatible climates for these species were projected to endure through end‐of‐century, an additional climatic niche overlap analysis compares the spatial overlap between Kaua’i endemics and current native species on prospective destination islands. Suitable climate‐based ranges exist on Maui and Hawai'i for these Kaua'i endemics that offer climatically distinct areas compared to niche distributions of destination island endemics. While we recognize that any decision to translocate birds will include assessing numerous additional social, political, and biological factors, our focus on locations of enduring and ecologically compatible climate‐based ranges represents the first step to evaluate this potential conservation option. Our approach considering baseline and future distributions of species with climatic niche overlap metrics to identify undesirable range overlap provides a method that can be utilized for other climate‐vulnerable species with disjointed compatible environments beyond their native range.     

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 ‘Akikiki (Oreomystis bairdi) and ‘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 1994–1997 and again during 2007–2013, 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 ‘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.     

Large-Scale Range Collapse of Hawaiian Forest Birds under Climate Change and the Need 21st Century Conservation Options

Hawaiian forest birds serve as an ideal group to explore the extent of climate change impacts on at-risk species. Avian malaria constrains many remaining Hawaiian forest bird species to high elevations where temperatures are too cool for malaria’s life cycle and its principal mosquito vector. The impact of climate change on Hawaiian forest birds has been a recent focus of Hawaiian conservation biology, and has centered on the links between climate and avian malaria. To elucidate the differential impacts of projected climate shifts on species with known varying niches, disease resistance and tolerance, we use a comprehensive database of species sightings, regional climate projections and ensemble distribution models to project distribution shifts for all Hawaiian forest bird species. We illustrate that, under a likely scenario of continued disease-driven distribution limitation, all 10 species with highly reliable models (mostly narrow-ranged, single-island endemics) are expected to lose >50% of their range by 2100. Of those, three are expected to lose all range and three others are expected to lose >90% of their range. Projected range loss was smaller for several of the more widespread species; however improved data and models are necessary to refine future projections. Like other at-risk species, Hawaiian forest birds have specific habitat requirements that limit the possibility of range expansion for most species, as projected expansion is frequently in areas where forest habitat is presently not available (such as recent lava flows). Given the large projected range losses for all species, protecting high elevation forest alone is not an adequate long-term strategy for many species under climate change. We describe the types of additional conservation actions practitioners will likely need to consider, while providing results to help with such considerations.     

The effect of isolation,fragmentation, and population bottlenecks on song structure of a Hawaiian honeycreeper

Little is known about how important social behaviors such as song vary within and among populations for any of the endemic Hawaiian honeycreepers. Habitat loss and non‐native diseases (e.g., avian malaria) have resulted in isolation and fragmentation of Hawaiian honeycreepers within primarily high elevation forests. In this study, we examined how isolation of Hawai'i ‘amakihi (Chlorodrepanis virens) populations within a fragmented landscape influences acoustic variability in song. In the last decade, small, isolated populations of disease tolerant ‘amakihi have been found within low elevation forests, allowing us to record ‘amakihi songs across a large elevational gradient (10–1800 m) that parallels disease susceptibility on Hawai'i island. To understand underlying differences among populations, we examined the role of geographic distance, elevation, and habitat structure on acoustic characteristics of ‘amakihi songs. We found that the acoustic characteristics of ‘amakihi songs and song‐type repertoires varied most strongly across an elevational gradient. Differences in ‘amakihi song types were primarily driven by less complex songs (e.g., fewer frequency changes, shorter songs) of individuals recorded at low elevation sites compared to mid and high elevation populations. The reduced complexity of ‘amakihi songs at low elevation sites is most likely shaped by the effects of habitat fragmentation and a disease‐driven population bottleneck associated with avian malaria, and maintained through isolation, localized song learning and sharing, and cultural drift. These results highlight how a non‐native disease through its influence on population demographics may have also indirectly played a role in shaping the acoustic characteristics of a species.     

Local prevalence and transmission of avian malaria in the Alakai Plateau of Kauai,Hawaii, U.S.A.

Avian malaria is among the most important threats to native Hawaiian forest birds. It is caused by the parasite Plasmodium relictum and is transmitted by the introduced mosquito vector Culex quinquefasciatus. Temperature increases and precipitation declines due to climate change over the last decade may be responsible for the observed recent expansion in the range and prevalence of avian malaria on the Alakai Plateau, Kauai Island. To examine the hypothesis that conditions are now favorable for transmission of malaria on the Plateau, mosquitoes were sampled with CO₂ and Reiter oviposition traps at three sites (Kawaikoi, Halepa'akai, and Koke'e) on several occasions between October, 2013 and April, 2014. P. relictum infection was assessed by PCR or dissection under a microscope. We also surveyed mosquito larvae along Halepa'akai and Kawaikoi streams. We observed that Cx. quinquefasciatus is well established on the Alakai Plateau, as mosquitoes were caught on all field trips, except in April at Halepa'akai, and larvae were found throughout the year. We observed differences in adult abundance among sites and microhabitats (stream vs ridge lines).     

Modeling the impacts of global warming on predation and biotic resistance: mosquitoes, damselflies and avian malaria in Hawaii

Biotic resistance from native predators can play an important role in regulating or limiting exotic prey. We investigate how global warming potentially alters the strength and spatial extent of these predator–prey interactions in aquatic insect ecosystems. As a simple model system, we use rock pools in streams of rainforests of Hawaii, which contain the beautiful Hawaiian damselfly Megalagrion calliphya as predator and the invasive southern house mosquito Culex quinquefasciatus as prey. This abundant mosquito is the major vector of avian malaria transmission to native forest birds. We use mathematical modeling to evaluate the potential impacts of damselfly predation and temperature on mosquito population dynamics. We model this predator–prey system along an elevational gradient (749-1952 m elevation) and assess the effect of 1°C and 2°C climate warming scenarios as well as the effects of El Niño and La Niña oscillations, on predator–prey dynamics. Our results indicate that the strength of biotic resistance of native predators on invasive prey may decrease with increasing temperature because demographic rates of predator and prey are differentially affected by temperature. Future warming could therefore increase the abundance of invasive species by releasing them from predation pressure. If the invasive species is a disease vector, these shifts could increase the impact of disease on both humans and wildlife.     

Common European birds are declining rapidly while less abundant species' numbers are rising

Biodiversity is undergoing unprecedented global decline. Efforts to slow this rate have focused foremost on rarer species, which are at most risk of extinction. Less interest has been paid to more common species, despite their greater importance in terms of ecosystem function and service provision. How rates of decline are partitioned between common and less abundant species remains unclear. Using a 30‐year data set of 144 bird species, we examined Europe‐wide trends in avian abundance and biomass. Overall, avian abundance and biomass are both declining with most of this decline being attributed to more common species, while less abundant species showed an overall increase in both abundance and biomass. If overall avian declines are mainly due to reductions in a small number of common species, conservation efforts targeted at rarer species must be better matched with efforts to increase overall bird numbers, if ecological impacts of birds are to be maintained.     

Genetic structure and evolved malaria resistance in Hawaiian honeycreepers

Infectious diseases now threaten wildlife populations worldwide but population recovery following local extinction has rarely been observed. In such a case, do resistant individuals recolonize from a central remnant population, or do they spread from small, perhaps overlooked, populations of resistant individuals? Introduced avian malaria (Plasmodium relictum) has devastated low‐elevation populations of native birds in Hawaii, but at least one species (Hawaii amakihi, Hemignathus virens) that was greatly reduced at elevations below about 1000 m tolerates malaria and has initiated a remarkable and rapid recovery. We assessed mitochondrial and nuclear DNA markers from amakihi and two other Hawaiian honeycreepers, apapane (Himatione sanguinea) and iiwi (Vestiaria coccinea), at nine primary study sites from 2001 to 2003 to determine the source of re‐establishing birds. In addition, we obtained sequences from tissue from amakihi museum study skins (1898 and 1948–49) to assess temporal changes in allele distributions. We found that amakihi in lowland areas are, and have historically been, differentiated from birds at high elevations and had unique alleles retained through time; that is, their genetic signature was not a subset of the genetic variation at higher elevations. We suggest that high disease pressure rapidly selected for resistance to malaria at low elevation, leaving small pockets of resistant birds, and this resistance spread outward from the scattered remnant populations. Low‐elevation amakihi are currently isolated from higher elevations (> 1000 m) where disease emergence and transmission rates appear to vary seasonally and annually. In contrast to results from amakihi, no genetic differentiation between elevations was found in apapane and iiwi, indicating that slight variation in genetic or life‐history attributes can determine disease resistance and population recovery. Determining the conditions that allow for the development of resistance to disease is essential to understanding how species evolve resistance across a landscape of varying disease pressures.     

Bottlenecks and multiple introductions: population genetics of the vector of avian malaria in Hawaii

Avian malaria has had a profound impact on the demographics and behaviour of Hawaiian forest birds since its vector, Culex quinquefasciatus the southern house mosquito, was first introduced to Hawaii around 1830. In order to understand the dynamics of the disease in Hawaii and gain insights into the evolution of vector-mediated parasite-host interactions in general we studied the population genetics of Cx. quinquefasciatus in the Hawaiian Islands. We used both microsatellite and mitochondrial loci. Not surprisingly we found that mosquitoes in Midway, a small island in the Western group, are quite distinct from the populations in the main Hawaiian Islands. However, we also found that in general mosquito populations are relatively isolated even among the main islands, in particular between Hawaii (the Big Island) and the remaining Hawaiian Islands. We found evidence of bottlenecks among populations within the Big Island and an excess of alleles in Maui, the site of the original introduction. The mitochondrial diversity was typically low but higher than expected. The current distribution of mitochondrial haplotypes combined with the microsatellite information lead us to conclude that there have been several introductions and to speculate on some processes that may be responsible for the current population genetics of vectors of avian malaria in Hawaii.     

Arthropod-borne disease as a possible factor limiting the distribution of birds

Warner found that arthropod-borne disease in the Hawaiian islands (bird malaria and avian pox in particular) is a factor limiting the distribution of the Drepaniidae. Rowan considers it conceivable that the present distribution of certain African birds (and perhaps some other vertebrates showing similar patterns of occurrence) may have been determined in the same way. This suggestion is supported by the observations of the author in southern Africa in regard to bird malaria and avian pox.     

Fine-scale population genetic structure of a wildlife disease vector: the southern house mosquito on the island of Hawaii

The southern house mosquito, Culex quinquefasciatus, is a widespread tropical and subtropical disease vector. In the Hawaiian Islands, where it was introduced accidentally almost two centuries ago, it is considered the primary vector of avian malaria and pox. Avian malaria in particular has contributed to the extinction and endangerment of Hawaii's native avifauna, and has altered the altitudinal distribution of native bird populations. We examined the population genetic structure of Cx. quinquefasciatus on the island of Hawaii at a smaller spatial scale than has previously been attempted, with particular emphasis on the effects of elevation on population genetic structure. We found significant genetic differentiation among populations and patterns of isolation by distance within the island. Elevation per se did not have a limiting effect on gene flow; however, there was significantly lower genetic diversity among populations at mid elevations compared to those at low elevations. A recent sample taken from just above the predicted upper altitudinal distribution of Cx. quinquefasciatus on the island of Hawaii was confirmed as being a temporary summer population and appeared to consist of individuals from more than one source population. Our results indicate effects of elevation gradients on genetic structure that are consistent with known effects of elevation on population dynamics of this disease vector.     

Diversity,origins and virulence of <Emphasis Type="Italic">Avipoxviruses</Emphasis> in Hawaiian Forest Birds

We cultured avian pox (Avipoxvirus spp.) from lesions collected on Hawai‘i, Maui, Moloka‘i, and ‘Oahu in the Hawaiian Islands from 15 native or non-native birds representing three avian orders. Phylogenetic analysis of a 538 bp fragment of the gene encoding the virus 4b core polypeptide revealed two distinct variant clusters, with sequences from chickens (fowlpox) forming a third distinct basal cluster. Pox isolates from one of these two clusters appear closely related to canarypox and other passerine pox viruses, while the second appears more specific to Hawai‘i. There was no evidence that birds were infected simultaneously with multiple pox virus variants based on evaluation of multiples clones from four individuals. No obvious temporal or geographic associations were observed and strict host specificity was not apparent among the 4b-defined field isolates. We amplified a 116 bp 4b core protein gene fragment from an ‘Elepaio (Chasiempis sandwichensis) collected in 1900 on Hawai‘i Island that clustered closely with the second of the two variants, suggesting that this variant has been in Hawai‘i for at least 100 years. The high variation detected between the three 4b clusters provides evidence for multiple, likely independent introductions, and does not support the hypothesis of infection of native species through introduction of infected fowl. Preliminary experimental infections in native Hawai‘i ‘Amakihi (Hemignathus virens) suggest that the 4b-defined variants may be biologically distinct, with one variant appearing more virulent. These pox viruses may interact with avian malaria (Plasmodium relictum), another introduced pathogen in Hawaiian forest bird populations, through modulation of host immune responses.     

Was disease involved in the decimation of Guam's avifauna?

Between 1982 and 1986, 402 (290 live, 112 dead) exotic, migrant or native resident birds on Guam were surveyed for disease-causing agents to determine the role of disease in the decline of native forest bird populations on Guam. Traumatic injury, primarily from collisions with motor vehicles and predation, was the most prevalent (46%) cause of death. Thirty-eight percent of the carcasses examined were in poor body condition largely as a result of inadequate nutrition in captive native birds and poultry and adipose exhaustion in errant migrants. A variety of commensal or opportunistic bacteria, including Salmonella spp., were cultured from 220 birds, and nothing remarkable was found in 15 fecal samples. Lastly, no haematozoans, the suspected cause for the decline of the Hawaiian avifauna, were observed in blood slides examined from 260 birds. Based on the results of the survey and other lines of evidence presented in the discussion, we concluded there were no data implicating disease in the decline of Guam's avifauna.     

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 large A. koa trees 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 the A. koa population 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.     

Is Avian Malaria Playing a Role in Native Bird Declines in New Zealand? Testing Hypotheses along an Elevational Gradient

The mosquito-borne disease avian malaria (Plasmodium spp.) has impacted both captive populations and wild individuals of native New Zealand bird species. However, whether or not it is a cause of concern to their wild populations is still unclear. In Hawaii, the disease has been a major factor in the population declines of some native forest bird species, often limiting their elevational distribution due to an inverse relationship between force of infection and elevation. While studies have investigated latitudinal patterns of infection in New Zealand, elevational patterns are unexplored. To address this, a survey was conducted in Nelson Lakes National Park, a site experiencing native bird declines in which disease has been suggested as playing a role, to investigate whether there is a similar inverse relationship in New Zealand. Results from blood samples (n = 436) collected over three seasons across a broad elevational range (650–1400 m) support there being such a relationship. In addition, an overall higher prevalence in non-native (14.1%) versus native birds (1.7%) may indicate differential impacts on these two groups, while particularly high prevalence in non-native Turdus spp. supports previous suggestions that they are key reservoir hosts for the disease. Overall, these findings add weight to the hypothesis that avian malaria is playing a role in ongoing declines of native New Zealand birds.