Rapid environmental change in Alaska and other regions of the Arctic and sub-Arctic has raised concerns about increasing human exposure to ticks and the pathogens they carry. We tested a sample of ticks collected through a combination of passive and active surveillance from humans, domestic animals, and wildlife hosts in Alaska for a panel of the most common tick-borne pathogens in the contiguous United States to characterize the diversity of microbes present in this region. We tested 189 pooled tick samples collected in 2019-2020 for Borrelia spp., Anaplasma spp., Ehrlichia spp., and Babesia spp. using a multiplex PCR amplicon sequencing assay. We found established populations of Ixodes angustus Neumann (Acari: Ixodidae), Ixodes uriae White (Acari: Ixodidae), and Haemaphysalis leporispalustris Packard (Acari: Ixodidae) in Alaska, with I. angustus found on a variety of hosts including domestic companion animals (dogs and cats), small wild mammals, and humans. Ixodes angustus were active from April through October with peaks in adult and nymphal activity observed in summer months (mainly July). Although no known human pathogens were detected, Babesia microti-like parasites and candidatus Ehrlichia khabarensis were identified in ticks and small mammals. The only human pathogen detected (B. burgdorferi s.s.) was found in a tick associated with a dog that had recently traveled to New York, where Lyme disease is endemic. This study highlights the value of a combined passive and active tick surveillance system to detect introduced tick species and pathogens and to assess which tick species and microbes are locally established.

In the western United States, Ixodes pacificus Cooley & Kohls (Acari: Ixodidae) is the primary vector of the agents causing Lyme disease and granulocytic anaplasmosis in humans. The geographic distribution of the tick is associated with climatic variables that include temperature, precipitation, and humidity, and biotic factors such as the spatial distribution of its primary vertebrate hosts. Here, we explore (1) how climate change may alter the geographic distribution of I. pacificus in California, USA, during the 21(st) century, and (2) the spatial overlap among predicted changes in tick habitat suitability, land access, and ownership. Maps of potential future suitability for I. pacificus were generated by applying climate-based species distribution models to a multi-model ensemble of climate change projections for the Representative Concentration Pathway (RCP) 4.5 (moderate emission) and 8.5 (high emission) scenarios for two future periods: mid-century (2026-2045) and end-of-century (2086-2099). Areas climatically-suitable for I. pacificus are projected to expand by 23% (mid-century RCP 4.5) to 86% (end-of-century RCP 8.5) across California, compared to the historical period (1980-2014), with future estimates of total suitable land area ranging from about 88 to 133 thousand km(2), or up to about a third of California. Regions projected to have the largest area increases in suitability by end-of-century are in northwestern California and the south central and southern coastal ranges. Over a third of the future suitable habitat is on lands currently designated as open access (i.e. publicly available), and by 2100, the amount of these lands that are suitable habitat for I. pacificus is projected to more than double under the most extreme emissions scenario (from ~23,000 to >51,000 km(2)). Of this area, most is federally-owned (>45,000 km(2)). By the end of the century, 26% of all federal land in the state is predicted to be suitable habitat for I. pacificus. The resulting maps may facilitate regional planning and preparedness by informing public health and vector control decision-makers.

Ixodes pacificus Cooley & Kohls (Acari: Ixodidae), the primary vector of Lyme disease spirochetes to humans in the far-western United States, is broadly distributed across Pacific Coast states, but its distribution is not uniform within this large, ecologically diverse region. To identify areas of suitable habitat, we assembled records of locations throughout California where two or more I. pacificus were collected from vegetation from 1980 to 2014. We then employed ensemble species distribution modeling to identify suitable climatic conditions for the tick and restricted the results to land cover classes where these ticks are typically encountered (i.e., forest, grass, scrub-shrub, riparian). Cold-season temperature and rainfall are particularly important abiotic drivers of suitability, explaining between 50 and 99% of the spatial variability across California among models. The likelihood of an area being classified as suitable increases steadily with increasing temperatures >0 degrees C during the coldest quarter of the year, and further increases when precipitation amounts range from 400 to 800 mm during the coldest quarter, indicating that areas in California with relatively warm and wet winters typically are most suitable for I. pacificus. Other consistent predictors of suitability include increasing autumn humidity, temperatures in the warmest month between 23 and 33 degrees C, and low-temperature variability throughout the year. The resultant climatic suitability maps indicate that coastal California, especially the northern coast, and the western Sierra Nevada foothills have the highest probability of I. pacificus presence.

The distribution of I. scapularis, the tick vector of the bacteria that cause Lyme disease, has been expanding over the last two decades in the north-central United States in parallel with increasing incidence of human cases of Lyme disease in that region. However, assessments of residential risk for exposure to ticks are lacking from this region. Here, we measured the density of host-seeking I. scapularis nymphs in two suburban and two rural public recreational sites located in Washington County, Minnesota as well as in nearby residential properties. We sought to compare tick densities across land use types and to identify environmental factors that might impact nymphal density. We also assessed the prevalence of infection in the collected ticks with Lyme disease spirochetes (Borrelia burgdorferi sensu stricto, B. mayonii), and other I. scapularis-borne pathogens including B. miyamotoi, Babesia microti and Anaplasma phagocytophilum. Similar to studies from the eastern United States, on residential properties, I. scapularis nymphal densities were highest in the ecotonal areas between the forest edge and the lawn. Residences with the highest densities of nymphs were more likely to have a higher percentage of forest cover, log piles, and signs of deer on their property. In recreational areas, we found the highest nymphal densities both in the wooded areas next to trails as well as on mowed trails. Among the 303 host-seeking I. scapularis nymphs tested for pathogens, B. burgdorferi sensu stricto, A. phagocytophilum and B. miyamotoi were detected in 42 (13.8%), 14 (4.6%), and 2 (0.6%) nymphs, respectively.

The mosquitoes Aedes (Stegomyia) aegypti (L.)(Diptera:Culicidae) and Ae. (Stegomyia) albopictus (Skuse) (Diptera:Culicidae) transmit dengue, chikungunya, and Zika viruses and represent a growing public health threat in parts of the United States where they are established. To complement existing mosquito presence records based on discontinuous, non-systematic surveillance efforts, we developed county-scale environmental suitability maps for both species using maximum entropy modeling to fit climatic variables to county presence records from 1960-2016 in the contiguous United States. The predictive models for Ae. aegypti and Ae. albopictus had an overall accuracy of 0.84 and 0.85, respectively. Cumulative growing degree days (GDDs) during the winter months, an indicator of overall warmth, was the most important predictive variable for both species and was positively associated with environmental suitability. The number (percentage) of counties classified as environmentally suitable, based on models with 90 or 99% sensitivity, ranged from 1,443 (46%) to 2,209 (71%) for Ae. aegypti and from 1,726 (55%) to 2,329 (75%) for Ae. albopictus. Increasing model sensitivity results in more counties classified as suitable, at least for summer survival, from which there are no mosquito records. We anticipate that Ae. aegypti and Ae. albopictus will be found more commonly in counties classified as suitable based on the lower 90% sensitivity threshold compared with the higher 99% threshold. Counties predicted suitable with 90% sensitivity should therefore be a top priority for expanded mosquito surveillance efforts while still keeping in mind that Ae. aegypti and Ae. albopictus may be introduced, via accidental transport of eggs or immatures, and potentially proliferate during the warmest part of the year anywhere within the geographic areas delineated by the 99% sensitivity model.

The black-legged tick, Ixodes scapularis Say, is the primary vector in the eastern United States of the Lyme disease spirochetes Borrelia burgdorferi sensu stricto and B. mayonii, as well as Anaplasma phagocytophilum (anaplasmosis), Ehrlichia muris-like agent (ehrlichiosis), Babesia microti (babesiosis), and Powassan encephalitis virus (Goodman et al. 2005; Pritt et al. 2011, 2016). The documented distribution of this medically important tick has expanded substantially over the past two decades, paralleling the increase in the number and geographic distribution of reported human Lyme disease cases in the United States (Kugeler et al. 2015, Mead 2015, Eisen et al. 2016). Although these findings highlight the need for continued vector surveillance, resources for such work are often very limited. | Habitat suitability models can be useful for identifying areas of potential future expansion and thus aid in targeting limited public health resources. To better define the leading edge of the tick’s ongoing geographic expansion, recently we published a distribution map that identified counties that were classified as suitable by at least two statistical models, but where I. scapularis has not yet been documented to be established (Hahn et al. 2016). Peterson and Raghavan (2017) critiqued the distribution based on the cut-point used to dichotomize a continuous probability into a binary suitability surface. The authors used a simplified version of the Hahn et al. (2016) model to demonstrate a well-known outcome that arises when continuous probabilities are dichotomized into a binary response (Fielding and Bell 1997). That is, when the probability threshold is reduced, a broader geographic area is classified as suitable and model sensitivity increases. Indeed, when Peterson and Raghavan (2017) increased sensitivity from 78% (Hahn et al. 2016) to 99% (1% omission threshold), the predicted distribution of suitable I. scapularis habitat increased significantly; most areas east of 98.4° W longitude were considered suitable for I. scapularis (Peterson and Raghavan 2017). Ixodes scapularis, a primarily woodland-associated tick, is remarkably well-suited to survive a broad range of climatic conditions (Eisen et al. 2015, 2016), and it is likely that if introduced, could survive at low abundance in most of the eastern United States. However, the distribution map presented by Peterson and Raghavan (2017) provides little guidance on which counties are most likely to be suitable for I. scapularis to establish.

Aedes (Stegomyia) aegypti (L.) and Aedes (Stegomyia) albopictus (Skuse) are potential vectors of Zika, dengue, and chikungunya viruses in the United States. A Zika virus outbreak in Florida in the summer of 2016, driven by Ae. aegypti and resulting in > 200 locally acquired cases of human illness, underscored the need for up-to-date information on the geographic distribution of Ae. aegypti and Ae. albopictus in the United States. In early 2016, we conducted a survey and literature review to compile county records for presence of Ae. aegypti and Ae. albopictus in the United States from 1995 to 2016. Surveillance for these vectors was intensified across the United States during the summer and fall of 2016. At the end of 2016, we therefore conducted a follow-up survey of mosquito control agencies, university researchers, and state and local health departments to document new collection records for Ae. aegypti and Ae. albopictus. The repeated survey at the end of the year added Ae. aegypti collection records from 38 new counties and Ae. albopictus collection records from 127 new counties, representing a 21 and 10 percent increase, respectively, in the number of counties with reported presence of these mosquitoes compared with the previous report. Moreover, through our updated survey, 40 and 183 counties, respectively, added additional years of collection records for Ae. aegypti and Ae. albopictus from 1995 to 2016. Our findings underscore the continued need for systematic surveillance of Ae. aegypti and Ae. albopictus.

The Earth's ecosystems have been altered by anthropogenic processes, including land use, harvesting populations, species introductions and climate change. These anthropogenic processes greatly alter plant and animal communities, thereby changing transmission of the zoonotic pathogens they carry. Biodiversity conservation may be a potential win-win strategy for maintaining ecosystem health and protecting public health, yet the causal evidence to support this strategy is limited. Evaluating conservation as a viable public health intervention requires answering four questions: (i) Is there a general and causal relationship between biodiversity and pathogen transmission, and if so, which direction is it in? (ii) Does increased pathogen diversity with increased host biodiversity result in an increase in total disease burden? (iii) Do the net benefits of biodiversity conservation to human well-being outweigh the benefits that biodiversity-degrading activities, such as agriculture and resource utilization, provide? (iv) Are biodiversity conservation interventions cost-effective when compared to other options employed in standard public health approaches? Here, we summarize current knowledge on biodiversity-zoonotic disease relationships and outline a research plan to address the gaps in our understanding for each of these four questions. Developing practical and self-sustaining biodiversity conservation interventions will require significant investment in disease ecology research to determine when and where they will be effective.This article is part of the themed issue 'Conservation, biodiversity and infectious disease: scientific evidence and policy implications'.

Aedes (Stegomyia) aegypti (L.) and Aedes (Stegomyia) albopictus (Skuse) transmit arboviruses that are increasing threats to human health in the Americas, particularly dengue, chikungunya, and Zika viruses. Epidemics of the associated arboviral diseases have been limited to South and Central America, Mexico, and the Caribbean in the Western Hemisphere, with only minor localized outbreaks in the United States. Nevertheless, accurate and up-to-date information for the geographical ranges of Ae. aegypti and Ae. albopictus in the United States is urgently needed to guide surveillance and enhance control capacity for these mosquitoes. We compiled county records for presence of Ae. aegypti and Ae. albopictus in the United States from 1995-2016, presented here in map format. Records were derived from the Centers for Disease Control and Prevention ArboNET database, VectorMap, the published literature, and a survey of mosquito control agencies, university researchers, and state and local health departments. Between January 1995 and March 2016, 183 counties from 26 states and the District of Columbia reported occurrence of Ae. aegypti, and 1,241 counties from 40 states and the District of Columbia reported occurrence of Ae. albopictus During the same time period, Ae. aegypti was collected in 3 or more years from 94 counties from 14 states and the District of Columbia, and Ae. albopictus was collected during 3 or more years from 514 counties in 34 states and the District of Columbia. Our findings underscore the need for systematic surveillance of Ae. aegypti and Ae. albopictus in the United States and delineate areas with risk for the transmission of these introduced arboviruses.

In addition to serving as vectors of several other human pathogens, the black-legged tick, Ixodes scapularis Say, and western black-legged tick, Ixodes pacificus Cooley and Kohls, are the primary vectors of the spirochete (Borrelia burgdorferi) that causes Lyme disease, the most common vector-borne disease in the United States. Over the past two decades, the geographic range of I. pacificus has changed modestly while, in contrast, the I. scapularis range has expanded substantially, which likely contributes to the concurrent expansion in the distribution of human Lyme disease cases in the Northeastern, North-Central and Mid-Atlantic states. Identifying counties that contain suitable habitat for these ticks that have not yet reported established vector populations can aid in targeting limited vector surveillance resources to areas where tick invasion and potential human risk are likely to occur. We used county-level vector distribution information and ensemble modeling to map the potential distribution of I. scapularis and I. pacificus in the contiguous United States as a function of climate, elevation, and forest cover. Results show that I. pacificus is currently present within much of the range classified by our model as suitable for establishment. In contrast, environmental conditions are suitable for I. scapularis to continue expanding its range into northwestern Minnesota, central and northern Michigan, within the Ohio River Valley, and inland from the southeastern and Gulf coasts. Overall, our ensemble models show suitable habitat for I. scapularis in 441 eastern counties and for I. pacificus in 11 western counties where surveillance records have not yet supported classification of the counties as established.

West Nile virus (WNV) is a leading cause of mosquito-borne disease in the United States. Annual seasonal outbreaks vary in size and location. Predicting where and when higher than normal WNV transmission will occur can help direct limited public health resources. We developed models for the contiguous United States to identify meteorological anomalies associated with above average incidence of WNV neuroinvasive disease from 2004 to 2012. We used county-level WNV data reported to ArboNET and meteorological data from the North American Land Data Assimilation System. As a result of geographic differences in WNV transmission, we divided the United States into East and West, and 10 climate regions. Above average annual temperature was associated with increased likelihood of higher than normal WNV disease incidence, nationally and in most regions. Lower than average annual total precipitation was associated with higher disease incidence in the eastern United States, but the opposite was true in most western regions. Although multiple factors influence WNV transmission, these findings show that anomalies in temperature and precipitation are associated with above average WNV incidence. Readily accessible meteorological data may be used to develop predictive models to forecast geographic areas with elevated WNV disease risk before the coming season.

Flying foxes Pteropus spp. play a key role in forest regeneration as seed dispersers and are also the reservoir of many viruses, including Nipah virus in Bangladesh. Little is known about their habitat requirements, particularly in South Asia. Identifying Pteropus habitat preferences could assist in understanding the risk of zoonotic disease transmission broadly and, in Bangladesh, could help explain the spatial distribution of human Nipah virus cases. 2. We analysed characteristics of Pteropus giganteus roosts and constructed an ecological niche model to identify suitable habitat in Bangladesh. We also assessed the distribution of suitable habitat in relation to the location of human Nipah virus cases. 3. Compared to non-roost trees, P.giganteus roost trees are taller with larger diameters and are more frequently canopy trees. Colony size was larger in densely forested regions and smaller in flood-affected areas. Roosts were located in areas with lower annual precipitation and higher human population density than non-roost sites. 4. We predicted that 2-17% of Bangladesh's land area is suitable roosting habitat. Nipah virus outbreak villages were 2.6 times more likely to be located in areas predicted as highly suitable habitat for P. giganteus compared to non-outbreak villages. 5. Synthesis and applications. Habitat suitability modelling may help identify previously undocumented Nipah outbreak locations and improve our understanding of Nipah virus ecology by highlighting regions where there is suitable bat habitat but no reported human Nipah virus. Conservation and public health education is a key component of P.giganteus management in Bangladesh due to the general misunderstanding and fear of bats that are a reservoir of Nipah virus. Affiliation between Old World fruit bats (Pteropodidae) and people is common throughout their range, and in order to conserve these keystone bat species and prevent emergence of zoonotic viruses, it is imperative that we continue to improve our understanding of Pteropus resource requirements and routes of virus transmission from bats to people. Results presented here can be utilized to develop land management strategies and conservation policies that simultaneously protect fruit bats and public health.

Nipah virus has caused recurring outbreaks in central and northwest Bangladesh (the "Nipah Belt"). Little is known about roosting behavior of the fruit bat reservoir, Pteropus giganteus, or factors driving spillover. We compared human population density and ecological characteristics of case villages and control villages (no reported outbreaks) to understand their role in P. giganteus roosting ecology and Nipah virus spillover risk. Nipah Belt villages have a higher human population density (P < 0.0001), and forests that are more fragmented than elsewhere in Bangladesh (0.50 versus 0.32 patches/km2, P < 0.0001). The number of roosts in a village correlates with forest fragmentation (r = 0.22, P = 0.03). Villages with a roost containing Polyalthia longifolia or Bombax ceiba trees were more likely case villages (odds ratio [OR] = 10.8, 95% confidence interval [CI] = 1.3-90.6). This study suggests that, in addition to human population density, composition and structure of the landscape shared by P. giganteus and humans may influence the geographic distribution of Nipah virus spillovers.