History of the geographic distribution of the western blacklegged tick, Ixodes pacificus, in the United States.

Ixodes pacificus (the western blacklegged tick) occurs in the far western United States (US), where it commonly bites humans. This tick was not considered a species of medical concern until it was implicated in the 1980s as a vector of Lyme disease spirochetes. Later, it was discovered to also be the primary vector to humans in the far western US of agents causing anaplasmosis and hard tick relapsing fever. The core distribution of I. pacificus in the US includes California, western Oregon, and western Washington, with outlier populations reported in Utah and Arizona. In this review, we provide a history of the documented occurrence of I. pacificus in the US from the 1890s to present, and discuss associations of its geographic range with landscape, hosts, and climate. In contrast to Ixodes scapularis (the blacklegged tick) in the eastern US, there is no evidence for a dramatic change in the geographic distribution of I. pacificus over the last half-century. Field surveys in the 1930s and 1940s documented I. pacificus along the Pacific Coast from southern California to northern Washington, in the Sierra Nevada foothills, and in western Utah. County level collection records often included both immatures and adults of I. pacificus, recovered by drag sampling or from humans, domestic animals, and wildlife. The estimated geographic distribution presented for I. pacificus in 1945 by Bishopp and Trembley is similar to that presented in 2022 by the Centers for Disease Control and Prevention. There is no clear evidence of range expansion for I. pacificus, separate from tick records in new areas that could have resulted from newly initiated or intensified surveillance efforts. Moreover, there is no evidence from long-term studies that the density of questing I. pacificus ticks has increased over time in specific areas. It therefore is not surprising that the incidence of Lyme disease has remained stable in the Pacific Coast states from the early 1990s, when it became a notifiable condition, to present. We note that deforestation and deer depredation were less severe in the far western US during the 1800s and early 1900s compared to the eastern US. This likely contributed to I. pacificus maintaining stable, widespread populations across its geographic range in the far western US in the early 1900s, while I. scapularis during the same time period appears to have been restricted to a small number of geographically isolated refugia sites within its present range in the eastern US. The impact that a warming climate may have had on the geographic distribution and local abundance of I. pacificus in recent decades remains unclear.

Efficacy of unregulated minimum risk tick repellent products evaluated with Ixodes scapularis nymphs in a human skin bioassay.

BACKGROUND: The majority of vector-borne disease cases in the USA are caused by pathogens spread by ticks, most commonly the blacklegged tick, Ixodes scapularis. Personal protection against tick bites, including use of repellents, is the primary defense against tick-borne diseases. Tick repellents registered by the Environmental Protection Agency (EPA) are well documented to be safe as well as effective against ticks. Another group of tick repellent products, 25(b) exempt or minimum risk products, use alternative, mostly botanically derived, active ingredients. These are considered to pose minimal risk to human health and therefore are exempt from EPA registration; efficacy testing is not mandated for these products. METHODS: We used a finger bioassay to evaluate the repellency against I. scapularis nymphs for 11 formulated 25(b) exempt products together with two positive control DEET-based EPA registered products. Repellency was assessed hourly from 0.5 to 6.5 h after product application. RESULTS: The DEET-based products showed ≥ 97% repellency for all examined timepoints. By contrast, an average of 63% of ticks were repelled in the first 1.5 h after application across the 11 25(b) exempt products, and the average fell to 3% repelled between 2.5 and 6.5 h. Ten of the 11 25(b) exempt products showed statistically similar efficacy to DEET-based products at 30 min after application (repellency of 79-97%). However, only four 25(b) exempt products maintained a level of repellency similar to DEET-based products (> 72%) at the 1.5-h mark, and none of these products were effective in repelling ticks at the timepoints from 2.5 to 6.5 h after application. CONCLUSIONS: Neither the claims on the labels nor specific active ingredients and their concentrations appeared to predict the duration of efficacy we observed for the 25(b) exempt products. These products are not registered with the EPA, so the methods used to determine the application guidelines on their labels are unclear. Consumers should be aware that both the level of efficacy and the duration of repellency may differ among unregulated 25(b) exempt repellent products labeled for use against ticks. We encourage more research on these products and the 25(b) exempt active ingredients they contain to help determine and improve their efficacy as repellents under different conditions.

Evaluation of the association between climate warming and the spread and proliferation of Ixodes scapularis in northern states in the Eastern United States.

Ixodes scapularis (the blacklegged tick) is widely distributed in forested areas across the eastern United States. The public health impact of I. scapularis is greatest in the north, where nymphal stage ticks commonly bite humans and serve as primary vectors for multiple human pathogens. There were dramatic increases in the tick's distribution and abundance over the last half-century in the northern part of the eastern US, and climate warming is commonly mentioned as a primary driver for these changes. In this review, we summarize the evidence for the observed spread and proliferation of I. scapularis being driven by climate warming. Although laboratory and small-scale field studies have provided insights into how temperature and humidity impact survival and reproduction of I. scapularis, using these associations to predict broad-scale distribution and abundance patterns is more challenging. Numerous efforts have been undertaken to model the distribution and abundance of I. scapularis at state, regional, and global scales based on climate and landscape variables, but outcomes have been ambiguous. Across the models, the functional relationships between seasonal or annual measures of heat, cold, precipitation, or humidity and tick presence or abundance were inconsistent. The contribution of climate relative to landscape variables was poorly defined. Over the last half-century, climate warming occurred in parallel with spread and population increase of the white-tailed deer, the most important reproductive host for I. scapularis adults, in the northern part of the eastern US. There is strong evidence for white-tailed deer playing a key role to facilitate spread and proliferation of I. scapularis in the US over the last century. However, due to a lack of spatially and temporally congruent data, climate, landscape, and host variables are rarely included in the same models, thus limiting the ability to evaluate their relative contributions or interactions in defining the geographic range and abundance patterns of ticks. We conclude that the role of climate change as a key driver for geographic expansion and population increase of I. scapularis in the northern part of the eastern US over the last half-century remains uncertain.

Efficacy of Unregulated Minimum Risk Products to Kill and Repel Ticks.

Human-biting ticks threaten public health in the United States. Registration by the Environmental Protection Agency of products to kill host-seeking ticks or repel ticks contacting humans is indicative of their safety and effectiveness. Unregulated minimum risk products, exempt from Environmental Protection Agency registration and often based on botanical oils, are proliferating in the marketplace, but there is concern about their effectiveness to kill and repel ticks. Evaluations of such products are limited in the published literature. A review showed considerable variability among minimum risk products to kill host-seeking blacklegged ticks, with effectiveness similar to chemical pesticide products for some minimum risk products but minimal impact on the ticks for other products. Evaluations of minimum risk tick repellents have typically focused on individual active ingredients rather than formulated products, which often combine multiple active ingredients. Consumers should be aware that effectiveness to kill and repel ticks can differ among unregulated minimum risk products.

Acceptability of 4-poster deer treatment devices for community-wide tick control among residents of high Lyme disease incidence counties in Connecticut and New York, USA.

The 4-Poster Tick Control Deer Feeder (4-poster) device applies acaricide to white-tailed deer (Odocoileus virginianus) and can reduce populations of the blacklegged tick (Ixodes scapularis), which transmits the agents of Lyme disease, anaplasmosis, babesiosis, and Powassan virus disease in the Northeastern United States. While 4-poster devices have the potential to provide community-wide management of blacklegged ticks in Lyme disease endemic areas, no recent study has assessed their acceptability among residents. We conducted a survey of residents from 16 counties with high annual average Lyme disease incidence (≥ 10 cases per 100,000 persons between 2013 and 2017) in Connecticut and New York to understand perceptions and experiences related to tickborne diseases, support or concerns for placement of 4-poster devices in their community, and opinions on which entities should be responsible for tick control on private properties. Overall, 37% of 1652 respondents (5.5% response rate) would support placement of a 4-poster device on their own property, 71% would support placement on other private land in their community, and 90% would support placement on public land. Respondents who were male, rented their property, resided on larger properties, or were very or extremely concerned about encountering ticks on their property were each more likely to support placement of 4-poster devices on their own property. The primary reason for not supporting placement of a 4-poster device on one's own property was the need for weekly service visits from pest control professionals, whereas the top reason for not supporting placement on other land (private or public) was safety concerns. Most respondents (61%) felt property owners should be responsible for tick control on private properties. Communities considering 4-poster devices as part of a tick management strategy should consider targeting owners of larger properties and placing devices on public lands.

Community-based integrated tick management programs: cost and feasibility scenarios.

Numerous studies have assessed the efficacy of environmentally based control methods to suppress populations of the blacklegged tick (Ixodes scapularis Say), but few of these estimated the cost of control. We estimated costs for a range of tick control methods (including habitat management, deer exclusion or population reduction, broadcast of acaricides, and use of host-targeted acaricides) implemented singly or in combination and applied to a model community comprising 320 residential properties and parklands. Using the high end for cost ranges, tick control based on a single method was estimated to have mean annual costs per household in the model community ranging from $132 for treating only forest ecotone with a broadcast synthetic acaricide to kill host-seeking ticks (or $404 for treating all residential forested habitat) to >$2,000 for deployment of bait boxes (SELECT TCS) across all residential tick habitat to treat rodents topically with acaricide to kill infesting ticks. Combining different sets of multiple methods in an integrated tick management program placed the annual cost between $508 and 3,192 annually per household in the model community, underscoring the disconnect between what people in Lyme disease endemic areas say they are willing to pay for tick control (not more than $100-150 annually) and the actual costs for tick control. Additional barriers to implementing community-based tick management programs within residential communities are discussed.

Changes in the geographic distribution of the blacklegged tick, Ixodes scapularis, in the United States.

Ixodes scapularis (the blacklegged tick) was considered a species of no medical concern until the mid-1970s. By that time, the tick's geographic distribution was thought to be mainly in the southeastern United States (US), with additional localized populations along the Eastern Seaboard north to southern Massachusetts and in the Upper Midwest. Since 1975, I. scapularis has been implicated as a vector of seven human pathogens and is now widely distributed across the eastern US up to the border with Canada. Geographic expansion of tick-borne diseases associated with I. scapularis (e.g., Lyme disease, anaplasmosis, and babesiosis) is attributed to an expanding range of the tick. However, due to changes in tick surveillance efforts over time, it is difficult to differentiate between range expansion and increased recognition of already established tick populations. We provide a history of the documented occurrence of I. scapularis in the US from its description in 1821 to present, emphasizing studies that provide evidence of expansion of the geographic distribution of the tick. Deforestation and decimation of the white-tailed deer (Odocoileus virginianus), the primary reproductive host for I. scapularis adults, during the 1800s presumably led to the tick disappearing from large areas of the eastern US where it previously had been established. Subsequent reforestation and deer population recovery, together with recent climate warming, contributed to I. scapularis proliferating in and spreading from refugia where it had persisted into the early 1900s. From documented tick collection records, it appears I. scapularis was present in numerous locations in the southern part of the eastern US in the early 1900s, whereas in the north it likely was limited to a small number of refugia sites during that time period. There is clear evidence for established populations of I. scapularis in coastal New York and Massachusetts by 1950, and in northwestern Wisconsin by the late 1960s. While recognizing that surveillance for I. scapularis increased dramatically from the 1980s onward, we describe multiple instances of clearly documented expansion of the tick's geographic distribution in the Northeast, Upper Midwest, and Ohio Valley regions from the 1980s to present. Spread and local population increase of I. scapularis, together with documentation of Borrelia burgdorferi sensu stricto in host-seeking ticks, was universally followed by increases in Lyme disease cases in these areas. Southward expansion of northern populations of I. scapularis, for which the host questing behavior of the nymphal stage leads to substantially higher risk of human bites compared with southern populations, into Virginia and North Carolina also was followed by rising numbers of Lyme disease cases. Ongoing surveillance of ticks and tick-borne pathogens is essential to provide the data needed for studies that seek to evaluate the relative roles of land cover, tick hosts, and climate in explaining and predicting geographic expansion of ticks and tick-borne diseases.

Comparison of in vitro and in vivo repellency bioassay methods for Ixodes scapularis nymphs.

BACKGROUND: Numerous bioassay methods have been used to test the efficacy of repellents for ticks, but the comparability of results across different methods has only been evaluated in a single study. Of particular interest are comparisons between bioassays that use artificial containers (in vitro) with those conducted on a human subject (in vivo) for efficacy testing of new potential unregistered active ingredients, which most commonly use in vitro methods. METHODS: We compared four different bioassay methods and evaluated three ingredients (DEET [N,N-Diethyl-meta-toluamide], peppermint oil and rosemary oil) and a negative control (ethanol) over a 6-h period. Two of the methods tested were in vivo bioassay methods in which the active ingredient was applied to human skin (finger and forearm bioassays), and the other two methods were in vitro methods using artificial containers (jar and petri dish bioassays). All four bioassays were conducted using Ixodes scapularis nymphs. We compared the results using nymphs from two different tick colonies that were derived from I. scapularis collected in the US states of Connecticut and Rhode Island (northern origin) and Oklahoma (southern origin), expecting that ticks of different origin would display differences in host-seeking behavior. RESULTS: The results between bioassay methods did not differ significantly, even when comparing those that provide the stimulus of human skin with those that do not. We also found that tick colony source can impact the outcome of repellency bioassays due to differences in movement speed; behavioral differences were incorporated into the assay screening. DEET effectively repelled nymphs for the full 6-h duration of the study. Peppermint oil showed a similar repellent efficacy to DEET during the first hour, but it decreased sharply afterwards. Rosemary oil did not effectively repel nymphs across any of the time points. CONCLUSIONS: The repellency results did not differ significantly between the four bioassay methods tested. The results also highlight the need to consider the geographic origin of ticks used in repellency bioassays in addition to species and life stage. Finally, our results indicate a limited repellent efficacy of the two essential oils tested, which highlights the need for further studies on the duration of repellency for similar botanically derived active ingredients and for evaluation of formulated products.

Rodent-targeted approaches to reduce acarological risk of human exposure to pathogen-infected Ixodes ticks.

In the United States, rodents serve as important hosts of medically important Ixodes ticks, including Ixodes scapularis and Ixodes pacificus, as well as reservoirs for human pathogens, including Anaplasma phagocytophilum, Borrelia burgdorferi sensu stricto (s.s.), and Babesia microti. Over the last four decades, different methods to disrupt enzootic transmission of these pathogens between tick vectors and rodent reservoirs have been developed and evaluated. Early work focused on self-application of topical acaricide by rodents to kill infesting ticks; this resulted in two different types of commercial products based on (i) delivery of permethrin to rodents via impregnated cotton offered as nesting material or (ii) application of fipronil to rodents via an impregnated wick as they navigate through a bait box to reach a food source. More recent work has focused on approaches where acaricides, antibiotics, or a vaccine against Bo. burgdorferi s.s. are delivered orally via rodent food baits. Of these, the oral vaccine and oral acaricide are nearest to commercialization. Other approaches in early stages of development include anti-tick vaccines for rodents and use of heritable genome editing to engineer white-footed mice (Peromyscus leucopus) that are refractory to Bo. burgdorferi s.s. In this review, I first outline general benefits and drawbacks of rodent-targeted tick and pathogen control methods, and then describe the empirical evidence for different approaches to impact enzootic pathogen transmission and acarological risk of human exposure to pathogen-infected Ixodes ticks. Rodent-targeted methods remain promising components of integrated tick management approaches but there are concerns about the robustness of the impact of existing rodent-targeted products across habitats and variable tick host communities, and in some cases also for the implementation cost in relation to what homeowners in Lyme disease endemic areas say they are willing to pay for tick control.

Tick species infesting humans in the United States.

The data for human tick encounters in the United States (US) presented in this paper were compiled with the goals of: (i) presenting quantitative data across the full range of native or recently established human biting ixodid (hard) and argasid (soft) tick species with regards to their frequency of infesting humans, based on published records of ticks collected while biting humans or crawling on clothing or skin; and (ii) providing a guide to publications on human tick encounters. Summary data are presented in table format, and the detailed data these summaries were based on are included in a set of Supplementary Tables. To date, totals of 36 ixodid species (234,722 specimens) and 13 argasid species (230 specimens) have been recorded in the published literature to infest humans in the US. Nationally, the top five ixodid species recorded from humans were the blacklegged tick, Ixodes scapularis (n=158,008 specimens); the lone star tick, Amblyomma americanum (n=36,004); the American dog tick, Dermacentor variabilis (n=26,624); the western blacklegged tick, Ixodes pacificus (n=4,158); and the Rocky Mountain wood tick, Dermacentor andersoni (n=3,518). Additional species with more than 250 ticks recorded from humans included Ixodes cookei (n=2,494); the Pacific Coast tick, Dermacentor occidentalis (n=809); the brown dog tick, Rhipicephalus sanguineus sensu lato (n=714); the winter tick, Dermacentor albipictus (n=465); and the Gulf Coast tick, Amblyomma maculatum (n=335). The spinose ear tick, Otobius megnini (n=69), and the pajaroello tick, Ornithodoros coriaceus (n=55) were the argasid species most commonly recorded from humans. Additional information presented for each of the 49 tick species include a breakdown of life stages recorded from humans, broad geographical distribution in the US, host preference, and associated human pathogens or medical conditions. The paper also provides a history of publications on human tick encounters in the US, with tables outlining publications containing quantitative data on human tick encounters as well as other notable publications on human-tick interactions. Data limitations are discussed. Researchers and public health professionals in possession of unpublished human tick encounter data are strongly encouraged to publish this information in peer-reviewed scientific journals. In future papers, it would be beneficial if data consistently were broken down by tick species and life stage as well as host species and ticks found biting versus crawling on clothing or skin.

A comparison of horizontal and transovarial transmission efficiency of Borrelia miyamotoi by Ixodes scapularis.

Borrelia miyamotoi is a relapsing fever spirochete carried by Ixodes spp. ticks throughout the northern hemisphere. The pathogen is acquired either transovarially (vertically) or horizontally through blood-feeding and passed transtadially across life stages. Despite these complementary modes of transmission, infection prevalence of ticks with B. miyamotoi is typically low (<5%) in natural settings and the relative contributions of the two transmission modes have not been studied extensively. Horizontal transmission of B. miyamotoi (strain CT13-2396 or wild type strain) was initiated using infected Ixodes scapularis larvae or nymphs to expose rodents, which included both the immunocompetent CD-1 laboratory mouse (Mus musculus) and a natural reservoir host, the white-footed mouse (Peromyscus. leucopus), to simulate natural enzootic transmission. Transovarial transmission was evaluated using I. scapularis exposed to B. miyamotoi as either larvae or nymphs feeding on immunocompromised SCID mice (M. musculus) and subsequently fed as females on New Zealand white rabbits. Larvae from infected females were qPCR-tested individually to assess transovarial transmission rates. Tissue tropism of B. miyamotoi in infected ticks was demonstrated using in situ hybridization. Between 1 and 12% of ticks were positive (post-molt) for B. miyamotoi after feeding on groups of CD-1 mice or P. leucopus with evidence of infection, indicating that horizontal transmission was inefficient, regardless of whether infected larvae or nymphs were used to challenge the mice. Transovarial transmission occurred in 7 of 10 egg clutches from infected females. Filial infection prevalence in larvae ranged from 3 to 100% (median 71%). Both larval infection prevalence and spirochete load were highly correlated with maternal spirochete load. Spirochetes were disseminated throughout the tissues of all three stages of unfed ticks, including the salivary glands and female ovarian tissue. The results indicate that while multiple transmission routes contribute to enzootic maintenance of B. miyamotoi, transovarial transmission is likely to be the primary source of infected ticks and therefore risk assessment and tick control strategies should target adult female ticks.

Range Expansion of Native and Invasive Ticks: A Looming Public Health Threat.

Native and invasive tick species pose a serious public health concern in the United States. Range expansion of several medically important tick species has resulted in an increasing number of communities at risk for exposure to ticks and tick-borne pathogens.

Personal protection measures to prevent tick bites in the United States: Knowledge gaps, challenges, and opportunities.

Personal protection measures to prevent human tick encounters from resulting in bites are widely recommended as the first line of defense against health impacts associated with ticks. This includes using repellents, wearing untreated or permethrin-treated protective clothing, and conducting tick checks after coming inside, aided by removing outdoor clothing articles and running them in a dryer on high heat (to kill undetected ticks) and taking a shower/bath (to aid in detecting ticks on the skin). These measures have the benefit of incurring no or low cost, but they need to be used consistently to be most effective. In this paper, I review the level of use (acceptability combined with behavior) of the above-mentioned personal protection measures and their effectiveness to prevent tick bites and tick-borne disease. Studies on the level of use of personal protection measures to prevent tick bites have used different recruitment strategies, focused on different types of respondent populations, employed variable phrasings of survey questions relating to a given personal protection measure, and presented results based on variable frequencies of taking action. This complicates the synthesis of the findings, but the studies collectively indicate that members of the public commonly take action to prevent tick bites, most frequently by wearing untreated protective clothing or conducting tick checks (done routinely by 30 to 70% of respondents in most studies of the public), followed by showering/bathing after being outdoors or using repellents on skin/clothing (15 to 40% range), and with permethrin-treated clothing being the least frequently used tick bite prevention method (<5 to 20% range). A suite of experimental studies have shown that applying repellents or permethrin to coveralls or uniform-style clothing can result in decreased numbers of tick bites, but similar studies are lacking for members of the public wearing summer-weight clothing during normal daily activities. Moreover, a set of case-control and cross-sectional studies have explored associations between use of different personal protection measures to prevent tick bites and Lyme disease or other tick-borne infections. The results are mixed for each personal protection measure, with some studies indicating that regular use of the measure is associated with a reduction in tick-borne disease while other studies found no similar protective effect. One possible interpretation is that these personal protection measures can protect against tick-borne infection but the information gathered to date has not been sufficiently detailed to clarify the circumstances under which protection is achieved, especially with regards to frequency of use, parts of the body being protected, and use of combinations of two or more potentially protective measures. In conclusion, personal protection measures to prevent tick bites are used by the public and merit further study to better understand how they need to be used to have the greatest public health impact.

Designing an Intervention Trial of Human-Tick Encounters and Tick-Borne Diseases in Residential Settings Using 4-Poster Devices to Control Ixodes scapularis (Acari: Ixodidae): Challenges for Site Selection and Device Placement.

Blacklegged ticks, Ixodes scapularis Say, transmit Lyme disease spirochetes and other human pathogens in the eastern United States. White-tailed deer (Odocoileus virginianus) are key reproductive hosts for I. scapularis adults, and therefore control methods targeting deer have the potential for landscape-wide tick suppression. A topical acaricide product, containing 10% permethrin, is self-applied by deer to kill parasitizing ticks when they visit 4-Poster Tick Control Deer Feeders (hereafter, 4-Posters) Previous 4-Poster intervention studies, including in residential settings, demonstrated suppression of I. scapularis populations but did not include human-based outcomes. To prepare for a proposed 4-Poster intervention trial in residential areas of Connecticut and New York that would include human-tick encounters and tick-borne diseases as outcomes, we sought to identify areas (study clusters) in the 80-100 ha size range and specific locations within these areas where 4-Poster devices could be deployed at adequate density (1 device per 20-25 ha) and in accordance with regulatory requirements. Geographic Information System-based data were used to identify prospective study clusters, based on minimum thresholds for Lyme disease incidence, population density, and forest cover. Ground truthing of potential 4-Poster placement locations was done to confirm the suitability of selected clusters. Based on these efforts, we failed to identify more than a few residential areas fulfilling all criteria for a treatment cluster. We, therefore, reconsidered pursuing the intervention trial, which required inclusion of >30 treatment clusters to achieve adequate statistical power. The 4-Poster methodology may be more readily evaluated in natural or public areas than in residential settings in NY or CT.

Knowledge, attitudes, and behaviors regarding tick-borne disease prevention in Lyme disease-endemic areas of the Upper Midwest, United States.

Lyme disease and other tick-borne diseases are a major public health threat in the Upper Midwestern United States, including Michigan, Minnesota, and Wisconsin. To prevent tick bites and tick-borne diseases, public health officials commonly recommend personal protective measures and property management techniques. Adoption of tick-borne disease prevention behaviors and practices by individuals are, however, highly variable. We aimed to characterize current tick-borne disease knowledge, attitudes, and prevention behaviors (KAB) practiced by the public in these states, as well as their willingness to use specific tick control methods. We conducted a population-based survey in summer 2019 in 48 high-risk counties (those having a five-year average (2013-2017) Lyme disease incidence of ≥ 10 cases per 100,000 persons per year), in Michigan, Minnesota, and Wisconsin. A total of 2713 surveys were analyzed; survey weights were used to account for household selection probability and post-stratified to match county-level joint age and sex population distributions in population-level inference. An estimated 98% of the population had heard of Lyme disease, with most perceiving it as very or extremely serious (91%); however, only an estimated 25% perceived tick-borne diseases as very or extremely common in their community. Among those who spent time in places with ticks from April through October, an estimated 68% check themselves thoroughly for ticks most of the time or always and 43% use bug repellent on skin or clothing most of the time or always. An estimated 13% of the population had ever treated their property with a pesticide to kill ticks, and 3% had ever used devices that apply pesticide to rodents to kill ticks on their property. Willingness to practice tick bite prevention behaviors, however, was estimated to be much higher; with 82% being willing to perform tick checks at least once a day, and more than 60% willing to use bug repellent, tick control products on pets, or to bathe within two hours of being outdoors. We found that residents would likely be willing to support a county-wide tick control program to reduce the risk of tick-borne disease in their community (81%) or to apply tick control products to their property to reduce the risk of tick-borne disease in their household (79%). Tick checks were more likely to be practiced among participants who perceived tick-borne diseases to be highly prevalent in their community, if they or a household member had been previously diagnosed with a tick-borne disease?, or if they perceived tick exposure to be likely around their home, cabin, or vacation home. In addition, property-based tick control methods were associated with perceived risk of encountering ticks around the home, cabin, or vacation home. Participants who had seen information from state health departments were also more likely to practice preventive measures. The most common reported barriers to using any of these methods were forgetfulness, safety concerns, and lack of awareness. Our survey findings shed light on how residents from these Upper Midwest states may adopt tick control and tick bite prevention measures and how public health outreach may be most effective for this population.

Spatial Heterogeneity of Sympatric Tick Species and Tick-Borne Pathogens Emphasizes the Need for Surveillance for Effective Tick Control.

Three tick species that can transmit pathogen causing disease are commonly found parasitizing people and animals in the mid-Atlantic United States: the blacklegged tick (Ixodes scapularis Say), the American dog tick (Dermacentor variabilis [Say]), and the lone star tick (Amblyomma americanum [L.]) (Acari: Ixodidae). The potential risk of pathogen transmission from tick bites acquired at schools in tick-endemic areas is a concern, as school-aged children are a high-risk group for tick-borne disease. Integrated pest management (IPM) is often required in school districts, and continued tick range expansion and population growth will likely necessitate IPM strategies to manage ticks on school grounds. However, an often-overlooked step of tick management is monitoring and assessment of local tick species assemblages to inform the selection of control methodologies. The purpose of this study was to evaluate tick species presence, abundance, and distribution and the prevalence of tick-borne pathogens in both questing ticks and those removed from rodent hosts on six school properties in Maryland. Overall, there was extensive heterogeneity in tick species dominance, abundance, and evenness across the field sites. A. americanum and I. scapularis were found on all sites in all years. Overall, A. americanum was the dominant tick species. D. variabilis was collected in limited numbers. Several pathogens were found in both questing ticks and those removed from rodent hosts, although prevalence of infection was not consistent between years. Borrelia burgdorferi, Ehrlichia chaffeensis, Ehrlichia ewingii, and Ehrlichia "Panola Mountain" were identified in questing ticks, and B. burgdorferi and Borrelia miyamotoi were detected in trapped Peromyscus spp. mice. B. burgdorferi was the dominant pathogen detected. The impact of tick diversity on IPM of ticks is discussed.

Detection of Genetic Variability in Borrelia miyamotoi (Spirochaetales: Spirochaetaceae) Between and Within the Eastern and Western United States.

Borrelia miyamotoi is a hard tick-associated relapsing fever spirochete that is geographically widespread in Ixodes spp. (Acari: Ixodidae) ticks, but typically occurs at low prevalence. Genetic variability has been described among strains derived from Asia, Europe, and North America, and among tick species that carry the infection, but little variability has been described within foci or tick species. Capitalizing on access to B. miyamotoi nucleic acid extracted from host-seeking Ixodes scapularis Say or Ixodes pacificus Cooley & Kohls from 16 states, we explored genetic variability based on sequence analysis of four amplicons described herein. Consistent with previous studies, we detected significant genetic differences between strains derived from I. scapularis (eastern United States) and I. pacificus (western United States) and identified two distinct sequences in the western United States (Am-West-1 and Am-West-2). Unique to this study, we identified two distinct sequences in the eastern United States (Am-East-1 and Am-East-2). Based on the 161 samples we analyzed, Am-East-1 was the only type represented in 50 B. miyamotoi-infected ticks collected from the Northeast (Vermont, Maine, New York, Connecticut, and Rhode Island), whereas ticks collected from the North-Central and Mid-Atlantic states harbored B. miyamotoi comprised of both Am-East-1 and Am-East-2. Further studies are needed to better characterize the phylogeography of B. miyamotoi and to discern if there are biologically meaningful differences among sequence types. To facilitate further exploration, we developed a polymerase chain reaction (PCR) assay designed to differentiate Am-East-1, Am-East-2, and Am-West sequence types without having to sequence the amplicon.

Control of ixodid ticks and prevention of tick-borne diseases in the United States: The prospect of a new Lyme disease vaccine and the continuing problem with tick exposure on residential properties.

In the United States, exposure to human-biting ixodid ticks can occur while spending time on residential properties or in neighborhood green spaces as well as during recreational or occupational activities on public lands. Human-biting tick species collectively transmit >15 species of pathogenic microorganisms and the national burden of tick-borne diseases is increasing. The prospect of a new Lyme disease vaccine for use in humans provides hope for substantial reduction in the >450,000 estimated annual cases of Lyme disease but this breakthrough would not reduce cases of other tick-borne diseases, such as anaplasmosis, babesiosis, ehrlichiosis, spotted fever group rickettsiosis, and Powassan encephalitis. One intriguing question is to what extent a new Lyme disease vaccine would impact the use of personal protection measures acting broadly against tick-bites. The main tick vector for Lyme disease spirochetes in the eastern United States, Ixodes scapularis, also transmits causative agents of anaplasmosis, babesiosis, and Powassan encephalitis; and this tick species co-occurs with other human-biting vectors such as Amblyomma americanum and Dermacentor variabilis. It therefore is important that a new Lyme disease vaccine does not result in reduced use of tick-bite prevention measures, such as tick repellents, permethrin-treated clothing, and frequent tick checks. Another key issue is the continuing problem with tick exposure on residential properties, which represents a heavily used outdoor environment the residents cannot reasonably avoid and where they tend to spend large amounts of time outside. As it may not be realistic to keep up daily vigilance with personal protective measures against tick-bites on residential properties during many months of every year, homeowners may also consider the option to suppress host-seeking ticks by means of deer fencing, landscaping, vegetation management, and use of products to kill host-seeking ticks or ticks infesting rodents. When considering the full range of options for actions that can be taken to suppress host-seeking ticks on residential properties, it is clear that individual homeowners face a difficult and bewildering task in deciding what to do based on very general guidance from public health agencies (developed without the benefit of a strong evidence base) and often without ready access to local public health professionals experienced in tick control. This situation is not satisfactory but cannot be corrected without first addressing knowledge gaps regarding the impact of peridomestic tick control measures on host-seeking ticks, human tick-bites, and tick-borne diseases. In parallel with this effort, there also is a need to increase the local public health workforce with knowledge of and experience with tick control to provide better access for homeowners to sound and objective advice regarding tick control on their properties based on key characteristics of the landscaping, habitat composition, and use patterns by wild animal tick hosts as well as the residents.

Surveillance of Ticks and Tick-Borne Pathogens in Suburban Natural Habitats of Central Maryland.

Lyme and other tick-borne diseases are increasing in the eastern United States and there is a lack of research on integrated strategies to control tick vectors. Here we present results of a study on tick-borne pathogens detected from tick vectors and rodent reservoirs from an ongoing 5-yr tick suppression study in the Lyme disease-endemic state of Maryland, where human-biting tick species, including Ixodes scapularis Say (Acari: Ixodidae) (the primary vector of Lyme disease spirochetes), are abundant. During the 2017 tick season, we collected 207 questing ticks and 602 ticks recovered from 327 mice (Peromyscus spp. (Rodentia: Cricetidae)), together with blood and ear tissue from the mice, at seven suburban parks in Howard County. Ticks were selectively tested for the presence of the causative agents of Lyme disease (Borrelia burgdorferi sensu lato [s.l.]), anaplasmosis (Anaplasma phagocytophilum), babesiosis (Babesia microti), ehrlichiosis (Ehrlichia ewingii, Ehrlichia chaffeensis, and 'Panola Mountain' Ehrlichia) and spotted fever group rickettsiosis (Rickettsia spp.). Peromyscus ear tissue and blood samples were tested for Bo. burgdorferi sensu stricto (s.s), A. phagocytophilum, Ba. microti, and Borrelia miyamotoi. We found 13.6% (15/110) of questing I. scapularis nymphs to be Bo. burgdorferi s.l. positive and 1.8% (2/110) were A. phagocytophilum positive among all sites. Borrelia burgdorferi s.s. was found in 71.1% (54/76) of I. scapularis nymphs removed from mice and 58.8% (194/330) of captured mice. Results from study on tick abundance and pathogen infection status in questing ticks, rodent reservoirs, and ticks feeding on Peromyscus spp. will aid efficacy evaluation of the integrated tick management measures being implemented.