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In recent decades, tickborne disease (TBD) cases and established populations of medically important ticks have been reported over expanding geographic areas, and an increasing number of tickborne bacteria, viruses, and protozoans have been recognized as human pathogens, collectively contributing to an increasing burden of TBDs in the United States.The prevention and diagnosis of TBDs depend greatly on an accurate understanding by the public and healthcare providers of when and where persons are at risk for exposure to human-biting ticks and to the pathogens these ticks transmit. However, national maps showing the distributions of medically important ticks and the presence or prevalence of tickborne pathogens are often incomplete, outdated, or lacking entirely. Similar deficiencies exist regarding geographic variability in host-seeking tick abundance. Efforts to accurately depict acarological risk are hampered by lack of systematic and routine surveillance for medically important ticks and their associated human pathogens. In this review, we: 1) outline the public health importance of tick surveillance; 2) identify gaps in knowledge regarding the distributions and abundance of medically important ticks in the United States and the presence and prevalence of their associated pathogens; 3) describe key objectives for tick surveillance and review methods appropriate for addressing those goals; and 4) assess current capacity and barriers to implementation and sustainability of tick surveillance programs.

Background The vast majority of vector-borne diseases in the USA are associated with mosquitoes or ticks. Mosquito control is often conducted as part of community programs run by publicly-funded entities. By contrast, tick control focuses primarily on individual residential properties and is implemented predominantly by homeowners and the private pest control firms they contract. We surveyed publicly-funded vector control programs (VCPs), presumed to focus mainly on mosquitoes, to determine what tick-related services they currently offer, and their interest in and capacity to expand existing services or provide new ones. Methods We distributed a survey to VCPs in the Northeast, Upper Midwest and Pacific Coast states of the USA, where humans are at risk for bites by tick vectors ( Ixodes scapularis or Ixodes pacificus ) of agents causing Lyme disease and other tick-borne diseases. The data we report are based on responses from 118 VCPs engaged in vector control and with at least some activities focused on ticks. Results Despite our survey targeting geographic regions where ticks and tick-borne diseases are persistent and increasing public health concerns, only 11% (12/114) of VCPs reported they took direct action to suppress ticks questing in the environment. The most common tick-related activities conducted by the VCPs were tick bite prevention education for the public (70%; 75/107 VCPs) and tick surveillance (48%; 56/116). When asked which services they would most likely include as part of a comprehensive tick management program, tick bite prevention education (90%; 96/107), tick surveillance (89%; 95/107) and tick suppression guidance for the public (74%; 79/107) were the most common services selected. Most VCPs were also willing to consider engaging in activities to suppress ticks on public lands (68%; 73/107), but few were willing to consider suppressing ticks on privately owned land such as residential properties (15%; 16/107). Across all potential tick-related services, funding was reported as the biggest obstacle to program expansion or development, followed by personnel. Conclusions Considering the hesitancy of VCPs to provide tick suppression services on private properties and the high risk for tick bites in peridomestic settings, suppression of ticks on residential properties by private pest control operators will likely play an important role in the tick suppression landscape in the USA for the foreseeable future. Nevertheless, VCPs can assist in this effort by providing locally relevant guidelines to homeowners and private pest control firms regarding best practices for residential tick suppression efforts and associated efficacy evaluations. Publicly-funded VCPs are also well positioned to educate the public on personal tick bite prevention measures and to collect tick surveillance data that provide information on the risk of human encounters with ticks within their jurisdictions. Graphical Abstract

First identified in two Missouri farmers exhibiting low white-blood-cell and platelet counts in 2009, Heartland virus (HRTV) is genetically closely related to severe fever with thrombocytopenia syndrome virus (SFTSV), a tick-borne phlebovirus producing similar symptoms in China, Korea, and Japan. Field isolations of HRTV from several life stages of unfed, host-seeking Amblyomma americanum, the lone star tick, implicated it as a putative vector capable of transstadial transmission. Laboratory vector competence assessments confirmed transstadial transmission of HRTV, demonstrated vertical infection, and showed co-feeding infection between A. americanum. A vertical infection rate of 33% from adult females to larvae in the laboratory was observed, while only one of 386 pools of molted nymphs (1930) reared from co-feeding larvae was positive for HRTV (maximum-likelihood estimate of infection rate = 0.52/1000). Over 35 human HRTV cases, all within the distribution range of A. americanum, have been documented. Serological testing of wildlife in areas near the index human cases, as well as in widely separated regions of the eastern United States where A. americanum occur, indicated many potential hosts such as raccoons and white-tailed deer. Attempts, however, to experimentally infect mice, rabbits, hamsters, chickens, raccoons, goats, and deer failed to produce detectable viremia. Immune-compromised mice and hamsters are the only susceptible models. Vertical infection augmented by co-feeding transmission could play a role in maintaining the virus in nature. A more complete assessment of the natural transmission cycle of HRTV coupled with serosurveys and enhanced HRTV disease surveillance are needed to better understand transmission dynamics and human health risks.

Aedes (Stegomyia) aegypti (L.) and Ae. (Stegomyia) albopictus (Skuse) mosquitoes can transmit dengue, chikungunya, yellow fever, and Zika viruses. Limited surveillance has led to uncertainty regarding the geographic ranges of these vectors globally, and particularly in regions at the present-day margins of habitat suitability such as the contiguous United States. Empirical habitat suitability models based on environmental conditions can augment surveillance gaps to describe the estimated potential species ranges, but model accuracy is unclear. We identified previously published regional and global habitat suitability models for Ae. aegypti (n = 6) and Ae. albopictus (n = 8) for which adequate information was available to reproduce the models for the contiguous U.S. Using a training subset of recently updated county-level surveillance records of Ae. aegypti and Ae. albopictus and records of counties conducting surveillance, we constructed accuracy-weighted, probabilistic ensemble models from these base models. To assess accuracy and uncertainty we compared individual and ensemble model predictions of species presence or absence to both training and testing data. The ensemble models were among the most accurate and also provided calibrated probabilities of presence for each species. The quantitative probabilistic framework enabled identification of areas with high uncertainty and model bias across the U.S. where improved models or additional data could be most beneficial. The results may be of immediate utility for counties considering surveillance and control programs for Ae. aegypti and Ae. albopictus. Moreover, the assessment framework can drive future efforts to provide validated quantitative estimates to support these programs at local, national, and international scales.

Although Borrelia burgdorferi sensu lato (s.l.) are found in a great diversity of vertebrates, most studies in North America have focused on the role of mammals as spirochete reservoir hosts. We investigated the roles of birds as hosts for subadult Ixodes pacificus ticks and potential reservoirs of the Lyme disease spirochete B. burgdorferi sensu stricto (s.s.) in northwestern California. Overall, 623 birds representing 53 species yielded 284 I. pacificus larvae and nymphs. We used generalized linear models and zero-inflated negative binomial models to determine associations of bird behaviors, taxonomic relationships and infestation by I. pacificus with borrelial infection in the birds. Infection status in birds was best explained by taxonomic order, number of infesting nymphs, sampling year, and log-transformed average body weight. Presence and counts of larvae and nymphs could be predicted by ground- or bark-foraging behavior and contact with dense oak woodland. Molecular analysis yielded the first reported detection of Borrelia bissettii in birds. Moreover, our data suggest that the Golden-crowned Sparrow (Zonotrichia atricapilla), a non-resident species, could be an important reservoir for B. burgdorferi s.s. Of 12 individual birds (9 species) that carried B. burgdorferi s.l.-infected larvae, no birds carried the same genospecies of B. burgdorferi s.l. in their blood as were present in the infected larvae removed from them. Possible reasons for this discrepancy are discussed. Our study is the first to explicitly incorporate both taxonomic relationships and behaviors as predictor variables to identify putative avian reservoirs of B. burgdorferi s.l. Our findings underscore the importance of bird behavior to explain local tick infestation and Borrelia infection in these animals, and suggest the potential for bird-mediated geographic spread of vector ticks and spirochetes in the far-western United States.

We collected questing Haemaphysalis longicornis ticks from southeastern counties of Pennsylvania, USA. Of 263 ticks tested by PCR for pathogens, 1 adult female was positive for Borrelia burgdorferi sensu stricto, yielding a 0.4% infection rate. Continued monitoring of this invasive tick is essential to determine its public health role.

Tick-borne diseases are increasing in North America. Knowledge of which tick species and associated human pathogens are present locally can inform the public and medical community about the acarological risk for tick bites and tick-borne infections. Citizen science (also called community-based monitoring, volunteer monitoring, or participatory science) is emerging as a potential approach to complement traditional tick record data gathering where all aspects of the work is done by researchers or public health professionals. One key question is how citizen science can best be used to generate high-quality data to fill knowledge gaps that are difficult to address using traditional data gathering approaches. Citizen science is particularly useful to generate information on human–tick encounters and may also contribute to geographical tick records to help define species distributions across large areas. Previous citizen science projects have utilized three distinct tick record data gathering methods including submission of: 1) physical tick specimens for identification by professional entomologists, 2) digital images of ticks for identification by professional entomologists, and 3) data where the tick species and life stage were identified by the citizen scientist. We explore the benefits and drawbacks of citizen science, relative to the traditional scientific approach, to generate data on tick records, with special emphasis on data quality for species identification and tick encounter locations. We recognize the value of citizen science to tick research but caution that the generated information must be interpreted cautiously with data quality limitations firmly in mind to avoid misleading conclusions.

Background Ixodes scapularis and Ixodes pacificus are important vectors of multiple pathogens in the United States. However, their role in transmission of Bartonella spp., which are commonly reported in rodents and fleas, has been debated. Our previous investigation on Bartonella spp. in host-seeking I. scapularis and I. pacificus showed Bartonella spp. were absent in the ticks, suggesting the two species are unlikely to contribute to Bartonella transmission. It is unclear whether the absence of Bartonella spp. in the host-seeking ticks was attributable to ticks not being exposed to Bartonella in nature or being exposed but unable to acquire or transstadially transmit the bacterium. To assess the likelihood of exposure and acquisition, we tested Ixodes spp. ticks collected from rodents for Bartonella infections. Methods Blood-fed I. scapularis ticks ( n  = 792; consisting of 645 larvae and 147 nymphs), I. pacificus ticks ( n  = 45, all larvae), and Ixodes angustus ticks ( n  = 16, consisting of 11 larvae and 5 nymphs) collected from rodents from Minnesota and Washington were tested for Bartonella spp. using a quadruplex polymerase chain reaction (PCR) amplicon next-generation sequencing approach that targets Bartonella -specific fragments on gltA , ssrA , rpoB , and groEL. In parallel, rodents and fleas collected from the same field studies were investigated to compare the differences of Bartonella distribution among the ticks, fleas, and rodents. Results Bartonella spp. were commonly detected in rodents and fleas, with prevalence of 25.6% in rodents and 36.8% in fleas from Minnesota; 27.9% in rodents and 45.2% in fleas from Washington. Of all tested ticks, Bartonella DNA was detected by gltA in only one larval I. scapularis tick from Minnesota. Conclusions The high prevalence of Bartonella spp. in rodents and fleas coupled with extremely low prevalence of Bartonella spp. in blood-fed ticks suggests that although Ixodes ticks commonly encounter Bartonella in rodents, they rarely acquire the infection through blood feeding. Notably, ticks were at various stages of feeding on rodents when they were collected. Laboratory transmission studies are needed to assess acquisition rates in fully blood-fed ticks and to assess transstadial transmission efficiency if ticks acquire Bartonella infections from feeding to repletion. Graphical Abstract

Background Bartonella spp. infect a variety of vertebrates throughout the world, with generally high prevalence. Several Bartonella spp. are known to cause diverse clinical manifestations in humans and have been recognized as emerging pathogens. These bacteria are mainly transmitted by blood-sucking arthropods, such as fleas and lice. The role of ticks in the transmission of Bartonella spp. is unclear. Methods A recently developed quadruplex polymerase chain reaction (PCR) amplicon next-generation sequencing approach that targets Bartonella -specific fragments on gltA , ssrA , rpoB , and groEL was applied to test host-seeking Ixodes scapularis ticks ( n  = 1641; consisting of 886 nymphs and 755 adults) collected in 23 states of the eastern half of the United States and Ixodes pacificus ticks ( n  = 966; all nymphs) collected in California in the western United States for the presence of Bartonella DNA. These species were selected because they are common human biters and serve as vectors of pathogens causing the greatest number of vector-borne diseases in the United States. Results No Bartonella DNA was detected in any of the ticks tested by any target. Conclusions Owing to the lack of Bartonella detection in a large number of host-seeking Ixodes spp. ticks tested across a broad geographical region, our results strongly suggest that I. scapularis and I. pacificus are unlikely to contribute more than minimally, if at all, to the transmission of Bartonella spp. Graphical Abstract

The Asian longhorned tick, Haemaphysalis longicornis, an invasive tick species in the United States, has been found actively host-seeking while infected with several human pathogens. Recent work has recovered large numbers of partially engorged, host-seeking H. longicornis, which together with infection findings raises the question of whether such ticks can reattach to a host and transmit pathogens while taking additional bloodmeals. Here we conducted molecular blood meal analysis in tandem with pathogen screening of partially engorged, host-seeking H. longicornis to identify feeding sources and more inclusively characterize acarological risk. Active, statewide surveillance in Pennsylvania from 2020 to 2021 resulted in the recovery of 22/1,425 (1.5%) partially engorged, host-seeking nymphal and 5/163 (3.1%) female H. longicornis. Pathogen testing of engorged nymphs detected 2 specimens positive for Borrelia burgdorferi sensu lato, 2 for Babesia microti, and 1 co-infected with Bo. burgdorferi s.l. and Ba. microti. No female specimens tested positive for pathogens. Conventional PCR blood meal analysis of H. longicornis nymphs detected avian and mammalian hosts in 3 and 18 specimens, respectively. Mammalian blood was detected in all H. longicornis female specimens. Only 2 H. longicornis nymphs produced viable sequencing results and were determined to have fed on black-crowned night heron, Nycticorax nycticorax. These data are the first to molecularly confirm H. longicornis partial blood meals from vertebrate hosts and Ba. microti infection and co-infection with Bo. burgdorferi s.l. in host-seeking specimens in the United States, and the data help characterize important determinants indirectly affecting vectorial capacity. Repeated blood meals within a life stage by pathogen-infected ticks suggest that an understanding of the vector potential of invasive H. longicornis populations may be incomplete without data on their natural host-seeking behaviors and blood-feeding patterns in nature.