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BackgroundUnderstanding the geographic distribution ofRickettsia montanensisinfections inDermacentor variabilisis important for tick-borne disease management in the United States, as both a tick-borne agent of interest and a potential confounder in surveillance of other rickettsial diseases. Two previous studies modeled niche suitability forD. variabiliswith and withoutR. montanensis, from 2002-2012, indicating that theD. variabilisniche overestimates the infected niche. This study updates these, adding data since 2012.MethodsNewer surveillance and testing data were used to update Species Distribution Models (SDMs) ofD. variabilis, andR. montanensisinfectedD. variabilis, in the United States. Using random forest (RF) models, found to perform best in previous work, we updated the SDMs and compared them with prior results. Warren’s I niche overlap metric was used to compare between predicted suitability for all ticks and ‘pathogen positive niche’ models across datasets.ResultsWarren’s I indicated D. variabilisorR. montanensispositive niche. The updatedD. variabilisniche model overpredicted suitability compared to the updatedR. montanensispositive niche in key peripheral parts of the range, but slightly underpredicted through the northern and midwestern parts of the range. This reinforces previous findings of a more constrained pathogen-positive niche than predicted byD. variabilisrecords alone.ConclusionsThe consistency of predicted niche suitability forD. variabilisin the United States, with the addition of nearly a decade of new data, corroborates this is a species with generalist habitat requirements. Yet a slight shift in updated niche distribution, even of low suitability, included more southern areas, pointing to a need for continued and extended monitoring and surveillance. This further underscores the importance of revisiting vector and vector-borne disease distribution maps.

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.

Lyme disease is the most commonly reported vector-borne disease in the United States, and the number of cases reported each year continues to rise. The complex nature of the relationships between the pathogen (Borrelia burgdorferi sensu stricto), the tick vector (Ixodes scapularis Say), multiple vertebrate hosts, and numerous environmental factors creates challenges for understanding and predicting tick population and pathogen transmission dynamics. LYMESIM is a mechanistic model developed in the late 1990s to simulate the life-history of I. scapularis and transmission dynamics of B. burgdorferi s.s. Here we present LYMESIM 2.0, a modernized version of LYMESIM, that includes several modifications to enhance the biological realism of the model and to generate outcomes that are more readily measured under field conditions. The model is tested for three geographically distinct locations in New York, Minnesota, and Virginia. Model-simulated timing and densities of questing nymphs, infected nymphs, and abundances of nymphs feeding on hosts are consistent with field observations and reports for these locations. Sensitivity analysis highlighted the importance of temperature in host finding for the density of nymphs, the importance of transmission from small mammals to ticks on the density of infected nymphs, and temperature-related tick survival for both density of nymphs and infected nymphs. A key challenge for accurate modeling of these metrics is the need for regionally representative inputs for host populations and their fluctuations. LYMESIM 2.0 is a useful public health tool that downstream can be used to evaluate tick control interventions and can be adapted for other ticks and pathogens.

During September–December 2018, 25 live ticks were collected on-post at Fort Leavenworth, Kansas, in a home with a history of bat occupancy. Nine ticks were sent to the Army Public Health Center Tick-Borne Disease Laboratory and were identified as Carios kelleyi (Cooley and Kohls, 1941), a species that seldom bites humans but that may search for other sources of blood meals, including humans, when bats are removed from human dwellings. The ticks were tested for numerous agents of human disease. Rickettsia lusitaniae was identified by multilocus sequence typing to be present in two ticks, marking the first detection of this Rickettsia agent in the United States and in this species of tick. Two other Rickettsia spp. were also detected, including an endosymbiont previously associated with C. kelleyi and a possible novel Rickettsia species. The potential roles of C. kelleyi and bats in peridomestic Rickettsia transmission cycles warrant further investigation.

Adult females and males of Ixodes affinis and Ixodes scapularis are illustrated by focus stacking image photography, and morphological character states are described that reliably differentiate the two species. In conjunction with other environmental cues, such as the questing phenology of adults, these characteristics will enable the rapid identification of adults of either sex along the southern Coastal Plain of the United States, where these species are sympatric.

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.

The ixodid tick species Ixodes affinis is expanding its range northward, changing the tick community population dynamics in the Mid-Atlantic United States. We present five years of surveillance on newly established populations of I. affinis throughout southeastern Virginia and discuss the habitat and host associations of I. affinis in this northernmost extent of its range. We found that I. affinis populations tend to persist once they are established, and populations tend to increase as ecological succession progresses, provided a vegetated understory persists. Populations of I. affinis were never found in the smallest habitat fragments or in xeric dune habitats, and the highest densities of I. affinis were found in mixed pine-hardwood forests with an herbaceous understory. We also document several new mammalian hosts for I. affinis, including house mice (Mus musculus) and coyotes (Canis latrans) and discuss how these hosts may facilitate the continued dispersal of I. affinis and the maintenance of these newly established populations.

Small mammals are often parasitized by the immature stages of hard-bodied ticks (family Ixodidae) and may serve as reservoir hosts of tick-borne pathogens. Amblyomma maculatum, the Gulf Coast tick, is the primary vector of Rickettsia parkeri, the causative agent of R. parkeri rickettsiosis. This hard-bodied tick species is expanding its historical range from the Gulf Coast of the U.S. up the Mid-Atlantic coast. In Mid-Atlantic states, such as Virginia, R. parkeri prevalence is higher in these ticks than those found in its historical range. This high prevalence may be explained in part by small mammal populations. In this study, small mammals were trapped and checked for the presence of immature A. maculatum. The ticks as well as tissue samples from these mammals were tested for the presence of R. parkeri. This study found six rodent species acting as hosts to immature A. maculatum and three species that may play a role in the enzootic cycle of R. parkeri in Virginia.

In the early 1980s, Ixodes spp. ticks were implicated as the key North American vectors of Borrelia burgdorferi (Johnson, Schmid, Hyde, Steigerwalt and Brenner) (Spirocheatales: Spirochaetaceae), the etiological agent of Lyme disease. Concurrently, other human-biting tick species were investigated as potential B. burgdorferi vectors. Rashes thought to be erythema migrans were observed in patients bitten by Amblyomma americanum (L.) (Acari: Ixodidae) ticks, and spirochetes were visualized in a small percentage of A. americanum using fluorescent antibody staining methods, sparking interest in this species as a candidate vector of B. burgdorferi. Using molecular methods, the spirochetes were subsequently described as Borrelia lonestari sp. nov. (Spirocheatales: Spirochaetaceae), a transovarially transmitted relapsing fever Borrelia of uncertain clinical significance. In total, 54 surveys from more than 35 research groups, involving more than 52,000 ticks, have revealed a low prevalence of B. lonestari, and scarce B. burgdorferi, in A. americanum. In Lyme disease-endemic areas, A. americanum commonly feeds on B. burgdorferi-infected hosts; the extremely low prevalence of B. burgdorferi in this tick results from a saliva barrier to acquiring infection from infected hosts. At least nine transmission experiments involving B. burgdorferi in A. americanum have failed to demonstrate vector competency. Advancements in molecular analysis strongly suggest that initial reports of B. burgdorferi in A. americanum across many states were misidentified B. lonestari, or DNA contamination, yet the early reports continue to be cited without regard to the later clarifying studies. In this article, the surveillance and vector competency studies of B. burgdorferi in A. americanum are reviewed, and we conclude that A. americanum is not a vector of B. burgdorferi.

Two species of ixodid tick, Ixodes affinis Neumann and Amblyomma maculatum Koch, are simultaneously expanding their ranges throughout the mid-Atlantic region of the US. Although we have some understanding of the ecology and life history of these species, the ecological mechanisms governing where and how new populations establish and persist are unclear. To assess population connectivity and ancestry, we sequenced a fragment of the 16S mitochondrial rRNA gene from a representative sample of individuals of both species from populations throughout the eastern US. We found that despite overlapping host preferences throughout ontogeny, each species exhibited very different genetic and geographic patterns of population establishment and connectivity. Ixodes affinis was of two distinct mitochondrial clades, with a clear geographic break separating northern and southern populations. Both I. affinis populations showed evidence of recent expansion, although the southern population was more genetically diverse, indicating a longer history of establishment. Amblyomma maculatum exhibited diverse haplotypes that showed no significant relationship with geographic patterns and little apparent connectivity between sites. Heteroplasmy was also observed in the 16S mitochondrial rRNA gene in 3.5% of A. maculatum individuals. Genetic evidence suggests that these species rely on different key life stages to successfully disperse into novel environments, and that host vagility, habitat stability and habitat connectivity all play critical roles in the establishment of new tick populations.

The hispid cotton rat (Sigmodon hispidus) is a common field rodent in the southeastern United States, where volunteer loblolly pine trees (Pinus taeda) rapidly invade open space and accelerate ecological succession from field to pine forest. While using capture-mark-recapture methods in studies of small mammal populations, we observed dead pine trees in late summer, all killed by girdling, prompting us to determine the extent of damage on a 1.26 ha grid in a 5 y old successional pine forest in southeastern Virginia. We recorded damage to 65% of >15,000 trees, of which 2064 were killed by rodents. Areas with a high density of trees 11–40 mm in diameter (3–5 y old) were most actively damaged during late winter and early spring, following autumnal population peaks of cotton rats. Old field habitats undergoing succession to loblolly pine forest may briefly offer optimal habitat for cotton rats, and girdling by cotton rats may regulate the rate of succession depending on densities of trees and rodents.

Ticks are the most significant vectors of infectious diseases in the United States, inspiring many researchers to study aspects of their biology, ecology, and their effects on public health. However, regional differences in tick abundance and pathogen infection prevalence result in the inability to assume results from one area are relevant in another. Current local information on tick ranges, infection rates, and human cases is needed to assess tick-borne disease risk in any given region. The Mid-Atlantic Tick Summit III brought together over 100 area experts and researchers to share regional updates on ticks and their associated pathogens. We report some meeting highlights here. Regional meetings foster cross-disciplinary collaborations that benefit the community, and open novel lines of inquiry so that tick-bite risk can be reduced and tick-borne diseases can be treated effectively.

The predominant human-biting tick throughout the southeastern United States is Amblyomma americanum . Its ability to transmit pathogens causing Lyme disease-like illnesses is a subject of ongoing controversy. Results of previous testing by the Department of Defense Human Tick Test Kit Program and other laboratories indicated that it is highly unlikely that A. americanum transmits any pathogen that causes Lyme disease. In contrast, a recent publication by Clark and colleagues (K. L. Clark, B. Leydet, and S. Hartman, Int. J. Med. Sci. 10:915–931, 2013) reported detection of Lyme group Borrelia in A. americanum using a nested-flagellin-gene PCR. We evaluated this assay by using it and other assays to test 1,097 A. americanum ticks collected from humans. Using the Clark assay, in most samples we observed nonspecific amplification and nonrepeatability of results on subsequent testing of samples. Lack of reaction specificity and repeatability is consistent with mispriming, likely due to high primer concentrations and low annealing temperatures in this protocol. In six suspect-positive samples, Borrelia lonestari was identified by sequencing of an independent gene region; this is not a Lyme group spirochete and is not considered zoonotic. B. burgdorferi was weakly amplified from one pool using some assays, but not others, and attempts to sequence the amplicon of this pool failed, as did attempts to amplify and sequence B. burgdorferi from the five individual samples comprising this pool. Therefore, B. burgdorferi was not confirmed in any sample. Our results do not support the hypothesis that A. americanum ticks are a vector for Lyme group Borrelia infections.

The Gulf Coast tick Amblyomma maculatum Koch is increasingly relevant to medical and veterinary communities as human infection rates of Rickettsia parkeri rise, the risk of introduction of Ehrlichia ruminantium increases, and the range of this tick expands into the densely populated Mid-Atlantic region of the United States. We report on the results of five years of field surveillance to better describe the ecology of A. maculatum in newly established populations in southeastern Virginia. We document habitat preferences, host preferences, and the phenology of the adult human-biting life stage. We discuss key ecological factors needed for A. maculatum establishment and the influence of the successional process and anthropogenic activities on the persistence of A. maculatum populations in Virginia.

The predominant human-biting tick throughout the southeastern United States is Amblyomma americanum. Its ability to transmit pathogens causing Lyme disease-like illnesses is a subject of ongoing controversy. Results of previous testing by the Department of Defense Human Tick Test Kit Program and other laboratories indicated that it is highly unlikely that A. americanum transmits any pathogen that causes Lyme disease. In contrast, a recent publication by Clark and colleagues (K. L. Clark, B. Leydet, and S. Hartman, Int. J. Med. Sci. 10:915-931, 2013) reported detection of Lyme group Borrelia in A. americanum using a nested-flagellin-gene PCR. We evaluated this assay by using it and other assays to test 1,097 A. americanum ticks collected from humans. Using the Clark assay, in most samples we observed nonspecific amplification and nonrepeatability of results on subsequent testing of samples. Lack of reaction specificity and repeatability is consistent with mispriming, likely due to high primer concentrations and low annealing temperatures in this protocol. In six suspect-positive samples, Borrelia lonestari was identified by sequencing of an independent gene region; this is not a Lyme group spirochete and is not considered zoonotic. B. burgdorferi was weakly amplified from one pool using some assays, but not others, and attempts to sequence the amplicon of this pool failed, as did attempts to amplify and sequence B. burgdorferi from the five individual samples comprising this pool. Therefore, B. burgdorferi was not confirmed in any sample. Our results do not support the hypothesis that A. americanum ticks are a vector for Lyme group Borrelia infections.

Ixodes affinis Neumann (Acari: Ixodidae) is a hard-bodied tick species distributed throughout much of the southeastern United States. Although I. affinis does not parasitize humans, it is a competent vector of Borrelia burgdorferi sensu stricto, the causative-agent of Lyme disease, and thus contributes to the enzootic maintenance of this pathogen. This study presents evidence of I. affinis parasitizing five new host passerine species. During 2012-2014, 1,888 birds were captured and examined for ticks, and 18 immature I. affinis were collected from 12 birds-six Carolina Wrens (Thyrothorus ludovicianus); two Brown Thrashers (Toxostoma rufum); and one American Robin (Turdus migratorius), Eastern Towhee (Pipilo erythrophthalmus), Northern Cardinal (Cardinalis cardinalis), and White-throated Sparrow (Zonotrichia albicollis). Of 15 larvae and 3 nymphs collected, one nymph tested positive for B. burgdorferi DNA. I. affinis was found co-feeding on birds with immature Amblyomma americanum (L.), Ixodes brunneus Koch, Ixodes dentatus Marx, Ixodes scapularis Say, and Haemaphysalis leporispalustris Packard. The results of this research provide a better understanding of I. affinis hosts and identify avian taxa that may play a role in the maintenance and dispersal of this tick species.

We report evidence that Amblyomma maculatum tick populations are well established in southeastern Virginia. We found that 43.1% of the adult Gulf Coast ticks collected in the summer of 2010 carried Rickettsia parkeri, suggesting that persons living in or visiting southeastern Virginia are at risk for infection with this pathogen.

Background Since 1997, human-biting ticks submitted to the Department of Defense Human Tick Test Kit Program (HTTKP) of the US Army Public Health Command have been tested for pathogens by PCR. We noted differences in the phenology and infection prevalence among Ixodes scapularis ticks submitted from military installations in different geographic regions. The aim of this study was to characterize these observed differences, comparing the phenology and pathogen infection rates of I. scapularis submitted from soldiers at two sites in the upper Midwest (Camp Ripley, MN, and Ft. McCoy, WI) and one site in the northeastern US (Ft. Indiantown Gap, PA). Methods From 1997 through 2012, the HTTKP received 1,981 I. scapularis from the three installations and tested them for Anaplasma phagocytophilum, Babesia microti, Borrelia burgdorferi and the Ehrlichiamuris-like (EML) agent using PCR; pathogen presence was confirmed via sequencing or amplification of a second gene target. Pathogen and co-infection prevalence, tick engorgement status, and phenology were compared among installations. Results Greater rates of A. phagocytophilum and Ba. microti infections were detected in ticks submitted from installations in Minnesota than in Wisconsin or Pennsylvania, and the EML agent was only detected in ticks from Minnesota and Wisconsin. Midwestern ticks were also more likely to be co-infected than those from Pennsylvania. Both adult and nymphal ticks showed evidence of feeding on people, although nymphs were more often submitted engorged. Adult I. scapularis were received more frequently in June from Minnesota than from either of the other sites. Minnesota adult and nymphal peaks overlapped in June, and submissions of adults exceeded nymphs in that month. Conclusions There were clear differences in I. scapularis phenology, pathogen prevalence and rates of co-infection among the three military installations. Seasonal and temperature differences between the three sites and length of time a population had been established in each region may contribute to the observed differences. The synchrony of adults and nymphs observed in the upper Midwest has implications for pathogen infection prevalence. The EML agent was only detected in Minnesota and Wisconsin, supporting the previous assertion that this pathogen is currently limited to the upper Midwest.

Ticks and tick-borne diseases have been on the move throughout the United State over the past twenty years. We use an agent-based model, TICKSIM, to identify the key parameters that determine the success of invasion of the tick and if that is successful, the succees of the tick-borne pathogen. We find that if an area has competent hosts, an initial population of ten ticks is predicted to always establish a new population. The establishment of the tick-borne pathogen depends on three parameters: the initial prevalence in the ten founding ticks, the probability that a tick infects the longer-lived hosts and the probability that a tick infects the shorter-lived hosts. These results indicate that the transmission rates to hosts in the newly established area can be used to predict the potential risk of disease to humans.

The incidence of tick-borne rickettsial disease in the southeastern United States has been rising steadily through the past decade, and the range expansions of tick species and tick-borne infectious agents, new and old, has resulted in an unprecedented mix of vectors and pathogens. The results of an ongoing 4-year surveillance project describe the relative abundance of questing tick populations in southeastern Virginia. Since 2009, more than 66,000 questing ticks of 7 species have been collected from vegetation in a variety of habitats, with Amblyomma americanum constituting over 95% of ticks collected. Other species represented included Ixodes scapularis, Dermacentor variabilis, Amblyomma maculatum, Ixodes affinis, Haemaphysalis leporispalustris, and Ixodes brunneus. We found that 26.9–54.9% of A. americanum ticks tested were positive for Rickettsia amblyommii, a non-pathogenic symbiont of this tick species. We also found no evidence of R. rickettsii in D. variabilis ticks, although they did show low infection rates of R. montanensis (1.5–2.0%). Rickettsia parkeri and Candidatus R. andeanae were found in 41.8–55.7% and 0–1.5% A. maculatum ticks, respectively. The rate of R. parkeri in A. maculatum ticks is among the highest in the literature and has increased in the 2 years since R. parkeri and A. maculatum were first reported in southeastern Virginia. We conclude that tick populations in southeastern Virginia have recently undergone dramatic changes in species and abundance and that these populations support a variety of rickettsial agents with the potential for increased risk to human health.

The recent invasion of the hard tick Ixodes affinis, a sylvatic vector of Borrelia burgdorferi, into the Mid-Atlantic region of the United States has prompted questions as to the origin of these ticks, and the possibility of connectivity with presumed ancestral populations in Central and South America. To assess connectivity and ancestry of I. affinis throughout its range, a fragment of the 16S mitochondrial rRNA gene was sequenced from seven adult I. affinis ticks collected in Colon, Panama, and compared to 19 previously published 16S haplotypes in two clades from the US, seven previously published haplotypes from Belize, and one previously published haplotype from Colombia. This study did not find any overlapping haplotypes between the US, Panama, Belize, and Colombia, showing no evidence for connectivity between North, Central, and South American populations of I. affinis. Two new I. affinis clades associated with ticks from Panama and Belize were also identified. The node that gave rise to the Panama clade of I. affinis was more basal than the nodes leading to the clades containing the majority of ticks from Belize and the United States, and ticks from Belize were more closely related to US ticks than Panama ticks. Several possible competing migration pathways were identified, where ticks may have been introduced to the northern US from Belize, or to the southern US from Panama. Star clusters present in each clade suggest rapid mutation rates after arrival in new areas, and may be associated with the success of I. affinis in invading northern climates. This study provides preliminary evidence for the spread of I. affinis from Central America into the US, and serves as a first step in investigating the possibility of connectivity between Central and North America tick populations.