Modeling of historical and current distributions of lone star tick, Amblyomma americanum (Acari: Ixodidae), is consistent with ancestral range recovery.

The lone star tick, Amblyomma americanum L., is a three-host hard tick notorious for aggressive feeding behavior. In the early to mid-20th century, this species' range was mostly limited to the southern USA. Since the 1950s, A. americanum has been detected in many new localities in the western, northcentral, and northeastern regions of the country. To examine the influence of climate on this apparent expansion, we used historical (1748-1950) lone star locations from the literature and museum records to model areas suitable for this species based on past environmental conditions in the late 1800s - early 1900s. We then projected this model forward using present (2011-2020) climatic conditions and compared the two for evidence of climate-associated distributional shifts. A maximum entropy distribution or Maxent model was generated by using a priori selected climatic variables including temperature, precipitation, and vapor pressure deficit. Temperature and vapor pressure deficit were selected as the most important factors in creating a sensitive and specific model (success rate = 82.6 ± 6.1%) that had a good fit to the existing data and was significantly better than a random model [partial ROC (receiver operating characteristic) to AUC (area under the ROC curve) ratio = 1.97 ± 0.07, P < 0.001]. The present projected model was tested with an independent dataset of curated museum records (1952-2020) and found to be 95.6% accurate. Comparison of past and present models revealed > 98% A. americanum niche overlap. The model suggests that some areas along the western fringe are becoming less suitable for A. americanum, whereas areas in some Great Lakes and coastal northeastern regions are becoming more suitable, results that are compatible with possible effects of climate change. However, these changes are minor, and overall climate in North America does not appear to have changed in ways significant to A. americanum's distribution. These findings are consistent with an alternative hypothesis that recent changes in A. americanum's distribution are a result of this species re-occupying its historical range, driven predominantly by factors other than climate, such as shifts in land use and population densities of major hosts.

First Recognized Human Bite in the United States by the Asian Longhorned Tick, Haemaphysalis longicornis.

We present the case summary of the first human recognized to have been bitten by the Haemaphysalis longicornis tick in the United States, which occurred in New York State. Subsequent field studies confirmed that this tick was present in multiple geographic locations near the patient's residence, including on manicured lawns.

Distribution, Host-Seeking Phenology, and Host and Habitat Associations of Haemaphysalis longicornis Ticks, Staten Island, New York, USA.

Haemaphysalis longicornis, an invasive Ixodid tick, was recently reported in the eastern United States. The emergence of these ticks represents a potential threat for livestock, wildlife, and human health. We describe the distribution, host-seeking phenology, and host and habitat associations of these ticks on Staten Island, New York, a borough of New York City.

Multistate Infestation with the Exotic Disease-Vector Tick Haemaphysalis longicornis - United States, August 2017-September 2018.

Haemaphysalis longicornis is a tick indigenous to eastern Asia and an important vector of human and animal disease agents, resulting in such outcomes as human hemorrhagic fever and reduction of production in dairy cattle by 25%. H. longicornis was discovered on a sheep in New Jersey in August 2017 (1). This was the first detection in the United States outside of quarantine. In the spring of 2018, the tick was again detected at the index site, and later, in other counties in New Jersey, in seven other states in the eastern United States, and in Arkansas. The hosts included six species of domestic animals, six species of wildlife, and humans. To forestall adverse consequences in humans, pets, livestock, and wildlife, several critical actions are indicated, including expanded surveillance to determine the evolving distribution of H. longicornis, detection of pathogens that H. longicornis currently harbors, determination of the capacity of H. longicornis to serve as a vector for a range of potential pathogens, and evaluation of effective agents and methods for the control of H. longicornis.

The hitchhiker's guide to becoming invasive: exotic mosquitoes spread across a US state by human transport not autonomous flight.

Not all exotic species establish and expand aggressively (i.e. become invasive). As potential vectors of disease agents, invasive mosquitoes can have considerable impact on public health, livestock and wildlife; therefore, understanding the species characteristics and ecological circumstances promoting their invasiveness is important. The mosquito Aedes japonicus japonicus, originally from north-east Asia, was introduced at least two separate times to the north-eastern USA, as surmised from the initial existence of two populations with distinct nuclear and mitochondrial genetic signatures that later intermixed. Since these original introductions in the late 1990s, Ae. j. japonicus has expanded across 31 US states, two Canadian provinces and five European countries. Although some of the expanded range was due to other independent introductions, to understand what drove the postintroduction expansion of Ae. j. japonicus within the north-eastern USA, we performed a high-resolution landscape genetic analysis of 461 specimens collected across Virginia, a state south of the original introductions. All specimens were genotyped at seven pre-optimized microsatellite loci, and a subsample was sequenced at one mitochondrial locus. We concluded that throughout Virginia this species has primarily expanded in association with humans: genetic distance and distance along roads remained correlated after controlling for geographic distance, and proximity to I-95, a major interstate highway, strongly predicted nuclear ancestry. In contrast, there was very limited evidence of diffusion even at distances potentially suitable for autonomous mosquito flight. This implies that its association with humans (rather than innate species characteristics) is the single most important determinant of invasiveness in this mosquito.

Characterization of the doublesex gene within the Culex pipiens complex suggests regulatory plasticity at the base of the mosquito sex determination cascade.

BACKGROUND: The doublesex gene controls somatic sexual differentiation of many metazoan species, including the malaria mosquito Anopheles gambiae and the dengue and yellow fever vector Aedes aegypti (Diptera: Culicidae). As in other studied dipteran dsx homologs, the gene maintains functionality via evolutionarily conserved protein domains and sex-specific alternative splicing. The upstream factors that regulate splicing of dsx and the manner in which they do so however remain variable even among closely related organisms. As the induction of sex ratio biases is a central mode of action in many emerging molecular insecticides, it is imperative to elucidate as much of the sex determination pathway as possible in the mosquito disease vectors. RESULTS: Here we report the full-length gene sequence of the doublesex gene in Culex quinquefasciatus (Cxqdsx) and its male and female-specific isoforms. Cxqdsx maintains characteristics possibly derived in the Culicinae and present in the Aedes aegypti dsx gene (Aeadsx) such as gain of exon 3b and the presence of Rbp1 cis-regulatory binding sites, and also retains presumably ancestral attributes present in Anopheles gambiae such as maintenance of a singular female-specific exon 5. Unlike in Aedes aegypti, we find no evidence for intron gain in the female transcript(s), yet recover a second female isoform generated via selection of an alternate splice donor. Utilizing next-gen sequence (NGS) data, we complete the Aeadsx gene model and identify a putative core promoter region in both Aeadsx and Cxqdsx. Also utilizing NGS data, we construct a full-length gene sequence for the dsx homolog of the northern house mosquito Culex pipiens form pipiens (Cxpipdsx). Analysis of peptide evolutionary rates between Cxqdsx and Cxpipdsx (both members of the Culex pipiens complex) shows the male-specific portion of the transcript to have evolved rapidly with respect to female-specific and common regions. CONCLUSIONS: As in other studied insects, doublesex maintains sex-specific splicing and conserved doublesex/mab-3 domains in the mosquitoes Culex quinquefasciatus and Cx. pipiens. The cis-regulated splicing of Cxqdsx does not appear to follow either currently described mosquito model (for An. gambiae and Ae. aegypti); each of the three mosquito genera exhibit evidence of unique cis-regulatory mechanisms. The male-specific dsx terminus exhibits rapid peptide evolutionary rates, even among closely related sibling species.

First record of Ixodes keiransi (Acari: Ixodidae) in New Jersey, USA.

The hard tick, Ixodes keiransi Beati, Nava, Venzal, & Guglielmone, formerly the North American lineage of Ixodes affinis Neumann, is expanding its range northward along the US East Coast. In July 2023, we collected I. keiransi adult female and nymph in a single sampling event, suggesting its range now includes southern New Jersey. In this area, I. keiransi is sympatric with northern populations of Ixodes scapularis Say (Acari: Ixodidae), the primary vector of Lyme disease. Given its status as an enzootic vector of spirochaetes in the Borrelia burgdorferi sensu lato complex, proper differentiation of these 2 species will be critical for accurate estimates of entomological risk. Targeted surveillance should be implemented to monitor further I. keiransi expansion and to elucidate the phenology and enzootic role of this and other understudied Ixodes spp. in the northeastern United States.

Earliest records of the Asian longhorned tick (Acari: Ixodidae) in Staten Island, New York, and subsequent population establishment, with a review of its potential medical and veterinary importance in the United States.

Three Asian longhorned ticks (Haemaphysalis longicornis) were collected on Staten Island, Richmond County, New York, in 2014-2015 as part of a tick-borne disease surveillance program conducted by the New York City Department of Health and Mental Hygiene and the Defense Centers of Public Health - Aberdeen Tick-Borne Disease Laboratory. These records mark the earliest known occurrence of H. longicornis in New York State outside of quarantine areas, predating previously reported detections by several years. Robust populations of H. longicornis were collected in subsequent years at the Staten Island site where these few ticks were found, demonstrating that small infestations have the potential to proliferate quickly. Haemaphysalis longicornis is a 3-host ixodid tick native to eastern Asia but now established in the United States, as well as Australasia and several Pacific islands. Although H. longicornis has not yet been associated with human disease transmission in the United States, it warrants attention as a potential vector, as it is demonstrated to harbor various pathogens of medical and veterinary interest across its native and introduced range.

Five human pathogens detected by tick surveillance in New York City parks, 2014-2015.

A total of 2,504 ticks of 5 species (Ixodes scapularis, Dermacentor variabilis, Amblyomma americanum, Haemaphysalis leporispalustris, and H. longicornis) were collected over 2 yr (2014-2015) in New York City parks. Specimens were collected via tick-dragging, identified to species, and tested for pathogens of human diseases. The causative agents of 5 human diseases (Lyme borreliosis, ehrlichiosis, babesiosis, anaplasmosis, and Rocky Mountain spotted fever) were detected in a subset of samples. Results of this surveillance effort further illustrate that risk of tick-borne disease is considerable even in parks located adjacent to densely populated areas.

Baseline susceptibility of Haemaphysalis longicornis to organophosphate, carbamate, and pyrethroid acaricides.

BACKGROUND: The Asian longhorned tick, Haemaphysalis longicornis, continues to expand its range in North America, and synthetic acaricides are likely to play an increasing role in managing this species. Acaricide resistance is common in some tick species that infest livestock. However, baseline acaricide susceptibility has not previously been examined in this invasive tick. RESULTS: We used a standard larval packet test to evaluate the susceptibility of the Asian longhorned tick to acaricides currently or formerly used in tick control: propoxur, carbaryl, bifenthrin, permethrin, and coumaphos. Discriminating concentrations were estimated at 6.5, 27.9, 988, 2242, and 808 ppm, respectively. The half-maximal lethal concentration (LC50 ) values for propoxur, carbaryl, permethrin and coumaphos were compared with data available for other tick species and showed that H. longicornis was more susceptible to propoxur, carbaryl and coumaphos, and had a similar susceptibility to permethrin. CONCLUSIONS: The results indicate that resistance to these acaricides is not currently a concern for H. longicornis in the United States. However, responsible integrated management and early detection of resistance can help ensure the long-term efficacy of products used for controlling this tick species. © 2023 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Lone star ticks (Acari: Ixodidae) infected with Bourbon virus in New Jersey, USA.

Lone star ticks (Amblyomma americanum L.) are expanding within the northeast United States, a region historically focused on Ixodes scapularis-transmitted diseases. In Monmouth County, NJ, the shift has been dramatic, and lone star ticks now vastly outnumber blacklegged ticks. As a result, there is an enhanced need to focus on the potential health risks of A. americanum-transmitted pathogens, such as the emerging Heartland (HRTV) and Bourbon (BRBV) viruses. We screened 1,205 nymphal lone star ticks for HRTV and BRBV using RT-qPCR assays and detected BRBV in 3 ticks collected in Monmouth County, NJ, in 2021. Additionally, we sequenced a complete BRBV genome from a single infected specimen, finding 99.4% identity with human pathogenic isolates from the eastern-central United States. Our results have important public health implications for a region only recently becoming aware of public health risks posed by lone star ticks. Of note, we report successful detection of viral RNA in samples that were stored and intended for DNA preservation, for example, kept in ethanol at room temperature, which may reduce barriers for public health agencies seeking to expand their tick testing to include viruses.

Detection of multiple tick-borne pathogens in Ixodes scapularis from Hunterdon County, NJ, USA.

Several human pathogens vectored by the blacklegged tick (Ixodes scapularis Say; Acari: Ixodidae) are endemic in the state of New Jersey. Disease incidence data suggest that these conditions occur disproportionately in the northwestern portion of the state, including in the county of Hunterdon. We conducted active surveillance at three forested sites in Hunterdon County during 2020 and 2021, collecting 662 nymphal and adult I. scapularis. Ticks were tested for five pathogens by qPCR/qRT-PCR: Anaplasma phagocytophilum, Babesia microti, Borrelia burgdorferi, Borrelia miyamotoi, and Powassan virus (POWV) lineage 2. Over 2 years, 25.4% of nymphs and 58.4% of adults were found infected with at least one pathogen, with 10.6% of all ticks infected with more than one pathogen. We report substantial spatial and temporal variability of A. phagocytophilum and B. burgdorferi, with high relative abundance of the human-infective A. phagocytophilum variant Ap-ha. Notably, POWV was detected for the first time in Hunterdon, a county where human cases have not been reported. Based on comparisons with active surveillance initiatives in nearby counties, further investigation of non-entomological factors potentially influencing rates of tick-borne illness in Hunterdon is recommended.

Microhabitat modeling of the invasive Asian longhorned tick (Haemaphysalis longicornis) in New Jersey, USA.

The Asian longhorned tick (Haemaphysalis longicornis) is a vector of multiple arboviral and bacterial pathogens in its native East Asia and expanded distribution in Australasia. This species has both bisexual and parthenogenetic populations that can reach high population densities under favorable conditions. Established populations of parthenogenetic H. longicornis were detected in the eastern United States in 2017 and the possible range of this species at the continental level (North America) based on climatic conditions has been modeled. However, little is known about factors influencing the distribution of H. longicornis at geographic scales relevant to local surveillance and control. To examine the importance of local physiogeographic conditions such as geology, soil characteristics, and land cover on the distribution of H. longicornis we employed ecological niche modeling using three machine learning algorithms - Maxent, Random Forest (RF), and Generalized Boosting Method (GBM) to estimate probability of finding H. longicornis in a particular location in New Jersey (USA), based on environmental predictors. The presence of H. longicornis in New Jersey was positively associated with Piedmont physiogeographic province and two soil types - Alfisols and Inceptisols. Soil hydraulic conductivity was the most important predictor explaining H. longicornis habitat suitability, with more permeable sandy soils with higher hydraulic conductivity being less suitable than clay or loam soils. The models were projected over the state of New Jersey creating a probabilistic map of H. longicornis habitat suitability at a high spatial resolution of 90×90 meters. The model's sensitivity was 87% for locations sampled in 2017-2019 adding to the growing evidence of the importance of soil characteristics to the survival of ticks. For the 2020-2022 dataset the model fit was 57%, suggestive of spillover to less optimal habitats or, alternatively, heterogeneity in soil characteristics at the edges of broad physiographic zones. Further modeling should incorporate abundance and life-stage information as well as detailed characterization of the soil at collection sites. Once critical parameters that drive the survival and abundance of H. longicornis are identified they can be used to guide surveillance and control strategies for this invasive species.

Where have all the grouse ticks gone? Apparent decline in collections of Haemaphysalis chordeilis Packard.

Haemaphysalis chordeilis Packard, also known as the grouse or bird tick, is a three-host tick native to North America. Literature from the early 20th century reported a widespread distribution of this tick across the US and Canada. As its name implies, ground-dwelling birds such as grouse and quail were frequent hosts, and occasionally large infestations were reported in domestic flocks making it a pest of economic importance. However, after the mid-1900's records of this species appear scarce, and a number of more recent studies of ticks on birds (including ostensibly favored host species) did not detect it. To confirm this perception with data, we conducted a literature search for collection records of this species and compared the records across two eras (pre-1965 and post-1965), finding very few records of H. chordeilis in recent years, despite increased attention brought to this genus by the detection of exotic Haemaphysalis longicornis Neumann populations in the eastern US. We also compiled a list of studies after 1965 that examined appropriate hosts for ectoparasites but failed to find H. chordeilis. We interpret the apparent decline of ticks in the context of documented population declines in several major host species over the same time frame and discuss whether ectoparasite populations should be subject to the same conservation consideration as their hosts.

The Original Scientific Description of the Lone Star Tick (Amblyomma americanum, Acari: Ixodidae) and Implications for the Species' Past and Future Geographic Distributions.

Amblyomma americanum L. is an important vector in North America originally described by Linnaeus based on Pehr Kalm's 1754 report. While Kalm's 'Travels into North America' is well known, his 1754 report remains obscure. Some authors were skeptical that Kalm referred to A. americanum because he encountered them at sites farther north outside of the species' range. However, the details in 1754 report leave no doubt that Kalm described lone star ticks. In this historical review, we provide support for Kalm's identification using a modern translation of his 1754 report and other sources. We also delineate distributional changes of lone star ticks from the pre-colonization era to the present and interpret them in the context of large-scale anthropogenic changes in the landscape. In this framework, the lone star tick's current northward expansion is a recolonization of their former range. Extensive deforestation and extirpation of their principal host species, white-tailed deer, led to A. americanum's disappearance from the northern parts of its range by the 20th century. Subsequent recolonization by second-growth forest and increases in white-tailed deer populations by the mid-20th century is now allowing A. americanum to reclaim its former range. These changes in the land appear to be the driving force behind A. americanum's present expansion. Understanding this species' history and the factors contributing to its current expansion will enable better predictions about its future distribution and potential to transmit human pathogens.

Relevance of Spatial and Temporal Trends in Nymphal Tick Density and Infection Prevalence for Public Health and Surveillance Practice in Long-Term Endemic Areas: A Case Study in Monmouth County, NJ.

Tick-borne diseases are a growing public health problem in the United States, and the US northeast has reported consistently high case rates for decades. Monmouth County, New Jersey, was one of the earliest jurisdictions to report Lyme disease cases in 1979 and reports several hundred cases per year nearly 40 yr later. In the time since, however, tick-borne health risks have expanded far beyond Lyme disease to include a variety of other bacterial pathogens and viruses, and additional vectors, necessitating a continually evolving approach to tick surveillance. In 2017, Monmouth County initiated an active surveillance program targeting sites across three ecological regions for collection of Ixodes scapularis Say (Acari: Ixodidae) and Amblyomma americanum L. (Acari: Ixodidae) as well as testing via qPCR for associated bacterial pathogens. During the first five years of this program (2017-2021), we report high levels of spatiotemporal variability in nymphal density and infection prevalence in both species, limiting the granularity with which human risk can be predicted from acarological data. Nonetheless, broader patterns emerged, including an ongoing trend of A. americanum dominance, risks posed by Borrelia miyamotoi, and the frequency of coinfected ticks. We present some of the first county-level, systematic surveillance of nymphal A. americanum density and infection prevalence in the northeastern US. We also documented a temporary decline in Borrelia burgdorferi that could relate to unmeasured trends in reservoir host populations. We discuss the implications of our findings for tick-borne disease ecology, public health communication, and tick surveillance strategies in endemic areas.

The Gulf Coast Tick, Amblyomma maculatum (Ixodida: Ixodidae), and Spotted Fever Group Rickettsia in the Highly Urbanized Northeastern United States.

We report the multi-year collection of the Gulf Coast tick, Amblyomma maculatum Koch (Acaridae: Ixodida: Ixodidae) in Staten Island, New York City (NYC) as well as their detection in Brooklyn, NYC, and in Atlantic and Cumberland counties in southern New Jersey, USA. The first and most common detections were of adults, however in Freshkills Park on Staten Island larvae were also collected in a following year. The presence of larvae indicates that adults are successfully finding hosts in Staten Island. While it is still unknown how A. americanum reached Staten Island, immatures of this species often parasitize migratory birds, which are now often seen in Freshkills Park. We describe the landscape features of the area in Staten Island where populations were highest and larvae were detected, which could have facilitated the establishment of A. maculatum. Notably, we also report the presence of human pathogens Rickettsia parkeri in 5/10 (50%) of adults tested and R. felis in 1/24 (4.17%) of larvae tested. In addition to established populations in Staten Island we found evidence of A. maculatum in NJ and other NYC boroughs, suggesting current or future establishment is possible. The failure thus far to detect established populations in these areas may be due to inherent difficulties in detecting low density, spatially heterogeneous incipient populations, which could require targeted surveillance efforts for this species. We discuss the consequences to public health of the establishment of A. maculatum and detection of two additional rickettsial pathogens in the densely populated northeastern United States.

Comparative hologenomics of two Ixodes scapularis tick populations in New Jersey.

Tick-borne diseases, such as those transmitted by the blacklegged tick Ixodes scapularis, are a significant and growing public health problem in the US. There is mounting evidence that co-occurring non-pathogenic microbes can also impact tick-borne disease transmission. Shotgun metagenome sequencing enables sampling of the complete tick hologenome-the collective genomes of the tick and all of the microbial species contained therein, whether pathogenic, commensal or symbiotic. This approach simultaneously uncovers taxonomic composition and allows the detection of intraspecific genetic variation, making it a useful tool to compare spatial differences across tick populations. We evaluated this approach by comparing hologenome data from two tick samples (N = 6 ticks per location) collected at a relatively fine spatial scale, approximately 23 km apart, within a single US county. Several intriguing variants in the data between the two sites were detected, including polymorphisms in both in the tick's own mitochondrial DNA and that of a rickettsial endosymbiont. The two samples were broadly similar in terms of the microbial species present, including multiple known tick-borne pathogens (Borrelia burgdorferi, Babesia microti, and Anaplasma phagocytophilum), filarial nematodes, and Wolbachia and Babesia species. We assembled the complete genome of the rickettsial endosymbiont (most likely Rickettsia buchneri) from both populations. Our results provide further evidence for the use of shotgun metagenome sequencing as a tool to compare tick hologenomes and differentiate tick populations across localized spatial scales.