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.

Daily Variation in Sampled Densities of Ixodes scapularis and Amblyomma americanum (Acari: Ixodidae) Nymphs at a Single Site—Implications for Assessing Acarological Risk

The public health challenge posed by tick-borne disease (TBD) has increased efforts to characterize the spatial and temporal distribution of ticks and associated pathogens to better focus tick control strategies and personal protection measures. We describe variability in nymphal Ixodes scapularis Say and Amblyomma americanum (L.) density derived from daily drag sampling at a single location in New Jersey over 4 yr and explore how observed differences in daily collections might affect the estimation of acarological risk. We found significant variability in the density of host-seeking nymphs that could suggest substantially different rates of human-tick encounters depending on sampling date, habitat, and ambient weather conditions. The spatial and temporal variability in the distribution of 2 sympatric tick species with different host preferences and questing strategies, suggests that to produce results that are comparable among sites across the area sampled, surveillance efforts may be limited to shorter collection seasons, fewer sites or less sampling effort (fewer plots or fewer visits) per site, and a geographic scope that minimizes the potential temporal and spatial biases indicated here. Our results illustrate that evaluation of models of tick distribution or relative acarological risk based on surveillance data requires a full description of the diversity of habitats sampled and the conditions under which sampling is performed. The array of factors that affect tick host-seeking and that could bias interpretation of sampling results emphasizes the need to standardize sampling protocols and for more caution when interpreting tick sampling data collected over large temporal and spatial scales.

Northward Expansion of Amblyomma americanum (Acari: Ixodidae) into Southern Michigan

Amblyomma americanum (Linnaeus) (Acari: Ixodidae) (lone star tick) is an aggressive, generalist parasite that vectors numerous important human and animal pathogens. In recent decades its geographic range has been expanding northwards from endemic regions in the southeastern and southcentral US. In 2019 five questing A. americanum comprising two life stages were detected at one site in Berrien County, in southwestern Michigan, satisfying one CDC criterium for an established population for the first time in the state. To better characterize the northern extent of emerging A. americanum, we conducted active surveillance (i.e., drag sampling) in summer 2020 throughout Michigans southern counties and detected one adult A. americanum from each of six widespread sites, including where they had been detected in 2019. A larger established population was identified at another site in Berrien County, which yielded 691 A. americanum comprising three life stages. Questing tick phenologies at this site were similar to that reported for other regions. Statewide surveillance in 2021 revealed no A. americanum outside of Berrien County, but establishment criteria were met again at the two sites where established populations were first detected respectively in 2019 and 2020. These observations may represent the initial successful invasion of A. americanum into Michigan. Data from passive (1999-2020) and active surveillance (2004-2021) efforts, including a domestic animal sentinel program (2015-2018), are reported to provide context for this nascent invasion. Continued active surveillance is needed to help inform the public, medical professionals, and public health officials of the health risks associated with this vector.

Responses of ticks to immersion in hot bathing water: Effect of surface type, water temperature, and soap on tick motor control

Preventing bites from undetected ticks through bathing practices would benefit public health, but the effects of these practices have been researched minimally. We immersed nymphal and adult hard ticks of species common in the eastern United States in tap water, using temperatures and durations that are realistic for human hot bathing. The effect of (a) different skin-equivalent surfaces (silicone and pig skin), and (b) water temperature was tested on Amblyomma americanum, Dermacentor variabilis and Ixodes scapularis nymphs. Overall, the type of surface had a much larger effect on the nymphs’ tendency to stay in contact with the surface than water temperature did. Most nymphs that separated from the surface did so within the first 10 s of immersion, with the majority losing contact due to the formation of an air bubble between their ventral side and the test surface. In addition, adult Ixodes scapularis were tested for the effect of immersion time, temperature, and soap on tick responsiveness. Some individual adults moved abnormally or stopped moving as a result of longer or hotter immersion, but soap had little effect on responsiveness. Taken together, our results suggest that the surface plays a role in ticks’ tendency to stay in contact; the use of different bath additives warrants further research. While water temperature did not have a significant short-term effect on tick separation, ticks that have not attached by their mouth parts may be rendered unresponsive and eventually lose contact with a person’s skin in a hot bath. It should be noted that our research did not consider potential temperature effects on the pathogens themselves, as previous research suggests that some tickborne pathogens may become less hazardous even if the tick harboring them survives hot-water exposures and later bites the bather after remaining undetected.

Questing behavior of adult Amblyomma americanum (L.) in a laboratory setting

Tick-borne diseases are on the rise globally; however, information is lacking about tick questing behavior. In this laboratory study, we explored tick preferences for stem type (plastic grass, wooden, and metal), questing height, and head orientation. Using 60 Amblyomma americanum adults over three 72-hour replicates, we determined that 21.7% of ticks quested at any given time and that ticks exhibited a strong preference to quest with their heads oriented downwards, irrespective of stem type. Individual ticks tended to quest on only one stem in this study, and a maximum of three. Nonetheless, ticks appeared to prefer questing on wooden and plastic grass stems over metal stems. We did not find an effect of time of day on tick questing rates. Increased understanding of tick questing behavior can improve vector control efforts.

Effects of Climate on the Variation in Abundance of Three Tick Species in Illinois

The range of ticks in North America has been steadily increasing likely, in part, due to climate change. Along with it, there has been a rise in cases of tick-borne disease. Among those medically important tick species of particular concern are Ixodes scapularis Say (Acari: Ixodidae), Dermacentor variabilis Say (Acari: Ixodidae), and Amblyomma americanum Linneaus (Acari: Ixodidae). The aim of this study was to determine if climate factors explain existing differences in abundance of the three aforementioned tick species between two climatically different regions of Illinois (Central and Southern), and if climate variables impact each species differently. We used both zero-inflated regression approaches and Bayesian network analyses to assess relationships among environmental variables and tick abundance. Results suggested that the maximum average temperature and total precipitation are associated with differential impact on species abundance and that this difference varied by region. Results also reinforced a differential level of resistance to desiccation among these tick species. Our findings help to further define risk periods of tick exposure for the general public, and reinforce the importance of responding to each tick species differently.

Identification of public submitted tick images: A neural network approach

Ticks and tick-borne diseases represent a growing public health threat in North America and Europe. The number of ticks, their geographical distribution, and the incidence of tick-borne diseases, like Lyme disease, are all on the rise. Accurate, real-time tick-image identification through a smartphone app or similar platform could help mitigate this threat by informing users of the risks associated with encountered ticks and by providing researchers and public health agencies with additional data on tick activity and geographic range. Here we outline the requirements for such a system, present a model that meets those requirements, and discuss remaining challenges and frontiers in automated tick identification. We compiled a user-generated dataset of more than 12,000 images of the three most common tick species found on humans in the U.S.: Amblyomma americanum, Dermacentor variabilis, and Ixodes scapularis. We used image augmentation to further increase the size of our dataset to more than 90,000 images. Here we report the development and validation of a convolutional neural network which we call “TickIDNet,” that scores an 87.8% identification accuracy across all three species, outperforming the accuracy of identifications done by a member of the general public or healthcare professionals. However, the model fails to match the performance of experts with formal entomological training. We find that image quality, particularly the size of the tick in the image (measured in pixels), plays a significant role in the network’s ability to correctly identify an image: images where the tick is small are less likely to be correctly identified because of the small object detection problem in deep learning. TickIDNet’s performance can be increased by using confidence thresholds to introduce an “unsure” class and building image submission pipelines that encourage better quality photos. Our findings suggest that deep learning represents a promising frontier for tick identification that should be further explored and deployed as part of the toolkit for addressing the public health consequences of tick-borne diseases.

Alpha-Gal Syndrome: Involvement of Amblyomma americanum α-D-Galactosidase and β-1,4 Galactosyltransferase Enzymes in α-Gal Metabolism

Alpha-Gal Syndrome (AGS) is an IgE-mediated delayed-type hypersensitivity reaction to the oligosaccharide galactose-α-1, 3-galactose (α-gal) injected into humans from the lone-star tick (Amblyomma americanum) bite. Indeed, α-gal is discovered in salivary glands of lone-star tick; however, the tick’s specific intrinsic factors involved in endogenous α-gal production and presentation to host during hematophagy are poorly understood. This study aimed to investigate the functional role of two tick enzymes, α-D-galactosidase (ADGal) and β-1,4 galactosyltransferases (β-1,4GalT), in endogenous α-gal production, carbohydrate metabolism, and N-glycan profile in lone-star tick. The ADGal enzyme cleaves terminal α-galactose moieties from glycoproteins and glycolipids, whereas β-1,4GalT transfers α-galactose to a β1,4 terminal linkage acceptor sugars—GlcNAc, Glc, and Xyl—in various processes of glycoconjugate synthesis. An RNA interference approach was utilized to silence ADGal and β-1,4GalT in Am. americanum to examine their function in α-gal metabolism in tick and AGS onset. Silencing of ADGal led to the significant downregulation of genes involved in galactose metabolism and transport in Am. americanum. Immunoblot and N-glycan analysis of the Am. americanum salivary glands showed a significant reduction in α-gal levels in silenced tissues. However, there was no significant difference in the level of α-gal in β-1,4GalT-silenced tick salivary glands. A basophil-activation test showed a decrease in the frequency of activated basophil by ADGal-silenced salivary glands. These results provide an insight into the roles of ADGal and β-1,4GalT in α-gal production and presentation in ticks and the probable involvement in the onset of AGS.

Collaborating With Community Scientists Across Arkansas to Update Tick Distributions and Pathogen Prevalence of Spotted Fever Group Rickettsia and Ehrlichia

Tick-borne diseases (TBD) in humans have dramatically increased over recent years and although the bulk of cases are attributable to Lyme Disease in the Northeastern US, TBDs like spotted fever rickettsiosis and ehrlichiosis heavily impact other parts of the country, namely the mid-south. Understanding tick and pathogen distributions and prevalence traditionally requires active surveillance, which quickly becomes logistically and financially unrealistic as the geographic area of focus increases. We report on a community science effort to survey ticks across Arkansas to obtain updated data on tick distributions and prevalence of human tick-borne disease-causing pathogens in the most commonly encountered ticks. During a 20-mo period, Arkansans submitted 9,002 ticks from 71 of the 75 counties in the state. Amblyomma americanum was the most common tick species received, accounting for 76% of total tick submissions. Nearly 6,000 samples were screened for spotted fever group Rickettsia (SFGR) and Ehrlichia, resulting in general prevalence rates of 37.4 and 5.1%, respectively. In addition, 145 ticks (2.5%) were infected with both SFGR and Ehrlichia. Arkansas Department of Health reported 2,281 spotted fever and 380 ehrlichiosis cases during the same period as our tick collections. Since known SFGR vectors Dermacentor variabilis and Amblyomma maculatum were not the most common ticks submitted, nor did they have the highest prevalence rates of SFGR, it appears that other tick species play the primary role in infecting humans with SFGR. Our investigation demonstrated the utility of community science to efficiently and economically survey ticks and identify vector-borne disease risk in Arkansas.