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Amblyomma maculatum Koch sensu lato (s.l.) ticks are the vector of Rickettsia parkeri in Arizona, where nine cases of R. parkeri rickettsiosis have been identified since the initial case in 2014. The current study sought to better define the geographic ranges of the vector and pathogen and to assess the potential public health risk posed by R. parkeri in this region of the southwestern United States. A total of 275 A. maculatum s.l. ticks were collected from 34 locations in four counties in Arizona and one county in New Mexico and screened for DNA of Rickettsia species. Rickettsia parkeri was detected in 20.4% of the ticks, including one specimen collected from New Mexico, the first report of R. parkeri in A. maculatum s.l. from this state. This work demonstrates a broader distribution of A. maculatum s.l. ticks and R. parkeri in the southwestern United States than appreciated previously to suggest that R. parkeri rickettsiosis is underrecognized in this region.

Rickettsia parkeri, a tick-borne pathogen distributed throughout several countries of the Americas, causes a mild to moderately severe, eschar-associated spotted fever rickettsiosis. Although most U.S. cases of R. parkeri rickettsiosis are reported from southeastern states, some have been reported recently from remote regions of southern Arizona. These cases are linked to R. parkeri-infected ticks of the Amblyomma maculatum (Acari: Ixodidae) group found in several isolated mountain ranges of southern Arizona and New Mexico, referred to as ‘sky islands’. Archival records also document ticks of the A. maculatum group collected from domestic and wild animals in West Texas. We surveyed sites in two sky island chains of Jeff Davis and Brewster counties to document the off-host occurrence of these ticks and identify the presence of R. parkeri in the Trans-Pecos region of Texas. During August 2019, 43 adult A. maculatum group ticks were flagged from vegetation or removed from a road-killed, female mule deer. Of 39 samples evaluated by PCR, eight contained a partial sca0 sequence with complete identity to R. parkeri and two with complete identity to ‘Candidatus Rickettsia andeanae’, a species of undetermined pathogenicity. Four isolates of R. parkeri were obtained using cell culture. Persons at risk for R. parkeri rickettsiosis include those who work or recreate in these mountains, such as hikers, backpackers, research scientists, foresters, and border enforcement personnel. Additional investigations are needed to define the distribution of these medically important arthropods in other parts of the southwestern United States and northern Mexico.

The western black-legged tick (Ixodes pacificus) is the most frequently identified human-biting tick species in the western United States and the principal vector of at least three recognized bacterial pathogens of humans. A potentially pathogenic Rickettsia species, first described in 1978 and recently characterized as a novel transitional group agent designated as Rickettsia tillamookensis, also exists among populations of I. pacificus, although the distribution and frequency of this agent are poorly known. We evaluated DNA extracts from 348 host-seeking I. pacificus nymphs collected from 9 locations in five California counties, and from 916 I. pacificus adults collected from 24 locations in 13 counties, by using a real-time PCR designed specifically to detect DNA of R. tillamookensis. DNA of R. tillamookensis was detected in 10 (2.9%) nymphs (95% CI: 1.6–5.2%) and 17 (1.9%) adults (95% CI: 1.2–3.0%) from 11 counties of northern California. Although site-specific infection rates varied greatly, frequencies of infection remained consistently low when aggregated by stage, sex, habitat type, or geographical region. Four novel isolates of R. tillamookensis were cultivated in Vero E6 cells from individual adult ticks collected from Alameda, Nevada, and Yolo counties. Four historical isolates, serotyped previously as ‘Tillamook-like’ strains over 40 yr ago, were revived from long-term storage in liquid nitrogen and confirmed subsequently by molecular methods as isolates of R. tillamookensis. The potential public health impact of R. tillamookensis requires further investigation.

In the 1930s, R. A. Cooley noted that Dermacentor occidentalis (Acarina: Ixodidae) and Dermacentor andersoni were closely related and could hybridize. Decades later, James Oliver discovered that crosses of Dermacentor variabilis, D. andersoni, and D. occidentalis could, on occasion, produce hybrids. A recent molecular analysis (both mtDNA and nDNA) in our laboratory revealed that certain specimens of Dermacentor andersoni nested with Dermacentor parumapertus. Does this close relationship, along with the mito-nuclear discordance we have observed, mean D. andersoni and D. parumapertus are a single species? By contemporary taxonomic criteria, this seems improbable based on their distinctly different morphologies, host associations, and ecologies. This paper explores ideas related to mito-nuclear discordance, hybridization, and introgression (primarily) not only in these two species but also other members of the genus Dermacentor. Both D. andersoni and D. parumapertus can be found on the same hosts and have sympatric distributions, so introgression of genetic material by occasional cross-mating between these two species is possible. Further, the difficulty in applying specific species concepts in ticks has been recently pointed out and a unified agreement on an integrative species concepts could clearly be useful in this situation. With the discovery of D. parumapertus as a potential vector of Rickettsia parkeri and the historically recognized role of D. andersoni in transmission of Rickettsia rickettsii, understanding the specific status of each lineage of these species (and others in the genus) is extremely important from a public health perspective.

We identified an established population of the Gulf Coast tick (Amblyomma maculatum Koch) infected with Rickettsia parkeri in Connecticut, representing the northernmost range limit of this medically relevant tick species. Our finding highlights the importance of tick surveillance and public health challenges posed by geographic expansion of tick vectors and their pathogens.

The Pacific Coast tick (Dermacentor occidentalis Marx, 1892) is one of the most widely distributed and frequently encountered tick species in California. This tick is the primary vector of an unclassified spotted fever group rickettsial pathogen, designated currently as Rickettsia 364D, the etiologic agent of a recently recognized tick-borne rickettsiosis known as Pacific Coast tick fever. Despite intensified interest in this pathogen, important questions remain regarding its taxonomic status and possible variations in genotype among different strains that could influence its pathogenicity. Only the extensively passaged prototypical isolate (strain 364-D) is widely available to rickettsiologists and public health scientists worldwide. To achieve a larger, more geographically diverse, and contemporary collection of strains, 1,060 questing adult D. occidentalis ticks were collected from 18 sites across six counties in northern and southern California in 2016 and 2017. Fourteen ticks (1.3%) yielded DNA of Rickettsia 364D and from these, 10 unique isolates from Lake and Orange counties were obtained. Additionally, Rickettsia rhipicephali was detected in 108 (10.2%) ticks, from which eight isolates were obtained, and Rickettsia bellii in six (0.6%), from which three isolates were obtained. The panel of recently acquired, low-passage strains of Rickettsia 364D derived from this study could enhance opportunities for investigators to accurately determine the taxonomic standing of this agent and to develop specific diagnostic assays for detecting infections with Rickettsia 364D in ticks and humans.

Rickettsia parkeri is an emerging human pathogen transmitted by Amblyomma ticks in predominately tropical and subtropical regions of the western hemisphere. In 2014 and 2015, one confirmed case and one probable case of R. parkeri rickettsiosis were reported from the Pajarita Wilderness Area, a semi-arid mountainous region in southern Arizona. To examine more closely the potential public health risk of R. parkeri in this region, a study was initiated to investigate the pervasiveness of Amblyomma maculatum Koch group ticks in mountainous areas of southern Arizona and to ascertain the infection frequencies of R. parkeri in these ticks. During July 2016, a total of 182 adult ticks were collected and evaluated from the Pajarita Wilderness Area in Santa Cruz County and two additional sites in Cochise and Santa Cruz counties in southern Arizona. DNA of R. parkeri was detected in a total of 44 (24%) of these ticks. DNA of "Candidatus Rickettsia andeanae" and Rickettsia rhipicephali was detected in three (2%) and one (0.5%) of the samples, respectively. These observations corroborate previous collection records and indicate that established populations of A. maculatum group ticks exist in multiple foci in southern Arizona. The high frequency of R. parkeri in these tick populations suggests a public health risk as well as the need to increase education of R. parkeri rickettsiosis for those residing, working in, or visiting this area.

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.

Rickettsia bellii is a rickettsial species of unknown pathogenicity that infects argasid and ixodid ticks throughout the Americas. Many molecular assays used to detect spotted fever group (SFG) Rickettsia species do not detect R. bellii , so that infection with this bacterium may be concealed in tick populations when assays are used that screen specifically for SFG rickettsiae. We describe the development and validation of a R. bellii -specific, quantitative, real-time PCR TaqMan assay that targets a segment of the citrate synthase ( glt A) gene. The specificity of this assay was validated against a panel of DNA samples that included 26 species of Rickettsia , Orientia , Ehrlichia , Anaplasma , and Bartonella , five samples of tick and human DNA, and DNA from 20 isolates of R. bellii , including 11 from North America and nine from South America. A R. bellii control plasmid was constructed, and serial dilutions of the plasmid were used to determine the limit of detection of the assay to be one copy per 4 µl of template DNA. This assay can be used to better determine the role of R. bellii in the epidemiology of tick-borne rickettsioses in the Western Hemisphere.

Dermacentor parumapertus Neumann (Acari: Ixodidae) is a rather obscure tick found on jackrabbits in the western United States and parts of Canada and Mexico. Specimens from the northern part of their range are consistently different morphologically from ones found in southern and eastern parts of their range (particularly west Texas), leading some researchers to declare the southern form a variety or subspecies. This study examined field-collected adult D. parumapertus from two main locations—Utah and Texas—within its geographic distribution to ascertain the degree of genetic divergence in the two populations based upon both nuclear and mitochondrial DNA sequences. In total, 30 D. parumapertus were analyzed by PCR using both mtDNA and nDNA genes, and one D. nitens was included for comparison. Trees were constructed for all mtDNA genes individually, as well as after concatenating mtDNA (COI, COII, 12S) and nDNA (2 ITS2 primers), respectively. All constructed trees were exported to FigTree v1.4.3 and TreeGraph v2.14.1–771 beta for visualization. The majority of the Utah and Texas populations of D. parumapertus separated molecularly in both mtDNA and nDNA trees; however, analysis with mtDNA genes showed that 3/13 (23%) of Utah tick specimens were removed molecularly from other specimens collected at the same location. Thus, there was not enough evidence to declare these two disparate and morphologically different populations as distinct and separate species.

In the United States, the lone star tick, Amblyomma americanum (L.) (Acari: Ixodidae), is an aggressive southeastern species whose range has reportedly been steadily expanding northward. The number of A. americanum specimens submitted to the Tick Testing Laboratory (TTL) at the Connecticut Agricultural Experiment Station (CAES) increased by 58% from the period of 1996-2006 (n = 488) to 2007-2017 (n = 773), mainly from Fairfield County in the southwestern corner of the state. The greatest numbers of A. americanum submissions to the CAES-TTL were from the City of Norwalk and a few adjacent municipalities. We also report the discovery of a large infestation of adult and nymphal lone star ticks detected on a dead male white-tailed deer, Odocoileus virginianus (Zimmerman) (Artiodactyla: Cervidae), on Manresa Island, Norwalk, in June 2017, indicating a long established, undetected population along the southwestern coast. A sample of nymphal and adult host-seeking A. americanum collected July 2017 from Manresa Island were tested and a proportion were positive for Ehrlichia chaffeensis, Ehrlichia ewingii, and Anaplasma phagocytophilum. The A. americanum tick and its associated disease pathogens are expected to become an increasing public health concern in southern New England.

Dermacentor parumapertus Neumann (Acari: Ixodidae), a tick primarily associated with rabbits which occurs over much of the western United States, has a fairly large north-to-south distribution, being found from central Idaho southward into northern Mexico. This mostly obscure tick species has recently been the focus of attention due to the discovery of a unique strain of Rickettsia parkeri associated with it which appears closely related to a Rickettsia sp. found in the Atlantic rainforest of Brazil. Historically, a morphological variety of this species was reported in the literature based on significant variation in ornamentation of the tick throughout its range. This study examines several key morphological characters to determine if there are indeed more than one distinct population of this species throughout its range.

Amblyomma maculatum Koch, 1844 (also known as the Gulf Coast tick) is found in parts of the Americas, including the central and southern United States. Its primary importance is as the vector of Rickettsia parkeri, a spotted fever group rickettsia that causes an illness similar to, but milder than, Rocky Mountain spotted fever. A second spotted fever group rickettsia, "Candidatus Rickettsia andeanae," was detected in Gulf Coast ticks approximately 10 yr ago. However, the significance of this organism, including pathogenicity, has not yet been well-characterized. Here, we use transmission electron microscopy to describe bacteria within the tissues of A. maculatum ticks that were positive by polymerase chain reaction assay for "Ca. R. andeanae." In ultrathin sections of unfed A. maculatum adult females, we found evidence of bacteria with morphological features consistent with spotted fever group rickettsiae, including small size (≈0.3 by 0.9 μm), a halo zone (electron-lucent layer around the bacterium), and a trilaminar cell wall. In female ticks, bacteria were present in granular salivary glands and ducts, foregut, Malpighian tubules, nerve trunks, and reproductive tissue. These findings demonstrate evidence of "Ca. R. andeanae" in situ and contribute to our understanding of this novel rickettsia in A. maculatum.

Panola Mountain Ehrlichia (PME) has been suggested as an emerging pathogen of humans and dogs. Domestic goats and white-tailed deer (Odocoileus virginianus) are also susceptible and likely serve as reservoirs. Experimentally, both the lone star tick (Amblyomma americanum (L.)) and the Gulf Coast tick (Amblyomma maculatum Koch) can transmit PME among deer and goats. In the current study, we detected PME in adult wild-caught A. maculatum from the United States and Amblyomma variegatum (F.) from the Caribbean and Africa. This significantly expands the range, potential tick vectors, and risk for exposure to PME.

Amblyomma maculatum Koch (the Gulf Coast tick) is an aggressive, human-biting ixodid tick distributed throughout much of the southeastern United States and is the primary vector for Rickettsia parkeri, an emerging human pathogen. Amblyomma maculatum has diverse host preferences that include white-tailed deer, a known reservoir for Ehrlichia and Anaplasma species, including the human pathogens E. ewingii and E. chaffeensis. To examine more closely the potential role of A. maculatum in the maintenance of various pathogenic Ehrlichia and Anaplasma species, we screened DNA samples from 493 questing adult A. maculatum collected from six U.S. states using broad-range Anaplasmataceae and Ehrlichia genus-specific PCR assays. Of the samples tested, four (0.8%) were positive for DNA of Ehrlichia ewingii, one (0.2%) was positive for Anaplasma platys, and one (0.2%) was positive for a previously unreported Ehrlichia species closely related to Ehrlichia muris and an uncultivated Ehrlichia species from Haemaphysalis longicornis ticks in Japan. No ticks contained DNA of Ehrlichia chaffeensis, Ehrlichia canis, the Panola Mountain Ehrlichia, or Anaplasma phagocytophilum. This is the first identification of E. ewingii, A. platys, and the novel Ehrlichia in questing Gulf Coast ticks; nonetheless the low prevalence of these agents suggests that A. maculatum is not likely an important vector of these zoonotic pathogens.

Panola Mountain Ehrlichia (PME) has been suggested as an emerging pathogen of humans and dogs. Domestic goats and white-tailed deer (Odocoileus virginianus) are also susceptible and likely serve as reservoirs. Experimentally, both the lone star tick (Amblyomma americanum (L.)) and the Gulf Coast tick (Amblyomma maculatum Koch) can transmit PME among deer and goats. In the current study, we detected PME in adult wild-caught A. maculatum from the United States and Amblyomma variegatum (F.) from the Caribbean and Africa. This significantly expands the range, potential tick vectors, and risk for exposure to PME.

Amblyomma maculatum Koch, 1844 (also known as the Gulf Coast tick) is found in parts of the Americas, including the central and southern United States. Its primary importance is as the vector of Rickettsia parkeri, a spotted fever group rickettsia that causes an illness similar to, but milder than, Rocky Mountain spotted fever. A second spotted fever group rickettsia, "Candidatus Rickettsia andeanae," was detected in Gulf Coast ticks approximately 10 yr ago. However, the significance of this organism, including pathogenicity, has not yet been well-characterized. Here, we use transmission electron microscopy to describe bacteria within the tissues of A. maculatum ticks that were positive by polymerase chain reaction assay for "Ca. R. andeanae." In ultrathin sections of unfed A. maculatum adult females, we found evidence of bacteria with morphological features consistent with spotted fever group rickettsiae, including small size (≈0.3 by 0.9 μm), a halo zone (electron-lucent layer around the bacterium), and a trilaminar cell wall. In female ticks, bacteria were present in granular salivary glands and ducts, foregut, Malpighian tubules, nerve trunks, and reproductive tissue. These findings demonstrate evidence of "Ca. R. andeanae" in situ and contribute to our understanding of this novel rickettsia in A. maculatum.

Medical importance of the Gulf Coast tick, Amblyomma maculatum Koch, has been highlighted with the recent discovery of a new rickettsial pathogen associated with this tick. Accordingly, distribution and seasonal collection records of A. maculatum were assessed in Mississippi by using three sources of data: sampling of experimental field plots, national tick collection records, and Mississippi Department of Health human tick biting records. A. maculatum was collected in 17/82 (21%) Mississippi counties, mostly in central and southern regions. Estimates of seasonal activity were made using collection data for 2,217 adult, 11 nymphal, and 426 larval specimens. Adult A. maculatum were collected during March through November, with a peak during late July through early August. Nymphs were collected during February through August, and larvae were collected once during each of June, September, and October. Small sample sizes of the immature stages precluded determination of their peak activity.

Amblyomma maculatum Koch (the Gulf Coast tick) is a three-host, ixodid tick that is distributed throughout much of the southeastern and south-central United States, as well as several countries throughout Central and South America. A considerable amount of scientific literature followed the original description of A. maculatum in 1844; nonetheless, the Gulf Coast tick was not recognized as a vector of any known pathogen of animals or humans for >150 years. It is now identified as the principal vector of Hepatozoon americanum, the agent responsible for American canine hepatozoonosis, and Rickettsia parkeri, the cause of an emerging, eschar-associated spotted fever group rickettsiosis identified throughout much of the Western Hemisphere. Coincident with these discoveries has been recognition that the geographical distribution of A. maculatum in the United States is far more extensive than described 70 yr ago, supporting the idea that range and abundance of certain tick species, particularly those with diverse host preferences, are not fixed in time or space, and may change over relatively short intervals. Renewed interest in the Gulf Coast tick reinforces the notion that the perceived importance of a particular tick species to human or animal health can be relatively fluid, and may shift dramatically with changes in the distribution and abundance of the arthropod, its vertebrate hosts, or the microbial agents that transit among these organisms.

The lone star tick, Amblyomma americanum (L.), has increased in abundance in several regions of the northeastern United States, including areas of Long Island, NY. Adult and nymphal stage A. americanum collected from several sites on Long Island were evaluated for infection with Ehrlichia chaffeensis, the causative agent of human monocytic ehrlichiosis (HME), by using a nested polymerase chain reaction assay. Fifty-nine (12.5%) of 473 adults and eight of 113 pools of five nymphs each (estimated minimum prevalence of infection 1.4%) contained DNA of E. chaffeensis. These data, coupled with the documented expansion of lone star tick populations in the northeastern United States, confirm that E. chaffeensis is endemic to many areas of Long Island and that HME should be considered among the differential diagnoses of the many distinct tick-borne diseases that occur in this region.

One could not come away from a conversation with Michel Lavoipierre without knowing that he was an extraordinary person. He was truly a Renaissance man, a rara avis in an age of stifling overspecialization. He was most notably a gentle and loyal man, faithful to his religion, devoted to his children and students, and steadfast in his pursuit of scientific truth.