Pacificus Adults Research Articles

Surveillance for Ixodes scapularis and Ixodes pacificus ticks and their associated pathogens in Canada, 2019.

The primary vectors of the agent of Lyme disease in Canada are Ixodes scapularis and Ixodes pacificus ticks. Surveillance for ticks and the pathogens they can transmit can inform local tick-borne disease risk and guide public health interventions. The objective of this article is to characterize passive and active surveillance of the main Lyme disease tick vectors in Canada in 2019 and the tick-borne pathogens they carry. Passive surveillance data were compiled from the National Microbiology Laboratory Branch and provincial public health data sources. Active surveillance was conducted in selected sentinel sites in all provinces. Descriptive analysis of ticks submitted and infection prevalence of tick-borne pathogens are presented. Seasonal and spatial trends are also described. In passive surveillance, specimens of I. scapularis (n=9,858) were submitted from all provinces except British Columbia and I. pacificus (n=691) were submitted in British Columbia and Alberta. No ticks were submitted from the territories. The seasonal distribution pattern was bimodal for I. scapularis adults, but unimodal for I. pacificus adults. Borrelia burgdorferi was the most prevalent pathogen in I. scapularis (18.8%) and I. pacificus (0.3%). In active surveillance, B. burgdorferi was identified in 26.2% of I. scapularis; Anaplasma phagocytophilum in 3.4% of I. scapularis, and Borrelia miyamotoi and Powassan virus in 0.5% or fewer of I. scapularis. These same tick-borne pathogens were not found in the small number of I. pacificus tested. This surveillance article provides a snapshot of the main Lyme disease vectors in Canada and their associated pathogens, which can be used to monitor emerging risk areas for exposure to tick-borne pathogens.

Detection and Isolation of Rickettsia tillamookensis (Rickettsiales: Rickettsiaceae) From Ixodes pacificus (Acari: Ixodidae) From Multiple Regions of California.

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.

Phylogeography of Borrelia spirochetes in Ixodes pacificus and Ixodes spinipalpis ticks highlights differential acarological risk of tick-borne disease transmission in northern versus southern California.

The common human-biting tick, Ixodes pacificus, is the primary vector of the Lyme disease spirochete, Borrelia burgdorferi sensu stricto (ss) in western North America and has been found to harbor other closely-related spirochetes in the Borrelia burgdorferi sensu lato (sl) complex. Between 2008–2015, 11,066 adult and 3,815 nymphal I. pacificus and five adult and 144 nymphal Ixodes spinpalpis, a commonly collected wildlife tick, were collected from 42 California counties. Borrelia burgdorferi sl was detected in 1.2% and 3.8% I. pacificus adults and nymphs, respectively. Results from this study indicate genetic diversity and geographic structure of B. burgdorferi sl in California I. pacificus ticks, by sequence comparison of the16S rRNA gene, with B. burgdorferi ss, the agent of Lyme disease, found only in I. pacificus collected from the north and central coastal and Sierra Nevada foothill regions; B. burgdorferi ss was not detected in ticks tested from southern California. In contrast, Borrelia bissettiae, a member of the B. burgdorferi sl complex, was detected in both I. pacificus and I. spinipalpis, in the coastal region of both northern and southern California, but was absent from ticks in the Sierra Nevada foothills. In a similar pattern to B. bissettiae, Borrelia americana (a member of the B. burgdorferi sl complex) was detected in a single adult I. pacificus from the north coast and two I. spinipalpis nymphs from south-coastal California. This study highlights that the geographic area of Lyme disease acarological risk in California is the north-central and Sierra Nevada foothill regions of the state with little to no risk in the southern regions of the state.

Lyme disease risk in southern California: abiotic and environmental drivers of Ixodes pacificus (Acari: Ixodidae) density and infection prevalence with Borrelia burgdorferi

BackgroundTick-borne diseases, particularly Lyme disease, are emerging across the northern hemisphere. In order to manage emerging diseases and predict where emergence will likely occur, it is necessary to understand the factors influencing the distribution, abundance and infection prevalence of vector species. In North America, Lyme disease is the most common vector-borne disease and is transmitted by blacklegged ticks. This study aimed to explore the abiotic and environmental drivers of density and infection prevalence of western blacklegged ticks (Ixodes pacificus) in southern California, an understudied and densely populated region of North America.ResultsOver the course of this two-year study, densities of I. pacificus adults were consistently positively associated with host availability for juvenile ticks and dense oak woodland habitat. Densities of nymphal and larval I. pacificus, on the other hand were primarily predicted by host availability for juvenile ticks in the first year of the study, and by habitat characteristics such as dense leaf litter in the second year. Infection with the causative agent of Lyme disease, Borrelia burgdorferi (sensu stricto), and related spirochetes was not predicted by the abiotic conditions promoting I. pacificus populations, but rather by diversity of the tick community, and in particular by the presence of two Ixodes tick species that do not generally feed on humans (Ixodes spinipalpis and Ixodes peromysci). Borrelia spp. infection was not detected in the I. pacificus populations sampled, but was detected in other vector species that may maintain enzootic transmission of the pathogen on the landscape.ConclusionsThis study identified dense oak woodlands as high-risk habitats for I. pacificus tick encounter in southern California. The shift in relative importance of host availability to habitat characteristics in predicting juvenile tick abundance occurred as California’s historic drought intensified, suggesting that habitat providing suitable microclimates for tick survivorship became centrally important to patterns of abundance in the face of deleterious abiotic conditions. These results underscore the need for further investigation of the effects of climate change on tick-borne disease in California. Finally, despite low risk of human Lyme disease infection posed by I. pacificus in southern California, evidence of infection was found in other tick species, suggesting that enzootic transmission of tick-borne borreliae may be occurring in southern California, and involve parallel enzootic cycles with other tick and host species but not necessarily humans.

A host beetle pheromone regulates development and behavior in the nematode Pristionchus pacificus.

Nematodes and insects are the two most speciose animal phyla and nematode-insect associations encompass widespread biological interactions. To dissect the chemical signals and the genes mediating this association, we investigated the effect of an oriental beetle sex pheromone on the development and behavior of the nematode Pristionchus pacificus. We found that while the beetle pheromone is attractive to P. pacificus adults, the pheromone arrests embryo development, paralyzes J2 larva, and inhibits exit of dauer larvae. To uncover the mechanism that regulates insect pheromone sensitivity, a newly identified mutant, Ppa-obi-1, is used to reveal the molecular links between altered attraction towards the beetle pheromone, as well as hypersensitivity to its paralyzing effects. Ppa-obi-1 encodes lipid-binding domains and reaches its highest expression in various cell types, including the amphid neuron sheath and excretory cells. Our data suggest that the beetle host pheromone may be a species-specific volatile synomone that co-evolved with necromeny.

Eco-epidemiological factors contributing to the low risk of human exposure to ixodid tick-borne borreliae in southern California, USA

Little is known about the eco-epidemiology of Lyme disease in southern California, a region where the incidence is much lower than it is in northern California. Here, we sought to discover the previously unknown microhabitats of nymphs of the primary vector, the western black-legged tick (Ixodes pacificus), in 3 moderately to heavily-utilized state parks in the Santa Monica Mountains in Los Angeles County; to elucidate the seasonal distribution and abundance of adults of I. pacificus and another human-biter, the Pacific Coast tick (Dermacentor occidentalis); and to determine what Lyme-disease or relapsing-fever group borreliae are present in questing nymphs or adult ticks. I. pacificus nymphs were collected infrequently at various times of day in 2 chaparral or 7 woodland litter areas by dragging (combined mean=0.4 nymphs per hour). The western fence lizard (Sceloporus occidentalis) was a choice sentinel animal for detecting the presence of I. pacificus nymphs (and larvae) in diverse biotopes even when dragging litter in them was fruitless. The abundance and seasonality of I. pacificus and D. occidentalis adults resembled what had been documented previously for these ticks in northern California. Overall, zero of 27 free-living and 118 lizard-infesting I. pacificus nymphs, 7 (0.29%) of 2392 I. pacificus adults and 2 (0.22%) of 896 D. occidentalis adults were infected with Borrelia burgdorferi sensu lato (Bb sl), but none of them harbored B. miyamotoi, a relapsing-fever group spirochete implicated recently as a zoonotic pathogen in Russia. Borrelia americana and the human pathogen B. burgdorferi sensu stricto were each detected in one (0.04%), and uncharacterized Bb sl in 5 adult I. pacificus (0.21%) that clustered with B. americana. Both PCR-positive D. occidentalis adults contained B. burgdorferi sensu stricto. We conclude that the acarologic risk of being bitten by a B. burgdorferi sensu lato-infected ixodid tick in the habitats studied is slight, which offers a plausible explanation for the low reported incidence of Lyme disease in south coastal California.

Survival rates of immature Ixodes pacificus (Acari: Ixodidae) ticks estimated using field-placed enclosures

Survival rates of immature Ixodes pacificus (Acari: Ixodidae) ticks estimated using field-placed enclosures

ECOLOGY OF ANAPLASMA PHAGOCYTOPHILUM INFECTION IN GRAY FOXES (UROCYON CINEREOARGENTEUS) IN NORTHWESTERN CALIFORNIA

Although granulocytic anaplasmosis, caused by infection of Anaplasma phagocytophilum, is an emerging human and domestic animal disease, the ecology and natural history of the parasite is not well understood. Gray foxes (Urocyon cinereoargenteus) are relatively common, occasionally peri-urban mesocarnivores whose geographic distribution overlaps the reported distribution of granulocytic anaplasmosis in humans and domestic animals in North America. We evaluated the potential of foxes as hosts and reservoirs of A. phagocytophilum in both urban and backcountry habitats of the Hoopa Valley Indian Reservation, Humboldt County, California, USA. We trapped 54 individual foxes and had 16 recaptures for a total of 70 fox samples between June 2003 and October 2004 in delineated urban and backcountry zones. We collected 296 adult and 145 nymphal ticks from the 70 captured foxes including 193 Ixodes pacificus, 149 Ixodes texanus, 98 Dermacentor variabilis, and one Dermacentor occidentalis. There were seasonal differences in tick intensities, with most I. pacificus adults occurring in winter and spring (P < 0.001), most I. texanus nymphs in spring (P = 0.03), and most D. variabilis adults in spring and summer (P = 0.01). Thirty-six (51%) of the 70 fox sera had antibodies against A. phagocytophilum, with a higher (P = 0.24) prevalence in backcountry foxes (16 of 23) than in urban-zone foxes (12 of 31). Six (9%) of 70 fox samples were polymerase chain reaction-positive for A. phagocytophilum. Twenty-eight (31%) of 90 domestic dogs sampled from vaccine clinics within the study area were seropositive for A. phagocytophilum. There was a significant difference in prevalence between dogs and backcountry foxes (70%), but no differences were found between dogs and urban foxes (39%). We propose that gray foxes are a good sentinel species for A. phagocytophilum infections in northwestern California.

Acarologic Risk of Exposure to Emerging Tick-Borne Bacterial Pathogens in a Semirural Community in Northern California

An acarologic study was conducted in a semirural community in northern California to determine the relative abundance of, and the prevalence of infection with, three emerging bacterial pathogens in the western black-legged tick (Ixodes pacificus). These included the agents causing Lyme disease (Borrelia burgdorferi), human granulocytic ehrlichiosis [Ehrlichia phagocytophila (formerly Ehrlichia equi)], and human monocytic ehrlichiosis (Ehrlichia chaffeensis). The study area in Sonoma County consisted of two properties each with four residents and an uninhabited adjacent comparison area. Six of the eight residents had been either physician-diagnosed or serodiagnosed previously with Lyme disease, and, of these, one also had been serodiagnosed with human monocytic ehrlichiosis. Direct immunofluorescent/culture assays and bacterial species-specific polymerase chain reaction assays were used to test whole ticks individually for presence of B. burgdorferi and Ehrlichia spp., respectively. Overall, 6.5% of the nymphal (n = 589) and 1.6% of the adult ticks (n = 318) from the same generational cohort were found to contain B. burgdorferi. In contrast, none of 465 nymphs and 9.9% of 202 adults were infected with E. phagocytophila. Excised tissues from another 95 adult ticks yielded a comparable E. phagocytophila infection prevalence of 13.7%. E. chaffeensis was not detected in either nymphal or adult ticks. Using a combination of culture and polymerase chain reaction assays, coinfection of I. pacificus adults with B. burgdorferi and E. phagocytophila was demonstrated for the first time. The marked disparity in the infection prevalence of these pathogens in nymphal and adult ticks suggests that their maintenance cycles are inherently different.

Ecology of Borrelia burgdorferi in ticks (Acari: Ixodidae), rodents, and birds in the Sierra Nevada foothills, Placer County, California.

This study examined the prevalence of Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner in host-seeking adult and nymphal Ixodes pacificus Cooley & Kohls and estimated the I. pacificus infestation and B. burgdorferi infection of rodent and avian hosts in the western Sierra Nevada foothills of northern California. Additionally, we identified species likely to participate in an enzootic cycle for B. burgdorferi in this yellow pine transition habitat. Evidence of infection with B. burgdorferi was identified in 7.3 and 5.4% of host-seeking I. pacificus adults and nymphs, respectively. Mean numbers of I. pacificus observed on rodents were 1.15 for Neotoma fuscipes Baird and 0.18 for Peromyscus spp. One of 104 ear punch tissues obtained from woodrats and none from 49 Peromyscus spp. yielded B. burgdorferi. A total of 291 collected birds representing 34 species had a mean of 0.27 I. pacificus per bird. The mean I. pacificus infestation of ground-dwelling birds was 2.5 ticks per bird. Forty-nine of 92 (53%) blood smears collected from birds were reactive to a B. burgdorferi specific antibody. This study presents the identification of a B. burgdorferi-like spirochete in birds in western North America. The tick burden and spirochete infection of birds suggests that birds may be involved in a local B. burgdorferi enzootic cycle and likely participate in the transport of ticks and spirochetes to other locations while rodents from this site do not appear to be major contributors.

Spatial and Temporal Variation in the Density of &lt;I&gt;Ixodes pacificus&lt;/I&gt; (Acari: Ixodidae) Nymphs

The nymphal and adult life stages of the tick Ixodes pacificus Cooley & Kohls are the primary vectors of the causative agent(s) of Lyme disease in the far-western United States. In contrast to I. pacificus adults, data on the extent of spatial and temporal variation in the density of the nymphal stage have been scarce. Therefore, we compared the density of I. pacificus nymphs from 1997 to 1999 at a small rural community at high risk for Lyme disease (CHR) and the University of California Hopland Research and Extension Center (HREC), Mendocino County, CA, I. pacificus nymphs were collected readily by drag sampling in leaf litter but not from low vegetation. The mean number of questing nymphs per 100 m2 in leaf/fir-needle litter areas from late April to early June differed significantly among years at both the CHR and the HREC. Climatic data from the HREC suggested that yearly nymphal densities in this area may be positively correlated with the amount of rainfall and negatively correlated with maximum temperatures from March to May. Further, nymphal density was two to four times higher at the CHR than the HREC in all 3 yr. Yearly mean nymphal density generally differed two- to four-fold (HREC) and 10- to 20-fold (CHR) among individual sampling areas. Also, two- to three-fold differences in mean nymphal density were frequently observed between continuous litter and litter edges bordering on other habitats within sampling areas at the HREC, and between sampling areas <100 m apart at the CHR. The distributions of nymphs within individual areas were generally aggregated at both the HREC and the CHR, at the 15-m sampling transect scale we used. Environmental factors with some potential to predict the density of I. pacificus nymphs at different spatial scales included climatic conditions, topographic exposure, and presence of habitat edges or logs. However, stepwise multiple regression analysis revealed that only 6.2–18.0% (HREC) or 14.9–27.9% (CHR) of the yearly variation in nymphal densities was explained by the abiotic or biotic traits measured in this study, at the 15-m transect scale. Thus, other abiotic or biotic traits that we did not examine (e.g., local densities of tick hosts) must account for most of the spatial variation in nymphal density.

Density and spatial distribution of Ixodes pacificus (Acari: Ixodidae) in two recreational areas in north coastal California.

The density and distribution of Ixodes pacificus was assessed at 2 parks in north coastal California. The density of I. pacificus adults and nymphs varied significantly between years, trails, and sides of trails. Adult ticks occurred on vegetation along sun-exposed trails in January through March, their density (0-1.93 per 20 m) correlated with brush density, trail width, and presence of an uphill slope. Nymphs (0.06-5.10 per 20 m) occurred in leaf litter along shaded trails in May-July. Adult I. pacificus were rare at picnic sites (0.00-0.24 per 20 m), but nymphal densities (0.93-2.37 per 20 m) were comparable with those along some shaded trails. The prevalence of Borrelia burgdorferi in ticks (2.8% overall) did not differ significantly between locations, years, or stages. We conclude that the risk of acquiring Lyme disease in these sites is low, but varies among trails, seasons, and years.

Dusky-footed wood rats (Neotoma fuscipes) as reservoirs of granulocytic Ehrlichiae (Rickettsiales: Ehrlichieae) in northern California.

Dusky-footed wood rats (Neotoma fuscipes) and Peromyscus sp. mice (P. maniculatus and P. truei) were collected from one site in Placer County, one site in Santa Cruz County, and two sites in Sonoma County in northern California. Serum or plasma samples from 260 rodents were tested for antibodies to the agent of human granulocytic ehrlichiosis. Of these, samples from 25 wood rats (34% of those tested) and 10 (8%) Peromyscus sp. mice were found to be seropositive, but only those from one site. PCR assays targeting the groESL heat shock operon were conducted on all seropositive specimens and a subset of seronegative blood specimens. Ehrlichial DNA was identified in 17 (68%) of the 25 seropositive wood rat blood samples and in 1 of the 10 (10%) Peromyscus sp. specimens. None of 40 seronegative blood samples was PCR positive. Both seropositive and PCR-positive animals were collected during each trapping period. One male tick out of 84 Ixodes pacificus adults collected was PCR positive; samples of Dermacentor occidentalis nymphs and adults were negative. Nucleotide sequences of amplicons from three wood rat blood specimens and from the single PCR-positive tick differed by one and two bases, respectively, from a sequence previously obtained from Ehrlichia equi. At one site in Sonoma County, wood rats had a concurrent high prevalence of seropositivity and PCR positivity, while other sigmodontine rodents collected at the site were only occasionally infected. We suggest that dusky-footed wood rats serve as reservoirs of granulocytic ehrlichial agents in certain areas of northern California. The tick species involved in the transmission of granulocytic ehrlichiae among wood rats remains unknown.

Detection of the agents of human ehrlichioses in ixodid ticks from California.

A study was conducted in northern California to estimate the prevalence and distribution in ixodid ticks of the rickettsial agents of human monocytic (HME) and human granulocytic (HGE) ehrlichioses. More than 650 ixodid ticks were collected from 17 sites in six California counties over a 15-month period. Ehrlichia chaffeensis, the causative agent of HME, was detected by a nested polymerase chain reaction (PCR) in Ixodes pacificus (minimum infection rate [MIR] = 13.3%) and Dermacentor variabilis (infection rate=20.0%) from a municipal park in Santa Cruz County. The HGE agent was detected by nested PCR in I. pacificus adults from a heavily used recreational area in Alameda County (MIR = 4.7%) and a semirural community in Sonoma County (MIR = 6.7%). Evidence of infection with Ehrlichia spp. was not detected in D. occidentalis adults or I. pacificus nymphs. This study represents the first detection of E. chaffeensis in California ticks and the first report of infection in Ixodes spp. The competency of I. pacificus to be coinfected with and to transmit multiple disease agents, including those of human ehrlichioses and Lyme disease, has yet to be determined.

Temporal and Spatial Distribution of Ixodes pacificus and Dermacentor occidentalis (Acari: Ixodidae) and Prevalence of Borrelia burgdorferi in Contra Costa County, California

The seasonal activity and spatial distribution of adult and immature Ixodes pacificus Cooley & Kohls and Dermacentor occidentalis Marx were determined along trails and on hillsides in two parks in Contra Costa County, CA. I. pacificus and D. occidentalis adults were most numerous in January and May, respectively. Adult ticks were significantly more abundant along heavily vegetated trails than on open grassy hillsides, and on the uphill versus the downhill side of trails. Five species of rodents were captured, and numbers of I. pacificus and D. occidentalis larvae per rodent were highest in May-June and July, respectively. Few nymphs were recovered either by flagging or from captured rodents. An average of 2.2 and 2.8% of the I. pacificus adults collected from the two parks were infected with the Lyme disease spirochete, Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner. The greatest risk of contracting Lyme disease from adult I. pacificus in these two Contra Costa County parks is during the winter months, especially while hiking near the uphill side of trails.

Lyme borreliosis in California. Acarological, clinical, and epidemiological studies.

The relative abundance of, and spirochetal-infection rates in, adult ixodid ticks from eight Lyme borreliosis clinical-case areas and two comparison areas were investigated in northern California from late fall to early spring, 1984-87. The western black-legged tick (Ixodes pacificus) was the most abundant species at seven of nine sites yielding specimens as determined with a tick drag method. The Pacific Coast tick (Dermacentor occidentalis) was the most abundant species at two sites, and lesser numbers of this tick and the American dog tick (D. variabilis) were obtained from seven and two sites, respectively. Abundance of I. pacificus adults varied in clinical-case areas as well as in the comparison areas, and was not correlated significantly with spirochetal infection rates in this tick. Overall, spirochetes were detected in 1.4 and 1.0% of the adult I. pacificus collected from clinical case (n = 857) and comparison sites (n = 383), respectively, and in 0.8% of adult D. occidentalis (n = 253) from one comparison area. An additional 244 D. occidentalis adults from all other sites were tested with negative results. Five spirochetal isolates recovered from I. pacificus adults were identified as Borrelia burgdorferi with specific monoclonal antibodies. Seven of 10 patients interviewed reportedly contracted their infections in summer, and six presented with a history of tick bite. Nine patients owned or occasionally harbored one or more dogs, and at least one of the dogs contracted Lyme borreliosis at the same site as its owner. Clinical manifestations of the disease in human patients included erythema migrans (100%), arthritis/arthralgia (60%), neurologic abnormalities (60%), and cardiac involvement (20%). Four of nine patients whose sera were assayed by indirect immunofluorescence contained significant antibody titers to B. burgdorferi.