Far-western United States Research Articles

Comparative Microbiome Profiles of Sympatric Tick Species from the Far-Western United States.

Insight into the composition and function of the tick microbiome has expanded considerably in recent years. Thus far, tick microbiome studies have focused on species and life stages that are responsible for transmitting disease. In this study we conducted extensive field sampling of six tick species in the far-western United States to comparatively examine the microbial composition of sympatric tick species: Ixodes pacificus, Ixodes angustus, Dermacentor variabilis, Dermacentor occidentalis, Dermacentor albipictus, and Haemaphysalis leporispalustris. These species represent both common vectors of disease and species that rarely encounter humans, exhibiting a range of host preferences and natural history. We found significant differences in microbial species diversity and composition by tick species and life stage. The microbiome of most species examined were dominated by a few primary endosymbionts. Across all species, the relative abundance of these endosymbionts increased with life stage while species richness and diversity decreased with development. Only one species, I. angustus, did not show the presence of a single dominant microbial species indicating the unique physiology of this species or its interaction with the surrounding environment. Tick species that specialize in a small number of host species or habitat ranges exhibited lower microbiome diversity, suggesting that exposure to environmental conditions or host blood meal diversity can affect the tick microbiome which in turn may affect pathogen transmission. These findings reveal important associations between ticks and their microbial community and improve our understanding of the function of non-pathogenic microbiomes in tick physiology and pathogen transmission.

Evidence for transmission of the zoonotic apicomplexan parasite Babesia duncani by the tick Dermacentor albipictus

Babesiosis is a potentially fatal tick-borne zoonotic disease caused by a species complex of blood parasites that can infect a variety of vertebrates, particularly dogs, cattle, and humans. In the United States, human babesiosis is caused by two distinct parasites, Babesia microti and Babesia duncani. The enzootic cycle of B. microti, endemic in the northeastern and upper midwestern regions, has been well characterised. In the western United States, however, the natural reservoir host and tick vector have not been identified for B. duncani, greatly impeding efforts to understand and manage this zoonotic disease. Two and a half decades after B. duncani was first described in a human patient in Washington State, USA, we provide evidence that the enzootic tick vector is the winter tick, Dermacentor albipictus, and the reservoir host is likely the mule deer, Odocoileus hemionus. The broad, overlapping ranges of these two species covers a large portion of far-western North America, and is consistent with confirmed cases of B. duncani in the far-western United States.

Modeling Climate Suitability of the Western Blacklegged Tick in California.

Ixodes pacificus Cooley & Kohls (Acari: Ixodidae), the primary vector of Lyme disease spirochetes to humans in the far-western United States, is broadly distributed across Pacific Coast states, but its distribution is not uniform within this large, ecologically diverse region. To identify areas of suitable habitat, we assembled records of locations throughout California where two or more I. pacificus were collected from vegetation from 1980 to 2014. We then employed ensemble species distribution modeling to identify suitable climatic conditions for the tick and restricted the results to land cover classes where these ticks are typically encountered (i.e., forest, grass, scrub-shrub, riparian). Cold-season temperature and rainfall are particularly important abiotic drivers of suitability, explaining between 50 and 99% of the spatial variability across California among models. The likelihood of an area being classified as suitable increases steadily with increasing temperatures >0°C during the coldest quarter of the year, and further increases when precipitation amounts range from 400 to 800 mm during the coldest quarter, indicating that areas in California with relatively warm and wet winters typically are most suitable for I. pacificus. Other consistent predictors of suitability include increasing autumn humidity, temperatures in the warmest month between 23 and 33°C, and low-temperature variability throughout the year. The resultant climatic suitability maps indicate that coastal California, especially the northern coast, and the western Sierra Nevada foothills have the highest probability of I. pacificus presence.

Human Borrelia miyamotoi infection in California: Serodiagnosis is complicated by multiple endemic Borrelia species.

To determine whether human Borrelia miyamotoi infection occurs in the far-western United States, we tested archived sera from northwestern California residents for antibodies to this emerging relapsing fever spirochete. These residents frequently were exposed to I. pacificus ticks in a region where B. miyamotoi tick infection has been reported. We used a two-step B. miyamotoi rGlpQ assay and a B. miyamotoi whole-cell lysate (WCL) assay to detect B. miyamotoi antibody. We also employed Borrelia hermsii and Borrelia burgdorferi WCL assays to examine if these Borrelia induce cross reacting antibody to B. miyamotoi. Sera were collected from 101 residents in each of two consecutive years. The sera of 12 and 14 residents in years one and two, respectively, were B. miyamotoi rGlpQ seroreactive. Sufficient sera were available to test 15 of the 26 seropositive samples using B. miyamotoi and B. hermsii WCL assays. Two residents in year one and seven residents in year two were seroreactive to both Borrelia antigens. Although discernible differences in seroreactivity were evident between the B. miyamotoi and B. hermsii WCL assays, infection with one or the other could not be determined with certainty. Sera from two Borrelia burgdorferi /B. miyamotoi seropositive subjects reacted strongly against B. miyamotoi and B. hermsii WCL antigens. Ecological, epidemiological, and clinical data implicated B. miyamotoi as the probable cause of infection among those whose sera reacted against both antigens. Our findings suggest that human B. miyamotoi infection occurs in northern California and that B. hermsii and B. burgdorferi infections produce antibodies that cross-react with B. miyamotoi antigens. Health care professionals in the far-western United States should be aware that B. miyamotoi disease may occur throughout the geographic distribution of I. pacificus and that improved relapsing fever group spirochete antibody assays are urgently needed.

Human behaviors elevating the risk of exposure to Ixodes ricinus larvae and nymphs in two types of lowland coniferous forests in west-central Poland

Ixodes ricinus L. is the commonest tick encountered by humans in Central Europe and the most important vector of tick-borne diseases (TBD) in Europe. Foresters represent a group at elevated risk of exposure to I. ricinus throughout Europe. Here, we employed an experimental design similar to one used previously to determine what behavioral factors heighten exposure to questing Ixodes pacificus nymphs in a hardwood forest in the far-western United States. Five activities were evaluated systematically in a coniferous forest and a mixed broadleaved forest in west-central Poland: (i) walking in low-cut running shoes; (ii) sitting atop logs; (iii) sitting on leaf litter; (iv) sitting against tree trunks; and (v) gathering firewood. In total, 540 I. ricinus ticks were collected (198 larvae, 327 nymphs, 15 adults) in spring and late summer. Many more ticks were collected in spring than in late summer, and in the mixed broadleaved forest versus the coniferous forest. The riskiest behavior for acquiring nymphs in both spring and late summer was gathering firewood, though sitting atop logs was nearly as perilous in late summer. In contrast, the riskiest behaviors for encountering larvae in spring and late summer were sitting against trunks and sitting on leaf litter, respectively. Pairwise comparisons revealed only two significant associations among the variables explored for the nymphs: season and human gender, and season and type of forest. For larval ticks, the most significant associations were gender and sitting behaviors and for type of forest and sitting on leaf litter or against tree trunks. The most commonly infested body region was the lower legs.

Borrelia burgdorferi sensu lato spirochetes in wild birds in northwestern California: associations with ecological factors, bird behavior and tick infestation.

Although Borrelia burgdorferi sensu lato (s.l.) are found in a great diversity of vertebrates, most studies in North America have focused on the role of mammals as spirochete reservoir hosts. We investigated the roles of birds as hosts for subadult Ixodes pacificus ticks and potential reservoirs of the Lyme disease spirochete B. burgdorferi sensu stricto (s.s.) in northwestern California. Overall, 623 birds representing 53 species yielded 284 I. pacificus larvae and nymphs. We used generalized linear models and zero-inflated negative binomial models to determine associations of bird behaviors, taxonomic relationships and infestation by I. pacificus with borrelial infection in the birds. Infection status in birds was best explained by taxonomic order, number of infesting nymphs, sampling year, and log-transformed average body weight. Presence and counts of larvae and nymphs could be predicted by ground- or bark-foraging behavior and contact with dense oak woodland. Molecular analysis yielded the first reported detection of Borrelia bissettii in birds. Moreover, our data suggest that the Golden-crowned Sparrow (Zonotrichia atricapilla), a non-resident species, could be an important reservoir for B. burgdorferi s.s. Of 12 individual birds (9 species) that carried B. burgdorferi s.l.-infected larvae, no birds carried the same genospecies of B. burgdorferi s.l. in their blood as were present in the infected larvae removed from them. Possible reasons for this discrepancy are discussed. Our study is the first to explicitly incorporate both taxonomic relationships and behaviors as predictor variables to identify putative avian reservoirs of B. burgdorferi s.l. Our findings underscore the importance of bird behavior to explain local tick infestation and Borrelia infection in these animals, and suggest the potential for bird-mediated geographic spread of vector ticks and spirochetes in the far-western United States.

Detection of Borrelia burgdorferi Sensu Stricto ospC Alleles Associated with Human Lyme Borreliosis Worldwide in Non-Human-Biting Tick Ixodes affinis and Rodent Hosts in Southeastern United States

Comparative analysis of ospC genes from 127 Borrelia burgdorferi sensu stricto strains collected in European and North American regions where Lyme disease is endemic and where it is not endemic revealed a close relatedness of geographically distinct populations. ospC alleles A, B, and L were detected on both continents in vectors and hosts, including humans. Six ospC alleles, A, B, L, Q, R, and V, were prevalent in Europe; 4 of them were detected in samples of human origin. Ten ospC alleles, A, B, D, E3, F, G, H, H3, I3, and M, were identified in the far-western United States. Four ospC alleles, B, G, H, and L, were abundant in the southeastern United States. Here we present the first expanded analysis of ospC alleles of B. burgdorferi strains from the southeastern United States with respect to their relatedness to strains from other North American and European localities. We demonstrate that ospC genotypes commonly associated with human Lyme disease in European and North American regions where the disease is endemic were detected in B. burgdorferi strains isolated from the non-human-biting tick Ixodes affinis and rodent hosts in the southeastern United States. We discovered that some ospC alleles previously known only from Europe are widely distributed in the southeastern United States, a finding that confirms the hypothesis of transoceanic migration of Borrelia species.

Two Boundaries Separate Borrelia burgdorferi Populations in North America

Understanding the spread of infectious diseases is crucial for implementing effective control measures. For this, it is important to obtain information on the contemporary population structure of a disease agent and to infer the evolutionary processes that may have shaped it. Here, we investigate on a continental scale the population structure of Borrelia burgdorferi, the causative agent of Lyme borreliosis (LB), a tick-borne disease, in North America. We test the hypothesis that the observed population structure is congruent with recent population expansions and that these were preceded by bottlenecks mostly likely caused by the near extirpation in the 1900s of hosts required for sustaining tick populations. Multilocus sequence typing and complementary population analytical tools were used to evaluate B. burgdorferi samples collected in the Northeastern, Upper Midwestern, and Far-Western United States and Canada. The spatial distribution of sequence types (STs) and inferred population boundaries suggest that the current populations are geographically separated. One major population boundary separated western B. burgdorferi populations transmitted by Ixodes pacificus in California from Eastern populations transmitted by I. scapularis; the other divided Midwestern and Northeastern populations. However, populations from all three regions were genetically closely related. Together, our findings suggest that although the contemporary populations of North American B. burgdorferi now comprise three geographically separated subpopulations with no or limited gene flow among them, they arose from a common ancestral population. A comparative analysis of the B. burgdorferi outer surface protein C (ospC) gene revealed novel linkages and provides additional insights into the genetic characteristics of strains.

Molecular characterization reveals distinct genospecies of Anaplasma phagocytophilum from diverse North American hosts

Anaplasma phagocytophilum is an emerging tick-borne pathogen that infects humans, domestic animals and wildlife throughout the Holarctic. In the far-western United States, multiple rodent species have been implicated as natural reservoirs for A. phagocytophilum. However, the presence of multiple A. phagocytophilum strains has made it difficult to determine which reservoir hosts pose the greatest risk to humans and domestic animals. Here we characterized three genetic markers (23S-5S rRNA intergenic spacer, ank and groESL) from 73 real-time TaqMan PCR-positive A. phagocytophilum strains infecting multiple rodent and reptile species, as well as a dog and a horse, from California. Bayesian and maximum-likelihood phylogenetic analyses of all three genetic markers consistently identified two major clades, one of which consisted of A. phagocytophilum strains infecting woodrats and the other consisting of strains infecting sciurids (chipmunks and squirrels) as well as the dog and horse strains. In addition, analysis of the 23S-5S rRNA spacer region identified two unique and highly dissimilar clades of A. phagocytophilum strains infecting several lizard species. Our findings indicate that multiple unique strains of A. phagocytophilum with distinct host tropisms exist in California. Future epidemiological studies evaluating human and domestic animal risk should incorporate these distinctions.

Impact of the experimental removal of lizards on Lyme disease risk

The distribution of vector meals in the host community is an important element of understanding and predicting vector-borne disease risk. Lizards (such as the western fence lizard; Sceloporus occidentalis) play a unique role in Lyme disease ecology in the far-western United States. Lizards rather than mammals serve as the blood meal hosts for a large fraction of larval and nymphal western black-legged ticks (Ixodes pacificus--the vector for Lyme disease in that region) but are not competent reservoirs for the pathogen, Borrelia burgdorferi. Prior studies have suggested that the net effect of lizards is to reduce risk of human exposure to Lyme disease, a hypothesis that we tested experimentally. Following experimental removal of lizards, we documented incomplete host switching by larval ticks (5.19%) from lizards to other hosts. Larval tick burdens increased on woodrats, a competent reservoir, but not on deer mice, a less competent pathogen reservoir. However, most larvae failed to find an alternate host. This resulted in significantly lower densities of nymphal ticks the following year. Unexpectedly, the removal of reservoir-incompetent lizards did not cause an increase in nymphal tick infection prevalence. The net result of lizard removal was a decrease in the density of infected nymphal ticks, and therefore a decreased risk to humans of Lyme disease. Our results indicate that an incompetent reservoir for a pathogen may, in fact, increase disease risk through the maintenance of higher vector density and therefore, higher density of infected vectors.

Influence of Abiotic and Environmental Factors on the Density and Infection Prevalence ofIxodes pacificus(Acari: Ixodidae) WithBorrelia burgdorferi

The abiotic and biotic factors that govern the spatial distribution of Lyme disease vectors are poorly understood. This study addressed the influence of abiotic and biotic environmental variables on Ixodes pacificus Cooley & Kohls (Acari:Ixodidae) nymphs, because it is the primary vector of Borrelia burgdorferi Johnson, Schmidt, Hyde, Steigerwaldt & Brenner in the far-western United States. Three metrics of Lyme disease risk were evaluated: the density of nymphs, the density of infected nymphs, and the nymphal infection prevalence. This study sampled randomly located plots in oak (Quercus spp.) woodland habitat in Sonoma County, CA. Each plot was drag-sampled for nymphal ticks and tested for B. burgdorferi infection. Path analysis was used to evaluate the direct and indirect relationship between topographic, forest structure and microclimatic variables on ticks. Significant negative correlations were found between maximum temperature in the dry season and the density of infected ticks in 2006 and tick density in 2007, but we did not find a significant relationship with nymphal infection prevalence in either year. Tick density and infected tick density had an indirect, positive correlation with elevation, mediated through temperature. This study found that in certain years but not others, temperature maxima in the dry season may constrain the density and density of infected I. pacificus nymphs. In other years, biotic or stochastic factors may play a more important role in determining tick density.

A spatially-explicit model of acarological risk of exposure to Borrelia burgdorferi-infected Ixodes pacificus nymphs in northwestern California based on woodland type, temperature, and water vapor

In the far-western United States, the nymphal stage of the western black-legged tick, Ixodes pacificus, has been implicated as the primary vector to humans of Borrelia burgdorferi sensu stricto (hereinafter referred to as B. burgdorferi), the causative agent of Lyme borreliosis in North America. In the present study, we sought to determine if infection prevalence with B. burgdorferi in I. pacificus nymphs and the density of infected nymphs differ between dense-woodland types within Mendocino County, California, and to develop and evaluate a spatially-explicit model for density of infected nymphs in dense woodlands within this high-incidence area for Lyme borreliosis. In total, 4.9% (264) of 5431 I. pacificus nymphs tested for the presence of B. burgdorferi were infected. Among the 78 sampling sites, infection prevalence ranged from 0% to 22% and density of infected nymphs from 0 to 2.04 per 100 m 2. Infection prevalence was highest in woodlands dominated by hardwoods (6.2%) and lowest for redwood (1.9%) and coastal pine (0%). Density of infected nymphs also was higher in hardwood-dominated woodlands than in conifer-dominated ones that included redwood or pine. Our spatial risk model, which yielded an overall accuracy of 85%, indicated that warmer areas with less variation between maximum and minimum monthly water vapor in the air were more likely to include woodlands with elevated acarological risk of exposure to infected nymphs. We found that 37% of dense woodlands in the county were predicted to pose an elevated risk of exposure to infected nymphs, and that 94% of the dense-woodland areas that were predicted to harbor elevated densities of infected nymphs were located on privately-owned land.

Horizontal and vertical movements of host-seeking Ixodes pacificus (Acari: Ixodidae) nymphs in a hardwood forest

The nymph of the western black-legged tick (Ixodes pacificus) is an important bridging vector of the Lyme disease spirochete (Borrelia burgdorferi) to humans in the far-western United States. The previously unknown dispersal capabilities of this life stage were studied in relation to logs, tree trunks, and adjacent leaf-litter areas in a mixed hardwood forest using mark-release-recapture methods. In two spatially and temporally well-spaced trials involving logs, the estimated mean distances that nymphs dispersed ranged from approximately 0.04 to 0.20 m/day on logs vs 0.11 to 0.72 m/day in litter. Prior to recapture in either trial and within the confines of the sampling grids, the greatest estimated dispersal distances by individual nymphs released on logs, and in litter 0.5 m or 1.5 m from logs, were 2.4, 3.0, and 3.0 m, respectively. Nymphs released on logs or litter tended to remain within the same biotopes in which they were freed while host-seeking. In two simultaneous trials involving trunks spaced close-at-hand, nymphs released at the trunk/litter interface on all four aspects collectively dispersed a mean of 0.353 m/day on trunks vs 0.175 m/day in litter. In either trial, the greatest distances that recaptured nymphs climbed trunks, or dispersed in litter in an encircling 3-m grid, were 1.55 m and 2.97 m, respectively. Nymphs ascending trunks did not exhibit a preference for any one aspect, and the B. burgdorferi-infection prevalences in nymphs that climbed trunks (3.2-4.0%) did not differ significantly from those that moved horizontally into litter (10.5-17.6%). We conclude that I. pacificus nymphs use an ambush host-seeking strategy; that they disperse slowly in all biotopes studied; that they usually continue to host-seek in or on whatever substratum they access initially; and that B. burgdorferi-infected nymphs are as likely to move horizontally as vertically when offered a choice.

Effect of Prescribed Fire for Tick Control in California Chaparral

Prescribed fire was investigated as a method for controlling ixodid and argasid ticks in chaparral habitats in northern California. Two experimental and two adjacent control plots within a wildlife preserve were monitored for 1 yr postburn. Ticks were collected by flagging vegetation, by CO2-baited pitfall trap, and by live-trapping rodents. Twice as many rodents were caught at control sites compared with burn sites and no dusky-footed woodrats, Neotoma fuscipes Baird, were found in the treatment sites postburn. This species is known to be a reservoir of the agents of Lyme disease, Borrelia burgdorferi sensu stricto Johnson, Schmid, Hyde, Steigerwalt & Brenner, and human granulocytic anaplasmosis, Anaplasma phagocytophilum Dumler, Barbet, Bekker, Dasch, Palmer, Ray, Rikihisa, Rurangirwa. Six ixodid tick species were removed from rodents (Ixodes pacificus Cooley & Kohls, Ixodes jellisoni Cooley & Kohls, Ixodes spinipalpis Hadwen & Nuttall, Ixodes woodi Bishopp, Dermacentor occidentalis Marx, and Dermacentor parumapertus Neumann), two of which transmit bacterial zoonotic agents to people in the far-western United States. There was no decrease in number of ticks per animal trapped at either burn site compared with controls; in fact, the mean number of immature I. pacificus per rodent was significantly higher at one burn site than its control site. Soil refugia may protect ticks from fire-induced mortality; the argasid tick Ornithodoros coriaceus Koch, which lives in soil, was unaffected by the prescribed fire as were I. pacificus and D. occidentalis buried in packets 2.5 cm below ground. We conclude that although prescribed fires in chaparral habitats may diminish local rodent abundance, it does not decrease tick loads on rodents. Furthermore, burning chaparral does not result in a decreased abundance of adult ixodid ticks on vegetation and apparently does not affect argasid or ixodid ticks that are sheltered within soil refugia.

Host-seeking behavior of Ixodes pacificus (Acari: Ixodidae) nymphs in relation to environmental parameters in dense-woodland and woodland-grass habitats

In the far-western United States, the bacteria that cause Lyme disease (Borrelia burgdorferi, Bb) and human granulocytic anaplasmosis (Anaplasma phagocytophilum, Ap) are transmitted by the western black-legged tick (Ixodes pacificus). In a dense woodland, human behaviors involving contact with wood were recently found to pose greater risk for encountering I. pacificus nymphs than behaviors entailing exclusive exposure to leaf litter. A four-year follow-up study was undertaken in the same woodland and, as a comparison area in one year only, in a nearby woodland-grass habitat to explore the biotic and abiotic factors that might elevate human exposure to host-seeking nymphs. Nymphs were active in the dense woodland throughout the daytime, but no consistent pattern of activity was observed with respect to time of day, temperature, or relative humidity. Significantly more nymphs were collected from the southern aspects of dense-woodland trunks than from other aspects, and more nymphs quested at a height of < or = 1 m vs 1-2 m aboveground. The prevalence of bacterial infection in ticks from the dense woodland was highly variable among years, with maxima of 22.6% and 42.9% for Bb, and 15.6% and 1.8% for Ap, in nymphs from logs and trunks, respectively. The mean densities of nymphs, and of Bb- or Ap-infected nymphs, were typically higher on logs and trunks than in adjoining leaf litter or grass in both habitats. The acarologic risk of encountering an infected nymph on dense-woodland logs or trunks was 2.8 to 11 times higher for Bb than for Ap in two of three years, and it was usually higher in dense woodland than in woodland-grass for both agents. Coinfections were rare (0.27%, n = 369 nymphs from both habitats). Individuals having prolonged contact with logs or trunks in spring would be well advised to employ personal protective measures to minimize exposure to I. pacificus nymphs and their attendant bacterial zoonotic agents.

WILD TURKEY (MELEAGRIS GALLOPAVO) AS A HOST OF IXODID TICKS, LICE, AND LYME DISEASE SPIROCHETES (BORRELIA BURGDORFERI SENSU LATO) IN CALIFORNIA STATE PARKS

Rio Grande wild turkeys (Meleagris gallopavo intermedia) were evaluated as potential hosts of ixodid ticks, lice, and Lyme disease spirochetes (Borrelia burgdorferi sensu lato [s.l.]) in three state parks in Sonoma County, California, USA, during 2003 and 2004. In total, 113 birds were collected, 50 (44.2%) of which were found to be infested by 361 ixodid ticks representing three species: the western black-legged tick (Ixodes pacificus, n=248), the rabbit tick (Haemaphysalis leporispalustris, n=112), and one American dog tick (Dermacentor variabilis). Year-round the prevalence of all ticks combined was unrelated to the age or sex of turkeys, and the prevalence of infestation by I. pacificus (35.4%) was significantly higher than it was for either H. leporispalustris (14.2%) or D. variabilis (0.9%). The proportion of the two prevalent tick species differed significantly by life stage with 86.3% of the I. pacificus and 82.1% of the H. leporispalustris enumerated being nymphs and larvae, respectively. Three species of lice were collected, including the chicken body louse Menacanthus stramineus (12.5% of total), Chelopistes meleagridis (37.5% of total), and Oxylipeurus polytrapezius (50% of total). The records for all three ticks are the first ever from wild turkeys, and those for the lice are the first from this host in the far-western United States. Wild turkeys potentially were exposed to the feeding activities of I. pacificus nymphs infected with B. burgdorferi s.l. as 15% of host-seeking nymphs (n=200) collected in woodlands used by turkeys as roosting or foraging areas were infected mainly with B. burgdorferi sensu stricto (s.s.). However, only one (1%) of 90 turkey blood specimens tested by PCR contained B. burgdorferi s.s., and four in vitro, complement-protein assays demonstrated that domestic turkey serum is moderately bacteriolytic for this spirochete. Taken together, these findings indicate that wild turkeys are important avian hosts of I. pacificus nymphs, but they appear to be inconsequential hosts of B. burgdorferi s.l.

Western Gray Squirrel (Rodentia: Sciuridae): A Primary Reservoir Host of &lt;I&gt;Borrelia burgdorferi&lt;/I&gt; in Californian Oak Woodlands?

Abstract In California, dense woodlands have been recognized as important biotopes where humans are exposed to the nymphal stage of the western blacklegged tick, Ixodes pacificus Cooley & Kohls, the primary vector of the Lyme disease spirochete Borrelia burgdorferi sensu stricto (s.s.), in the far-western United States. To identify the principal reservoir host(s) of this spirochete, and of closely related spirochetes in the B. burgdorferi sensu lato (s.l.) complex, in dense woodlands in Mendocino County, California, ≈50 species of birds and mammals, including wood rats and kangaroo rats, were evaluated as potential hosts for vector ticks and borreliae in 2002 and 2003. Although polymerase chain reaction (PCR) and sequencing analyses revealed that many vertebrate species had been exposed to one or more members of the B. burgdorferi s.l. spirochetal complex, only the western gray squirrel, Sciurus griseus, fulfilled the major criteria for a reservoir host of B. burgdorferi s.s. Ear-punch biopsies from eight...

Western Gray Squirrel (Rodentia: Sciuridae): A Primary Reservoir Host ofBorrelia burgdorferiin Californian Oak Woodlands?

In California, dense woodlands have been recognized as important biotopes where humans are exposed to the nymphal stage of the western blacklegged tick, Ixodes pacificus Cooley & Kohls, the primary vector of the Lyme disease spirochete Borrelia burgdorferi sensu stricto (s.s.), in the far-western United States. To identify the principal reservoir host(s) of this spirochete, and of closely related spirochetes in the B. burgdorferi sensu lato (s.l.) complex, in dense woodlands in Mendocino County, California, approximately 50 species of birds and mammals, including wood rats and kangaroo rats, were evaluated as potential hosts for vector ticks and borreliae in 2002 and 2003. Although polymerase chain reaction (PCR) and sequencing analyses revealed that many vertebrate species had been exposed to one or more members of the B. burgdorferi s.l. spirochetal complex, only the western gray squirrel, Sciurus griseus, fulfilled the major criteria for a reservoir host of B. burgdorferi s.s. Ear-punch biopsies from eight of 10 squirrels collected from five separate woodlands were PCR-positive for B. burgdorferi s.s., 47% of I. pacificus larvae (n = 64) and 31% of nymphs (n = 49) removed from squirrels contained B. burgdorferi s.l., and the engorgement status of I. pacificus larvae was associated positively with acquisition of spirochetes. Overall, 83 and 100% of the amplicons sequenced from PCR-positive I. pacificus larvae and nymphs, respectively, were identified as B. burgdorferi s.s, Among the five remaining positive I. pacificus larvae, three contained B. bissettii and two had uncharacterized B. burgdorferi s.l. Borrelia burgdorferi s.s. was detected in one of five larvae and zero of two nymphs of the Pacific Coast tick, Dermacentor occidentalis Marx, that likewise had been removed from squirrels. The rickettsial agent of human anaplasmosis, Anaplasma phagocytophilum, was detected in the blood or ear biopsies of two squirrels and in one (1.6%) of 64 I. pacificus larvae and two (4.1%) of 49 nymphs obtained from squirrels. The one rickettsial-positive larva was coinfected with B. burgdorferi s.s. The apparently high reservoir potential of S. griseus for B. burgdorferi s.s., plus the fact that the geographic distribution of this squirrel coincides well with that of most reported human cases of Lyme disease in this region, indicated that it may be essential for maintaining foci of B. burgdorferi s.s. in certain types of woodlands. The findings with respect to A. phagocytophilum, although of less certain significance, suggest that S. griseus could serve as a secondary host of this rickettsia.

Remote Sensing (Normalized Difference Vegetation Index) Classification of Risk Versus Minimal Risk Habitats for Human Exposure toIxodes pacificus(Acari: Ixodidae) Nymphs in Mendocino County, California

In California, Ixodes pacificus Cooley & Kohls nymphs have been implicated as the primary bridging vectors to humans of the spirochetal bacterium causing Lyme disease (Borrelia burgdorferi). Because the nymphs typically do not ascend emergent vegetation, risk of human exposure is minimal in grasslands, chaparral, and woodland-grass. Instead, woodlands with a ground cover dominated by leaf litter (hereinafter referred to as woodland-leaf) have emerged as a primary risk habitat for exposure to B. burgdorferi-infected nymphs. As a means of differentiating woodland-leaf habitats from others with minimal risk (e.g., chaparral, grassland, and woodland-grass), we constructed a maximum likelihood model of these habitat types within a 7,711-ha area in southeastern Mendocino County based on the normalized difference vegetation index derived from Landsat 5 Thematic Mapper imagery (based on a 30 by 30-m pixel size) over four seasons. The overall accuracy of the model to discriminate woodland-leaf, woodland-grass, open grassland, and chaparral was 83.85% (Kappa coefficient of 0.78). Validation of the accuracy of the model to classify woodland-leaf yielded high values both for producer accuracy (93.33% of validated woodland-leaf pixels correctly classified by the model) and user accuracy (96.55% of model-classified validation pixels correctly categorized as woodland-leaf). Woodland-leaf habitats were found to be highly aggregated within the examined area. In conclusion, our model successfully used remotely sensed data as a predictor of habitats where humans are at risk for Lyme disease in the far-western United States.

Research on animal-borne parasites and pathogens helps prevent human disease

A large body of basic and applied research conducted at the UC Hopland Research and Extension Center (HREC) has focused on micro- or macroparasites that infest humans, domestic animals and wildlife, including viruses, bacteria, protozoans, helminths and blood-sucking arthropods. Approximately 17 microorganisms detected at the HREC are transmitted by ticks, and of these, at least five cause zoonotic diseases — animal-borne diseases that can be transmitted to people. Scientists working at the HREC identified the basic transmission cycle of the Lyme disease spirochete in the far-western United States. This information now is being used to develop and implement risk-assessment and preventive strategies.