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.     

Abundance of Ticks (Acari: Ixodidae) Infesting the Western Fence Lizard, Sceloporus Occidentalis, in Relation to Environmental Factors

We examined the impact of environmental characteristics, such as habitat type, topographic exposure and presence of leaf litter, on the abundance of Ixodes pacificus ticks infesting the western fence lizard (Sceloporus occidentalis) at the University of California Hopland Research and Extension Center (HREC), Mendocino County, California. A total of 383 adult lizards were slip-noosed and examined for tick infestation in April and May 1998. At least 94% of the lizards were infested by ticks and at least 20% of the females and 33% of the males carried > 15 ticks. This intensive utilization of western fence lizards (which do not serve as natural reservoirs for Lyme disease spirochetes) by subadult ticks, is probably the primary reason for the low prevalence of infection with Borrelia burgdorferi in I. pacificus nymphs and adults previously recorded at the HREC. Tick loads were higher on male than female lizards. Also, male lizards were generally more heavily infested in late April than in late May. The prevalence of tick infestation exceeded 88% in all habitat types but males collected in woodland and grass/woodland edges had higher tick loads than those collected in open grassland. Male lizards captured in open, exposed grassland tended to carry heavier tick loads in northern/eastern, as compared to southern/western, exposures, and when leaf litter was present.     

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.     

Habitat-related variation in infestation of lizards and rodents with Ixodes ticks in dense woodlands in Mendocino County, California

During the spring and early summer of 2002, we examined the relative importance of Borrelia-refractory lizards (Sceloporus occidentalis, Elgaria spp.) versus potential Borrelia burgdorferi sensu lato (s.l.)-reservoirs (rodents) as hosts for Ixodes pacificus immatures in 14 woodland areas (six oak, five mixed oak/Douglas fir, and three redwood/tanoak areas) distributed throughout Mendocino County, California. Lizards were estimated to serve as hosts for 93-98% of all larvae and > or =99.6% of all nymphs infesting lizards or rodents in oak woodlands and oak/Douglas fir sites in the southern part of the county. In redwood/tanoak woodlands and oak/Douglas fir sites in northern Mendocino County, the contribution of rodents to larval feedings reached 36-69% but lizards still accounted for 94-100% of nymphal bloodmeals. From late April to mid-June, I. pacificus larvae were recovered from 95 to 96% of lizards and dusky-footed woodrats (Neotoma fuscipes) and from 59% of Peromyscus spp. mice. In contrast, 99% of lizards but few woodrats (15%) and none of the mice were infested by nymphs. Comparisons of tick loads for 19 lizard-Peromyscus spp. mouse pairings, where the lizard and mouse were captured within 10m of each other, revealed that the lizards harbored 36 times more larvae and >190 times more nymphs than the mice. In oak woodlands, loads of I. pacificus larvae decreased from late April/early May to late June for S. occidentalis lizards but increased for Peromyscus spp. mice. We conclude that the relative utilization of Borrelia-refractory lizards, as compared to rodents, by I. pacificus larvae was far higher in dry oak woodlands than in moister habitats such as redwood/tanoak and oak/Douglas fir woodlands in northern Mendocino County. Non-lizard-infesting potential enzootic vectors of B. burgdorferi s.l. (I. angustus and I. spinipalpis) were recorded from rodents in three of six oak woodland areas, two of five oak/Douglas fir woodland areas, and two of three redwood/tanoak woodland areas.     

Lyme disease and migrating birds in the Saint Croix River Valley

During a study of migrating land birds in 1987, we examined over 9,200 individual birds representing 99 species from the Saint Croix River Valley, a Lyme disease-endemic area of east central Minnesota and northwestern Wisconsin. We found that 250 deer tick (Ixodes dammini) larvae and nymphs infested 58 birds from 15 migrant species; 56 ticks (22.4%) were positive for the Lyme disease spirochete Borrelia burgdorferi. Five ground-foraging migrant bird species favoring mesic habitats, veery (Catharus fuscescens), ovenbird (Seiurus aurocapillus), northern waterthrush (S. novaboracensis), common yellowthroat (Geothlypis trichas), and swamp sparrow (Melospiza georgiana), accounted for nearly three-quarters of parasitized individuals. Nearly half of the spirochete-positive ticks were removed from migrating birds taken in a riparian floodplain forest. Recaptured migrants with infected ticks indicate that they transmit B. burgdorferi to hexapod larvae. We suggest that birds may be both an important local reservoir in the upper Mississippi Valley and long-distance dispersal agents for B. burgdorferi-infected ticks to other regions of the continent.     

Lyme disease and migrating birds in the Saint Croix River Valley.

During a study of migrating land birds in 1987, we examined over 9,200 individual birds representing 99 species from the Saint Croix River Valley, a Lyme disease-endemic area of east central Minnesota and northwestern Wisconsin. We found that 250 deer tick (Ixodes dammini) larvae and nymphs infested 58 birds from 15 migrant species; 56 ticks (22.4%) were positive for the Lyme disease spirochete Borrelia burgdorferi. Five ground-foraging migrant bird species favoring mesic habitats, veery (Catharus fuscescens), ovenbird (Seiurus aurocapillus), northern waterthrush (S. novaboracensis), common yellowthroat (Geothlypis trichas), and swamp sparrow (Melospiza georgiana), accounted for nearly three-quarters of parasitized individuals. Nearly half of the spirochete-positive ticks were removed from migrating birds taken in a riparian floodplain forest. Recaptured migrants with infected ticks indicate that they transmit B. burgdorferi to hexapod larvae. We suggest that birds may be both an important local reservoir in the upper Mississippi Valley and long-distance dispersal agents for B. burgdorferi-infected ticks to other regions of the continent.     

The roles of birds, lizards, and rodents as hosts for the western black-legged tick Ixodes pacificus.

We compared the infestation by ixodid ticks of lizards, rodents, and birds collected simultaneously within areas representing common habitat types in Mendocino County, CA. Lizards were infested only by Ixodes pacificus Cooley and Kohls, birds by I. pacificus and Haemaphysalis leporispalustris (Packard), and rodents by I. pacificus, I. spinipalpis Hadwen and Nuttall, I. woodi Bishopp, Dermacentor occidentalis Marx, and D. variabilis (Say). Infestation by I. pacificus larvae and nymphs of lizards (Sceloporus occidentalis Baird and Girard; Elgaria spp.) and western gray squirrels (Sciurus griseus Ord) (means of 9-35 larvae and 5-6 nymphs per animal) was several times greater than for Neotoma fuscipes Baird woodrats, Peromyscus spp. mice, and birds (means of 0.9-3.5 larvae and 0-0.3 nymphs). Overall, Borrelia-refractory lizards accounted for 84% of I. pacificus larvae and 91% of nymphs collected from animals in dense woodlands. Bird species frequently utilizing tick-questing substrates such as leaf litter (guild I birds) were more heavily infested by I. pacificus subadults (5.2 larvae and 1.0 nymphs per bird) than guild IV birds with minimal perceived contact with tick-questing substrates (0.08 larvae and 0.06 nymphs per bird). Notably, guild I birds carried similar larval loads and at least 20-fold higher nymphal loads relative to woodrats and mice. Only guild IV birds carried as few I. pacificus nymphs as did these rodents. The ratios of larvae to nymphs suggest that, relative to birds, lizards, and squirrels (infested by 1.3-6.0 larvae per nymph), nocturnally active ground-dwelling rodents such as woodrats and mice are underutilized by the nymphal stage (69 to >100 larvae per nymph). The western gray squirrel and guild I-II birds (e.g., the dark-eyed junco, Junco hyemalis [L.]) were the only potential reservoirs of Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt, and Brenner (the causative agent of Lyme disease in North America) that were frequently infested with both I. pacificus larvae and nymphs and commonly utilized dense woodland habitats.     

Relative importance of lizards and mammals as hosts for ixodid ticks in northern California

Lizards and mammals were trapped and examined for ticks from August 1992 to June 1993 in two habitat types, chaparral and woodland-grass, in northern California. Five tick species were collected from mammals (Dermacentor occidentalis, Haemaphysalis leporispalustris, Ixodes pacificus, I. spinipalpis, I. woodi), but only I. pacificus was found on lizards. Dermacentor occidentalis, I. pacificus, and I. woodi occurred in both habitats, whereas H. leporispalustris and I. spinipalpis were found only on animals trapped in chaparral. The tick species most commonly encountered on mammals was D. occidentalis in chaparral and I. pacificus in woodland-grass. Peak infestation of mammals occurred in spring for I. pacificus immatures and H. leporispalustris, summer for D. occidentalis immatures, fall through spring for I. woodi immatures, and fall through winter for I. spinipalpis. The primary aim of the study was to quantify the relative importance of the western fence lizard (Sceloporus occidentalis), which is reservoir-incompetent for Borrelia burgdorferi sensu lato (s.l.), and mammalian B.burgdorferi s.l.-reservoirs as hosts for the immature stages of I. pacificus in spring. The estimated relative utilization by I. pacificus of the western fence lizard versus mammals was 88% for larvae and 99% for nymphs in chaparral in May. When tick infestation data were corrected for a two-fold lower efficiency of field examinations for rodents than for lizards, the western fence lizard still accounted for 78% of larval and 98% of nymphal feedings. In woodland-grass, 46% of 100 I. pacificus larvae and 100% of 52 nymphs recovered from mammals or western fence lizards during May-June were collected from the lizards. However, this may represent an underestimate of the importance of the western fence lizard as a larval host in this habitat because inclement weather during the late May sampling period doubtless resulted in significantly decreased lizard activity. In conclusion, the western fence lizard was more heavily utilized by I. pacificus immatures, especially nymphs, than were rodents. This revised version was published online in July 2006 with corrections to the Cover Date.     

Prevalence and abundance of Ixodes pacificus immatures (Acari: Ixodidae) infesting western fence lizards (Sceloporus occidentalis) in northern California: temporal trends and environmental correlates.

The prevalence and abundance of immature Ixodes pacificus ticks on western fence lizards (Sceloporus occidentalis) were examined in relation to time of year, host attributes (i.e., age, gender, and presence or absence of blood parasites), and 5 environmental characteristics, including topographic exposure and ground cover substrate, over a 2-year period in northern California. Lizards were infested with subadult ticks from early March until late July or early August, with peak median numbers of larvae and nymphs recorded in late April and early May of both years. Peak larval and nymphal abundances differed between years. The overall ratio of larvae to nymphs on adult male lizards was low, ranging from 0.80 in 1999 to 2.41 in 2000. Such intensive feeding of nymphs versus larvae on these lizards, which are reservoir-incompetent for Borrelia burgdorferi spirochetes, may explain previous observations of decreasing spirochetal infection prevalence from the nymphal to adult stage in northwestern California. Adult male lizards were more likely to be infested with nymphs and harbored greater abundances of larvae and nymphs than adult females. Lizards uninfected with blood parasites had more nymphs than infected lizards. The measured environmental characteristics could explain only a small percentage of the total variation observed in larval prevalence (22%) and in larval and nymphal abundance (12 and 3%, respectively).     

Prevalence of Borrelia burgdorferi sensu lato in ticks collected from migratory birds in Switzerland

The prevalence of ticks infected by Borrelia burgdorferi sensu lato on birds during their migrations was studied in Switzerland. A total of 1,270 birds captured at two sites were examined for tick infestation. Ixodes ricinus was the dominant tick species. Prevalences of tick infestation were 6% and 18.2% for birds migrating northward and southward, respectively. Borrelia valaisiana was the species detected most frequently in ticks, followed by Borrelia garinii and Borrelia lusitaniae. Among birds infested by infected ticks, 23% (6/26) were infested by B. lusitaniae-infected larvae. Migratory birds appear to be reservoir hosts for B. lusitaniae.     

Seasonal patterns of abundance and host relationships of the Australian paralysis tick,Ixodes holocyclus Neumann (Acarina: Ixodidae), in southeastern Queensland

Estimates of seasonal abundance of larvae, nymphs and females of the paralysis tick, Ixodes holocyclus, were obtained by collecting ticks that engorged on small mammals and birds trapped in two localities in southeastern Queensland: Brisbane (wet sclerophyll forest) and Tamborine Mountain (cleared rain forest). The long-tailed short-nosed bandicoot, Isoodon macrourus, was the most common mammal trapped but small numbers of other marsupials, rodents and ground frequenting birds were also captured. Small numbers of five other tick species were also collected. In both habitats there was clearly one dominant generation of I. holocyclus;7er year, although the presence of all stages at most times of the year indicated overlapping of smaller cohorts. Females were most abundant in spring and early summer, larvae in summer-autumn, and nymphs in autumn-winter. I. holocyclus was abundant on I. macrourus and rare on most other mammals and birds captured. At the peak of abundance of each instar, each bandicoot dropped from 500 to 2000 engorged larvae, 100 to 200 engorged nymphs, and four to six engorged females. Life tables were compiled for the tick in both habitats and these indicate that there was relatively high survival from engorged larva to engorged nymph and thence to engorged female and that most mortality occurred between detaching of the engorged female and the detaching of the engorged larva. The tick was more abundant on bandicoots from cleared rain forest and rain forest edge, than on those from sclerophyll woodland. The survival of engorged larvae and nymphs of I. holocyclus and the larval productivity of engorged females were examined in a warm moist climate where the tick was abundant (Tamborine Mountain) and in a hotter dryer climate where the tick was rare or absent (Amberley). In both localities engorged larvae and nymphs survived to the next instar in all seasons of the year. On most occasions engorged females produced eggs which hatched. Mature bandicoots from tick infested areas showed little or no resistance to infestation with larvae or nymphs of I. holocyclus, whereas other small mammals from the same area showed an appreciable degree of resistance to the immature stages of the tick. Feeding larvae and nymphs exposed to normal light-dark cycles in the laboratory detached during the afternoon and early evening. This behaviour and host resistance are discussed in relation to the daily activity cycles of host species, their habitat preferences, and their role as hosts for I. holocyclus.     

Tick infestation on birds in an urban Atlantic Forest fragment in north-eastern Brazil

Birds are important hosts for various tick species, playing a significant role in their biological life cycle and dispersion. In this study, we investigated tick infestations on birds trapped in an urban remnant of Atlantic Forest in Pernambuco state, Brazil. From February 2015 to March 2017, 541 birds belonging to 52 species were trapped with mist nets and examined for ectoparasites. Birds trapped in the late successional forest were significantly more infested than birds trapped in the early successional forest. In the same way, ectoparasite infestation varied significantly according to bird weight and collection plot. Overall, 198 birds (36.6%) belonging to 27 species were parasitized by ectoparasites (i.e., ticks, lice and/or mites). Ectoparasites were effectively collected from 111 birds, of which 99 belonging to 20 species were infested by ticks (n?=?261), namely, Amblyomma longirostre (13 nymphs), Amblyomma nodosum (21 nymphs), Amblyomma varium (one nymph), and Amblyomma spp. (five nymphs and 221 larvae). Most of the ticks (>?90%) were collected from Passeriformes. This study provides the second record of A. varium in Pernambuco state and confirms that birds, especially Passeriformes, are important hosts for larvae and nymphs of Amblyomma spp. in the Atlantic Forest biome of Pernambuco.

     

Bird ticks in an area of the Cerrado of Minas Gerais State, southeast Brazil

In the present study the tick prevalence, mean intensity of infestation and species were recorded on birds captured between January 2009 and December 2010 in the Ecological Station Pirapitinga-ESEC from Minas Gerais State, Brazil. A total of 967 birds, from 15 families and 40 species were captured and 165 (17.1 %) individuals were parasitized by ticks. Of these 160 (97 %) belonged to the order Passeriformes. Five tick species were identified: Amblyomma longirostre (n = 274) was the most common species followed by Amblyomma parvum (n = 43), Amblyomma nodosum (n = 39), Amblyomma ovale (n = 24) and Riphicephalus sanguineus (n = 7). None of 61 unengorged larvae molted to nymph. The mean intensity of infestation was 2.7 ± 2.4 ticks per bird (448 ticks/165 birds) ranging from 1 to 10. Only 19 (11.4 %) birds were infested with one species of tick. The remaining birds were infested by two, three or four species of ticks. Also new hosts for all five ticks were recorded. Only nymphs were recorded throughout the year with two similar peaks during autumn and winter 2009 and 2010.     

Migratory songbirds disperse ticks across Canada, and first isolation of the Lyme disease spirochete, Borrelia burgdorferi, from the avian tick, Ixodes auritulus

During a 3-yr comprehensive study, 196 ixodid ticks (9 species) were collected from 89 passerine birds (32 species) from 25 localities across Canada to determine the distribution of avian-associated tick species and endogenous Lyme disease spirochetes, Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt, and Brenner. We report the following first records of tick parasitism on avian hosts: the rabbit-associated tick, Ixodes dentatus Marx, from Manitoba and Ontario; the mouse tick, Ixodes muris Bishopp and Smith, from British Columbia; and the blacklegged tick, Ixodes scapularis Say, from New Brunswick. Moreover, we provide the first record of the Neotropical tick, Amblyomma humerale Koch (1 nymph), in Canada and its parasitism of any bird. This tick was compared morphologically with nymphs of other Neotropical Amblyomma spp., and genetically, using a 344-bp fragment of the 12S rDNA sequence of 41 New World Amblyomma species. The first collections of the western blacklegged tick, Ixodes pacificus Cooley and Kohls, from passerine species in Alberta and British Columbia, are also reported. Notably, we further report the first isolation of B. burgdorferi from the bird tick, Ixodes auritulus Neumann, collected from an American robin, Turdus migratorius L., on Vancouver Island. Furthermore, B. burgdorferi-positive I. auritulus larvae were collected from a reservoir-competent fox sparrow, Passerella iliaca (Merrem). Our findings indicate that ground-dwelling passerines, in particular, are parasitized by certain ixodid ticks and play an important role across Canada in the wide dispersal of B. burgdorferi-infected ticks and increased risk of Lyme disease exposure.     

The role of medium-sized mammals as reservoirs of Borrelia burgdorferi in southern New York

The ability of raccoons (Procyon lotor), striped skunks (Mephitis mephitis) and opossums (Didelphis virginiana) to serve as reservoirs of Borrelia burgdorferi, the spirochetal agent of Lyme disease, was compared with that of white-footed mice (Peromyscus leucopus). Twenty-eight (28) medium-sized mammals and 34 white-footed mice were captured in Westchester County, New York (USA) in summer 1986. Animals were caged over pans of water for 1 to 2 days to recover engorged tick larvae (Ixodes dammini) that detached from the hosts after feeding. With the exception of mice, numbers of engorged tick larvae recovered exceeded those counted during initial examinations of the hosts by 30% (opossums) to nearly 90% (raccoons). Newly-molted nymphal ticks derived from the engorged larvae were examined for the presence of spirochetes by darkfield microscopy. Percentage infection was 5% (n = 22) for ticks from skunks and 14% (n = 191) for ticks from raccoons. None of 24 nymphs from larvae that fed on opossums survived long enough for spirochete examination. By comparison, 40% (n = 72) of nymphs from larvae which fed on white-footed mice were infected. Of the individual hosts from which molted nymphs had fed as larvae, 67% of mice, 33% of skunks, and 55% of raccoons produced spirochete-positive ticks.     

Borrelia Burgdorferi Sensu Lato in Ixodes Ricinus Ticks and Rodents in a Recreational Park in South-Western Ireland

Ixodes ricinus ticks infected with Borrelia burgdorferi sensu lato were numerous on the edges of paths and roads in a recreational park in south-western Ireland. The abundance of ticks at different sites was related to the presence of deer, but a negative relationship was shown between tick abundance and tick infection rates. This is thought to be due to the deposition of large numbers of uninfected ticks by deer, which are apparently not good reservoir hosts of B. burgdorferi s.l. Blood meal analysis only detected deer DNA in uninfected nymphs. Reservoir competent rodents, Apodemus sylvaticus and Clethrionomys glareolus, were abundant at all sites and a high proportion of captured specimens were infested with larval ticks. However, very few rodents were infected with B. burgdorferi s.l. and none of the unfed infected nymphs analysed for the identity of their larval blood meal had fed on rodents. The spirochaetes detected in I. ricinus in the study area may be poorly adapted to rodents or are not transmitted readily because of the absence of nymphal infestation. The majority of spirochaetes in these ticks were apparently acquired from non-rodent hosts, such as birds.     

Role of small mammals in the ecology of Borrelia burgdorferi in a peri-urban park in north coastal California

The role of small mammals other than woodrats in the enzootiology of the Lyme disease spirochete, Borrelia burgorferi, was assessed in a peri-urban park. Mammals were collected monthly from September through to April. Following tick removal, the animals were tested for B. burgdorferi by culture of ear-punch biopsies. Larvae and nymphs that were intermediate in morphology between Ixodes spinipalpis and Ixodes neotomae occurred on several species of rodents (Peromyscus truei, Peromyscus californicus, Microtus californicus, Rattus rattus and Reithrodontomys megalotis) and the brush rabbit (Sylvilagus bachmani). Morphometric analyses of these I. spinipalpis-like ticks and the offspring from two I. neotomae females from this site suggest that I. neotomae may be conspecific with I. spinipalpis. Borrelia burgdorferi was isolated from eight out of 109 (7.3%), three out of 16 (18.8%), two out of 38 (5.3%) and two out of six (33.3%) P. truei, P. maniculatus, M. californicus and R. rattus, respectively. One brush rabbit yielded the first isolate of B. burgdorferi from a lagomorph in western North America. This isolate and three others derived from unfed I. spinipalpis-like nymphs failed to produce infection when inoculated intradermally into 11–12 P. maniculatus each. Likewise, no spirochetes were detected in 420 Ixodes pacificus nymphs derived from larvae fed on animals inoculated with these isolates. An additional isolate, derived from an I. spinipalpis-like nymph, was recovered by ear-punch biopsies from five out of 12 (42%) needle-inoculated P. maniculatus. However, spirochetes were not detected in 20 I. pacificus nymphs fed as larvae on each of five mice (two infected and three uninfected) inoculated with this isolate. We conclude that brush rabbits and several species of rodents besides woodrats may contribute to the maintenance of B. burgdorferi because they harbour the spirochete and are fed upon by competent enzootic and bridge vectors.     

Widespread dispersal of Borrelia burgdorferi-infected ticks collected from songbirds across Canada

Millions of Lyme disease vector ticks are dispersed annually by songbirds across Canada, but often overlooked as the source of infection. For clarity on vector distribution, we sampled 481 ticks (12 species and 3 undetermined ticks) from 211 songbirds (42 species/subspecies) nationwide. Using PCR, 52 (29.5%) of 176 Ixodes ticks tested were positive for the Lyme disease spirochete, Borrelia burgdorferi s.l. Immature blacklegged ticks, Ixodes scapularis , collected from infested songbirds had a B. burgdorferi infection prevalence of 36% (larvae, 48%; nymphs, 31%). Notably, Ixodes affinis is reported in Canada for the first time and, similarly, Ixodes auritulus for the initial time in the Yukon. Firsts for bird-parasitizing ticks include I. scapularis in Quebec and Saskatchewan. We provide the first records of 3 tick species cofeeding on passerines (song sparrow, Swainson's thrush). New host records reveal I. scapularis on the blackpoll warbler and Nashville warbler. We furnish the following first Canadian reports of B. burgdorferi-positive ticks: I. scapularis on chipping sparrow, house wren, indigo bunting; I. auritulus on Bewick's wren; and I. spinipalpis on a Bewick's wren and song sparrow. First records of B. burgdorferi-infected ticks on songbirds include the following: the rabbit-associated tick, Ixodes dentatus, in western Canada; I. scapularis in Quebec, Saskatchewan, northern New Brunswick, northern Ontario; and Ixodes spinipalpis (collected in British Columbia). The presence of B. burgdorferi in Ixodes larvae suggests reservoir competency in 9 passerines (Bewick's wren, common yellowthroat, dark-eyed junco, Oregon junco, red-winged blackbird, song sparrow, Swainson's thrush, swamp sparrow, and white-throated sparrow). We report transstadial transmission (larva to nymph) of B. burgdorferi in I. auritulus. Data suggest a possible 4-tick, i.e., I. angustus, I. auritulus, I. pacificus, and I. spinipalpis, enzootic cycle of B. burgdorferi on Vancouver Island, British Columbia. Our results suggest that songbirds infested with B. burgdorferi-infected ticks have the potential to start new tick populations endemic for Lyme disease. Because songbirds disperse B. burgdorferi-infected ticks outside their anticipated range, health-care providers are advised that people can contract Lyme disease locally without any history of travel.     

Variations in Ixodes ricinus Density and Borrelia Infections Associated with Cattle Introduced into a Woodland in The Netherlands

The effect of introduced large herbivores on the abundance of Ixodes ricinus ticks and their Borrelia infections was studied in a natural woodland in The Netherlands. Oak and pine plots, either ungrazed or grazed by cattle, were selected. Ticks were collected weekly by blanket dragging. Borrelia infections were determined by PCR and restriction fragment length polymorphism. Rodent densities were estimated using mark-release-recapture methods. On occasion, the cattle were inspected for tick infestations. Meteorological data were recorded for each habitat. Significantly more ticks were collected in the ungrazed woodland than in the grazed woodland. The ungrazed oak habitat had higher tick densities than the pine habitat, while in the grazed habitats, tick densities were similar. Borrelia infection rates ranged from zero in larvae to 26% in nymphs to 33% in adult ticks, and B. afzelii, B. burgdorferi sensu stricto, B. garinii, and B. valaisiana were the species involved. Coinfections were found in five ticks. There was no effect of the presence of cattle on Borrelia infections in the ticks. In the ungrazed area, Borrelia infections in nymphs were significantly higher in the oak habitat than in the pine habitat. More mice were captured in the ungrazed area, and these had a significantly higher tick burden than mice from the grazed area. Tick burden on cattle was low. The results suggest that grazing has a negative effect on small rodents as well as on ticks but not on Borrelia infections. Implications of these results for management of woodland reserves and risk of Lyme disease are discussed.     

Risk factors in habitats of the tick Ixodes ricinus influencing human exposure to Ehrlichia phagocytophila bacteria

Ixodes ricinus L. (Acari: Ixodida) were sampled during 1996-99 in southern Scotland, on vegetation using cloth drags, on humans by removal from clothing and on roe deer (Capreolus capreolus L.) by searching legs of culled deer. Developmental microclimate was recorded by automatic recorders and questing microclimate by portable instruments during tick collections. Ticks and deer were examined for infection with Ehrlichia phagocytophila bacteria (Rickettsiales) using microscopy and polymerase chain reaction. This pathogen causes tick-borne fever of sheep in Europe and human granulocytic ehrlichiosis in North America, but in Europe human clinical ehrlichiosis due to E. phagocytophila has not been recorded despite serological evidence of exposure. Among three types of habitat, coniferous woodland was most infested with questing ticks (560 ticks/km of drag; mean numbers collected on long trousers: 24.3 larvae, 13.5 nymphs and 0.8 adult ticks/km walked), deciduous woodland had slightly lower infestation (426 ticks/km drag) and upland sheep pasture had much lower infestation (220 ticks/km drag). Of the three main vegetation types, bracken was least infested (360 ticks/km drag), ericas most (430 ticks/km drag) and grassland had intermediate infestation density (413 ticks/km drag). Questing and developmental microclimates were poor predictors of exposure within these habitats, except lower infestation of pastures was attributed to greater illumination there. Collectors who walked a total of 300 km through all habitats (taking 360 h in all seasons), wearing cotton trousers hanging outside rubber boots, were bitten by only four nymphs and 11 larvae of I. ricinus (but no adult ticks). There was a negative correlation between densities of deer and ticks collected, although presence of deer remains a major indicator of exposure. The proportion of infected ticks was fairly uniform at four sites studied. Overall prevalence of E. phagocytophila in I. ricinus was 3.3% in nymphs (40/1203) but only approximately 1.5% in adults of both sexes (although males do not bite). It was estimated that nymphs of I. ricinus gave 4.4% probability of one infected bite/person/year (for occupational exposure during this research) due to presence in all seasons and habitats, their human biting rate of 0.011 nymphs/h or 0.013 nymphs/km and widespread infection with E. phagocytophila. The frequency distribution of intensity of infection in ticks was approximately normal (mean 98 morulae/nymph infected), thus there is a high risk of receiving a high dose from any one infected tick bite.