Flagging versus dragging as sampling methods for nymphal Ixodes scapularis (Acari: Ixodidae)

The nymphal stage of the blacklegged tick, Ixodes scapularis (Acari: Ixodidae), is responsible for most transmission of Borrelia burgdorferi, the etiologic agent of Lyme disease, to humans in North America. From 2010 to fall of 2012, we compared two commonly used techniques, flagging and dragging, as sampling methods for nymphal I. scapularis at three sites, each with multiple sampling arrays (grids), in the eastern and central United States. Flagging and dragging collected comparable numbers of nymphs, with no consistent differences between methods. Dragging collected more nymphs than flagging in some samples, but these differences were not consistent among sites or sampling years. The ratio of nymphs collected by flagging vs dragging was not significantly related to shrub density, so habitat type did not have a strong effect on the relative efficacy of these methods. Therefore, although dragging collected more ticks in a few cases, the numbers collected by each method were so variable that neither technique had a clear advantage for sampling nymphal I. scapularis.     

Questing behaviour of Ixodes ricinus ticks (Acari: Ixodidae)

The vertical distribution in the vegetation of questing Ixodes ricinus ticks was investigated in two different vegetation types (high and low vegetation) at two localities in south-central Sweden during 1992-1993 (Toro) and 1995 (Bogesund). Significant correlations were found between the vertical distribution of immature ticks and the height of the vegetation. The greatest mean availabilities of the larvae and nymphs in low vegetation were in the intervals 0-9 and 30-39 cm, respectively. The larval numbers were greatest close to the ground (0-29 cm) in both high and low vegetation. The larval : nymphal ratio, at ground level at localities free of ground vegetation, varied between 8 : 1 and 32 : 1. In high vegetation, the greatest mean numbers of nymphal and adult ticks were at height intervals of 50-59 and 60-79 cm, respectively. These ranges are within the estimated height interval (40-100 cm) of the main part of the body surface of their preferred host, the roe deer (Capreolus capreolus). The presence of most questing I. ricinus larvae at ground level would favour the transmission of Borrelia burgdorferi s.l., since this is where the highly reservoir-competent rodents and shrews usually occur.     

Sexual transmission of Borrelia garinii by male Ixodes persulcatus ticks (Acari, Ixodidae)

We investigated the transmission of Borrelia garinii and Borrelia afzelii between male and female Ixodes persulcatus ticks. For this purpose the infection rate of partners from tick couples was determined by polymerase chain reaction and reverse line blot. In couples, where the male tick was infected with B. garinii, four out of nine female partners carried B. garinii. In eight couples, male ticks had a dual infection of B. afzelii and B. garinii and three female partners were infected by Borrelia spirochetes. Two female ticks carried B. garinii, and one female tick had a dual infection. No evidence for transmission of B. afzelii from male to female ticks was found among seven couples. In 45 couples where the female tick was infected, not one male tick carried spirochetes. The difference in the B. garinii infection rate between male and female ticks among these couples is highly significant. Our data suggest that transmission of B. garinii from male ticks to female ticks does occur. Sexual transmission of this pathogen may play an important role in the maintenance of B. garinii in I. persulcatus. © Rapid Science Ltd. 1998     

Vector competence of Ixodes angustus (Acari: Ixodidae) for Borrelia burgdorferi sensu stricto

The vector competence of Ixodes angustus for Borrelia burgdorferi sensu stricto (s.s.) was investigated in the laboratory. The larval progeny of female ticks from Washington State were placed on Swiss-Webster mice that had been inoculated intravenously with 10⁸ spirochetes each of a Californian isolate of B. burgdorferi. Spirochetes were detected in 6 (12%) of 50 nymphs derived from larvae that had fed on these animals. Ten nymphs from the same cohort of experimentally infected ticks were placed on each of 4 naive deer mice (Peromyscus maniculatus). One of the mice seroconverted to B. burgdorferi and spirochetes were isolated from its ear tissues 4 weeks after exposure to ticks. Further vector competence trials were conducted with I. angustus ticks from California. Larvae were fed on deer mice that had been inoculated intradermally with B. burgdorferi along with larvae of I. spinipalpis as a comparison group. There was no significant difference in the prevalence of infection in nymphs of I. angustus (8.2%) versus those of I. spinipalpis (12.1%). We conclude that I. angustus is a competent experimental vector of B. burgdorferi s.s. and its efficiency for acquiring and transstadially passing such spirochetes is similar to that of I. spinipalpis.     

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 ≈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.     

Detection of Lyme disease spirochete,Borrelia burgdorferi sensu lato,including three novel genotypes in ticks (Acari: Ixodidae) collected from songbirds (Passeriformes) across Canada

Lyme disease is reported across Canada, but pinpointing the source of infection has been problematic. In this three‐year, bird‐tick‐pathogen study (2004–2006), 366 ticks representing 12 species were collected from 151 songbirds (31 passerine species/subspecies) at 16 locations Canada‐wide. Of the 167 ticks/pools tested, 19 (11.4%) were infected with Borrelia burgdorferi sensu lato (s.l.). Sequencing of the rrf‐rrl intergenic spacer gene revealed four Borrelia genotypes: B. burgdorferi sensu stricto (s.s.) and three novel genotypes (BC genotype 1, BC genotype 2, BC genotype 3). All four genotypes were detected in spirochete‐infected Ixodes auritulus (females, nymphs, larvae) suggesting this tick species is a vector for B. burgdorferi s.l. We provide first‐time records for: ticks in the Yukon (north of 60° latitude), northernmost collection of Amblyomma americanum in North America, and Amblyomma imitator in Canada. First reports of bird‐derived ticks infected with B. burgdorferi s.l. include: live culture of spirochetes from Ixodes pacificus (nymph) plus detection in I. auritulus nymphs, Ixodes scapularis in New Brunswick, and an I. scapularis larva in Canada. We provide the first account of B. burgdorferi s. l. in an Ixodes muris tick collected from a songbird anywhere. Congruent with previous data for the American Robin, we suggest that the Common Yellowthroat, Golden‐crowned Sparrow, Song Sparrow, and Swainson's Thrush are reservoir‐competent hosts. Song Sparrows, the predominant hosts, were parasitized by I. auritulus harboring all four Borrelia genotypes. Our results show that songbirds import B. burgdorferi s.l.‐infected ticks into Canada. Bird‐feeding I. scapularis subadults were infected with Lyme spirochetes during both spring and fall migration in eastern Canada. Because songbirds disperse millions of infected ticks across Canada, people and domestic animals contract Lyme disease outside of the known and expected range.     

Optimal sampling and spatial distribution of Ixodes pacificus, Dermacentor occidentalis and Dermacentor variabilis ticks (Acari: Ixodidae)

A common method for sampling tick populations is flagging, which is a method of dragging a white cloth over a plant substrate for a fixed distance along a transect. Flagging over rough physical surfaces or using long subtransect lengths could lead to the underestimation of tick densities. Using estimates of the drop-off rates of adult Ixodes pacificus in flag sampling, optimal sampling schemes (the length and number of subtransects) were examined using the relationships between the tick drop-off rates (c), the tick density and distribution patterns and the roughness of the sampled plant substrate. It was found that the optimal number of subtransects and Lmax, the longest subtransect length which did not significantly underestimate the tick density from c, were affected by the tick density, substrate roughness and tick distribution pattern. This study also showed that the density and distribution of I. pacificus and Dermacentor occidentalis varied greatly over time in the populations sampled, while the Dermacentor variabilis densities were low and showed no significant changes over time. Both I. pacificus and D. occidentalis had clumped distributions along trails and these clumps were aggregated. However, the clump sizes (or individual clump areas) changed significantly over time because of density fluctuation or the movement of ticks. Finally, a positive association between the number of I. pacificus and D. occidentalis adults was observed from 2 m subtransect flag collections in March 1995; no relationship was found in 1994. © Rapid Science Ltd. 1998     

Climate change and the potential for range expansion of the Lyme disease vector Ixodes scapularis in Canada

We used an Ixodes scapularis population model to investigate potential northward spread of the tick associated with climate change. Annual degree-days >0 degrees C limits for I. scapularis establishment, obtained from tick population model simulations, were mapped using temperatures projected for the 2020s, 2050s and 2080s by two Global Climate Models (the Canadian CGCM2 and the UK HadCM3) for two greenhouse gas emission scenario enforcings 'A2'and 'B2' of the Intergovernmental Panel on Climate Change. Under scenario 'A2' using either climate model, the theoretical range for I. scapularis establishment moved northwards by approximately 200 km by the 2020s and 1000 km by the 2080s. Reductions in emissions (scenario 'B2') had little effect on projected range expansion up to the 2050s, but the range expansion projected to occur between the 2050s and 2080s was less than that under scenario 'A2'. When the tick population model was driven by projected annual temperature cycles (obtained using CGCM2 under scenario 'A2'), tick abundance almost doubled by the 2020s at the current northern limit of I. scapularis, suggesting that the threshold numbers of immigrating ticks needed to establish new populations will fall during the coming decades. The projected degrees of theoretical range expansion and increased tick survival by the 2020s, suggest that actual range expansion of I. scapularis may be detectable within the next two decades. Seasonal tick activity under climate change scenarios was consistent with maintenance of endemic cycles of the Lyme disease agent in newly established tick populations. The geographic range of I. scapularis-borne zoonoses may, therefore, expand significantly northwards as a consequence of climate change this century.     

Infestation of mammals by Ixodes ricinus ticks (Acari: Ixodidae) in south-central Sweden

Infestation by Ixodes ricinus ticks on rodents, hares and cervids was examined at Bogesund, 10 km north of Stockholm, in south-central Sweden during 1991-1994 and on varying hares (Lepus timidus) at Stora Karlso and Gotska Sandon in the Baltic Sea during 1992-1993. At Bogesund, there were great differences between two consecutive years in the number of I. ricinus larvae infesting bank voles (Clethrionomys glareolus). The seasonal pattern of infestation by I. ricinus larvae and nymphs on bank voles was unimodal in 1991, with peaks in June-July and bimodal in 1992, with peaks in June and August. Male bank voles, compared to females and older voles, compared to young voles, harboured greater numbers of I. ricinus ticks. Apodemus mice, compared to bank voles, harboured greater numbers of I. ricinus ticks. Ixodes ricinus larvae engorged on Apodemus mice were heavier than larvae engorged on bank voles and resulted in larger nymphs. However, there was no difference in the proportions of viable nymphs resulting from larvae engorged on mice or voles. The ranges in the numbers of I. ricinus ticks infesting individual hosts were 1-451 for rodents, 16-2374 for hares and 428-2072 for roe deer (Capreolus capreolus). These ranges of tick numbers are estimated to represent potential blood losses from individual hosts of approximately 0.2-65% for rodents, 0.2-13% for hares and 0.3-9.0% for roe deer. Within the populations of all host species examined, the distributions of all stages of I. ricinus were clumped, with most host individuals harbouring few ticks and only a few individuals harbouring many ticks. The data suggest that, even though a small proportion of tick hosts may be severely affected, the direct effects of feeding by I. ricinus are unlikely to play an important role on mammal population dynamics.     

Development of the salivary glands in embryos of Ixodes ricinus (Acari: Ixodidae)

We examined Ixodes ricinus embryos between 18 and 28 days of development with light, scanning and transmission electron microscopy. The differences in inner structure attested to establish three successive developmental stages: days 18–20, day 23, and days 26–28. Between 18 and 20 days the embryos are at early stages of organogenesis. Salivary glands cannot be identified at that stage. In 23-day-old embryos salivary glands are already outlined but the structure of alveoles is still different from that in larvae in which the embryonic development has been completed. Gland cells start to form alveoles and become active between 26 and 28 days of the development. This revised version was published online in July 2006 with corrections to the Cover Date.     

Regeneration of Haller's sensory organ in the tick, Ixodes rubicundus (Acari: Ixodidae)

A study of regeneration in nymphs and adults of the South African tick Ixodes (Afrixodes) rubicundus, from which the forelegs had been amputated during the previous instar, revealed that the structural changes in regenerated Haller's sensory organs resemble those observed in other ixodid ticks, in particular in another prostriate tick, Ixodes (Ixodes) ricinus. The adult regenerates re-establish their atavistic features in terms of the increased number of different sensilla on the distal knoll, in the anterior pit and the capsule. The nymphal regenerates, in contrast, re-establish the features of the previous instar through a reduction in the number of some sensilla on the distal knoll and in the post-capsular area. The structural changes in different compartments of the organ appear independent. The phenomenon of regenerative induction through the appearance of specific changes in Haller's organ of the contralateral non-treated foreleg is probably characteristic only of prostriate ticks. A unique modification in the regenerated Haller's organ as revealed by duplication of the Haller's organ capsule was discovered in both I. rubicundus nymphs and adult ticks.     

Borrelia burgdorferi sensu lato in the female cement plug of Ixodes persulcatus ticks (Acari,Ixodidae)

Borrelia burgdorferi sensu lato was detected in one out of five cement plugs of female Ixodes persulcatus ticks. The spirochetes were found by dark field microscopy as early as 18 h after attachment of the ticks to the skin of a white mouse. The relevance of this finding is discussed in relation to the epidemiology of Lyme Borreliosis.     

Geographical distribution and density of Ixodes dammini (Acari: Ixodidae) and relationship to Lyme disease transmission in New Jersey

As part of continuing studies of Lyme disease, deer were surveyed during three hunting seasons in 1981 to obtain information on geographic distribution and density of I. dammini in New Jersey. I. dammini occurred throughout central and southern New Jersey. Four deer management zones (DMZs) were shown to have high tick densities. Geographical distribution and density data were independently regressed against 25 environmental and physical factors. Elevation was shown to be the most important factor in explaining the variability in both I. dammini distribution and density. Lyme disease cases were closely associated with the distribution of I. dammini and 57.3 percent of 117 Lyme disease cases occurred in the four DMZs previously identified as having the highest tick density.     

Geographic occurrence of Ixodes scapularis and Amblyomma americanum (Acari: Ixodidae) infesting white-tailed deer in North Carolina

A state-wide survey to determine the occurrence and comparative numbers of ticks infecting white-tailed deer (Odocoileus virginianus) was conducted in North Carolina (USA). One thousand six hundred twenty nine deer were examined in 60 of 100 counties; with the exception of one county in the piedmont region, all tick-infested deer occurred in the coastal plain. Ixodes scapularis (46%) and Amblyomma americanum (53%) were the most prevalent species encountered and accounted for more than 98% of the 4,286 ticks collected. Some specimens of Dermacentor albipictus and Amblyomma maculatum also were collected.     

Differences in Borrelia infections in adult Ixodes persulcatus and Ixodes ricinus ticks (Acari: Ixodidae) in populations of north-western Russia

Dark-field microscopy was used to determine the number of Borrelia spirochetes in 630 standard preparations obtained from adult ixodid ticks (344 Ixodes persulcatus and 286 I. ricinus) collected in 1989-92 in the Leningrad region of Russia. The average numbers of Borrelia in I. persulcatus and I. ricinus preparations were 34.7 and 23.3 per 100 microscopic fields, respectively. The maximal individual values registered each year for ticks of both species were several hundred times greater than the minimal values. Ticks carrying relatively small numbers of Borrelia generally predominated. Proportions of more heavily infected ticks varied considerably from year to year. These parameters were significantly higher in foci with predominance of I. persulcatus ticks. As a consequence, risk to acquire Lyme borreliosis in such foci is considered greater than in foci where I. ricinus predominates.     

Hosts and Pathogen Detection for Immature Stages of Ixodes ricinus (Acari: Ixodidae) in North-Central Spain

To determine hosts of the immature stages of a southern population of Ixodes ricinus, we trapped rodents and birds in an area of north-central Spain in May–June and August–September of 1998 and 1999. The most frequently trapped rodents were Apodemus sylvaticus (230 specimens) and Clethrionomys glareolus (99), with a larval infestation prevalence of 49% and 81%, respectively (in spring) and 21% and 41% (in summer). C. glareolus was always more heavily parasitized by larvae (mean numbers 19.8 in spring, 3.4 in summer) than A. sylvaticus (6.1 and 1.2, respectively). No nymphs were collected from the rodents. The larval distribution pattern differed considerably between rodent host species but not according to the season of the year. The most commonly trapped birds were members of the Turdidae, which had infestation incidences of about 70% (larvae) and 10% (nymphs) in spring and 20–15% in summer. We also used molecular methods for the identification of reservoir hosts of the larval ticks and of the pathogens they contained, from nymphs collected in the same site in June of 2002 and 2003. The study showed that most of the ticks had fed on birds and few on rodents and ruminants. Pathogens detected in these ticks included Babesia microti, Borrelia garinii, B. valaisiana and B. afzelii. Borrelia afzelii was detected in a tick that had apparently fed on a wild boar.