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.     

Bodies of liquid water as a source of water gain for Ixodes ricinus ticks (Acari: Ixodidae)

It is well established that free-living unfed ticks can compensate for their inevitable body water losses by active water vapour absorption in periods of ambient relative humidity (RH) of greater than 75-90%. Whilst many species of terrestrial arthropods are known to be capable of locating liquid water and drinking when in need, the existing knowledge concerning the ecological significance of bulk water for rehydration in ticks is scarce. In the present laboratory study batches of unfed Ixodes ricinus (larvae, nymphs, and adults) in varying states of (de)hydration were supplied with tapwater either as droplets for 45 min in a Petri dish or in a small trough for 24 h. The body mass of each tick was gravimetrically determined before and after exposure. Though fully hydrated ticks were not usually attracted to liquid water, the response became increasingly positive with a growing body water deficit (p > 0.05). No tick was ever found with its mouthparts inserted into the water, nor had any tick residues of orange G in its alimentary tract when supplied with orange G-coloured water. Linear regression analysis demonstrated that water intake took place in a regulated manner in adult ticks, i.e. the more severely the ticks had been dehydrated the more net water they subsequently gained (p > 0.01). The maximum uptake rates were <20% of the original body mass per day. These findings indicate that unfed I. ricinus do not drink liquid water but are well able to use the high RH in the immediate vicinity of bulk water to actively take up vapour.     

Genetic diversity of Ixodes ricinus (Ixodida: Ixodidae) ticks in sympatric and allopatric zones in Baltic countries

Ixodes ricinus (Linnaeus 1758) and Ixodes persulcatus (Schulze 1930) ticks are involved in the transmission of a wide variety of pathogens with considerable impact on human and animal health. The co‐distribution zone of these two tick species is situated in the Baltic countries, which provides a special setting for the population studies. In the present study, genetic variability of I. ricinus ticks collected in allopatric and sympatric locations in the Baltic countries has been investigated using a sequence analysis of the mitochondrial DNA control region, 16S rRNA and cytb genes. There were 32 haplotypes (Hd: 0.8551) and 27 haplotypes (Hd:0.8213) of control region sequences from ticks in allopatric and sympatric zones detected, respectively. Out of 47 16S rRNA gene haplotypes, 32 haplotypes (Hd: 0.7213) were found in the allopatric zone and 27 (Hd:0.9572) in the sympatric zone. The Cytb gene was very conserved and monomorphic in ticks from the allopatric zone, whereas three unique haplotypes were observed in the sympatric zone. The higher number of unique haplotypes of the control region was detected in the allopatric zone. Median joining network and Fst analysis did not reveal a clear separation between ticks from the two zones.     

Risk indicators for the tick Ixodes ricinus and Borrelia burgdorferi sensu lato in Sweden

The distributional area of the tick Ixodes ricinus (L.), the primary European vector to humans of Lyme borreliosis spirochaetes (Borrelia burgdorferi sensu lato) and tick‐borne encephalitis virus, appears to be increasing in Sweden. It is therefore important to determine which environmental factors are most useful to assess risk of human exposure to this tick and its associated pathogens. The geographical distribution of I. ricinus in Sweden was analysed with respect to vegetation zones and climate. The northern limit of I. ricinus and B. burgdorferi s.l. in Sweden corresponds roughly to the northern limit of the southern boreal vegetation zone, and is characterized climatically by snow cover for a mean duration of 150 days and a vegetation period averaging 170 days. The zoogeographical distribution of I. ricinus in Sweden can be classified as southerly–central, with the centre of the distribution south of the Limes Norrlandicus. Ixodes ricinus nymphs from 13 localities in different parts of Sweden were examined for the presence of B. burgdorferi s.l. and found to be infected with Borrelia afzelii and Borrelia garinii. Tick sampling localities were characterized on the basis of the density of Borrelia‐infected I. ricinus nymphs, presence of specific mammals, dominant vegetation and climate. Densities of I. ricinus nymphs and Borrelia‐infected nymphs were significantly correlated, and nymphal density can thus serve as a general indicator of risk for exposure to Lyme borreliosis spirochaetes. Analysis of data from this and other studies suggests that high densities of Borrelia‐infected nymphs typically occur in coastal, broadleaf vegetation and in mixed deciduous/spruce vegetation in southern Sweden. Ixodes ricinus populations consistently infected with B. burgdorferi s.l. can occur in: (a) biotopes with shrews, rodents, hares and birds; (b) biotopes with shrews, rodents, hares, deer and birds, and (c) island locations where the varying hare (Lepus timidus) is the only mammalian tick host.     

Ticks in the ecotone: the impact of agri-environment field margins on the presence and intensity of Ixodes ricinus ticks (Acari: Ixodidae) in farmland in southern England

The present study aimed to assess whether agri-environment field margins provide a habitat for the sheep/deer tick Ixodes ricinus. Field studies were conducted in arable farmland in southern England in both extant and newly constructed field margins. The presence and intensity (i.e. the mean number of nymphs per transect, excluding zeros) of questing nymphs and adult I. ricinus were compared between field margins with three adjacent habitats: woodland, hedgerow and arable land. The presence and intensity of ticks within a field margin was also compared between three ecozones: the ecotone, the margin and the crop. It was found that field margins do support I. ricinus, although the intensity of ticks was associated with field margins with adjacent woodland, with a higher tick intensity along the ecotonal ecozone, compared with the rest of the margin or the crop edge. The presence of a hedge also increased the likelihood of finding questing nymphs in a field margin compared with a margin adjacent to arable land. This effect, however, was less pronounced than in field margins with adjacent woodland. The provision of footpaths within the margin (at least 1–2 m from the ecotone), or on the edge of the crop where paths run next to woodland known to be an important tick habitat, could be promoted to minimize tick exposure. In addition, based on the results of the present study, raising awareness that walking alongside woodlands also constitutes a tick risk could be promoted.     

Geostatistics as Predictive Tools to Estimate Ixodes ricinus (Acari: Ixodidae) Habitat Suitability in the Western Palearctic From AVHRR Satellite Imagery

Geostatistics (cokriging) were used to model the cross-correlated information between satellite-derived vegetation and climate variables and actual records of the tick Ixodes ricinus in the western Palearctic. The output was used to map the habitat suitability for I. ricinus on a continental scale. A database of collecting localities of I. ricinus was built up from a total of 812 records. This database has been cross-tabulated with satellite NOAA AVHRR pictures obtained from 1982 to 1994 over the Palearctic at 10 day intervals, with a resolution of 8 km per pixel. A cokriging system was generated to exploit satellite-derived data and to estimate the distribution of I. ricinus. Three vegetation (standard NDVI values) and four temperature variables output the habitat suitability prediction with a sensitivity of 0.98 and a specificity of 0.92. Results obtained with the model closely agreed with actual records of the tick, with 4 and 3% of false-positive and false-negative sites, respectively. Such statistical analysis can guide field work towards the correct interpretation of the distributional limits of ticks and may also be used to make predictions about the impact of global change on tick distributional ranges.     

Densities of Ixodes ricinus ticks (Acari: Ixodidae) on moorland vegetation communities in the UK

Unfed (questing) Ixodes ricinus ticks were collected by blanket dragging on a monthly basis from heather-dominated, Vaccinium-dominated and bracken-dominated vegetation communities from two different biogeographical regions of the UK (the Quantock Hills in Somerset, south west England and the North York Moors, north east England) throughout the spring and summer months of 1991 and 1992. Eighteen sites were monitored across the two regions and a total of 1920 blanket drags were carried out. Vaccinium sites showed high tick densities at all life stages, as did bracken sites. Significantly lower numbers of larval and nymphal ticks per drag were collected on heather sites than were collected on either Vaccinium (bilberry/whortleberry) or bracken sites, while similar numbers of adult ticks per drag were collected from each of the three vegetation communities. There was no significant difference between the mean numbers of any tick life stage collected on the Quantock Hills and those collected on the North York Moors on these vegetation communities or between the mean numbers of any tick life stage collected in 1991 and those collected in 1992 on these vegetation communities.     

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.     

Borreliae in Ixodes ricinus ticks feeding on humans

A total of 298 Ixodes ricinus (L.) ticks (Acari: Ixodidae) feeding on humans in the Czech Republic were tested for borreliae (Borrelia burgdorferi sensu lato) by darkfield microscopy between 1997 and 2003. A majority (68%) of the supplied I. ricinus ticks were nymphs, 25% were females and 7% were larvae. Overall, 20% of 74 examined females and 9% of 203 examined nymphs (but none of 21 examined larvae) were infected with borreliae. The proportion of ticks with a high infection load (>100 spirochetes) was 4% in females and 2% in nymphal I. ricinus. During the year, the highest numbers and proportions of infected nymphal and female ticks were taken from humans in June. Detection of borreliae in the ticks feeding on humans might be helpful in the prophylaxis of Lyme borreliosis.     

Interstadial variation in the attachment sites of Ixodes ricinus ticks on sheep

The spatial aggregation of ticks feeding on vertebrate hosts has been recognized for some time but, for hosts supporting more than one stage of the tick, observations of interstadial variation in the site of attachment have not previously been quantified. This study showed that all three parasitic stages of Ixodes ricinus ticks feeding on sheep attach most commonly to the hair-covered areas of the head and limbs while few ticks attach to the fleeced region of the body. However, significant differences were observed in the site of attachment of the three feeding stages of the tick. Larvae attached to distal limbs and rostral areas of the head and adult females attached to the proximal areas of the limbs and around the neck and ears, while nymphs attached in locations between the larvae and adults. The importance of the spatial aggregation of the ticks and interstadial variation in their distribution on the host, for the transmission of tick-borne pathogens and the epidemiology of the diseases they cause, is discussed. © Rapid Science Ltd. 1998     

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.     

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     

Seasonal incidence of Ixodes ricinus ticks (Acari: ixodidae) on rodents in western France

Data collected from a longitudinal survey carried out over 2 years on four farms in western France were used to assess the incidence and infestation of Ixodes ricinus on rodents. Once a month, on each farm, 25 Sherman live traps were set in hedges bordering selected pastures. A total of 799 micromammals were examined, including Apodemus sylvaticus, Clethrionomys glareolus, Microtus agrestis, Microtus arvalis, and Crocidura spp. Larvae and nymphs of I. ricinus were found. Small numbers of Ixodes (Exopalpiger) trianguliceps were also recovered from each farm. The mean infestation rate of the I. ricinus larvae (1.6–5.9) among all animals examined varied between farms Most animals were infested by only a single tick, but one M. agrestis harboured 43 I. ricinus larvae. Larvae or nymphs were found throughout the year, with peaks from March to October.     

Infestation of urban populations of the Northern white‐breasted hedgehog,Erinaceus roumanicus,by Ixodes spp. ticks in Poland

Infestation by the nest‐dwelling Ixodes hexagonus Leach and the exophilic Ixodes ricinus (Linnaeus) (Ixodida: Ixodidae) on the Northern white‐breasted hedgehog, Erinaceus roumanicus (Erinaceomorpha: Erinaceidae), was investigated during a 4‐year study in residential areas of the city of Poznań, west‐central Poland. Of 341 hedgehogs, 303 (88.9%) hosted 10 061 Ixodes spp. ticks encompassing all parasitic life stages (larvae, nymphs, females). Ixodes hexagonus accounted for 73% and I. ricinus for 27% of the collected ticks. Male hedgehogs carried significantly higher tick burdens than females. Analyses of seasonal prevalence and abundance of I. hexagonus revealed relatively stable levels of infestation of all parasitic stages, with a modest summer peak in tick abundance noted only on male hosts. By contrast, I. ricinus females and nymphs peaked in spring and declined steadily thereafter in summer and autumn, whereas the less abundant larvae peaked in summer. This is the first longterm study to evaluate the seasonal dynamics of both tick species on populations of wild hedgehogs inhabiting urban residential areas.     

Mosaic Pattern of Borrelia Infection in a Continuous Population of the Tick Ixodes Ricinus (Acari: Ixodidae)

An array of 12 20 × 20 m quadrats in a mixed forest near Poteply, Central Bohemia, Czech Republic, was investigated for the abundance and spatial distribution of host-questing Ixodes ricinus nymphs and their infection with borreliae. Tick densities were estimated by flagging and their borrelia infection status was determined by direct immunofluorescence. While the tick population appeared to be continuous and homogeneous based on quadrat counts, their infection with borreliae evinced a mosaic pattern. The nymphal infection rates ranged between 1.6 and 10.5% (mean = 6.0%) with significant differences between adjacent quadrats. Home ranging of small rodents (Apodemus flavicollis and Clethrionomys glareolus) inhabiting the forest seems to be responsible for the spatial pattern of borrelial infection.     

Abundance and seasonal activity of questing Ixodes ricinus ticks in their natural habitats in southern Germany in 2011

Questing ticks were sampled monthly over a period of 11 months from February, 2011 to December, 2011 at 13 sites in southern Germany using the flagging method. The ticks were identified to species, gender, and stadium. Although both I. ricinus and D. reticulatus were sampled, this study concentrated on I. ricinus, since it was the most abundant tick to be found. Additional weather data (air and soil temperature, relative air humidity, precipitation, sunshine duration) were recorded on each sampling site and the local vegetation described. A total of 14, 394 ticks was collected (7,862 larvae, 5,568 nymphs, 964 adults) and their activity was recorded in order to determine the seasonal activity pattern over different periods of the year. In contrast to the widely accepted pattern of a bimodal seasonal activity in moderate areas with a dominant peak in spring and a minor peak in autumn, a unimodal activity pattern was found for all development stages on six of the 12 sampling sites. Tick abundance was compared to weather variables. Tick host‐seeking activity was found to be significantly dependent on the temperature at ground level, precipitation, and sunshine duration as well as relative air humidity. Adult ticks showed a positive correlation with the duration of sunshine, whereas nymphs were mostly unaffected by this phenomenon.     

Lyme borreliosis spirochetes in Ixodes ricinus and Haemaphysalis punctata ticks (Acari: Ixodidae) on three islands in the Baltic Sea

The prevalence of infection with Lyme borreliosis spirochetes in nymphal Ixodes ricinus and Haemaphysalis punctata was examined on three Swedish islands in the Baltic Sea. The proportion of H. punctata nymphs infected with Borrelia spirochetes (2%) was lower than that of I. ricinus nymphs (11–16%), even on the island of Stora Karlsö where both tick species feed on the single mammalian host present, the varying hare, Lepus timidus. Ten of the 12 infected questing H. punctata nymphs harboured very few spirochetes, whereas the remaining two harboured as many as 30–40 spirochetes. However, it remains to be seen whether H. punctata nymphs are capable of transmitting spirochetes during their blood meal and, thus, serving as vectors for Lyme borreliosis spirochetes. On Gotland and Fårö, I. ricinus was more abundant than H. punctata in deciduous and coniferous forest, whereas H. punctata was equally or more abundant than I. ricinus in juniper brush areas on open land. Host preference and biotope usage of H. punctata seems, in general, to prevent this tick from feeding on highly infective Borrelia reservoirs, such as Apodemus mice.     

Sex-biased genetic structure in the vector of Lyme disease,Ixodes ricinus

Abstract.— We analyzed 725 Ixodes ricinus ticks (the principal vector of Lyme disease in Europe) collected in Switzerland in 1995 and 1996 (three and eight samples, respectively) and in Tunisia in 1996 (one sample) with five microsatellite markers. We found highly significant genetic differentiation between Swiss and Tunisian samples but detected almost no differentiation within Switzerland, even between those samples separated by the Alps. Interestingly, we found that I. ricinus females were more genetically related to one another than were males at a local scale, which would indicate a higher dispersal rate of immature males. Possible explanations for these findings in terms of sex-specific association of ticks with certain hosts (e.g., birds) and their epidemiological consequences are discussed.