Spiracular transpiration in ticks: a passive diffusion barrier in three species of Ixodidae (Metastigmata: Acarina)

In ixodid ticks the morphology of the spiracle, and in particular that of the labyrinth, differs between species. A reduction in transpiration of water vapour by the 'mutual interference' effect previously described in Ixodes ricinus (L.) is not significant in many species. Instead, transpiration is retarded by very restricted air movement within the labyrinth and by a high resistance to diffusion generated by numerous small aeropyles. These adaptations may be particularly useful in arid environments.     

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.     

Variability in cuticular hydrocarbons and phenotypic discrimination of Ixodes ricinus populations (Acarina: Ixodidae) from Europe

The cuticular hydrocarbon composition and a stepwise discriminant analysis are used to elucidate the phenotypic relationships of 66 populations of Ixodes ricinus in Europe. The method correctly allocates Ixodes persulcatus (outgroup) populations away from the main cluster of I. ricinus samples and separates the samples into ten relatively defined clusters of specimens. Populations from Poland are inseparable from samples collected in Germany, Switzerland and the Italian Alps, while individuals from Slovakia and the Czech Republic come into separate groups of phenotypic similarity. Irish and British specimens are separated but highly related and Spanish populations show an unexpectedly high distance from the remaining clusters.     

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.     

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.     

The effect of feeding status on sexual attractiveness of Ixodes ricinus (Acari: Ixodidae) females

Sexual attractiveness of unengorged, semiengorged and fully engorged females of both field and laboratory populations of Ixodes ricinus was studied under laboratory conditions by means of a computerised video tracking system. A male and a female were allowed to walk freely in a glass arena during 1 h and their behavioural interactions were observed. Obtained results revealed that the feeding status of I. ricinus females affects their sexual attractiveness. The highest attractiveness was observed in engorged females, the lowest in unengorged females of the field population. Copulation occurred in all experimental groups with the highest frequency observed in the unengorged virgin laboratory females. Despite engorged females being highly attractive for males on distance, the frequency of copulation was low. Differences between field and laboratory populations were found in both behavioural parameters and frequency of copulation. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

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.     

Genetic variation of ticks (Ixodes ricinus L.) in the Lithuanian and Norwegian populations

RAPD markers were used to measure the genetic diversity of 119 individuals of Ixodes ricinus collected from Lithuania and Norway. The samples were analysed within and also between the populations. We analysed 74 loci in each of 6 populations. Our results show high levels of diversity within the populations. The percentage of polymorphic loci of the six analysed populations: Birzai, Vilnius, Kretinga, Tjore, Kjosvik and Odderoya were 68.9%, 58.1%, 78.38%, 62.2%, 44.6% and 68.9%, respectively. The percentage of polymorphic loci in the Lithuanian populations was 93.2%, and in the Norwegian populations 81.08%. The genetic distance ranged from 0.019 to 0.079 within Norwegian populations and from 0.005 to 0.0967 within Lithuanian populations and between the countries from 0.022 to 0.146. The genetic variation of I. ricinus among Norwegian populations was lower than among Lithuanian populations. The highest part of genetic variation in I. ricinus ticks depends on variation within Kretinga (Lithuania) and Odderoya (Norway) populations situated in coastal areas where many migratory and sea birds are aggregated.     

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.     

Seasonal incidence and ecology of the tick Ixodes ricinus (Acari: Ixodidae) on grazing pastures in Western France

A longitudinal survey was carried out during a 2 year period in Western France to assess the infestation level of grazing pastures byIxodes ricinus ticks. Four farms were visited once a month and each of the grazing pastures was sampled in the centre and at the border using the blanket dragging method. A total of 3562I. ricinus (34 adults, 900 nymphs and 2628 larvae) were collected and the infestation was significantly higher during the first year (p<0.0001). The infestation level byI. ricinus varied between grazing pastures and farms. Grazing pastures in the vicinity of forest were more infested than the others, all through the study. The seasonal distribution of ticks showed peaks, with low fluctuations between farms, years and stages. Tick abundance could not be related to vegetation, but only to the vicinity of woods.     

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.     

Functional morphology of the spiracles in two species of Argasidae (Metastigmata: Acarina), with particular reference to responses to desiccation

Spiracle morphology of Argas persicus (Oken) and Omithodorus acinus Whittick is described and the differences between the two species are related to biological factors. Argasid and ixodid spiracle structures and functions are compared.     

The longevity of the tick Ixodes ricinus (Acari: Ixodidae) in Central Russia

The longevity of the tick Ixodes ricinus has been studied on 27 samples from a wild population of the Moscow Region. Ticks were kept in small cages and glass tubes under conditions close to natural. About 2/3 of the specimens collected in spring survived till the next spring and later died out gradually during the second season of their activity. Very few specimens reached the beginning of the third winter in their life, but apparently none of them survived the end of the winter. The life span of males was the same as that of females. No mass activation of adult ticks was recorded in the autumn of the year of their molting. It seems that the rise of the autumnal density of adult ticks depends mainly on the increased activity of the females which molted from nymphs last year.     

Recent discovery of widespread Ixodes affinis (Acari: Ixodidae) distribution in North Carolina with implications for Lyme disease studies

Ixodes affinis, which is similar morphologically to Ixodes scapularis, is widely distributed in North Carolina. Collections have documented this species in 32 of 41 coastal plain counties, but no piedmont or mountain counties. This coastal plain distribution is similar to its distribution in Georgia and South Carolina, where it is considered an enzootic vector of Borrelia burgdorferi sensu stricto. An updated list of hosts for I. affinis in the U.S.A. is included, increasing the number to 15 mammal and one bird species. The presence of questing adults of I. affinis from April to November reinforces the need for confirmed identifications of suspected tick vectors of Borrelia spirochetes collected during warm months.     

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.