The tick Ixodes ricinus: distribution and climate preferences in the western Palaearctic

In this study, multivariate spatial clustering on monthly normalized difference vegetation index (NDVI) maps is used to classify ecological regions over the western Palaearctic. This classification is then used to delineate the distribution and climate preferences of populations (clades) of the tick Ixodes ricinus L. (Acari: Ixodidae) from a geographically extensive dataset of tick records and a gridded 2.5-km resolution climate dataset. Using monthly layers of the NDVI, regions of similar ecological attributes were defined and nine populations with significant differences in critical climate parameters (P< 0.005) were detected. Grouping of tick records according to other categories, such as political divisions, a 4 degrees x 4 degrees grid overlying the study area, or the CORINE) and USGS) vegetation classification schemes did not provided significantly separated populations (P = 0.094-0.304). Factor analysis and hierarchical tree clustering provided an ecological overview of these tick clades: two Mediterranean and one Scandinavian (western) clades are clearly separated from a node that includes clades of different parts of central Europe and the British Isles, with contrasting affinities between the different clades. The capture records of these ecologically separated clades produce a clear bias when bioclimate envelope modelling is applied to the mapping of habitat suitability for the tick in the western Palaearctic. The best-performing methods (Cohen's kappa = 0.834-0.912) use partial models developed with data from each ecoregion, which are then overlapped over the region of study. It is concluded that the use of ecologically derived ecoregions is an objective step in assessing the presence of ecologically different clades, and provides a guide in the development of data partitioning for habitat suitability modelling.     

Climate warming and changes in habitat suitability for Boophilus microplus (Acari: Ixodidae) in Central America.

Remote sensing on the basis of AVHRR (advanced very high resolution radiometer) satellite imagery was used, together with standard geostatistical methods (cokriging), to estimate the distribution of habitat suitability (HS) for the tick Boophilus microplus (Canestrini) in Central America. Most attention was paid to the expected limits of tick distribution as well as the temperature and vegetation values responsible for different HS zones and their variability within the year. The highest HS extended through wide areas of southeastern United States, much of Mexico, and other countries of Central America. Areas with higher HS had relatively stable temperatures (within 20-25 C) throughout the year, and had a NDVI (normalized derived vegetation index) of around 0.4. These areas need to be targeted to ensure acaricide usage at optimum intervals and to avoid the formation of areas with enzootic instability. A seasonal analysis of the climate trend through the study period (1982-1999) revealed a warming cycle, together with a rise in NDVI index values. The main consequence of this trend is the gradual increase in unsuitability in areas where the tick is already established, with the establishment of new foci in zones currently too cold to support tick populations. The cooling periods of 1-2 yr that were observed between warming cycles may also lead to enzootic instability when warm zones cool to within the suitability range. As the model is remotely sensed, a continuous update of the B. microplus distribution could be performed, assuring maximum efficiency in developing management strategies.     

The relationships between habitat topology, critical scales of connectivity and tick abundance Ixodes ricinus in a heterogeneous landscape in northern Spain

The habitat mosaic was used to quantify connectivity between patches of different tick density of the notorious tick species Ixodes ricinus in an attempt to determine the cause of variations in tick abundance among apparently homogeneous sites in northern Spain. The analysis revealed that patches with high tick abundance are "stepping-stone" territories that, when removed from the landscape, cause large changes in connectivity. Sites with medium tick abundance do not cause such a critical transition in connectivity. Patches with low tick abundance, but optimal abiotic conditions for survival, are located within the minimum cost corridors network joining the patches, while those sites where the tick has been intermittently collected are located at variable distances from this network. Sites where the tick is consistently absent, but where the habitat is predicted to be suitable (old, heterogeneous forests of Quercus spp.) for the tick, are very separated from this main network of connections. These results suggest that tick distribution in a zone is highly affected not only by abiotic variables (vegetation and weather) but also by host movements. Dispersal of the tick is a function of how the hosts perceive the habitat, and the habitat's permeability to host movement. Permanent tick populations seem to be supported by the existence of these critical, high density patches, located at significant places within the habitat network.     

Correlation of Borrelia burgdorferi sensu lato prevalence in questing Ixodes ricinus ticks with specific abiotic traits in the western palearctic

This meta-analysis of reports examining ticks throughout the Western Palearctic region indicates a distinct geographic pattern for Borrelia burgdorferi sensu lato prevalence in questing nymphal Ixodes ricinus ticks. The greatest prevalence was reported between the 5°E and 25°E longitudes based on an analysis of 123 collection points with 37,940 nymphal tick specimens (87.43% of total nymphs; 56.35% of total ticks in the set of reports over the target area). Climatic traits, such as temperature and vegetation stress, and their seasonality correlated with Borrelia prevalence in questing ticks. The greatest prevalence was associated with mild winter, high summer, and low seasonal amplitude of temperatures within the range of the tick vector, higher vegetation indices in the May-June period, and well-connected vegetation patches below a threshold at which rates suddenly drop. Classification of the target territory using a qualitative risk index derived from the abiotic variables produced an indicator of the probability of finding infected ticks in the Western Palearctic region. No specific temporal trends were detected in the reported prevalence. The ranges of the different B. burgdorferi sensu lato genospecies showed a pattern of high biodiversity between 4°W and 20°E, partially overlapping the area of highest prevalence in ticks. Borrelia afzelii and Borrelia garinii are the dominant species in central Europe (east of ～25°E), but B. garinii may appear alone at southern latitudes and Borrelia lusitaniae is the main indicator species for meridional territories.     

Multi-criteria Decision Analysis to Model <Emphasis Type="Italic">Ixodes ricinus</Emphasis> Habitat Suitability

Tick-borne diseases present a major threat to both human and livestock health throughout Europe. The risk of infection is directly related to the presence of its vector. Thereby it is important to know their distribution, which is strongly associated with environmental factors: the presence and availability of a suitable habitat, of a suitable climate and of hosts. The present study models the habitat suitability for Ixodes ricinus in Ireland, where data on tick distribution are scarce. Tick habitat suitability was estimated at a coarse scale (10 km) with a multi-criteria decision analysis (MCDA) method according to four different scenarios (depending on the variables used and on the weights granted to each of them). The western part of Ireland and the Wicklow mountains in the East were estimated to be the most suitable areas for I. ricinus in the island. There was a good level of agreement between results from the MCDA and recorded tick presence. The different scenarios did not affect the spatial outputs substantially. The current study suggests that tick habitat suitability can be mapped accurately at a coarse scale in a data-scarce context using knowledge-based methods. It can serve as a guideline for future countrywide sampling that would help to determine local risk of tick presence and refining knowledge on tick habitat suitability in Ireland.     

Distribution,density and migration dynamics of Ixodes ricinus in an area of the Jurassic mountains of Switzerland

In an area of the Jurassic mountains of Switzerland ecological aspects of the tick species Ixodes ricinus were studied. In particular, the influence of temperature, humidity and vegetation parameters on tick density and distribution were investigated several times in six different habitat types. This was carried out on two scales; on a small scale, within a habitat type and on a larger scale, where all six habitat types were included. By a marking experiment, aspects of the migration dynamics of the ticks were studied. The densities of the six habitat types differed significantly in most of the cases. An analysis of the distribution pattern within a habitat type showed a contagious distribution mostly for the forests and the forest edge, whereas a random distribution was found as well, mainly for the meadows. But these distribution patterns tend towards randomness with time. On a small scale none of the factors showed an influence on tick distribution, therefore the most important factor on this scale might be the hosts of Ixodes ricinus. On a larger scale, temperature and humidity do in some cases influence tick densities. The marking experiment suggests an extremely high turnover of the ticks in an area with an average host density, probably due to passive transportation by hosts and to activation of resting individuals.     

Using satellite data to forecast the occurrence of the common tick Ixodes ricinus (L.)

To forecast the incidence of the common tick, Ixodes ricinus, the authors used the finding that some types of vegetation may indicate the presence of the tick. To obtain the necessary information in a scope which would enable its practical application in medicine to prevent populational exposure to ticks, remote sensing data were utilized as obtained from the Multispectral Scanner operating aboard Landsat 5. A file was selected from the full scene representing a territory which measured 41 by 41 kilometers and in the centre of which there was an area known to have consistently high tick numbers, as revealed in a 25-year continuous study of I. ricinus, and which proved to be a natural focus of tick-borne encephalitis. This area (Potepl) was taken as a model and was compared with its surroundings. Six landscape classes were examined (1--coniferous forest, 2--leaved forest, 3--mixed forest, 4--water basins, 5--glades, 6--housing developments), the former three being of crucial importance to evaluate the probability of tick presence. The 6th category is significant in assessing human exposure to ticks. Data processing was carried out by supervised classification using the Baye's decision rule of maximum likelihood. The findings were obtained both in a graphic form and in the form of statistical reviews as regards the presence of appropriate landscape categories.     

Effect of forest clearing on the abundance of Ixodes ricinus ticks and the prevalence of Borrelia burgdorferi s.l

Questing Ixodes ricinus L. (Acari: Ixodidae) ticks were collected on a forest trail that had been completely cleared of shrubs and ground vegetation in winter 2002 and on a nearby control uncleared forest transect in South Moravia (Czech Republic). Samples were collected each May in 2003, 2004 and 2005. Nymphal ticks were 3.4 times, 1.9 times and 1.2 times less frequent on cleared forest than on uncleared forest trails in the three respective years, whereas adult tick abundance was 27.2 times, 4.0 times and 2.2 times lower, respectively. The ticks were examined for borreliae by dark-field microscopy: prevalence of nymphal ticks infected with Borrelia burgdorferi sensu lato (12.6% to 20.0%) did not differ significantly between the cleared and uncleared trail during the 3 years. In conclusion, the habitat modification appeared to result in a decreased abundance of I. ricinus as well as a reduced frequency of infected ticks (and thus indirectly a lower potential risk of Lyme borreliosis), which lasted, however, for only 2 years. Eight cultures of borreliae isolated from the ticks were all identified as the 'ornithophilic' genomic species Borrelia garinii, possibly indicating a greater role of forest birds than that of forest rodents as the hosts of immature I. ricinus in the tick (and borrelial) colonization of the cleared part of the forest.     

Lacerta bilineata (Reptilia,Lacertidae) as a host of Ixodes ricinus (Acari,Ixodidae) in a protected area of northern Italy

A survey on the importance of Lacerta bilineata as host of Ixodes ricinus was conducted in Northern Italy over a three-year period. A total of 202 western green lizards were captured and a total of 2349 ticks were collected. All ticks were identified as I. ricinus; 53.2% and 46.7% were at the larval and nymphal stages, respectively. Tick number and prevalence were higher in males than in females, especially from April to June during the host breeding period. The level of tick infestation increased with lizard age and size. The number of ticks collected on adult lizards peaked in June and in August. Infestation levels appear to be related to lizard activity patterns and behaviour. Tick number and prevalence also varied in relation to host habitat, infestation being higher in lizards from areas with hard vegetation cover.     

Factors driving the abundance of ixodes ricinus ticks and the prevalence of zoonotic I. ricinus-borne pathogens in natural foci

Environmental factors may drive tick ecology and therefore tick-borne pathogen (TBP) epidemiology, which determines the risk to animals and humans of becoming infected by TBPs. For this reason, the aim of this study was to analyze the influence of environmental factors on the abundance of immature-stage Ixodes ricinus ticks and on the prevalence of two zoonotic I. ricinus-borne pathogens in natural foci of endemicity. I. ricinus abundance was measured at nine sites in the northern Iberian Peninsula by dragging the vegetation with a cotton flannelette, and ungulate abundance was measured by means of dung counts. In addition to ungulate abundance, data on variables related to spatial location, climate, and soil were gathered from the study sites. I. ricinus adults, nymphs, and larvae were collected from the vegetation, and a representative subsample of I. ricinus nymphs from each study site was analyzed by PCR for the detection of Borrelia burgdorferi sensu lato and Anaplasma phagocytophilum DNA. Mean prevalences of these pathogens were 4.0% ± 1.8% and 20.5% ± 3.7%, respectively. Statistical analyses confirmed the influence of spatial factors, climate, and ungulate abundance on I. ricinus larva abundance, while nymph abundance was related only to climate. Interestingly, cattle abundance rather than deer abundance was the main driver of B. burgdorferi sensu lato and A. phagocytophilum prevalence in I. ricinus nymphs in the study sites, where both domestic and wild ungulates coexist. The increasing abundance of cattle seems to increase the risk of other hosts becoming infected by A. phagocytophilum, while reducing the risk of being infected by B. burgdorferi sensu lato. Controlling ticks in cattle in areas where they coexist with wild ungulates would be more effective for TBP control than reducing ungulate abundance.     

Tick species infesting ruminants in Italy: ecological and bio-climatic factors affecting the different regional distribution

The distribution of tick species able to infest ruminants in Italy is briefly reviewed on the basis of published records and new data obtained in several surveys carried out mainly in northern Italy. The most frequent and abundant tick species found on ruminants throughout Italy are Ixodes ricinus and Dermacentor marginatus, the first gradually replacing its association with sheep by the association with wild ruminants. Rhipicephalus bursa was mainly found on cattle while R. turanicus, which is considered one of the most sheep-associated species, was found only on wild boar. Haemaphysalis punctata, H. sulcata and H. parva are the other tick species quite frequently found on small ruminants when they are reared on pastures mainly in central-southern Italy. However, the land exploitation of these last decades has dramatically reduced the diversity of Italian environment and significantly modified the distribution and the abundance of ticks causing a heavy spread of the tick species characterized by a strong adaptability such as I. ricinus and R. sanguineus. If this trend continues, it is possible to hypothesise that few "resistant" tick species will gradually replace most of the others. This hypothesis seems to be confirmed by the high percentage observed for I. ricinus (57.4%) and R. sanguineus (35.73%) out of 8,712 specimens collected from several different hosts and from the environment.     

Distribution of Ixodes ricinus in the British Isles: investigation of historical records

Ixodes ricinus Linnaeus (Acari: Ixodidae) is the most abundant and widely distributed tick in the British Isles, and is a vector for a number of bacterial, viral and protozoal pathogens of both medical and veterinary importance. This report provides an update to the historical distribution data of I. ricinus, published by the Biological Records Centre (BRC), Monks Wood in The Provisional Atlas of the Ticks (Ixodidae) of the British Isles by K. P. Martyn (1988), and is supplemented with additional BRC records since 1988, additional data from published scientific literature and unpublished field studies, and enhanced with spatial and temporal information on tick stages collected and their host associations. Records have been mapped at 10 km resolution and enhanced to 5 km, 1 km and 0.1 km. Differentiation between records representing one-off collections from those representing populations of I. ricinus has been achieved through the classification of the records into either reported or established populations. Detailed seasonality and host associations of records are investigated, highlighting the value in obtaining additional detailed contemporary data to aid risk assessments and research within this field.     

Using Species Distribution Models to Predict Potential Landscape Restoration Effects on Puma Conservation

A mosaic of intact native and human-modified vegetation use can provide important habitat for top predators such as the puma (Puma concolor), avoiding negative effects on other species and ecological processes due to cascade trophic interactions. This study investigates the effects of restoration scenarios on the puma’s habitat suitability in the most developed Brazilian region (São Paulo State). Species Distribution Models incorporating restoration scenarios were developed using the species’ occurrence information to (1) map habitat suitability of pumas in São Paulo State, Southeast, Brazil; (2) test the relative contribution of environmental variables ecologically relevant to the species habitat suitability and (3) project the predicted habitat suitability to future native vegetation restoration scenarios. The Maximum Entropy algorithm was used (Test AUC of 0.84 ± 0.0228) based on seven environmental non-correlated variables and non-autocorrelated presence-only records (n = 342). The percentage of native vegetation (positive influence), elevation (positive influence) and density of roads (negative influence) were considered the most important environmental variables to the model. Model projections to restoration scenarios reflected the high positive relationship between pumas and native vegetation. These projections identified new high suitability areas for pumas (probability of presence >0.5) in highly deforested regions. High suitability areas were increased from 5.3% to 8.5% of the total State extension when the landscapes were restored for ≥ the minimum native vegetation cover rule (20%) established by the Brazilian Forest Code in private lands. This study highlights the importance of a landscape planning approach to improve the conservation outlook for pumas and other species, including not only the establishment and management of protected areas, but also the habitat restoration on private lands. Importantly, the results may inform environmental policies and land use planning in São Paulo State, Brazil.     

Normalized difference vegetation index (NDVI) as a predictor of forage availability for ungulates in forest and field habitats

Quantifying available plant biomass is a crucial step towards improving our understanding of herbivore ecology and trophic interactions. Thanks to the development of satellite-derived vegetation indices such as the normalized difference vegetation index (NDVI), ecologists have been provided with indirect estimates of primary production at various temporal and spatial scales. When it comes to forested ecosystems, most mammalian herbivores predominantly rely on the ground vegetation, yet little is known regarding the suitability of NDVI to predict this component of forest vegetation cover. This study compares the relationship between NDVI and ground vegetation biomass in two contrasting habitats (field and forest) in Eastern Poland over the spring and summer seasons (2007–2008). Results indicate that seasonality shapes the relationship between NDVI and ground vegetation biomass for each habitat type. In the field habitat, NDVI and ground vegetation biomass were positively related, with a stronger correlation between the two variables occurring in summer. In the forest habitat, a switch in the direction of the correlation between biomass and NDVI (from positive in spring to negative in summer) was detected. The timing of the switch was related to the timing of full development of tree and shrub leaves (late May–early June). This suggests that the usefulness of NDVI as a predictor of ground vegetation biomass is dependent upon the habitat considered and the targeted season.     

The geographic distribution and ecological preferences of the American dog tick,Dermacentor variabilis (Say), in the U.S.A.

Equine piroplasmosis (EP), caused by two parasitic organisms, Theileria equi and Babesia caballi, is a tick‐borne disease of recent concern in horses in the U.S.A. Outbreaks of EP have been detected in Florida, Missouri, Kansas and Texas. In 2009, EP transmission in Texas occurred through the adults of two tick species, Amblyomma mixtum [formerly known as Amblyomma cajennense (Fabricius, 1787)] Koch (Ixodida: Ixodidae) and Dermacentor variabilis (Say) (Ixodida: Ixodidae), the American dog tick (ADT). In this study, we developed a continent‐scale map for the distribution of the EP vector species D. variabilis, using a presence‐only modelling approach to assess the habitat preferences of this tick. We used identification records from our tick geodatabase of locations in which the presence of the ADT had been noted. The potential distribution of the ADT in the U.S.A. was estimated from environmental factors using the maximum entropy approach based on localities in which there is a high probability of occurrence according to habitat suitability. Elevation and temperature were found to be biologically significant environmental variables influencing the presence of this tick species. Properly designed and constructed probability surfaces using maximum entropy offer one useful approach to the mapping of distribution ranges of tick species based on suitable habitat in the U.S.A.     

IgG responses to salivary gland extract of Ixodes ricinus ticks vary inversely with resistance in naturally exposed sheep

Enzyme-linked immunosorbent assay (ELISA) was used to investigate the antibody responses of control sheep, and sheep naturally exposed to Ixodes ricinus Linné (Acari: Ixodidae) ticks, to salivary gland extract (SGE) proteins of partially fed, adult I. ricinus. Comparisons between responses of control sheep and naturally infested sheep by Western blot analysis suggested that variations in IgG responses of I. ricinus-exposed sheep were mostly associated with specific responses to I. ricinus SGE antigens. Sheep IgG responses were positively related to the numbers of adult ticks feeding per sheep at the time samples were collected, were greater during the spring than the autumn periods of I. ricinus activity and were inversely related to sheep resistance to ticks measured by the weights of nymphal I. ricinus that engorged on the sheep. These findings suggest that sheep lose their resistance to ticks due to polarization of a Th1 type response to some tick antigens towards a Th2 type response when sheep are exposed to high, natural tick infestations, or to seasonal conditions of relative nutritional stress. Potential consequences for the epidemiology of tick-borne diseases are discussed.     

Acarological studies in two protected areas of Central Italy

In the present note are reported the results of preliminary studies on tick distribution in two wildlife areas of Abruzzo (National Park of Abruzzo, a mountainous area, approximately 40,000 ha, in Central Apennines, interesting Abruzzo, Latium and Molise regions) and Latium (Capocotta-Castel Porziano Presidential reserve, on the Tyrrhenian coast, 30 km from Rome). Sampling of ticks from domestic and wild mammals as well as from vegetation, was performed in three different areas of the National Park in 1995. Tick species identified include Rhipicephalus bursa, R. sanguineus, Hyalomma marginatum, Haemaphysalis punctata, Dermacentor marginatus, Ixodes ricinus, I. hexagonus. The presence of I. ricinus was discontinuous and sporadic. In Capocotta estate, samplings were performed bimonthly from March to November 1997 in an restricted area (1 ha) with typical Mediterranean flora and fauna. The following species were collected: I. ricinus, Haemaphysalis concinna, D. marginatus, R. bursa, Hy. marginatum. There was a high dominance of I. ricinus and H. concinna.     

Habitat and Vegetation Variables Are Not Enough When Predicting Tick Populations in the Southeastern United States

Two tick-borne diseases with expanding case and vector distributions are ehrlichiosis (transmitted by Amblyomma americanum) and rickettiosis (transmitted by A. maculatum and Dermacentor variabilis). There is a critical need to identify the specific habitats where each of these species is likely to be encountered to classify and pinpoint risk areas. Consequently, an in-depth tick prevalence study was conducted on the dominant ticks in the southeast. Vegetation, soil, and remote sensing data were used to test the hypothesis that habitat and vegetation variables can predict tick abundances. No variables were significant predictors of A. americanum adult and nymph tick abundance, and no clustering was evident because this species was found throughout the study area. For A. maculatum adult tick abundance was predicted by NDVI and by the interaction between habitat type and plant diversity; two significant population clusters were identified in a heterogeneous area suitable for quail habitat. For D. variabilis no environmental variables were significant predictors of adult abundance; however, D. variabilis collections clustered in three significant areas best described as agriculture areas with defined edges. This study identified few landscape and vegetation variables associated with tick presence. While some variables were significantly associated with tick populations, the amount of explained variation was not useful for predicting reliably where ticks occur; consequently, additional research that includes multiple sampling seasons and locations throughout the southeast are warranted. This low amount of explained variation may also be due to the use of hosts for dispersal, and potentially to other abiotic and biotic variables. Host species play a large role in the establishment, maintenance, and dispersal of a tick species, as well as the maintenance of disease cycles, dispersal to new areas, and identification of risk areas.