Figure 1

Figure 1. Geographic distribution of study sites ranked by abundance of Ixodes scapularis in Wisconsin, northern Illinois, and Menominee County in Michigan.

The locations of the 138 sites that were sampled in Wisconsin, Illinois, and Michigan are shown in Figure 1. Among the four categories, 56 sites were classified as negative, 24 were ranked as 1, 32 as 2, and 26 as 3. Most negative sites were in northeastern Wisconsin. In the southeastern part of Wisconsin, sites were negative except those situated in the Kettle Moraine State Forests (Sheboygan, Fond du Lac, Jefferson, Walworth, and Waukesha counties), which are located on the terminal glacial moraines. Negative sites in Illinois were at Blackhawk Nature Preserve (Rock Island County), located in a suburban area, and White Pines State Park (Ogle County), which has large stands of secondary growth pine forest. In Wisconsin, positive high-density sites were found in the southwestern driftless area and in the central sandy uplands, as well as in the well-recognized northwest part of the state (and across the state line into Minnesota).

In Michigan, where only a small area of the Upper Peninsula was sampled, all sites had very dense tick populations, except for a site that was classified as excessively drained (>99% sand). The sites classified in the other two abundance categories (1 and 2) did not appear to cluster in any areas. In Illinois, the two parks that have been infested for at least a decade, Castle Rock State Park (Ogle County) and Loud Thunder Forest Preserve (Rock Island County), were classified as having dense tick populations, with lower populations in some sites along the Illinois River.

Figure 2

Figure 2. Soil particle size analysis of samples from positive and negative sites. Soil texture is expressed as the sum of percent sand, silt, and clay.

Particle size analysis, which is a function of the proportions of sand, silt, and clay, was performed at 82 sites (Figure 2). The positive sites were clustered in the sand/loamy sand texture classes. Individual percentages of sand, silt, and clay per sample were not correlated with tick abundance; however, texture class, which is a combination of these three percentages, correlated significantly (r=0.42, p<0.05) with greater tick densities found in soils with a greater proportion of sand. The soil texture class of samples determined from the soil analysis correlated significantly (r=0.46, p<0.001) with the soil texture class of each site as obtained from the STATSGO database.

Figure 3

Figure 3. Categories of environmental variables and number of positive and negative sites.

Figure 4

Figure 4. Map of soil orders in Wisconsin and northern Illinois, overlaid with tick study sites.

The univariate analysis detected significant associations (p<0.25) between tick presence and land cover, soil order, bedrock geology, quaternary geology, soil texture, forest type, spring, summer, fall and winter precipitation, snowfall, and elevation (Figure 3). The results of the discriminant analysis are listed in Table 1. When negative and positive sites were contrasted, the variables forest type, soil order, land cover, soil texture and bedrock were significant. Tick presence was positively associated with deciduous (Figure 3a), dry/mesic and dry forests (Figure 3b), fertile soils such as alfisols (Figures 3c, 4), sand and loamy/sand soil texture (Figures 2, 3d), and sedimentary bedrock (Figure 3f). There was a negative association with grasslands and conifer forests (Figure 3a), wet and wet/mesic forests (Figure 3b), acidic soils such as spodosols (Figure 3c), clay soil texture (Figure 3d), and Precambrian bedrock (Figure 3e). Elevation was not an important discriminator in the model, nor was Quaternary geology (Figure 3f) important even though sites located on the plateaus and loess-covered areas were all positive. However, the distribution of the sites among the categories of Quaternary deposits was skewed because a large proportion of the state parks were located on terminal glacial moraines. The discriminate model was able to correctly classify 85.7% of the sites. The canonical correlation coefficient was 0.69, and the eigenvalue was close to 1 (0.92), indicative of a strong discriminant function. When the single stage category (1) was included with the negative group, only two variables, forest type and soil order, were significant. Most of the sites (78.6%) were still correctly classified; however, the eigenvalue decreased to 0.43. These same variables were significant when all the groups were considered separately; but the model only correctly classified 51.8% of the sites. Even though only 4/33 in the negative group were misclassified, there was very poor discrimination among the tick positive groups. No significant variables resulted from the discriminant analysis performed using the satellite data. Since all sites were located in forested areas, TM imagery may not have been able to discriminate well among suitable and unsuitable forested habitats. The precipitation variable was also not a significant discriminator between positive and negative sites in the model.

Figure 5

Figure 5. Predictive risk map of habitat suitability for Ixodes scapularis in Wisconsin and Illinois.

The results of the logistic regression analysis were in agreement with the discriminant analysis model in the positive versus negative group as seen in Table 2. The same variables were significant (p<0.05), and the model correctly classified 83.9% of the sites. The predictive risk map generated from the logistic regression model is shown in Figure 5. The higher probabilities indicate increased suitability of habitat for I. scapularis. In Wisconsin, the areas of moderate suitability (26%-40%) are located in the western half of the state. Patchy areas of higher probability (60%-100%) are found in the central and northern portion (Juneau, Adams, Waushara, and Marquette counties.) and along the border with Minnesota (Vernon and Crawford counties). In Illinois, the positive sites that were sampled corresponded to areas of increased suitability (60%-100%). Castle Rock State Park, where the highest tick densities are found, had a 90%-100% probability of suitable habitat. The areas bordering the Illinois River appear to be adequate habitat for I. scapularis, especially on the western side. Shawnee National Forest in the extreme southern portion of the state also appears to have a high probability (60%-80%), even though I. scapularis populations have not been detected (42).

Ixodes scapularis may be introduced into new areas by several routes. Adult I. scapularis are carried into new areas primarily by deer (43), which are capable of ranging over wide areas, especially along riparian corridors. However, infected adult ticks have limited potential for spreading Lyme disease since transovarial transmission of B. burgdorferi is rare. Small mammals are efficient disease reservoirs, and juveniles tend to disperse during the spring and summer when tick larvae and nymphs are questing. However, the potential for long-range dispersal of Lyme disease by rodents is limited, since they occupy much smaller home ranges than deer (44). Birds have a high potential for introducing infected immature stages of I. scapularis into distant areas (45–47), especially during spring and fall migration.

To become successfully established in a new area, I. scapularis requires available hosts for feeding, which is not a limiting factor in our study area, and a suitable habitat for questing, molting, diapause, and oviposition. The vegetation, soil, topography, and climate are interrelated, and extremes of any one factor may adversely affect the tick’s ability to survive.

The environmental characteristics vary throughout the two states, and certain combinations may determine whether introduced I. scapularis populations can become established. Tick abundance is an indicator of the suitability of environmental conditions for reproduction and survival. Finding only one stage of the tick may indicate either a poor microenvironment or a recent introduction. Finding all three stages at one site strongly suggests that a population has become established. A less than optimal habitat may account for low density in an established I. scapularis population, or it may indicate a recent introduction. Errors in classification may occur in an extensive field survey, as reported here, and a dynamic situation (i.e., the process of invasion of a new site) may mask the occurrence of some positive or potentially positive sites. By including a large number of sites and conducting repeat visits, we have tried to minimize such confounding effects.

Environmental factors such as bedrock geology, quaternary deposits, soils, vegetation, and climate influence each other directly and indirectly to create unique habitats. This is why we included risk factors that are not necessarily independent in a model that is most unbiased. The soil orders in the region (Figure 4) are influenced by the type of underlying bedrock and by quaternary deposits. The soils, in turn, influence the type of vegetation overlying them. Soil texture is the component of soil that influences the extent of drainage. The soil texture classes are independent of soil order and are usually a function of the degree of soil weathering and the parent material (bedrock or quaternary deposit). The tree composition of a forest is determined by a moisture gradient involving soil aeration, soil nutrient supply, and microclimatic features (38), and this gradient functions as a continuum. The forest types classified as dry and dry/mesic have oaks and jack pines as the dominant species that prefer well-drained, sandy soils. Oak forests also have a dense canopy layer that provides protection for the underlying vegetation. Wet and wet/mesic forests are composed of trees that have a high tolerance for very moist soils. The factors interacting at the microclimatic level within the topsoil and leaf litter appear to have an important influence on tick survival. Excessively moist conditions at the soil level were negatively associated with the establishment of I. scapularis. Soil texture, in addition to the topography, determines the extent of drainage, and the level of moisture of the ground layer, regardless of the amount of precipitation. However, given the effect of weather on tick abundance (19), associations between tick presence and amount of yearly precipitation or snowfall need to be analyzed further.

Our findings suggest that abiotic factors play a major role in determining whether populations of I. scapularis can become established in an environment. Precambrian bedrock of volcanic origin results in the formation of acidic soils that are found mainly under coniferous forests, the forest type least likely to support tick populations. Soils containing increased acidity (spodosols) and a high proportion of clay that can retain excess moisture (48) were also more frequently present in negative sites. Excessive moisture in the soil may be deleterious to tick survival since they overwinter in the topsoil and leaf litter. It may also enhance the growth of organisms, such as fungi and entomophagous nematodes, which may have adverse effects on the tick population (49). Leaf litter is a necessary component for the survival of immature stages of I. scapularis (50). However, the type and quantity may determine the densities of ticks in a specific habitat. Tick densities were highest in forests dominated by oak, followed by maple, and lowest in coniferous forests that produce minimal amounts of leaf litter (38). Tick densities were also highest in areas with underlying sedimentary bedrock, which is associated with alfisol and mollisol soil orders and soil textures of increased particle size (38).

The statistically significant risk factors derived from the logistic regression analysis were in agreement with those obtained from the discriminant analysis, and allowed us to quantify and predict the environmental risk for the presence of I. scapularis. Several environmental factors must be evaluated simultaneously to assess the combination of factors required for successful establishment. Determining the environmental factors that limit survival can facilitate the development of measures for the control of the tick in the environment.

Using a GIS, we generated a risk map (Figure 5) to predict the presence of the tick vector, I. scapularis. The areas of suitable habitat for I. scapularis in Wisconsin corresponded to areas of increased incidence of human Lyme disease and known areas of tick endemicity. The extensive area of suitable habitat in the western portion of the state can explain the rapid expansion of the tick from the original northwestern focus to the southwestern portion of the state (2–5). While initial studies of tick distribution (3) and human granulocytic ehrlichiosis (51) point to the risk of tick-borne disease transmission in Northwest Wisconsin, our study points also to the sandy barrens of Central Wisconsin as most suitable habitats. Indeed, the highest numbers of ticks were collected in Council Ground State Park (Lincoln County), Fort McCoy (Monroe County), Hartman Creek State Park (Waupaca County), and Wildcat Mountain State Park (Vernon County), as well as in sites in the long-recognized Spooner area (Washburn County). Further, the highest prevalence of canine seropositivity to B. burgdorferi in northern Illinois and Wisconsin was found in dogs in the west-central counties of Wisconsin (52). Based on the risk map, most of the north-central and northeastern portions of Wisconsin have a <25% probability for tick presence. These are areas where our sampled sites were consistently negative for I. scapularis. In the eastern half of the state, the main areas of increased suitability were along the glacial terminal moraines, which is where the positive sites in the Kettle Moraine State Forest were located. There was also a higher probability in the northeastern corner of the state bordering Menominee County, Michigan, where positive sites were located.

In Illinois, areas of increased suitability corresponded to the same areas where the positive sites were located in Ogle, Rock Island, and Jo Daviess counties. The risk map indicated there is adequate habitat for I. scapularis populations to become established along the Illinois River, as well as the Mississippi River. However, in Illinois, tick populations may be limited to river corridors since extensive areas are used for agriculture. Where forested habitat is sparse, tick establishment may be restricted, even though geologic and soil factors are favorable. In southern Illinois, where climatic conditions may differ and other reservoir hosts may be present, the inclusion of additional parameters to the model may result in reduced risk probabilities. In contrast, the risk factor model and predictive map may be valid for other north-central areas that have similar environmental characteristics, particularly in parts of Minnesota, Michigan, northern Indiana, and Ohio. The model may be applied to other areas of the United States by using local geographic coverages.

In conclusion, this model can be used to help determine the risk of acquiring Lyme disease and other diseases transmitted by I. scapularis by predicting which locations may be currently infested with the tick. It can also be used to assess whether habitats that are currently nonendemic for I. scapularis would have the necessary combination of environmental factors to allow new populations of I. scapularis to become established. The model can thus be continuously refined based on findings from new areas. The risk of Lyme disease transmission could be predicted in areas capable of sustaining I. scapularis populations if ticks harboring B. burgdorferi are introduced by migrating deer or birds. The results obtained from these field studies can also form the basis for controlled experimental studies under field and laboratory conditions to further elucidate the preferred microenvironment of I. scapularis.