Habitat Suitability Model for the Distribution of Ixodes scapularis (Acari: Ixodidae) in Minnesota.

Ixodes scapularis Say, the black-legged tick, is the primary vector in the eastern United States of several pathogens causing human diseases including Lyme disease, anaplasmosis, and babesiosis. Over the past two decades, I. scapularis-borne diseases have increased in incidence as well as geographic distribution. Lyme disease exists in two major foci in the United States, one encompassing northeastern states and the other in the Upper Midwest. Minnesota represents a state with an appreciable increase in counties reporting I. scapularis-borne illnesses, suggesting geographic expansion of vector populations in recent years. Recent tick distribution records support this assumption. Here, we used those records to create a fine resolution, subcounty-level distribution model for I. scapularis using variable response curves in addition to tests of variable importance. The model identified 19% of Minnesota as potentially suitable for establishment of the tick and indicated with high accuracy (AUC = 0.863) that the distribution is driven by land cover type, summer precipitation, maximum summer temperatures, and annual temperature variation. We provide updated records of established populations near the northwestern species range limit and present a model that increases our understanding of the potential distribution of I. scapularis in Minnesota.

Climate-based model predicting acarological risk of encountering the human-biting adult life stage of Dermacentor andersoni (Acari: Ixodidae) in a key habitat type in Colorado.

We exploited an elevation (climate) gradient ranging from 1,700 to 2,500 m in Poudre Canyon of Larimer County, CO, to determine climatic correlates of abundance per 15-s drag sampling time unit (hereafter referred to as abundance) of the human-biting adult life stage of the Rocky Mountain wood tick, Dermacentor andersoni Stiles (Acari: Ixodidae), in a key risk habitat for tick exposure: south/west-facing, rocky hillsides with mixed grass-brush-conifer vegetation. The relationship between elevation and abundance was parabolic, with peak tick abundances occurring at mid-range elevations (2,200-2,400 m) and tick abundances approaching zero at approximately 2,100 and 2,500 m. Regression modeling demonstrated that abundance of host-seeking adult ticks in south/west-facing exposures was accurately predicted by several climate variables related to temperature (e.g., mean annual minimum temperature, maximum temperature, and base 10 degrees C growing degree-days, and median length of annual freeze-free period; r2 values ranging from 0.771 to 0.864), whereas mean annual precipitation, snowfall, or relative humidity were uninformative in this respect (r2 values ranging from 0.020 to 0.316). Abundance of D. andersoni adults peaked at a mean annual maximum temperature of approximately 10 degrees C and a mean annual growing degree-day value of approximately 650. Relationships between climate variables and abundance of D. andersoni adults were used to create geographic information system (GIS)-based models for predicted tick abundance in south/west-facing exposures in Larimer County. This is the first GIS-based model developed for spatial patterns of abundance of D. andersoni. Finally, preliminary data from Poudre Canyon indicate a shift toward peak abundances of D. andersoni adults occurring in sheltered northern/eastern exposures, rather than in drier and hotter southern/ western exposures, at elevations below 2,100 m.

Refractoriness of the western fence lizard (Sceloporus occidentalis) to the Lyme disease group spirochete Borrelia bissettii.

The western fence lizard, Sceloporus occidentalis, is refractory to experimental infection with Borrelia burgdorferi sensu stricto, one of several Lyme disease spirochetes pathogenic for humans. Another member of the Lyme disease spirochete complex, Borrelia bissettii, is distributed widely throughout North America and a similar, if not identical, spirochete has been implicated as a human pathogen in southern Europe. To determine the susceptibility of S. occidentalis to B. bissettii, 6 naïve lizards were exposed to the feeding activities of Ixodes pacificus nymphs experimentally infected with this spirochete. None of the lizards developed spirochetemias detectable by polymerase chain reaction for up to 8 wk post-tick feeding, infected nymphs apparently lost their B. bissettii infections within 1-2 wk after engorgement, and xenodiagnostic L. pacificus larvae that co-fed alongside infected nymphs did not acquire and maintain spirochetes. In contrast, 3 of 4 naïve deer mice (Peromyscus maniculatus) exposed similarly to feeding by 1 or more B. bissettii-infected nymphs developed patent infections within 4 wk. These and previous findings suggest that the complement system of S. occidentalis typically destroys B. burgdorferi sensu lato spirochetes present in tissues of attached and feeding I. pacificus nymphs, thereby potentially reducing the probability of transmission of these bacteria to humans or other animals by the resultant adult ticks.

Geographical distribution patterns and habitat suitability models for presence of host-seeking ixodid ticks in dense woodlands of Mendocino County, California.

We used drag sampling to examine the geographical distribution patterns of ixodid ticks engaging in open (non-nidicolous) host-seeking behavior in dense woodland habitats of the climatically and ecologically diverse Mendocino County in north coastal California. The findings based on this sampling methodology reflect risk of human exposure to host-seeking ticks rather than the true distribution of the ticks. Drag sampling in 78 sites yielded 7,860 nymphal or adult Ixodes pacificus Cooley & Kohls, 220 Dermacentor occidentalis Marx, 150 Ixodes spinipalpis Hadwen & Nuttall, 15 Hemaphysalis leporispalustris (Packard), 12 Ixodes angustus Neumann, 12 Ixodes auritulus Neumann, and a single Dermacentor variabilis (Say). I. pacificus, which is the primary vector of the Lyme disease spirochete Borrelia burgdorferi to humans in California, occurred in all 78 sites examined. D. occidentalis, another tick species commonly biting humans in California, and H. leporispalustris typically were encountered in oak-associated woodlands in the central or eastern parts of the county. In contrast, three species of Ixodes ticks (I. angustus, I. auritulus, and I. spinipalpis) most commonly were found questing openly in woodlands with redwood present in the western part of the county. I. angustus and I. spinipalpis are occasional human biters and known experimental vectors of B. burgdorferi. Our study represents the first collection of large numbers of openly host-seeking I. spinipalpis ticks. Univariate tests of associations between presence of ticks (D. occidentalis, H. leporispalustris, I. angustus, I. auritulus, or I. spinipalpis) and environmental geographical information systems-remote sensing (GIS/ RS)-based data indicated that elevation, number of growing degree-days, and tasseled cap brightness, greenness, and wetness are especially useful predictors of presence of openly hostseeking ticks. Combinations of the above-mentioned GIS/RS-based data yielded significant logistic regression models for habitat suitability of host-seeking ticks for all five above-mentioned species. The model equations were used to create spatial surfaces of predicted presence of suitable habitat for openly host-seeking ticks in Mendocino County dense woodlands.

Acarologic risk of exposure to Borrelia burgdorferi spirochaetes: long-term evaluations in north-western California, with implications for Lyme borreliosis risk-assessment models.

Over a 5-year period (1997-2001) the population densities of Ixodes pacificus Cooley & Kohls (Acari: Ixodidae) nymphs infected with spirochaetes of Borrelia burgdorferi sensu lato (s.l.) were evaluated in areas of 2000 ha at two localities (CHR, nine sites; HREC, seven sites) 25 km apart in Mendocino County, north-western California. The 5-year median density of infected nymphs was significantly higher at CHR than at HREC (0.51 vs. 0.09 per 100 m(2) and site-specific yearly densities exceeding one infected nymph per 100 m2 were 10-fold more likely to occur at CHR than at HREC. The importance of long-term data in acarologic risk assessment was demonstrated by significantly higher median yearly densities of infected nymphs at CHR from 1997 to 1999, whereas both areas had similar densities during 2000-2001. Overall, the causative agent of Lyme borreliosis in North America, B. burgdorferi Johnson et al. sensu stricto (s.s.) accounted for 76% of 46 genetically characterized B. burgdorferi s.l. infections from I. pacificus nymphs. Tremendous variability in acarologic risk was recorded within both areas: yearly densities of infected nymphs varied 11-97-fold between sites at CHR and 8-30-fold at HREC. Part of this variation could be explained by environmental traits, most notably deer usage. However, correlations between environmental factors and density of infected nymphs (for CHR and HREC combined) did not necessarily apply when these areas were considered separately. Thus, a Lyme borreliosis ecology model developed in one of these areas needs testing in the other area.

Seasonal activity patterns of Ixodes pacificus nymphs in relation to climatic conditions.

In western North America, the tick Ixodes pacificus Cooley & Kohls (Acari: Ixodidae) is the primary vector to humans and domestic animals of the disease agents causing Lyme disease and granulocytic ehrlichiosis. We examined the seasonal activity patterns of I. pacificus nymphs over a 4-year period, including the wet and cold El Niño winter/spring of 1998, in a dry oak/madrone woodland, and for one year in a cooler and moister redwood/tanoak woodland in Mendocino County, California. Linear regressions were used to estimate when nymphal densities first exceeded and then fell below 25, 50 and 75% of the recorded yearly peak densities. In oak/madrone woodland, nymphs typically were active by mid-March, reached 50% of their yearly peak densities in early to mid-April, peaked by early May, fell below 50% of their peak densities by early to mid-June, and were absent by late July to mid-August. The lengths of the periods with nymphal densities exceeding 50 and 75% of the recorded yearly peaks in oak/madrone woodland were associated positively with rainfall and negatively with maximum air temperatures during April-May. Moreover, nymphal numbers typically reached 50% of their peak 10-15 days later, remained at levels above 50% of the peak 1.3-1.5 times longer, and started declining 4-6 weeks later under cooler, moister climatic conditions (oak/madrone woodland in 1998 and redwood/tanoak woodland in 2000) relative to warmer, drier conditions (oak/madrone woodland in 2000-2001). In oak/madrone woodland, nymphal densities typically started to decline when mean maximum daily air temperatures exceeded 23 degrees C. Nymphal densities were higher in dry oak/madrone relative to moist redwood/tanoak woodland from mid-March to late May 2000, similar in both habitat types in early June, but higher in redwood/tanoak woodland from late June onwards. We conclude that large-scale studies of the density of I. pacificus nymphs in California need to consider spatial variation in the length of nymphal activity periods and select temporal sampling regimens that yield representative data for all included habitat types.

Landscape features associated with infection by a malaria parasite (Plasmodium mexicanum) and the importance of multiple scale studies.

In a 3-year study, we examined landscape features (aspect, slope, sun exposure, canopy cover, type of ground cover, and nearest water source) that were potentially related to prevalence of infection with Plasmodium mexicanum in fence lizards (Sceloporus occidentalis) within a 4.5 ha study area in northern California, USA. Logistic regression analysis showed that ground cover type was the primary mediator of the probability of P. mexicanum infection. Infected lizards were captured more often in rock and/or leaf litter locations than in grassy ones. In another experiment, the study area was divided into 9 sites (0.07-0.33 ha), and infection prevalence was calculated for each. Three sites with high (> 30%) infection prevalence had significantly more rocky outcrops and leaf litter than those with low (< 20%) or moderate (20-30%) infection prevalence (N = 3 sites each). We conclude that lizard site selection may influence the probability of exposure to infected vectors and thus the likelihood of P. mexicanum infection. We also demonstrate that studies at different spatial scales may be required to understand fully the relationship between landscape features and parasite distribution.

Prevalence and abundance of Ixodes pacificus immatures (Acari: Ixodidae) infesting western fence lizards (Sceloporus occidentalis) in northern California: temporal trends and environmental correlates.

The prevalence and abundance of immature Ixodes pacificus ticks on western fence lizards (Sceloporus occidentalis) were examined in relation to time of year, host attributes (i.e., age, gender, and presence or absence of blood parasites), and 5 environmental characteristics, including topographic exposure and ground cover substrate, over a 2-year period in northern California. Lizards were infested with subadult ticks from early March until late July or early August, with peak median numbers of larvae and nymphs recorded in late April and early May of both years. Peak larval and nymphal abundances differed between years. The overall ratio of larvae to nymphs on adult male lizards was low, ranging from 0.80 in 1999 to 2.41 in 2000. Such intensive feeding of nymphs versus larvae on these lizards, which are reservoir-incompetent for Borrelia burgdorferi spirochetes, may explain previous observations of decreasing spirochetal infection prevalence from the nymphal to adult stage in northwestern California. Adult male lizards were more likely to be infested with nymphs and harbored greater abundances of larvae and nymphs than adult females. Lizards uninfected with blood parasites had more nymphs than infected lizards. The measured environmental characteristics could explain only a small percentage of the total variation observed in larval prevalence (22%) and in larval and nymphal abundance (12 and 3%, respectively).

Life history of a malaria parasite (Plasmodium mexicanum) in its host, the western fence lizard (Sceloporus occidentalis): host testosterone as a source of seasonal and among-host variation?

The course of infection of a malaria parasite (Plasmodium mexicanum) is highly variable in its host, the fence lizard (Sceloporus occidentalis). However, a seasonal trend is superimposed on this variation such that gametocyte production is intensified during mid- to late summer. Host testosterone levels follow a similar seasonal fluctuation and are variable among individual lizards. We sought to determine if testosterone levels affect seasonal and among-host variation in 11 P. mexicanum life history traits: rate of increase in level of infection (3 measures), peak parasitemia (3 measures), duration of increase (3 measures), time to detectable infection, and timing of production of gametocytes. We followed the course of infection in 125 male S. occidentalis, each randomly assigned to 1 of 4 treatment groups: castrated, castrated and implanted with exogenous testosterone, sham implanted, and unmanipulated controls. Median values for the 11 life history traits did not differ among treatment groups, and variances were homogeneous among the treatment groups for 10/11 traits. However, elevated testosterone significantly reduced the variation in timing of the onset of gametocyte production. Therefore, testosterone does not appear to be a primary regulator of P. mexicanum life history, yet testosterone may have some effect on when gametocytes first become detectable.

Life history of a malaria parasite (Plasmodium mexicanum): independent traits and basis for variation.

Plasmodium mexicanum, a malaria parasite of lizards, exhibits substantial variation among infections in the life-history traits which define its blood-dwelling stages. Such variation in life histories among infections is common in Plasmodium and may influence the ecology and evolution of the parasite's transmission success and virulence. Insight into these issues requires identification of independent traits (some traits may be bound by developmental trade-offs) and the importance of genetic versus host effects producing the variation. We studied 11 life-history traits in 120 induced infections of P. mexicanum in its natural lizard host (20 each from six donor infections). The traits varied among infections and fell into three clusters: rate/peak (rate of increase and peak parasitaemia of asexuals and gametocytes), time (duration of pre-patent period and the infection's growth) and maturity (timing of first gametocytes). Thus, few life-history traits define an infection in the lizard's blood. Donor effects were significant for ten traits and two trait clusters (maturity was the exception) suggesting genetic differences among infections may influence the rate of increase and peak parasitaemia, but not the timing of the first production of gametocytes.

Abundance of ticks (Acari: Ixodidae) infesting the western fence lizard, Sceloporus occidentalis, in relation to environmental factors.

We examined the impact of environmental characteristics, such as habitat type, topographic exposure and presence of leaf litter, on the abundance of Ixodes pacificus ticks infesting the western fence lizard (Sceloporus occidentalis) at the University of California Hopland Research and Extension Center (HREC), Mendocino County, California. A total of 383 adult lizards were slip-noosed and examined for tick infestation in April and May 1998. At least 94% of the lizards were infested by ticks and at least 20% of the females and 33% of the males carried > 15 ticks. This intensive utilization of western fence lizards (which do not serve as natural reservoirs for Lyme disease spirochetes) by subadult ticks, is probably the primary reason for the low prevalence of infection with Borrelia burgdorferi in I. pacificus nymphs and adults previously recorded at the HREC. Tick loads were higher on male than female lizards. Also, male lizards were generally more heavily infested in late April than in late May. The prevalence of tick infestation exceeded 88% in all habitat types but males collected in woodland and grass/woodland edges had higher tick loads than those collected in open grassland. Male lizards captured in open, exposed grassland tended to carry heavier tick loads in northern/eastern, as compared to southern/western, exposures, and when leaf litter was present.