Deer density and the abundance of Ixodes scapularis (Acari: Ixodidae)

The abundance of Ixodes scapularis Say (Ixodes dammini Spielman, Clifford, Piesman & Corwin), the vector tick of the Lyme disease spirochete and other human pathogens, is related to the presence of its primary reproductive stage host, white-tailed deer (Odocoileus virginianus Zimmerman). However, this relationship has not been quantified in terms that would guide wildlife management in areas in which the public is, or is likely to become, exposed to infected ticks. In this study, deer density and tick abundance were measured in an emergent area for Lyme disease at three spatial scales using estimation methods appropriate for each. Simple linear regression was used to relate (1) the number of ticks found on deer at tagging stations in southern Maine to harvest-derived estimates of the density of deer within the towns in which they were killed, (2) tick densities estimated from fall flagging counts to deer densities estimated from pellet group counts made within multiple transects distributed through 5.2-km2 study sites, and (3) tick counts to pellet group counts within the individual transects. At the broadest scale, ticks on deer decreased with elevation and distance from the coast and increased with deer density, although deer and tick presence were only weakly related. Among the 5.2-km2 study sites and within individual transects, tick abundance related more strongly to deer pellet group counts. Few ticks were collected at deer densities <7/km2.     

Influence of deer abundance on the abundance of questing adult Ixodes scapularis (Acari: Ixodidae).

Nymphal and adult Ixodes scapularis Say were sampled by flagging at 2 sites on a barrier island, Fire Island, NY, and at 2 sites on the nearby mainland. Nymphal densities did not differ consistently between island and mainland sites, but adult densities were consistently lower on the island. We tested whether lower adult densities on the island resulted from greater nymphal mortality on the island than the mainland, or whether adult ticks on the island were poorly sampled by flagging because they had attached abundantly to deer, which were common on Fire Island. Differential nymphal mortality on islands versus mainland did not explain this difference in adult densities because survival of flat and engorged nymphs in enclosures was the same at island and mainland sites. Ticks were infected by parasitic wasps on the island and not the mainland, but the infection rate (4.3%) was too low to explain the difference in adult tick densities. In contrast, exclusion of deer by game fencing on Fire Island resulted in markedly increased numbers of adult ticks in flagging samples inside compared with samples taken outside the exclosures. Therefore, the scarcity of adult ticks in flagging samples on Fire Island resulted, at least in part, from the ticks being unavailable to flagging samples because they were on deer hosts. Differences in the densities of flagged ticks inside and outside the exclosures were used to estimate the percentage of questing adults on Fire Island that found deer hosts, excluding those that attached to other host species. Approximately 56% of these questing adult ticks found deer hosts in 1995 and 50% found deer hosts in 1996. Therefore, in areas where vertebrate hosts are highly abundant, large proportions of the questing tick population can find hosts. Moreover, comparisons of tick densities at different sites by flagging can be potentially biased by differences in host densities among sites.     

Pathogenicity of Bacillus thuringiensis variety kurstaki to Ixodes scapularis (Acari: Ixodidae)

Pathogenicity of the entomopathogenic bacterium Bacillus thuringiensis variety kurstaki de Barjac & Lemille was tested against the black-legged tick, Ixodes scapularis Say. Engorged larvae dipped in a solution of 10(8) spores per milliliter showed 96% mortality 3 wk after infection. The LC50 value for engorged larve (concentration required to kill 50% of ticks) was 10(7) spores per milliliter. B. thuringiensis shows considerable potential as a microbial control agent for the management of I. scapularis.     

Ixodes scapularis (Acari: Ixodidae) distribution surveys in the Chicago metropolitan region

Considering recent studies confirming an increased risk of contracting Lyme disease near metropolitan Chicago, we surveyed a more comprehensive area to assess whether the geographical distribution and establishment of Ixodes scapularis (Say) populations across northeast Illinois are widespread or limited in occurrence. From May through October 2008 and from April through October 2009, 602 I. scapularis ticks of all three life stages (larva, nymph, adult) were collected from sites in Cook, DuPage, Lake, and McHenry counties in northeast Illinois. The surveys were conducted by drag sampling vegetation in public-access forested areas. I. scapularis comprised 56.4% of ticks collected (n = 1,067) at 17 of 32 survey sites. In addition, four other tick species were incidentally collected: Dermacentor variabilis (Say), Haemaphysalis leporispalustris (Packard), Ixodes dentatus (Marx), and Amblyomma americanum (L.). This study updates the I. scapularis distribution in northeast Illinois. Our random sampling of suitable tick habitats across a large geographic area of the Chicago metropolitan area suggests a widespread human exposure to I. scapularis, and, potentially, to their associated pathogens throughout the region. These results prompt continued monitoring and investigation of the distribution, emergence, and expansion of I. scapularis populations and Borrelia burgdorferi transmission within this heavily populated region of Illinois.     

Ixodes scapularis (Acari: Ixodidae) in Kansas: a new distribution record.

The blacklegged tick, Ixodes scapularis Say, previously known to occur only in the extreme southeastern corner of Kansas, has been collected in Douglas and Jefferson counties in the northeast. The new collections extend the northern range of this species in Kansas by approximately 240 km. The role of I. scapularis as a vector of the Lyme borreliosis spirochete, Borrelia burgdorferi, makes this a noteworthy extension of its known range. The proximity of these new collection sites to large centers of human population is of special significance to health-care providers in that region.     

Ixodes scapularis (Acari: Ixodidae): abundance and rate of infection with Borrelia burgdorferi in four state parks in Wisconsin

Four state parks located in Lyme disease endemic regions of Wisconsin were surveyed for the presence of Ixodes scapularis Say during May and June of 1998 by drag sampling along hiking trails. Nymphal abundance varied between parks, with the average number of nymphs encountered in 1 h ranging from 6.2 +/- 3.8-47.1 +/- 36.3 (mean +/- SD). Questing nymphs were tested for the presence of Borrelia burgdorferi by culture in BSK medium and 7-12% was found to be infected. The average risk of encountering an infected nymph (entomologic risk index) ranged from 0.5 to 5.2 infected nymphs per hour. The highest entomological risk index was recorded from a small island park in northwestern Wisconsin during the last week in May (8.0 infected nymphs per hour). These results indicate a lower risk for human Lyme disease exposure in Wisconsin state parks in comparison with highly endemic areas of the northeastern United States.     

Lack of spatial autocorrelation in fine-scale distributions of Ixodes scapularis (Acari: Ixodidae)

Spatial patterns of Ixodes scapularis Say, the vector of the Lyme disease agent, have been examined at various geographic scales, demonstrating that distributions of these ticks are spatially autocorrelated at both national and state scales. We tested the hypothesis that distributions of nymphal I scapularis ticks at the fine scale of an endemic community also are spatially autocorrelated. Nymphal tick densities were determined by collecting ticks from 51 and 47 wooded residential properties in a southern Rhode Island town in 2002 and 2003, respectively. The average tick density at residences during 2002 was 51.17 ( +/- 46.04) nymphs per hour, with a range of 3-297 and median of 40.82. In 2003, the average tick density was 44.48 (+/- 38.31) nymphs per hour, with a range of 3-153 and median of 36. Semivariance analysis revealed no spatial autocorrelation in tick densities between residences, likely due to the high variability of tick distributions at this scale. Further analysis of drag-sampling data at individual residences by using Lloyd's patchiness index (m*/m) demonstrated a patchy distribution of nymphs. High variability of nymphal I. scapularis densities may greatly affect predicting spatial patterns of ticks at a fine scale.     

A technique for longitudinally sampling individual adult Ixodes scapularis (Acari: Ixodidae)

The first technique for repeatedly sampling individual Ixodes scapularis adult ticks was developed and validated. Gut samples from the same individual ticks were removed and analyzed at weekly intervals. Multiple analyses were conducted on each gut sample (e.g., total protein concentration, presence of viable B. burgdorferi spirochetes, and concentration of outer surface protein A [OspA]). Female I. scapularis survived for up to 25 d after gut sampling. Seventy-five percent of females oviposited after the sampling procedure, with 14% of ticks laying >1,500 eggs. No significant differences in either fecundity or length of survival were found between B. burgdorferi-infected and uninfected I. scapularis. This technique will enable longitudinal studies on both tick-pathogen interactions and physiological studies that have hitherto not been attempted.     

Seasonal activity and host associations of Ixodes scapularis (Acari: Ixodidae) in southeastern Missouri

Based on tick collections recovered from wild vertebrates and by dragging, the seasonal occurrence of adult blacklegged ticks, Ixodes scapularis Say, extended from October through May in southeastern Missouri. Adult activity was bimodal with the higher peak occurring in November followed by a lower peak in February. The activity of immature I. scapularis had the general pattern of that found in the Northeast where Lyme disease is hyperendemic, with larval activity (July) peaking after that of nymphs (May and June). Vertebrates varied in their importance as hosts of I. scapularis. White-tailed deer, Odocoileus virginanus (Zimmerman), and coyotes, Canis latrans Say, were the primary hosts of adult I. scapularis. Broad-headed skinks, Eumeces laticeps (Schneider), and eastern fence lizards, Sceloporus undulatus (Latreille), were the primary hosts of nymphal I. scapularis. The broad-headed skink, 5-lined skink, Eumeces fasciatus (L.), and Carolina wren, Thryothorus ludovicianus (Latham), were the primary hosts of larval I. scapularis. Homeotherms were important hosts of immature I. scapularis, accounting for 30% of nymphs and 39% of larvae collected. The eastern cottontail rabbit, Sylvilagus floridanus (Allen), may play an important role in the epidemiology of Lyme disease in Missouri. Isolates of Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner were made from ticks recovered from rabbits, making the cottontail rabbit a key species for further study of the epidemiology of Lyme borreliosis in Missouri.     

Immune response of lizards and rodents to larval Ixodes scapularis (Acari: Ixodidae).

The house mouse (laboratory strain), Mus musculus (L.), the cotton mouse, Peromyscus gossypinus (LeConte), the broad-headed skink, Eumeces laticeps (Schneider), and the guinea pig, Cavia porcellus (L.), were successively infested five times with larvae of the blacklegged tick, Ixodes scapularis Say. Tick feeding success, engorgement weight, and subsequent molting success were measured after each infestation. A greater percentage of ticks (P less than 0.05) fed on M. musculus and E. laticeps than on P. gossypinus or C. porcellus. P. gossypinus expressed a transitory partial resistance, measured in percentage of ticks feeding, during the third infestation but showed increased tolerance during the fourth and fifth infestations. Ticks fed on E. laticeps were heavier than those fed on any other host (P less than 0.05). Those fed on M. musculus were heavier than those fed on P. gossypinus, but the difference was not statistically significant. On C. porcellus, only 1.6% of larvae from the third infestation and none thereafter engorged; weights of larvae from first and second infestations were higher (P less than 0.05) than those fed on M. musculus and P. gossypinus and lower (P less than 0.05) than those fed on E. laticeps. A greater percentage of larvae from E. latticeps and M. musculus (P less than 0.05) molted to nymphs compared with those from P. gossypinus and C. porcellus. Molting success was the same for ticks fed on P. gossypinus and on C. porcellus during the first and second infestations. M. musculus, P. gossypinus, and E. laticeps expressed no resistance (measured as percentage feeding, engorgement weight, and molting) to feeding by I. scapularis larvae after five infestations. Host serum was tested using an ELISA for detection of antibodies against I. scapularis salivary gland antigens before and after each infestation. Antibodies were detected after the second infestation and thereafter in C. porcellus, the only species to show antibodies or to express and maintain an acquired resistance to tick feeding throughout five infestations.     

Spatiotemporal patterns of host-seeking Ixodes scapularis nymphs (Acari: Ixodidae) in the United States

The risk of Lyme disease for humans in the eastern United States is dependent on the density of host-seeking Ixodes scapularis Say nymphal stage ticks infected with Borrelia burgdorferi. Although many local and regional studies have estimated Lyme disease risk using these parameters, this is the first large-scale study using a standardized methodology. Density of host-seeking I. scapularis nymphs was measured by drag sampling of closed canopy deciduous forest habitats in 95 locations spaced among 2 degrees quadrants covering the entire United States east of the 100th meridian. Sampling was done in five standardized transects at each site and repeated three to six times during the summer of 2004. The total number of adults and nymphs of the seven tick species collected was 17,972, with 1,405 nymphal I. scapularis collected in 31 of the 95 sites. Peak global spatial autocorrelation values were found at the smallest lag distance (300 km) and decreased significantly after 1,000 km. Local auto-correlation statistics identified two significant high-density clusters around endemic areas in the northeast and upper Midwest and a low-density cluster in sites south of the 39th parallel, where only 21 nymphs were collected. Peak nymphal host-seeking density occurred earlier in the southern than in the most northern sites. Spatiotemporal density patterns will be combined with Borrelia prevalence data as part of a 4-yr survey to generate a nationwide spatial risk model for I. scapularis-borne Borrelia, which will improve targeting of disease prevention efforts.     

Direct detection of Borrelia burgdorferi in Ixodes scapularis (Acari: Ixodidae) nymphs by hybridization to ribosomal RNA

A method for direct detection of Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner has been developed. Cells are lysed to facilitate release of ribosomal RNA. Lysates are filtered onto nylon membranes that are hybridized with probes specific for sequences in B. burgdorferi 23S rRNA. The technique is rapid and does not require any enzymatic amplification steps. With the use of a cocktail containing five different probes, approximately 1,000 organisms could be detected. The assay was successfully applied to direct detection of B. burgdorferi in Ixodes scapularis Say nymphs.     

Microclimate and habitat in relation to Ixodes scapularis (Acari: Ixodidae) populations on Long Point, Ontario, Canada.

The impact of microclimate and density of hosts for adult ticks on the density of Ixodes scapularis Say was evaluated within 4 habitats on Long Point, Ontario, from 1989-1992. During the period from May to September, mean weekly vapor pressure deficits were greater within the oak savannah and cottonwood dune habitats than at the maple forest and white pine habitats, which were similar. Vapor pressure deficit was likely the major factor affecting the survivorship of eggs and immature tricks in these habitats. Based on drag sampling, I. scapularis adults demonstrated peak activity in April and October of each year. The mean number of I. scapularis adults collected by dragging during the fall or in the spring did not differ significantly within each habitat. The mean number of adults collected also did not differ among tick cohorts within each habitat; however, significantly more adults were collected within the maple forest than in the white pine habitat. The mean number of I. scapularis adults per white-tailed deer, Odocoileus virginianus (Zimmerman), increased from 1989 to 1991 and then decreased in 1992. Significantly more adult I. scapularis infested deer were observed in 1990 than in 1989. Removal of deer in 1989 and 1990 resulted in a calculated decrease of > 100,000 fed female ticks. Although seasonal variation in microclimate within habitats was closely linked with tick survival and partly explains the differences in abundance of I. scapularis among habitats on Long Point, habitat utilization by deer was also a primary factor governing the local abundance of I. scapularis populations.     

Meteorologically mediated diurnal questing of Ixodes scapularis and Amblyomma americanum (Acari: Ixodidae) nymphs

Questing Ixodes scapularis Say and Amblyomma americanum (L.) nymphs were collected in the field by drag sampling to determine whether ticks would be collected in greater numbers during certain times of the day and under certain ambient meteorological conditions. Ticks were collected hourly for 16 h on 4 d and counts were contrasted with simultaneous measurements of ambient temperature and relative humidity, and with similar measurements made within the leaf litter. Peak numbers of questing I. scapularis and A. americanum nymphs were collected at distinctly different times of day, suggesting that the two species were responding to different environmental conditions. Both species demonstrated responses to diel changes in local meteorological conditions. Numbers of ticks were correlated with ambient temperature and humidity and with conditions recorded in the leaf litter that appeared to play a significant role in mediating questing behavior of both species. The time of day when tick sampling is conducted might significantly bias population estimates for sympatric species and the assessment of tick-borne disease transmission risk.     

Reduced abundance of Ixodes scapularis (Acari: Ixodidae) and the tick parasitoid Ixodiphagus hookeri (Hymenoptera: Encyrtidae) with reduction of white-tailed deer

The principal vector for the pathogens of Lyme disease, human granulocytic ehrlichiosis, and human babesiosis is the tick Ixodes scapularis Say. A chalcid wasp, Ixodiphagus hookeri, in the family Encyrtidae parasitizes populations of the tick on several islands or other geographically isolated sites in New England with high densities of these ticks and white-tailed deer (Odocoileus virginianus), the principal host for adult I. scapularis. Deer densities were reduced at a forested tract in Bridgeport and the Bluff Point Coastal Reserve in Groton, Connecticut, from levels exceeding 90 animals per km2 in 1992 (Bridgeport) and 1994 (Bluff Point) to 17 and 10 animals per km2, respectively, by fall 2001. Tick densities declined with sustained reductions in the population of white-tailed deer. Similarly, prevalence of tick parasitism by Ixodes hookeri declined at both sites from 30 to 25% to <1.0% and was significantly correlated with previous year's deer density at both sites (r(s) = 0.933 and r(s) = 0.867, P < or = 0.0001) and with nymphal tick densities at Bridgeport (r(s) = 0.867, P < or = 0.0001), but was not as well correlated with tick densities in Groton. The virtual disappearance of I. hookeri in this study corresponds with a lack of I. hookeri in mainland I. scapularis at comparable deer and tick densities, suggesting that there is a threshold deer density of approximatley 10-20/km2, with corresponding tick densities necessary for I. hookeri to successfully parasitize I. scapularis.     

Defining plasmids required by Borrelia burgdorferi for colonization of tick vector Ixodes scapularis (Acari: Ixodidae)

Maintenance in nature of Borrelia burgdorferi, the pathogenic bacterium that causes Lyme disease, requires transmission through an infectious cycle that includes a tick vector and a mammalian host. The genetic requirements for persistence in these disparate environments have not been well defined. B. burgdorferi has a complex genome composed of a chromosome and >20 plasmids. Previous work has demonstrated that B. burgdorferi requires two plasmids, lp25 and lp28-1, in the mammalian host. To investigate the requirement for these same two plasmids during tick infection, we experimentally infected larval ticks with B. burgdorferi lacking either lp25 or lp28-1 and then assessed the spirochete load in ticks at different points of the infection. Whereas plasmid lp28-1 was dispensable in ticks, plasmid lp25 was essential for tick infection. Furthermore, we investigated the requirement in ticks for the lp25 gene bbe22, which encodes a nicotinamidase that is necessary and sufficient for mammalian infection by B. burgdorferi clones lacking lp25. This gene was also sufficient in ticks to restore survival of spirochetes lacking lp25. This is the first study to investigate the requirement for specific plasmids by B. burgdorferi within the tick vector, and it begins to establish the genomic components required for persistence of this pathogen throughout its natural infectious cycle.     

Responses of adult Ixodes scapularis (Acari: Ixodidae) to urine produced by white-tailed deer of various reproductive conditions

The responses of adult female blacklegged ticks, Ixodes scapularis Say, to urine from white-tailed deer, Odocoileus virginianus (Zimmermann), belonging to 4 reproductive categories (doe in estrous, doe out of season, reproductive [dominant] buck, young buck) and to a mixture of urine from nondominant bucks in rut, young bucks out of rut, and nonestrous does were studied in laboratory behavioral bioassays. In high humidity (approximately 95% RH) in a glove box there were no statistically significant arrestment responses to any of the 5 types of urine, but an avoidance response was observed to urine from dominant reproductive bucks. When ticks were tested at approximately 50% RH, with samples of all 5 types of urine in the glove box, significant arrestant responses by the ticks were elicited by urine from does in estrous and by dominant reproductive bucks. When tested without other types of urine in the glove box, the urine mixture elicited an arrestant response at 50% RH. In some circumstances, adult I. scapularis may possibly use deer urine as a chemical cue in selecting host-ambush sites.     

Efficacy of a permethrin-based acaricide to reduce the abundance of Ixodes dammini (Acari: Ixodidae)

Permethrin-impregnated cotton was distributed to reduce abundance of immature Ixodes dammini Spielman, Clifford, Piesman & Corwin feeding upon white-footed mice (Peromyscus leucopus) and questing on vegetation at a private resort site (The Crane Reservation of The Trustees of Reservations in Ipswich, Mass.) located in coastal New England. This test constituted the first independent evaluation of the efficacy of this commercial product (Damminix). Over a 3-yr period, 2,000 applicator tubes containing treated cotton were distributed over the 7.3-ha site in 5 regularly scheduled applications. Mice removed treated fiber from the tubes and transported it to their nests. Within 3 wk of the first application, virtually all mice in the treated site were rendered free of ectoparasites. After the first season of application, no nymphal host-seeking I. dammini could be found on vegetation. Visitor and employee complaints about deer tick bites or ticks found on skin and clothing had been attributed to the site before treatment, but not thereafter. We confirmed the efficacy of Damminix for reduction of the abundance of vector ticks and thereby contributed to the protection of humans against Lyme disease at this site.     

Abundance of Ixodes scapularis (Acari: Ixodidae) larvae and nymphs in relation to host density and habitat on Long Point, Ontario.

Ixodes scapularis Say populations were evaluated within 4 habitats on Long Point, Ontario, from 1990 to 1992 to ascertain whether differences in density of mouse populations within and among habitats were correlated with that of immature I. scapularis populations. I. scapularis immatures were rarely collected by dragging within the cottonwood dune habitat. Significantly more larvae (P < or = 0.05) were collected by drag sampling within the maple forest habitat than in the oak savannah or white pine habitats for the 1989, 1990, and 1991 cohorts, whereas the size of the 1992 larval cohort did not differ significantly among these habitats. Significantly more nymphs were collected by dragging within the maple forest than in the other 2 habitats for all 4 cohorts. Nymphs from the 1989 and 1990 cohort were more abundant within the oak savannah than the white pine habitat, whereas the 1991 and 1992 cohorts were similar. With few exceptions, I. scapularis immatures were most prevalent on white-footed mice, Peromyscus leucopus (Rafinesque), captured within the maple forest > oak savannah > white pine > cottonwood dune, although differences were not significant in all years and in all habitats. The number of mice captured within the 4 habitats was not correlated with the number of I. scapularis larvae or nymphs infesting them. Likewise, the minimum number of mice alive was not significantly correlated with conversion indices of larvae to nymphs or nymphs to adults. Lack of association between mouse availability and relative size of subsequent cohorts of host-seeking ticks suggests that factors other than the size of the mouse populations were responsible for the observed differences in tick abundance among habitats.     

Ixodes dammini (Acari: Ixodidae) in Indiana

Ixodes dammini Spielman, Clifford, Piesman and Corwin is reported from Indiana for the first time. Four specimens were taken from deer and one from a dog. All specimens are adult females and all were collected in 1987. The significance of this finding is discussed.