Effects of reduced deer density on the abundance of Ixodes scapularis (Acari: Ixodidae) and Lyme disease incidence in a northern New Jersey endemic area

We monitored the abundance of Ixodes scapularis Say (Acari: Ixodidae) and the Lyme disease incidence rate after the incremental removal of white-tailed deer, Odocoileus virginianus Zimmermann, within a suburban residential area to determine whether there was a measurable decrease in the abundance of ticks due to deer removal and whether the reduction in ticks resulted in a reduction in the incidence rate within the human population. After three seasons, the estimated deer population was reduced by 46.7%, from the 2002 postfawning estimate of 2,899 deer (45.6 deer per km2) to a 2005 estimate of 1,540 deer (24.3 deer per km2). There was no apparent effect of the deer culling program on numbers of questing I. scapularis subadults in the culling areas, and the overall numbers of host-seeking ticks in the culling areas seemed to increase in the second year of the program. The Lyme disease incidence rate generated by both passive and active surveillance systems showed no clear trend among years, and it did not seem to vary with declining deer density. Given the resources required to mount and maintain a community-based program of sufficient magnitude to effectively reduce vector tick density in ecologically open situations where there are few impediments to deer movement, it may be that deer reduction, although serving other community goals, is unlikely to be a primary means of tick control by itself. However, in concert with other tick control interventions, such programs may provide one aspect of a successful community effort to reduce the abundance of vector ticks.     

Highly variable acquisition rates of Ixodes scapularis (Acari: Ixodidae) by birds on an Atlantic barrier island

Acquisition of ticks by bird hosts is a central process in the transmission cycles of many tick-borne zoonoses, but tick recruitment by birds has received little direct study. We documented acquisition of Ixodes scapularis Say on birds at Fire Island, NY, by removing ticks from mist-netted birds, and recording the number of ticks on birds recaptured within 4 d of release. Eight bird species acquired at least 0.8 ticks bird(-1) day(-1) during the seasonal peak for at least one age class of I. scapularis. Gray Catbirds, Eastern Towhees, Common Yellowthroats, and Northern Waterthrushes collectively accounted for 83% of all tick acquisitions; and six individuals apportioned among Black-billed Cuckoo, Gray Catbird, Eastern Towhee, and Common Yellowthroat were simultaneously infested with both larvae and nymphs. Bird species with the highest acquisition rates were generally ground foragers, whereas birds that did not acquire ticks in our samples generally foraged above the ground. Tick acquisition by birds did not differ between deciduous and coniferous forests. Among the 15 bird species with the highest recruitment rates, acquisition of nymphs was not correlated with acquisition of larvae. Tick acquisition rates by individual bird species were not correlated with the reservoir competence of those species for Lyme borreliae. However, birds with high tick acquisition rates can contribute large numbers of infected ticks, and thus help maintain the enzootic cycle, even if their levels of reservoir competence are relatively low.     

Ixodes scapularis ticks (Acari: Ixodidae) from Louisiana are competent to transmit Borrelia burgdorferi, the agent of Lyme borreliosis

The principal vector of Borrelia burgdorferi, the Lyme borreliosis spirochete, in the Northeast and Midwestern regions of the United States is the blacklegged tick Ixodes scapularis. Because of a favorable environment, I. scapularis is also plentiful in the South; however, a correlation with Lyme borreliosis cases does not exist in this region of the United States. Concern existed that something intrinsic to ticks found in Louisiana could mitigate their ability to transmit B. burgdorferi. Therefore, we set out to assess the ability of I. scapularis ticks from Louisiana to become infected with and transmit B. burgdorferi using mice as hosts. In the laboratory, mating adult female ticks collected in southeastern Louisiana were fed on the ears of rabbits. After oviposition and egg hatching, the resulting larvae were fed on mice that had been needle-inoculated with two different strains of B. burgdorferi sensu stricto, B31 and JD1. Larvae were found to be positive for spirochetes. Additional fed larvae were allowed to molt into the nymphal stage. Flat nymphs remained infected with B. burgdorferi. Infected nymphs were allowed to feed on na?ve mice, all of which became infected as shown by culture of ear biopsy specimens. Na?ve larvae were then fed on these same mice to assess transmissibility. The resulting engorged larvae harbored spirochetes. We have demonstrated that the I. scapularis ticks found in Louisiana are fully competent to carry and transmit B. burgdorferi infection.     

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.     

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.     

Feasibility of controlling Ixodes scapularis ticks (Acari: Ixodidae), the vector of Lyme disease, by parasitoid augmentation

A theoretical analysis of the feasibility of controlling tick populations (Ixodidae) by the release of reared Ixodiphagus parasitoids in tick ecosystems yielded promising results. The analysis suggested that if reasonable progress could be made in mass-rearing the parasitoids, it would be possible to control the blacklegged tick, Ixodes scapularis (Say), the vector of Lyme disease, by this biological control procedure. Lyme disease has become the most important vector-borne disease in the United States. In a field-release experiment conducted in Africa by members of the International Center for Insect Physiology and Ecology, effective control of Amblyomma variegatum (F.) was obtained by the release of Ixodiphagus parasitoids in tick habitats. Encouraging theoretical results along with the encouraging results of a field-release experiment indicate the need for civil and political leaders in countries where ticks are a major problem to sponsor strong and well-coordinated research initiatives focused on the development of this new method of dealing with tick problems.     

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.     

Populations of Ixodes scapularis (Acari: Ixodidae) are modulated by drought at a Lyme disease focus in Illinois

From 1990 through 1997, Ixodes scapularis Say larvae and nymphs were sampled between May and October along a 400-m segment of a nature trail in a Lyme disease endemic site in northern Illinois. Ticks were removed from Peromyscus leucopus mice and collected via tick drags at approximately 3-wk intervals. Mouse population estimates along the trail varied from 2, in the spring of 1996 following a year of drought, to > 200 in 1993, the wettest year on record. During the 8-yr period, there were major droughts during the summers of 1991 and 1995. Cumulative degree-days were positively correlated with the number of ticks collected on drags in the same year and negatively correlated with larval tick populations for the following year (P < 0.05). Cumulative rainfall was positively correlated with larval tick abundance for the following year. This was most readily apparent by examination of the larval density on captured mice. In the year following each of two drought years, larval densities were significantly depressed compared with the 8-yr average at the site.     

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.     

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.     

County-level surveillance of white-tailed deer infestation by Ixodes scapularis and Dermacentor albipictus (Acari: Ixodidae) along the Illinois River

From 1998 to 2003, 4,935 hunter-killed deer in northern and central Illinois were examined for ticks; 4,066 blacklegged ticks, Ixodes scapularis Say, and 6,530 winter ticks, Dermacentor albipictus (Packard) (Acari: Ixodidae), were collected. I. scapularis was the predominant tick species in the northern portion of the study area, with a decreasing north-to-south prevalence gradient. In contrast, D. albipictus was more common in the south with a decreasing south-to-north prevalence gradient. Compared with previous studies, the geographic range for both species expanded into the central portion of the Illinois River. Prevalence and intensity of both tick species were greater on bucks, and infested bucks were geographically more widespread than infested does and fawns. These findings indicate that blacklegged tick and winter tick distributions remain dynamic in the north central United States     

Attempt to control ticks (Acari: Ixodidae) on deer on an isolated island using ivermectin-treated corn

We report an attempt by an offshore island community to control the vector tick of Lyme disease by providing ivermectin-treated corn to an isolated herd of free-ranging white-tailed deer, Odocoileus virginianus Zimmerman. Medicated corn was supplied in troughs within the island village and from automatic feeders at remote sites during 5 consecutive fall and spring adult tick questing seasons. Acaricide consumption was monitored by assaying its presence in fresh deer pellets and its concentration in deer sera. Its effectiveness was evaluated by recording the number of adult ticks collected from the hides of deer, the number of females becoming sufficiently engorged to oviposit, and the success of subsequent oviposition and eclosion. Entomologic risk was monitored by collecting immature ticks from hosts and adult ticks from vegetation. Estimates based on a subsequent deer reduction program indicated that up to twice as many deer had been present during the project as originally presumed. For this and other reasons related to deer behavior, target levels of serum ivermectin were achieved in a minority of deer. Nevertheless, > 90% control of female tick infestation, subsequent oviposition, and larval eclosion was obtained in those 8 of 16 sampled deer with serum ivermectin levels of > or = 15 ng/ml. In addition, the ratio of females to males, the numbers of females engorging > 10 mg body weight, and the numbers of those eventually hatching, were all significantly less among ticks from island deer in comparison with ticks from untreated deer. No consistent changes in the numbers of ticks found on immature-stage hosts or removed from vegetation were noted within 3 yr of the cessation of treatment.     

Protection against lyme disease spirochete transmission provided by prompt removal of nymphal Ixodes scapularis (Acari: Ixodidae)

Public health recommendations for Lyme disease prevention generally include daily tick checks and prompt removal of attached ticks as a means of decreasing the risk of acquiring Lyme disease in highly endemic regions. In the current study, we determined whether crushing nymphal ticks during removal with forceps increased the risk of B. burgdorferi transmission, what degree of protection from transmission of B. burgdorferi was provided by removal of nymphal Ixodes scapularis Say at specific intervals, and whether commercial devices marketed for tick removal worked when tested against nymphal I. scapularis. Both removal via gentle pressure (26% transmission) or crushing the tick (30% transmission) caused a significant decrease in transmission as compared with the sham control (70% transmission). The degree of protection provided via tick removal decreased steadily up to 60 h of attachment; between 60 and 66 h, a dramatic falloff in protection occurred to the point where no protection was observed at 66 h. Finally, commercial tick removal devices varied widely in their efficacy for the removal of attached nymphal I. scapudaris.     

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.     

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.     

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.     

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.     

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.     

Distribution, density, and Lyme disease spirochete infection in Ixodes dammini (Acari: Ixodidae) on white-tailed deer in Maryland.

A Statewide survey of ticks parasitizing white-tailed deer was carried out in Maryland during November 1989 to assess the status of the deer tick, Ixodes dammini Spielman, Clifford, Piesman & Corwin, the major vector of Lyme disease in the northeastern United States. Ticks were collected from deer carcasses brought in by hunters at 23 check stations (one per county). A total of 3,437 I. dammini were collected from 538 of 1,281 deer (42%), together with 2,013 Dermacentor albipictus (Packard) and 23 Amblyomma americanum (L.) from 34 and 0.5% of deer respectively. I. dammini prevalence ranged from 0 to 79% of deer and mean abundance from 0 to 7.3 ticks per deer at different check stations. Lyme spirochete, Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner, infection rates in ticks ranged from 0 to 21%, with a mean of 8%. Deer-tick density and spirochete infection rates varied with physiographic region and were low in the Appalachian, intermediate in the Piedmont, and high in the Western and Eastern Coastal Plains regions. County-based human case rates correlated positively with I. dammini abundance. We concluded that I. dammini was well established except in the mountainous western region of Maryland and was involved in Lyme disease transmission.