Distribution and abundance of Ixodes scapularis (Acari: Ixodidae) in North America: ecological processes and spatial analysis

Conditions influencing the distribution and abundance of Ixodes scapularis Say in North America have been the object of considerable study during the past 2 decades, resulting in new information about its expanding range, population structure, host associations, and environmental constraints. Variation in the methods, timing, or extent of sampling, the type of population estimate developed, and the spatial scale of analysis makes comparisons of results difficult. This report summarizes research on I. scapularis habitat, including the diversity of hosts and physical conditions in the environment where they are abundant, and how this may influence dispersal and range expansion. Recent efforts to analytically address spatial distributions are considered. Approaches that may be useful in clarifying and resolving unanswered questions are suggested, and a conceptual framework for analyzing this vector's expanding abundance and distribution is offered.     

Determining the duration of Ixodes scapularis (Acari: Ixodidae) attachment to tick-bite victims

The duration of tick attachment is one factor associated with risk for human infection caused by several tick-borne pathogens. We measured tick engorgement indices at known time intervals after tick attachment and used these indices to determine the length of time that ticks were attached to tick-bite victims in selected Rhode Island and Pennsylvania communities where the agents of Lyme disease and human babesiosis occur. The total body length and width as well as the length and width of the scutum were measured on nymphal and adult female Ixodes scapularis Say removed from laboratory animals at 0, 12, 24, 36, 48, 60, and 72 h after their attachment. Three engorgement indices were calculated at each time interval. In addition, engorgement indices measurements were recorded for 504 ticks submitted to a commercial laboratory for pathogen detection testing between 1990 and 1992. No detectable change was observed in the average engorgement indices for either nymphal or adult ticks between 0 and 24 h of attachment using any of the engorgement indices. After 24 h of tick attachment, all engorgement indices continuously increased: average indices for nymphs attached 36, 48, and 60 h were significantly different from those attached < or = 24 h and from each other. Similarly, average engorgement indices for adult ticks attached < or = 36 h were significantly different from those attached for 48 h or more. More than 60% of tick-bite victims removed adult ticks by 36 h of attachment, but only 10% found and removed the smaller nymphal ticks within the first 24 h of tick feeding. The duration of tick attachment may serve as a useful predictor of risk for acquiring various infections, such as Lyme disease and babesiosis, transmitted by I. scapularis. Regression equations developed herein correlate tick engorgement indices with duration of feeding. A table containing specific engorgement index prediction intervals calculated for both nymphs and adults will allow the practitioner or clinical laboratory to use easily measured tick engorgement indices to predict transmission risk by determining the duration of feeding by individual ticks.     

Impact of controlled burns on the abundance of Ixodes scapularis (Acari: Ixodidae)

Information on the effect of vegetative destruction by controlled burns in reducing the abundance of the blacklegged tick, Ixodes scapularis Say, the vector for the agents of Lyme disease, human babesiosis, and human granulocytic ehrlichiosis, is limited. Therefore, the abundance of nymphal, larval, and adult I. scapularis was monitored by dragging the vegetation at 2 separate 4-ha tracts in Cockaponset State Forest in Connecticut following a single controlled burn on 15 April or 21 May 1992. The burn at the April burn site was rated as light to moderate with a flame height of 0.3 m and consumed approximately 67% of the surface leaf litter. The burn at the May burn site was rated moderate to severe with a flame height of 0.6-0.9 m., which consumed vegetation < 5 cm in diameter and approximately 100% of the surface leaf litter. The impact of the burn was strongly influenced by the intensity and timing of the burn. Burning of the vegetation resulted in a reduction of the abundance of nymphal I. scapularis by 74% at the moderately burned site and 97% at the severely burned site, compared with adjacent unburned woodland. No larvae were recovered later in the summer from the severely burned tract. However, judging by the comparable abundance of adult I. scapularis in the fall at the burned and unburned woodlands, the effect of the burns was temporary. Burning the vegetation for the control of I. scapularis appears limited in effect and could be applied only on a large scale in areas with little or no human habitations.     

Experimental transmission of Powassan virus (Flaviviridae) by Ixodes scapularis ticks (Acari:Ixodidae)

Systematic inflammatory response syndrome, SIRS, is one of the biological responses to surgical and anesthetic invasions. SIRS itself is a defense response, but excessive SIRS may result in post-operative failure of major organs. There is evidence that inflammatory cytokines play an important role in SIRS. Therefore it may be treated by regulating these inflammatory cytokines. Changes in inflammatory cytokines during the perisurgical period were discussed in relation to post-operative organ injury in patients who had undergone highly invasive procedures such as open heart surgery or resection of esophageal cancer to explore the possibility of anticytokine therapy.     

Pathogenicity of the entomopathogenic fungus Metarhizium anisopliae (Deuteromycetes) to Ixodes scapularis (Acari: Ixodidae)

The entomopathogenic fungus Metarhizium anisopliae is highly pathogenic to the black-legged tick, Ixodes scapularis. Spore concentrations of 10(8)/ml for engorged larvae and 10(7)/ml for engorged females resulted in 100% tick mortality, 2 wk postinfection. The LC50 value for engorged larvae (concentration to kill 50% of ticks) was 10(7) spores/ml. Metarhizium anisopliae shows considerable potential as a microbial control agent for the management of Ixodes scapularis.     

Reduced abundance of Ixodes scapularis (Acari: Ixodidae) with exclusion of deer by electric fencing

To assess the effect of deer exclusion on populations of Ixodes scapularis Say (formerly I. dammini Spielman, Clifford, Piesman & Corwin) in the northeastern United States, host-seeking ticks and ticks on white-footed mice, Peromyscus leucopus (Rafinesque), were monitored inside and outside a wooded, residential deer exclosure (approximately 3.5 ha) in Lyme, CT, in 1991 and 1992. Another deer exclosure was added in Lyme (approximately 7.4 ha) during 1992. Additional sample sites at other residences served as secondary controls. A seven-wire, slanted, high-tensile electric deer fence was used at both areas. Larvae of I. scapularis were 81.5% (1991) and 97.8% (1992) less abundant within the exclosure than immediately outside the deer exclosures. Nymphs of I. scapularis were 47.4% (1991) and 55.8% (1992) less abundant within the deer fence. The effect on adult ticks was mixed. No difference in tick abundance was seen at the 3.5-ha site. However, larvae, nymphs, and adults were 100, 83.8, and 74.1% less abundant, respectively, in plots at the 7.4-ha exclosure > or = 70 m from the deer fence and isolated from woodlands outside the fence by lawns, driveways, and buildings. The recovery of larvae and nymphs of I. scapularis from mice captured within the deer exclosures indicates that infestations of nymphs and adults are probably, at least in part, a result of the movement of these rodents. Based upon the number of nymphs per 100 m2 infected with Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner, the causal agent of Lyme disease, there were 73 and 82% fewer infected nymphs within the deer exclosures in 1992 in comparison with the number of infected ticks outside the fence and at the secondary control sites, respectively. The exclusion of deer in conjunction with other tick control strategies in large areas could substantially reduce populations of I. scapularis and the risk of acquiring Lyme disease.     

Cotton rats and other small mammals as hosts for immature Dermacentor variabilis (Acari: Ixodidae) in central Oklahoma

Eight species of small mammals were evaluated as potential hosts for American dog ticks, Dermacentor variabilis (Say), in an upland, tallgrass prairie study site in central Oklahoma. Only hispid cotton rats, Sigmodon hispidus, and deer mice, Peromyscus maniculatus, were found to be important hosts for immature D. variabilis. Although D. varibilis larvae and nymphs frequently infested both cotton rats and deer mice, cotton rats were the most important host species for both immature stages in the study area. Cotton rats constituted 63.2% of the total 530 small mammals captured and were hosts to 85.2% of all larvae and 88.7% of all nymphs. Deer mice accounted for 19.8% of all small mammals captured and were hosts for 14.5% of the larvae and 10.8% of the nymphs recovered. The remaining small mammal species were hosts for less than 1% of the immature ticks collected. Larval infestations peaked during summer, whereas summer and spring peaks were noted for the nymphal infestations. The relative importance of cotton rats and deer mice as hosts for immature ticks could be largely, but not completely, explained by cotton rats being more than three times as abundant as deer mice. Attachment site data indicated that differences in grooming behavior also might be partially responsible for the larger infestations observed on cotton rats. Other possible ecological and behavioral explanations of the heavy infestations observed on cotton rats are discussed.     

Geographic distribution of ticks (Acari: Ixodidae) in Michigan, with emphasis on Ixodes scapularis and Borrelia burgdorferi

A 12-yr (1985-1996) passive survey in Michigan based upon tick submissions from citizens yielded 4,755 ticks of 21 species, 16 of which were probably indigenous in the state. Three species of Dermacentor [most common, D. variabilis Say and D. albipictus (Packard)]; 2 species of Amblyomma [most common, A. americanum (L.)]; and 12 species of Ixodes (most common, I. cookei Packard and I. scapularis Say), as well as Haemaphysalis leporispalustris (Packard), Rhipicephalus sanguineus Latreille, and the soft ticks Ornithodoros kelleyi Cooley & Kohls, and Otobius megnini (Duges) were submitted. New state records were I. kingi Bishopp, I. texanus Banks, I. sculptus Neumann, and I. baergi Cooley & Kohls. Examination of gut smears from dissections of 1,037 ticks of 13 species by indirect immunofluorescent assay, using murine monoclonal H9724 as the primary antibody, revealed that 11 of 175 I. scapularis were infected with Borrelia spp. All positive I. scapularis were from Menominee County in the upper peninsula of the state, which also provided 79.8% of all submitted I. scapularis. Surveys for ticks on 5,449 hunter-killed white-tailed deer were conducted from 1988 to 1990, encompassed deer taken from 65 of the state's 83 counties, and showed that although D. albipictus was distributed widely in the northern part of the state, I. scapularis occurred only on deer taken from southern townships of Menominee County. Of 1,218 canine sera tested for antibodies to B. burgdorferi in 1992 and 1993, 25 of 299 (8.0%) from Menominee County were positive but only 1 of 919 sera submitted from 5 counties in the lower peninsula was positive.     

Microclimate-dependent survival of unfed adult Ixodes scapularis (Acari:Ixodidae) in nature: life cycle and study design implications

We report that acetyl-N-Ser-Asp-Lys-Pro (AcSDKP), which removes progenitor cells from cell cycle, in combination with granulocyte-colony stimulating factor (G-CSF) can significantly improve myelorestoration following irradiation (7 Gy). Peripheral blood, spleen and bone marrow (BM) cell recovery and progenitor cell reconstitution [IL-3-responsive colony-forming cells (CFC) and high proliferative potential colony-forming cells (HPP-CFC)] were studied. Studies on the optimal schedule of AcSDKP administration revealed maximal effects on progenitor cells when AcSDKP was administered as a continuous infusion for 3 d starting 24 h prior to irradiation and used in combination with G-CSF. The numbers of CFC and HPP-CFC in the BM were significantly increased following irradiation in mice receiving AcSDKP and G-CSF as compared to either drug alone. The numbers of CFC in the spleen were significantly increased in mice receiving AcSDKP and G-CSF on days 10 and 14 as compared to AcSDKP alone, but not G-CSF. Similarly, CFC and HPP-CFC in the spleen were significantly increased in mice receiving AcSDKP and G-CSF on day 18 as compared to mice receiving PBS and G-CSF. These studies suggest that AcSDKP in combination with G-CSF may have potential for the protection of progenitor cells in patients undergoing intensive chemo- and/or radiotherapy.     

Kairomonal activity of white-tailed deer metatarsal gland substances: a more sensitive behavioral bioassay using Ixodes scapularis (Acari: Ixodidae)

OBJECTIVE: The objective was to determine if maximal cytoreductive surgery could carry any benefit in pelvic and abdominal recurrent endometrial carcinoma. METHODS: Twenty women at their first large pelvic or abdominal recurrence from endometrial carcinoma were treated with maximal cytoreductive surgery. Women were classified as R1 (residual tumor) or R0 (no residual tumor) by tumor left at the end of surgery. Adjuvant postoperative therapy was undertaken upon clinical judgement. Progression-free, overall, and cancer-related survivals were analyzed with the product-limit method and compared with the log-rank test. The Cox regression model was used to study the variables involved in progression-free and overall survival. RESULTS: Complete macroscopic resection of tumor was feasible in 13 women (65%). R0 group women had a significant both progression-free (median reached at 9.1 months) and overall survival (median reached at 11.8 months) compared to R1 group women. There were 2 (10%) perioperative deaths. Eight women died of cancer, 5 in the R1 group and 3 in the R0 group. There were four intercurrent deaths in women still free from the disease. Local control of neoplasia was achieved in 84.6% of R0 women and their survival was affected mostly by distant recurrences or intercurrent deaths. Residual tumor at the end of surgery was the only significant variable to affect both progression-free and overall survival. CONCLUSION: Intensive surgery is a valid treatment option in women with large pelvic or abdominal recurrence from endometrial carcinoma. Tumor can be completely resected and local control of the disease can be achieved in most of the patients, although survival could be affected by distant recurrence and intercurrent deaths.     

Life cycle of Ixodes minor (Acari: Ixodidae) in the laboratory

OBJECTIVE: It is commonly believed that the full moon exerts an influence on violence and aggression in psychiatric settings. The literature to date is contentious. This study used a robust methodology to examine the hypothesis that there was an increased frequency of violent and aggressive behaviour among hospitalised psychiatric clients at the time of the full moon. METHOD: Prospective data were collected in five inpatient psychiatric settings across the Northern Sydney Area Health Service. Morrison's hierarchy of violence and aggression was used to rate behaviour. Lunar phases were clearly defined and Poisson regression used to examine relationships between lunar phase and violence. Extraneous temporal variation was considered. RESULTS: No significant relationship was found between total violence and aggression or level of violence and aggression and any phase of the moon. CONCLUSION: Future research could profitably examine the implications of a belief in the lunar effect among health workers in the face of evidence that no relationship exists between violence, aggression and the lunar cycle.     

Effects of the feeding process of Ixodes scapularis (Acari: Ixodidae) on embryonic development of its parasitoid, Ixodiphagus hookeri (Hymenoptera: Encyrtidae)

The chalcid wasp Ixodiphagus hookeri (Howard) is a parasitoid of several ixodid ticks including the blacklegged tick, Ixodes scapularis Say. We evaluated effects of the feeding process of nymphal I. scapularis on the embryonic development of I. hookeri. Potentially wasp-parasitized nymphal I. scapularis were collected on Prudence Island, RI. Subsamples of the questing nymph cohort were allowed to feed on laboratory white mice. Both the body length and the scutal length of ticks were measured individually for questing nymphs and for feeding nymphs that were removed from hosts at time intervals of 12, 24, 36, 48, and 72 h after attachment. The diameters of wasp eggs they contained were also measured for each designated time interval. There was a positive relationship between the mean scutal index (ratio between body length and scutal length) of ticks and the mean diameter of wasp eggs during 72 h of tick feeding (P < 0.05). Moreover, it appeared that within 24 h of tick attachment, the scutal index of ticks remained unchanged. However, after that period, the scutal index increased significantly (P < 0.05). Diameters of wasp eggs increased continuously during tick feeding and at 72 h after attachment, enclosed eggs and completely formed larvae were found in several ticks. We conclude that factors related to the feeding process of nymphal I. scapularis are necessary to initiate the embryonic development of wasps.     

Isolation of aerobic microbes from Ixodes scapularis (Acari: Ixodidae), the vector of Lyme disease in the eastern United States

The spirochete Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Benner is transmitted by Ixodes scapularis Say, a vector of Lyme disease. As a 1st step into investigating the possibility of biocontrol of the tick, we identified the microbiota associated with the ticks. We collected, identified, and determined the sex of ticks from foliage and deer. Seventy-three initial bacterial isolates were recovered from 43 ticks (27 adults and 16 nymphs). The bacteria isolated from nymphs were qualitatively different (mainly gram-negative cocci) from the bacteria isolated from adult ticks (gram-negative and gram-positive rods). To determine long-term viability, these isolates were stored for 6 mo under laboratory conditions. After storage, 63 surviving bacterial isolates were characterized using the Biology System of identification by substrate utilization. Forty-four isolates were identified to the species level. Our characterization efforts focused on the 40 spore-forming bacteria, which could prove useful in the biocontrol of ticks. Eleven species of Bacillus were identified. Bacillus thuringiensis-B. cereus was the predominant species group isolated. Six isolates from this group formed crystals.     

Vegetational association of host-seeking adult blacklegged ticks, Ixodes scapularis Say (Acari: Ixodidae), on dairy farms in northwestern Wisconsin

Non-obese diabetic NOD/SCID mice have been used to grow human leukaemia as a systemic disease. The animals were inoculated with leukaemic cells obtained from a 36-year-old male with early B-cell precursor acute lymphoblastic leukaemia and on day 15 were given the first of three weekly injections of 1 mg/kg vincristine or equimolar liposomal vincristine. The development of leukaemia in the mice was monitored by taking weekly blood samples and measuring the cell content by flow cytometry. The median time to 50% human cells in the peripheral blood of mice treated with free vincristine was 41 d from the start of treatment compared with 49 d for mice treated with liposomal vincristine (P < 0.01). The median day of death for mice treated with free vincristine was 47 d from the start of treatment and 57 d for mice receiving liposomal vincristine (P<0.01), thus providing a 21% increase in lifespan for animals treated with the liposomal preparation. There was slightly greater weight loss in mice treated with free vincristine than those given liposomal vincristine. Measurement of in vitro colony forming bone marrow progenitor cells in similarly treated, tumour-free mice, showed no difference in progenitor cell survival between mice that received either type of vincristine. We conclude that encapsulating vincristine in liposomes improves the therapeutic index of this drug measured in mice bearing human leukaemia. This may lead to use of the drug in conventional combination chemotherapy with greater safety or, in this setting, at higher dosage.     

Optimal sampling and spatial distribution of Ixodes pacificus, Dermacentor occidentalis and Dermacentor variabilis ticks (Acari: Ixodidae)

A common method for sampling tick populations is flagging, which is a method of dragging a white cloth over a plant substrate for a fixed distance along a transect. Flagging over rough physical surfaces or using long subtransect lengths could lead to the underestimation of tick densities. Using estimates of the drop-off rates of adult Ixodes pacificus in flag sampling, optimal sampling schemes (the length and number of subtransects) were examined using the relationships between the tick drop-off rates (c), the tick density and distribution patterns and the roughness of the sampled plant substrate. It was found that the optimal number of subtransects and Lmax, the longest subtransect length which did not significantly underestimate the tick density from c, were affected by the tick density, substrate roughness and tick distribution pattern. This study also showed that the density and distribution of I. pacificus and Dermacentor occidentalis varied greatly over time in the populations sampled, while the Dermacentor variabilis densities were low and showed no significant changes over time. Both I. pacificus and D. occidentalis had clumped distributions along trails and these clumps were aggregated. However, the clump sizes (or individual clump areas) changed significantly over time because of density fluctuation or the movement of ticks. Finally, a positive association between the number of I. pacificus and D. occidentalis adults was observed from 2 m subtransect flag collections in March 1995; no relationship was found in 1994.     

Mark and recapture of adult Ixodes pacificus (Acari: Ixodidae) to determine the effect of repeated removal sampling on tick abundance

The effect of biweekly flagging on adult Ixodes pacificus Cooley & Kohls abundance was determined by using mark-recapture techniques. Ticks were recaptured up to three times during the 12-wk study, and increasingly higher proportions of marked ticks were recaptured as the study progressed. By week 6 of the study, 46% of the collected ticks had been marked, and on the final sampling date, all ticks had been captured previously. Removal sampling would have substantially reduced estimates of the abundance of the questing I. pacificus adult population. For ecological studies that require the repeated sampling of a given habitat, the most representative data are obtained by returning flagged ticks to their collection site.     

Kinetics of male pheromone production by Amblyomma variegatum (Acari: Ixodidae)

Males of Amblyomma variegatum (F.), when attached on their host, produce a pheromonal blend composed of methyl salicylate, ortho-nitrophenol, and nonanoic acid, which acts as an aggregation-attachment pheromone. Using olfactometry assays, assays on hosts, and quantification of the compounds by capillary gas chromatography, the kinetics of pheromone production has been studied. Males engorged for at least 10 d elicit attachment from most of the tested females. Attractiveness of males for females is significant only between days 14 and 23 of engorgement. Before 10 d of attachment, the amounts of pheromones detected in hexanic male body extracts are very low (< 10 ng); they then increase to a maximum of 2 micrograms for the major compound per male. Large amounts can be detected until 80 d of engorgement. The inter-individual variation is very large throughout the engorgement period. The interest and the limitations of the different techniques used are discussed.     

Ticks (Acari: Ixodidae) collected from small and medium-sized Kansas mammals

Seven species of hard-bodied ticks were collected from 20 species of small and medium-sized mammals in Kansas; Amblyomma americanum L., Dermacentor variabilis (Say), Haemaphysalis leporispalustris (Packard), Ixodes cookei Packard, I. kingi Bishopp, I. sculptus Neumann, and I. texanus Banks. Dermacentor variabilis was found statewide, A. americanum only in the eastern one-third of the state, and the Ixodes spp. and H. leporispalustris were widely scattered. The most common tick found was D. variabilis, both by itself and in association with other ticks. Mammals that ticks were collected from included Canis latrans Say, Cynomys ludovicianus ludovicianus (Ord), Didelphis virginianus Kerr, Geomys bursarius (Shaw), Lynx rufus (Schreber), Marmota monax bunkeri Black, Mephitis mephitis (Schreber), Microtus ochrogaster (Wagner), Mus musculus L., Peromyscus leucopus (Rafinesque), P. maniculatus (Wagner), Procyon lotor hirtus Nelson and Goldman, Reithrodontomys megalotis (Baird), Sciurus niger rufiventer Geoffroy, Sigmodon hispidus texianus (Audubon and Bachman), Sylvilagus floridanus (J. A. Allen), Taxidea taxus taxus (Schreber), and Vulpes velox velox (Say).     

Experimental transmission of Ehrlichia canis (Rickettsiales: Ehrlichieae) by Dermacentor variabilis (Acari: Ixodidae)

Four trials were conducted in which laboratory-reared Dermacentor variabilis nymphs were exposed to Ehrlichia canis by feeding on experimentally infected dogs as soon as classical morulae were detected in peripheral blood monocytes. After molting 25, 50 or 90 adult tick pairs were permitted to feed on 7 Ehrlichia-naive dogs. Transmission occurred in trials 1 (1/1 dog), 3 (1/1 dog) and 4 (2/2 dogs) but not in trial 2 (0/3 dogs), with 4 of 7 dogs becoming infected. Successful transstadial transmission was demonstrated by detection of morulae in peripheral blood lymphocytes and by seroconversion to Ehrlichia canis 30 d post-exposure. Incubation periods ranged between 17 and 22 days (mean = 19). Clinical signs, typical of ehrlichiosis, included mucopurulent ocular discharge, lymphadenopathy and malaise with accompanying pyrexia, leukopenia and thrombocytopenia. Pyrexia, thrombocytopenia and erythrophagocytosis and vacuolization of the cytoplasm of monocytic cells were observed 1-4 d prior to detection of morulae. This is the first demonstration that a tick other than Rhipicephalus sanguineus is capable of transstadial transmission of this important pathogen of dogs.     

Establishment of the foreign partheonogenetic tick Amblyomma rotundatum (Acari: Ixodidae) in Florida

The parthenogenetic tick Amblyomma rotundatum, a Central and South American species, has become established in southern Florida. The date of introduction is unknown, but it is suspected to be either during the 1930s, when 1 of its natural hosts, the giant or marine toad, Bufo marinus, was introduced to southern Florida as a potential biological control of pest beetles in sugar cane fields, or between 1955 and 1964 when specimens of B. marinus were accidentally or deliberately released in the greater Miami area. Several museum specimens of this toad collected in the Miami area 25 April 1979 had nymphal and adult A. rotundatum attached. Subsequent examination of living giant toads collected at another Miami area site from 1983 through 1985 revealed larval, nymphal, and adult A. rotundatum and confirmed colonization of this tick. Under laboratory conditions, another neotropical amphibian and reptile tick, Amblyomma dissimile, is capable of transmitting Cowdria ruminantium, the causative agent of heartwater, a disease present in the Caribbean area. Therefore, we suggest that A. rotundatum should also be tested for vectorial competence.