Invasive potential of cattle fever ticks in the southern United States.

ABSTRACT' BACKGROUND: For >100 years cattle production in the southern United States has been threatened by cattle fever. It is caused by an invasive parasite-vector complex that includes the protozoan hemoparasites Babesia bovis and B. bigemina, which are transmitted among domestic cattle via Rhipicephalus tick vectors of the subgenus Boophilus. In 1906 an eradication effort was started and by 1943 Boophilus ticks had been confined to a narrow tick eradication quarantine area (TEQA) along the Texas-Mexico border. However, a dramatic increase in tick infestations in areas outside the TEQA over the last decade suggests these tick vectors may be poised to re-invade the southern United States. We investigated historical and potential future distributions of climatic habitats of cattle fever ticks to assess the potential for a range expansion. METHODS: We built robust spatial predictions of habitat suitability for the vector species Rhipicephalus (Boophilus) microplus and R. (B.) annulatus across the southern United States for three time periods: 1906, present day (2012), and 2050. We used analysis of molecular variance (AMOVA) to identify persistent tick occurrences and analysis of bias in the climate proximate to these occurrences to identify key environmental parameters associated with the ecology of both species. We then used ecological niche modeling algorithms GARP and Maxent to construct models that related known occurrences of ticks in the TEQA during 2001-2011 with geospatial data layers that summarized important climate parameters at all three time periods. RESULTS: We identified persistent tick infestations and specific climate parameters that appear to be drivers of ecological niches of the two tick species. Spatial models projected onto climate data representative of climate in 1906 reproduced historical pre-eradication tick distributions. Present-day predictions, although constrained to areas near the TEQA, extrapolated well onto climate projections for 2050. CONCLUSIONS: Our models indicate the potential for range expansion of climate suitable for survival of R. microplus and R. annulatus in the southern United States by mid-century, which increases the risk of reintroduction of these ticks and cattle tick fever into major cattle producing areas.

Analysis of doramectin in the serum of repeatedly treated pastured cattle used to predict the probability of cattle fever ticks (Acari: Ixodidae) feeding to repletion.

Analysis of doramectin concentration in blood serum of pastured cattle injected repeatedly (12 treatments) at two different dosage rates and 28-day intervals throughout the year was used to predict the probability that cattle fever ticks could successfully feed to repletion during the interval between any two consecutive treatments. Treatment at ~270 µg/kg indicated that serum doramectin concentration dropped below the baseline concentration estimated for tick survival (8 ppb) in 7 of the 12 treatments. However, the longest period between any two treatments during which the doramectin concentration remained below the 8 ppb baseline level for successful tick feeding was 15 days, making it virtually impossible for any ticks to reach ovipositional status prior to a subsequent treatment. At a dosage rate of ~540 µg/kg, the concentration dropped below the baseline tick survival level (8 ppb) only once, following the initial treatment, and the duration during which the concentration remained below the baseline level prior to the subsequent treatment was only 6 days. Thus, at the high dosage rate results indicated, with absolute certainty, that no ticks could successfully feed to repletion between any two consecutive treatments. Based on the data obtained in the study it was concluded that analysis of doramectin concentration in serum of treated animals would be a reliable predictor for assessing the probability that ticks could successfully develop to repletion. More importantly, results demonstrated that the trial policy, instituted by the Cattle Fever Tick Eradication Program, of repeatedly treating cattle with doramectin injections at 25-28 day intervals for eliminating cattle fever ticks would produce little or no risk of any viable ticks developing to repletion and re-infesting the field between treatment applications.

Efficacy and blood sera analysis of a long-acting formulation of moxidectin against Rhipicephalus (Boophilus) microplus (Acari: Ixodidae) on treated cattle.

The therapeutic and persistent efficacy of a single subcutaneous injection of a long-acting formulation of moxidectin at a concentration of 1 mg/kg body weight was determined against Rhipicephalus (Boophilus) microplus (Canestrini), along with the concentration-time blood sera profile in treated cattle. The therapeutic efficacy against ticks of all parasitic stages on cattle at the time of treatment was >99.9%, and the mean tick number, index of fecundity, engorgement weight, and egg mass weight of ticks recovered from treated animals were all significantly lower than ticks from untreated animals. The index of fecundity, engorgement weight of females, and egg mass weight of ticks recovered from treated animals infested at weekly (7-d) intervals between 14 and 63 d posttreatment were significantly lower than for ticks on untreated animals, whereas the number of ticks per animal recovered from treated cattle remained lower than that of untreated cattle for up to 49 d posttreatment. The percentage control remained >99% at weekly intervals between 14 and 49 d posttreatment, which is the minimum level of efficacy considered acceptable for use in the United States Cattle Fever Tick Eradication Program. The serum concentration of moxidectin in treated cattle increased to 25.6 ppb (parts per billion) within 1 d after treatment, and peaked at 47.3 ppb at 8 d posttreatment. Moxidectin sera levels remained above the estimated 100% threshold level for elimination of feeding ticks (5-8 ppb) for 44-53 d after treatment. The label claim of 50 d of prevention against reinfestation for the long-acting moxidectin formulation used in the study was supported by the efficacy and sera concentration data obtained. Based on these results, cattle could be treated at 63-d intervals with minimal risk of viable ticks detaching from treated animals. This treatment interval would be 4.5-fold longer than the presently required treatment interval of 14 d, thus leading to approximately 75% reduction in gathering and handling costs of cattle incurred by producers.

Therapeutic and persistent efficacy of a long-acting (LA) formulation of ivermectin against Rhipicephalus (Boophilus) microplus (Acari: Ixodidae) and sera concentration through time in treated cattle.

The concentration-time profile, therapeutic, and persistent efficacy of a single subcutaneous injection of cattle with a long-acting (LA) formulation of ivermectin at a concentration of 630microg/kg of body weight were determined against Rhipicephalus (Boophilus) microplus. Ivermectin sera concentration in treated cattle increased to 13.0ppb within 1d after treatment, and peaked at 26.2ppb at 11d post-treatment. Ivermectin sera levels remained above the threshold level for control of feeding ticks (>or=8ppb) for 42.6d after treatment. Therapeutic efficacy of ticks on treated animals was >99.9%, and tick number, index of fecundity, engorgement weight, and egg mass weight of ticks from treated animals remained dramatically less than ticks from untreated animals. Tick number and reproductive capacity of ticks infested on treated animals at 14 and 28d post-treatment were less than for ticks on untreated animals, whereas engorgement weight and egg mass weight of treated ticks remained lower than that of untreated ticks 49d post-treatment. However, the level of control against ticks infested at 14d after treatment (99.9%) was the only post-treatment infestation interval that provided the required 99% control necessary for use in the U.S. tick eradication program. The 14d post-treatment infestation was also the only interval at which infested ticks were exposed to ivermectin levels above the threshold level of 8ppb for the entire parasitic development period. Cattle would have to be treated at intervals of no more than 31d apart to ensure that no viable ticks could reach repletion and detach from the host. Although this treatment interval is >2-fold longer than the present treatment requirement (14d), it is dramatically less than the label claim for the LA ivermectin formulation of 75d of prevention against re-infestation.

Topical treatment of white-tailed deer with an acaricide for the control of Ixodes scapularis (Acari: Ixodidae) in a Connecticut Lyme borreliosis hyperendemic Community.

The 4-Poster device for the topical treatment of white-tailed deer, Odocoileus virginianus (Zimmermann), against ticks using the acaricide amitraz, was evaluated in a Lyme borreliosis endemic community in Connecticut. As part of a 5-year project from 1997 to 2002, 21-24 of the 4-Posters were distributed at residential sites in Old Lyme, CT, in a core treatment area of approximately 5.2 km(2) in fall 1997. The 4-Posters were active October to mid-December and March into May, corresponding to the peak periods of activity for adult Ixodes scapularis in this particular area. Corn consumption ranged from 361 to 4789 kg/month for October and November and 696-3130 kg/month during April. Usage of 4-Posters by deer generally was high (>90%), except during acorn masts in fall 1998 and 2001. Amitraz was applied by rollers at the estimated rate of 1.3 g active ingredient/ha/year. The abundance of host-seeking I. scapularis nymphs declined significantly (p < 0.001) in the core treatment area, as compared to a control community in Old Saybrook, CT, through 2004, over the project period from 1998 to 2003, from 9.3/100m(2) to 0.97/100m(2), rising to 1.90/100m(2) in 2004. From 1999 through 2003, there were 46.1%, 49.6%, 63.4%, 64.6%, and 70.2% reductions, respectively, in the nymphal tick population in comparison with the untreated community and initial tick abundance in 1998. Control of I. scapularis adults declined to only 19.1% in 2004; 2 years after the treatment of deer was discontinued. Differences in nymphal tick abundance between the control and core treatment area were significant in 1999 (p = 0.042) and highly significant in 2001 (p < 0.001) and 2002 (p = 0.002). The passive topical application to deer of the acaricide amitraz resulted in a significant decrease in the population of free-living I. scapularis nymphs in the treated core in Connecticut.

Acaricidal treatment of white-tailed deer to control Ixodes scapularis (Acari: Ixodidae) in a New York Lyme disease-endemic community.

The efficacy of topically treating white-tailed deer with an acaricide was evaluated in a Lyme disease-endemic community of southern New York State. Twenty-four 4-Poster feeders were placed in a 5.2 km(2) treatment area in Bedford, NY, while a site in Lewisboro, NY, 4.8 km distant, served as control. Treatment periods ran from 15 September to 15 December each fall from 1997 to 2001, and from 15 March to 15 May each spring from 1998 to 2002. Corn consumption averaged 15,779 kg in fall sessions and 9054 kg in spring sessions, and a mean of 89.6% of deer in the study area showed evidence of using the feeders. Deer densities, estimated by aerial snow counts, averaged 22 and 28 deer per km(2) in Bedford and Lewisboro, respectively, over a 3-year period. Significant reductions in tick numbers on deer captured in the treatment area were noted in fall 1999 compared to deer captured at the control site. Drag sampling for nymphal host-seeking ticks indicated 63.6% control in 2001, which dropped to 54.8% the following year, but reached 80% in 2003. Higher-than-normal acorn production in 2001 that likely caused a drop in deer visitation to the feeders may have reduced efficacy against larval ticks in 2002. The 4-Poster effectively reduced the density of Ixodes scapularis, though the level of control is dependent on environmental factors that affect feeding behavior of white-tailed deer.

The impact of 4-Poster deer self-treatment devices at three locations in Maryland.

From 1998-2002 twenty-five deer self-treatment devices (4-Posters), using 2% amitraz, were operated at three locations in Maryland to determine their effectiveness in controlling blacklegged ticks, Ixodes scapularis Say, and lone star ticks, Amblyomma americanum (L.). Each treatment site was approximately 518 ha and paired with a similar site lacking 4-Posters. Locations varied in deer density, tick abundance, and land use. Flagging for host-seeking ticks showed declines in tick populations at all treatment sites compared to control sites by the third year. By 2002, control of I. scapularis nymphs attributable to the 4-Poster intervention at the three sites was 69.0%, 75.8%, and 80%. Control of A. americanum nymphs at the two sites where they occurred was 99.5% and 95.3%. In 2003, the first posttreatment year, control of I. scapularis remained around 2001-2002 levels, but by 2004, an upward trend in nymphal numbers was detectable. Populations of A. americanum showed no increase posttreatment. These results demonstrate that control of these tick species is locally possible with 4-Poster intervention.

Sustained control of Gibson Island, Maryland, populations of Ixodes scapularis and Amblyomma americanum (Acari: Ixodidae) by community-administered 4-Poster deer self-treatment bait stations.

In 1998, twenty-five 4-Poster deer treatment bait stations were deployed on Gibson Island (GI), Maryland, as part of the U.S. Department of Agriculture (USDA) Northeast Area-Wide Tick Control Project. Treatments concluded in June 2002, having achieved 80% and 99.5% control of blacklegged ticks, Ixodes scapularis, and lone star ticks, Amblyomma americanum, respectively. No area-wide tick control was attempted again on the island until 2003, when 15 Dandux-manufactured 4-Posters were purchased by the GI Corporation and operated until the present. Annual flagging at sites on the island and a similar untreated area on the nearby mainland in May and June from 1998 to 2007 has demonstrated that populations of host-seeking nymphs of both tick species have remained at consistently low levels on the island during GI Corporation administration of the 4-Posters, in spite of 40% fewer 4-Posters and increased deer density during 2003-2007.

Evaluation of the United States Department Of Agriculture Northeast Area-wide Tick Control Project by meta-analysis.

As part of the Northeast Area-wide Tick Control Project (NEATCP), meta-analyses were performed using pooled data on the extent of tick-vector control achieved through seven concurrent studies, conducted within five states, using U.S. Department of Agriculture "4-Poster" devices to deliver targeted-acaricide to white-tailed deer. Although reductions in the abundance of all life-stages of Ixodes scapularis were the measured outcomes, this study focused on metrics associated with I. scapularis nymphal tick densities as this measure has consistently proven to directly correlate with human risk of acquiring Lyme disease. Since independent tick sampling schemes were undertaken at each of the five environmentally distinct study locations, a meta-analytic approach permitted estimation of a single true control-effect size for each treatment year of the NEATCP. The control-effect is expressed as the annual percent I. scapularis nymphal control most consistent with meta-analysis data for each treatment year. Our meta-analyses indicate that by the sixth treatment year, the NEATCP effectively reduced the relative density of I. scapularis nymphs by 71% on the 5.14 km(2) treatment sites, corresponding to a 71% lower relative entomologic risk index for acquiring Lyme disease.

Depletion rate of doramectin from blood serum of penned female white-tailed deer (Artiodactyla: Cervidae).

Female white-tailed deer, Odocoileus virginianus (Zimmerman), were held in small pens and administered doramectin by free choice of doramectin-coated whole kernel corn, Zea mays L., fed ad libitum with 19% protein deer pellets also being fed ad libitum in a separate container. The mean concentration of doramectin in the serum during treatment was 72.8 ppb. The mean doramectin concentration in the serum decreased to <2 ppb, the lower limit of detection by high-pressure liquid chromatography, by day 14 after termination of treatment after withdrawal of doramectin-treated corn from the diet.

Depletion rates of injected and ingested ivermectin from blood serum of penned white-tailed deer, Odocoileus virginianus (Zimmermann) (Artiodactyla: Cervidae).

Penned female and male white-tailed deer, Odocoileus virginianus (Zimmerman), were administered ivermectin both by direct subcutaneous injection and by ingestion of ivermectin-medicated whole kernel corn. Depletion rates of ivermectin were determined by biweekly and weekly assays of blood serum. No statistical differences were observed between mean peak ivermectin serum concentrations in deer (data of sexes combined) from injection and ingestion studies, and ivermectin concentrations decreased to below detectable within 21 d after injection and 14 d after ingestion.

Efficacy of the ivomec SR bolus for control of horn flies (Diptera: Muscidae) on cattle in South Texas.

The concentration of ivermectin in the serum of Hereford heifers treated with a single Ivomec SR bolus reached a maximum of 8.8 +/- 0.9 ppb at 2 wk post-treatment. The single bolus treatment resulted in <10% mortality of adult horn flies feeding on the blood of the treated animals over the 21-wk trial. Bioassays of the manure from treated cattle showed complete inhibition of development of immature horn flies through week 19 post-treatment. When the trial was repeated using two Ivomec SR boluses/heifer, the concentration of ivermectin in the serum of the treated cattle reached a maximum level of 31.2 +/- 3.9 ppb at week 13 post-treatment. During the first 17 wk of treatment, the use of two boluses/heifer resulted in 96.2 and 81.2% mortality of adult male and female horn flies feeding on the blood of treated animals, respectively. From these studies, we conclude that a single Ivomec SR bolus used as an anthelmintic treatment can be expected to provide significant control of immature horn flies developing in the manure, but not of adults feeding on the treated cattle.

Introduced ticks and tick-borne diseases: the threat and approaches to eradication.

Exotic tick species and tick-borne diseases are serious threats to live-stock, companion animals, and wildlife in the United States. Recurring introductions of exotic tick species into the United States are a significant indicator of the degree of risk. Successful tick-eradication campaigns, such as the national program that eradicated Boophilus annulatus and B. microplus from the United States, the Cattle Fever Tick Eradication Program of the US Department of Agriculture's Veterinary Services that protects against the re-entry and dissemination of Boophilus ticks from Mexico back into their former haunts in the southern states, and the eradication action that eliminated Rhipicephalus evertsi from a game park in Florida, are sources of useful information that aid in elucidating essential elements of successful eradication programs. Examples of failed eradication programs in places such as Puerto Rico and St. Croix also have heuristic value. Among the varieties of tick species and related infectious agents that threaten the United States, Boophilus ticks and bovine babesiosis, Amblyomma species (especially the tropical bont tick) and heartwater, and equine babesiosis, for which endemic vectors exist, are of special concern. Risk assessments to accumulate, evaluate, and synthesize information needed to appraise risks, consequences, and preparedness are necessary not just to inform federal, state, and local officials, as well as producers and stakeholders, but also to facilitate the creation of emergency response plans.

Control of Ixodes scapularis and Amblyomma americanum through use of the '4-poster' treatment device on deer in Maryland.

Deer self-treatment devices ('4-posters') were evaluated for their efficacy in reducing populations of blacklegged ticks, Ixodes scapularis, and lone star ticks, Amblyomma americanum. At each of three locations in Maryland, 25 '4-posters' were operated in study areas of approximately 5.18 km2. Populations of host-seeking ticks were monitored by flagging of treated areas and similar untreated control areas without '4-posters.' From 1998 to 2002 the percent mortalities achieved were 69, 75.8 and 80 at the three study sites infested with I. scapularis nymphs, and 99.5 and 95.3 for A. americanum nymphs at the two sites where this species occurred.

Synganglial and neurosecretory morphology of female Ornithodoros parkeri (cooley) (acari: Argasidae).

Single esophageal and paired cheliceral, palpal, pedal (I-IV), and opisthosomal nerves enter the synganglion and form specific neuropilar ganglia. The ganglia are integrated by a complex series of commissures and connectives. Eighteen paraldehyde-fuchsin-positive neurosecretory regions, which vary greatly in size and amount of granular neurosecretory material, are each associated (one or more) with neuropilar ganglia. Presumably transport of neurosecretory materials to target tissues occurs through axonal pathways, perineurial-neural lamella associations, and the neurohemal retrocerebral organ complex.

The mouthparts of female Corethrella brakeleyi and C. wirthi (Diptera: Chaoboridae).

The mouthparts of female Corethrella brakeleyi and C. wirthi were studied using light and electron microscopy. Mandibles, hypopharynx and labium are highly sclerotized and are modified for obtaining blood meals. All structures were larger in C. brakeleyi than in C. wirthi except mandibular and hypopharyngeal teeth; these were smaller and more numerous in C. brakeleyi. The labium of both species terminates in peg-like structures which are similar to those reported from several genera of mosquitoes. Sensillae on the second segment of the maxillary palps appear to be identical to those described in both biting and nonbiting male and female blackflies.

Reproductive morphology and Spermatogenesis in Dermanyssus gallinae (DeGeer) (Acari: Dermanyssidae).

Sperm induction pores, tubuli annulati, rami sacculi, sacculus foemineus, cornu sacculus and acantho-membranous hood in female Dermanyssus gallinae support the probability of coxal insemination as reported for some other mesostigmatid species. Development of the circular testis in males (from fusion of two anterolaterally projecting protonymphal testicular arms) and formation of haploid (n = 3) spermatids from haploid (n = 3) spermatogonial cells is entirely mitotic. Spermatogenesis begins within one hour after protonymphal feeding and continues through adult ecdysis in normally fed individuals. Nutritional deficit causes cessation of spermatogenic divisions in deutonymphs starved 23 days, but is rapidly rejuvenated upon feeding. No rejuvenation of division was observed in adult males given the opportunity to feed, thus adult males might not feed.