Acaricide resistance profiles of single- and multi-host ticks from communal and commercial farming areas in the Eastern Cape and North-West Provinces of South Africa

A field study (February 2000 to August 2001) was conducted on communal and commercial farms in the Eastern Cape and North-West Provinces of South Africa to detect the levels of tick resistance to commonly used acaricides. The larvae obtained from engorged females of the one-host tick Boophilus decoloratus, the two-host tick Rhipicepalus evertsi evertsi and the three-host ticks Amblyomma hebraeum and Rhipicephalus appendiculatus were tested against various concentrations of amitraz, chlorfenvinphos and cypermethrin using the Shaw Larval Immersion Test method. Ticks from the communal farms showed higher levels of resistance to cypermethrin and some resistance to chlorfenvinphos whilst no resistance was detected against amitraz. However, ticks from commercial farms were equally resistant to amitraz, chlorfenvinphos and cypermethrin. The B. decoloratus populations tested were considerably more resistant to all the acaricides tested than the R. evertsi evertsi, A. hebraeum and R. appendiculatus populations. This supports the hypothesis that single-host ticks develop resistance faster than multi-host ticks. This trend was recorded on most of the farms where single- and multi-host ticks co-existed. It was concluded that the use of acaricides at high frequencies and high concentrations was one of the main causes of tick resistance in the study areas. Possible factors which caused the resistance problems are discussed and acaricide management strategies recommended.     

Acaricide efficiency of amitraz/cypermethrin and abamectin pour-on preparations in game

The efficacy of an amitraz/cypermethrin pour-on preparation (1% w/v each) was tested against natural tick infestations of buffaloes, eland and blesbok in three separate trials. The eland were also treated with a 0.02% abamectin (w/v) acaricidal pour-on preparation. The amitraz/cypermethrin pour-on was effective against Amblyomma hebraeum, Rhipicephalus evertsi evertsi, Rhipicephalus appendiculatus and Hyalomma marginatum rufipes on the buffaloes. Both acaricides were effective against R. appendiculatus and Rhipicephalus (Boophilus) decoloratus in the eland. The amitraz/cypermethrin acaricide was effective against R. (Boophilus) decoloratus in the blesbok. Ticks can cause damage to the skins, secondary infections, abscesses, anaemia, loss of condition, tick toxicosis and act as vectors of infectious diseases. Introduction of hosts and/or ticks from endemic to non-endemic areas because of translocation of game, may lead to severe losses. The pouron acaricides tested were effective against natural tick infestations and should always be used according to the manufacturer's instructions and efficacy claims.     

Reliminary survey of ticks (Acari: Ixodidae) on cattle in Central Equatoria State, Southern Sudan

In a preliminary survey conducted in 2005, the species composition and seasonality of ticks infesting cattle in Central Equatoria State, Southern Sudan was determined. Three locations were selected (Gumbo, Khor Rumla and Nyaing) and surveyed every 3 months. Two cattle herds in each of the three locations were visited four times during the study period. Total body collections of ticks were made from each of five cattle (Nilotic Zebu breed) kept in six different herds. Four tick genera and ten species were identified. The tick species identified were Amblyomma lepidum, Amblyomma variegatum, Boophilus annulatus, Boophilus decoloratus, Hyalomma marginatum rufipes, Hyalomma truncatum, Rhipicephalus appendiculatus, Rhipicephalus evertsi evertsi, Rhipicephalus praetextatus and Rhipicephalus sanguineus group. The highest number of ticks was collected in October during the rainy season. A finding of great significance was that R. appendiculatus, vector of East Coast fever, has now firmly established itself throughout the year with possible implications for cattle production in Central Equatoria State.     

Influence of exposure period and management methods on the effectiveness of chickens as predators of ticks infesting cattle.

Experiments undertaken on Rusinga Island, Kenya, indicated that Rhipicephalus appendiculatus (Neumann) was the predominant tick consumed on tick-infested cattle by chickens, followed by Amblyomma variegatum (Fabricius), Boophilus decoloratus (Koch) and Rhipicephalus evertsi evertsi (Neumann). The number of ticks consumed by chickens released with cattle in a 3 h release period was not significantly different from ticks consumed in a 4 h release period. In a free management system, chickens 1-3 months of age consumed more ticks than older ones; in a confined management system however, chickens 4-6 months of age consumed more ticks than younger birds. Although the numbers of ticks consumed by local and exotic breeds of chickens were not significantly different, a period of 8-10 weeks was necessary for the exotic breeds to become behaviorally adapted to tick predation on cattle. Local chickens bred in environments where cattle were absent took a shorter period of 4-6 weeks to adapt. It was concluded that the relatively large number of ticks consumed by each chicken during the 3 or 4 h release period is an indication that a good reduction of ticks on cattle can be achieved through tick predation by chickens. The preponderance of R. appendiculatus over other tick species consumed was a reflection of its relative abundance on cattle on the island rather than a selective preference for tick species.     

Seroprevalence of Babesia bigemina in smallholder dairy cattle in Tanzania and associated risk factors

Variations in the seroprevalence of antibody to Babesia bigemina infection by farm and animal level risk factors were investigated for 2 contrasting regions of Tanga and Iringa in Tanzania. Tanga is situated in the eastern part of the country and has typical tropical coast climate while Iringa is situated in the Southern Highlands and has a tropical highland climate. Two hundred farms from each region were selected using simple random sampling procedure and visited once between January 1999 and April 1999. Blood samples were collected from 1329 smallholder dairy animals on selected farms for harvesting serum which was subsequently used for serodiagnosis of B. bigemina using an indirect enzyme linked immuno-sorbent assay (ELISA). Of the 1329 sera samples screened, 34.9 % were positive for B. bigemina. The prevalence was higher in Iringa Region [43%, 95% confidence intervals (CI) = 39.5-47.3] than in Tanga Region (27%, CI = 23.6-30.5). Using a logistic binomial regression model as an analytical method for predicting the likelihood of animal seropositivity, we found (in both regions) that the risk of positive reaction varied with the animal's age, history of grazing and geographical location. Seroprevalence increased with age (beta = 0.01 and 0.01 per year of age, P < 0.005 in Tanga and Iringa, respectively). Animals located in Lushoto and Iringa urban district were associated with increased risk of seropositivity [Odds ratio (OR) = 4.24, P = 0.001, for Lushoto, and OR = 1.81, P = 0.040, for Iringa Urban, respectively). Animals grazed 3 months prior to sampling had higher odds for seropositivity than zero/semi-grazed, despite farmer-reported high frequency of tick control (OR = 2.71, P = 0.0087, for Tanga, and OR = 4.53, P = 0.001, for Iringa). Our study suggests that even though herd sizes are small, B. bigemina infection is widespread in many smallholder dairy farms and endemic stability with respect to this disease has not yet been attained, but the observed levels are sufficiently high to ensure that clinical disease would be a risk.     

Species composition and geographic distribution of ticks infesting cattle, goats and dogs in a temperate and in a subtropical region of south-east Africa

The species and distribution of ticks infesting cattle, goats and dogs in the eastern region of the Eastern Cape Province, South Africa and Maputo Province, Mozambique were determined from collections made from these animals at 72 localities in the former region and 30 in the latter. Eleven ixodid and one argasid species were recovered in the Eastern Cape Province and 15 ixodid species in Maputo Province. The most common ticks infesting cattle and goats in both provinces were Amblyomma hebraeum, Rhipicephalus (Boophilus) microplus, Rhipicephalus appendiculatus and Rhipicephalus evertsi evertsi. The dominant species on dogs were Haemaphysalis elliptica and Rhipicephalus simus. The geographic distributions of the major species and some of the minor species in both regions were plotted. The partial or complete displacement of the indigenous tick Rhipicephalus (Boophilus) decoloratus by the introduced species R. (B.) microplus was a major feature of both surveys.     

Efficacy of ivermectin delivered from an intraruminal sustained-release bolus against natural infestations of five African tick species on cattle

The efficacy of ivermectin delivered by an orally administered prototype sustained-release bolus over approximately 90 days was evaluated against natural infestations of five African tick species. Twenty cattle, allocated by restricted randomization based on counts of standard Boophilus decoloratus, were allocated to two groups and were either given an ivermectin bolus or designated as non-medicated controls. All cattle grazed a single pasture of native grasses for 20-40 days before treatment and until trial termination. Starting on Days 27, 40, 68 and 82 after bolus administration, four replicates were confined to individual tick-collection stanchions for 4 to 5-day periods. Ticks recovered from these cattle were counted by species, sex, and stage and degree of repletion; engorged females were weighed and incubated to determine the number which oviposited. For the other replicates, half-body counts of adult ticks (classified by species, sex and degree of repletion by females) were made at 1- and 2-week intervals through Day 90. Among replicates confined to stanchions periodically, fewer (P less than 0.05) engorged adult female B. decoloratus, Hyalomma spp., Rhipicephalus appendiculatus and Rhipicephalus evertsi evertsi were collected from bolus-treated cattle than from controls. Numbers of engorged adult female Amblyomma hebraeum were reduced, but differences were not statistically significant (P greater than 0.10). Among cattle maintained continuously on pasture, tick numbers were reduced on the ivermectin-treated groups. A significant (P less than 0.05) treatment by linear time effect was seen for all adult ticks counted except R. appendiculatus. A significant (P less than 0.05) treatment by quadratic time effect was seen for A. hebraeum, B. decoloratus and R. evertsi evertsi, and overall treatment differences were significantly different (P less than 0.05) for these species. The differences tended to increase with time. Except for Boophilus, reductions in tick numbers on treated animals relative to controls were not readily apparent. There were no adverse reactions attributable to ivermectin treatment or the presence of the bolus. Each treated animal retained its bolus throughout the trial, based on metal detection.     

Predation on livestock ticks by chickens.

Chickens were shown to be natural predators of ticks. Rhipicephalus appendiculatus were recovered in large numbers from the crops and gizzards of chickens which had scavenged for 30 min-1 h among tick-infested cattle. Other ticks recovered were Amblyomma variegatum and Boophilus decoloratus. The numbers of ticks recovered ranged from 3 to 331, with an average of 81 per chicken. Cattle facilitated the predation of ticks by certain behavioural actions. Chickens also picked up both engorged and unengorged ticks seeded on vegetation, but unengorged ticks were preferred. The possibility of tick predation by chickens forming a viable biological control component in an integrated tick management system in Africa is discussed.     

Delayed-type hypersensitivity test for assessing tick-immune status of cattle in Zambia

Delayed-type hypersensitivity skin reactions were used to assess the tick resistance status of Tonga calves in Zambia. The antigen used in the tests was a homogenate of unfed nymphal Rhipicephalus appendiculatus which had been shown to give protective immunity in guinea pigs to adult female R appendiculatus. There was a significant negative correlation between the intensity of the reactions and the total number of ticks (Amblyomma variegatum, R appendiculatus, Hyalomma truncatum, Boophilus decoloratus and Rhipicephalus species) on the animals.     

Seroprevalence Estimation and Risk Factors for <Emphasis Type="Italic">A. marginale</Emphasis> on Smallholder Dairy Farms in Tanzania

A cross-sectional serological survey of A. marginale was conducted on 200 randomly selected smallholder farms in each of the Tanga and Iringa Regions of Tanzania between January and April 1999. Sera, from dairy cattle of all ages, sexes and breeds were tested for antibodies against A. marginale using an indirect enzyme-linked immunosorbent assay. Antibodies to A. marginale were present in cattle throughout the study areas and the overall prevalence was 20% for Tanga and 37% for Iringa. The forces of infection based on the age seroprevalence profile were estimated at 8 for Tanga and 15 for Iringa per 100 cattle years-risk, respectively. In both regions, seroprevalence increased with age (β = 0.01 and 0.017 per year of age, p < 0.005, in Tanga and Iringa, respectively). Older animals in Iringa were significantly and negatively associated with decreased seropositivity (β = −0.002, p = 0.0029). Further results of logistic regression models reveal that geographic location of animals in Tanga was associated with seropositivity (odds ratio (OR) = 2.94, p = 0.005, for Tanga Rural and OR = 2.38, p = 0.066, for Muheza). Animals acquired as a gift in Iringa had higher odds for seropositivity than brought-in cattle (OR = 2.44, p = 0.005). Our study has identified and quantified some key risk factors that can guide planners devising disease control strategies.     

Tick infestations on livestock in the yemen arab republic and their potential as vectors of livestock diseases

A survey of ticks on cattle, camels, sheep, goats and donkeys in four different geographical locations of the Yemen Arab Republic (YAR) was carried out to provide more information on the possible risk of tick-borne diseases to imported exotic cattle included in the YAR's plans for livestock development. The most abundant ticks were Hyalomma spp. particularly on camels. Ticks found on cattle included Hyalomma spp., Amblyomma variegatum, Boophilus annulatus and Rhipicephalus spp. In general with the exception of camels tick burdens on all species of domestic livestock were very low. Two hundred and ninety eight serum samples from miscellaneous adult cattle throughout the country were negative to a test for Anaplasma marginale antibodies. It is speculated that tick burdens in the YAR are too low for significant disease transmission and the implications of the findings are discussed.     

Factors related to cattle infestation level and resistance to acaricides in Boophilus microplus tick populations in New Caledonia

Boophilus microplus, even in the absence of babesiosis, is a major disease of the cattle in New Caledonia where the particular farming system associates continental European breeds and a tropical climate tempered by the Pacific Ocean. In order to have a better understanding of the factors involved in cattle tick infestation, to decrease the possible wastage and use of chemicals and to increase the lifetime of the acaricides, the veterinary authorities investigated the conditions of the chemical treatments. A survey among 148 cattle farms of the whole of New Caledonia was carried out and factors that explain the development of tick resistance and cattle infestation have been determined. From this survey, three typologies for the main characteristics of the farms have been set up, the technical practices of the farmers and the tick control measures applied by the farmers, respectively. Some variables are significantly associated with the tick resistance to deltamethrin but their contribution to the explanation model is always moderate: farms in the south, with a positive resistance gradient from east to west, absence of bush fire and membership to a cattle farmers organization. The more the farmers have intensified their breeding-male castration, weaning, heifer separation, drenching, etc.-and pasture-high stock rate, mowing, extra feeding of the cows, many paddocks, etc.-techniques, the higher was the probability for the ticks in their farm to be resistant to deltamethrin. The technical details of the acaricide treatment had a low contribution to the explanation model. However, the use of a spray generated more resistance than a dip. Furthermore, there is a negative resistance gradient when the farmers increased the treatment interval average. Considering infestation, none of the variables from the three typologies were associated with the two infestation variables (1: semi-engorged tick females and 2: other ticks) at the herd level. However, the seven studied variables-the three typologies, breed, age, body condition score and breeding status-affected significantly the two infestation variables at the cow level, but their predictive ability remained very low (R(2)<3.5%). This result-individual effect more important than herd effect on the infestation-is confirmed by the importance of the variance of the intra-farm factors (99%) when compared with inter-farm factors (1%). Cows of Charolais breed, in poor body condition, old, pregnant or lactating, and those of the farms with irrational and high pressure control of ticks are the most infested.     

Molecular identification of tick-borne pathogens in Nigerian ticks

A molecular epidemiology investigation was undertaken in two Nigerian states (Plateau and Nassarawa) to determine the prevalence of pathogens of veterinary and public health importance associated with ticks collected from cattle and dogs using PCR, cloning and sequencing or reverse line blot techniques. A total of 218 tick samples, Amblyomma variegatum (N=153), Rhipicephalus (Boophilus) decoloratus (N=45), and Rhipicephalus sanguineus (N=20) were sampled. Pathogens identified in ticks included piroplasmids (Babesia spp., Babesia bigemina and Babesia divergens), Anaplasma marginale and Rickettsia africae. Piroplasmids were identified in A. variegatum, A. marginale was found in R. decoloratus, while R. africae was detected in all tick species examined. Ehrlichia spp. and Theileria spp. were not identified in any of the ticks examined. Of the 218 ticks examined, 33 (15.1%) contained pathogen DNA, with the presence of B. divergens and R. africae that are zoonotic pathogens of public health and veterinary importance. The variety of tick-borne pathogens identified in this study suggests a risk for the emergence of tick-borne diseases in domestic animals and humans, especially amongst the Fulani pastoralists in Plateau and Nassarawa states of Nigeria.     

Laboratory and field investigations into the Theileria parva carrier-state in cattle in Zimbabwe.

The Theileria parva carrier-state in cattle on commercial farms on Zimbabwe was investigated using parasitological and serological methods. The proportion of cattle showing Theileria piroplasms on two farms, which had recent histories of disease outbreaks, were 64% (n = 106, total of heifers and weaned calves examined) and 71.5% (n = 60) while the proportion of T. parva antibodies for the same animals were 59% and 98.5%, respectively. On four farms where no cases of the disease occurred for over 10 years, the average proportion of animals showing piroplasms and antibodies were 55.4% (range 32-82, n = 223) and 73% (range 47-91, n = 223), respectively. However, on another three farms which had no history of theileriosis outbreaks these proportions were very low, being 11.4% (0-24, n = 157) for piroplasms and 12.2% (5-23, n = 157) for antibodies. The mean infection rate in unfed Rhipicephalus appendiculatus adults collected from farms with a high prevalence of cattle which were carriers of Theileria piroplasms during the tick activity season was 29% (range 12-60%) with 9.3 (range 2-18.7) mean infected acini per infected tick. The infectivity of different tick batches to susceptible cattle produced a wide spectrum of theileriosis reactions. Laboratory controlled experiments were carried out to study the persistence of T. parva (Boleni) piroplasms in cattle immunized with this strain as well as its infectivity for ticks and its subsequent transmissibility to cattle. Examination of the salivary glands of 15 batches of ticks collected from six immunized cattle on three different occasions over 18 months showed that none were infected with Theileria parasites. However, the infectivity of other ticks in the same batches to susceptible animals was demonstrated 6, 10 and 18 months after cattle had been immunized with Boleni stabilate.     

A ten-year review of commercial vaccine performance for control of tick infestations on cattle

Ticks are important ectoparasites of domestic and wild animals, and tick infestations economically impact cattle production worldwide. Control of cattle tick infestations has been primarily by application of acaricides which has resulted in selection of resistant ticks and environmental pollution. Herein we discuss data from tick vaccine application in Australia, Cuba, Mexico and other Latin American countries. Commercial tick vaccines for cattle based on the Boophilus microplus Bm86 gut antigen have proven to be a feasible tick control method that offers a cost-effective, environmentally friendly alternative to the use of acaricides. Commercial tick vaccines reduced tick infestations on cattle and the intensity of acaricide usage, as well as increasing animal production and reducing transmission of some tick-borne pathogens. Although commercialization of tick vaccines has been difficult owing to previous constraints of antigen discovery, the expense of testing vaccines in cattle, and company restructuring, the success of these vaccines over the past decade has clearly demonstrated their potential as an improved method of tick control for cattle. Development of improved vaccines in the future will be greatly enhanced by new and efficient molecular technologies for antigen discovery and the urgent need for a tick control method to reduce or replace the use of acaricides, especially in regions where extensive tick resistance has occurred.     

A SURVEY FOR RESISTANCE IN CATTLE TICKS TO ACARICIDES

SUMMARY: A survey was made from late 1976 to late 1977 to determine the extent of resistance to acaricides in the cattle tick Boophilus microplus in Queensland. Questionnaires and requests for samples of ticks were forwarded to more than 900 randomly selected stock owners in the tick infested area which had been divided into 4 regions, Far North, Coastal North, Coastal Central and South East. The response measured by the number of tick samples tested was 43%. The prevalence of resistance to organophosphorus (OP) compounds was highest in South East where 96% of the farms had OP-resistant ticks and 95% had the Biarra strain and lowest in Far North where 12% of the farms had OP-resistant ticks and 10% had the Biarra strain which was the predominant one in all regions. The highest percentages of the Ridgelands strain (35%) and the Tully strain (30%) occurred in Coastal North. The South East had the highest percentage of Mt Alford (30%).
Although chlorinated hydrocarbons were banned for use in control of ticks in 1962, 49% of the farms in South East had some ticks resistant to dieldrin but in Far North it was only 2%. DDT-resistant ticks, which because of cross resistance to synthetic pyrethroids will affect the future use of this group, were present on 8% of the farms in both Coastal North and Coastal Central and 3% in South East and Far North.
No resistance to the amidines, chlordimeform, chloromethiuron or amitraz was found.     

Histopathology of attachment sites of Amblyomma variegatum and Rhipicephalus appendiculatus on zebu cattle of varying resistance to ticks.

Mature, unimproved East African Zebu cattle (Bos indicus) naturally exposed to ticks from birth were ranked for resistance to ticks by repeated whole-body counts of infesting ticks. Four cattle of high resistance, four of low resistance and two of intermediate resistance were artificially infested with nymphs of Amblyomma variegatum and Rhipicephalus appendiculatus. Two Friesian cattle (Bos taurus) with less exposure to ticks were similarly infested. Biopsies of tick attachment sites were examined histologically. All attachment sites showed acute inflammatory lesions, and sites of both tick species on high resistance cattle showed delayed hypersensitive reactions with intra-epidermal pustulation and significant increases in the numbers of granulocytes. The predominant cells infiltrating attachment sites on high resistance cattle were eosinophils with A. variegatum and neutrophils with R. appendiculatus. Such differences need to be taken into account in developing immunological tests for selecting cattle for resistance but there are sufficient common features of reactions to the two infesting tick species to justify dermal hypersensitivity tests.     

Ticks on livestock in Barbados

Cattle, sheep, goats, and horses in most areas of Barbados were examined for ticks. About 68% of the cattle, mostly Holstein dairy cows, and 23% of the sheep, mostly purebred or crossbred black belly sheep, were infested with the southern cattle tick, Boophilus microplus (Canestrini), a vector of bovine babesiosis and anaplasmosis. About 13% of the horses were infested with the tropical horse tick, Anocentor nitens (Neumann), a vector of equine babesiosis. Cattle owners used a variety of acaricides as dusts, sprays, or washes on their cattle for the control of ticks.     

Epidemiological observations of Zimbabwean theileriosis: disease incidence and pathogenicity in susceptible cattle during Rhipicephalus appendiculatus nymphal and adult seasonal activity.

Fifty-nine Hereford cattle susceptible to tick-borne diseases were used as tracer animals to assess the tick challenge and pathogenicity of Theileria parva under field conditions in Zimbabwe. They were moved periodically in groups of five to three commercial farms (one group consisted of four) during seasons of Rhipicephalus appendiculatus nymphal and adult activity. All tracer cattle were herded together with the farm cattle but were not dipped. The nymphal tick counts were high on two of the farms (up to 2000 per animal) but were very low on the third farm (less than ten per animal). On the three farms, 19 out of 24 (76%) tracers had patent Theileria schizonts. There was a range of clinical manifestations of theileriosis with acute and fatal infections occurring on one farm. The adult R. appendiculatus infestations during the wet season numbered 120-800 per animal on the three farms. The disease transmitted by the adults was very pathogenic on the three farms; 30 out of 35 (86%) had severe theileriosis infections. Cattle, which survived the nymphal diseases challenge, showed various degrees of immunity to subsequent T. parva challenge transmitted by adult ticks. Therefore, 13 out of 18 (72%) of these cattle had a second disease episode and the case fatality rate on the three farms was 46%. The factors which determined the epidemiological status of Theileria challenge on the farms, such as the farming systems and presence of wild animals, are discussed.     

In vitro and in vivo evaluation of deltamethrin and amitraz mixtures for the control of Rhipicephalus (Boophilus) microplus (Acari: Ixodidae) in New Caledonia

Acaricide resistance is a major problem that hinders the control of the tropical cattle tick, Rhipicephalus (Boophilus) microplus (Canestrini), in many parts of the world where cattle production continues to suffer severe economic losses to tick infestation. Deltamethrin and amitraz have been used alone to control R. microplus in New Caledonia for the past decade, and tick populations have developed resistance to both acaricides. A study was conducted to evaluate the effectiveness of deltamethrin and amitraz mixtures, through in vitro laboratory bioassays and in vivo on-animal efficacy trials, for the control of resistant R. microplus on cattle at two dairy farms in New Caledonia. Results of laboratory bioassays using modified larval packet tests (LPT) revealed up to 16.59-fold resistance to deltamethrin, and up to 5.86-fold resistance to amitraz. Significant synergism was observed when amitraz was used as a synergist in deltamethrin bioassays. Amitraz significantly increased deltamethrin toxicity to tick larvae, while deltamethrin was much less effective on amitraz toxicity. Synergism of amitraz by deltamethrin only occurred when the deltamethrin concentration was relatively high. Results of on animal efficacy trials of deltamethrin and amitraz alone and mixtures of both at different concentrations revealed a similar pattern of synergism. Adding amitraz to a deltamethrin formulation led to dramatic increases of percent reduction of both immature and adult ticks. In contrast, adding deltamethrin to an amitraz formulation did not increase control efficacy. Results from this study may lead to the adoption of an acaricide mixture strategy for the control of pyrethroid-resistant R. microplus in New Caledonia and elsewhere.