Progress Towards the Eradication of Amblyomma variegatum from the Caribbean

Progress in the Caribbean Amblyomma Program (CAP) is reviewed since its inception in 1995 when regional eradication activities for Amblyomma variegatum ticks were initiated using Bayticol pour-on. Technical achievements in the various islands were slow initially, and showed a wide diversity in attaining eradication targets. St. Kitts, considered as a model program, eliminated the tropical Bont tick (TBT) from most of the island in less than 3 years. However, the elimination of very low numbers of residual adult TBT that persisted in three ‘hot-spots’ took another 3 years. A similar problem was faced in St. Lucia, although the approach there was to cull the wild cattle in the last remaining hot-spot. Both islands were certified as ‘Provisionally free from TBT’ in November 2001. In 2002, Anguilla and Montserrat attained the same status, and Barbados and Dominica also qualify for certification. Certification is based on strict, three-monthly, surveillance criteria: essentially, two successive TBT-free surveillance rounds on a statistically acceptable sample of the livestock population. Within the responsibility of the CAP, three islands remain TBT-infested, Antigua, Nevis, and St. Martin/St. Maarten. Nevis is of concern because it has been operational since late 1995, and appears unable to resolve the problem of treating stray small ruminants. Current funding should be adequate to complete eradication on Nevis and in St. Martin/St. Maarten. The major outstanding challenge is Antigua. In comparison to other islands they have the largest number of livestock, and have much larger populations than originally reported: sheep and goats are 4–5-fold higher, and cattle are 1.35-fold higher. The cost for Bayticol is, therefore, almost double and an additional US$ 1.5 million is required for that commodity alone. This revised version was published online in July 2006 with corrections to the Cover Date.     

Rickettsia africae in Amblyomma variegatum and domestic ruminants on eight Caribbean islands

We used PCRs with omp A primers to determine if spotted fever group rickettsiae occurred in Amblyomma variegatum from 6 Caribbean islands. Positive amplicons were obtained from ticks from the U.S. Virgin Islands (9/18; 50%), Dominica (39/171; 30%), Montserrat (2/5; 40%), Nevis (17/34; 50%), St. Kitts (46/227; 20%), and St. Lucia (1/14; 7%). Sequences for a convenience sample of reaction products obtained from A. variegatum on St. Kitts (7), American Virgin Islands (4), Montserrat (2), and St. Lucia (1) were 100% homologous with that of Rickettsia africae , the agent of African tick-bite fever. To determine if transmission of R. africae occurred, we used Rickettsia rickettsii antigen in IFA tests and found positive titers (≥ 1/80) with sera from cattle, goats, and sheep from Dominica (24/95 [25%], 2/136 [2%], 0/58 [0%]), Nevis (12/45 [27%], 5/157 [3%], 0/90 [0%]), St. Kitts (2/43 [5%], 1/25 [4%), 1/35 [3%]), and St. Lucia (6/184 [3%] cattle), respectively. No seropositive animals were found in Grenada (0/4, 0/98/, 0/86), Montserrat (0/12, 0/26, 0/52), or Puerto Rico (0/80 cattle). Our study indicates that R. africae and African tick-bite fever are widespread in the Caribbean.     

Attractivity of cattle infested by Amblyomma variegatum (Acari: Ixodidae) for conspecific adult ticks from the field in Guadeloupe

Four field experiments were conducted with creole cattle in Guadeloupe to study the attractivity for adult ticks of animals previously infested with males of Amblyomma variegatum producing aggregation attachment pheromones (AAPs). In each experiment, two groups of cattle were used; the first group was infested with adult A. variegatum and the second group was kept free of ticks and pheromones by acaricidal treatment and by washing the hair and skin of previously infested sites. Both groups were then exposed to natural infestation by ticks present in the environment or by ticks bred in the laboratory and released on the pasture. The number of additional ticks infesting animals were compared between the treated and control groups. There were no significant differences between both groups in the numbers of new male ticks infesting the animals during the first days following the beginning of the experiment. Thereafter, comparison of the slopes of infestation indicated that infested cattle were no more attractive for male ticks than uninfested hosts. Uninfested cattle were spontaneously and rapidly infested by male ticks. It is concluded that carbon dioxide or another host component was a major attractant for A. variegatum males. Female ticks attached on predilection sites exclusively in the presence of AAP-producing males, i.e. 5–10 days after the attachment of the first males. Based on these results we postulate that slow-release devices impregnated with pheromones have no additional attractive effect on A. variegatum males under Caribbean conditions. However, they may be useful in inducing the attachment of females, but only in the first days preceding the arrival of pioneer AAP-producing males. © Rapid Science Ltd. 1998     

Efficacy of tags impregnated with pheromone and acaricide for control of Amblyomma variegatum

The efficacy of tags impregnated with pheromone and acaricide for control of Amblyomma variegatum on cattle in Guadeloupe was determined for a 13-week trial. Comparisons were made between untreated cattle and cattle with tags containing either pheromones alone (o-nitrophenol, methyl salicylate, 2,6-dichlorophenol and phenylacetaldehyde), pheromones plus acaricide (cyfluthrin or deltamethrin), or acaricide alone. Tags were fastened to cattle both on collars on the neck and with adhesive to tail hairs, with over 98 and 90% of tags retained, respectively, during the trial. By the end of the trial, tick infestations increased on untreated cattle (311.7%) and cattle with pheromone tags (154.8%) but decreased on cattle with pheromone/cyfluthrin (–45.0%), cyfluthrin (–42.8%), pheromone/deltamethrin (–68.7%) and deltamethrin tags (–87.6%). Cattle with pheromone tags had greater proportions of ticks on the hind regions (81%) compared to untreated cattle (62.5%) and on the front regions (18.2%) compared to untreated cattle (8.2%) indicating that ticks aggregated in response to pheromones. Analysis of hair samples by gas chromatography indicated that, within 2 days of placement, cyfluthrin spread rapidly over the body from the tags. Both cyfluthrin and deltamethrin were present in detectable levels on all eight body regions examined throughout the trial. Analysis of the tags indicated that they all still contained pheromone components after 13 weeks under field conditions.     

Footbath acaricide treatment to control cattle infestation by the tick Amblyomma variegatum

Previous studies have shown that about 90% of adult Amblyomma variegatum Fabricius (Acari: Ixodidae) picked up daily by grazing cattle are still attached to the interdigital areas in the evening, when the animals return from pasture. It was therefore postulated that a targeted treatment, designed to kill the ticks attached to the feet, would limit infestation of the predilection sites. Footbaths filled with various pyrethroid formulations were used over 3 years, at the beginning of the rainy season (from mid-May to the end of July), to assess the efficacy of such a control method. It proved efficient in preventing the ticks from attaching to the predilection sites. Although five to 12 A. variegatum adults attached to each treated animal daily, and although the tick burden of the predilection sites of control cattle increased each day by four to 10 ticks, the average infestation of the predilection sites of treated cattle that were initially highly infested (over 100 ticks/animal) continuously decreased to reach a level of about 10-30 ticks/animal after 6-8 weeks of treatment. In herds with a lower initial tick burden (40-70 ticks/animal) this level was obtained within 2-3 weeks and the mean infestation subsequently remained consistently low. Footbath treatment carried out every other day during the adult peak infestation period should therefore greatly limit losses due to ticks. This method was appreciated by traditional livestock farmers, essentially because it is not time-consuming and because it requires only c. 200 mL aqueous formulation per animal at each passage. The cost of the acaricide needed to treat one animal during the peak infestation period was assessed at c. euro 0.20. This control method might also have an impact on some species of tsetse flies and mosquitoes, thereby contributing to trypanosomiasis and malaria control.     

Morphogenetic diapause in Amblyomma variegatum (Acari: Ixodidae)

In southern Africa, Amblyomma variegatum Fabricius is characterized by a strict seasonal activity. Experiments were carried out to determine whether a diapause mechanism regulates this seasonality. Engorged A.variegatum females were exposed to controlled laboratory conditions or natural field conditions at different times of the year. Females exposed in a natural environment in September-October (short day) had significantly longer pre-oviposition periods than females exposed from November to March. The season in which the previous instar fed had no apparent effect on the engorgement or pre-oviposition periods of the females. Furthermore, artificial changes in photoperiod during and after female engorgement had no significant effects on pre-oviposition periods. It is tentatively concluded that the unfed female is the responsive stage to photoperiodic changes which induce diapause. Diapause could be terminated and oviposition induced by exposing females to a short period of chilling (18 degrees C for 48 h). It is concluded that a morphogenetic diapause mechanism exists in A.variegatum, which is probably induced by short day responses and terminated following rainfall and a concomitant decrease in soil temperature. The diapause, which occurs in females which fed early in the season, causes a delay in oviposition and therefore effectively synchronizes the life-cycle to ensure that eggs and larvae occur at a climatically favourable period.     

Distributions of the vectors of heartwater, Amblyomma hebraeum and Amblyomma variegatum (Acari: Ixodidae), in Zimbabwe

The tick vectors of heartwater (Cowdria ruminantium infection) in Zimbabwe, Amblyomma hebraeum and Amblyomma variegatum, historically were believed to be confined to the low-lying regions of the south and north-west of the country. However, country-wide surveys performed in 19751980 and 19881991 demonstrated that both species were also established in western parts of the highveld plateau and had started to encroach on the predominantly heartwater-free central and eastern highveld regions. To determine the current distributions of both the vectors and evaluate the potential threat of heartwater to animals in the highveld, a survey of ticks infesting cattle was performed in 1996 at 2994 locations in small-holder and large-scale commercial farming areas throughout Zimbabwe. Amblyomma hebraeum was collected at 1329 locations, A. variegatum at 72 locations and both A. hebraeum and A. variegatum at 13 locations. The results demonstrated that A. hebraeum was present, as previously recorded, throughout the southern half of the country and appeared to have undergone further limited spread into the central and eastern highveld regions. Only the northern-most region of the country appeared to be free of this species. Amblyomma variegatum was collected mainly in the north-west, as previously recorded, but was also found at isolated locations across the central highveld region and along the eastern border with Mozambique. This species was, however, still absent from the southern half and the northern-most regions of the country. An overlap of the distributions of the two species existed within a zone along the southern and eastern regions of the distribution of A. variegatum. These results suggest that the vectors of heartwater are spreading and threaten to introduce heartwater into intensive livestock-producing regions of the country.Exp Appl Acarol 22: 725740 © 1998 Kluwer Academic Publishers     

Model for survival of unfed female Amblyomma variegatum (Acari: Ixodidae) in Kenya

Experiments were conducted in threc different ecozones in Kenya to measure the survival of unfed females of the ixodid tick Amblyomma variegatum under field conditions. Throughout the study, temperature, relative humidity, and rainfall were recorded every hour at each of the threc sites. Using a multiple linear regression procedure, a model was produced to describe survival of unfed females in terms of high temperature at ground level (cumulative day-degrees >34°C) and age. The various ways in which age and temperature can influence survival are discussed.     

Acaricide susceptibility ofAmblyomma variegatum (Acari: Ixodidae) from Puerto Rico and Guadeloupe

Base-line data on acaricide susceptibility of larvae and of larvae and nymphs of the tropical bont tick,Amblyomma variegatum (Fabricius), from Puerto Rico and Guadeloupe were collected on 14 and six compounds, respectively. Nymphs 7–8 weeks old from Puerto Rico and Guadeloupe, and larvae 3–4 weeks old from Guadeloupe, were exposed for 24 or 48 h to residues of acetone dilutions of technical-grade or commercial formulations of acaricides on disposable glass pipettes. Effectiveness of the acaricides in killing nymphs (lc ₅₀) from Guadeloupe and Puerto Rico ranged from 0.0001% for deltamethrin to 0.269% for malathion. For Puerto Rico, the susceptibility to toxaphene and amitraz, but not to dioxathion, increased with increasing age of nymphs. Nymphs from Guadeloupe were 9.5 times more susceptible to chlorpyrifos than nymphs from Puerto Rico. The susceptibility of larvae from Guadeloupe to acaricides tested ranged fromlc ₅₀ 0.000012% for deltamethrin to 0.0141% for ethion. There was no evidence of development of acaricide resistance in ticks from Guadeloupe or Puerto Rico.     

Vector capacity of Rhipicephalus appendiculatus and Amblyomma variegatum for Thogoto and Dhori viruses

ABSTRACT. Experimental studies were undertaken to ascertain the vector potential of Rhipicephalus appendiculatus Neumann and Amblyomma variegatum Fabricius for Thogoto (THO) and Dhori (DHO) viruses, candidate members of the Orthomyxoviridae. In the first set of experiments, ticks were infected orally by feeding on viraemic hamsters. THO virus replicated in R.appendiculatus and A.variegatum, persisted trans-stadially in both tick species, and was transmitted to susceptible hosts during feeding. In contrast, both R. appendiculatus and A. variegatum were refractory to per os infection by DHO virus. In the second set of experiments, engorged R. appendiculatus and A. variegatum nymphs were parenterally inoculated with DHO virus. The virus persisted transstadially in both tick species and was transmitted by bite to susceptible hosts. These results indicate that the midgut acts as a barrier to per os infection of R.appendiculatus and A. variegatum by DHO virus. However, when this barrier is bypassed, i.e. by parenteral inoculation, both R.appendiculatus and A.variegatum can serve as efficient vectors of DHO virus.     

Field sampling of the tick Amblyomma variegatum (Acari: Ixodidae) on pastures in Guadeloupe; attraction of CO2 and/or tick pheromones and conditions of use

Field experiments were conducted in Guadeloupe to evaluate the attraction of different chemicals on a Guadeloupean strain of Amblyomma variegatum. Such tick sampling techniques may help to assess the size of the tick population on pastures before, during and after an attempted tick eradication campaign. CO2 was absolutely necessary to activate ticks, including the larvae. In addition to activation, our data suggested that CO2 itself is attractive for ticks. The reactivity of the ticks varied greatly with the period of the year, the adults being insensitive to CO2 alone or combined with pheromones for a period extending from December to June. For the whole period, the total captures were equal for the males and females, but the males were relatively more sensitive at the beginning of the season of activity (June–July), while the females dominated, but not significantly, for the rest of the period. There was also a gradient of reactivity depending on the distance from the trap; 28% of the ticks were captured at 0.5 m from the CO2 traps and 1.2% at a distance of 6 m. The ticks arrived rapidly at the trap. After 1 h of trapping, 55, 68 and 81% of the active nymphs, males and females, respectively, had reached the trap. Synthetic pheromones (methyl salicylate, o-nitrophenol and nonanoic acid in the proportions 1:2:8) or freshly detached males added to CO2, greatly increased the efficacy of the CO2 traps. Depending on the concentration of the pheromones, the attraction was four to 70 times higher than with CO2 alone. The effect depended on the sex of the ticks, males being attracted in a greater proportion by small amounts of pheromones, while the females were activated by higher doses. From a practical point of view, sampling should be conducted from June to December, with CO2 and high concentrations of pheromones, the traps being in operation for approximately 1 h at least, the area concerned by effluents being located downwind and covering an area of approximately 36 m2.     

Predators of the tickAmblyomma variegatum (Acari: Ixodidae) in Guadeloupe,French West indies

Some vertebrate species in Guadeloupe are predators of free or parasitic stages ofAmblyomma variegatum (Fabricius). Among birds, 1.9% of the 421 identified animals found in the stomachs of grackles (Quiscalus lugubris), 1.6% of the 364 animals found in the stomachs of free-ranging chickens, and 0.3% of the 4642 animals found in the stomach of cattle egrets (Bubulcus ibis) wereA. variegatum ticks. The most efficient predator of ticks was the tropical fire ant,Solenopsis geminata, which was observed to only attack engorged stages of ticks. An average of 8% of the 564 individual ticks or batch of ticks released on the ground, mainly in a grass environment, were attacked by this ant species. Mice (Mus musculus) and mongooses (Herpestes auropunctatus) feed on engorged nymphs and female ticks. None of the 15 strains of entomophagous nematodes, generaSteinernema andHeterorhabditis, experimentally put in contact with engorged larvae and nymphs were able to parasitize them. In the laboratory, the insectsMegaselia scalaris (Diptera) andTineola sp. (Lepidoptera) were occasionally found to feed on engorged and unfed ticks, respectively.     

Footrot control and eradication (elimination) strategies

Footrot is a major welfare problem and cause of economic losses in most sheep-keeping countries. The main reasons why this is the case are reviewed, particularly the effects of weather, environment and husbandry methods on the spread of the disease. The main methods for control and elimination (eradication) are briefly described. Attention is drawn to the fact that control is particularly difficult in temperate countries without predictable non-transmission periods.     

Crimean-Congo haemorrhagic fever virus replication in adult Hyalomma truncatum and Amblyomma variegatum ticks

The kinetics of the replication of the Crimean-Congo haemorrhagic fever virus (CCHFV) was studied in intra-anally inoculated adult Hyalomma truncatum and Amblyomma variegatum ticks. The virus was re-isolated by suckling mouse inoculation and revealed by antigen capture with ground ticks and indirect immunofluorescence of haemolymph. The virus was detected in ticks in the first hours post-inoculation (p.i.) and its replication was observed from 36 h p.i. onwards. Virus titre reached a maximum within 3–5 days then decreased slowly to a level of at 2 log LD₅₀/ml for several months until the end of observations.Several specific, non-identified factors seem to favour CCHFV replication in H. truncatum. Long-term virus persistence seems to occur in CCHFV-infected adult ticks.     

Novel control strategies for mosquito-borne diseases

Mosquito-borne diseases are an increasing global health challenge, threatening over 40% of the world''s population. Despite major advances in malaria control since 2000, recent progress has stalled. Additionally, the risk of Aedes-borne arboviruses is rapidly growing, with the unprecedented spread of dengue and chikungunya viruses, outbreaks of yellow fever and the 2015 epidemic of Zika virus in Latin America. To counteract this growing problem, diverse and innovative mosquito control technologies are currently under development. Conceptually, these span an impressive spectrum of approaches, from invasive transgene cassettes with the potential to crash mosquito populations or reduce the vectorial capacity of a population, to low-cost alterations in housing design that restrict mosquito entry. This themed issue will present articles providing insight into the breadth of mosquito control research, while demonstrating the requirement for an interdisciplinary approach. The issue will highlight mosquito control technologies at varying stages of development and includes both opinion pieces and research articles with laboratory and field-based data on control strategy development.This article is part of the theme issue ‘Novel control strategies for mosquito-borne diseases''.     

The ultrastructure of somatic neuromuscular junctions in the tick Amblyomma variegatum (Fabr.)

Summary Injections of large doses of horseradish peroxidase (HRP) into the telencephalon of the squirrel fish (Holocentrus rufus) revealed the first anatomical evidence for a visual thalamo-telencephalic projection in a teleost. The central optic nucleus of the thalamus appears to be the only visual thalamic nucleus projecting to the telencephalon in this species. Since the central optic nucleus has a large tectal input but not a direct one from the retina, it is suggested that a retino-geniculo-telencephalic pathway does not exist in this species. Acknowledgements. The author is grateful to Drs. J. Maldonado, Dietrich Meyer and Henning Scheich for encouragement and support in this endeavor. The study was supported by: National Institutes of Health grant EY-02014 and EY-03264, a NIH grant to Dr. José del Castillo and the German Science Foundation (SFB 45)     

Performance of a Prototype Baited-trap in Attracting and Infecting the Tick Amblyomma variegatum (Acari: Ixodidae) in Field Experiments

Investigations were commenced to study the potential use of the fungi, Beauveria bassiana, Metarhizium anisopliae, and the attraction-aggregation-attachment pheromone (AAAP) for the control of Ambloyomma variegatum as an environmentally friendly technology. The objective of the study was to develop and test a device, which could be used for pheromone and carbon dioxide delivery and infection of ticks with the fungi in an attempt to control the tick populations in the vegetation. Using a pheromone-baited device treated with the fungi mixture, 79% of the ticks released were attracted and exposed to the fungi and of these, 78% died during incubation in the laboratory. In another set of experiments, of the released ticks that were similarly exposed to fungi using the pheromone-baited device and left in the vegetation, 33.8% were recovered compared to recoveries of between 76 and 84% in the controls. These results were significantly different at the 5% level, an indication that the pheromone/fungi mixtures had significant effect in reducing the tick population in the field.     

Factors affecting the distributions of the ticks Amblyomma hebraeum and A. variegatum in Zimbabwe: implications of reduced acaricide usage

The ticks Amblyomma hebraeum and A. variegatum are the main vectors of heartwater, a disease of ruminants caused by Cowdria ruminantium, in the agricultural areas of Zimbabwe. At present, A. hebraeum is widely distributed in the dry southern lowveld, and occurs in at least seven foci in the higher rainfall highveld. Amblyomma variegatum occurs in the Zambezi valley and surrounding dry lowveld areas in the northwest. The distribution of A. hebraeum has changed considerably over the past 70 years, while that of A. variegatum appears to have remained fairly static. The distribution patterns of both species in Zimbabwe display anomalous features; the ticks occur in areas of lowest predicted climatic suitability for survival and development and in areas where the densities of cattle, the most important domestic host, are lowest. The only factor favouring the survival of the species in the lowveld habitats in which they occur is the presence of alternative wildlife hosts for the adult stage. Their absence from more climatically favourable highveld habitats appears to have been the result of intensive acaricide treatment of cattle over a long period and a historic absence of significant numbers of wildlife hosts. Eradication of A. hebraeum and A. variegatum by intensive acaricide treatment of cattle can be achieved in the absence of significant numbers of alternative hosts, because of the long attachment and feeding periods of the adults of these tick species. However, eradication becomes impossible when alternative hosts for the adult stage are present, because a pheromone emitted by attached males attracts the unfed nymphal and adult stages to infested hosts. The unfed ticks are not attracted to uninfested hosts, such as acaricide-treated cattle.Regular acaricide treatment of cattle is expensive and so, for economic reasons, the Government of Zimbabwe is no longer enforcing a policy of strict tick control. It is likely that reduced tick control will result in the spread of Amblyomma ticks to previously uninfested areas. Added to this, recent introductions of various wildlife species to highveld commercial farming areas have created conditions in which the ticks could become established in higher rainfall areas. Amblyomma hebraeum is more likely to spread than A. variegatum, because its adults parasitize a wider range of wildlife hosts (warthogs, medium to large-sized antelope, giraffe, buffalo and rhinoceros), whereas adults of A. variegatum appear to be largely restricted to one wildlife species (buffalo) in Zimbabwe, the distribution of which is now confined to very limited areas of the country, as part of foot and mouth disease control measures. A model to predict the rate of spread of A. hebraeum through the highveld is described.Possible control options for dealing with the spread of Amblyomma ticks and heartwater to previous unaffected highveld areas, include (1) continuation of intensive acaricide treatment of cattle to prevent the spread, (2) establishment of a buffer zone of intensive tick control around affected areas to contain the spread and (3) allow the spread to occur and control heartwater by means of immunization. An economic analysis to determine the costs and benefits of the control options, which takes into account the development of Amblyomma-specific tick control technologies and improved heartwater vaccines, is recommended.Deceased.