Biology and control of ticks infesting dogs and cats in North America

At least 15 different infectious agents and diseases are transmitted or produced by ticks parasitizing dogs and cats in North America. Those infectious agents and diseases are distributed among eight different tick species that commonly infest dogs and cats. All but one of these species are hard ticks with a three-host development cycle in which each motile stage (larva, nymph, and adult) feeds on a different host after molting. Tick species, disease occurrence, and peak activity of each tick life stage can vary dramatically depending on geographic and climatic conditions. Gaining an understanding of tick distribution, tick ecology, and seasonal occurrence of different tick life stages can help with the management of tick infestations and reduce the incidence of tick-transmitted diseases in dogs and cats. Control should be based on an understanding and management of ecologic factors responsible for tick infestations and selection of appropriate acaricides. Occasionally, topical acaricides will appear not to have prevented tick infestations. This lack of control may be real or perceived based on reinfestation rates and/or pet owner expectations of product performance.     

The effect of East Coast fever immunisation and different acaricidal treatments on the productivity of beef cattle

A trial was performed on a farm in the Coast Province of Kenya to study the effects of East Coast fever immunisation and different acaricidal treatments on the productivity of immunised and unimmunised beef cattle. Eighty cattle were immunised against Theileria parva parva (Marikebuni) by the infection and treatment method and a similar group was left as an unimmunised control. Immunisation had no deleterious effect on the cattle. After immunisation, the immunised and control groups were each subdivided into four groups of 20 and each subgroup was managed under a different tick control regimen. The tick control regimen were, acaricidal spraying twice a week or once every three weeks, the application of acaricide-impregnated ear-tags, and no tick control. During a nine-month exposure period there were 18 cases of East Coast fever among the 80 immunised cattle, three which were severe and the others mild. Among the 80 unimmunised cattle there were 57 cases of East Coast fever, 50 of which were severe. The highest morbidity and mortality occurred in the groups under limited tick control or without tick control. Overall weight gain in the immunised cattle, irrespective of the tick control regimen, was better than the weight gain in the unimmunised groups. Within the immunised groups, the weight gain of the cattle sprayed twice weekly was comparable to the weight gain of the animals with acaricidal ear-tags and was significantly higher than the weight gains in the groups sprayed once every three weeks or with tick control. Preliminary cost/benefit analysis showed that it was uneconomical to maintain unimmunised cattle under limited or no tick control.(ABSTRACT TRUNCATED AT 250 WORDS)     

Life cycle of the tick Amblyomma parvum Aragão, 1908 (Acari: Ixodidae) and suitability of domestic hosts under laboratory conditions

Amblyomma parvum is a widespread Neotropical tick found on several domestic animals and is known to harbor a Rickettsia species of yet unknown pathogenicity. However its life cycle on, and suitability of, several of these hosts has not been described. In this work the biology of A. parvum is presented when fed on seven domestic hosts (chicken, dog, rabbit, horse, guinea pig, cattle and goat). The complete life cycle of the tick varied from 96.8 to 102 days. Highest engorgement weight of larvae was from ticks fed on horses and that of nymphs from guinea pigs. Highest larval yield was from guinea pigs and that of nymphs from horses. Engorged female and egg mass weights, yield and conversion of female weight to eggs rates were the highest in dog ticks and the lowest in goat ticks. The highest egg hatching rate was seen in ticks from dogs and the lowest in ticks from cattle. Overall it was seen that dogs were the best host for adult A. parvum ticks, and guinea pigs for immatures. Horses were also shown to be a good host for all tick stages. It can thus be affirmed that A. parvum is a host generalist tick, and its distribution is probably determined by environmental requirements rather than by hosts.     

Disease at the wildlife-livestock interface: Acaricide use on domestic cattle does not prevent transmission of a tick-borne pathogen with multiple hosts

Several prominent and economically important diseases of livestock in East Africa are caused by multi-host pathogens that also infect wildlife species, but management strategies are generally livestock focused and models of these diseases tend to ignore the role of wildlife. We investigate the dynamics of a multi-host tick-borne disease in order to assess the efficacy of tick control from an ecological perspective. We examined the efficacy of a widespread measure of tick control and developed a model to explore how changes in the population of ticks due to control measures on cattle impact dynamics of Theileria parva infection in a system with two primary host species, cattle and Cape buffalo (Syncerus caffer). We show that the frequency of acaricide application has a significant impact on the tick population both on the host and in the environment, which can greatly reduce the pathogen load in cattle. We also demonstrate that reducing the tick population through cattle-related control measures is not sufficient to diminish disease transmission in buffalo. Our results suggest that under current control strategies, which target ticks on cattle only, T. parva is likely to remain a significant problem in East Africa, and require the continued use of acaricides, which has significant economic and ecological consequences.     

Ecological preferences of exophilic and endophilic ticks (Acari: Ixodidae) parasitizing wild carnivores in the Iberian Peninsula

Ticks parasitizing wild carnivores and the tick-borne pathogens (TBPs) that they transmit may affect domestic carnivores and humans. Thus, investigating the role of wild carnivores as tick hosts is of relevance for understanding the life cycle of ticks in natural foci and the epidemiology of TBPs shared with domestic animals and humans. Therefore, the main objective of this study was to determine the ixodid tick fauna of wild carnivores in Peninsular Spain and the environmental factors driving the risk of wild carnivores to be parasitized by ixodid ticks. We hypothesized that the adaptation of tick species to differing climatic conditions may be reflected in a similar parasitization risk of wild carnivores by ticks between bioclimatic regions in our study area. To test this, we surveyed ixodid ticks in wild carnivores in oceanic, continental-Mediterranean, and thermo-Mediterranean bioclimatic regions of Peninsular Spain. We analyzed the influence of environmental factors on the risk of wild carnivores to be parasitized by ticks by performing logistic regression models. Models were separately performed for exophilic and endophilic ticks under the expected differing influence of environmental conditions on their life cycle. We found differences in the composition of the tick community parasitizing wild carnivores from different bioclimatic regions. Modelling results partially confirmed our null hypothesis because bioclimatic region was not a relevant factor influencing the risk of wild carnivores to be parasitized by exophilic ticks. Bioclimatic region was however a factor driving the risk of wild carnivores to be parasitized by endophilic ticks. Spanish wild carnivores are hosts to a relevant number of tick species, some of them being potential vectors of pathogens causing serious animal and human diseases. Information provided herein can be of help to understand tick ecology in Spanish wildlife, the epidemiology of tick-borne diseases, and to prevent the risks of TBPs for wildlife, domestic animals, and humans.     

Cellular immunity against Theileria parva and its influence on parasite diversity

Theileria parva, a tick-borne parasite of African cattle, causes a fatal disease known as East Coast fever. Cattle that recover from the disease develop strong parasite-specific MHC-class I-restricted cytotoxic T-lymphocyte responses. Protection can be transferred between immune and na?ve calves in the CD8+ T cell fraction emanating from a responding lymph node. In vitro studies suggest that this response requires input from activated CD4+ T cells. The T parva life cycle involves developmental stages in mammalian and tick hosts and can lead to a number of different endemic scenarios for the disease. These range from a stable situation with high prevalence of herd infection, but low fatality rates, to a low prevalence/high fatality scenario. The impact on endemic stability is an important consideration for the design of vaccine implementation strategies. For subunit vaccines targeted at T parva schizonts, the principal issue in this regard is whether development of the piroplasm stage is blocked by immunity.     

A simulation model for environmental population densities, survival rates and prevalence of Boophilus decoloratus (Acari: ixodidae) using remotely sensed environmental information.

A simulation model for the African tick Boophilus decoloratus is presented. This model is based on the use of a dynamic life table that is driven by abiotic variables (temperature and vegetation status) remotely sensed (AVHRR sensor of the NOAA series of satellites) over time. The model incorporates temperature-dependent rates of egg production and development, climate-driven density-independent mortality rates, and density-dependent regulation of on-host stages. The model successfully describes both the seasonality and annual variation in the numbers of questing larvae and engorging females observed in eight sites throughout sub-Saharan Africa. Climate data from 1983 to 1999 in 10-day intervals are used as the basic input for modelling the dynamic patterns of activity at four different sites in Africa and to understand how abiotic factors can modulate the long-term life cycle of B. decoloratus.     

Retrospective observations on the geographical relationship between Rhipicephalus appendiculatus and East Coast fever in southern Africa

East Coast fever appeared in southern Africa in 1902, after the importation of cattle from East Africa, and spread throughout the range of Rhipicephalus appendiculatus until it reached the southern limit of distribution of this tick in 1916. It was progressively eradicated by quarantine, destocking, slaughter and frequent dipping in arsenic. The last case occurred in Swaziland in 1960. A computer based system for matching climates and ecology (CLIMEX) revealed a direct correlation between the climatic favorability of different areas for the tick and the time required to eradicate the disease.     

Estimating the distribution and abundance of Rhipicephalus appendiculatus in Africa

The brown ear tick Rhipicephalus appendiculatus is responsible for transmitting the parasite Theileria parva in eastern, central and southern Africa, where it causes East Coast fever, Corridor disease and January disease in cattle. In an effort to assess the impact of these diseases and their control on livestock production in the region, studies are underway to model the factors controlling the distribution of the vector tick. Three recent studies have attempted to quantify and evaluate the variables influencing the distribution of R. appendiculatus, and these are reviewed and discussed in this paper.

The relationship between the distribution of an ecoclimatic index of suitability for the tick, calculated by the model CLIMEX on a 25 km² interpolated climate database for Africa, and of recorded observations of the tick itself, are evaluated. The sensitivity and specificity of the ecoclimatic index were calculated as 70.2–82.6% and 69.4–84.7%, respectively. In addition, addition, satellite-derived mean maximum normalized difference vegetation index (NDVI) for 1987 was assessed as a predictor of R. appendiculatus habitat in eastern Africa, by comparing the distribution of NDVI values in areas in which the presence of the tick has and has not been recorded. Some visual correlation was observed between mean maximum NDVI values of 0.15 or greater and known R. appendiculatus distribution in Kenya. In addition, this range of NDVI values corresponded closely to areas in Ethiopia where the tick does not occur but for which climatic suitability was predicted by CLIMEX. However, these NDVI values should not be considered universal predictors of habitat suitable for R. appendiculatus, and further study of the NDVI with ground truthing of habitat is considered necessary.     

Ixodes frontalis and avian tick-related syndrome in the United Kingdom

OBJECTIVES: This study aimed to characterise tick species responsible for avian tick infestations in the UK, to analyse various risk factors for tick-related syndrome in tick-infested birds and to test samples for the presence of certain tick-transmitted pathogens. METHODS: Ticks, blood, splenic tissue and tick attachment site tissue from birds with attached ticks were requested from veterinarians and wildlife sanctuaries around the UK. Ticks were identified according to standard keys, and samples were analysed via DNA PCR test for Borrelia burgdorferi sensu lato, Babesia species, Bartonella species and Ehrlichia species. RESULTS: Ixodes frontalis was the most commonly identified tick, and an association of adult female I frontalis with tick-related syndrome in birds was demonstrated. Tick infestation was markedly seasonal. I frontalis was found on 32 species of birds. DNA PCR testing was uniformly negative. Of the birds known to have been treated, 75 per cent (nine of 12) survived. CLINICAL SIGNIFICANCE: Tick-related syndrome is a poorly understood syndrome, with sporadic distribution, both geographically and seasonally. This study confirms I frontalis as the most common cause of this syndrome in the UK and identifies some features of the tick life cycle in this country. The benefit of treatment in affected birds is highlighted. Risk factors for tick-related syndrome are examined and preventive strategies discussed.     

Effect of different methods of maintenance on the development and morphology of Babesia bigemina in the gut of Boophilus microplus

The development and morphology of Babesia bigemina in the gut of Boophilus microplus ticks were studied during laboratory maintenance of the babesia by four methods: tick transmission in unsplenectomised (intact) calves; tick transmission in splenectomised calves; syringe passage at intervals of five days or less in splenectomised calves; and syringe passage at intervals of six weeks or more in intact calves. The first method had no apparent effect on the development of B bigemina in ticks compared to that of the original isolate. The other three methods had obvious effects, the most pronounced being increased numbers of babesial forms with processes, particularly during early stages of development. The findings emphasise the importance of maintaining laboratory strains of parasites by natural means in life cycle studies.     

Climate change and the genus Rhipicephalus (Acari: Ixodidae) in Africa

The suitability of present and future climates for 30 Rhipicephalus species in Africa are predicted using a simple climate envelope model as well as a Division of Atmospheric Research Limited-Area Model (DARLAM). DARLAM's predictions are compared with the mean outcome from two global circulation models. East Africa and South Africa are considered the most vulnerable regions on the continent to climate-induced changes in tick distributions and tick-borne diseases. More than 50% of the species examined show potential range expansion and more than 70% of this range expansion is found in economically important tick species. More than 20% of the species experienced range shifts of between 50 and 100%. There is also an increase in tick species richness in the south-western regions of the sub-continent. Actual range alterations due to climate change may be even greater since factors like land degradation and human population increase have not been included in this modelling process. However, these predictions are also subject to the effect that climate change may have on the hosts of the ticks, particularly those that favour a restricted range of hosts. Where possible, the anticipated biological implications of the predicted changes are explored.     

A note on the transmission of Babesia bovis (syn B argentina) by the one-host tick, Boophilus microplus.

Boophilus microplus infected with Babesia bovis were transferred artificially from one splenectomised calf to another during each moult in the parasitic life cycle of the tick. Eggs from the engorged female ticks recovered at the end of the cycle were incubated and the resulting larvae used to infest more splenectomised calves. Babesia bovis was transmitted only by the original larvae used at the commencement of the experiment and it was concluded that the protozoan parasite did not persist in an infective form in the ticks beyond the larval stage.     

Applications of a cell culture system for studying the interaction of Anaplasma marginale with tick cells

A cell culture system for the tick-borne rickettsia Anaplasma marginale offers new opportunities for research on this economically important pathogen of cattle. A. marginale multiplies in membrane-bound inclusions in host cells. Whereas erythrocytes appear to be the only site of infection in cattle, A. marginale undergoes a complex developmental cycle in ticks and transmission occurs via the salivary glands during feeding. We recently developed a cell culture system for A. marginale using a cell line derived from embryos of Ixodes scapularis. Here we review the use of this cell culture system for studying the interaction of A. marginale with tick cells. Several assays were developed using the A. marginale/tick cell system. An adhesion assay was developed for the identification of proteins required by A. marginale for adhesion to tick cells. The effect of antibodies against selected major surface proteins in inhibiting A. marginale infection was tested in an assay that allowed further confirmation of the role of surface proteins in the infection of tick cells. A drug screening assay for A. marginale was developed and provides a method of initial drug selection without the use of cattle. The culture system was used to test for enhancing effects of tick saliva and saliva components on A. marginale infection. The tick cell culture system has proved to be a good model for studying A. marginale-tick interactions. Information gained from these studies may be applicable to other closely related tick-borne pathogens that have been propagated in the same tick cell line.     

The Impact of Tick Control on the Productivity of Indigenous Cattle under Ranch Conditions in Uganda

The impact of tick control on the productivity of indigenous cattle was investigated in three herds of East African shorthorn Zebu and Zebu×Ankole (Nganda) breeds, maintained under three tick management systems practised under ranch conditions: twice-a-week dipping, once-a-month dipping, and no tick control. The milk production and the growth rates of pre-weaned and post-weaned calves were monitored once a month over a period of 34 months. Milk offtake was 23% higher during the heavy rainy seasons than in the long dry seasons. Twice-a-week dipping increased the milk offtake by 21% in the second year of study and prolonged the duration of lactation in cows. Similarly, twice-a-week dipping increased the pre-weaning growth rate by 39% in the second year of study, but had no significant effect on the post-weaning growth rate. Generally, the growth rate of calves were greatest during the heavy rainy seasons and least during the long dry seasons. These results provide a basis for assessing the losses under different tick management systems in the various production systems.     

Babesia and its hosts: adaptation to long-lasting interactions as a way to achieve efficient transmission

Babesia, the causal agent of babesiosis, are tick-borne apicomplexan protozoa. True babesiae (Babesia genus sensu stricto) are biologically characterized by direct development in erythrocytes and by transovarial transmission in the tick. A large number of true Babesia species have been described in various vertebrate and tick hosts. This review presents the genus then discusses specific adaptations of Babesia spp. to their hosts to achieve efficient transmission. The main adaptations lead to long-lasting interactions which result in the induction of two reservoirs: in the vertebrate host during low long-term parasitemia and throughout the life cycle of the tick host as a result of transovarial and transstadial transmission. The molecular bases of these adaptations in vertebrate hosts are partially known but few of the tick-host interaction mechanisms have been elucidated.     

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.     

A weather-based prediction model for the life-cycle of the sheep tick, Ixodes ricinus L.

The incidence of sheep tick activity depends not only on the climatic conditions within the tick habitat, but also on the rates of fecundity, development, activity, engorgement and mortality of each stage of the life-cycle. Use of existing experimental results on the effect of these factors enables a model of the life-cycle to be formulated for the purpose of predicting the occurrence of tick activity in the field where climatic conditions vary. The components of such a model are described and the predicted results compared with field studies carried out in Ireland. It is hoped that the model presented will show the potential of formulating a system for predicting tick activity. Such a system could be used to enhance the control of tick-transmitted diseases.     

Treatment of field cases of East Coast fever with halofuginone lactate

In a series of field trials 48 (55%) of 88 field cases of East Coast fever treated with halofuginone lactate recovered and survived and 40 (45%) died while 31 (86%) of 36 untreated control animals died of East Coast fever and five (14%) recovered. For cases diagnosed and treated early a 100% recovery rate was achieved. A single dose at 1.2 mg/kg body weight per os was adequate. Recovery rate was only 36% for cases diagnosed and treated at an advanced stage. Such cases required two doses at 1.2 mg/kg or a combination of 1.2 and 2.4 mg/kg body weight given at about two days interval. The temperature dropped to normal levels within 48 h of treatment and schizonts started degenerating as early as 24 h after treatment and had disappeared by 48 to 72 h. Piroplasms were not affected by the drug. Recovered animals remained free of East Coast fever for periods up to 12 months of observation despite continuous tick challenge. It was concluded that, with early detection and treatment augmented with proper tick control, outbreaks of East Coast fever can be effectively controlled with halofuginone lactate.     

Functional genomics and evolution of tick–Anaplasma interactions and vaccine development

The genus Anaplasma (Rickettsiales: Anaplasmataceae) includes several tick-transmitted pathogens that impact veterinary and human health. Tick-borne pathogens cycle between tick vectors and vertebrate hosts and their interaction is mediated by molecular mechanisms at the tick–pathogen interface. These mechanisms have evolved characteristics that involve traits from both the tick vector and the pathogen to insure their mutual survival. Herein, we review the information obtained from functional genomics and genetic studies to characterize the tick–Anaplasma interface and evolution of A. marginale and A. phagocytophilum. Anaplasma and tick genes and proteins involved in tick–pathogen interactions were characterized. The results of these studies demonstrated that common and Anaplasma species-specific molecular mechanism occur by which pathogen and tick cell gene expression mediates or limits Anaplasma developmental cycle and trafficking through ticks. These results have advanced our understanding of the biology of tick–Anaplasma interactions and have opened new avenues for the development of improved methods for the control of tick infestations and the transmission of tick-borne pathogens.