Focus of human parasitism by the brown dog tick, Rhipicephalus sanguineus (Acari: Ixodidae)

Throughout most of its range, the brown dog tick, Rhipicephalus sanguineus (Latreille), prefers dogs as a host, but human bites occasionally occur in the Mediterranean region and Central America; historically, this species has rarely bitten humans in the United States. A focus of 15 human bite cases by this tick is reported from four Air Force bases located within 200 miles of each other in north central Texas and southwestern Oklahoma. The sudden appearance of numerous documented bite cases indicates that either the species is becoming more anthropophilic in this area or than an introduction of a more human adapted population of R. sanguineus has occurred. The species is a known reservoir and vector of Ehrlichia canis, the causative agent of canine ehrlichiosis in dogs. The presence of an anthropophilic strain of R. sanguineus in the United States may lead to increased exposure to E. canis and thus an increase in the incidence of human ehrlichiosis.     

The dog factor in brown dog tick Rhipicephalus sanguineus (Acari:Ixodidae) infestations in and near human dwellings

Three cases of the tick Rhipicephalus sanguineus infiltration in or near human dwellings caused by dogs, and their influence on epidemiological features of human habitats have been investigated, (a) The observation of dogs kept indoors proved that single tick females could engorge and oviposit inside apartments followed by the development of subadults. (b) Abundant micropopulations of ticks were formed in small yards or gardens near the dwellings where dogs lived in kennels, (c) A huge field population of R. sanguineus was observed on a farm where watchdogs constantly patrolled along the farm perimeter. Tick abundance near the kennels and in the permanent resting sites of the dogs reached more than 30 adults per 10 min of collecting, while the number of adults on a dog reached 100. Unfed adult females under conditions of constant dog availability had a larger scutal index than females collected in the control field site. On the basis of circumstantial evidence it is possible to conclude that under the above conditions tick development may change from the normal 3-host cycle to a 2-host cycle. Ticks in the field had one complete generation per year. Ticks on the farm, as well as ticks in kennels, developed faster and a significant part of their population had two complete generations per year. R. sanguineus is the main vector and reservoir of a pathogen from the Rickettsia conorii complex, the causative agent of Israeli tick typhus. The described conglomerations of R. sanguineus create a great risk to humans who can be attacked by infected ticks in and around their homes, even in large towns. Such a feature of the tick life history most likely exists not only in Israel but in other countries as well.     

Computer simulation of population dynamics of the American dog tick (Acari: Ixodidae)

A comprehensive computer model was developed for simulation of the population dynamics of the American dog tick (ADT), Dermacentor variabilis Say, in North America. The model simulates the effects of major environmental variables, such as ambient temperature, saturation deficit, kind of habitat, and host density, on ADT population dynamics in ecosystems with small mammals as hosts for immature ticks and medium-sized mammals or domestic dogs as hosts for adult ticks. General validity of the model was established by comparisons between simulated and actual population densities for a series of years at locations in Virginia, Maryland, and Massachusetts using actual weekly weather data for each year as a model input. Using historical-average weather data for 11 locations within the known geographic range of ADT and 3 locations outside this range, the model produced acceptable values for initial population growth rate and generation time, as well as realistic equilibrium population densities and seasonal activity patterns. This model can be used as a framework for additional modeling efforts to simulate the transmission of Rocky Mountain spotted fever and to study various strategies for management of ADT populations.     

Cloning and sequencing of putative acetylcholinesterase cDNAs from the American dog tick, Dermacentor variabilis, and the brown dog tick, Rhipicephalus sanguineus (Acari: Ixodidae)

Two putative cDNAs of acetylcholinesterase (AChE), one from Dermacentor variabilis, and the other from Rhipicephalus sanguineus, were amplified and sequenced. The deduced amino acid sequences have high amino acid identities (between 70 and 94%) to known tick AChE sequences deposited in GenBank. Furthermore, these two AChEs also possess common features in their primary AChE structure such as catalytic active sites. A 2,220-bp contiguous sequence, containing a 1,791-bp open reading frame encoding an AChE precursor with 596 amino acid residues, was obtained from D. variabilis. The deduced proteins of R. sanguineus are different in size by 6 amino acids because of alternative splicing at the 5' end. A gene tree deduced from phylogenetic analysis indicates that there are at least three lineages of AChE in arthropods.     

Computer simulation of Rocky Mountain spotted fever transmission by the American dog tick (Acari: Ixodidae)

A computer model was developed for simulation of the transmission of Rickettsia rickettsii, the causative agent of Rocky Mountain spotted fever (RMSF), by the American dog tick, Dermacentor variabilis (Say). The model of RMSF was combined with a model for population dynamics of the American dog tick and included simulation of infection and transmission of rickettsiae between ticks and host mammals and transmission of RMSF to humans. The model simulated the effects of biotic and environmental variables such as weather, host density, habitat, transovarial transmission, fecundity of infected ticks, and infectivity level of ticks and mammals. Some parameters in the model were fitted by iterative simulations to produce realistic rates of R. rickettsii infection in adult ticks and small and medium-sized mammal hosts. Parameters also were fitted to yield the historical average number of RMSF cases for Virginia. Comparisons of the simulated and actual number of cases for nine other states indicated a reasonable level of validity for the model. A theoretical tick density threshold of 252 unfed adult ticks/ha for transmission of RMSF was determined from a relationship between rate of transmission to humans and density of ticks. The transmission threshold can be used for additional modeling efforts to study the effects of management technologies on tick densities and RMSF human cases. The model can serve as a framework for modeling other tick-borne diseases such as Lyme disease, babesiosis, and heartwater.     

Novel Ehrlichia organism (Rickettsiales: Ehrlichieae) in white-tailed deer associated with lone star tick (Acari: Ixodidae) parasitism

Polymerase chain reaction (PCR) evidence of a novel Ehrlichia organism was found recently in wild white-tailed deer, Odocoileus virginianus Zimmermann, and lone star ticks, Amblyomma americanum L., from the southeastern United States. To evaluate whether lone star tick parasitism was associated with the presence of this novel Ehrlichia organism in deer, 2 retrospective studies were conducted using specific nested PCR to test archived deer serum samples. The 1st study of 150 serum samples collected from a single deer population over a 15-yr period examined the temporal association between the presence of the Ehrlichia organism in deer and parasitism by lone star ticks. The deer Ehrlichia was not detected in serum samples collected before 1986, when lone star ticks were absent or rare, but was detected in samples collected in 1986 and every year thereafter, when lone star ticks became increasingly abundant. In the 2nd study, serum samples from 120 deer from 24 sites in 14 southeastern states were tested to evaluate if a site-specific, spatial association existed between the presence of the deer Ehrlichia and lone star ticks. All 60 serum samples from the 12 deer populations without evidence of lone star tick infestation were negative for the deer Ehrlichia, whereas 83% of the 12 populations infested by lone star ticks had PCR evidence of infection. These data suggest that lone star ticks may be a vector of the deer Ehrlichia; however, they do not preclude the involvement of other arthropods in maintaining infection with this organism in deer populations.     

Resistance between stages of the tick Rhipicephalus appendiculatus (Acari: Ixodidae)

The ability of one stage of Rhipicephalus appendiculatus Neumann to induce resistance in a host to the feeding of the next stage in the life cycle was studied using Bos taurus cattle and rabbits. Simple resistance within a stage and cross-resistance between stages was generally less than 50%. The cumulative effect of simple resistance and cross-resistance over the whole life cycle in this experiment was estimated at 97% resistance measured as reduction in engorgement mass, numbers detached, and survival. Potential antigens in salivary glands of the three stages were compared by microscopy and electrophoresis and found closely similar. Production of serum antibodies against salivary antigens was examined by Western blotting of the electrophoretically separated salivary peptides. Cross reactions occurred but with little consistency. Hypersensitivity reactions were observed directly and tested immunologically. Only delayed hypersensitive reactions were directly involved in simple resistance and cross-resistance. These results can be applied in the selection of cattle for resistance to ticks, and this is discussed.     

Computer simulation of Boophilus cattle tick (Acari: Ixodidae) population dynamics

A comprehensive computer model was developed for simulation of the population dynamics of the cattle ticks, Boophilus microplus (Canestrini) and B. annulatus (Say). The model is deterministic and based on a dynamic life table with weekly time steps. The model simulates the effects of major environmental variables, such as ambient temperature, saturation deficit, precipitation, type of pasture, type of cattle, and cattle density on Boophilus cattle tick population dynamics. General validity of the model is established by comparing simulated and observed yearly densities of standard female ticks/host/day. B. microplus population comparisons were made for a series of years using weekly weather data from two locations in Queensland, Australia. The model also produced acceptable values for initial population growth rate, generation time, and 3-yr population density when historical weather at 7 locations in Australia and 23 locations in the Americas were used. This model provides a framework for the study of Babesia transmission by Boophilus ticks, and can be used to study the effects of control technologies and to develop more efficient and environmentally acceptable eradication strategies for Boophilus ticks.     

Production of pheromones by artificially fed males of the tick Amblyomma maculatum (Acari: Ixodidae)

Aggregation-attachment pheromones are produced only by male ticks of the genus Amblyomma that have imbibed blood for at least 8 d from their mammalian hosts. This report demonstrates that production and release of aggregation-attachment pheromones by Amblyomma maculatum (Koch) males can be induced in vitro by using nonblood media, artificially introduced through capillary tubes, into the male ticks. The presence of these pheromones is demonstrated by using both biological observations on bovine hosts and in vitro experimentation with petri dishes. The attraction of unfed female ticks to the artificially fed males (using the petri dish method) was significantly higher (P < 0.001) than the attraction of unfed females to unfed males. Similarly, attraction and attachment were recorded on bovine hosts. Only 62.8 +/- 17.5% of the females released attached to the bovine host. Of these, 61.9 +/- 19.37% attached around artificially fed males that were placed on the bovine 24 h earlier. This percentage did not differ significantly from the 81.4 +/- 7.1% of the total that attached around males that fed naturally for 8 d. In comparison, only 33.3 +/- 21.9% of the total number of females placed on the host did so after being fed 1 d (control group). The possibility of using capillary feeding as a new tool for investigating the physiology and reproductive behavior of blood-sucking arthropods is discussed.     

A dispersal model for the range expansion of blacklegged tick (Acari: Ixodidae)

The blacklegged tick, Ixodes scapularis Say, a vector for the agents of Lyme borreliosis and other diseases, has expanded its range dramatically over the past 20 yr. However, the relative contributions of different vertebrate host species to this expansion have remained largely unexplored. To address this issue, we simulated the expansion of a theoretical tick population across a simple landscape by using a deterministic, spatially explicit, cellular automata model. The model incorporates the ecology of ticks and three vertebrate hosts: white-tailed deer, Odocoileus virginianus Zimmermann; white-footed mouse, Peromyscus leucopus Rafinesque; and American robin, Turdus migratorius L. Host contribution to tick dispersal is modeled as a function of tick burden, home range size, and population density. These parameters were determined using published and unpublished data. Our results suggest that 1) hosts with high tick burdens and large home ranges (e.g., deer) play a critical role in I. scapularis range expansion; 2) hosts with small home ranges (e.g., mice) can limit range expansion if they divert a sufficient number of ticks from feeding on more mobile hosts; and (3) birds that migrate annually (e.g., robins) can play a crucial role in tick range expansion.     

Transmission efficiency of Francisella tularensis by adult american dog ticks (Acari: Ixodidae)

The American dog tick, Dermacentor variabilis (Say) (Acari: Ixodidae), has been implicated as a potential bridging vector to humans of Francisella tularensis, the etiological agent of tularemia. Since the initial studies evaluating vector competency of D. variabilis were conducted, F. tularensis has been subdivided into subspecies and clades that differ in their geographical distribution in the United States and in the severity of infections caused in humans. Here, we demonstrate that D. variabilis nymphs efficiently acquire, transtadially maintain, and transmit each of the strains tested (clades A1b and A2, and type B). Transmission efficiency by adult females was similarly high among infection groups and ranged from 58% for type B to 89% for A2 infections. In addition, we demonstrated that transmission can occur shortly after tick attachment. These findings support the concept that D. variabilis adults may play a significant role in epizootic transmission of F. tularensis, and as a bridging vector to humans.     

Mitotic activity of the hemocytes in the tick Ixodes ricinus (Acari; Ixodidae)

 The blood cells, or hemocytes, of Ixodes ricinus have been shown to recognize, attack, and phagocytose microorganisms invading the body cavity, or hemocoel, of this tick. Regulated proliferation and differentiation of hemocytes, also referred to as immunocytes, is basic to an effective immune response to invading microorganisms. Therefore, this study dealt with hemopoiesis in I. ricinus, the vector tick of the Lyme disease spirochete Borrelia burgdorferi. Histological evidence for the presence of hemopoietic tissue, a preferential proliferation site of hemocytes, is presented. Mainly the mitotic activity of free-floating hemocytes was examined. By means of microscopical photometry and flow cytometry, all three types of hemocytes in engorging female I. ricinus were found in different stages of the cell cycle. In the engorging tick, up to 40% of the hemocytes counted were in the S phase or the G₂/M phase. From this study we conclude that the differentiated hemocyte types do not differentiate from stem cells in the adult tick. Moreover, microorganisms entering the hemocoel of engorging ticks are confronted with high numbers of hemocytes and, therefore, with an effective cellular immune response. Received: 20 October 1995 / Accepted: 30 January 1996