Variation in body size in the tick complex Rhipicephalus appendiculatus/Rhipicephalus zambeziensis.

We examined the relationship between body size and the phenology of the tick complex Rhipicephalus appendiculatus/Rhipicephalus zambeziensis. These ticks transmit Theileria parva in cattle. In Africa, the body size of R. appendiculatus increases with latitude while the body size of the morphologically similar Rhipicephalus zambeziensis is constant at two different latitudes. A larger body size is necessary once survival becomes a constraint. The most plausible explanation for the smaller R. appendiculatus in equatorial Africa is the cost to produce a larger egg. The consequences of these findings for the introduction of R. appendiculatus in new environments are discussed. New field observations from southern Zambia indicate that R. appendiculatus body size does not vary seasonally as compared to eastern Zambia. This is an additional indication of the presence of a single diapausing population of larger ticks.     

A survey of the ixodid ticks parasitising cattle in the Eastern Province of Zambia

The results of tick surveys carried out in the Eastern province of Zambia between December 1982 and February 1996 were principally in agreement with the findings of earlier surveys conducted during the period 1965–72. Boophilus decoloratus has almost been replaced by Boophilus microplus . Hyalomma truncatum was found in small numbers throughout the province and Hyalomma marginatum rufipes was only rarely encountered in collections made from cattle. Rhipicephalus evertsi evertsi was largely confined to the southern part of the plateau and the valleys of the Zambezi tributaries. The main difference between the present survey and the previous one concerns the status of Rhipicephalus appendiculatus . This species is currently expanding its range in a westward direction, whereas it was virtually absent from the southern part of the province during the period 1965–72. The majority of specimens collected are morphologically intermediate between R. appendiculatus and the closely allied Rhipicephalus zambeziensis . The available evidence indicates that R. zambeziensis ( sensu stricto ) is absent from the province. The phenology of R. appendiculatus is aberrant in the province: at lower altitudes a second generation of adult ticks is recorded on the hosts at the start of the dry season.     

Genetic variation in Rhipicephalus appendiculatus (Acari: Ixodidae) from Zambia: correlating genetic and ecological variation with Rhipicephalus appendiculatus from eastern and southern Africa.

Based on their ecology, Rhipicephalus appendiculatus ticks from eastern and southern Africa have been divided into three groups. We investigated how two geographic genetically differentiated stocks of R. appendiculatus from the southern and the eastern provinces of Zambia, representing two ecological groups, i.e., southern African and transition groups, respectively, genetically compare to stocks from east Africa (Rwanda) and southern Africa (Zimbabwe and South Africa). The ITS2 and two mitochondrial genes segments, 12s rDNA and COI, were used in the investigations. The ITS2 tree did not show support for differentiation into any groups, while the two mitochondrial genes trees (12s rDNA and COI) showed two genetically differentiated groups: an east African genetic group which included specimens from Rwanda and the plateau area of the eastern province of Zambia, and a southern African genetic group represented by specimens from South Africa, Zimbabwe and specimens collected on the fringes of the eastern province plateau in the Nyimba district of Zambia. This suggests that the two geographically differentiated stocks of the southern and eastern provinces of Zambia might be part of two wider geographic genetically differentiated R. appendiculatus groups that extend beyond Zambia. Stocks of "transition" ecology (eastern province) belong to the east African genetic group and the differences in ecology within this genetic grouping may be due to genetic polymorphism, phenotypic plasticity, and other local factors.     

A study of the diapausing behaviour of Rhipicephalus appendiculatus and R. zambeziensis under quasi-natural conditions in Zambia

Abstract. The diapausing behaviour of Rhipicephalus appendiculatus was studied under quasi-natural conditions in the Eastern Province of Zambia. Newly-moulted adults of ticks indigenous to the study area entered a behavioural diapause when exposed to daylengths below a critical photoperiod, provisionally a daylength between 11 h 20min and 11 h 45min. In the Eastern Province of Zambia the diapause was apparently not terminated by a long-day signal, but by a weakening of the photoperiodic maintenance of the diapause because of increasing age of the ticks. Adults of a reference stock indigenous to Kenya also entered a diapause when exposed to daylengths below the same threshold and maintained this diapause for the same length of time. Adults of a reference stock of Rhipicephalus zambeziensis indigenous to Zimbabwe did not enter a diapause when exposed to the same daylengths. The relevance of the findings is discussed in relation to the distributions of the two species.     

Survival of unfed Rhipicephalus appendiculatus and Rhipicephalus zambeziensis adults

Two ixodid tick species, Rhipicephalus appendiculatus (Neumann) and Rhipicephalus zambeziensis (Walker et al.) (Acari: Ixodidae), both originating from the southern province of Zambia, were used to study the survival time of adults at a range of different humidities and temperatures. In general, the experiment clearly demonstrates the different survival times of the two species in relation to the climatic conditions tested: survival of R. zambeziensis was better under more extreme conditions of temperature and humidity. These findings offer an explanation for the different distribution ranges of the two tick species. Rhipicephalus appendiculatus is more confined to cooler and wetter conditions, whereas R. zambeziensis is adapted to hotter and drier areas.     

Genetic variation in the body weight of ticks of the Rhipicephalus appendiculatus/zambeziensis complex

Sixteen strains of ticks of the Rhipicephalus appendiculatus/zambeziensis complex were created by cross-breeding for one to three generations. The body-weights of adult male ticks of these consanguineous lines were compared. A significant genetic variation was distinguished between the groups. Three classes whose mean weights varied around, respectively, 2, 3 and 4 mg/adult male tick, were found. This variation in weight may be explained by hybridization between R. appendiculatus Neumann and R. zambeziensis Walker, Norval & Corwin. Hybrid ticks were undersized whereas R. appendiculatus and R. zambeziensis groups had significantly higher weights.     

Ecological aspects of cattle tick control in central Zambia

In ecological studies in central Zambia, both climate and ecotype affected population dynamics of tick species. Below average rainfall for several years caused a suppression in numbers of Rhipicephalus appendiculatus Neumann adults. Reduction in rainfall leading to changes in grazing patterns is thought to have been responsible for an increase in numbers of Amblyomma variegatum Fabricius adults in a grassland habitat. There were reasonable correlations between numbers of each tick species on individual hosts over 1 year old. However, there were no relationships between numbers of ticks and bovine lymphocyte antigens (BoLA).     

Monthly dynamics of ticks (Acari: Ixodida) infesting N'Dama cattle in the Republic of Guinea

The occurrence of ixodid ticks on N'Dama cattle was studied in the Republic of Guinea between June 1994 and May 1995. Monthly tick collections were performed on 80 animals from 14 villages located in Dabola, Kouroussa and Dinguiraye prefectures. A total of 19,804 ticks was collected and classified using standard taxonomic keys. The following tick species were identified: Amblyomma variegatum, Hyalomma marginatum rufipes, Hyalomma trunctum, Hyalomma nitidum, Rhipicephalus lunulatus, Rhipicephalus muhsamae, Rhipicephalus senegalensis, Rhipicephalus sulcatus, Rhipicephalus turanicus, Boophilus annulatus, Boophilus geigyi. Boophilus spp. were the most numerous adult ticks (57.1%), Am. variegatum adults constituted 27.4%, while 12.4% were Rhipicephalus spp. and 2.5% Hyalomma spp. Rhipicephalus turanicus and Hyalomma nitidum were recorded for the first time in the country. Am. variegatum and Boophilus spp. were present throughout the year, whereas Am. variegatum adults showed a peak during the rainy season between April and September. Immature stages collected belonged exclusively to the genera Amblyomma and Boophilus. Am. variegatum larvae and nymphs showed a peak during the dry season (October-March); no significant variation between seasons was observed for Boophilus immatures. A significantly higher infestation of cattle by Rhipicephalus spp. was found in Dabola and Kouroussa prefectures, located in the southern part of the study area, with similar climatic, vegetation and rainfall characteristics. Possible options for the control of ticks in the study area are briefly discussed.     

Further evidence for geographic differentiation in R. appendiculatus (Acari: Ixodidae) from Eastern and Southern provinces of Zambia

Studies in the biology, ecology and behaviour of R. appendiculatus in Zambia have shown considerable variation within and between populations often associated with their geographical origin. We studied variation in the mitochondrial COI (mtCOI) gene of adult R. appendiculatus ticks originating from the Eastern and Southern provinces of Zambia. Rhipicephalus appendiculatus ticks from the two provinces were placed into two groups on the mtCOI sequence data tree. One group comprised all haplotypes of specimens from the Eastern province plateau districts of Chipata and Petauke. The second group consisted of a single haplotype of specimens from the Southern province districts and Nyimba, an Eastern province district on the fringes of the valley. This variation provides additional evidence to the earlier observations in the 12S rDNA and ITS2 data for the geographic subdivision of R. appendiculatus from Southern province and Eastern province plateau. The geographic subdivision further corresponds with differences in body size and diapause between R. appendiculatus from these geographic areas. The possible implications of these findings on the epidemiology of East Coast fever (ECF) the disease for which R. appendiculatus is one of the vectors are discussed. Nucleotide sequence data reported in this paper are available in the GenBank^TM under the Accession Numbers DQ859259–DQ859266     

Factors influencing the distribution of questing ticks and the prevalence estimation of T. parva infection in brown ear ticks in the Tanga region, Tanzania.

Questing ticks from various districts and agro-ecological zones (AEZ) in the Tanga Region of Tanzania were studied for a two-year period between September 1999 and July 2001. Collections of both nymphal and adult ticks occurred at 29 sites using a blanket or white cloth dragging technique. The species recorded in the order of decreasing abundance were Rhipicephalus appendiculatus, Boophilus spp., and Ambylomma variegatum. Rhipicephalus appendiculatus field infestation levels varied across administrative districts and seasons, with Amani and the cool-to-dry season having lower tick counts ( = -2.09, SE = 0.71, P < 0.001 and = -1.54, SE = 0.56, P = 0.006 for Amani and cool to dry season, respectively). Based on the polymerase chain reaction technique, Theileria parva infection prevalence in adult R. appendiculatus was estimated to be 2.6%. Implications of these findings in light of the spatial and temporal distribution of ticks are discussed for the accurate diagnosis of multiple tick-borne diseases, the infected vector tick, the endemic status of T. parva in the region, and the implemention of control strategies.     

Skin test to detect resistance of cattle to Rhipicephalus appendiculatus ticks

The evaluation of a skin test to detect acquired resistance to Rhipicephalus appendiculatus Neumann ticks is described. An extract of salivary glands of partially fed female R. appendiculatus was prepared by dissection, sonication and filtration and used for intra-dermal injection. Tests were carried out on twelve calves of known resistance to R. appendiculatus and on twelve naive calves to establish threshold values for positive reactions. Four rabbits naive to ticks were skin tested repeatedly to assess immunogenicity of the test. Reactions to the test at 1 h and at 24 h after injection were significantly correlated with resistance. The correlation was higher with the reactions at 24 h. Rabbits were immunized by the test but the reactions never exceeded the positive threshold. Further development by field testing is recommended.     

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.     

A novel SINE family occurs frequently in both genomic DNA and transcribed sequences in ixodid ticks of the arthropod sub-phylum Chelicerata

Reassociation kinetics and flow cytometry data indicate that ixodid tick genomes are large, relative to most arthropods, containing>or=10(9) base pairs. The molecular basis for this is unknown. We have identified a novel small interspersed element with features of a tRNA-derived SINE, designated Ruka, in genomic sequences of Rhipicephalus appendiculatus and Boophilus (Rhipicephalus) microplus ticks. The SINE was also identified in expressed sequence tag (EST) databases derived from several tissues in four species of ixodid ticks, namely R. appendiculatus, B. (R.) microplus, Amblyomma variegatum and also the more distantly related Ixodes scapularis. Secondary structure predictions indicated that Ruka could adopt a tRNA structure that was, atypically, most similar to a serine tRNA. By extrapolation the frequency of occurrence in the randomly selected BAC clone sequences is consistent with approximately 65,000 copies of Ruka in the R. appendiculatus genome. Real time PCR analyses on genomic DNA indicate copy numbers for specific Ruka subsets between 5800 and 38,000. Several putative conserved Ruka insertion sites were identified in EST sequences of three ixodid tick species based on the flanking sequences associated with the SINEs, indicating that some Ruka transpositions probably occurred prior to speciation within the metastriate division of the Ixodidae. The data strongly suggest that Class I transposable elements form a significant component of tick genomes and may partially account for the large genome sizes observed.     

The effects of the brown ear-tick, Rhipicephalus appendiculatus, on milk production in dairy cattle

Abstract. The effect is reported of artificially controlled levels of infestation with adults of Rhipicephalus appendiculatus on the milk yield of twenty commercial Bos taurus dairy cattle on a high plane of nutrition and eighteen crossbed B.taurus × Sanga cattle on a lower plane of nutrition in the highveld of Zimbabwe. The results showed no significant effect on milk yield of infestations averaging twenty engorging ticks per animal per day, despite severe ear damage in some animals. They indicate that milk production of dairy cattle under commercial management is not sensitive to infestation with R.appendiculatus. The results have important implications for management of ticks in Africa, but need to be interpreted within the context of the control of tick-borne diseases.     

An assessment of host resistance to ticks on cross-bred cattle in Burundi

Abstract. Groups of cattle of four different cross-breeds (Ankole × Friesian, Ankole × Brown-Swiss, Ankole × Guernsey, Ankole × Sahiwal) plus a group of pure Ankole cattle were immunized against tick-borne diseases: East Coast Fever, using a Theileria parva trivalent vaccine, and anaplasmosis, babesiosis and, subsequently, heartwater. With the exception of two small subgroups of Ankole and Ankole × Sahiwal which were treated by weekly spraying using Delnav, all the groups were exposed to natural field challenge of ticks to quantify the development of host resistance to ticks. The results indicate resistance to the most abundant species, Rhipicephalus appendiculatus , as follows: Ankole > Ankole × Sahiwal > Ankole × Brown Swiss > Ankole × Friesian > Ankole × Guernsey. In both Amblyomma variegatum and Rhipicephalus evertsi evertsi , the manifestation of different levels of resistance between breeds is inapparent. Correlations between total ticks and standard female ticks were good for R. appendiculatus (r = 0.73) and R. evertsi (r - 0.51) but poor for A. variegatum (r = 0.31). Correlations between species were reasonably consistent (r = 0.43-0.59). There was no significant correlation between tick burdens and daily liveweight gain (DLWG) over the. whole period of the study. Of the cross-bred groups, Ankole × Sahiwal, which had the highest level of resistance, had the highest DLWG. In contrast, the Ankole × Guernsey which developed the lowest level of resistance had only marginally lower DLWG, the difference being only 8 g/day. During the first period of exposure to ticks, however, the untreated Ankole × Sahiwal group had markedly lower DLWG, which may indicate that the development and manifestation of resistance is at the expense of productivity.     

Detection of Babesia spp. in free-ranging Pukus, Kobus vardonii, on a game ranch in Zambia

Babesia spp. were detected from 4 asymptomatic pukus captured on a game ranch in central Zambia in October 2008. Blood smears were examined in 4 species of aymptomatic free-ranging antelopes, namely the puku (Kobus vordanii), reedbuck (Redunca arundinum), bushbuck (Tragelaphus sylvaticus), and kudu (Tragelaphus strepsiceros), and showed the presence of Babesia parasites only in the puku. In the puku, the prevalence of babesiosis was estimated at 33.3% (n = 12), while the overall prevalence in all examined animals was 8.5% (n = 47). The parasites showed morphological characteristics of paired ring-like stages with the length varying between 1.61 μm and 3.02 μm (mean = 2.12 μm, n = 27; SD = 0.76 μm). Both the infected and non-infected pukus showed good body condition scores (BCS), while the dominant tick species detected from all animals were Rhipicephalus appendiculatus, Rhipicephalus spp., and Boophilus spp. To our knowledge this is the first report of Babesia spp. infection in pukus in Zambia. These findings suggest that wildlife could play an important role in the epidemiology of babesiosis in Zambia.     

Population dynamics of ticks on indigenous cattle in a pastoral dry to semi-arid rangeland zone of Uganda

Studies on seasonality and population dynamics of ticks on indigenous cattle and their crosses (calves) were carried out in Buruli Ranching Scheme, Nabiswera, Luwero district of Uganda on three treatment groups of animals: group 1 (twice a week dipping), group 2 (once a month dipping) and group 3 (no tick control). During this study, four major species of ticks of economic importance were recorded in decreasing order of abundance: Rhipicephalus appendiculatus, Rhipicephalus evertsi evertsi, Amblyomma variegatum and Boophilus decoloratus. Of these ticks, a seasonal pattern of activity was only observed in R. appendiculatus ticks, with peak activities occurring during rainy seasons. Greater numbers of ticks were recorded on cows than calves in the three treatment groups, with the exception of A. variegatum where the reverse occurred. The mean numbers of ticks per animal were highly significantly different (p < 0.01) when group 1 animals were compared with group 2 and 3 animals. However, no significant differences (p > 0.05) were observed in mean tick numbers between group 2 and 3 animals. Highly significant differences (p < 0.01) were observed in mean tick numbers on cows and calves (more than 12 months old) in different calving seasons. The state of lactation only affected tick counts on cows in group 1; significantly more ticks (p < 0.01) were observed in lactating than non-lactating cows. Furthermore, significantly greater (p < 0.05) numbers of ticks (with the exception of B. decoloratus) were recorded during the second year of study (March 1992-May 1993) than the first year (January 1991-February 1992), despite lower rainfall during the former period.     

Seasonal dynamics of the brown dog tick,Rhipicephalus sanguineus,on a confined dog population in Italy

This study evaluated the seasonal dynamics of Rhipicephalus sanguineus (Latreille) (Acari: Ixodidae) on naturally infested dogs in a private shelter in southern Italy. From March to May 2008, 39 autochthonous mixed‐breed young dogs and 10 beagles were enrolled in the study. From March 2008 until March 2009, every 21 ± 2 days, 11 body sites of each dog were checked for ticks. At each follow‐up, the number of ticks, their developmental stage, sex and location on the dog's body were recorded. Adult ticks were found throughout the year, but immatures were absent in January and February. The adult tick population increased from July to August, whereas the load of immatures increased in early July and peaked in September, which suggests that R. sanguineus develops one generation per year in this area. The mean number of immature ticks per infested dog was higher than that of adults from March to October 2008. Ears, interdigital areas and armpits were the most frequent attachment sites of adult ticks. At the last follow‐up, a total of 2266 ticks were collected and identified as R. sanguineus. The results suggest that R. sanguineus develops one generation per year in the study area, but that it infests dogs in all seasons. This information should be taken into account when planning control programmes against this tick species and the pathogens it transmits.     

Synonymy of Boophilus Curtice, 1891 with Rhipicephalus Koch, 1844 (Acari: Ixodidae)

Recent molecular and morphological studies of the genera Rhipicephalus Koch, 1844 and Boophilus Curtice, 1891 revealed that the five species of Boophilus make the genus Rhipicephalus paraphyletic. Thus, Rhipicephalus Koch, 1844 is not a monophyletic (natural) lineage and some species of Rhipicephalus are more closely related to the species of Boophilus than to other species of Rhipicephalus. Here, we revise these genera: Boophilus is synonymised with Rhipicephalus, and Rhipicephalus (sensu lato) (including Boophilus) is redefined. By synonymising Boophilus with Rhipicephalus, we have changed the nomenclature so that it reflects our understanding of the phylogeny of these ticks. Boophilus is retained as a subgenus of Rhipicephalus, so the synonymy of Boophilus with Rhipicephalus does not result in the loss of the name Boophilus. In addition, Rhipicephalus is a well-known genus and the change proposed is simple -- all five species of Boophilus become members of Rhipicephalus (Boophilus).     

Abiotic and biotic determinants of the seasonal dynamics of the tick Rhipicephalus appendiculatus in South Africa

Abstract. The creation of a generic population model for the tick Rhipicephalus appendiculatus requires a detailed, quantified understanding of the interactions of these ticks with their biotic and abiotic environment in the different parts of their range, from the tropical regions of East Africa to the temperate regions of South Africa. The much greater seasonal variation in climatic conditions, particularly temperature, further from the equator introduces variable development rates and diapause into the life cycle. Estimates of natural temperature-dependent interstadial development periods, derived from a combination of published laboratory and field data, were applied to published data on the seasonal abundance of R.appendiculatus on three farms in the Eastern Cape Province of South Africa. This enabled an assessment of which ticks of one stage give rise to which ticks of the next stage, from which (a) the onset and duration of diapause in unfed adults, and (b) seasonal interstadial mortality indices in the form of fc-values, could be estimated. The contribution of biotic (tick density) and abiotic (climatic) factors as predictors of mortality at each life stage was investigated by step-wise multiple regression. Density-independent mortality at the female-to-larval stage is correlated with geographically variable climatic factors, minimum temperature at two farms and minimum relative humidity at the third. The other two stages are governed by density-dependent mortality, which, it is argued, may be caused largely by the hosts' acquired resistance to ticks. As expected on theoretical grounds, this density dependence is weaker nearer to the edge of the tick's range and for the more vulnerable immature stages.