Hard ticks (Acari: Ixodidae) parasitizing bushbabies (Mammalia: Galagidae) in a biodiversity hotspot of northern South Africa.

South Africa has six species of primates, three of which are bushbabies (family Galagidae). Very little information is available on their parasites due to the lack of longitudinal studies, although Rhipicephalus appendiculatus, Amblyomma hebraeum and Haemaphysalis elliptica were previously reported from the brown greater galago (Otolemur crassicaudatus) in South Africa. During 2014-2019, 83 O. crassicaudatus (70 live-trapped and 13 deceased animals) were checked for the presence of hard ticks, all from Limpopo Province, South Africa. Seventy-three of 83 (88 %) galagos were found to be tick-infested. Among ixodid genera, Haemaphysalis had the highest prevalence (46 % of the bushbabies), followed by Rhipicephalus (25 %) and Ixodes (18 %). In total, ten tick species were identified. Importantly, all infestations were monospecific. Ticks occurred on various body parts of bushbabies, thus no predilection site was noted. In conclusion, while previously only three ixodid species were known to infest bushbabies in South Africa, the present study showed that these animals can be parasitized by a much broader range of hard ticks.

Molecular-phylogenetic analyses of Ixodes species from South Africa suggest an African origin of bird-associated exophilic ticks (subgenus Trichotoixodes).

BACKGROUND: Among hard ticks (Acari: Ixodidae), the genus Ixodes comprises the highest number of species, which in turn are most numerous in the Afrotropical zoogeographic region. In South Africa extensive morphological studies have been performed on Ixodes species but only few reports included molecular analyses. METHODS: In this study, 58 Ixodes spp. ticks, collected from ten mammalian and eight avian host species in South Africa, were molecularly and phylogenetically analyzed. In addition, a newly collected sample of the Palearctic Ixodes trianguliceps was included in the analyses. RESULTS: Among the ticks from South Africa, 11 species were identified morphologically. The majority of ticks from mammals represented the Ixodes pilosus group with two species (n = 20), followed by ticks resembling Ixodes rubicundus (n = 18) and Ixodes alluaudi (n = 3). In addition, single specimens of Ixodes rhabdomysae, Ixodes ugandanus, Ixodes nairobiensis and Ixodes simplex were also found. Considering bird-infesting ticks, Ixodes theilerae (n = 7), Ixodes uriae (n = 4) and ticks most similar to Ixodes daveyi (provisionally named I. cf. daveyi, n = 2) were identified. Molecular analyses confirmed two species in the I. pilosus group and a new species (I. cf. rubicundus) closely related to I. rubicundus sensu stricto. Phylogenetic trees based on concatenated mitochondrial or mitochondrial and nuclear gene sequences indicated that the subgenus Afrixodes forms a monophyletic clade with bird-associated exophilic ticks (subgenus Trichotoixodes). Ixodes trianguliceps clustered separately whereas I. alluaudi with their morphologically assigned subgenus, Exopalpiger. CONCLUSIONS: Phylogenetic analyses shed new lights on the relationships of Ixodes subgenera when including multiple sequences from subgenus Afrixodes and African as well as Palearctic species of subgenera Trichotoixodes and Exopalpiger. Subgenera Afrixodes and bird-associated Trichotoixodes share common ancestry, suggesting that the latter might have also originated in Africa. Regarding the subgenus Exopalpiger, I. alluaudi is properly assigned as it clusters among different Australian Ixodes, whereas I. trianguliceps should be excluded.

Compositional turnover in ecto- and endoparasite assemblages of an African bat, Miniopterus natalensis (Chiroptera, Miniopteridae): effects of hierarchical scale and host sex.

We studied the compositional turnover in infracommunities and component communities of ecto- and endoparasites infesting a bat, Miniopterus natalensis (Chiroptera, Miniopteridae), across seven sampling sites using the zeta diversity metric (measuring similarity between multiple communities) and calculating zeta decline and retention rate (both scales) and zeta decay (component communities). We asked whether the patterns of zeta diversity differ between (a) infracommunities and component communities; (b) ecto- and endoparasites and (c) subsets of communities infecting male and female bats. The pattern of compositional turnover differed between infracommunities and component communities in endoparasites only. The shape of zeta decline for infracommunities indicated that there were approximately equal probabilities of ecto- and endoparasitic species to occur on/in any bat individual within a site. The shape of zeta decline for component communities suggested the stochasticity of ectoparasite turnover, whereas the turnover of endoparasites was driven by niche-based processes. Compositional turnover in component communities of ectoparasites was more spatially dependent than that of endoparasites. Spatial independence of compositional turnover in endoparasites was due to subcommunities harboured by female bats. We conclude that the patterns of compositional turnover in infracommunities were similar in ecto- and endoparasites, whereas the patterns of turnover in component communities differed between these groups.

Mitochondrial and nuclear multilocus phylogeny of Rhipicephalus ticks from Kenya.

The morphological diversity of African ticks of the genus Rhipicephalus and subgenus Boophilus have been studied in detail. However, their taxonomy remains poorly resolved with limited molecular studies performed to improve inter-species discrimination. Herein, ribosomal cytochrome c oxidase I (COI), 12S ribosomal DNA (12S rDNA) and nuclear ribosomal DNA internal transcriber spacer 2 (ITS2) were analyzed in Rhipicephalus tick populations in Kenya. While the morphological and molecular criteria separated R. e. evertsi, R. pulchellus and R. appendiculatus from other members of the genus, except the morphologically similar sibling species R. zambeziensis, this was not the case for other tick populations. COI sequences of Rhipicephalus ticks from Ruma National Park (RNP) in Southwestern Kenya, that were morphologically similar to R. praetextatus/R. simus, a formed distinct clade and barcode gap group. 12S rDNA haplotypes of this population were 99% identical to a GenBank accession of R. muhsamae which is thought to be endemic in West and Central Africa. However, the ITS2 locus indicated that the RNP samples were genetically closest to ticks identified morphologically as R. praetextatus. The COI and 12S rDNA haplotype sequences of R. praetextatus clustered closely with R. simus reference sequences though the two species occurred in distinct barcode gap groups. Our results suggest that the R. simus/R. praetextatus/R. muhsamae comprise a closely related tick species complex found across sub-Saharan Africa and includes the yet to be described RNP population. More studies on the biology, ecology and genomics of all life stages of tick species in the complex may clarify their taxonomic status. A continent-wide study that combines morphology, DNA marker sequencing and emerging methods, such as mass spectrometry and whole-genome resequencing may reveal the diversity and distribution of taxa within the genus Rhipicephalus in sub-Saharan Africa.

Argasid and ixodid systematics: Implications for soft tick evolution and systematics, with a new argasid species list.

The systematics of the genera and subgenera within the soft tick family Argasidae is not adequately resolved. Different classification schemes, reflecting diverse schools of scientific thought that elevated or downgraded groups to genera or subgenera, have been proposed. In the most recent classification scheme, Argas and Ornithodoros are paraphyletic and the placement of various subgenera remains uncertain because molecular data are lacking. Thus, reclassification of the Argasidae is required. This will enable an understanding of soft tick systematics within an evolutionary context. This study addressed that knowledge gap using mitochondrial genome and nuclear (18S and 28S ribosomal RNA) sequence data for representatives of the subgenera Alectorobius, Argas, Chiropterargas, Ogadenus, Ornamentum, Ornithodoros, Navis (subgen. nov.), Pavlovskyella, Persicargas, Proknekalia, Reticulinasus and Secretargas, from the Afrotropical, Nearctic and Palearctic regions. Hard tick species (Ixodidae) and a new representative of Nuttalliella namaqua (Nuttalliellidae), were also sequenced with a total of 83 whole mitochondrial genomes, 18S rRNA and 28S rRNA genes generated. The study confirmed the utility of next-generation sequencing to retrieve systematic markers. Paraphyly of Argas and Ornithodoros was resolved by systematic analysis and a new species list is proposed. This corresponds broadly with the morphological cladistic analysis of Klompen and Oliver (1993). Estimation of divergence times using molecular dating allowed dissection of phylogeographic patterns for argasid evolution. The discovery of cryptic species in the subgenera Chiropterargas, Ogadenus and Ornithodoros, suggests that cryptic speciation is common within the Argasidae. Cryptic speciation has implications for past biological studies of soft ticks. These are discussed in particular for the Ornithodoros (Ornithodoros) moubata and Ornithodoros (Ornithodoros) savignyi groups.

First records of Hyalomma rufipes and Ixodes neitzi (Acari: Ixodidae) found on large carnivores in South Africa.

Ixodid ticks (Acari: Ixodidae) are important disease vectors for large carnivores, but the composition of the tick communities that parasitize carnivores is poorly understood. We collected ticks from leopards (Panthera pardus) and brown hyenas (Hyaena brunnea) in the Soutpansberg Mountains, South Africa, to determine which species feed on these carnivores. We identified a total of eight tick species belonging to six genera, and recorded Ixodes neitzi and Hyalomma rufipes on P. pardus for the first time.

Detection of zoonotic agents and a new Rickettsia strain in ticks from donkeys from South Africa: Implications for travel medicine.

BACKGROUND: In rural South Africa, people are in close contact with tick-infested donkeys. This study aimed to investigate the presence of spotted fever group Rickettsia, Anaplasma, Ehrlichia and Coxiella species in these arthropods. METHOD: 376 ticks (7 species) from donkeys from Limpopo Province (South Africa) were pooled and analyzed using PCR assays for the bacterium detection. RESULTS: Rickettsia africae was amplified in 6 Amblyomma hebraeum, 1 Rhipicephalus appendiculatus and 5 Rhipicephalus evertsi evertsi pools. Rickettsia aeschlimannii was found in 1 Hyalomma rufipes, 1 Rh. appendiculatus and 2 Rh. e. evertsi pools. Three Rhipicephalus simus specimens were infected with a new Rickettsia strain that showed low identity with any validated Rickettsia species. Ehrlichia canis was detected in 2 Rh. e. evertsi pools and in one of them Anaplasma bovis was amplified. An Am. hebraeum pool showed infection with Anaplasma ovis and another with Coxiella burnetii. CONCLUSION: South African donkeys are involved in the epidemiology of tick-borne pathogens and other associated agents such as C. burnetii with Health importance. A potential new Rickettsia species, with unknown pathogenic potential, has been detected in the anthropophilic Rh. simus.

Parasites of domestic and wild animals in South Africa. L. Ixodid ticks infesting horses and donkeys.

The aim of the study was to determine the species spectrum of ixodid ticks that infest horses and donkeys in South Africa and to identify those species that act as vectors of disease to domestic livestock. Ticks were collected opportunistically from 391 horses countrywide by their owners or grooms, or by veterinary students and staff at the Faculty of Veterinary Science, University of Pretoria. Ticks were also collected from 76 donkeys in Limpopo Province, 2 in Gauteng Province and 1 in North West province. All the ticks were identified by means of a stereoscopic microscope. Horses were infested with 17 tick species, 72.1% with Rhipicephalus evertsi evertsi, 19.4% with Amblyomma hebraeum and 15.6% with Rhipicephalus decoloratus. Rhipicephalus evertsi evertsi was recovered from horses in all nine provinces of South Africa and R. decoloratus in eight provinces. Donkeys were infested with eight tick species, and 81.6% were infested with R. evertsi evertsi, 23.7% with A. hebraeum and 10.5% with R. decoloratus. Several tick species collected from the horses and donkeys are the vectors of economically important diseases of livestock. Rhipicephalus evertsi evertsi is the vector of Theileria equi, the causative organism of equine piroplasmosis. It also transmits Anaplasma marginale, the causative organism of anaplasmosis in cattle. Amblyomma hebraeum is the vector of Ehrlichia ruminantium, the causative organism of heartwater in cattle, sheep and goats, whereas R. decoloratus transmits Babesia bigemina, the causative organism of babesiosis in cattle.

Investigation of Rickettsia, Coxiella burnetii and Bartonella in ticks from animals in South Africa.

Ticks are involved in the epidemiology of several human pathogens including spotted fever group (SFG) Rickettsia spp., Coxiella burnetii and Bartonella spp. Human diseases caused by these microorganisms have been reported from South Africa. The presence of SFG Rickettsia spp., C. burnetii and Bartonella spp. was investigated in 205 ticks collected from domestic and wild animals from Western Cape and Limpopo provinces (South Africa). Rickettsia massiliae was detected in 10 Amblyomma sylvaticum and 1 Rhipicephalus simus whereas Rickettsia africae was amplified in 7 Amblyomma hebraeum. Neither C. burnetii nor Bartonella spp. was found in the examined ticks. This study demonstrates the presence of the tick borne pathogen R. massiliae in South Africa (Western Cape and Limpopo provinces), and corroborates the presence of the African tick-bite fever agent (R. africae) in this country (Limpopo province).

Multi-locus genotyping reveals absence of genetic structure in field populations of the brown ear tick (Rhipicephalus appendiculatus) in Kenya.

Rhipicephalus appendiculatus is an important tick vector of several pathogens and parasitizes domestic and wild animals across eastern and southern Africa. However, its inherent genetic variation and population structure is poorly understood. To investigate whether mammalian host species, geographic separation and resulting reproductive isolation, or a combination of these, define the genetic structure of R. appendiculatus, we analyzed multi-locus genotype data from 392 individuals from 10 geographic locations in Kenya generated in an earlier study. These ticks were associated with three types of mammalian host situations; (1) cattle grazing systems, (2) cattle and wildlife co-grazing systems (3) wildlife grazing systems without livestock. We also analyzed data from 460 individuals from 10 populations maintained as closed laboratory stocks and 117 individuals from five other species in the genus Rhipicephalus. The pattern of genotypes observed indicated low levels of genetic differentiation between the ten field populations (FST=0.014±0.002) and a lack of genetic divergence corresponding to the degree of separation of the geographic sampling locations. There was also no clear association of particular tick genotypes with specific host species. This is consistent with tick dispersal over large geographic ranges and lack of host specificity. In contrast, the 10 laboratory populations (FST=0.248±0.015) and the five other species of Rhipicephalus (FST=0.368±0.032) were strongly differentiated into distinct genetic groups. Some laboratory bred populations diverged markedly from their field counterparts in spite of originally being sampled from the same geographic locations. Our results demonstrate a lack of defined population genetic differentiation in field populations of the generalist R. appendiculatus in Kenya, which may be a result of the frequent anthropogenic movement of livestock and mobility of its several wildlife hosts between different locations.

Distribution of endemic and introduced tick species in Free State Province, South Africa.

The distributions of endemic tick vector species as well as the presence of species not endemic to Free State Province, South Africa, were determined during surveys or opportunistic collections from livestock, wildlife and vegetation. Amongst endemic ticks, the presence of Rhipicephalus appendiculatus was confirmed in the north of the province, whilst Rhipicephalus decoloratus was collected at 31 localities mostly in the centre and east, and Ixodes rubicundus at 11 localities in the south, south-west and centre of the province. Amongst the non-endemic species adult Amblyomma hebraeum were collected from white rhinoceroses (Ceratotherium simum) on four privately owned farms, whilst the adults of Rhipicephalus microplus were collected from cattle and a larva from vegetation at four localities in the east of the province. The collection of Rhipicephalus evertsi mimeticus from a sheep in the west of the province is the second record of its presence in the Free State, whereas the presence of Haemaphysalis silacea on helmeted guineafowl (Numida meleagris) and vegetation in the centre of the province represents a first record for this species in the Free State. The first collection of the argasid tick, Ornithodoros savignyi, in the Free State was made from a domestic cow and from soil in the west of the province. The localities at which the ticks were collected have been plotted and the ticks' role in the transmission or cause of disease in domestic livestock and wildlife is discussed.

First description of the immature stages and redescription of the adults of Cosmiomma hippopotamensis (Acari: Ixodidae) with notes on its bionomics.

Cosmiomma hippopotamensis (Denny, 1843) is one of the most unusual, beautiful, and rare tick species known to the world. All stages of this species possess a unique morphology, on the one hand making them easy to identify, while on the other they exhibit similarities to certain species of Amblyomma Koch, 1844, Dermacentor Koch, 1844, and Hyalomma Koch, 1844. Adults of C. hippopotamensis have been collected on only two occasions from their hosts, namely Hippopotamus amphibius L. and Diceros bicornis (L.), and have been recorded from only a few widely separated localities in East and southern Africa. Here, the larva and nymph are described and illustrated for the first time, while the male and female are illustrated and redescribed. Data on hosts, geographic distribution, and life cycle of C. hippopotamensis are also provided.

Micro- and minisatellite-expressed sequence tag (EST) markers discriminate between populations of Rhipicephalus appendiculatus.

Biological differences, including vector competence for the protozoan parasite Theileria parva have been reported among populations of Rhipicephalus appendiculatus (Acari: Ixodidae) from different geographic regions. However, the genetic diversity and population structure of this important tick vector remain unknown due to the absence of appropriate genetic markers. Here, we describe the development and evaluation of a panel of EST micro- and minisatellite markers to characterize the genetic diversity within and between populations of R. appendiculatus and other rhipicephaline species. Sixty-six micro- and minisatellite markers were identified through analysis of the R. appendiculatus Gene Index (RaGI) EST database and selected bacterial artificial chromosome (BAC) sequences. These were used to genotype 979 individual ticks from 10 field populations, 10 laboratory-bred stocks, and 5 additional Rhipicephalus species. Twenty-nine markers were polymorphic and therefore informative for genetic studies while 6 were monomorphic. Primers designed from the remaining 31 loci did not reliably generate amplicons. The 29 polymorphic markers discriminated populations of R. appendiculatus and also 4 other Rhipicephalus species, but not R. zambeziensis. The percentage Principal Component Analysis (PCA) implemented using Multiple Co-inertia Analysis (MCoA) clustered populations of R. appendiculatus into 2 groups. Individual markers however differed in their ability to generate the reference typology using the MCoA approach. This indicates that different panels of markers may be required for different applications. The 29 informative polymorphic micro- and minisatellite markers are the first available tools for the analysis of the phylogeography and population genetics of R. appendiculatus.

The dynamics of questing ticks collected for 164 consecutive months off the vegetation of two landscape zones in the Kruger National Park (1988-2002). Part III. The less commonly collected species.

Despite many studies regarding tick ecology, limited information on long-term changes in tick populations exist. This study assessed the long-term population dynamics of the less frequently collected questing ixodid ticks in the Kruger National Park (KNP). From 1988 to 2002, monthly dragging of the vegetation was performed in three habitats (grassland, woodland and gully) at two sites in the KNP (Nhlowa Road, Landscape Zone 17, and Skukuza, Landscape Zone 4). Amblyomma marmoreum and Rhipicephalus evertsi evertsi were collected as larvae most commonly. Most A. marmoreum larvae were collected at Skukuza and numbers peaked from March to July. More R. evertsi evertsi larvae were collected at Nhlowa Road and numbers peaked in summer and in winter, while at Skukuza there was a single peak in spring. Haemaphysalis elliptica, Rhipicephalus simus and Rhipicephalus turanicus were collected as adults most commonly. More Ha. elliptica and R. turanicus were collected at Nhlowa Road than at Skukuza, while R. simus numbers from the two sites were approximately equal. Ha. elliptica were collected most often between February and June, and R. simus and R. turanicus during February and March. All three species were collected more frequently in gullies than in grassland or woodland. Their numbers increased in 1994/1995 following an eruption of rodents, the preferred hosts of the immature stages. The different host-seeking strategies of ticks largely determine the development stage at which they are likely to be collected during vegetation dragging and reflect a complex interaction between ticks, their hosts and the environment.

Parasites of domestic and wild animals in South Africa. XLVIII. Ticks (Acari: Ixodidae) infesting domestic cats and wild felids in southern Africa.

Ticks collected from domestic cats (Felis catus), cheetahs (Acinonyx jubatus),caracals (Caracal caracal), African wild cats (Felis lybica), black-footed cats (Felis nigripes), a serval (Leptailurus serval), lions(Panthera leo), and leopards (Panthera pardus) were identified and counted. Thirteen species of ixodid ticks and one argasid tick were identified from domestic cats and 17 species of ixodid ticks from wild felids. The domestic cats and wild felids harboured 11 ixodid species in common. The adults of Haemaphysalis elliptica, the most abundant tick species infesting cats and wild felids, were most numerous on a domestic cat in late winter and in mid-summer, during 2 consecutive years. The recorded geographic distribution of the recently described Haemaphysalis colesbergensis, a parasite of cats and caracals, was extended by 2 new locality records in the Northern Cape Province, South Africa.

Parasites of domestic and wild animals in South Africa. XLVII. Ticks of tortoises and other reptiles.

A total of 586 reptiles, belonging to 35 species and five subspecies, were examined in surveys aimed at determining the species spectrum and geographic distribution of ticks that infest them. Of these reptiles 509 were tortoises, 28 monitor or other lizards, and 49 snakes. Nine ixodid tick species, of which seven belonged to the genus Amblyomma, and one argasid tick, Ornithodoros compactus were recovered. Seven of the ten tick species are parasites of reptiles. Amongst these seven species Amblyomma marmoreum was most prevalent and numerous on leopard tortoises, Geochelone pardalis; Amblyomma nuttalliwas present only on Bell's hinged tortoises, Kinixys belliana; and most Amblyomma sylvaticum were collected from angulate tortoises, Chersina angulata. Amblyomma exornatum (formerly Aponomma exornatum) was only recovered from monitor lizards, Varanus spp.; most Amblyomma latum (formerly Aponomma latum) were from snakes; and a single nymph of Amblyomma transversale (formerly Aponomma transversale) was collected from a southern African python, Python natalensis. All 30 Namaqualand speckled padloper tortoises, Homopus signatus signatus, examined were infested with O. compactus. The seasonal occurrence of A. sylvaticum and the geographic distribution of this tick and of A. marmoreum, A. nuttalli, A. exornatum, A. latum and O. compactus are illustrated.

Hosts, seasonality and geographic distribution of the South African tortoise tick, Amblyomma marmoreum.

The tortoise tick Amblyomma marmoreum was collected from large numbers of reptiles and other animals during the course of numerous surveys conducted in South Africa. A total of 1229 ticks, of which 550 were adults, were recovered from 309 reptiles belonging to 13 species, with leopard tortoises, Geochelone pardalis being the most heavily infested. The 269 birds sampled harboured 4901 larvae, 217 nymphs and no adult ticks, and the prevalence of infestation was greatest on helmeted guineafowls, Numida meleagris. Only two larvae were recovered from 610 rodents, including 31 spring hares, Pedetes capensis, whereas 1144 other small mammals yielded 1835 immature ticks, of which 1655 were collected from 623 scrub hares, Lepus saxatilis. The 213 carnivores examined harboured 2459 ticks of which none were adult. A single adult tick and 6684 larvae and 62 nymphs were recovered from 656 large herbivores, and a total of 4081 immature ticks and three adults were collected from 1543 domestic animals and 194 humans. Adult male and female A. marmoreum were most numerous on reptiles during January and February, and larvae during March. The largest numbers of larvae were present on domestic cattle and helmeted guineafowls in the Eastern Cape Province during March or April respectively, whereas larvae were most numerous on helmeted guineafowls, scrub hares and the vegetation in north-eastern Mpumalanga Province during May. In both provinces nymphs were most numerous between October and December. Amblyomma marmoreum appears to be most prevalent in the western regions of the Western and Eastern Cape and Free State provinces, and the north-eastern regions of the Northern Cape, KwaZulu-Natal, Mpumulanga and Limpopo provinces.

Prevalence of theileriosis in Red Hartebeest (Alcelaphus buselaphus caama) in Namibia.

A total of 23 blood samples from Red Hartebeest and 28 ticks of the subspecies Rhipicephalus evertsi were collected in Namibia during the summer 2003. For a Piroplasmida species, Theileria sp 95.7% of blood and 50.0% of tick samples (all R. e. mimeticus, none of R. e. evertsi were carrying the pathogen) were PCR positive. Sequencing showed infection from a re-emerging mild species of Theileria phylogenetically different from other African Theileria species such as Theileria parva, T. buffeli, T. annulata, T. taurotragi or T. mutans.

Ixodid ticks feeding on humans in South Africa: with notes on preferred hosts, geographic distribution, seasonal occurrence and transmission of pathogens.

This paper records the identities of 558 ixodid ticks feeding on 194 humans in South Africa. These ticks belonged to 20 species in six genera and those most frequently encountered were AmblYomma hebraeum, Haemaphysalis leachi, Hyalomma marginatum rufipes, Hyalomma truncatum, Rhipicephalus appendiculatus, Rhipicephalus gertrudae and Rhipicephalus simus. With the exception of the larvae of R. appendiculatus, the incidents of these ticks feeding on humans correlated well with their seasonal occurrences on preferred hosts. Ticks were also collected at monthly intervals, for 14 consecutive months, from the clothing of a game-guard providing protection for field-workers engaged in the collection, by means of flannel strips, of free-living ticks from the vegetation of four localities in the southern region of the Kruger National Park. In addition, with the exception of 3 months when the particular worker was absent, ticks that fed on one of the field-workers were collected over the same 14-month period. A total of 54,429 free-living ticks belonging to 14 species and six genera were collected from the vegetation at the four localities during this time and 3751 ticks belonging to 11 species and six genera from the clothing of the game-guard. The larvae of A. hebraeum and Boophilus decoloratus were the most numerous of the immature ticks, and H. leachi and R. simus of the adults on both the vegetation and the guard's clothing. Ticks fed on the field-worker on six occasions and 14 were collected, all of which were A. hebraeum larvae.