Horak_231-242.indd INTRODUCTION Although numerous collections of ticks have been made from giraffes, Giraffa camelopardalis, African buffaloes, Syncerus caffer and eland, Taurotragus oryx (Theiler 1962; Yeoman & Walker 1967; Baker & Keep 1970; Walker 1974; Norval 1983), it would seem that few attempts have been made to deter- mine the total tick burdens of these large animals. This is perhaps not surprising considering the amount of assistance required to collect ticks from dead or immobilized animals of this size, and the time that has to be spent identifying and counting immature ticks in representative samples of these collections and often all the several thousand adult ticks that a single large herbivore may harbour. Amongst those who have attempted to do so are Horak, Potgieter, Walker, De Vos & Boomker (1983), Rechav, Zeederberg & Zeller (1987), Horak, Fourie, Novellie & Williams (1991a), Horak, Anthonissen, Krecek & Boomker (1992a) and Zieger, Horak, Cauldwell & Uys (1998). Prior as well as subsequent to these published studies the opportunity to sam- 231 Onderstepoort Journal of Veterinary Research, 74:231–242 (2007) Ticks associated with the three largest wild ruminant species in southern Africa I.G. HORAK1*, H. GOLEZARDY2 and A.C. UYS2 ABSTRACT HORAK, I.G., GOLEZARDY, H. & UYS, A.C. 2007. Ticks associated with the three largest wild rumi- nant species in southern Africa. Onderstepoort Journal of Veterinary Research, 74:231–242 The objective of this study was to assess the host status of the three largest southern African wild ruminants, namely giraffes, Giraffa camelopardalis, African buffaloes, Syncerus caffer, and eland, Taurotragus oryx for ixodid ticks. To this end recently acquired unpublished data are added here to already published findings on the tick burdens of these animals, and the total numbers and species of ticks recorded on 12 giraffes, 18 buffaloes and 36 eland are summarized and discussed. Twenty-eight ixodid tick species were recovered. All stages of development of ten species, namely Amblyomma hebraeum, Rhipicephalus (Boophilus) decoloratus, Haemaphysalis silacea, Ixodes pilosus group, Margaropus winthemi, Rhipicephalus appendiculatus, Rhipicephalus evertsi evertsi, Rhipicephalus glabroscutatum, Rhipicephalus maculatus and Rhipicephalus muehlensi were collected. The adults of 13 species, of which the immature stages use small mammals as hosts, namely Haemaphysalis aciculifer, Hyalomma glabrum, Hyalomma marginatum rufipes, Hyalomma truncatum, Ixodes rubi- cundus, Rhipicephalus capensis, Rhipicephalus exophthalmos, Rhipicephalus follis, Rhipicephalus gertrudae, Rhipicephalus lounsburyi, Rhipicephalus lunulatus, Rhipicephalus pravus group and Rhipi- cephalus simus, were also collected. Keywords: African buffaloes, eland, Giraffa camelopardalis, giraffes, ixodid ticks, Syncerus caffer, Taurotragus oryx, tick burdens * Author to whom correspondence is to be directed. E-mail: ivan.horak@up.ac.za 1 Department of Veterinary Tropical Diseases, Faculty of Vet- erinary Science, University of Pretoria, Onderstepoort, 0110 South Africa; Division of Parasitology, Onderstepoort Veter- inary Institute, Onderstepoort, 0110 South Africa; and Depart- ment of Zoology and Entomology, University of the Free State, P.O. Box 339, Bloemfontein, 9300 South Africa 2 Department of Veterinary Tropical Diseases, Faculty of Veter- inary Science, University of Pretoria, Onderstepoort, 0110 South Africa. A.C. Uys’ present address: P.O. Box 652, New- lands, 0049 South Africa Accepted for publication 23 March 2007—Editor 232 Ticks associated with largest wild ruminant species in southern Africa ple more of these large herbivores has arisen and the present paper reports the results of the latter collections. However, since we thought it unlikely that, in the near future, similar exhaustive tick col- lections would be made from such large animals in southern Africa, we decided to combine the earlier published data with the more recent findings in or- der to give a more comprehensive overview of the numbers and species of ticks that may infest gi- raffes, African buffaloes and eland in a variety of habitats. Giraffes occur in several associations of dry savan- na, varying from scrub to woodland. They do not occur in forest or desert and generally are not found in open plains. They are present in north-eastern Limpopo and Mpumalanga Provinces, South Africa, and in the north-eastern parts of Namibia (Skinner & Smithers 1990), but in recent times have been introduced into a number of regions in South Africa in which they previously did not occur. African buffaloes prefer savanna-type habitats and need a plentiful supply of grass, shade and water. They occur in herds, which increase in size in the dry season, but become more fragmented during the wet season because of the usual plentiful sup- ply of water and grazing (Skinner & Smithers 1990). The current distribution of African buffaloes in South Africa is mostly patchy. Large numbers are present in the Kruger National Park (KNP) and the Hluhluwe- iMfolozi Park in the north-eastern regions of the Lim- popo, Mpumalanga and KwaZulu-Natal Prov inces, with smaller populations in national, provincial and privately owned reserves in these and nearly all other provinces of South Africa. Eland are gregarious animals usually occurring in small herds, and are as at home in arid semi-desert scrub associations as they are in montane grass- land (Skinner & Smithers 1990). Historically they were present virtually throughout South Africa, but overexploitation reduced their numbers. However, because they are a sought after species, reintroduc- tions have resulted in their now occupying virtually the same regions in which they originally occurred. One of the serious shortcomings of the past and current studies is that they are biased towards lo- calities and seasons at which animals were, or have been, made available for survey purposes and are thus not necessarily representative of the entire ixodid tick fauna present within the geographic dis- tribution of the host species being examined. Nevertheless a remarkably large number of tick species were recovered. Moreover, because of the large size of the hosts, large numbers of adult ticks were collected (Gallivan & Horak 1997), and the greater certainty with which the adult ticks can be identified has augmented the accuracy of our iden- tification of the immature stages, and hence our re- sults. MATERIALS AND METHODS The localities at which animals were examined, includ ing those of the earlier studies of Horak et al. (1983, 1991a, 1992a), Rechav et al. (1987) and Zie- ger et al. (1998), are summarized in Table 1. With the exception of Free State and Gauteng Provinces at least one of the three large wild ruminant species was sampled in each province of the Republic of South Africa. In addition, six of the giraffes were exam ined in the Etosha National Park in northern Namibia (Horak et al. 1992a), and two of the eland on a game ranch in Central Province, Zambia (Zie- ger et al. 1998). Ticks were recovered from the animals in the earlier surveys as described by the authors of those stud- ies. The carcasses of giraffes, African buffaloes and eland in the present surveys were processed for tick recovery and the ticks identified and counted as de- scribed by Horak, Boomker, Spickett & De Vos (1992b). When animals had been immobilized, and not killed, an attempt was made to collect as many adult ticks as possible, with particular attention be- ing paid to the preferred sites of attachment of the various species. A total of 12 giraffes, 18 African buffaloes and 36 eland were examined, but in the tables devoted to each host species only the results for those animals sampled within a particular tick species’ distribution range are given for that particular tick. RESULTS AND DISCUSSION The species and numbers of ticks collected from the giraffes, buffaloes and eland are summarized in Tables 2, 3 and 4, and the total numbers of male and female ticks recovered and, where it was pos- sible to calculate this, the ratio of males to females for each species are summarized in Table 5. A calculated total of 450 709 ticks, belonging to 28 species and three subspecies, were collected from the 66 animals examined. Of these 57 688 were adults, of which at least half were identified and counted. Although this may seem like a large num- ber of ticks, probably three times as many would 233 I.G. HORAK, H. GOLEZARDY & A.C. UYS have been recovered had the digestion technique of Van Dyk & McKenzie (1992) been used. However, the fine structures of tick larvae and nymphs are of- ten damaged during the period of digestion required in the latter technique, and thus, although tick re- covery may be more complete, it is achieved at the expense of accurate species determination. Two of the 28 species recovered are one-host ticks, namely Margaropus winthemi and Rhipicephalus (Boophilus) decoloratus. Five are two-host ticks, these being Hyalomma glabrum, Hyalomma margi- natum rufipes, Hyalomma truncatum, Rhipicephalus glabroscutatum and Rhipicephalus evertsi of which both subspecies of the latter, namely evertsi and mi- meticus were collected. The remaining 21 species are all three-host ticks. All stages of development of Amblyomma hebraeum, R. (Boophilus) decoloratus, Haemaphysalis silacea, Ixodes pilosus group, M. winthemi, Rhipicephalus appendiculatus, R. evertsi evertsi, R. glabroscutatum, Rhipicephalus maculatus and Rhipicephalus muehlensi were collected. Only the adults of 13 species, namely Haemaphysalis aci- culifer, H. glabrum, H. marginatum rufipes, H. trun- catum, Ixodes rubicundus, Rhipicephalus capensis, Rhipicephalus exophthalmos, Rhipicephalus follis, Rhipicephalus gertrudae, Rhipicephalus lounsburyi, Rhipicephalus lunulatus, Rhipicephalus pravus group and Rhipicephalus simus, of which the imma- ture stages use small mammals as hosts, were re- covered. In addition the larvae and nymphs of the South African tortoise tick, Amblyomma marmoreum, nymphs and adults of the tropical bont tick, Ambly- omma variegatum, a single male Rhipicephalus longiceps, 30 Rhipicephalus supertritus males and 16 Rhipicephalus zambeziensis nymphs were re- TABLE 1 Localities at which large wild ruminants were examined for ixodid ticks Country and province Locality Co-ordinates Vegetation type (Van der Merwe 1983; White 1983; Acocks 1988) South Africa Western Cape West Coast National Park 33°06’ S, 17°59’ E Strandveld and patches of Coastal Fynbos Eastern Cape Thomas Baines Nature Reserve 33°23’ S, 26°28’ E False Macchia, Eastern Province Thornveld and Valley Bushveld Andries Vosloo Kudu Reserve 33°07’ S, 26°40’ E Valley Bushveld Mountain Zebra National Park 32°15’ S, 25°27’ E Karroid Merxmeullera Mountain Veld, replaced by Karoo on the higher slopes and Karroid Broken Veld in the north Northern Cape Kgalagadi Transfrontier Park 24°45’–26°28’ S, 20°00’–20°50’ E Lightly wooded grassland on dune crests and grassland in depres- sions between dunes KwaZulu-Natal Eastern Shores Park 28°08’ S, 32°30’ E Zululand Palm Veld, subdivision of Coastal Thornveld and Coastal communities iMfolozi Nature Reserve 28°15’ S, 31°57’ E Zululand Thornveld and Lowveld North West SA Lombaard Reserve 27°35’ S, 25°29’ E Dry Cymbopogon-Themeda veld Mpumalanga Pretoriuskop (KNP) 25°10’ S, 31°16’ E Lowveld Sour Bushveld Lower Sabie (KNP) 25°07’ S, 31°55’ E Lowveld Mtethomusha Nature Reserve 25°29’ S, 31°17’ E Lowveld Namibia Okaukuejo, Etosha National Park 19°11’ S, 15°55’ E Mopane savanna Zambia Central Mtendere Game Ranch 15°05’ S, 28°16’ E Miombo Woodland KNP = Kruger National Park 234 Ticks associated with largest wild ruminant species in southern Africa covered. Ten of the 28 tick species recovered occur only in South Africa. They are H. silacea, H. glabrum, I. pilosus, I. rubicundus, M. winthemi, R. capensis, R. follis, R. glabroscutatum and R. lounsburyi. With the exception of ticks of the I. pilosus group and I. rubicundus, of which more females than males were recovered, considerably more males than fe- males of all other species, of which both sexes were present, were collected. Mating in Ixodes species may occur in the preparasitic phase of the life cycle (Fourie & Horak 1994), and many males thus never attach. This behaviour is probably largely responsi- ble for the female-biased parasitic populations of the two ticks in this genus in the present study. The male-biased populations of the other tick species is due to the propensity of parasitic male ixodid ticks of several species to spend extended periods on their hosts (Londt 1976; Jordaan & Baker 1981), whereas the females may require only a few days to two weeks to mate and engorge before detaching. Furthermore the larger size of engorging females is likely to make them more susceptible to removal by grooming or predation than the considerably smaller males. Amblyomma species The larvae of Amblyomma hebraeum infest a wide variety of hosts, including ungulates, carnivores, lago morphs and birds, and all stages of develop- ment, but particularly the adults, infest very large hosts (Horak, MacIvor, Petney & De Vos 1987a). The males may remain attached for several months (Jordaan & Baker 1981), thus augmenting the al- ready male-biased population of adult ticks. Ambly- TABLE 2 Giraffes examined and the number infested within the distribution range of each tick Tick species Hosts examined (infested) Life stage Total No. of ticks Locality Amblyomma hebraeum 6 (6) Larvae Nymphs Males Females 773 554 2 406 512 KNP Hyalomma marginatum rufipes 12 (8) Males Females 374 77 KNP, Etosha Hyalomma truncatum 12 (12) Males Females 1 649 619 KNP, Etosha Rhipicephalus (Boophilus) decoloratus 6 (6) Larvae Nymphs Males Females 2 472 2 538 1 503 821 KNP Rhipicephalus appendiculatus 6 (6) Larvae Nymphs Males Females 168 682 49 4 KNP Rhipicephalus evertsi evertsi 6 (6) Larvae Nymphs Males Females 452 190 106 20 KNP Rhipicephalus evertsi mimeticus 6 (5) Males Females 14 5 Etosha Rhipicephalus longiceps 6 (1) Males 1 Etosha Rhipicephalus pravus group 6 (3) Males Females 32 20 KNP Rhipicephalus simus 6 (4) Males Females 22 6 KNP KNP = Kruger National Park 235 I.G. HORAK, H. GOLEZARDY & A.C. UYS omma hebraeum is present in the Bushveld and Low veld regions of northern, eastern and south- eastern South Africa (Walker & Olwage 1987). Amblyomma marmoreum is known as the South African tortoise tick, and all stages of development may be found on a variety of these reptiles, with leo- pard tortoises, Geochelone pardalis, generally the most heavily infested (Horak, McKay, Heyne & Spick- ett 2006). The larvae infest a wide variety of mam- mals and birds, and are often found on mammals and birds of several species that are present within the tick’s distribution range (Horak et al. 2006). Amblyomma variegatum does not occur in South Africa, but is present in most sub-Saharan countries TABLE 3 African buffaloes examined and the number infested within the distribution range of each tick Tick species Hosts examined (infested) Life stage Total No. of ticks Locality Amblyomma hebraeum 18 (18) Larvae Nymphs Males Females 47 815 6 437 6 154 1 764 KNP, Eastern Shores, iMfolozi, Thomas Baines, Mtethomusha Amblyomma marmoreum 18 (4) Larvae Nymphs 8 26 KNP, Eastern Shores, iMfolozi, Thomas Baines, Mtethomusha Haemaphysalis silacea 15 (7) Larvae Nymphs Males Females 81 357 135 78 Eastern Shores, iMfolozi, Thomas Baines Hyalomma truncatum 17 (2) Males Females 6 0 KNP, iMfolozi, Mtethomusha Rhipicepalus (Boophilus) decoloratus 18 (11) Larvae Nymphs Males Females 1 640 290 248 160 Eastern Shores, iMfolozi, Thomas Baines, Mtethomusha Rhipicephalus appendiculatus 18 (17) Larvae Nymphs Males Females 139 389 32 715 1 360 774 KNP, Eastern Shores, iMfolozi, Thomas Baines, Mtethomusha Rhipicephalus evertsi evertsi 18 (15) Larvae Nymphs Males Females 721 35 96 45 KNP, Eastern Shores, iMfolozi, Thomas Baines, Mtethomusha Rhipicephalus follis 1 (1) Males Females 40 32 Thomas Baines Rhipicephalus maculatus 14 (14) Larvae Nymphs Males Females 15 379 16 772 946 398 Eastern Shores, iMfolozi Rhipicephalus muehlensi 16 (15) Larvae Nymphs Males Females 11 775 495 193 141 Eastern Shores, iMfolozi, Mtethomusha Rhipicephalus simus 18 (9) Males Females 121 64 KNP, Eastern Shores, iMfolozi, Mtethomusha KNP = Kruger National Park 236 Ticks associated with largest wild ruminant species in southern Africa TABLE 4 Eland examined and the number infested within the distribution range of each tick Tick species Hosts exam- ined (infested) Life stage Total No. of ticks Locality Amblyomma hebraeum 4 (4) Larvae Nymphs Males Females 32 394 2 405 3 488 530 KNP, Andries Vosloo Kudu Reserve, Thomas Baines NR Amblyomma marmoreum 15 (8) Larvae 167 KNP, Mountain Zebra NP, AV Kudu Reserve, Thomas Baines NR Amblyomma variegatum 2 (2) Nymphs Males Females 4 60 26 Mtendere Game Ranch Haemaphysalis aciculifer 2 (1) Males 2 KNP Haemaphysalis silacea 4 (4) Larvae Nymphs Males Females 7 686 933 1 409 386 Andries Vosloo Kudu Reserve, Thomas Baines NR Hyalomma glabrum 11 (11) Males Females 955 191 Mountain Zebra NP Hyalomma marginatum rufipes 16 (16) Males Females 907 470 Kgalagadi Transfrontier Park, Thomas Baines NR, SA Lombaard NR Hyalomma truncatum 32 (27) Males Females 1 702 650 Mountain Zebra NP, Kgalagadi Transfrontier Park, West Coast NP, SA Lombaard NR Ixodes pilosus group 4 (4) Larvae Nymphs Males Females 2 817 177 22 64 Andries Vosloo Kudu Reserve, Thomas Baines NR Ixodes rubicundus 11 (7) Males Females 34 81 Mountain Zebra NP Margaropus winthemi 11 (9) Larvae Nymphs Males Females 12 792 6 874 1 915 893 Mountain Zebra NP Rhipicephalus (Boophilus) decoloratus 8 (5) Larvae Nymphs Males Females 3 106 3 988 3 581 848 KNP, AV Kudu Reserve, Thomas Baines NR, Mtendere Game Ranch Rhipicephalus appendiculatus 8 (8) Larvae Nymphs Males Females 18 025 1 489 8 106 3 345 KNP, AV Kudu Reserve, Thomas Baines NR, Mtendere Game Ranch Rhipicephalus capensis 2 (2) Males Females 1 506 408 West Coast NP Rhipicephalus evertsi evertsi 34 (31) Larvae Nymphs Males Females 6 900 2 128 1 574 392 KNP, Mountain Zebra NP, AV Kudu Reserve, Thomas Baines NR, West Coast NP, SA Lombaard NR, Mtendere Game Ranch 237 I.G. HORAK, H. GOLEZARDY & A.C. UYS to the north (Walker & Olwage 1987). Its host spec- trum is similar to that of A. hebraeum and its pres- ence on the two eland examined in Zambia is thus to be expected. Haemaphysalis species The two male Haemaphysalis aciculifer recovered from an eland in the KNP during September 1979 (Horak et al. 1983; Table 4), were the only speci- mens of this species reported in the park until 2000, when Horak, Braack, Fourie & Walker (2000) re- corded 13 males and two females from a honey badger, Mellivora capensis. This tick is seldom re- covered in large numbers but appears to be present in north-eastern Mpumalanga Province and thence along the eastern and southern coastal and adjacent inland regions to the south-western region of the Cape Province (Theiler 1962; Horak, Keep, Spickett & Boomker 1989; Horak & Boomker 1998). Haemaphysalis silacea is present in north-eastern KwaZulu-Natal and in the Eastern Cape Provinces (Walker 1991), and is associated with vegetation classified as Valley Bushveld (Acocks 1988) and with greater kudus, Tragelaphus strepsiceros, com- mon duikers, Sylvicapra grimmia, Cape grysbok, Rhaphicerus melanotis, and helmeted guineafowls, Numida meleagris (Horak & Knight 1986; MacIvor & Horak 2003). Seven of the 18 buffaloes and all four eland examined in Valley Bushveld-type vegetation in the north-eastern KwaZulu-Natal and the Eastern Cape Provinces were infested. Hyalomma species Delpy (1949) described the tick now known as Hya- lomma glabrum as Hyalomma rufipes glabrum, and the latter name persisted until Theiler (1956) raised it to species level as Hyalomma glabrum. In the same year Hoogstraal (1956) synonymized it with Asian Hyalomma marginatum turanicum, and since then it has been considered identical to the latter tick. Recent studies have, however, revealed that it is a separate taxon, and Apanaskevich & Horak (2006) have subsequently reinstated it as Hyalomma glabrum. Howell, Walker & Nevill (1978) have illus- trated its distribution (as H. m. turanicum) in the central and western regions of South Africa. The adults prefer large animals such as eland and Cape mountain zebras, Equus zebra zebra, and its imma- ture stages infest scrub hares, Lepus saxatilis, and ground-frequenting birds (Apanaskevich & Horak 2006). Tick species Hosts exam- ined (infested) Life stage Total No. of ticks Locality Rhipicephalus exophthalmos 11 (1) Males Females 0 2 Mountain Zebra NP Rhipicephalus follis 11 (11) Males Females 983 447 Mountain Zebra NP, Thomas Baines NR Rhipicephalus gertrudae 2 (1) Males Females 4 2 West Coast NP Rhipicephalus glabroscutatum 14 (13) Larvae Nymphs Males Females 6 510 2 846 868 499 Mountain Zebra NP, Andries Vosloo Kudu Reserve, West Coast NP Rhipicephalus lounsburyi 11 (2) Males 3 Mountain Zebra NP Rhipicephalus lunulatus 2 (2) Males Females 218 48 Mtendere Game Ranch Rhipicephalus simus 6 (4) Males Females 39 5 KNP, AV Kudu Reserve, Thomas Baines NR Rhipicephalus supertritus 2 (2) Males 30 Mtendere Game Ranch Rhipicephalus zambeziensis 2 (1) Nymphs 16 KNP KNP = Kruger National Park NP = National Park NR = Nature Reserve TABLE 4 Cont. 238 Ticks associated with largest wild ruminant species in southern Africa The distribution of Hyalomma marginatum rufipes is more extensive than that of H. glabrum, which it al- most entirely overlaps (Howell et al. 1978). Its host preferences are the same as those of the latter tick (Walker 1991), but include giraffes and buffaloes, which are generally not present within the distribu- tion range of H. glabrum (Tables 2 and 3). With the exception of the eastern coastal regions and some adjacent inland areas and the southern coastal regions, Hyalomma truncatum is present vir- tually throughout the country (Howell et al. 1978). The adults are found on the same large hosts as the former two ticks, and its immature stages infest scrub hares and rodents (Walker 1991). Ixodes species Ticks of the Ixodes pilosus group are present in the southern coastal and adjacent regions of the Western and Eastern Cape Provinces and in south- eastern KwaZulu-Natal Province, where all stages of development may be encountered on antelopes, caracals, Caracal caracal and domestic dogs (Horak, Jacot Guillarmod, Moolman & De Vos 1987b; Horak et al. 1989; Horak & Boomker 1998; Horak & Mat- thee 2003). The colloquial name given to Ixodes rubicundus is Karoo paralysis tick, because of the paralysis as- sociated with infestation of small domestic livestock in the Karoo regions of the country (Howell et al. 1978; Fourie & Horak 1994). The hosts of the adults are wild and domestic ruminants, caracals and do- mestic dogs (Horak et al. 1987b, 1991a; Fourie & Horak 1993; Horak & Matthee 2003). The immature stages infest Smith’s red rock rabbits, Pronolagus ru pestris, caracals and rock elephant shrews, Ele- phan tulus myurus (Horak et al. 1991a; Fourie, Horak & Woodall 2005). Margaropus winthemi This tick is commonly known as the winter horse tick, and its distribution in the cooler higher-lying re- gions of the country suggests that its original hosts TABLE 5 Male to female ratios of tick species collected from giraffes, African buffaloes and eland in southern Africa Tick species Total No. of adult ticks Male:female ratio Male Female Male Female Amblyomma hebraeum Amblyomma variegatum Haemaphysalis aciculifer Haemaphysalis silacea Hyalomma glabrum Hyalomma marginatum rufipes Hyalomma truncatum Ixodes pilosus group Ixodes rubicundus Margaropus winthemi Rhipicephalus (Boophilus) decoloratus Rhipicephalus appendiculatus Rhipicephalus capensis Rhipicephalus evertsi evertsi Rhipicephalus evertsi mimeticus Rhipicephalus exophthalmos Rhipicephalus follis Rhipicephalus gertrudae Rhipicephalus glabroscutatum Rhipicephalus longiceps Rhipicephalus lounsburyi Rhipicephalus lunulatus Rhipicephalus maculatus Rhipicephalus muehlensi Rhipicephalus pravus group Rhipicephalus simus Rhipicephalus supertritus 12 048 60 2 1 544 955 1 281 3 357 22 34 1 915 5 332 9 515 1 506 1 776 14 0 1 023 4 868 1 3 218 946 193 32 182 30 2 806 26 0 464 191 547 1 269 64 81 893 1 829 4 123 408 457 5 2 479 2 499 0 0 48 398 141 20 75 0 4.29 2.31 – 3.33 5.00 2.34 2.65 0.34 0.42 2.14 2.92 2.31 3.69 3.89 2.80 – 2.14 2.00 1.74 – – 4.54 2.38 1.37 1.60 2.43 – 1.0 1.0 – 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 – 1.0 1.0 1.0 – – 1.0 1.0 1.0 1.0 1.0 – Total (excluding Ixodes spp.) 42 805 14 682 2.92 1.0 239 I.G. HORAK, H. GOLEZARDY & A.C. UYS were Cape mountain zebras, on which it still occurs in very large numbers during the winter months (Horak et al. 1991a). The burdens of the eland (Ta- ble 4), examined at the same time as zebras in the Mountain Zebra National Park, although large, did not nearly approach those of the latter animals. Rhipicephalus species Rhipicephalus (Boophilus) decoloratus. Using molec- ular analyses supported by an analysis of morpho- logical characters Murrell, Campbell & Barker (2000) and Beati & Keirans (2001) decided that the genus Rhipicephalus is paraphyletic with respect to the genus Boophilus and consequently Murrell & Barker (2003) proposed the use of the above nomenclature. We have chosen to follow their proposal, although many people prefer to retain Boophilus as a generic name. Giraffes and eland can generally be consid- ered as good hosts of this tick, whereas Afri can buf- faloes appear to be resistant to infestation, with more than half of the adult ticks reported here com- ing from a single, approximately 6-month-old calf. Norval (1984) and Horak, Golezardy & Uys (2006) have discussed the apparent resistance of African buffaloes to artificial and natural infestations with this tick, and it would seem that this resistance is acquired, rather than innate. Adult Rhipicephalus appendiculatus are common parasites of African buffaloes, eland, greater kudus, male nyalas, Tragelaphus angasii, and domestic cat- tle in wooded savannas from south-eastern South Africa in the south to Kenya and Uganda in the north (Walker, Keirans & Horak 2000). The immature stages are found on the same hosts as the adults, but also infest smaller antelopes and scrub hares (Walker et al. 2000). The eland included in the pres- ent study and more particularly those from Zambia, were heavily infested with adult ticks (Table 4). The buffaloes, which were examined mainly during the early winter months, harboured very large numbers of larvae (Table 3). Rhipicephalus capensis, Rhipicephalus follis, Rhipi- cephalus gertrudae and Rhipicephalus simus have several morphological characteristics in common and also have similar life cycles during which the immature stages infest murid rodents and the adults the larger species of antelopes (Walker et al. 2000). Adult R. gertrudae also infest domestic dogs and primates, including humans (Brain & Bohrmann 1992; Horak, Fourie, Heyne, Walker & Needham 2002; Horak & Matthee 2003), while those of R. simus infest domestic and wild equids, suids and carnivores (Horak et al. 1987b, 2000; Walker et al. 2000). Rhipicephalus capensis occurs almost exclu- sively in the western coastal regions of the West ern Cape Province, South Africa; R. gertrudae is pres- ent in the same western regions, but its distribu tion extends east to the southern and southern-central regions of the country and north into Namibia; R. follis is only found in South Africa and is present to the east and north-east of R. gertrudae’s distribu- tion range and generally in mountainous terrain; while, with the exception of the more arid regions of central and western South Africa and southern Na- mibia, the distribution range of R. simus effectively overlays those of the other three species and ex- tends north into Africa to a latitude of approximately 9° south. Rhipicephalus evertsi evertsi and Rhipicephalus evertsi mimeticus are two-host ticks and all stages of development may infest the same host species. Excluding the deserts and regions of high rainfall, the former tick is found throughout sub-Saharan Africa, whereas the latter is confined to the arid and semi-arid regions of Namibia, western Botswana and parts of Angola (Walker et al. 2000). The adults of Rhipicephalus exophthalmos parasit- ize antelopes, domestic ruminants and scrub hares in the south-eastern and north-western regions of South Africa and in a broad, central band from the south to the north of Namibia (Walker et al. 2000). Its immature stages prefer elephant shrews, El ephan- tulus spp., as hosts (Fourie et al. 2005). The two-host tick, Rhipicephalus glabroscutatum, is present in the fynbos and Karoo regions of the Western Cape Province, and also in the Karoo and the Valley Bushveld regions of the Eastern Cape Province (Walker et al. 2000). It is a common para- site of the feet and lower legs of small and large antelope in these regions (Horak & Boomker 1998; MacIvor & Horak 2003). Infestation with its adults is a contributory cause of foot abscess in domestic goats in the Valley Bushveld regions of the Eastern Cape Province (MacIvor & Horak 1987). The distribution in South Africa of Rhipicephalus maculatus and Rhipicephalus muehlensi is confined to the coastal bush and adjacent inland regions of north-eastern KwaZulu-Natal (Walker et al. 2000). The recovery of an R. maculatus nymph from the vegetation and adults from an elephant, Loxodonta africana, in the southern regions of the KNP (Braack, Maggs, Zeller & Horak 1995), and of R. muehlensi adults from one of the buffaloes examined in the Mtethomusha Nature reserve, just south of the KNP, suggest that they have been introduced into these 240 Ticks associated with largest wild ruminant species in southern Africa reserves on animals translocated from north-east- ern KwaZulu-Natal. All stages of development of both ticks may infest the same host species, but the adults of R. maculatus prefer thick-skinned animals such as buffaloes and bush pigs, Potamochoerus larvatus, and those of R. muehlensi nyalas (Horak, Boomker & Flamand 1991b; 1995). Rhipicephalus longiceps is a rarely encountered tick and apparently is present only in certain regions of Namibia and Angola (Walker et al. 2000). Rhipicephalus lounsburyi prefers the higher moun- tainous regions of the Eastern Cape Province (Walker et al. 2000) and two eland in the Mountain Zebra National Park were infested. The adults attach around the feet of their antelope and sheep hosts, while the only known hosts of its immature stages are four-striped grass mice, Rhabdomys pu milio, from which a larva and two nymphs have been col- lected (Walker et al. 2000; Horak, Fourie & Braack 2005). Although Rhipicephalus lunulatus is fairly wide- spread in sub-Saharan Africa its distribution in South Africa is limited to the eastern regions (Walker et al. 2000). The adults have a wide host range and would seem to attach around the feet and lower legs of their hosts (Walker et al. 2000). The eland exam- ined in Zambia had fairly large burdens of adult ticks (Table 4), while a multimammate mouse, Mastomys sp., and a scrub hare examined on the same ranch as the eland were infested with nymphs (Zieger et al. 1998). Ticks belonging to the Rhipicephalus pravus group have been collected from scrub hares in the KNP (Horak, Spickett, Braack & Penzhorn 1993), and were assigned to Rhipicephalus sp. near pravus by Walker et al. (2000). However, the ticks collected in the same park from three of the giraffes, appear to us to be very similar, if not identical to the true R. pravus of East Africa. The immature stages of Rhipicephalus supertritus are unknown, but are assumed to be similar in ap- pearance to those of R. appendiculatus. It is appar- ently commonest in Central Africa, including north- ern Zimbabwe and Mozambique, parts of Zambia and southern Tanzania (Walker et al. 2000), and both eland examined in Zambia were infested. ACKNOWLEDGEMENTS We are most grateful to Ezemvelo KZN Wildlife, SANParks, and the Provincial Division of Nature Conservation of the Eastern Cape Province for plac- ing the animals included in the recent surveys at our disposal, and for providing assistance and facilities to process them for tick recovery. We are particu- larly indebted to Messrs Johan Sithole and Eddie Williams for their assistance in the latter respect. The University of Pretoria and the National Research Foundation provided funds for the conduct of this project. 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