PARASITES OF DOMESTIC AND WILD ANIMALS IN SOUTH AFRICA. XXI. ARTHROPOD PARASITES OF VAAL RIBBOK, BONTEBOK AND SCRUB HARES IN THE WESTERN CAPE PROVINCE

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1 OnderstepoonJ. vet. Res., 53, (1986) PARASITES OF OMESTIC AN WIL ANIMALS IN SOUTH AFRICA. XXI. ARTHROPO PARASITES OF VAAL RIBBOK, BONTEBOK AN SCRUB HARES IN THE WESTERN CAPE PROVINCE I. G. HORAK >, K. SHEPPEY< 2 >, M. M. KNIGHT >, and C. L. BEUTHIN< 3 > ABSTRACT _ HORAK, I. G., SHEPPEY, K., KNIGHT, M. M. & BEUTHIN, C. L., Parasites of domesti and wild animals in South Afria. XXI. Arthropod parasites of vaal ribbok, bontebok and srub hares in the western.cape Provine. Onderstepoort Journal of Veterinary Researh, 53, (1986) Vaal ribbok (Pelea apreolus), bontebok (amalisus doras doras) and srub hares (Lepus saxatilis) were slaughtered in the Bontebok National Park at regular intervals and ex;unined for arthropod parasites. Twelve speies of ixodid tiks were reovered; the antelope eah harboured 9 speies and the hares 11. Rhipiephalus nitens was the most prevalent and most abundant tik. The vaal ribbok also harboured large numbers of Ixodes pilosus and the bontebok fairly large numbers of Rhipiephalus glabrosutatum. The vaal ribbok were infested with 2 lie speies and the larvae of 3 oestrid fly speies. The bontebok harboured 2 lie speies and the larvae of 2 oestrid flies, and the srub hares were infested with a louse speies and a flea speies. The host preferene and seasonal abundane of some of the parasites were determined. INTROUCTION The parasites of 3 antelope speies found in the Bontebok National Park (34 2' S, 2 25' E, Alt. 9-2 m) situated in the south-western Cape Provine, have been the subjets of onsiderable study in the past few years (Verster, Imes & Smit, 1975; Boomker, Horak & e Vos, 1981; Horak, Meltzer & e Vos, 1982; Horak, Brown, Boomker, evos & Van Zyl, 1982; Horak, e Vos & e Klerk, 1982; Boomker, Horak, Gibbons & e Vos, 1983). This researh was aimed at determining the speies omposition and abundane of the arthropod and helminth fauna of the bontebok, amalisus doras doras; the vaal ribbok, Pelea apreolus, and the springbok, Antidoras marsupia/is, in the park. However, it did not give an indiation of the seasonal abundane of the various parasites reovered. A year-long study of the biology of the vaal ribbok within the park neessitated the regular slaughter of some animals. So as not to waste valuable researh material, these animals were proessed for the reovery of arthropod and helminth parasites. At the same time bontebok, srub hares (Lepus saxatilis), striped mie (Rhabdomys pumilio), and vlei rats (Otomys irroratus) were killed and examined for parasites. MATERIALS AN METHOS The physiography of the Bontebok National Park has been desribed by Boomker et al. (1981), but sine that date the springbok population, whih then numbered 83, has been redued to 5 animals. Four vaal ribbok and 2 bontebok were shot at 2- monthly intervals from February 1983 until February On eah oasion an attempt was made to shoot a juvenile, a sub-adult and an adult male and female vaal ribbok and a juvenile and adult bontebok. One vaal ribbok killed by a ar just outside the park and 2 additional vaal ribbok and 2 bontebok from within the park were also examined. Parasites from these animals were reovered, ounted and identified as desribed by Malan, Reineke & Sialdo (1981); Horak, Brown, Boomker, (I) Tik Researh Unit, Rhodes University, Grahamstown 614 < 2 ) epartment of Zoology and Entomology, Rhodes University, Grahamstown 614 <J) Computer Centre, Rhodes University, Grahamstown 614 Reeived 2 July Editor evos & VanZyl (1982) and Horak, Potgieter, Walker, e Vos & Boomker (1983)! One eye of eah animal was plaed in formalin and later examined under a stereosopi mirosope for I st stage Gedoelstia sp. larvae, and the subsutaneous side of the skin of eah animal was examined for the presene of Strobiloestrus sp. lar _vae. The immature stages of the ixodid tiks Hyalomma trunatum, Ixodes pilosus and Rhipiephalus nitens were identified by omparison with laboratoryreared speimens. Exept for April 1983, when only 1 srub hare was obtained, 2 srub hares were shot at night. at 2-monthly intervals from April1983 until February Etoparasites were reovered from these animals by plaing the whole hare, with the skin intat, in a sturdy plasti bag immediately after it had been shot and adding suffiient tik-detahmg agent* to immerse the animal. The hare was left in the bag until the following morning, then thoroughly srubbed with a brush with 2 mm long steel bristles and washed, partiular attention being paid to the external ear anals, nek and the feet. The srubbings and washings plus the ontents of the plasti bag were poured onto a sieve with 15 -tm apertures. The material that was retained was then olleted, preserved with formalin and stored. A total of 19 striped mie olleted in the park from April-Otober 1983, and 9 v lei rats olleted in the park from April-eember 1983, were examined in a similar fashion to the srub hares. Rainfall and minimum and maximum atmospheri temperatures were reorded daily in the park. RESULTS The total numbers of arthropod J'arasites reovered from the vaal ribbok, bontebok an srub hares, their relative abundane and the perentage of animals infested are summarized in Tables 1-3. A total of 12 tik speies was reovered, of whih eah antelope speies harboured 9 and the hares 11. The larvae of 3 oestrid fly S_P:eies were reovered, the vaal ribbok being infested wtth all 3 and the bontebok with 2- of these. The vaal ribbok and the bontebok were eah infested with 2 louse speies and the srub hares with 1 speies. The srub hares also harboured 1 flea speies. The vaal ribbok killed outside the park harboured small Triatix: Coopers SA (Pty) Ltd 187

2 PARASITES OF OMESTIC AN WIL ANIMALS IN SOUTH AFRICA. XXI TABLE 1 The arthropod parasites reovered from 3 vaal ribbok in the Bontebok National Park Arthropod speies Total numbers of arthropods reovered Adults Ixodid tiks Larvae Nymphae Total abundane Males Females Amblyomma marmoreum ,75 Boophilus sp. 2 2,2 Haemaphysalis aiulifer 5(1) 5,4 Ixodes pilosus (22) ,79 Rhipiephalus evertsi evertsi 2 2 4,3 Rhipiephalus gertrudae 4 2 6,4 Rhipiephalus glabrosutatum ,73 Rhipiephalus nitens (6) ,16 Rhipiephalus sp ,44 Total , Lie Nymphae Adults Total abundane amalinia pelea ,59 Linognathus peleus ,41 Total , Oestrid fly larvae 1st stage 2nd stage 3rd stage Total abundane Gedoelstia Sf * ,2 Oestrus ovis * 14 6[1] [5] 2[6] 9,13 Strobiloestrus sp ,67 Total 7 117[1] 32[5] 219[6] 1, Perentage of vaal ribbok infested 36,7 3,3 1, 1, 6,7 6,7 63,3 1, 3, 1, 3,3 1,7 26,9 6, ( ) = Number of maturing female tiks, i.e. idiosoma of H. aiulifer > 4,5 mm and of I. pilosus andr. nitens > 5, mm in length [ ] = ead larvae.. = Eyes of 28 animals examined... = 26 animals examined TABLE 2 The arthropod parasites reovered from 16 bontebok in the Bontebok National Park Arthropod speies Total numbers of arthropods reovered Adults Ixodid tiks Larvae Nymphae Males Females Total Amblyomma marmoreum Haemaphysalis aiulifer 4 4(1) 8 Hyalomma trunatum 1 1 Ixodes pilosus Rhipiephalus evertsi evertsi Rhipiephalus gertrudoe Rhipiephalus glabrosutatum (4) Rhipiephalus nitens (12) Rhipiephalus sp Total Lie Nymphae Adults Total amalinitl sp Litwgnathus sp Total Oestrid fly larvae 1st stage 2nd stage 3rd stage Total Gedoelstitl sp Strobiloestrus sp. 1 1 Total abundane,25,6,1 2,66,11,3 12,69 83,96,23 1, abundane 63,9 36,1 1, abundane 99,94,6 1, Perentage ofbontebok infested ( ) = Number of maturing female tiks, i.e. idiosomaofh. aiulifer and of R. glabrosutatum > 4,5 nun and of R. nitens > 5, nun in length ,5 25, 6,3 93,8 6,3 6,3 93,8 1, 37,5 93,8 56,3 1, 6,3

3 TABLE 3 The arthropod parasites reovered from 11 srub hares in the Bontebok National Park I. G. HORAK, K. SHEPPEY, M. M. KNIGHT & C. L. BEUTHIN Arthropod speies Total numbers of arthropods reovered Perentage Adults of hares Ixodid tiks Larvae Nymphae Total abundane infested Males Females Amblyomma marmoreum ,63 45,5 Boophilus sp. 5 5,39 9,1 Haemaphysalis aiulifer 2 2,16 9,1 Haemaphysalis leahi 7 1 8,62 9,1 Hyalomma trunatum ,16 27,3 Ixodes pilosus I 9(1) ,22 72,7 Ixodes sp. 1 1,8 9,1 Rhipiephalus evertsi evertsi 1 1 2,16 18,2 Rhipiephalus g/abrosutatum 7 1 8,62 36,4 Rhipiephalus nitens (5) ,57 1, Rhipiephalus sp. 5 5,39 18,2 Total , Lie Nymphae Adults Total abundane Haemodipsus sp , 54,5 Fleas Adults Total abundane Ctenoephalides felis damarensis , 72,7 ( ) = Number of maturing female tiks, i.e. idiosoma of I. pilosus and R. nit ens > 5, mm in length TABLE 4 The ixodid tiks reovered from 19 striped mie in the Bontebok National Park Tik speies Total numbers of tiks reovered Perentage of mie Larvae Nymphae Males Females Total infested Haemaphysalis leahi ,8 Rhipiephalus glabrosutatum 1 1 5,3 Rhipiephalus sp ,8 Total ,6 TABLE 5 The ixodid tiks reovered from 9 vlei rats in the Bontebok National Park Tik speies Larvae Boophilus sp. 1 Haemaphysalis leahi 5 3 Ixodes pilosus 1 Rhipiephalus sp Total Total numbers of tiks reovered Perentage of rats Nymphae Males Females Total infested 1 11,1 8 33,3 1 11, , ,6 numbers of Boophilus deoloratus, Boophilus miroplus, I. pilosus, Rhipiephalus evertsi evertsi, R. nitens, amalinia pelea and 26 2nd stage Strobiloestrus sp. larvae. The total numbers of ixodid tiks reovered from the striped mie and vlei rats are summarized in Tables 4 and 5. The mie were infested with 3 tik speies and the rats with4. Ixodid tiks Beause vaal ribbok, bontebok and srub hares were shot on eah oasion during April, June, August and eember 1983 and during February 1984, the mean tik burdens of these 3 host speies during an 11-month period are omparable. Table 6 is a summary of the mean burdens and the overall ratios of larvae to nymphae to adults of the 3 major tik speies reovered from the 3 host speies from Aprill983- February Ixodes pilosus All the vaal ribbok were infested with I. pilosus and they harboured more larvae, nymphae and adults of this tik than did the other 2 hosts, on whih the mean tik burdens were similar in most respets. More than 11 of female I. pilosus on all vaal ribbok were maturing (Table 1). More female than male I. pilosus were reovered from eah of the host speies. The ratio of parasiti larvae to nymphae to adults was 14,7:5,6:1, (Table 6). The seasonal abundane of I. pilosus on the 3 hosts separately as well as the ombined mean burdens are graphially presented in Fig. 1. No lear pattern of seasonal abundane an be diserned on the individual host speies. The ombined mean burdens reveal that larvae peaked during June, nymphae during August and adults during Otober and eember. 189

4 PARASITES OF OMESTIC AN WIL ANIMALS IN SOUTH AFRICA. XXI TABLE 6 The mean numbers of tiks reovered from vaal ribbok, bontebok and srub hares in the Bontebok National Park during the period Aprili983-February 1984 Tik and host speies Mean numbers of tiks reovered Perentage of hosts Larvae Nymphae Males Females Total infested Ixodes pilosus Vaalribbok 112,4 29,2 2, 1 5,4(,8) 149, 1 1, Bontebok 13,4 1,2,,7 24,3 91,7 Srub hares 7,6 11,6,1,8 2,1 72,7 Total 133,4 51, 2,2 6,9 193,5 Ratios 14,7 :5,6 :1, Rhipiephalus glabrosutatum Vaalribbok 2,9 4,3,3,4 25,9 66,7 Bontebok 79,7 32,8 5,3 2,5(,3) 12,3 91,7 Srub hares,6,1,,,7 36,4 Total 11,2 37,2 5,6 2,9 146,9 Ratios 11,9 :4,4 :1, Rhipiephalus nitens Vaalribbok 27, 35,5 5,9 6,7(1,5) 318,1 1, Bontebok 548,7 237,3 13,2 13,5(1,) 812,7 1, Srub hares 5,5 25,2 7,4 6,6(,4) 89,7 1, Total 869,2 298, 26,5 26,8 I 22,5 Ratios 16,3 :5,6 :1, ( ) = Number of maturing female tiks, i.e. idiosorna of R. glabrosutatum > 4,5 mrn and of I. pilosus and R. nitens > 5, mrn in length Most of the bontebok were infested with R. glabrosutatum, and they also harboured more tiks in eah stage of development than did the other 2 hosts. Only the bontebok harboured maturing female R. glabrosutatum (Table 2), while the srub hares arried no adults of this tik. Approximately twie as many males as females were reovered. The ratio of parasiti larvae to nymphae to adults was 11,9:4,4:1, (Table 6). Peak numbers of immature R. glabrosutatum were present on the vaal ribbok and bontebok from April-August. Too few adults were reovered from the vaal ribbok to aurately determine seasonal abundane on this host, while on the bontebok they reahed a peak from August-eember (Fig. 2). The ombined mean burdens of the vaal ribbok and bontebok revealed the same pattern of seasonal abundane as that observed on the bontebok. Rhipiephalus nitens All the vaal ribbok, bontebok and srub hares were infested with R. nitens. The bontebok harboured the greatest mean numbers of larvae, nymphae and adults. The vaal ribbok harboured onsiderably more larvae than did the srub hares, but the nymphal and adult burdens of these 2 hosts were similar. More than 24 of female R. nitens on the vaal ribbok were maturing ompared with 7,1 on the bontebok and 6,8 on the srub hares (Tables 1-3). The mean total burdens onsisted of slihtly more female than male tiks. The ratio ofj>arasitt larvae to nymphae to adults was 16,3:5,6:1, (Table 6). Larval numbers of R. nitens peaked during April and nymphae during August or Otober on eah of the 3 hosts (Fig. 3). The largest numbers of adults were present on the vaal ribbok and bontebok during February and eember 1983 and February Small numbers of adults were reovered from the srub hares on eah oasion, peak numbers ourring during eember 1983 and February Lie Amongst the vaal ribbok examined there were 6 juveniles and an adult female with bilateral pneumonia. Some of the juveniles and the sik female harboured onsiderably more. pelea than the other vaal ribbok. Table 7 is a summary of the mean lie burdens of these 3 groups of vaal ribbok. The juveniles and the sik female exluded, the seasonal abundane of. pelea on the other animals is graphially illustrated in Fig. 4. Peak burdens of. pelea were reovered during June. The mean amalinia sp. burdens of the 7 juvenile bontebok onsisted of 124 nymphae and 7 adults ompared with 65 nymphae and 22 adults on the 9 ad.ult animals. With the exeption of 1 adult female whth harboured a large number of Linognathus sp., the juvenile bontebok always arried more lie of thts genus than did the adult animals. No pattern of seasonal prevalene ould be determined for the amalinia sp. or Linognathus sp. on the bontebok. Few lie were reovered from the srub hares (Table 3). '. Oestrid fly larvae Gedoelstia sp. The orneas of the eyes of 3 of the 28 vaal ribbok examined for Gedoelstia larvae were infested. Only 1st stage larvae were reovered from these animals. All the bontebok were infested with Gedoelstia larvae, whose mature 3rd stage was larger than those of either Gedoelstia ristata or Gedoelstia hassleri and differed from the latter speies in ertain aspets of its ventral spinulation. Of the 845 1st stage larvae reovered 19 were present on the orneas, 237 in the heart hambers, 589 in the nasal passages and none on the dura of the ranial avity. First stage larvae were reovered from the eyes during April, June, August and eember 1983 and, 19

5 25,_,_, A Vaal ribbok I. G. HORAK, K. SHEPPEY, M. M. KNIGIIT & C. L. BEUTHIN 2 Larvae Nymphae A--4 Adults Q- 1 llll/pkepijiiju,.,._,,,., A Vaal ribbok w ".g 6 4 rn 2 Larvae o--o Nymphae A--4 Adulls- 12 [ w u. ::;; w u 6. 4 ::;; 15 [ Co w "; u 5 ::;; Combined burdens FIG. 2 The seasonal abundane of Rhipiephalus glabrosutatum on A. Vaal ribbok B. Bontebok, and C. The ombined burdens of the 2 hosts in the Bontebok National Park FIG. I The seasonal abundane of b:odes pilosus on A. Vaal ribbok B. Bontebok C. Srub hares, and. The ombined burdens of the 3 hosts in the Bontebok National Park during February 1984, but no lear pattern of seasonal abundane was evident. Oestrus ovis Seven of the 26 vaal ribbok examined for the larvae of. ovis were infested. Although these larvae developed to mature 3rd stage larvae in the vaal ribbok, all the 3rd stage larvae reovered were dead. Strobiloestrus sp. The seasonal abundane of the 3 larval stages is graphially presented in Fig. 5. First stage larvae were reovered from the subutaneous tissue of the vaal ribbok during eember 1983, 2nd stage larvae from the subutaneous tissue and hides from February-June 1983 and during eember 1983 and February Third stage larvae, of whih several 191

6 PARASITES OF OMESTIC AN WIL ANIMALS IN SOUTH AFRICA. XXI ,...,.,. A Vaal ribbok Larvae o---o Nymphae.tr----! Adults -O 5 4 o3 amalinia pelea Vaal ribbok Nymphae -<:l Adults -O too 5 B Bontebot FlO. 4 Tbe seasonal abundane of amalinia pelea on sub-adult and adult vaal ribbok in the Bontebok National Park 4 g3 2 ill ::;; to t 2 1 Strobiloestru:s sp. v aal ribbok 1st stage o----o 2nd stage l----a 3rd stage -O ::;; M Combined burdens FlO. 3 The seasonal abundane of Rhipiephalus nitens on A. Vaal ribbok B. Bontebok C. Srub hares, and. The ombined burdens of the 3 hosts in the Bontebok National Park FlO. S The seasonal abundane of the larvae of Strobiloestrus sp. in the subutaneous tissue and hides of vaal ribbok in the Bontebok National Park appeared to be nearly mature, were reovered from warble-like lesions in the hides during June No larvae were reovered during August and Otober The larvae were reovered mainly from the subutaneous tissue and hides of the upper half of the thorax and loin region. Fewer were present on the shoulders and rump. Larvae were not found in the viinity of the spine. Only 1 bontebok was infested, and that with a single 2nd stage larva. Climate The mean monthly minimum and maximum atmospheri temperatures and total monthly rainfall in the park from February 1983-February 1984 are graphially represented in Fig. 6. The lowest minimum temperatures were reorded during July and August 1983, and the highest maxima during February and Marh 1983 and from eember 1983-February Although the numbers of days on whih rain fell during the 1st and 2nd halves of 1983 were similar (38 as opposed to 39), the total rainfall for the 1st half of the year totalled only 169,9 mm ompared with 29,5 mm during the latter half. ISCUSSION Host preferene uring eember 1979, 5 vaal ribbok and 8 bontebok were examined for parasites in the Bontebok National 192

7 12 1 E' ao. _ :! 8 g j 6 i z 2 FIG. 6 Monthly mean minimum and maximum atmospheri temperaitllres and monthly rainfall in the Bontebok Nattonal Park from February 1983-February 1984 TABLE 7 The mean burdens of amalinia pelea on 3 groups of vaal ribbok Group Number Mean number of lie reovered of animals Nymphae Adults Total Juveniles Sik female Sub-adults and adults TABLE 8 The definitive arthropod parasites of vaal ribbok, bontebok and srub hares in the bontebok National Park Arthropod speies Host speies Vaalribbok Bontebok Srub hare Ixodid tiks Ixodes pilosus X Rhipiephalus glabrosutatum X Rhipiephalus nitens X X X Lie amalinia pelea Fleas Ctenoephalides felis damarensis Park (Horak, Brown, Boornker, e Vos & Van Zr.I, 1982; Horak, evos & e Klerk, 1982). The vaal nbbok slaughtered then harboured only 2 ixodid tik speies and 1 louse speies. No fly larvae were reovered, although lesions suggesting earlier Gedoelstia sp. and Strobiloestrus sp. infestation were present. The bontebok harboured 3 ixodid tik speies, a lie speies and Gedoelstia sp. larvae. The larger variety of arthrotx.>ds re>vered i te preset survey.is not indiatiye of e1ther,an mrease m mfestat1n or an Improvement m reovery X X I. G. HORAK, K. SHEPPEY, M. M. KNIGHT & C. L. BEliTlflN methods, whih were basially similar in both investigations, but is partly due to the larger number of animals proessed, and more partiularly to the examination of animals at regular intervals during period of 13 months. This demonstrates the neessity of regular examinations in order to obtain aurate host-parasite reords for a host speies in a partiular environment. It also eliminates the possibility of parasites that may have been aidentally aquired by a host speies bemg regarded as definitive parasites of that speies. Table 8 is a summary of the arthropods whih we onsider to be definitive parasites of vaal ribbok, bontebok and srub hares in the Bontebok National Park. Only those parasites whih have been speifially identified are inluded in the table. A great advantage of examining several host speies within a partiular environment at regular intervals is that, where ertain parasites utilize more than 1 of these hosts, the ontribution made by eah to the total parasite. population an be assessed. This has been demonstrated by Randolph (1975a, b) in her studies on Ixodes triangulieps on small mammals in Sussex, England and by MaLeod (197) and Minshull (1981), who studied the tiks of several large mammals in Zambia and Zimbabwe respetively. In addition, the effet that eah host speies, its sex or its habits may have on the abundane of a parasite an also be determined (Randolph, 1975b). Ixodid tiks Sine the redution in springook numbers;- vaal ribbok, bontebok and srub hares omprise more than 9 of the total population of larger mammals reorded in the park by Boornker et al. (1981). They are also the major hosts of the 3 most prevalent tik speies, namely I. pilosus, R. glabrosutatum and R. nitens. The 28 mie and rats examined harboured only 1 larva of eah of the former 2 tiks (Tables 4, 5). Consequently, the antelope and hares' ombined tik burdens will aount for the bulk of tiks parasiti on animals in the park, and the ratios of the parasiti life stages of the 3 major tik speies summarized in Table 6 will be reasonably aurate. However, it must be borne in mind that the various life stages probably feed for different lengths of time and that the 2 month mtervals between examinations ould have led to major peaks of ativity of any of the life stages being m1ssed. Both these fators ould have a marked effet on the life stage ratios. The ratios of parasiti larvae to nymphae to adults for the 3 tik speies were fairly similar. In all ases they indiated that a greater loss ourred between the nymphal and adult stages than between the larval and nymphal stages. Randolph (1975a), after areful examination of a number of host speies, onluded that in the ase of I. triangulieps there was better survival from nymphs to adults than from larvae to nymphs. We are inlined to agree with her findings and feel that, despite the thorough methods of tik reovery employed in the present survey, it is likely that more larvae than other stages were not olleted. espite this shortoming it is worth noting that at least parasiti larvae of the 3 speies under disussion are required to ensure the later attahment of I adult (Table 6). Amblyomma marmoreum The adults of this tik prefer tortoises, while the immature staes have been reovered from a variety of hosts (Theiler & Salisbury 1959; Norval, 1975). More than 35 of all vaal ribbok, bontebok and srub hares examined in the present survey were infested with the immature stages of this tik. By ombining the burdens of the 3 hosts and then alulating the mean burden for eah month of examination a fairly lear pattern of seasonal abundane emerges 193

8 PARASITES OF OMESTIC AN WIL ANIMALS IN SOUTH AFRICA. XXI despite the small numbers. The greatest number of larvae were reovered from February-June 1983 and during February 1984, and the greatest number of nymphae during Otober and eember 1983 (spring to early summer). 11ris pattern of seasonal abundane seems to indiate that the greatest numbers of adults would be present on tortoises from eember-marh (summer), provided that the life yle i!i ompleted in I year. No tortoises were examined, but Norval (1975) states that tortoises in the eastern Cape Provine harboured the greatest numbers of A. marmoreum larvae during April (autumn), nymphae during spring and adults during mid-summer. Boophilus spp. Cattle are the preferred hosts of both B. de olora/us and B. miroplus (Hoogstraal, 1956; Howell, Walker & Neville, 1978), a fat verified by the tik burden of the vaal ribbok whih was killed outside the park after it had been grazing in lose proximity to attle. This animal harboured tiks of both these speies in eater numbers than the ombined burdens of all the ammals examined from within the park. Horak, Potgieter, Walker, evos & Boomker (1983) and Horak, evos & Brown (1983) have found that B. deoloratus prefers the larer antelope speies while most animals below blue wlldebeest size (bontebok and vaal ribbok fall within the ategory) do not beome heavily infested. This ould be one of the reasons for the small Boophilus burdens of the park animals. HaemaphhaduHkr This tik has a wider distribution than that given by Theiler (1962), who stated that it had been found in the warmer, more humid areas of the northern and eastern Transvaal and in the Umfolozi and Pietermaritzburg regions of Natal. It had apparently been introdued into South Afria from East Afria at the tum of the entury (Theiler, 1962) and onsequently it may still be spreading. We did not reover it m large numbers and it IS thus diffiult to determine its preferred hosts. espite the small number of adults reovered in the present survey, they did exhibit a pattern of seasonal abundane, being present during February 1983 and from August 1983-February Rodents and hares have been reorded as hosts of the immature stages of this tik (Hoogstraal & El Kammah, 1972) and 1 of the srub hares in this survey was infested with 2 nymphae (Table 3). Haemaphysalh leahi Some of the srub hares, mie and rats were infested with immature tiks and one of the mie also arried an adult tik. Hoogstraal (1956) lists domesti dogs and the larger wild arnivores as hosts of adult tiks and field rodents as hosts of the immature stages. The soure of infestation in the park was probably araal, and domesti dogs and ats whih oasionally get into the park Hyalomma trunatum A single male tik was reovered from 1 of the bontebok, but no other antelqj?e were infested. Hares are fre-' quently infested by the Immature stages of Hyalomma spp. (Clifford, Flux & Hoostraal, 1976), and 3 of the srub hares examined were mfested with the larvae and nymj?hae of H. trunatum. These hares had probably aquired their infestation outside the park, wnere they had used pasture previously utilized by attle, as these animals and large antelope are the preferred hosts of the adult Hyalomma spp. ourring in South Afria (Howell etal., 1978). Ixodes pilosus Adult tiks were reovered from the heads of the antelope, partiularly around the fae, and from the under- 194 side of the body. The immature stages were present on the legs, bodies and heads of the antelope. The total numbers of male and female /. pilosus reovered from the 3 major host speies were 63 and 25 respetively (Tables 1-3), a ratio of 1:3,3. Norval (1974) olleted 29 males and 12 females (a ratio of 1:3,5) from bushbuk and duiker in the eastern Cape Provine. He found that slightly more than half the males were attahed to the host, while equal proportions of the remainder were in opula or attahed to females. We do not know how many of the males we olleted were attahed to the host but a large number were in opula. None, however, were attahed to the integument of females. On the evidene of the male to female ratio, Norval (1974) suggests that mating may our on the host or on the ground. Althouh we have listed the vaal ribbok as a definitive host of this tik, it ould be argued that the bontebok and srub hares also qualify, as they too harboured both adult and immature tiks. Furthermore, onsideration should be given to the fat that the habitat preferenes of the animals and of the tik ould determine the level of infestation on a partiular host speies. This tik is found in sourveld areas all along the oast from Port Shepstone to Cape Town (Howell et al., 1978), and sourveld isi usually assoiated with the ooler southern slopes within a region. In the park, the vaal ribbok are frequently enountered on the gravel slopes, while the bontebok and hares prefer the plain and other level areas. Thus, the habitat preferenes of the animals and not neessarily the host preferene of/. pilosus ould ditate their levels of infestation. The seasonal abundane, as revealed by the ombined mean burdens of the vaal ribbok, bontebok and srub liares, indiates that a single life yle is ompleted annually. Peak burdens of larvae durin June give rise to peak nymphal burdens in August whih in tum are responsible for the peak in adult numbers during Otober and eember. However, it is possible that a separate.winter ohort of adult tiks, as evidened by the inrease in adult burdens during June, ould be responsible for the slight rise in larval numbers during eember (Fig. 1). No orresponding rise in nymphal burdens is evident, however. RhipiCephalus e ertsi e ertsi This tik prefers attle and equids as hosts (Hoogstraal, 1956; Norval, 1981; Horak, 1982; Horak, evos & e Klerk, 1984). The progenitors of the small numbers reovered in the present survey had probably been brought into the park originally by hares whih had aquired their infestations on the surrounding farms, Of by the Cape buffalo and eland that had been introdued and subsequently removed from the park (Boomker et al., 1981). Rhipiephalus gertrudae Very few adult tiks were reovered and then only from the vaal ribbok and bontebok. Rhipiephalus glabrosutatum All stages of development of. this 2-host tik were reovered mainly from around the feet and on the lower legs of the antelope. The fat that more males than f males were reovered is in our experiene normal for many members of the genus Rhipiephalus in South Afria. It is possible that host habitat preferene influened the levels of infestation on the 2 antelope speies, but nevertheless we have not aorded the vaal ribbok the status of preferred host. The reason for this is that on the bontebok more than 113 of the larvae suessfully

9 I. G. HORAK, K. SHEPPEY, M. M. KNIGIIT & C. L. BEUTHIN moulted into nym.e_hae, while on the vaal ribbok only about 115 did so (Tables 1, 2). The vaal ribbok is manifestly not a good host of the immature stages. The seasonal prevalene indiates that one life yle a year an be ompleted, with the immature stages that ate present in autumn and winter giving rise to adults during spring and early summer (Fig. 2). This pattern of seasonal abundane is virtually idential with that desribed for this tik on kudu and domesti goats in Valley Bushveld in the eastern Cape Provine (Knight & Rehav, 1978; Maivor & Horak, 1984) and mountain zebra in the Karoo (Horak, Knight & e Vos, 1986). In both regions, this tik has to ope with hot and usually omparatively dry summers. Rhipiephalus nitens Adult tiks were reovered mainly from the heads of the antelope, and partiularly lfrom the outer ear and the lower edge of the mandible. On the hares they ourred on the ears. The immature stages were reovered mainly from around the feet and on the lower legs of the antelope. Fair numbers of nymphae were also present on the heads of the bontebok. The fat that more female than male tiks were reovered from the antelope, and that male tiks only slightly exeeded the number of female tiks on the hares is unusual for a tik of the genus Rhipiephalus in whih, in our experiene, males usually onsiderably outnumber females (Horak, 1982; Horak, Potgieter, Walker, e Vos & Boomker, 1983; Horak et al., 1984; Horak & Fourie, 1986; Horak et al., 1986). However, the 5 vaal ribbok and 8 bontebok examined in the park in eember 1979 harboured more male than female R. nit ens (864 and 62 respetively) (Horak, Brown, Boomker, evos & Van Zyl, 1982), while the 6 vaal ribbok and 2 bontebok examined during eember 1983 in the present survey also harboured more males than females (171 and 128 respetively). In ontrast, the 4 vaal ribbok and 2 bontebok examined during February 1983 and during February 1984 harbojred a total of 69 male:94 female and 136 male:191 fmale R. nitens respetively. Apparently, therefore, raales outnumber females on the host animals early in me season of adult ativity, while the onverse is truf', later in the season. If this is not just a fortuitous ob:o-ervation, additional Ion term studies planned for t.ile park might eluidate thts phenomenon. Three vaal ribbok and 3 bontebok that were examined during Feruary 1985 harboured total burdens of 75 male: Ill female R. nit ens. We t.ave listed both antelope speies and the srub hare a, preferred hosts on the basis of the number of imma',ure and adult tiks reovered on eah as well as the fat hat female tiks matured on all of them. We are unaware of other Rhipiephalus speies in South Afria tha; utilize both ruminants and lagomorphs as preferred hosts for all stages of development. R. nitens has a gwgraphial distribution limited to the south-western Cape Provine (Morel, 1969), and by makin use of a wid range of hosts within its onfined distribution its hanes of survival are enhaned. Jud ing from the number of tiks reovered, the bontebok ts apparently a better host of R. nitens than the vaal ribbok. However, an important fator in the determination of ltost suitability is the number of female tiks that are able to engorge and mature on the host. Thus, although the bontebok may be the preferred host of R. nitens, the vaal ribbok appears to be the effetive host, as it allows more females to mature. The latter fat may be entirely due to the physial attributes of the 2 antelope speies. The bontebok has short, fairly oarse hair whih would probably failitate the dislodging of engorging female tiks during grooming. In ontrast, the vaal ribbok has fairly long, fine, dense hair. The seasonal ourrene and morphology of R. nitens is similar to that of Rhipiephalus appendiulatus. Howell.et al. (1978) makmg use of Theiler' s (1962) distribution data, show R. appendiulatus as ourring in the south-western Cape Provine and thus overlapping the distribution of R. nitens. In our experiene, R. appendiulatus does not our in this region, and Its southern distribution probably does not extend west muh past 25 East. Rhipiephalus sp. Adult tiks of this speies were reovered mainly from around the ft and on the lower legs of vaal ribbok and bontebok and were present from April-August Lie The lie on the bontebok and srub hares have been identified only to generi level. Aording to Ledger ( 198), there ts still doubt as to the identity of the amalinia and Linognathus speies infesting bontebok, and he has listed no lie from the srub hare. In both antelope speies, the juveniles generally harboured more lie than the older animals. Chalmers & Charleston (198) reorded the same phenomenon for amalinia bovis and Linognathus vituli on attle in New Zealand, and Horak, e Vos & Brown (1983) made similar observations on young blue wildebeest alves and those a year older. Only amalinia pelea on the older vaal ribbok exhibited a lear pattern of seasonal abundane being more numerous from April-August with a peak in June. Horak, e Vos & Brown (1983) found that amalinia theileri and Linognathus speies peaked on blue wildebeest in the Kruger National Park during autumn and during spring. This preferene for the ooler months of the year onfirms Murray & Gordon's ( 1969) observations for amalinia ovis on sheep in Australia and those of Chalmers & Charleston (198) for attle lie in New Zealand. Oestrid fly larvae Gedoelstia sp. Flies of this genus normally deposit their larvae on the eyes of their hosts, whene they migrate by various routes to the nasal_passages and sinuses (Basson, 1966; Horak & Butt, 1977). Aidental infestations of domesti stok with larvae of this genus may result in severe oular lesions (Bas son, 1962). The lst stage larvae reovered from the vaal ribbok must be regarded as aidental infestations of an abnormal host and, although oular lesions were present in several of these animals, larvae were reovered from the orneas of only 3. It is the experiene of one of us (I.G.H.) that Gedoelstia sp. larvae are seldom reovered from severely affeted eyes. Some of the young bontebok exhibited oular lesions but none of the older ones did. The reovery of 1st stage Gedoelstia sr. larvae from the eyes, the heart hambers and the nasa passages of the bontebok, while none were found on the dura mater, indiates that the infestation followed an oulo-vasular route to the nasal avity rather than an oulo-ranial route (Basson, 1966; Horak & Butt, 1977). The small number of larvae reovered from the eyes ompared with the fairly large number in the heart suggests that the larvae may spend some time in the heart. A similar phenomenon has been notied with Gedoelstia speies infesting blue wildebeest, where the I st stage larvae seemed to aumulate on the dura mater before migrating to the nasal avity (Horak, e Vos & Brown, 1983). The reovery of 1st stage larvae from the eyes ofbontebok during most months of the survey indiates that in 195

10 PARASITES OF OMESTIC AN WIL ANIMALS IN SOUTH AFRICA. XXI the Bontebok National Park the life yle ontinues throughout the year and thus no lear pattern of seasonal abundane an be determined. It is our experiene, and that of the park warden, however, that most eye problems our 1 vaal ribbok durin eember eah year, an indiation of a peak of fly ativtty at this time. Oestrus ovis Bedford (1926) lists sheep and goats as hosts of. ovis, and he also mentions that he had reently reeived some larvae of this fly olleted from vaal ribbok. Zumpt (1965) states that only domesti sheep and goats and ertain of their lose wild relatives are hosts of this parasite. We agree with Zumpt: although one of us (I.G.H.) has examined hundreds of antelope of several speies for oestrid larvae, the vaal ribbok is the only animal he has found to be infested with. ovis. We feel that the vaal ribbok is only an aidental host of these larvae beause, although the larvae are able to mature they all seem to die before leaving the nasal passages. Thts is possibly due to the ranial anatomy of the vaal ribbok whih has very small frontal sinuses, not extend ing into the horns, and omparatively large maxillary and palatine sinuses. The 3rd stage larvae, whih normally mature in the frontal sinuses of sheep (Cobbett & Mithell, 1941), raw I into the maxillary and palatine sinuses of the vaal ribbok probably beause of the lak of. spae in the frontal sinuses. Here they mature, but being then unable to esape, subsequently die. All the dead larvae were reovered from these sinuses. The infestation in the vaal ribbok oriinates from sheep whih often graze in large numbers nght up to the perimeter of the park. Strobiloestrus sp. Aording to Zumet's (1965) host list, the larvae reovered from the vaal nbbok are likely to be Strobiloestrus larkii. Only 2 adult flies of this speies are known, both of whih were aught in the Cape Provine during the last entury. No speifi identifiation of the larvae reovered from the hides of the vaal ribbok ould be made, as we were unable to rear flies from the 3rd stage larvae we had olleted. The vaal ribbok are natural hosts of larvae of this fly, as not only were most of them infested, but the larvae were able to mature and leave their hides to pupate. We assume this proess took plae between June and August beause of the large number of nearly mature 3rd stae larvae reovered in June. The animals examined 1 August harboured no larvae but had lesions where the larvae had previously been loated. The bontebok is an aidental host of this parasite and beomes infested only beause of its lose assoiation with vaal ribbok in the park. Aidental infestations have been reorded in attle (Horak & Boomker, 1981) and Merino sheep (Brain, Van der Merwe & Horak, 1983) running with klipspringers and mountain reedbuk respetively, both of whih are hosts of Strobiloestrus larkii (Zumpt, 1965). The life yle and seasonal ourrene of Strobiloestrus sp. infesting the vaal ribbok in the Bontebok National Park is similar to that of Hypoderma lineatum infesting attle in the Northern Hemisphere. H. lineatum adults lay egs attahed to the hair of attle during the northern spnng and summer. The 1st stage larvae, whih hath from these eggs, migrate by various routes to the subutaneous tissue of the baks of the attle, whih they reah about 4 weeks after hathing. Here they moult to the 2nd and 3rd stages before leaving the skins during spring to pupate in the soil (Zumpt, 1965). Strobiloestrus sp. flies probably attah their eggs to the hair of the vaal ribbok during September and Otober 196 (spring) and I st stage larvae are found in the subutaneous tissue of the bak during eember. We were unable to determine whether these larvae had miated there from other sites or had penetrated the skin drretly above them, after hathing from the egg. These larvae moult to the 2nd stage during eember and by February all are in this stage of development. The 2nd stage larvae moult to the 3rd stage during May and June (autumn and winter), and the latter larvae leave the skin to pupate during June and July (winter), so that the animals are free of infestation in August. One of us (I.G.H.) has found that in regions with relatively old winters the 1st oestrid flies to hath after winter (. ovis, Oestrus variolosus and G. hassleri) do so towards the end of September and at the beginning of Otober (Horak, 1977; Horak & Butt, 1977). We suggest that the same ours with Strobiloestrus sp. in the park. Only I life yle a year is possible. ACKNOWLEGEMENTS We wish to thank the National Parks Board for plaing the animals and failities in the Bontebok National Park at our disposal. The researh was funded by the Meat Board, Rhodes University, the Counil for Sientifi and Industrial Researh and the National Parks Board. We gratefully aknowledge the assistane of the Park Warden, Mr E. Fourie and his staff and Mr P. Beukes of the University of Stellenbosh with the proessing of the neropsies, also the tehnial assistane of Mrs Valerie A. HorakandMrE. J. Williams. r Jane B. Walker assisted with the identifiation of the tiks, partiularly R. gertrudae and R. nitens, and drew our attention to the work of Morel on the distribution of the latter tik. She also very kindly supplied laboratory-reared speimens of the immature stages of H. trunatum and/. pilosus. Mr I. T. P. Pajor assisted with the identifiation of the fleas. 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