Onderstepoort 1. vet. Res., 58,81-105 (1991)

A REVIEW OF THE IXODID (, ) OCCURRING IN SOUTHERN AFRICA JANE B. WALKER, Veterinary Research Institute, Onderstepoort 0110

ABSTRACT WALKER, JANE B., 1991. A review of the ixodid ticks (Acari, Ixodidae) occurring in southern Africa. Onderstepoort Journal of Veterinary Research, 58,81-105 (1991). Eighty-three of ixodid ticks, as well as several entities that have yet to be described, occur in the Republic of plus the 4 independent states lying within its borders (Bophuthatswana, Venda, Transkei and Ciskei) and in , , and Swaziland. They belong to the following genera: Amblyomma (8 spp.); Aponomma (3 spp.); Boophilus (2 spp.); Cosmiomma (1 sp.); Dermacentor (1 sp.); (10 sp_p.); Hyalomma (2 spp., one of them with 2 subspp.); Ixodes (25 spp. ); Margaropus (1 sp. ); Rhipicentor (2 spp. ), and (28 spp ). The history of research in this region is reviewed briefly and advances made by the major contributors to our knowledge are highlighted. Short comments on each are given. These are followed by information on every species known to occur in the region, presented under the following headings: Species diagnosis, under which references to relevant descriptiOns are listed and, where appropriate, notes on commonly used syno­ nyms, related species and identification problems are included; hosts, indicating whenever possible the preferences of both the adults and the immature stages, as well as differentiating between common and mcidental hosts, and distribution in terms of political (not ecological) divisions.

INTRODUCTION pests of crops and orchards, then in 1898 he Historical background began the detailed, painstaking observations on ticks and tickborne diseases for which he became famous. Descriptions of ticks that occur in South Africa In 1899 he wrote to Neumann and began sending began to appear in the literature over 200 years ago. him specimens. In 1901 Neumann published a In 1778 Baron C. de Geer described 2 species: "Aca­ description of Rhipicephalus appendiculatus, fol­ rus silvaticus", based on a <:? tick taken from a lowed in 1904 by descriptions of Rhipicephalus nitens "Schildkrote" (i.e. a tortoise) at the Cape of Good and Ixodes rubicundus, based on some of Louns­ Hope by Sparrmann, and "Acarus rhinocerotis" bury's collections. Besides these 3 species Neumann from a rhinoceros, also at the Cape of Good Hope. described another 16 that occur in South Africa from Over the years there has been much confusion and specimens obtained from various sources in other controversy about the entities to which his names parts of the continent. apply. The current view is that they are the valid names for Amblyomma sylvaticum and Amblyomma Lounsbury also sent many ticks, including live rhinocerotis respectively (Theiler, 1943a; Hoog­ specimens for experimental purposes, to G . H. F. straal, 1956). Nuttall in England (Keirans, 1985). Nuttall and his Cambridge colleague, C. Warburton, with their During the following 66 years another 8 South collaborators W. F. Cooper and L. E. Robinson of African ticks were described. Most were common, the Cooper organization, made considerable contri­ well-known species, for example Rhipicephalus butions to the systematics of African ticks. Between sanguineus (Latreille, 1806) and Haemaphysalis them they described 9 species that occur in southern leachi (Audouin, 1827), but one, Cosmiomma Africa, among them Haemaphysalis silacea and the hippopotamensis (Denny, 1843), has rarely been genus Rhipicentor with its 2 species R. bicornis and found since it was first discovered. R. nuttalli. Many of their findings were included in In 1844 C. L. Koch published his historic work on books on the and on the genera Ixodes, ticks from different parts of the world in which he Haemaphysalis and Amblyomma, published under laid a large part of the foundation of modern tick the general title "Ticks-A monograph of the Ixo­ systematics, mcluding establishing 5 genera. Eleven doidea" (Nuttall, Warburton, Cooper & Robinson, ixodids that occur in southern Africa feature in this 1908-1926), that remain essential references to this work, amongst them such important species as Amb­ day. lyomma hebraeum, Boophilus decoloratus (as Rhipi­ Another renowned tick systematist during the cephalus decoloratus), Hyalomma marginatum ru­ early part of this century was W. Donitz of Berlin, to fipes (as H. rufipes), Hyalomma truncatum, Ixodes whom many tick collections from "Deutsch-Siid­ pilosus, Rhipicephalus capensis and Rhipicephalus west-afrika" (Namibia) and "Deutsch-Ostafrika" simus (Theiler, 1962). () were sent. He published a number of In terms of the number of new tick species he valuable papers on his findmgs, of which those on described Koch's contribution to our knowledge has the genus Amblyomma (Donitz, 1909) and on been exceeded only by that of the great French para­ various southern African species (Donitz, 1910) are sitologist L. G. Neumann, of the Veterinary School especially relevant. Subsequently the ticks of Nami­ at Toulouse, in southern France. His interest in bia also received attention from Trommsdorff (1914) South African ticks was stimulated through his close and Hans Sigwart (1915). and cordial association with C. P. Lounsbury, who The first overall review of southern African ticks emigrated from the United States of America to take was published in 1908 by C. W. Howard who, like up an appointment as Government Entomologist to Lounsbury, had emigrated from the United States. the Department of Agriculture, Cape of Good It was a particularly remarkable achievement be­ Hope, in 1895. Initially Lounsbury worked on the cause he was appointed as Assistant Entomologist, Transvaal Department of Agriculture, only in 1905 and by 1908 had moved to Portuguese East Africa Received 31 January 1991-Editor (). 81 A REVIEW OF THE IXODID TICKS OCCURRING IN SOUTHERN AFRICA In February 1912 G. A. H. Bedford arrived from available. England to take up a post as Entomologist at Onder­ The transmission of human and pathogens stepoort, where he worked for 26 years. Referred to by the ixodid tick species reviewed here is men­ later by Theiler (1975) as "a taxonomist par excel­ tioned under the various genera, but only briefly. lence ", he studied virtually all the parasitic arthro­ References on this subject have been kept to a mini­ pods occurring in South Africa. These included mum. In particular those cited by Neitz (1956) in his ticks, of which he described 3 new species (Haema­ comprehensive review of this subject have been physalis cooleyi, Ixodes elongatus and Rhipicephalus omitted. theileri). In 1932 he published a valuable checklist and host list of the ectoparasites found on South GENUS AMBLYOMMA KOCH , 1844 African reptiles, birds and mammals, to which he Generic characteristics: later added a supplement (Bedford, 1932, 1936). He also began what was obviously intended to be a (a) Ornate, i.e. a coloured pattern is usually present series of papers on South African ticks, of which on the scutum only the first part was published (Bedford, 1934). (b) Legs banded Following Bedford's early death in 1938 R . du (c) Hypostome and palps long Toit assumed responsibility for work on ticks, and in (d) Eyes present 1941 described Rhipicephalus glabroscutatum. His involvement with these parasites was relatively brief (e) Festoons present because Gertrud Theiler was appointed at Onderste­ (f) Adanal plates in the male absent poort in 1940 and immediately took over the basic This genus, the third largest in the family Ixodi­ tick research. She undertook this task with enthu­ dae, is distributed world-wide. The majority of these siasm and during the following quarter of a century bont ticks, which are mostly large, colourful species, carried out numerous fundamental studies on the feature in a monograph by Robinson (1926). This systematics of African ticks, assisted for short work remains a basic reference even today for most periods by Britha Robinson and Lois Salisbury. She African members of the genus. maintained close and cordial links with other tick workers worldwide and became the doyenne of Eight species of Amblyomma occur in southern those in Africa. Her studies culminated in her review Africa. These fall into 2 groups: 4 species whose of all the known tick species occurring in the Afro­ known hosts are primarily mammals (hebraeum, rhi­ tropical region (Theiler, 1962). Finally in 1975 she nocerinus, tholloni and variegatum) and 4 that are published an interesting synopsis of tick research in basically, but not always exclusively, parasites of Africa which amplifies this brief account. Further reptiles (marmoreum, nuttalli, sparsum and sylvati­ details regarding the contributions made by many cum). The latter group was reviewed in detail in a other people to our current knowledge of southern paper by Theiler & Salisbury (1959) that supersedes African ticks, for example H . Hoogstraal, D. R. the earlier findings on some of these ticks by Robin­ Arthur, F. Zumpt and J. A. T. Santos Dias, will son (1926). also be found later in the present paper under the Economically A. hebraeum and A. variegatum are accounts of individual genera and spectes. 2 of the most important tick parasites of livestock in Scope of this review Africa, mainly because they are the principal vectors of Cowdria ruminantium, the causative agent of All the genera and species of ixodid ticks presently heartwater in , and goats. The ecology of recognized in southern Africa are included. Short these ticks, and of other Amblyomma spp. that can comments on each genus are followed by informa­ transmit C. ruminantium, was recently reviewed by tion on the individual species presented under the Petney, Horak & Rechav (1987). In East Africa subheadings species diagnosis, hosts, and distribu­ A. variegatum is also a vector of Nairobi sheep dis­ tion. ease virus. Both A . hebraeum and A. variegatum The sections on species diagnosis include refer­ can transmit Rickettsia conori, which causes human ences to descriptions plus notes on commonly used tick-bite fever (tick-borne typhus). synonyms, related species and taxonomic problems. Adult bont ticks, and sometimes their nymphae, In these sections the term "undescribed" indicates inflict severe bites on . These bites result in that the stage(s) referred to have been identified but discomfort and extensive tissue damage, especially not as yet formally described, whereas "unknown" to the udders and teats of cows and to hides and indicates that they have not even been recognized. skins in general. Often such bites act as routes of The sections on hosts include, whenever possible, infection for various pathogenic organisms (Yeoman information on the preferences of both the adults & Walker, 1967). For example, the bites of numer­ and the immature stages. Their preferred and inci­ ous A. hebraeum adults and nymphae round the dental hosts are also differentiated. Host nomencla­ hooves of goats caused skin damage and abscessa­ ture is according to that given in the following publi­ tion resulting in lameness (Maclvor & Horak, 1984, cations: reptiles-Broadley (1983) , Patterson & 1987). Bannister (1987), Boycott & Bourquin (1988) ; birds-Maclean (1985) , and mammals-Meester, Amblyomma hebraeum Koch, 1844 Rautenbach, Dippenaar & Baker (1986). Species diagnosis: Robinson (1926), Arthur The information on distribution is given primarily (1973), Walker & Olwage (1987). according to political, not ecological, divisions. As Hosts: Cattle are regarded as the primary do­ used here the term "southern Africa" implies the mestic hosts of the adults. They also feed readily on Republic of South Africa and the 4 independent sheep, goats, horses and other equines, and some­ states lying within its borders (Bophuthatswana, times on pigs and dogs. In addition adults have been Venda, Transkei and Ciskei), plus Namibia, collected from many species of wild mammals, Botswana and Swaziland. Some records from Leso­ especially the larger ungulates (Theiler, 1962; Pet­ tho are also included, though no formal tick survey ney eta!., 1987). Larvae and nymphae often feed on has ever been carried out there and few data are the same hosts as the adults. They also parasitize 82 JANE B. WALKER many smaller animals, among them various carni­ Amblyomma nuttalli Donitz, 1909 vores (Carnivora, Canidae, Viverridae and Felidae) Species diagnosis: Theiler & Salisbury (1959) , and hares, especially the scrub hare (Lepus saxatilis) Arthur (1975a, b). This tick was sometimes confused (Lagomorpha, Leporidae) (Horak, Maclvor, Petney with A. marmoreum by Robinson (1926). & De Vas, 1987b; Horak, Jacot Guillarmod, Mool­ man & De Vas, 1987c). Birds are important hosts of Hosts: All stages feed on reptiles. The most com­ monly recorded hosts are tortoises, including the ~he immature stages, espec~ally ground-feeding spec­ Ies such as the helmeted gumea fowl , Numida melea­ hinged tortoise, Kinixys belliana, and the leopard gris (Galliformes) (Theiler, 1962; Horak & Wil­ tortoise, Geochelone pardalis (Chelonia, Testudini­ liams, 1986). Sometimes the leopard tortoise, Geo­ dae). Leguaans (Varanus spp.) are also favoured, as chelone pardalis (Reptilia, Testudinae) harbours are some of the larger snakes such as the python quite large numbers of nymphae (Walker & Schulz (Pytho'! sebae ~ ,. puff ad?er (Bitis arietans) and ga­ 1984). ' boon viper (BztlS gabomca) (Squamata, Varanidae, Boidae and Viperidae). Theiler & Salisbury (1959) Distribution: In South Africa A. hebraeum occurs and Theiler (1962) also list various birds and mam­ in the Transvaal bushveld; much of Swaziland; most mals as hosts, often of the immature stages only, but of the thornveld and coastal areas of KwaZulu, their significance in the maintenance of this species Natal and the Transkei; in the coastal areas of the has yet to be established. eastern Cape Province as far as Humansdorp, and in the Mossel Bay area (Theiler, 1948). Since Theiler's Distribution: In South Africa A . nuttalli is com­ original survey the boot tick has apparently spread in monest in KwaZulu and Natal. It has also been re­ both Swaziland and Natal (Baker & Ducasse, 1967; corded in the Transvaal at Onderstepoort; in the Howell, Walker & Nevill, 1978; Jagger, Wedder­ north-eastern Cape Province at Kuruman, and in the burn & McCartan, 1987; Walker & Olwage, 1987). eastern Cape Province at Grahamstown. It is widely In Botswana A. hebraeum is widespread in North­ distributed extralimitally in the Afrotropical region. eastern and Kgatleng Districts and in the eastern Amblyomma rhinocerotis (De Geer, 1778) pa~ts of Central, Kweneng and Southern Districts. Species diagnosis: Robinson (1926, as Amb­ Pame (1982) commented that the boot tick was be­ lyomma petersi) , Hoogstraal (1956). Immature lieved locally to have spread during the previous stages unknown. d~cade. H.e had collected a single C) in Ghanzi Dis­ tnct but did not know whether the tick was actually Hosts: Adults feed primarily on the white and the black rhinoceros, Ceratotherium simum and Diceros ~sta~lished there. Extralimitally A . hebraeum occurs m and southern Mozambique. bicornis (Perissodactyla, Rhinocerotidae). It has only occasionally been collected from other animals Amblyomma marmoreum Koch, 1844 (Hoogstraal, 1956; Yeoman & Walker, 1967). Species diagnosis: Theiler & Salisbury (1959) ; Distribution: In South Africa this species has been Arthur (1975a, b); Walker & Olwage (1987) . This recorded in northern KwaZulu and Natal in Ndumu, species was confused with A . sparsum, and some­ Mkuze, Hluhluwe and Umfolozi Game Reserves times with A. nuttalli, by Robinson (1926). plus the Corridor area between the latter 2 reserves Hosts: All stages feed on reptiles, most commonly (Baker & Keep, 1970). Theiler (1962) also lists an tortoises (Chelonia, Testudimdae) but also some of old record from the eastern Cape Province in "Kaf­ the larger snakes, especially the puff adder, Bitis fraria". Extralirnitally it has been widely recorded in ariet~ns , varanids and other lizards (Squamata) East and Central Africa. (Theiler, 1962; Walker & Schulz, 1984). Unlike the Amblyomma sparsum Neumann, 1899 adults, which are specific parasites of reptiles (Hoog­ straal. & Aeschlimann, 1982), the immature stages, Species diagnosis: Theiler & Salisbury (1959), especially the larvae, also feed on a wide range of Walker & Olwage (1987). Prior to the definitive other animals. They have been collected from cattle, study of the Amblyomma marmoreum group by sheep, goats and dogs; various carnivores (Canidae Theiler & Salisbury this tick was frequently misiden­ Viverridae and Felidae), especially black-backed tified as A. marmoreum. jackals, Canis mesomelas, and caracals, Felis cara­ Hosts: Adults of A. sparsum favour 2 distinct cal; a few ungulates (Artiodactyla, Bovidae), the groups of animals as hosts-some of the larger scrub hare, Lepus saxatilis, and Smith's red hare, species of wild mammals on the one hand and Pro no fagus rupestris (Lagomorpha, Leporidae) various reptiles on the other. Their commonest (Norval, 1975b; Horak & Knight, 1986; Horak, Pot­ mammalian hosts are the black rhinoceros, Diceros gieter, Walker, De Vas & Boomker, 1983b; Horak bicornis (Perissodactyla, Rhinocerotidae) and the eta/., 1987 b, c). Various ground-feeding birds have buffalo, Syncerus caffer (Artiodactyla, Bovidae). also bee~ recorded as hosts, particularly the hel­ They have also been collected from various other meted gumeafowl, Numida meleagris (Galliformes) wild mammals, mostly the larger species, but such (Norval, 1975b; Horak & Williams, 1986). records are comparatively rare. Amongst reptiles, Distribution: In South Africa A . marmoreum is tortoises are most commonly parasitized, including widely distributed (Theiler & Salisbury, 1959; specifically the hi.nged tortoise, Kinixys belliana, and Walker & Olwage, 1987). In Namibia, though, there the leopard tortoise, Geochelone pardalis (Chelonia, are as yet scattered records only, from Outjo, Oka­ Testudmidae). Water and rock leguaans (Varanus handja, Windhoek and Gobabis, and from Karas­ niloticus and Varanus exanthematicus), agamid burg District (Theiler & Salisbury, 1959; Heloise lizards (Agama spp.), pythons (Python sebae) and Heyne, unpublished data). In Botswana it has been puff adders (Bitis arietans) (Squamata, Varanidae, collected on Chief's Island in the Okavango; 100 km Agamidae, Boidae and Viperidae) may also be in­ east of Maun on the Francistown road, and at Tshe­ fested (Theiler, 1962; Yeoman & Walker, 1967; sebe and S~rowe (Theiler & Salisbury, 1959; Walker, 1974; Petney eta/., 1987). Walker, Mehhtz & Jones, 1978; Paine, 1982). Extra­ Distribution: Thus far A . sparsum has been found limitally it occurs in Zimbabwe and southern Mo­ in southern Africa only in northern Namibia. OneS? zambique. was recorded in Outjo District (Heloise Heyne, un- 83 A REVIEW OF THE IXODID TICKS OCCURRING IN SOUTHERN AFRICA published data) and ad was collected in 1933 by the throughout the eastern Caprivi Strip (Theiler, 1962). Government Veterinary Officer, Grootfontein In the neighbouring parts of northern Botswana it (Theiler & Salisbury, 1959). Extralimitally it is has been found in Ngamiland in the Kwando R iver widely distributed in the Afrotropical region, area, also in Chobe District at Kavimba, Kazungulu especially in eastern and central Africa. (Kazungula), Leshomo (Lesomo) and Pandama­ tenga (Paine, 1982). Extralimitally it is the most Amblyomma sylvaticum (De Geer, 1778) widely distributed of the Afrotropical Amblyomma Species diagnosis: Theiler (1943a). species. It has also extended its range considerably Hosts: Reptiles. All stages parasitize tortoises, outside Africa, eastwards to the Yemen Arab especially the angulate tortoise, Chersina angulata, Republic, Madagascar and various islands in the also the common padloper, Homopus areolatus, and Indian Ocean, and westwards to the Cape Verde the tent tortoise, Psammobates tentorius (Chelonia, islands and islands in the eastern Caribbean (Walker Testudinidae). The immature stages only have been & Olwage, 1987). collected from the spiny agama, Agama hispida, and Knox's desert lizard, Meroles knoxi (Squamata, GENUS APONOMMA NEUMANN, 1899 Agamidae, Lacertidae), and adults plus nymphae Generic characteristics: from the mole snake, Pseudaspis cana (Squamata, Colubridae). (al Hypostome and palps long (b Eyes absent Distribution: This species is known only from ( c Festoons present South Africa, Cape Province, where it has been ( d Adanal plates in the male absent recorded in the coastal areas from Port Elizabeth This genus of small, eyeless ticks, almost all of westwards to the Cape peninsula and up the west which are parasites of snakes and/or varanid lizards Cape coast as far as Hondeklip Bay. It may well (leguaans or monitors) (Hoogstraal & Aeschlimann, occur throughout the range of the angulate tortoise 1982), was revised by Kaufman (1972). Sub­ (Boycott & Bourquin, 1988). Warburton (1927) also lists 3 collections of A. sylvaticum (syn. Amblyomma sequently Santos Dias (1985) has disagreed with latum Koch, 1884) in the Vienna Museum from Kaufman's findings regarding the synonymy of some "Natal" and " Cape Zelabor". As Theiler (1962) species. pointed out, these collections should be restudied; Three Aponomma species occur in southern this tick has not been recorded in Natal in recent Africa. One (exornatum) is ornate while the other 2 times and the whereabouts of Cape Zelabor is un­ (tatum and transversale) are inornate. known. Aponomma exornatum (Koch, 1844) Amblyomma tholloni Neumann, 1899 Species diagnosis: Theiler (1945a), Kaufman Species diagnosis: Robinson (1926), Hoogstraal (1972). (1956), Van der Borght-Elbl (1977), Walker & Hosts: Primarily the water and rock leguaans Olwage (1987). Varanus niloticus and V. exanthematicus (Squamata, Hosts: This species occurs erimarily on the Varanidae). The adults often attach in the leguaan's elephant, Loxodonta africana (Proboscidea, El­ nasal passages and sometimes even suffocate captive ephantidae), from which adults, and sometimes specimens (Young, 1965). Theiler (1962) also lists nymphae, have often been collected. If cattle, sheep this tick from a wide range of other animals, includ­ and goats are kept in areas inhabited by elephants ing different reptiles, a bird, and various mammals, they too may be parasitized by the immature stages but these are thought to be incidental hosts only. of this tick (MacKenzie & Norval, 1980). Various Distribution: In South Africa A. exornatum is very reptiles, birds and other wild mammals have also widely distributed. In Namibia it is recorded in been listed as hosts (Theiler, 1962; Petney et al. , Etosha Game Reserve and at various places in 1987) but they are probably of secondary importance Grootfontein, Omaruru and Windhoek Districts, in its maintenance. and in Botswana in Ngamiland and at Kanye. It is Distribution: A . tholloni is restricted to areas in probably present throughout the range of its varanid which elephants occur. In South Africa it has been hosts. Apart from the Namib Desert and the south­ recorded m the Transvaal, in Sibasa and the Kruger western Cape Province V. exanthematicus occurs National Park, and KwaZulu, in Ndumu Game Re­ throughout southern Africa, while V. niloticus is serve and Tongaland. Extralimitally it is very widely found along the east flowing rivers, in the Okavango distributed in the Afrotropical region. Swamps, and in the lower reaches of the Kunene and Orange Rivers (Patterson & Bannister, 1987). Ex­ Amblyomma variegatum (Fabricius, 1794) tralimitally A. exornatum is widely distributed in Species diagnosis: Hoogstraal (1956), Van der East and parts of Central Africa. Borght-Elbl (1977), Walker & Olwage (1987). Hosts: All stages feed on cattle, which are major Aponomma latum (Koch, 1844) hosts of this species. Sheep, goats and other Species diagnosis: Theiler (1945b), Kaufman domestic animals are infested to a lesser extent (Pet­ (1972). ney et al. , 1987). On wild animals adults are most Hosts: Many species of snakes (Squamata, sub­ prevalent on medium-sized to large . order Ophidia), especially the larger ones (Theiler, Many species of the order Artiodactyla, in parti­ 1962; Kaufman, 1972). The occasional records that cular, have been recorded as hosts. Herbivores these authors list from various lizards, and even belonging to other orders are much less commonly from a few mammals, are thought to represent inci­ parasitized by this tick. The immature stages have dental infestations only. been found on a few reptiles, various species of birds Distribution: It is widely distributed in South and many different mammals (Theiler, 1962; Petney Africa, but in Namibia has as yet been recorded only eta/., 1987). at Okahandja and Gobabis, and in Botswana on Distribution: In Namibia A. variegatum occurs Xhaga Island (not located, but presumably in the

84 JANE B. WALKER Okavango swamps). Extralimitally it is very widely Hosts: The blue tick is primarily a parasite of the distributed in the Afrotropical region. larger domestic and wild ungulates (Artiodactyla, Bovidae, Perissodactyla and Equidae) (Theiler, Aponomma transversale (Lucas, 1845) 1962) . Cattle are its main domestic hosts, and very Species diagnosis: Theiler (1945b ), Kaufman heavy infestations may also develop on horses (1972). Larva unknown. (Theiler, 1911; Hoogstraal, 1956). Other domestic Hosts: Pythons (Sq_uamata, Boidae), of which animals appear to be much less important as hosts Python sebae, the Afncan python, is the only host (Hoogstraar, 1956; Baker & Ducasse, 1968). In the species recorded in southern Africa so far. Kruger National Park it occurs in large numbers on Distribution: In South Africa 4 collections only Burchell's zebra, Equus burchellii (Horak, De Vos have been recorded, of which 2 were from captive & De Klerk, 1984a), and was the most abundant and pythons in the Johannesburg Zoo and the Cape most prevalent tick recorded on blue wildebeest, Town Snake Park. It has also been collected in the Connochaetes taurinus (Horak, De Vos & Brown, eastern Transvaal lowveld, in the Manyeleti Game 1983c). Numerous blue ticks have also been found Reserve (Heloise Heyne, unpublished data), and in on giraffe, Giraffa camelopardalis; kudu, nyala and Natal at Pietermaritzburg. The true range of this tick bushbuck, Tragelaphus strepsiceros, T. angasii and probably coincides with that of the African python, T. scriptus respectively, and eland, Taurotragus oryx which in South Africa occurs in parts of the (Horak et al., 1983b), as well as impala, Aepyceros Transvaal, Swaziland, KwaZulu, Natal, and the melampus. The few species of birds, carnivores and north-eastern and eastern Cape Province, also in lagomorphs listed as hosts by Theiler (1962) are not northern Namibia and in parts of Botswana (Broad­ thought to be of any significance in the maintenance ley, 1983). The paucity of records of this tick pos­ ofthis tick. sibly reflects collectors' lack of opportunity, and per­ Distribution: In South Africa B . decoloratus is ~aps ~nthusiasm , to examine its apparent predilec­ widely distributed in the Transvaal, Swaziland, Kwa­ tion Site on these large, somewhat unmanageable Zulu and Natal, the northern and eastern Orange and irascible snakes, their eyelids. Extralimitally it is Free State, north-eastern and eastern Cape Pro­ widely, though discontinuously, recorded in the vince, in the southern coastal belt and in the winter Afrotropical region. rainfall areas of the western Cape Province (Theiler, 1949b; Howell et al., 1978; Jagger et al., 1987). In GENUS BOOPHILUS CURTICE, 1891 Namibia it has been recorded only in localized areas Generic characteristics: in the north. It is prevalent in much of eastern and south-eastern Botswana and also occurs in Ngami­ (a) Inornate land around the Okavango swamps and in north­ (b) Hypostome and palps short eastern Chobe District (Paine, 1982). Extralimitally (c) Eyes present, though they are sometimes diffi­ it is very widely distributed in the Afrotropical re­ cult to see gion. (d) Festoons absent Boophilus microplus (Canestrini, 1887) (e) Adanal plates in the male present Species diagnosis: Hoogstraal (1956) , Landt & Members of this small, but economically impor­ Arthur (1975), Heyne (1986). It was referred to as tant, ~enus are commonly known as blue ticks. They Boophilus (Uroboophilus) fallax by Theiler (1962), are Widely distributed in the Afrotropical and Orien­ but later she became convinced that this species is a tal regions, Australia and the New World. Two synonym of B. microplus (Hoogstraal, 1956; Ger­ species only have been recorded in southern Africa. trud Theiler, unpublished data). In a series of papers published during the 1930s Hosts: Cattle are the primary hosts of this tick. It W. Minning divided the genus into 3 subgenera, has only occasionally been collected from sheep, Boophilus sensu stricto, Uroboophilus and Palpo­ goats and horses (Theiler, 1943b; Hoogstraal, 1956; boophilus, and described several new species. His Mason & Norval, 1980). Records from wild animals classification was initially accepted, and some of the are rare. They include the lion, Panthera leo (Carni­ names he proposed appear m the South African vora, Felidae); grey rhebok, Pelea capreolus; sable literature, but it is not now regarded as valid antelope, Hippotragus niger, and buffalo, Syncerus (Hoogstraal, 1956). caffer (Artiodactyla, Bovidae) (Theiler, 1962, Ho­ The boophilids are one-host ticks. They are im­ rak, Sheppey, Knight & Beuthin, 1986b). The indi­ portant both as vectors of various pathogens and cations are, therefore, that its potential host range also because they have over the years developed re­ resembles that of B. decoloratus . sistance to a wide range of acaricides. The common Distribution: According to Howell et al. (1978) blue tick, Boophilus decoloratus, is the species that and Baker, Jordaan & Robertson (1979), in South is most frequently implicated in the transmission of 3 Africa B. microplus occurs in the Transvaal to the cattle parasites: Babesia bigemina, causing African north and east of Pretoria and in the Witbank, Bel­ redwater, also Anaplasma marginale and A. cen­ fast, White River, Barberton, Carolina, Ermelo, trale, causing gallsickness. The pantropical blue tick, Standerton and Wakkerstroom Districts, also in Boophilus microplus, is a vector not only of B. bige­ KaNgwane. In Natal it has been found in the north mina but also of Babesia bovis, causing Asiatic red­ in Ingwavuma, Newcastle, Hlabisa, lower Umfolozi water, and of A. marginale. In addition both these and Mtunzini Districts and at the southern end of the ticks can transmit Borrelia theileri, the cause of spi­ province in Camperdown, Richmond, Ixopo, rochaetosis in various domestic animals. Umzinto, Port Shepstone and Alfred Districts. It also occurs in the Umzimkulu, Bizana, Flagstaff, Ta­ Boophilus decoloratus (Koch, 1844) bankulu, Lusikisiki, Libode, Tsolo, Ngqeleni, Species diagnosis: Hoogstraal (1956), Arthur & Elliotdale, Willowvale and Kentani areas of the Landt (1973), Heyne (1986). It was referred to as Transkei. In the eastern Cape Province it is present Boophilus (Palpoboophilus) decoloratus by Theiler in Stutterheim and Victoria East Districts, in the (1949b). Ciskei at Keiskammahoek, and in isolated pockets 85 A REVIEW OF THE IXODID TICKS OCCURRING IN SOUTHERN AFRICA along the southern Cape coast in the districts of circumguttatus, an elephant parasite that occurs Humansdorp, Knysna, George, Mossel Bay, Heidel­ from West Africa eastwards across the continent to berg, Swellendam, Caledon and Robertson. It has western and Tanzania, and D . rhinocerinus been collected at several places in Swaziland (Jagger (see below). et al., 1987). As yet, through, B. microplus has not Morel (1969) placed the 2 African Dermacentor been recorded in either Namibia or Botswana. spp. in the genus Amblyocentor Schulze, 1932 but Extralimitally it has been found in parts of East and th1s classification is not universally accepted. Santos Central Africa. Dias (1963) regarded Amblyocentor merely as a sub­ GENUS COSMIOMMA SCHULZE, 1919 genus of Dermacentor. Generic characteristics: Dermacentor rhinocerinus (Denny, 1843) (a) Ornate Species diagnosis: Hoogstraal (1956), Arthur (1960), Clifford & Anastos (1964). The adults are (b) Hypostome and palps long large ticks with a striking black and deep gold scutal (c) Eyes present pattern. Larva undescribed. (d) Festoons present Hosts: Adults feed primarily on both the white (e) Adanal plates in the male present and the black rhinoceros, Ceratotherium simum and Diceros bicornis (Perissodactyla, Rhinocerotidae). An extremely rare, monospecific Afrotropical Hoogstraal (1956) also listed it from various genus. domestic animals, a jackal and eland (Taurotragus Cosmiomma hippopotamensis (Denny, 1843) oryx), to which Theiler (1962) added the monitor Species diagnosis: Donitz (1910, as Hyalomma lizard (Varanus sp.), elep!Iant (Loxodonta africana) , hippopotamense), Arthur (1960). Originally Denny buffalo (Syncerus caffer) and roan ant~?lope (1843) described the d as Ixodes bimaculatus and the (Hippotragus equinus) , but none of these ammals are thought to be significant hosts of this species. ~ as Ixodes hippopotamensis. It has also featured in the literature as an Amblyomma and a Dermacentor. Clifford & Anastos (1964) noted that adults and The adults are large ticks with a black pattern on nymphae were collected from the nests of their pale yellowish scutum &nd light-coloured and a macroscelid, an indication of the hosts of the mottling on the dorsal surfaces of their legs. Imma­ immature stages. ture stages undescribed. Distribution: In South Africa D. rhinocerinus has Hosts: The types were recorded from Hippopota­ been recorded in the Transvaal in the Kruger Natio­ mus amphibius (Artiodactyla, Hippopotamidae). nal Park. In northern Natal it occurs in Ndumu, Adults have since been found on the black rhinoce­ Hluhluwe and Umfolozi Game Reserves plus the ros, Diceros bicornis (Perissodactyla, Rhinoceroti­ Corridor area between the latter 2 reserves (Baker dae), which is now regarded as its most likely host & Keep, 1970), also at a few points in adjacent (Bezuidenhout & Schneider, 1972). [In a report areas. In Namibia 1 d was collected from a black accompanying 6 adults sent to Onderstepoort for rhinoceros at Grootfontein in 1933 (OP 2683 iii) but identification in 1960 it was stated that these ticks this species has apparently not been found there had been collected from cattle, although they since. It has been widely recorded extralimitally in preferred small stock, especially goats. Subsequent eastern, central, and other parts of southern Africa. enquiries, though, revealed that this information was incorrect (State Veterinarian, Ohopoho, 1960, GENUS HAEMA PHYSALIS KOCH , 1844 personal communication; Theiler, 1962; Bezuiden­ Generic characteristics: hout & Schneider, 1972)]. (a) Inornate Distribution: The type locality for this species is (b) Hypostome and palps short recorded merely as "South Africa". Adults have since been collected in southern Africa only in (c) Eyes absent Namibia, Kaokoland, in 1959 at Ohopoho and Otji­ (d) Festoons present janjasemo, and in 1971 at Ondjarrakagha, Otjibo­ (e) Adanal plates in the male absent ronbonga, Otjipembi and Ekoto (Bezuidenhout & Much of our knowledge of this genus, the second Schneider, 1972). C. hippopotamensis has also been largest in the family Ixodidae, is based on research recorded extralimitally in . A morphologi­ carried out from 1955-85 by H . Hoogstraal and his cally slightly different population occurs in . colleagues. Their findings on the Afrotropical spe­ cies were published in a series of papers under the GENUS DERMACENTOR KOCH, 1844 general heading "Notes on African Haemaphysalis Generic characteristics: ticks". In these papers individual species were de­ scribed, or redescribed, and Hoogstraal progress­ (a) Ornate ively developed his ideas regarding the subgenera, (b) Hypostome and palps relatively short and species groupings and relationships of the African broad haemaphysalids. His final conclusions were incorpo­ (c) Eyes present rated in a major study on tick and mammal coevolu­ tion with particular reference to Haemaphysalis (d) Festoons present species throughout the world (Hoogstraal & Kim, (e) Adanal plates in the male absent 1985). This relatively large genus is widely distributed in Ten species of these small, light brown, eyeless parts of North and Central America, Eurasia and ticks are currently known to occur in southern Africa. Several species are extremely important hu­ Africa. These fall into 4 of the 14 subgenera dis­ man and animal parasites and some are vectors of cussed by Hoogstraal & Kim (1985), as follows: Or­ various pathogens. Neither of the 2 species present nithophysalis (hoodi); Haemaphysalis (silacea); Kai­ in the Afrotropical region, though, is known to be of seriana (aciculifer, parmata), and Rhipistoma (cool­ any economic importance. These are Dermacentor eyi, hyracophila, leachi, pedetes, spinulosa, zumpti). 86 JANE B. WALKER Thus far the only Haemaphysalis species in sou­ both non-passerine and passerine species. thern Africa known to be a vector of any pathogens Distribution: In South Africa it is recorded in the is H. leachi. It transmits Babesia canis, causing ca­ Transvaal at Bloemhof, Tzaneen, Newington and in nine biliary fever, a disease that is frequently fatal. It the Kruger National Park; in Natal at Pietermaritz­ also transmits Rickettsia conori, which causes human burg, and in the eastern Cape Province at _East Lon­ tick-bite fever, and can harbour Coxiella burneti, the don; also in Botswana at Maun (Santos Dms, 1955). causative agent of Q fever in animals and man. It is very widely distributed extralimitally in the Haemaphysalis aciculifer Warburton, 1913 Afrotropical region. Species diagnosis: Hoogstraal & El Kammah Haemaphysalis houyiNuttall & Warburton, 1915 (1972). This tick is not now thought to occur in southern Hosts: Adults occur on cattle, sheep and goats. Africa. It is a specific parasite ~f Geoffroy'_s ground The commonest wild animal hosts are various large squirrel, Xerus erythropus, whtch occurs m s_outh­ and small antelopes, especially bushbuck, Tragela­ eastern Morocco and from across Afnca to phus scriptus, and common duiker, Sylvicapra grim­ western and Kenya (Corbet & Hill, .1?86). mia, and occasionally buffalo, Syncerus caffer Theiler (1962) listed H. houyi from a few locahhes m (Artiodactyla, Bovidae). They have also been col­ the Transvaal, Orange Free State and Cape Province lected, but less commonly, from various carnivores but commented: "The South African records need (Carnivora, Canidae, Viverridae and Felidae) and to be confirmed." Unfortunately the specimens on from the Cape hare, Lepus capensis (Lagomorpha, which these records were based apparently no longer Leporidae). The immature stages apparently feed exist. primarily on various rodents (Rodentia, ) and to a lesser extent on other small mammals, es­ Haemaphysalis hyracophila Hoogstraal, Walker & pecially the smaller carnivores and hares (Hoog­ Neitz, 1971 straal & El Kammah, 1972; Horak et al., 1986b; Species diagnosis: Hoogstraal et al. (1971). Lik~ Horak, Keep, Spickett & Boomker, 1989). H. cooleyi this tick belongs to the Ha_emaphY_salzs Distribution: In South Africa H. aciculifer has (Rhipistoma) orienta/is subgroup of dasste parasttes. been recorded from scattered localities in the nor­ Hosts: It is known only from the rock dassie, Pro­ thern and eastern Transvaal, including the Kruger cavia capensis, and one unidentified hyrax (Hyracoi­ National Park; at various places in Natal (Baker & dea, Procaviidae) (Hoogstraal et al., 1971; Norval, Keep, 1970; Horak et al., 1989), and in the western 1985). Cape (Horak et al., 1986b). Theiler (1962) considers Distribution: In South Africa it is recorded in the that this tick was introduced into South Africa on Cape Province, Clanwilliam District, in the Cedar­ cattle imported from East Africa after the Boer War berg; Montagu District, Kogman's Kloof, and Cra­ and the rinderpest pandemic. Although widely dis­ dock District, Mountain Zebra National Park; also tributed it is never encountered in large numbers. in the Transvaal, Letaba District, at Gravelotte. It Extralimitally it is widely distributed in East and Central Africa. occurs extralimitally in Zimbabwe. Haemaphysalis leachi Haemaphysalis cooleyi Bedford, 1929 (Audouin, 1827) Species diagnosis: Hoogstraal & Wassef (1981). Species diagnosis: Hoogstraal (1958); Matthysse & Larva unknown. It is one of the 4 African species Colbo (1987) . It is now accepted that "leachi" is the constituting the Haemaphysalis (Rhipistoma) orien­ correct spelling of this specific name. No subspecies ta/is subgroup that all parasitize dassies (hyraxes) of this tick are currently recognized (see also below (Hoogstraal, 1956; Hoogstraal, Walker & Neitz, under Haemaphysalis spinulosa). The information 1971). compiled under the name H. leachii leachii by Thetler (1962) undoubtedly includes several differ­ Hosts: The type specimens (3 cf'cf', 1 ~, 1 N) were ent entities. Hoogstraal (1964) commented: "The collected from a rock dassie, Procavia capensis leachii group is the most polymorphic of any in this (Hyracoidea, Procaviidae) (Hoogstraal & Wassef, genus and, to date, the more it has been studied, the 1981). Adults will also feed on the yellow-!potted more confused the taxonomic status of its various rock dassie, Heterohyrax brucei (Norval, 1985 ). The components has become". Subsequently Hoostraal record from Procavia habessinica in Theiler (1962), and his colleagues J. L. Camicas and K. El Kammah though, almost certainly refers to another species in clarified the situation somewhat by describing seve­ the orienta/is subgroup since this dassie occurs in the ral new species in this group, among them Haema­ Ethiopian highlands (Corbet & Hill, 1986) whereas physalis pedetes and Haemaphysalis zumpti (see H. cooleyi is apparently confined to southern Africa. below). Various taxonomic problems are, however, Distribution: Currently the only confirmed record still outstanding. Only recently Horak et al. (1987c) for H. cooleyi in South Africa is the type collection noted that they had been unable to differentiate the from the Transvaal, near Onderstepoort (OP 2454i). immature stages of the 2 commonest species in the Theiler (1962) also listed it from several other locali­ group in southern Africa, H. leachi and H. spinu­ ties in the Transvaal, Natal and eastern Cape, but of losa, which .emphasizes the need for further research these collections only the specimens from the east­ on their morphology. The following comment by ern Cape, New Bethesda (OP2974i), still exist. On Hoogstraal (1964) should be remembered by anyone re-examination these have proved to be Haemaphy­ trying to understand this difficult group of ticks: salis hyracophila (see below). H. cooleyi has been "The high aegree of host specificity displayed by recorded extralimitally in Zimbabwe. almost all species in the genus Haemaphysalis sug­ gests the advisability of commencing any study Ghana, Uganda probably do refer to this species sensu stricto because and the . In their original description of this their findings were based primarily on collections tick Hoogstraal & El Kammah (1972) also included from dogs. They distinguished this dog tick from a the following information: "Questionable record similar species referred to in their 1953a publication (South Africa). From Felis ( = Leptailurus) s. serval, as H. leachi var. indica , and in Theiler (1962) as H. Pietermaritzburg, 2,500 ft alt., Natal Province, date leachii muhsami, collected from small wild carni­ not stated but earlier than 1936, R. F. Lawrence: 3 cf' vores (see below under H. spinulosa). Theiler (1962) (HH 42,565) (Onderstepoort 2446ii). [This lot, commented that in southern Africa H. leachi is plen­ which also contains 1 ~ H. (K.) aciculifer (HH 42, tiful in areas with a mean annual "Tainfall of 635 mm 564), is either mixed and mislabelled or represents a ~25 inches) or more, but occurs less frequently, and remarkable discontinuity in H. (K.) rugosa distribu­ m smaller numbers, in areas with rainfalls of 508--{)35 tion. Unless it can be proven that H. (K.) rugosa mm (20--25 inches) . She said it is absent where the occurs in South Africa, we are inclined to disregard rainfall is less than 508 mm per.annum, i.e. in the this record]". I endorse this statement. north-western Cape, western _.and north-western Haemaphysalis silacea Robinson, 1912 Orange Free State, western Transvaal, the Kgala­ Species diagnosis: Hoogstraal (1963). gadi (Kalahari) and southern Namibia. She also found it to be absent in areas with over 120 days of Hosts: Sheep, goats, cattle and various antelopes frost annually. More recent studies have confirmed (Artiodactyla, Bovidae) act as hosts of all stages the presence of H. leachi in Namibia, both in the (Norval, 1975a; Knight & Rechav, 1978; Horak et north .and once in the e?'treme south at Karasburg al., 1983b). Kudu, Tragelaphus strepsiceros; bush­ (Helmse Heyne, unpubhshed data). In Botswana it buck, Tragelaphus scriptus, and eland, Taurotragus occurs primarily around the Okavango swamps, in ~ryx, are preferred and may be particularly heavily the eastern part of the country and at various points mfested. The common duiker, Sylvicapra grimmia, along its southern border (Walker et al., 1978; is also a good host. Immature stages only have been Paine, 1982). It is very widely distributed in the Af­ found on dogs; on some of the smaller wild carni­ rotropical region. vores, especially the caracal, Felis caracal (Carni­ vora, Canidae, Viverridae and Felidae), and on Haemaphysalis parmata Neum~, 1905 occasional rodents and hares. Various birds (Aves) Species diagnosis: Theiler (1945c); Matthysse & may also be infested by the immature stages, parti­ Colbo (1987). cularly the hadeda ibis, Hagedashia hagedash, and the helmeted guineafowl, Numida meleagris (Nor­ Hosts: The adults, which are extremely small and val, 1975a; Horak & Williams, 1986). easily overlooked, have been recorded from cattle and other domestic animals but are primarily para­ Distribution: H. silacea occurs only in South sites of antelopes (Hoogstraal, 1956; Horak, Keep, Africa, where it is found primarily in the eastern Flamand & Boomker, 1988a). The immature stages Cape Province in localized areas of Fish River Bush, often feed on the same animals as the adults. They a xerophytic scrub thicket vegetation that is charac­ have also been recorded from a wide range of other teristic of the hot dry ravines and river valleys in this hosts, mostly smaller mammals such as mongooses region. It is also established in northern Zululand, in and genets (Carnivora, Viverridae), various rodents, the Mkuze, Hluhluwe, Umfolozi and St Lucia areas hares, and insectivores, but also birds (Aves), (Baker & Keep, 1970; Horak et al., 1988a). Further especially ground-feeding species such as ­ south it has been collected in the Umgeni Valley fowls and francolins (Theiler, 1962). Nature Reserve; in the Bayne's Drift area north of Pietermaritzburg, and in the Vernon Crooks Nature Distribution: In South Africa&: parmata has been Reserve, Umzinto. recorded only in Natal, in the Charter's Creek Nature Reserve (Horak et. al., 1988a) and Durban. Haemaphysalis spinulosa Neumann, 1906 Theiler (1962) questioned whether it was a recent Species diagnosis: Hoogstraal (1964) , Hussein & introduction but its prevalence on red duikers Mustafa (1983). It was recorded by Theiler (1962) 'as (Cephalophus natalensis) in the'Charter's Creek area H. Leachii muhsami Santos Dias, 1954, with H. Lea­ s~gg~sts that this is ~ot so. Extr~li~itally it is widely chii var. indica sensu Theiler (1943b) given d1stnbuted from Sierra Leorie m West Africa provisionally as its synonym. In 1964 Hoogstraal re­ eastwards across the continent to Ethiopia and the described H. spinulosa, though he still had reserva­ East African countries. tions about its status. He commented: 'Study of both Haemaphysalis pedetes Hoogstraal, 1972 syntypes of H. spinulosa reveals that this is in fact a "small form" of H. Leachii that has been obscured by Species diagnosis: Hoogstraal (1972). Nymph inadequate description and incorrect illustration. undescribed; larva unknown. The question of whether H. spinulosa is a synonym, f!osts: Adults and nymphae are parasites of the a subspecies, or a full species closely related to H . spnnghare, Pedetes capensis (Rodentia, Pedetidae) Leachii must await further study'. Finally Camicas, (Hoogstraal, 1972; Norval, 1985). According to Hoogstraal & El Kammah (1972) recognized H. spi- 88 JANE B. WALKER nulosa as a full species, with H. muhsamae as its ground squirrel, Xerus inauris. This squirrel and P. junior synonym. As in the case of H. leachi the infor­ cepapi are both known hosts of the immature stages. mation given by Theiler (1962) about H. spinulosa Some of the records from these hosts listed under undoubtedly covers more than one entity (see es­ the name H. leai:hii muhsami by Theiler (1962) pro­ pecially H. zumpti). Horak et a/. (1987c) were un­ bably refer to this·species. able to differentiate the immature stages of H. leachi Distribution: It occurs in the eastern part of South and H. spinulosa. Africa, from scattered localities in the Transvaal, The entity originally described by Warburton Orange Free State and eastern Cape Province, and (1910) as H. leachi var. indica is now regarded a full in Botswana at Francistown and Chobe River species, H. indica, occurring in the Oriental and (Hoogstraal & El Kammah, 1974) . It also occurs Palaearctic regions (Hoogstraal & Kim, 1985), not in extralimitally in . Africa. Hosts: Adults of H. spinulosa have been found GENUS HYALOMMA KOCH, 1844 occasionally on domestic dogs and cats (Horak eta/., 1987c; J. B. Walker, unpublished data). They occur Generic characteristics: most frequently on wild carnivores, primarily on the (a) Scutum inornate smaller species such as mongooses and genets (Vi­ (b) Legs banded verridae) though they have also been collected from various members of the families Canidae and Feli­ (c) Hypostome and palps long dae (Horak eta/., 1987c). I have great reservations (d) Eyes present about some of the hosts listed for this species by (e) Festoons present, but sometimes not as well Theiler (1962), in particular those from various das­ defined as in other genera sies (Hyracoidea) since their parasites are usually (f) Adanal, accessory adanal and subanal plates in very host specific. the male present Distribution: In South Africa Theiler & Robinson Members of this genus, one of the smaller ones in (1953a) commented, with reference to their H. lea­ the family Ixodidae, occur in the Palaearctic, Orien­ chi var. indica, that ". . . the wild carnivores and tal and Afrotropical regions. Individual species often their tick variety, are present throughout South show a great range of morphological variation. The Africa extending even into the more arid areas . . . " failure of most workers to appreciate this, plus the This statement cannot at present be confirmed be­ fact that for a long time the species described in 1844 cause many of the ticks on which they based it are no by Koch (cited by Hoogstraal, 1956) largely re­ longer available and existing collections of H. spinu­ mained unrecognised, resulted in the nomenclatorial losa by no means cover the whole area. Specimens chaos that prevailed in the genus for many years. have been seen from the Transvaal, Kruger National Much of th1s confusion was resolved some 40 years Park; Natal, Vernon Crookes Nature Reserve, ago, primarily through painstaking research by S. Umzinto; Orange Free State, and eastern Cape Pro­ Adler and B. Feldman-Muhsam in Israel and by L. vince from various places (J. B. Walker, unpub­ Delpy in Iran. The genus was reviewed in detail by lished data; I. G. Horak, unpublished data). In Na­ Hoogstraal (1956). mibia a single cJ only has been recorded in Windhoek District. Extralimitally H. spinulosa is There are indications in various species in this very widely distributed in the Afrotropical region. genus that the type of life cycle followed is in­ fluenced by the prevailing environmental conditions Haemaphysalis tauffliebi Morel, 1965 and hosts (Hoogstraal, 1956; Balashov, 1972). Although this tick was recorded from South Under controlled laboratory conditions, at 26 ± Africa by Hoogstraal & Wassef (1973) it now seems 1 °C, when the immature stages were fed on rabbits virtually certain that it does not in fact occur here. and the adults on calves, Hyalomma marginatum These authors, who stated that its hosts are "various rufipes behaved as a 2-host species (Knight, Norval birds and small mammals", gave its range as "Ethio­ & Rechav, 1978) . pia to West Africa and South Africa". The only Three entities belonging to this genus are wide­ known record from South Africa, though, is of 3 spread in the drier parts of southern Africa. The ~ ~, 2 NN. from the plain backed pipit, adults' habit of actively seeking potential hosts de­ Anthus leucophrys (Aves, Passeriformes), East Lon­ tected in their vicinity, including humans, is well­ don, eastern Cape Province, October 1966, collected known to many field workers. Hyalomma spp. are by S. van der Merwe (HH 7301, RML 79689). Un­ regarded with particular caution by most people be­ fortunately the present whereabouts of this collec­ cause they are the primary vectors of the virus caus­ tion is unknown (J. E. Keirans, personal commu­ ing Crimean-Congo haemorrhagic fever in man nication, 1991). (Hoogstraal, 1979; Swanepoel, Shepherd, Leman, Haemaphysalis zumpti Hoogstraal & El Kammah, Shepherd, McGillivray, Erasmus, Searle & Gill, 1974 1987). H. marsinatum rufipes (syn. H. aegyptium var. impressum) has also been shown to harbour a Species diagnosis: Hoogstraal & El Kammah Rickettsia sp. identical with strains that produce tick (1974). A small tick in the H. leachi group, closely typhus in man. Recently H. truncatum has been related to H. pedetes (Hoogstraal & Kim, 1985) . Its found to be an efficient vector of Babesia caballi, the immature stages are difficult to distinguish morpho­ causative agent of equine babesiosis (De Waal, logically from those of H. leachi and H. spinulosa. 1990). Some strains of H. truncatum produce a der­ Hosts: Adults of H. zumpti have been collected motropic toxin that causes sweating sickness in from a wide range of small carnivores (Mustelidae, cattle, especially calves (Neitz, 1959). Two cases of Viverridae, Protelidae and Felidae) and the tree paralysis in man caused by this tick are on record squirrel, Paraxerus cepapi (Rodentia, Sciuridae). (Swanepoel, 1959). The bites of adult Hyalomma The most commonly recorded hosts are the suricate, spp. cause wounds that may be invaded either by Suricata suricatta, and the yellow mongoose, Cynic­ bacteria, resulting in the formation of abscesses, tis penicillata, which share the burrow systems of the sloughing of teats and tailtips, lameness and footrot, 89 A REVIEW OF THE IXODID TICKS OCCURRING IN SOUTHERN AFRICA or by larvae of the screwworm, Chrysomya bez­ Hyalomma marginatum turanicum Pomerantsev, ziana. In dogs large necrotic lesions, thought to be 1946 caused by a dermotropic toxin, sometimes develop Species diagnosis: Hoogstraal (1956, as H. turani­ around the bites of H. truncatum adults (Burr 1983). ' cum). Immature stages undescribed. It was listed by Theiler (1962) as H. glabrum. Further comments on Hyalomma marginatum ruflpes Koch, 1844 its nomenclature appear above under H.m. rufipes. . Species diagnosis: Hoogstraal (1956, as H. ru­ Hosts: This species apparently favours the same fzpes), Arthur (1975a, b). Doubt still exists regarding hosts as H. m. rufipes. The adults feed on "domestic the c<;mect name o~ this tick. Koch (1844) originally stock, mainly sheep" (Theiler, 1962). Horak & desc~Ibed H. margmatum and H. rufipes as separate Maclvor (1987) found numerous adults on the Cape species and they have been treated thus by many mountain zebra, Equus zebra zebra (Perissodactyla, people ever since, for example Theiler (1962) and Equidae) and eland (Taurotragus oryx), but few on Ke!rans (1985). In 1960, though, Hoogstraal & the springbok, Antidorcas marsupialis, and black Kaiser stated: "After exhaustive study of large wildebeest, Connochaetes gnou (Artiodactyla, Bovi­ numbers of H. marginatum, rufipes, and turanicum dae), present in the same area. They collected the from numerous areas in Europe, Africa, and Asia, it immature stages in large numbers from scrub hares, has become apparent that each of these forms is a Lepus saxatilis, but larvae only, in smaller numbers, geographical but closely related subspecies . . . The from Smith's red hares, Pronolagus rupestris (Lago­ study of this subject is nearing completion and will morpha, Leporidae). Both larvae and nymphae be presented shortly". Unfortunately, though, their were found on spring hares, Pedetes capensis re~ults J?.ever were presented in greater detail. De­ (Rodentia, Pedetidae) and various birds (Aves). spite this, many researchers, following the lead of Distribution: This subspecies is thought to have J:Ioogstraal himself, subsequently referred to this been introduced from the Near East (Theiler, 1962). tick as H. marginatum rufipes, a name that is now Its distribution in South Africa is restricted to "the well established in the more recent South African Karoo areas of the Eastern Cape, Western and literature. To revert immediately to the name H. South-western Cape, and the Cape Midlands as also rufipes would seem to I. G. Horak (personal com­ in the Brokenveld of the Southern Orange Free m~nic~~ion, 1990) an? ~he writer to negate, perhaps State" (Theiler, 1956). It has not been recorded else­ ~OJ~Stifiably ~ the validity of Hoogstraal & Kaiser's where in the Afrotropical region. fmdmgs. This could cause even greater confusion later. We suggest, therefore, that someone should Hyalomma truncatum Koch, 1844 look at this problem again and, after due consider­ Species diagnosis: Hoogstraal (1956). Immature ation, resolve it formally. stages undescribed. Theiler (1962) lists many syno­ ~heiler (1962) lists many other synonyms under nyms for this tick, including H. transiens, a name which this tick has featured in the literature from under which it often appears in the older South Afri­ time to time. can literature. Hosts: The primary domestic hosts of the adults of Hosts: The adults feed on cattle and other do­ H. m. rufipes are cattle; other domestic animals are mestic animals, including dogs, but usually not in rarely infested. They also parasitize the larger wild great numbers (Yeoman & Walker, 1967; Horak & ungulates, such as zebra, Equus spp. (Perissodac­ Maclvor, 1987). They have been recorded from tyla, Equidae); giraffe, Giraffa camelopardalis; buf­ many wild animals, among them various large and falo, Syncerus caffer, and eland, Taurotragus oryx small carnivores (Carnivora, Canidae and Felidae) (Artiodactyla, Giraffidae and Bovidae) (Theiler, and a wide range of ungulates (Perissodactyla and 1962). The immature stages feed very commonly on Artiodactyla). Although, like H. m. rufipes adults, birds (Aves), of which many species have been they apparently prefer the larger ungulates they recorded as hosts (Hoogstraal, 1956; Hoogstraal, have afso been found on some of the medium-sized Kaiser, Traylor, Gaber & Guindy, 1961; Theiler, and smaller species (Theiler, 1962; Norval, 1982). 1962). Both the Cape hare, Lepus capensis, and the The immature stages have a strong predilection for scrub hare, Lepus saxatilis (Lagomorpha, Lepori­ the Cape hare, Lepus capensis, and the scrub hare, dae) are also favoured by the immature stages Lepus saxatilis, and to a lesser extent for Smith's red (Rechav, 1986; Horak & Maclvor, 1987). Theiler hare, Pronolagus rupestris (Lagomorpha, Lepori­ (1962) lists various other small mammals, particu­ dae). They have also been collected from various larly insectivores and rodents, as hosts of the imma­ rodents (Theiler, 1962; Rechav, 1986; Horak & ~ur~s, but their role in the ma.intenance of this spec­ Maclvor, 1987). The latter authors failed to obtain Ies IS at present open to questiOn. Horak & Maclvor this species from 42 birds, belonging to 6 species, (1987) found that the rodents they examined were that they examined in the same area. This observa­ mfested by H. truncatum, not this species. tion corroborates that of Hoogstraal et al. (1961), Distribution: H.m. rufipes is very widely distrib­ who did not include H. truncatum among the species uted in southern Africa. In South Africa it does not collected from the 340 northward-migrating birds, occur in the easternmost parts of the Transvaal, belonging to 22 "host forms", that they listed as tick­ Swaziland, KwaZulu, Natal, Orange Free State, infested in Egypt. These findings raise doubts about Transkei and eastern Cape Province, nor in most the accuracy of the records of H. truncatum imma­ parts of the southern and south-western Cape Pro­ tures from birds in Theiler (1962). Her records of vince, but is prevalent elsewhere. In Namibia it is this species from reptiles are thought to represent precariously established or absent in the driest areas accidental infestations. and dies out in the moister, more tropical parts of Distribution: In South Africa H. truncatum occurs northern Owambo, but occurs throughout the rest of almost throughout the country east, north and west the country (Theiler, 1956, 1962). It has also been of a line drawn from Barberton via Lydenburg to collected in many parts of Botswana (Walker et al., Pretoria and Krugersdorp, then southwards via 1978; Paine, 1982). Extralimitally it is widely distrib­ Bloemfontein to the coast at Alexandria. In the east uted in the Afrotropical region. its distribution zone extends from Barberton south- 90 JANE B. WALKER wards through eastern Swaziland and northern Kwa­ Ixodes bedfordi Arthur, 1959 Zulu/Natal as far as Nkandla (Theiler, 1956; 1962). Species diagnosis: Arthur (1965). Immature stages Apart from the dry coastal areas it is present unknown. throughout Namibia and in most parts of Botswana Hosts: The adults feed on rodents ( Otomys and (Theiler, 1962; Walker et al., 1978; Paine, 1982). Extralimitally it is widely distributed in the Afro­ Rhabdomys spp.) (Rodentia, Muridae). tropical region. Distribution: Lesotho, several localities. Theiler (1962) sugge~t.ed that South ~fric~n c?llections pre­ GENUS IXODES LATREILLE, 1796 viously identlfied as Ixodes nazrobzenszs , from Natal, Generic characteristics: Cathedral Peak, and Cape Province, Lady Gray, may also belong to this speci~s ; this cannot be con­ (a) Inornate firmed because these specimens apparently no (b) Hypostome and palps long longer exist. (c) Eyes absent Ixodes catherinei Keirans, Clifford & Walker, 1982 (d) Festoons absent Species diagnosis: Keirans et al. (1982). Only the (e) Anal groove anterior to the anus. (In all other type collection of 6 ~ ~ is known. genera the anal groove is posterior to the anus) Host: The scrub hare, Lepus saxatilis (Lagomor­ This is the second largest genus in southern Africa pha, Leporidae). (and in the world context by far the largest genus in Distribution: South Africa, eastern Cape Pro­ the family Ixodidae), containing 25 species plus vince, Clark's Siding, near Dordrecht. several entities in the pilosus group that are as yet undescribed. Ixodes cavipalpus Nuttall & Warburton, 1908 In general the Ixodes species remain poorly Species diagnosis: Arthur (1965). Nymph understood. Four species (catherinei, myotomys, spi­ unknown. culae, and transvaalensis) are known only from the Hosts: Adults have been collected from a wide few specimens in the type collections. Only one range of animals, inclu<;iing Prii?-ates, ~oth large an~ species in southern Africa, Ixodes rubicundus, is cur­ small species of domestic and wild carmvores (C~rni ­ rently known to be economically important as the vora, Canidae, Viverridae and Felidae) and vanous females cause tick l?aralysis in various species of antelopes (Artiodactyla, Bovidae). domestic and wild ammals. Distribution: It is probably the most widespread The identity of many collections of Ixodes spp. in Ixodes sp. in the Afrotropical region. In .south the Onderstepoort Tick Collection was confirmed by Africa, Transvaal, it has been recorded near Pieters­ C. M. Clifford (unpublished data, 1969). burg; at Nylsvley Nature Reserve, near Naboom­ spruit, and in Suikerbosrand Nature Reserve. In Ixodes alluaudiNeumann, 1913 south-east Botswana there are records from Macha­ Species diagnosis: Theiler (1941); Arthur (1965) . neng Mines near Kanye, and Otse. Hosts: Both the adults and immature stages prefer Ixodes corwini Keirans, Clifford & Walker, 1982 shrews, especially the greater musk shrew, Croci­ dura flavescens (Insectivora, Soricidae), also various Species diagnosis: Keirans et al. (1982). rodents (Rodentia, Bathyergidae and Muridae) Hosts: The adults and immature stages prefer Car­ (Theiler, 1962). nivora, including domestic dogs (Cani~ae), also the Distribution: Scattered localities throughout clawless otter, Aonyx capensis (Mustehdae); genets South Africa and in Lesotho. It also occurs extrali­ (Geizetta genetta, G. tigrina), and ~ongRwanda and Zai:re. Aliwal North and Grahamstown, origmally identi­ Ixodes pterodromae Arthur, 1960 fied as Ixodes nairobiensis, might belong to this spe­ cies. This cannot be confirmed because these ticks Species diagnosis: Arthur (1965). The holotype ~ apparently no longer exist. It also occurs extralimi- is said to have been deposited "in the Onderstepoort tally. . Research Station" but is not present now, and ts not recorded in the Onderstepoort Tick Collection cata­ Ixodes myotomys Clifford & Hoogstraal, 1970 logue. Larva unknown. Species diagnosis: Clifford & Hoogstraal (1970). Hosts: Primarily marine birds (wandering alba­ Only the type'? is known. tross, petrels, fleshfooted shearwater) (Aves) Host: The bush Karoo rat, Otomys unisulcatus (Arthur, 1965). It is also recorded from a feral cat. (syn. Myotomys unisulcatus) (Rodentia, Muridae). Distribution: Marion Island. It is widely distri­ Distribution: South Africa, western Cape Pro­ buted extralimitally. vince, Little Namaqualand, Port Nolloth. Ixodes rhabdomysae Arthur, 1959 Ixodes neitzi Clifford, Walker & Keirans, 1977 Species diagnosis: Arthur (1965). The holotype Species diagnosis: Clifford et al. (1977) . Immature '? "Onderstepoort Research Station collection stages unknown. No. 609", is said to have been deposited at the Hosts: Mountain reedbuck, Redunca fulvorufula, " Onderstepoort Research Station", but it is not pre­ and impala, Aepyceros melampus (Artiodactyla, sent now and is not listed in the Onderstepoort Tick Bovidae) (Clifford et al., 1977; Norval, Spickett & Collection Catalogue. Clifford, 1987). Host: The striped mouse, Rhabdomys pumilio Distribution: South Africa, Transvaal, Loskop (Rodentia, Muridae). Dam Nature Reserve (type locality). It also occurs Distribution: South Africa, eastern Cape Pro­ extralimitally in Zimbabwe. vince, Howison's Poort (given as Howieson's Poort), Ixodes pilosus group and Hellensbord (not located). A. M. Spickett (personal communication, 1990) considers that ticks recorded as Ixodes pilosus Koch, Ixodes rubicundus Neumann, 1904 1844 by Theiler (1950c, 1962) , plus other specimens Species diagnosis: Arthur (1965). obtained subsequently, can be separated into several Hosts: The adults are important parasites of closely related species which together constitute the sheep, cattle and goats, and to a lesser extent of Ixodes pilosus group. These ticks have been other domestic animals. They also parasitize wild recorded from a wide range of mammals, including animals, particularly the caracal, Felis caracal (~ar­ Insectivora, Primates, Carnivora, Artiodactyla, nivora, Felidae) and various antelopes, espectally Lagomorpha and Rodentia. the mountain reedbuck, Redunca fulvorufula, and The various members of this group occur pri­ eland, Taurotragus oryx (Artiodactyla, Bovidae ). marily in South Africa, where they have been The immature stages are most prevalent on Smith's recorded from widely separated areas in the red hare (red rock rabbit), Pronolagus rupestris (La­ Transvaal, Swaziland, Natal, Transkei and Cape gomorpha, Leporidae) and on caracal (Theiler, Province. One species has been recorded extralimi­ 1962; Horak, Moolman & Fourie, 1987a). tally in Zimbabwe (Norval et al. , 1987). Distribution: It is confined to South Africa, with Ixodes pilosus Koch, 1844 its main focus in "the southern parts of the Orange Species diagnosis: Nuttall & Warburton (1911). Free State and in the heart of the Karoo" (Spickett & Heyne, 1988). It occurs south-west of this main Hosts: All stages often feed on the same hosts. focus in Murraysburg, Aberdeen and Jansenville Cattle and dogs appear to be the most favoured do­ Districts, and southwards in Cradock District. It is mestic animals. Amongst wild animals various ante­ also present in a number of places in the western lopes have been recorded as hosts, particularly the Cape, notably in Sutherland, Worcester and Mon­ grey rhebok, Pelea capreolus; bushbuck, Tragela­ tagu Districts. North-east of its main focus I. rubi­ phus scriptus, and common duiker, Sylvicapra grim­ cundus appears to be established in the Transvaal in mia (Artiodactyla, Bovidae) (Horak et al. 1986b, both Heidelberg and Belfast Districts, and in Natal 1987c, 1989). The immature stages have also been in the Dargle (Lion's River) area. collected from several smaller animals, esrecially the caracal, Felis caracal; black-backed jacka , Canis Ixodes simplex simplex Neumann, 1906 mesomelas, and bat-eared fox, Otocyon megalotis Species diagnosis: Arthur (1965) . (Carnivora, Felidae and Canidae), and the scrub hare, Lepus saxatilis (Lagomorpha, Leporidae). Hosts: Bats, commonly on Schreiber's long-fin­ gered bat, Miniopterus schreibersi and Miniopterus Distribution: I. pilosus sensu stricto has been re­ sp., also on horseshoe bats, Rhinolophus spp. corded in southern Natal, Alfred District, in Weza ( Chiroe_tera, Vespertilionidae and Rhinolophidae). State Forest (Horak et al., 1989), and in Cape Pro­ Only ~ '? , nymphae and larvae are present m vince, both in the east, at various places in Albany the Onderspoort Collection. Males probably do not District, and in the south-west in the Bontebok feed, or feed rapidly and then detach again (Arthur, National Park (Horak et at., 1986b, 1987c). 1965). Ixodes procaviae Arthur & Burrow, 1957 Distribution: Recorded in South Africa, Species diagnosis: Arthur (1965). Larva unknown. Transvaal, from Makapansgat; Zebediela; Rooi- 92 JANE B. WALKER berg, near Warmbaths; Sterkstroom Cave; Pretoria, Ixodes ugandanus Neumann, 1906 and Zwartkop Cave; and in western Cape Province, Species diagnosis: Arthur (1965). from Kalk Bay and Bredasdorp. It is very widely distributed extralimitally. Host: The greater cane-rat, Thryonomys swinde­ rianus (Rodentia, Thryonomyidae). Ixodes simplex african us Arthur, 1956 Distribution: South Africa, "Zululand" (precise Species diagnosis: Arthur (1965). Only the type locality unknown), 1 ~ confirmed. Other speci­ ~ was previously known, from Kenya, Mt. mens from KwaZulu, Umfolozi, and Transvaal, Menengai. Tzaneen are regarded as questionable. [A ~ col­ Hosts: A long-fingered bat, Miniopterus sp. (1 lected at Louis Trichardt, originally listed under I. ~) (Chiroptera, Vespertilionidae) and "bat" (2 ugandanus by G. Theiler (1962), was reidentified nymphae). merely as "Ixodes sp." by C. M. Clifford (unpub­ lished data, 1969)]. It also occurs extralimitally. Distribution: In South Africa, Transvaal, it is recorded from Krugersdorp (Kromdraai, Grobbe­ Ixodes uriae White, 1852 laar's Lower Cave), and Sandspruit (possibly at Species diagnosis: Arthur (1965). Rooiberg, near Warmbaths). It also occurs extrali­ mitally in Kenya. Hosts: The females and immature stages feed on marine birds (Aves). In southern African waters Ixodes spiculae Arthur, 1956 various species of penguins, albatrosses, petrels, the Species diagnosis: Arthur (1956, 1965). His broadbilled prion, Cape hen, and kelp gull are description is based on a single d from South recorded as hosts. The males have rudimentary Africa, host and locality unknown. Arthur (1956, p. mouthparts, so probably do not feed. They have 184, last paragraph), in an apparent reference to this been collected in copula with ~ ~ in the species, not to Ixodes vespertilionis, commented: hosts' breeding places (Arthur, 1965). "The structure of Haller's organ on tarsus I most Distribution: Recorded in South Africa, eastern strongly suggests that it is a species associated with Cape Province, at Kleinemonde, and in western bats ... It is very probable that this male will prove Cape Province from Table Bay; also from Marion to be that of an already described female, but it is Island. It is very widely distributed extralimitally. unsafe at present to attribute it to any known spec­ ies". Ixodes vespertilionis Koch, 1844 Species diagnosis: Arthur (1965). Ixodes spinae Arthur, 1958 Hosts: Bats (Chiroptera). In southern Africa it has Species diagnosis: Arthur (1965). Immature stages been collected from Sundevall's leaf-nosed bat, Hip­ unknown. The holotype d and allotype ~, posideros caffer (Hipposideridae) and Temminck's from Procavia capensis, Pietersburg, South Africa, hairy bat, Myotis tricolor (Vespertilionidae). 20.IX. 1954, are said to have been deposited at the "Onderstepoort Veterinary Research Department" Distribution: Recorded in South Africa, but they are not present now, and are not recorded Transvaal, from Sterkfontein Cave, and in eastern in the Onderstepoort Tick Collection Catalogue. Cape Province, at Lootsberg. It is very widely dis­ The present whereabouts of these types, if they still tributed extralimitally. exist, is unknown. Host: The rock dassie, Procavia capensis (Hyra­ GENUS MARGAROPUS KARSCH, 1879 coidea, Procaviidae). Generic characteristics: Distribution: South Africa, Transvaal, Pieters­ (a) Inornate burg, and western Car,e Province, Cedarberg, from (b) Hypostome and palps short the farm "Middelburg '. It also occurs extralimitally. (c) Eyes present, but often extremely difficult to Ixodes theilerae Arthur, 1958 see in the males Species diagnosis: Arthur (1965). Only the ~ (d) Festoons absent. Conspicuous setae, either in is known. tufts or in a fringe, round the posterior end of Hosts: An avian parasite, collected from various the body of the male species of Passeriformes. (e) Adanal plates in the male present Distribution: A southern African species, (f) Leg segments in the male greatly expanded recorded in South Africa from the Transvaal, Preto­ This Afrotropical genus contains only 3 species, of ria and Onderstepoort; KwaZulu, Ndumu; Orange which one (winthemi)) occurs in South Africa and Free State, Ficksburg and Bothaville; western Cape the other 2 (reidi and wileyi) in eastern Africa. At Province, Cape Town; also in Namibia, Marienthal one time Margaropus winthemi was thought to have District (formerly Gibeon) at Marienthal itself. been introduced from Valparaiso, Chile, but this Ixodes transvaalensis Clifford & Hoogstraal, 1966 theory has now been discounted (Theiler, 1962). Species diagnosis: Clifford & Hoogstraal (1966). Margaropus is closely related to Boophilus and, Only the type collection of 2 ~ ~ is known. like the blue ticks, M. winthemi is a one-host species. Host: Found in the nest of a Namaqua rock Margaropus winthemi Karsch, 1879 mouse, Aethomys namaquensis (Rodentia, Muri­ Species diagnosis: Walker & Laurence (1973), dae). Gothe (1967), Arthur (1975a). Distribution: Collected in South Africa, western Hosts: As its common name, the winter horse tick, Cape Province, at Citrusdal. [The specific name indicates, its commonest domestic hosts are horses, transvaalensis is a misnomer, based on inadvertent though it also infests cattle. Massive infestations mislabelling of the ty_Ee collection at some point by have also been found on the Cape mountain zebra an unknown person (C. M. Clifford, personal com­ Equus zebra zebra (Perissodactyla, Equidae) munication, 1989)]. (Young, Zumpt, Boomker, Penzhorn & Erasmus, 93 A REVIEW OF THE IXODID TICKS OCCURRING IN SOUTHERN AFRICA 1973; Horak, Knight & DeVos, 1986a), and smaller "hyaena" (Hyaenidae); and cheetah, Acinonyx ju­ numbers on the gemsbok, Oryx gazella (Artiodac­ batus; leopard, Panthera pardus; lion, Panthera leo, tyla, Bovidae) (Horak eta!. 1983b). Another bovid, and "wild cat" (Felidae). Hosts of the immature the eland, Taurotrasus oryx, was recorded as a host stages are unknown. by Gertrud Theiler (unpublished data). Distribution: South Africa, Transvaal, in a belt Distribution: M. winthemi has been recorded only stretching from the Waterberg southwards to the in southern Africa. In South Africa it has been found Pretoria area and beyond as far as Potchefstroom in the Transvaal at a few places in the west and and Heidelberg Districts; Cape Province, in Rich­ north-west, also in the south-eastern Highveld; in mond, Hanover, Grahamstown and especially Clan­ Lesotho; at scattered points in the Orange Free william Districts; Namibia, at various places from State, especially in the south and west; in the higher Windhoek northwards, and Botswana at Nokanen, parts of southern Natal and the Transkei; at places in Ngamiland; Lobatse and in the Tati Concession. It the north-eastern and eastern Cape Province, in the occurs extralimitally in Zimbabwe. Cape Midlands and in areas bordering on the Karoo in the southern and western Cape Province (Theiler GENUS RHIPICEPHALUS KOCH, 1844 & Salisbury, 1958; Theiler, 1962; Horak et al., 1986a). Records in Namibia from Windhoek and Generic characteristics: Rehoboth District, and in Botswana from Francis­ (a) Most species inornate but 4 ornate, i.e. with a town, are thought to represent introductions only colour pattern on their scutum (maculatus and, (Theiler, 1962). in East Africa, pulchellus, humeralis, and dux) (b) Hypostome and palps short GENUS RHIPICENTOR NUTTALL & WARBURTON, 1908 (c) Basis capituli usually hexagonal (d) Eyes present Generic characteristics: (e) Festoons present (a) Inornate (f) Adanal plates in the male present (b) Hypostome and palps short This is the largest genus in southern Africa, com­ (c) Eyes present prising 28 species plus one entity that has yet to be (d) Festoons present described and several others whose taxonomic status (e) Adanal plates in the male absent is at present uncertain. Some of the information pre­ (f) Males with coxa IV much larger than the oth­ sented below is based on my re-examinations of ers and bearing 2 spurs specimens in the Onderstepoort Tick Collection. I. G. Horak (personal communications) has also This exclusively Afrotropical genus, containing kindly allowed me to incorporate some of his unpub­ only 2 species, was reviewed by Theiler (1961). Care lished data on the following species: arnoldi; capen­ is necessary to avoid confusing these ticks with Rhi­ sis sensu stricto; follis; glabroscutatum; lounsburyi; picephalus spp. oculatus; a new species near oculatus, and punctatus. Rhipicentor nuttalli can cause paralysis, which may It presents many taxonomic problems, especially be fatal, in dogs (Theiler, 1962; Norval & Colborne, to the beginner. As early as 1912 Cecil Warburton, 1985). in the opening paragraph of one of the most percep­ Rhipicentor bicornisNuttall & Warburton, 1908 tive papers ever published on this genus, summa­ rized the situation thus: "The identification of spec­ Species diagnosis: Theiler (1961). Nymph unknown. ies of Rhipicephalus is likely to give more trouble Hosts: Adults parasitize domestic dogs, and some­ than is the case with any other genus of Ixodidae, for times cattle. Wild hosts are primarily Carnivora, in­ while, on the one hand, there are few species which cluding the bat-eared fox, Otocyon megalotis, and depart greatly from the general type, on the other jackal, Canis sp. (Canidae); genets, Genetta sp. (Vi­ hand the range of variation withm the species is verridae), and the cats, especially the cheetah, Ac­ extremely great". Often this variability is dependant' inonyx jubatus; leopard, Panthera pardus, and lion, on nutrition, as was shown by Warburton's col­ Panthera leo, also 2 smaller species, the serval, Felis leagues G. H. F. Nuttall (1913) and N. Cunliffe serval, and African wild cat, Felis lybica (Felidae). It (1914) in their classic studies on Rhipicephalus has been recorded once from the eland, Taurotragus appendiculatus and R. sanguineus respectively. oryx (Artiodactyla, Bovidae). Hosts of the imma­ Recently it has also been shown that, under labora­ ture stages unknown. tory conditions, hybridization between certain clo­ Distribution: South Africa, Transvaal, in the Wa­ sely related species can occur (Zivkovic, Pegram, terberg and at Onderstepoort, and Namibia, where Jongejan & Mwase, 1986; Wouters, 1989; Wouters, it is widely distributed in the northern part of the Berkvens & Gomes, 1989). country, and is also recorded in the south from Be­ Considerable advances in our knowledge of this thanien District. Extralimitally it is widely distrib­ genus were made by Gertrud Theiler. In 1949 she uted in Central Africa. produced a valuable review of many little known Rhipicentor nuttam Cooper & Robinson, 1908 rhipicephalids, then in 1953 she and Britha Robin­ son published a detailed study of 6 of the lesser Species diagnosis: Theiler (1961). Nymph undes­ known species. Some of Theiler's taxonomic deci­ cribed. sions are not now regarded as valid, but it must be Hosts: Adults are recorded from domestic dogs, remembered that when she did much of her early and a donkey. The most commonly recorded wild taxonomic research it was difficult, if not impossible, hosts are the South African hedgehog, Atelerix fron­ for her to study the numerous type specimens that talis (Insectivora, Erinaceidae) and porcupine, Hys­ had been deposited in museum coilectwns in Britain trix africaeaustralis (Rodentia, Hystricidae). It has and Europe. This factor undoubtedly contributed also been found on a genet, Genetta sp. (Carnivora, largely to the erroneous conclusions that she some­ Viverridae); brown hyaena, Hyaena brunnea, and times reached. In later years she herself queried 94 JANE B. WALKER several of her earlier decisions, either in papers or in Felidae); Proboscidea (occasional adult ticks); Peris­ discussions with the writer. sodactyla (especially Equidae); and Rodentia (a few During the 1940s F. Zumpt also published a series species are recorded as hosts of the immatures, but of papers in which he discussed many Rhipicephalus sometimes all stages are present on the Hystricidae). spp. His interpretation of interspecific relationships Immature stages only are recorded from various was, however, seriously hampered by the fact that species of lnsectivora, Hyracoidea, the smaller he did not include the immature stages in his studies. species of Rodentia; the Lagomorpha, Leporidae, and birds (Aves), including both non-passennes and In recent years the use of scanning electron passerines (Theiler, 1962; Norval, Walker & microscopy has greatly facilitated our understanding Colborne, 1982). of this genus. Distribution: In southern Africa R. appendiculatus Several species of Rhipicephalus are known vec­ occurs from south-eastern Botswana eastwards tors of pathogens affecting domestic and wild ani­ across the Limpopo River, through the Transvaal mals, and to a lesser extent man, in Africa. By far Bushveld south of the Soutpansberg and into parts the most important of these is the brown ear tick, R. of the Lowveld, including the southern end of the appendiculatus, which is the primary vector of Thei­ Kruger National Park. It also occurs in a salient of leria parva parva, the protozoon causing East Coast the Bushveld extending into the northern end of the fever of cattle in East and Central Africa. It can also Kruger National Park at Pafuri, near the Zimbabwe transmit other pathogens, including Theileria parva and Mozambique borders. From the Transvaal Low­ bovis, Theileria parva lawrencei, Theileria taurotragi, veld its distribution extends southwards through Ehrlichia bovis, and Nairobi sheep disease virus, as Swaziland, most parts of Natal below c. 1500 m, the well as Rickettsia conori, the causative agent of tick­ Transkei, and the coastal areas of the eastern Cape bite fever in man (De Vas, 1981). Other vectors of as far as Port Elizabeth. Thus it is present in various the Theileria spp. affecting cattle are R. zambezien­ types of bushland and thornveld, but not in open sis, which can transmit all the T. parva group para­ grassland (Lessard, L'Eplattenier, Norval, Kundert, sites plus T. taurotragi (Lawrence, Norval & Uilen­ Dolan, Craze, Walker, Irvin & Perry, 1990). berg, 1983); R. duttoni, which transmits T. parva lawrencei in Angola (Da Gra~a & Serrano, 1971), The distribution of R. appendiculatus as shown by and R. evertsi evertsi, which can sometimes transmit Theiler (1949c) and Howell et al. (1978, Map 7) is in T. parva parva, though it is apparently not a very part incorrect. This tick is not now thought to occur efficient vector of this organism. m the north-western and northern Transvaal, between the Soutpansberg and the Limpopo River, Recently R. simus, which was shown to be capable where it was in the past confused with R. zambezien­ of transmitting Anaplasma marginale by Sir Arnold sis. It is sympatric with R. zambeziensis in parts of Theiler, has also proved to be a vector of Anaplasma the Transvaal. In the Fauresmith area the tick re­ centrale (Potgieter & Van Rensburg, 1987), and R. ferred to by Theiler (1949c) as a separate "variety or evertsi evertsi, long known to carry Babesia equi, has subspecies of R. appendiculatus" has now been iden­ now been established as a vector of Babesia caballi tified as R. punctatus (see below). Furthermore, R. (De Waal & Potgieter, 1987). R. sanguineus, the appendiculatus does not occur in the coastal areas of most cosmopolitan member of the genus, is a vector Cape Province, between Port Elizabeth and Cape of Babesia canis, Ehrlichia canis, and Hepatozoon Town, where it was confused with R. nitens (see canis in dogs, as well as R. conori in man. below). Some Rhipicephalus spp. are known to secrete Extralimitally R. appendiculatus is widely distri­ toxins that can have deleterious effects on their buted in East and Central Africa (Lessard et al., hosts. For example, cattle that are heavily infested with R. appendiculatus may develop a syndrome 1990). known as brown ear tick toxicosis. Other species Rhipicephalus arnoldi Theiler & Zumpt, 1949 have been associated with paralysis of animals and man. Among these are R. evertsi evertsi, which Species diagnosis: Theiler & Zumpt (1949). sometimes causes spring lamb paralysis, and R. Hosts: All stages prefer hares, especially red simus, which has occasionally been reported as caus­ hares, Pronolagus spp., (sometimes called red rock ing paralysis in man. Recently Fourie, Horak & rabbits) (Horak & Fourie, 1986), 9ccasionally also Marais (1988a) described cases of paralysis in the scrub hare, Lepus saxatilis, and Cape hare, Le­ Angora goats caused by an R. pravus-like tick that is pus capensis (Lagomorpha, Leporidae). Immature now believed to be R. punctatus (see below). stages only have been collected from the caracal, Felis caracal (Carnivora, Felidae) (Horak et al., Rhipicephalus appendiculatus Neumann, 1901 1987c); rock dassie, Procavia capensis (Hyracoidea) Species diagnosis: Walker, Norval & Corwin (Horak & Fourie, 1986), also the striped mouse, (1981). Rhabdomys pumilio, and a "field mouse" (Roden­ tia, Muridae). Hosts: All stages often feed on the same animals, and this tick has an extremely wide host range. It will Distribution: South Africa, Transvaal, at Maka­ parasitize all species of domestic animals, especially pan Caves, Naboomspruit, Onderstepoort and cattle, on which very large infestations may occur. Potchefstroom; Orange Free State, Fauresmith; and Sheep and goats are of secondary importance only. Cape Province, at various points in Richmond, Dogs and cats are rarely infested. It has also been Murraysburg, Graaff-Reinet and Cradock Districts. recorded from numerous species of wild animals. Its Extralimitally it has been recorded in Zimbabwe. preferred hosts are Artiodactyla; many species carry [Records from the Sudan (Hoogstraal, 1956) are all stages but the smaller antelopes are usually now considered incorrect]. infested by the immature stages only. It has been found less commonly on Primates (usually small Rhipicephalus capensis group infestations only); Carnivora (mostly immature Theiler (1962), in her review of the distribution of stages on the smaller species of Canidae, Viverridae R. capensis in South Africa, commented that this and Felidae, and adults on the Hyaenidae and larger specific name 'appears to be a catchall for "capensis- 95 A REVIEW OF THE IXODID TICKS OCCURRING IN SOUTHERN AFRICA like" ticks ..... ' Subsequent studies have confirmed and Schoemansdrift; Natal, Hluhluwe/Umfolozi this. In southern Africa the group at present com­ complex (Baker & Keep, 1970, as R. simpsoni) and prises 3 species: R. capensis sensu stricto, R. follis Weenen District at Muden; Orange Free State, at sensu stricto, and R. gertrudae. The status of at least Fauresmith; Cape Province, at various points in the one other entity occurring in Namibia, identified north-eastern and eastern parts of the province, also merely as a member of this group, has yet to be in the western Cape at Hout Bay; Namibia, mostly determined. north of Windhoek, at Opuwo (formerly Ohopoho), Earlier Theiler (1950a) reviewed and mapped the Sesfontein, Kamanjab, Kalkveld, Omaruru, Oka­ distribution of the ticks she was then identifying as hanja, Osona and Otjongombe, also south-west of R. capensis. Unfortunately many of the collections Windhoek at Naukluft; and Botswana, only in the on which her paper was based are no longer avail­ east at Tshesebe and Kanye. Widely distributed able. The accounts given below of the 3 currently extralimitally, mostly in East and Central Africa. recognized species in this group are based on exist­ Rhipicephalus duttoniNeumann, 1907 ing specimens only. Species diagnosis: Theiler (1949a). Immature Rhipicephalus capensis Koch, 1844 stages undescribed. Species diagnosis: Feldman-Muhsam (1960). Hosts: The adults apparently prefer ungulates Immature stages of this species sensu stricto undes­ (Artiodactyla, Bovidae), though few data are avail­ cribed. able. In southern Africa the sole recorded host is the black-faced impala, Aepyceros melampus petersi. In Hosts: Adults feed on cattle and horses. The only other parts of its range this tick occurs on most spec­ definite wild animals hosts recorded to date are the ies of domestic animals (Sousa Dias, 1950). mountain zebra, Equus zebra (Perissodactyla, Equi­ dae) and the eland, Taurotragus oryx (Artiodactyla, Distribution: In southern Africa it is recorded only Bovidae). This host list is undoubtedly very incom­ in north-western Namibia, Kaokoland 1, at Swart­ plete. Hosts of the immature stages of this particular booisdrift on the Cunene River and at Enyandi species have not as yet been recorded, although (Heloise Heyne, unpublished data). [The records in Theiler (1962) lists various rodents as hosts of imma­ Howard (1908) from the Transvaal, and from ture R. capensis sensu latu. Mozambique, based on a few specimens only, can­ not be checked because the ticks no longer exist; Distribution: It is recorded in South Africa only, these records are thought to be incorrect]. It occurs in Cape Province. With one exception, in eastern Cape Province, Cradock District, Mountain Zebra extralimitally in Angola and Zaire. National Park, all records are from western Cape Rhipicephalus evertsi evertsi Neumann, 1897 Province, in Namakwaland, Vanrhynsdorp, Clanwil­ Species diagnosis: Theiler (1943b ); Hoogstraal liam, Hopefield, Simonstown and Robertson Dis­ (1956); Howell et al. (1978) . tricts. Hosts: All stages frequently feed on the same Rhipicephalus deltoideus Neuman, 1910 hosts. This species parasitizes all domestic herbiv­ Species diagnosis: Theiler (1949a). The type col­ ores, especially horses, donkeys and cattle, but lection from Lesotho, originally comprising 1 C), 3 rarely dogs, and not cats. Amongst wild animals its preferred hosts include Burchell's zebra, Equus bur­ 2 ~, is the only one recorded from southern Africa. 6f this collection only 1 ~ syntype (Neumann No. chellii (Perissodactyla, Equidae) (Horak et a!., 1721, Nuttall Collection No. 2894, RML 111767) 1984a), and . other ungulates (Artiodactyla). It has now remains, deposited in the British Museum also been recorded from a Primate, the , Papio ursinus; occasionally from the large ~Natural History), London (Keirans, 1985) . Accord­ mg to Gertrud Theiler (personal communication), cats (Carnivora, Felidae); from an elephant, Loxo­ the types from the Neumann Collection were lost in donta africana (Proboscidea) and frequently from transit between South Africa and France after they wild hares, Lepus spp. (Lagomorpha, Leporidae). had been sent to her on loan. The immature stages have occasiOnally been col­ lected from various species of Insectivora; some of Host: Not recorded. the smaller Carnivora (Canidae, Viverridae and Fe­ Distribution: Lesotho, precise locality not lidae) (Horak et al., 1987c); Hyracoidea and Roden­ recorded, but said to be at an altitude of 2 135m. tia, also various birds (Aves), both non-passerines Rhipicephalus distinctus Bedford, 1929 and passerines. Species diagnosis: Theiler (1949a) . Immature Distribution: In South Africa R. evertsi evertsi oc­ stages undescribed. curs throughout the Transvaal; Swaziland; Natal and Zululand; the Transkei; the Orange Free State; and Hosts: All stages prefer the rock dassie, Procavia in Cape Province in the north-eastern and eastern capensis (Horak & Fourie, 1986) and "dassies" border districts, all along the southern coastal strip (species not recorded) (Hyracoidea). It has not as and up the west coast as far as Vanrhynsdorp. yet been collected from the tree dassie, Dendrohy­ Theiler (1950b) also recorded 2 populations further rax arboreus, in South Africa. One collection of north in Namakwaland, though m general she con­ adults from a klipspringer, Oreotragus oreotragus sidered this district to be free of the red-legged tick. (Artiodactyla, Bovidae), listed as R. simpsoni by It is not known whether these isolated populations Baker & Keep (1970), has proved on re-examination still exist. In Namibia R. evertsi evertsi is present in to be R. distinctus. A few Immature specimens have the moister northern part of the country and been recorded from caracal, Felis caracal (Carni­ throughout the Caprivi strip. In Botswana it has vora, Felidae) (Horak et al. 1987c). The record in been recorded at many places in the northern, east­ Theiler (1962) from the bushpig, Potamochoerus ern and southern parts of the country but more porcus (Artiodactyla, Suidae), IS based on an incor­ rarely in the western Kgalagadi (Walker et al., 1978; rect identification. Paine, 1982) . Theiler (1950b) noted that its spread is Distribution: In South Africa, Transvaal, recorded apparently limited by increasing aridity, the critical from Pietersburg, Gnderstepoort, Potchefstroom rainfall level being about 250 mm. Extralimitally it is 96 JANE B. WALKER one of the most widely distributed species in the widely distributed in Cape Province, including parts Afrotropical region. of the Karoo (Williston and Fraserburg Districts). In the east it extends into the southern and central Rhipicephalus evertsi mimeticus Donitz, 1910 Orange Free State, and in the west northwards Species diagnosis: Sousa Dias (1950). This subspe­ through Namakwaland into Namibia (J. B. Walker, cies has banded legs, so can easily be mistaken for a unpublished data; Heloise Heyne, unpublished Hyalomma by the unwary. Otherwise it closely data). It is not thought to occur extralimitally. resembles R. e. evertsi morphologically. Immature stages undescribed. Rhipicephalus glabroscutatum Du Toit, 1941 Hosts: All stages frequently feed on the same ani­ Species diagnosis: Du Toit (1941) mals. It apparently parasitizes the same hosts as Hosts: All stages feed on goats, and to a lesser R. e. evertsi, i.e. all the domestic herbivores and, extent on sheep and cattle (Macivor, 1985). Their amongst wild animals, Hartmann's mountain zebra, wild animal hosts are primarily various large and Equus zebra hartmannae (Perissodactyla, Equidae) small antelopes, especially browsers such as kudu, (Horak, Biggs & Reinecke, 1984b), and various Tragelaphus strepsiceros, and eland, Taurotragus species of Artiodactyla .. It is also .recorded from the oryx (Artiodactyla, Bovidae), _Elu~ the Cape mom~­ han, Pantheta leo (Carnivora, Fehdae). tain zebra, Equus zebra zebra (Penssodactyla, Eqm­ Distribution: Namibia, mostly north of the Tropic dae) (Young et al., 1973; Maclvor 1985, Horak et of Capricorn (Horak et al., 1984b; Heloise Heyne, al., 1986a). The immature stages also feed on the unpublished data, 1989), and western and southern scrub hare, Lepus saxatilis (Lagomorpha, Lepori­ Botswana (Walker et al., 1978; Paine, 1982). Extrali­ dae) (Horak & Knight, 1986). mitally it occurs in Angola and western Zaire. Distribution: This species is recorded only in Rhipicephalus Iollis Donitz, 1910 South Africa, Cape Province, primarily in the south­ Species diagnosis: Theiler (1949a). [The descrip­ east but with a few records from the central and tion of R. follis by Theiler & Robinson (1953b) ref­ southern parts of the province (Maclvor, 1985), also ers to another species: seeR. lounsburyi]. In the past from the west coast at Langebaan, Hopefield Dis­ this tick has frequently been misidentified as R. ca­ trict. pensis (Theiler, 1950a, 1962). Immature stages Rhipicephalus kochi Donitz, 1905 undescribed. Species diagnosis: Clifford, Walker & Keirans Hosts: The adults feed almost exclusively on the (1983). [Note that Yeoman & Walker (1967) resur­ large herbivores. Amongst domestic animals cattle rected the name Rhipicephalus jeanneli Neumann, are the most frequently recorded hosts. Most wild 1913 for the East African highland tick listed as R. hosts are antelopes, especially the eland, Taurotra­ kochi in Theiler (1962)]. gus oryx (Artiodactyla, Bovidae). Adults have also been recorded once from a cheetah, Acinonyx juba­ Hosts: In South Africa adults and nymphae have tus, and larvae from the caracal, Felis caracal (Carni­ been recorded from kudu, Tragelaphus strepsiceros; vora, Felidae) (Horak et al., 1987c). The preferred nyala, Tragelaphus angasii, and bushbuck, Tragela­ hosts of the immature stages are, however, rodents phus scriptus (Artiodactyla, Bovidae) (Horak et al., (Rodentia, Muridae). 1983b), also the warthog, Phacochoerus aethiopicus (Suidae) (Horak, Boomker, De Vas & Potgieter, Distribution: Present indications are that R. follis 1988b), and scrub hare, Lepus saxatilis (Lagomor­ occurs only in South Africa, where it is widely dis­ pha, Leporidae). Extralimitally it has been recorded tributed in the south-eastern Transvaal; central from a very wide range of both domestic and wild Orange Free State, Natal, and Cape Province, pri­ animals, mostly Artiodactyla, Bovidae and Suidae, marily in the south-east but with a few records from but including various species of Carnivora, Felidae; the southern coastal areas [Horak et al., 1986a, Proboscidea; Perissodactyla, Rhinocerotidae and 1987c, both as Rhipicephalus sp. (near Equidae; Rodentia, Hystricidae; Lagomorpha, and R. capensis); Horak et al., 1989; J. B. Walker, un­ Macroscelidea, and even a ground-feeding bird, the published data]. black-bellied korhaan, Lissotis melanogaster (Aves) Rhipicephalus gertrudae Feldman-Muhsam, 1960. (Clifford et al., 1983). Species diagnosis: Feldman-Muhsam (1960). This Distribution: In South Africa R. kochi has been tick was included under R. capensis by Theiler found at Pafuri, at the northern end of the Kruger (1950a, 1962). Biggs & Langenhoven (1984), who National Park (Horak et al. , 1983b, 1988b), and studied the seasonal incidence of ticks on cattle on a Ndumu, in KwaZulu. Extralimitally it is widely dis­ farm in Windhoek District, Namibia, were unable to tributed in East and Central Africa. distinguish R. gertrudae consistently from another R. capensis group tick occurring in the same area. Rhipicephalus longiceps Warburton, 1912 Immature stages undescribed. Species diagnosis: · Theiler (1949a). Immature Hosts: The adults are almost exclusively parasites stages unknown. of the larger herbivores. Amongst domestic animals Hosts: This rare tick has been recorded once from this species has been recorded from cattle, sheep, cattle. Wild animal hosts all belong to the Artiodac­ goats, horses and a donkey. Wild animal hosts tyla: the klipspringer, Oreotragus oreotragus, and include the mountain zebra, Equus zebra (Perisso­ gemsbok, Oryx gazella (Bovidae), also warthog, dactyla, Equidae), various large antelope species Phacochoerus aethiopicus (Suidae) (Horak, Biggs, (Artiodactyla, Bovidae), and the porcupine, Hystrix Hanssen & Hanssen, 1983a). africaeaustralis (J. B. Walker, unpublished data). Distribution: Namibia, on the farm "Okonjima" Hosts of the immature stages have not as yet been (20° 51' S, 16° 40' E); at Omandumba, Omaruru; recorded but are likely to be small mammals, pro­ Okahandja (Santos Dias, 1958); near Sukses, and on bably rodents. the farm "Lichtenstein-Sud" (22° 51' S, 17° 01' E), Distribution: In South Africa R. gertrudae is near Windhoek. Extralimitally it occurs in Angola. 97 A REVIEW OF THE IXODID TICKS OCCURRING IN SOUTHERN AFRICA Rhipicephalus lounsburyi Walker, 1990 lensi. It occurs extralimitally in eastern Africa, Species diagnosis: Walker (1990). It was originally mainly in the coastal regions of Mozambique, Tan­ described as Rhipicephalus follis by Theiler & Rob­ zania and Kenya. inson (1953b). Rhipicephalus muehlensi Zumpt, 1943 Hosts: The only domestic animals recorded thus far are sheep. It has been collected from the follow­ Species diagnosis: Salisbury (1959) . ing wild ungulates (Artiodactyla, Bovidae): bonte­ Hosts: All stages often feed on the same hosts. bok, Damaliscus dorcas , and grey rhebok, Pelea This tick sometimes occurs, in very small numbers, capreolus (Horak et al., 1986b), also the black wilde­ on cattle, and very occasionally on sheep and goats. beest, Connochaetes gnou; eland, Taurotragus oryx, It is commonest on various species of large and small and mountain reedbuck, Redunca fulvorufula. The wild antelopes, especially nyala, Tragelaphus anga­ hosts of the immature stages are still unknown. sii, and bushbuck, Tragelaphus scriptus (Artiodac­ Distribution: It has been recorded only in South tyla, Bovidae) (Horak et al., 1983b, 1988a), and on Africa, most commonly in eastern Cape Province, the wild pigs (Suidae). It has also been recorded especially in the mountainous areas of Barkly East, from various small carnivores (Canidae and Musteli­ Dordrecht and Cradock Districts. In western Cape dae); the elephant, Loxodonta africana (Probosci­ Province it has been found near Swellendam (Horak dea), both species of rhinoceros, Diceros bicornis et al., 1986b) and between Clan william and Graaf­ and Ceratotherium simum, and Burchell's zebra, water, and once in Natal, in the Impendle area. Equus burchellii (Perissodactyla, Rhinocerotidae and Equidae) (Baker & Keep, 1970). Rhipicephalus lunulatus Neumann, 1907 Distribution: South Africa, in the coastal areas of Species diagnosis: Walker, Keirans, Pegram & northern Natal, often in the same places as R. macu­ Clifford (1988). It was synonymized with Rhipice­ latus. It occurs extralimitally in the coastal areas of phalus tricuspis by Theiler (1949a, 1962), but this eastern Africa. finding was never universally accepted. Rhipicephalus neumanni Walker, 1990 Hosts: The adults will parasitize almost all species of domestic animals, especially cattle and often Species diagnosis: Walker (1990) . dogs. Amongst wild animals R . lunulatus has an ex­ Hosts: Primarily sheep, and to a lesser extent tremely wide host range. The commonest hosts in­ goats. It has been found once on a horse, and once clude the African buffalo, Syncerus caffer, and many on a gemsbok, Oryx gazella (Artiodactyla, Bovidae) species of large and small antelopes (Artiodactyla, (J.B. Walker, unpublished data). Bovidae), also the bushpig, Potamochoerus porcus, Distribution: South Africa, Cape Province, in and warthog, Phacochoerus aethiopicus (Suidae). It scattered localities in the Karoo, and Namibia, has been collected quite frequently from various car­ mainly in Bethanien and Keetmanshoop Districts, nivores (Canidae, Viverridae and Felidae). A few plus one record from north of Windhoek at Oman­ records, probably representing accidental records dumba, Omaruru (J.B. Walker, unpublished data). only, exist from other mammals, including man, and It is not known to occur extralimitally. 2 from water birds (Aves). Hosts of the immature stages are unknown. Rhipicephalus nitens Neumann, 1904 Distribution: In South Africa it occurs in the nor­ Species diagnosis: Neumann (1904). Immature thern and eastern Transvaal and Natal, and in stages undescribed. It was listed by Zumpt (1949) as Botswana mainly in and around the Okavango delta, a synonym of R. appendiculatus. Subsequently also in the east at Tshesebe. Extralimitally it is very Theiler (1962) commented: "Present day findings widely distributed in the Afrotropical region. tend to show that this is a valid species ...... ". Keirans (1985) also regarded it as valid. Rhipicephalus maculatus Neumann, 1901 Hosts: All stages will feed on the same host Species diagnosis: Theiler & Robinson (1953b). (Horak et al. , 1986b). Amongst domestic animals it Hosts: All stages often feed on the same hosts. strongly favours sheep, sometimes feeds on goats, This tick has sometimes been collected from cattle, and occasionally occurs on dogs (Horak et a/., and very occasionally from sheep and goats, but not 1987c). Known wild animal hosts are mostly ante­ from other domestic animals. The adults, and some­ lopes: bontebok, Damaliscus dorcas dorcas; spring­ times the immature stages, occur on many of the buck, Antidorcas marsupia/is, and grey rhebok, larger game animals, e.g. the large cats and hyaenas Pelea capreolus (Artiodactyla, Bovidae) (Horak, (Carnivora, Felidae and Hyaenidae); the elephant, Meltzer & De Vos, 1982a; Horak, Brown, Boom­ Loxodonta africana (Proboscidea); both species of ker, DeVos & Van Zyl, 1982b; Horak, DeVos & rhinoceros, Diceros bicornis and Ceratotherium De Klerk, 1982c; Horak et al. , 1986b). Others simum, and Burchell's zebra, Equus burchellii include the mountain zebra, Equus zebra (Perisso­ (Perissodactyla, Rhinocerotidae and Equidae); the dactyla, Equidae) (J.B. Walker, unpublished data) buffalo, Syncerus caffer, and various large antelopes and especially the scrub hare, Lepus saxatilis (Lago­ (Artiodactyla, Bovidae), and the wild pigs (Suidae) morpha, Leporidae) (Horak et al., 1986b). (Baker & Keep, 1970; Horak et al., 1983b). The Distribution: South Africa, Cape Province, mainly immature stages have been collected from some of in the southern coastal strip from Port Elizabeth the smaller carnivores, including the civet, Civettictis westwards, in association with Cape shrubland (fyn­ civetta (Viverridae) and serval, Felis serval (Feli­ bos) vegetation (White, 1983). It has also been dae); and the smaller antelopes, impala, Aepyceros found, again in association with fynbos, between melampus, and various duikers (Baker & Keep, Grahamstown and Paterson. Howell et al. (1978) 1970); also the rock dassie, Procavia capensis (Hyra­ erroneously included the distribution zone of R. coidea), and scrub hare, Lepus saxatilis (Lagomor­ nitens with that of R. appendiculatus. pha, Leporidae). Distribution: South Africa, in the coastal areas of Rhipicephalus oculatus Neumann, 1901 northern Natal, often in the same places as R. mueh- Species diagnosis: Neumann (1901). Immature 98 JANE B. WALKER stages undescribed. According to C. M. Clifford & phalus pravus-like tick (Fourie, Horak & Marais, J. E. Keirans (personal communication), their 1988a, b). examination of the syntype C) (Zoological Museum, Hosts: In South Africa adults are recorded from Berlin 17613) and ~ (ZMB 17614) of R. oculatus cattle, sheep, goats, and various wild. ungulates, from "Damaraland, D.S.W. Afrika" revealed that including giraffe, Giraffa camelopardabs; tsessebe, these ticks do not belong to the same species as the Damaliscus lunatus; steenbok, Raphicerus campes­ series described under this name (OP 2810) by tris · impala, Aepyceros melampus; gemsbok, Oryx Theiler & Robinson (1953b). Theiler (1962) listed R. galella, and eland, Taurotragus oryx (Artiodactyla, oculatus from a wide range of hosts and from various Giraffidae and Bovidae). All stages feed on scrub countries in East and Central Africa, though she hares, Lepus saxatilis (Lagomorpha, Leporidae) and gueried its presence in Kenya, Uganda and Tanzania the immature stages only on rock elephant shrews, (formerly Tanganyika). Her records are now known Elephantulus myurus (Macroscelidea). to include at least 3, possibly 4, related species: R. oculatus sensu stricto; R. sp. near oculatus (see Distribution: In South Africa R. punctatus is below); R. pravus in East Africa (Walker, 1974), widely distributed in the western, northern and and possibly R. punctatus in Central and southern north-eastern Transvaal, plus the southern Orange Africa. Only host and distribution records presently Free State and adjacent districts south of the Orange known to refer to R. oculatus sensu stricto are in­ River in north-eastern Cape Province. In Botswana cluded below. it is recorded in the eastern parts of Central and Ghanzi Districts. Extralimitally it occurs in parts of Hosts: Thus far this species has been recorded East and Central Africa. virtually exclusively from hares, especially the scrub hare, Lepus saxatilis, and sometimes the Cape hare, Rhipicephalus sanguineus (Latreille, 1806) Lepus capensis, and Smith's red hare, Pronolagus Species diagnosis: Pegram, Keirans, Clifford & rupestris (Lagomorpha, Leporidae). R. H. N. Walker, 1987bf These authors, who re-examined Smithers (personal communication) commented that numerous collections originally identified as R. "Lepus timidus", from which the Damaraland types sanguineus, found that many records listed under were recorded by Neumann (1901), was probably L. this name by Theiler (1962) refer to R. turanicus, a capensis, though it could possibly have been L. saxa­ species not then known to occur in the Afrotropical tilis. region (see below). Distribution: South Africa, Orange Free State, Hosts: It is primarily a parasite of dogs, and is only Fauresmith, and Cape Province, from scattered occasionally found on other hosts. If not controlled localities in the Karoo and eastern Cape Province; it will readily establish itself and breed in kennels Namibia, between Windhoek and Walvis Bay, also (Horak, 1982) and human habitations. in Damaraland. It is not known to occur extralimi­ tally. Distribution: Theiler (1962) considers that in South Africa R. sanguineus "is prevalent through­ Rhipicephalus sp. near oculatus out, though somewhat rare in the drier parts". This is probably true but it requires confirmation. The Species diagnosis: Theiler & Robinson (1953b) as only specimens present in the Onderstepoort Collec­ R. oculatus; see above under R. oculatus sensu tion are from the Pretoria area and Johannesburg. In stricto. Only host and distribution records for this Namibia it is widely distributed (Heloise Heyne, un­ species that have been confirmed recently are listed published data). It is the only cosmopolitan species below. m this genus. Hosts: Domestic animal hosts recorded for the adults are most commonly cattle and sheep, some­ Rhipicephalus simpsoni Nuttall, 1910 times goats, and only occasionally other species. Species diagnosis: Adults (Theiler, 1949a); nymph Known wild animal hosts are mostly antelopes, (Clifford & Anastos, 1964); larva undescribed. especially gemsbok, Oryx gazella, and kudu, Trage­ Hosts: A specific parasite of the greater cane ra!, laphus strepsiceros, also springbuck, Antidorcas Thryonomys swinderianus (Rodentia, Thryonomyl­ marsupia/is (Artiodactyla, Bovidae). Only a few dae). Other hosts are rare. The record in Baker & . records exist from the Cape hare, Lepus capensis, Keep (1970) from the klipspringer, Oreotragus area­ and scrub hare, Lepus saxatilis (Lagomorpha, Lepo­ tragus (Artiodactyla, Bovidae), was based on a mis­ ridae). identification (see above under R. distinctus). Distribution: In South Africa this species has as Distribution: South Africa, Transvaal, Nylstroom yet been recorded only in eastern Cape Province, and Komatipoort, and KwaZulu, Ndumu. It is wi­ south of the line from Jansenville-Mountain Zebra dely distributed extralimitally. National Park-Bucklands farm (33° 05' S, 26° 43' E), north-east of Grahamstown. In Namibia it is Rhipicephalus simus Koch, 1844 widely distributed, especially south of Windhoek. Species diagnosis: Pegram, Walker, Clifford & Keirans (1987c). At least 3 species were included Rhipicephalus punctatus Warburton, 1912. under this name by Theiler (1962), i.e. R. simus Species diagnosis: Clifford et al. (1983.) It is one of sensu stricto (southern and central Africa); R. prae­ the 3 closely related species included under the name textatus (eastern Africa), and R. muhsamae (western Rhipicephalus pravus Donitz, 1910 in Theiler (1962): Africa). The precise boundaries between the ranges R. pravus sensu stricto, which occurs in eastern of these species have not as yet been determined. Africa; R. kochi sensu stricto, whose range extends Hosts: Adults feed readily on cattle and dogs but from eastern to southern Africa (see above), and R. apparently rarely on other domestic animals. Adults punctatus, which is now thought to range from east­ have also been recorded from many wild animals, ern to southern Africa. It features in the southern especially Carnivora (Canidae, Viverridae, Proteli­ African literature as an atypical strain of R. appendi­ dae, Hyanidae and Felidae); Perissodactyla, particu­ culatus occurring in the Fauresmith area (Theiler, larly Burchell's zebra, Equus burchellii (Equidae); 1949c); as R. pravus (Paine, 1982), and as a Rhipice- Artiodactyla, especially the warthog, Phacochoerus 99 A REVIEW OF THE IXODID TICKS OCCURRING IN SOUTHERN AFRICA aethiopicus (Suidae), and the larger species of Bovi­ accepted (seeR. lunulatus). Only confirmed records dae, particularly the buffalo, Syncerus caffer, and are reviewed below. Rodentia, primarily the porcupine, Hystrix afri­ Hosts: Adults have been collected from cattle, caeaustralis (Hystricididae). Other recorded hosts goats and dogs but are recorded more frequently include Insectivora (Erinaceidae, the southern Afri­ from relatively small wild animals. These include can hedgehog, Atelerix frontalis); Tubulidentata (the wild antelopes, mostly the smaller species, especially aardvark, Orycteropus afer) and Pholidota (the the steenbok, Raphicerus campestris (Artiodac­ pangolin, Manis temminckii) (Norval & Mason, tyla, Bovidae); hares, Lepus spp. (Lagomorpha, 1981; Horak et al., 1983b, 1984b, 1987c, 1988b ). The Leporidae); the spring hare, Pedetes capensis immature stages parasitize various species of rats (Rodentia, Pedetidae), and various jackals and and mice (Rodentia, Muridae). other wild canids (Carnivora, Canidae). Hosts of the Distribution: In the absence of evidence to the immature stages are unknown; they are probably contrary it has been assumed that the South African small mammals. records of R. simus given by Theiler (1962) refer to Distribution: South Africa, at scattered localities this species sensu stricto. She indicated that it is in the Transvaal and north-eastern Cape Province; prevalent in the north-eastern and eastern northern Namibia, in Kavango District, and Transvaal, and in Marico, Brits and Pretoria Dis­ Botswana, primarily in Ngamiland and in the south­ tricts, but is either sparsely distributed or absent eastern part of the country. It also occurs extralimi­ elsewhere in this rrovince. It is also prevalent in tally. Swaziland, much o Natal, and in the coastal districts of the Transkei, Ciskei and the eastern Cape as far Rhipicephalus turanicus Pomerantsev, 1936 as Port Elizabeth. There are scattered records from Species diagnosis: Pegram et al. (1987a) . Theiler parts of the Karoo and western Cape but R. simus (1962) did not list this species, and the re-examina­ has not been recorded from much of western and tion of specimens in the Onderstepoort Tick Collec­ northern Cape Province, the inland districts of the tion has shown that she included it with R. sangui­ eastern Cape nor the Orange Free State. In Namibia neus (see above). this species is recorded from scattered localities, mostly north of Windhoek, and in Botswana mainly Hosts: Adults of this species have been recorded from the eastern part of the country and from Nga- from a very wide range of both domestic and wild miland. It also occurs extralimitally. ' animals. They occur on cattle, but rarely in large numbers. Norval, Daillecourt & Pegram (1983), Rhipicephalus sulcatus Neumann, 1908 who recorded it as a Rhipicephalus sp., considered Species diagnosis: Pegram, Clifford, Walker & that sheep and goats are the more usual domestic Keirans (1987a). Only records confirmed by these hosts, and their ears may become very heavily in­ authors are included below. fested. Dogs are somewhat less favoured than sheep and goats. Amongst wild animals R . turanicus has Hosts: The commonest recorded hosts of the been collected from hares, Lepus spp. (Lagomor­ adults are dogs and hares, Lepus spp. (Lagomorpha, pha, Leporidae); many species of carmvores (Carni­ Leporidae). Adults are also listed from cattle; vora, Canidae, Viverridae and Felidae); various various wild carnivores (Carnivora, Canidae, Viver­ ungulates (Perissodactyla and Artiodactyla), and ridae and Felidae) and a few antelopes (Artiodac­ sundry ground-feeding birds (Aves). Ostriches tyla, Bovidae) (Pegram et al., 1987a). The hosts of (Struthio camelus) may be infested by large numbers the immature stages are unknown. of this tick. Hosts of the immature stages unknown. Distribution: South Africa, "Masekwa" [r.resu­ mably Masekwaspoort (22° 53' S, 29° 53' E)] and Distribution: South Africa, Transvaal, from scat­ tered localities, and Natal, Bergville; northern Botswana, Sekoma Pan. It is widely distributed Namibia, from Etosha National Park and Grootfon­ extralimi tally. tein. It is widely distributed extralimitally. Rhipicephalus theileri Bedford & Hewitt, 1925 Species diagnosis: Theiler (1949a). Immature Rhipicephalus zambeziensis Walker, Norval & Cor­ stages undescribed. win, 1981 Hosts: Primarily a group of small mammals that Species diagnosis: Walker et al. (1981). often inhabit the same burrow systems: the suricate, Hosts: A wide range, with all stages often feeding Suricata suricatta, and yellow mongoose, Cynictis on the same hosts. Domestic animal hosts comprise penicillata (Carnivora, Viverridae), plus the ground cattle and, less commonly, horses and dogs. Wild squirrel, Xerus inauris, (Rodentia, Sciuridae). It animal hosts include various carnivores, particularly also parasitizes the tree squirrel, Paraxerus cepapi. It the large cats (Canidae, Viverridae, Hyaenidae and is occasionally recorded from other hosts (Theiler, Felidae) and numerous ungulates (mostly Artiodac­ 1962). tyla, especially the Suidae and Bovidae) (Norval et Distribution: South Africa, Transvaal, in the Mes­ al., 1982; Horak, 1987; Horak et al., 1983b, c, sina, Naboomspruit and Bloemhof areas, plus 1984b, 1987c, 1988b). Incidental hosts include the various places in the Orange Free State and in the antbear, Orycteropus afer (Tubulidentata, Oryctero­ eastern and north-western Cape Province; Namibia, podidae); spring hare, Pedetes capensis (Rodentia, from scattered localities in the central part of the Pedetidae), and Cape hare, Lepus capensis (Lago­ country, ranging from Outjo and Grootfontein morpha, Leporidae). Districts southwards to Mariental (formerly Gi­ Distribution: South Africa, at scattered localities beon), and Botswana, Ngamiland, at Toteng and in the Transvaal, particularly at the southern end of Shoshong. Itis not known to occur extralimitally. the Kruger National Park; northern Namibia, in Kaokoland and Grootfontein Districts, and Rhipicephalus tricuspis Donitz, 1906 Botswana, in Ngamiland and in the south-eastern Species diagnosis: Walker et al. (1988). Theiler corner of the country (Walker et al., 1978; Norval et (1949a, 1962) synonymized R. lunulatus with this al., 1982). It also occurs extralimitally in parts of species but her finding was never universally East and Central Africa. 100 JANE B. WALKER

ACKNOWLEDGEMENTS BIGGS, H. C. & LANGENHOVEN, J. W., 1984. Seasonal preva­ lence of ixodid ticks on cattle in the Windhoek District of South I am most grateful to I.G. Horak for his generous West Africa!Namibia. Onderstepoort Journal of Veterinary permission to quote some of his unpublished data, Research, 51 , 175-182. and for his many constructive comments on the BOYCOTT, R. C. & BOURQUIN, 0., 1988. The South African manuscript in general; to J. E. Keirans for checking Tortoise Book. Johannesburg: Southern Book Publishers (Pty) various data for me; to Heloise Heyne for the use of Ltd. her unpublished data, especially from Namibia; for BROADLEY, D . G., 1983. FitzSimons' snakes of Southern Africa. drawing my attention to Haemaphysalis tauffliebi at 2nd edn. Johannesburg: Delta Books. BURR, E. W., 1983. Tick toxicosis in a crossbred terrier caused by the 11th hour, and for other useful information; to Hyalomma truncatum. Veterinary Record, 113,260-261. A. M. Spickett and T. N. Petney for their advice on CAMICAS, J. L., HOOGSTRAAL, H. & EL KAMMAH, K. , 1972. some of the Ixodes spp.; toW. H. Stoltsz for clarify­ Notes on African Haemaphysalis ticks. VIII. H. (Rhipistoma) ing some aspects of disease transmission by Rhipice­ moreli sp. n., a carnivore parasite of the H. (R.) leachi group phalus spp., and to W. D. Haacke for information (Ixodoidea, Ixodidae). Journal of Parasitology, 58, 1185-1196. on the nomenclature of reptiles. I also sincerely CLIFFORD, C. M. & ANASTOS , G ., 1964. Ticks. Exploration du thank Mrs C. A. Smith, and Mrs N. van Staden, for Pare National de Ia Garamba-Mission H. de Saeger, Fascicule their efficient processing of the manuscript for me. 44,40 pp. CLIFFORD, C. M. & HOOGSTRAAL, H ., 1966. Ixodes transvaalen­ REFERENCES sis n. sp. from a nest in the Republic of South Africa (Acarina: Ixodidae). Journal of Medical Entomology, 2, AR1HUR, D. R., 1956. The Ixodes ticks of Chiroptera (Ixodoi· 372- 373. dea, Ixodidae). 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FOURIE, L. J ., HORAK, I. G. & MARAIS, L ., 1988a. An unde­ HORAK, I. G ., 1982. Parasites of domestic and wild animals in scribed Rhipicephalus species associated with field paralysis of South Africa. XIV. The seasonal prevalence of Rhipicephalus Angora goats. Journal of the South African Veterinary Associa­ sanguineus and Ctenocephalides spp. on kennelled dogs in Pre­ tion, 59, 47--49. toria North. Onderstepoort Journal of Veterinary Research, 49, FOURIE, L. J., HORAK, I. G. & MARAIS, L., 1988b. The seasonal 6~8 . abundance of adult ixodid ticks on Merino sheep in the south­ HORAK, I. G., BROWN, MOIRA R., BOOMKER, J., DEVos, V. & western Orange Free State. Journal of the South African Veteri­ VAN ZYL, ELSA A., 1982b. Helminth and parasites nary Association, 59, 191-194. of blesbok, Damaliscus dorcas phillipsi, and of bontebok, Dam­ GOTHE , R. , 1967. Ticks in the South African Zoological Survey aliscus dorcas dorcas. Onderstepoort Journal of Veterinary Re­ Collection. Part XII. 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Onderste­ America, 53,445--457. poort Journal of Veterinary Research, 50,243-255. HOOGSTRAAL, H. , KAISER, M. N., TRAYLOR , M.A., GABER , HORAK, I. G ., DEVos, V. & DE KLERK, B. D ., 1984a. Parasites S. & GUINDY, E ., 1961. Ticks (Ixodoidea) on birds migrating of domestic and wild animals in South Africa. XVII. Arthropod from Africa to Europe and Asia. Bulletin of the World Health parasites of Burchell's zebra, Equus burchelli, in the eastern Organization, 24, 197-212. Transvaal Lowveld. Onderstepoort Journal of Veterinary Research, 51, 145-154. HOOGSTRAAL , H., 1963. Notes on African Haemaphysalis ticks. HORAK , I. G., BIGGS, H. C. & REINECKE, R . K. , 1984b. Arth­ V. Redescription and relationships of H. silacea Robinson, 1912 , from South Africa (Ixodoidea, Ixodidae). Journal of Pa­ ropod parasites of Hartmann's mountain zebra. Equus zebra rasitology, 49, 830-837. hartmannae, in South West Africa/Namibia. Onderstepoort Journal of Veterinary Research, 51, 183-187. HOOGSTRAAL, H., 1964. Notes on African Haemaphysalis ticks. VI. H. spinulosa Neumann, and its relation to biological and HORAK , I. G. & WILLIAMS, E. J., 1986. Parasites of domestic nomenclatorial problems in the H. leachii group of Africa and and wild animals in South Africa. XVIII. The crowned guinea Asia (Ixodoidea, Ixodidae). Journal of Parasitology, 50, fowl (Numida meleagris) , an important host of immature ixodid 786-791. ticks. Onderstepoort Journal of Veterinary Research, 53, HOOGSTRAAL, H. , WALKER, JANE B. & NEITZ, W. 0 ., 1971. 119-122. Notes on African Haemaphysalis ticks. VII. H. (Rhipistoma) HORAK, I. G. & FOURIE, L. J ., 1986. Parasites of domestic and hyracophila sp. n. (Ixodoidea: Ixodidae), a parasite of hyraxes wild animals in South Africa. XIX. Ixodid ticks and fleas on in South Africa. Journal of Parasitology, 57, 417--425. rock dassies (Procavia capensis) in the Mountain Zebra HOOGSTRAAL , H . & EL KAMMAH , K. M., 1972. Notes on Afri­ National Park. Onderstepoort Journal of Veterinary Research, can Haemaphysalis ticks. X. H. (Kaiseriana) aciculifer Warbur­ 53, 123-126. ton and H. (K.) rugosa Santos Dias, the African representa­ HORAK, I. G. , KNIGHT, M. M. & DEVos, V., 1986a. Parasites of tives of the spinigera subgroup. Journal of Parasitology, 58, domestic and wild animals in South Africa. XX. Arthropod 960-978. parasites of the Cape mountain zebra (Equus zebra zebra). HOOGSTRAAL, H., 1972. Notes on African Haemaphysalis ticks. Onderstepoort Journal of Veterinary Research, 53, 127-132. IX. H. (Rhipistoma) pedetes sp. n., a relict parasite of the HORAK, I. G ., SHEPPEY, K., KNIGHT, M. M. & BEUTHIN,C. L., springhare (Rodentia, Pedetidae) in Transvaal (Ixodoidea, Ixo­ 1986b. Parasites of domestic and wild animals in South Africa. didae). Journal of Parasitology, 58,979-983. XXI.Arthropod parasites of vaal ribbok, bontebok and scrub HOOGSTRAAL , H. & WASSEF , HILDA Y., 1973. The Haemaphy­ hares in the western Cape Province. Onderstepoort Journal of salis ticks (Ixodoidea: Ixodidae) of birds. 3. H. (Ornithophysa­ Veterinary Research, 53, 187-197. lis) subgen. n.: definition, species, hosts, and distribution in the HORAK, I. G. & KNIGHT, M. M., 1986. A comparison of the tick Oriental, Palearctic, Malagasy, and Ethiopian faunal regions. burdens of wild animals in a nature reserve and on an adjacent Journal of Parasitology, 59, 1099-1117. farm where tick control is practised. Journal of the South Afri­ HOOGSTRAAL, H. & EL KAMMAH, K. M., 1974. Notes on Afri­ can Veterinary Association, 57,199-203. can Haemaphysalis ticks. XII. H. (Rhipistoma) zumpti sp. n., a HORAK, I. G., MOOLMAN, L. C. & FOURIE, L. J., 1987a. Some parasite of small carnivores and squirrels in southern Africa. wild hosts of the Karoo paralysis tick, Ixodes rubicundus Neu­ Journal of Parasitology, 60, 188-197. mann, 1904 (Acari: Ixodidae). Onderstepoort Journal of HOOGSTRAAL, H. , 1979. The epidemiology of tick-borne Cri­ Veterinary Research, 54,49-51. mean-Congo hemorrhagic fever in Asia, Europe and Africa. HORAK, I. G. & MACIVOR, K. M. DE F., 1987. The scrub hare, a Journal of Medical Entomology, 15, 307--417 . reliable indicator of the presence of Hyalomma ticks in the HOOGSTRAAL , H. & WASSEF, HILDA Y., 1981. Notes on African Cape Province. Journal of the South African Veterinary Asso­ Haemaphysalis ticks. XIII. Identity of H. (Rhipistoma) cooleyi, ciation, 58, 15-19. a parasite of the rock hyrax in South Africa (Acarina: Ixodi­ HORAK, I. G ., MACIVOR, K. M. DE F., PETNEY, T. N. & DE dae). Onderstepoort Journal of Veterinary Research, 48, Vos, V., 1987b. Some avian and mammalian hosts of 135-140. Amblyomma hebraeum and Amblyomma marmoreum (Acari: HOOGSTRAAL, H. & AESCHLIMANN, A., 1982. Tick-host specif­ Ixodidae). Onderstepoort Journal of Veterinary Research, 54, icity. Bulletin de la Societe Entomologique Suisse, 55,5-32. 397--403. HOOGSTRAAL , H. & KIM, K. C., 1985. Chapter 10. Tick and HORAK, I. G ., JACOT GUILLARMOD, AMY, MOOLMAN, L. C. & mammal coevolution, with emphasis on Haemaphysalis. In: DEVos, V., 1987c. Parasites of domestic and wild animals in KIM , K.C. (ed.} Coevolution of parasitic and mam­ South Africa. XXII. Ixodid ticks on domestic dogs and on wild mals, 505-568. New York: John Wiley and Sons. carnivores. Onderstepoort Journal of Veterinary Research, 54, HORAK. I. G., MELTZER, D . G. A. & DEVos, V., 1982a. Hel­ 573-580. minth and arthropod parasites of springbok, Antidorcas marsu­ HORAK, I. G ., 1987. Arthropod parasites of some wild animals in pia/is, in the Transvaal and western Cape Province. Onderste­ South Africa and Namibia. Journal of the South African poort Journal of Veterinary Research, 49,7-10. Veterinary Association, 58, 207-211. 102 JANE B. WALKER

HORAK, I. G., KEEP,M. E ., FLAMAND,J. R. B. & BOOMKER,J. , MASON, C. A. & NORVAL. R. A . 1. , 1980. The ticks of Zim­ 1988a. Arthropod parasites of common reedbuck, Redunca babwe. 1. The genus Boophilus. Zimbabwe Veterinary Journal, arundinum, in Natal. Onderstepoort Journal of Veterinary 11 , 36-43. Research, 55, 19--22. MATTHYSSE, J. G . & COLBO, M. H., 1987. The ixodid ticks of HORAK, I. G., BOOMKER, J., DEVos, V. & POTGIETER, F. T ., Uganda, together with species pertinent to Uganda because of 1988b. Parasites of domestic and wild animals in South Africa. their present known distribution. Entomological Society of XXIII. Helminth and arthropod parasites of warthogs, Phaco­ Arnenca, College Park, Md., USA. choerus aethiopicus, in the eastern Transvaal Lowveld. Onder­ MEESTER , J. A . J ., RAUTENBACH , I. L. , DIPPENAAR. N. J. & stepoort Journal of Veterinary Research, 55, 145-152. BAKER, C. M. , 1986. Classification of Southern African Mam­ HORAK , I. G., KEEP, M. E., SPICKETT, A.M. & BOOMKER, J. , mals. Transvaal Museum, Pretoria, Monograph No. 5, [x] + 359 1989. Parasites of domestic and wild animals in South Africa. pp. XXIV. Arthropod parasites of bush buck and common duiker in MOREL , P. C., 1966. Sur quelques larves d'lxodes Latreille, 1796, the Weza State Forest, Natal. Onderstepoort Journal of Veterin­ d'Afrique (Acariens: Ixodoidea). A carologia, 8, 208-221. ary Research, 56, 6~6. MOREL. P. C. , "!969 . Contribution a Ia connaissance de Ia distri­ HOWARD, C. W., 1908. A list of the ticks of South Africa, with bution des tiques (Acariens, Ixodidae et Arnblyomrnidae) en descriptions and keys to all the forms known. Annals of the Afrique ethiopienne continentale, 388 pp. + annexe cartogra­ Transvaal Museum, 1, 73-169 + 16 plates. phique, cartes 1-62. D.Sc. thesis, University of Paris. HOWELL, C. J. , WALKER, JANE B. & NEVILL, E. M., 1978. Ticks, NEITZ, W. 0 ., 1956. A consolidation of our knowledge of the and infesting domestic animals in South Africa. transmission of tick-borne diseases. Onderstepoort Journal of Part 1. Descriptions and biology. Department of Agricultural Veterinary Re-search, 27,115-163. · Technical Services, Republic of South Africa, Science Bulletin, No. 393, 69 pp. NEITZ, W. 0., 1959. Sweating sickness: The present state of our knowledge. Onderste'poort Journal of Veterinary Science, 28 , HUSSEIN, H. S. & MUSTAFA, B. E., 1983. Haemaphysalis (Rhi­ 3-38. pistoma) spinulosa Neumann, 1906: Description of immature stages, adult structural variations, and notes on biology (Ixo­ NEUMANN. L. G., 1901. Revision de Ia famille de Ixodides. (4' . doidea: Ixodidae). Journal of Parasitology, 69,405-412. Memoire). Memoires de Ia Societe Zoologique de France, 14, JAGGER, T. D., WEDDERBURN, P. A. & MCCARTAN, B., 1987. 249--372. University of Edinburgh, Royal (Dick) School of Veterinary NEUMANN , L. G ., 1904. Notes sur les Ixodides-II. Archives de Studies, Veterinary Expedition to Swaziland 1985. Centre for Parasitologie, 8, 444-464. Tropical Veterinary Medicine, Easter Bush, Roslin, Midlo­ NOR VAL , R. A . l. , 1975a. Studies on the ecology of Haemaphysa­ thian, Scotland. lis silacea Robinson 1912 (Acarina, Ixodidae). Journal of Para­ KAUFMAN. T. S. , 1972. A revision of the genus Aponomma Neu­ sitology, 61,730-736. mann, 1899 (Acarina, Ixodidae). Ph.D . thesis, University of NORVAL, R. A. 1. , 1975b. Studies on the ecology of Amblyomma Maryland. rnarmoreum Koch 1844 (Acarina: Ixodidae). Journal of Parasi­ KEIRANS , J. E., 1985. George Henry Falkiner Nuttall and the tology, 61, 737-742. Nuttall Tick Catalogue. United States Department of Agricul­ NORVAL, R. A. I. & MASON , C. A ., 1981. The ticks of Zim­ ture, Agricultural Research Service, Miscellaneous Publication babwe. II. The life cycle, distribution and hosts of Rhipicepha­ No. 1438, 1785 pp. Washington D .C.: U.S. Government Print­ lus simus Koch, 1844. Zimbabwe Veterinary Journal, 12, 2-9. ing Office. NORVAL, R. A. I. , 1982. The ticks of Zimbabwe. IV. The genus KEIRANS , J. E., CLIFFORD , C. M. & WALKER, JANE B., 1982. Hyalomma. Zimbabwe Veterinary Journal, 13 , 2-10. The Ixodes (Afrixodes) oldi group (Acari: Ixodidae) from sub­ NORVAL , R. A . I. , WALKER, JANE B. & COLBORNE. J., 1982. Saharan Africa with descriptions of five new species. Journal of The ecology of Rhipicephalus zambeziensis and Rhipicephalus Medical Entomology, 19, 309--329 . appendiculatus (Acarina, Ixodidae) with particular reference to KNIGHT, M. M., NORVAL, R. A. I. & RECHAV , Y. , 1978. The life Zimbabwe. Onderstepoort Journal of Veterinary Research, 49, cycle of the tick Hyalomma marginatum rufipes Koch (Acarina: 181-190. Ixodidae) under laboratory conditions. Journal of Parasitology, NORVAL, R. A . 1. , DAILLECOURT. T. & PEGRAM . R. G., 1983. 64, 143-146. The ticks of Zimbabwe. VI. The Rhipicephalus sanguineus KNIGHT, M. M. & RECHAV, Y., 1978. Ticks associated with kudu group. Zimbabwe Veterinary Journal, 13, 38-46. in the eastern Cape: Prelimanary report. Journal of the South NORVAL, R. A. l. & COLBORNE, J., 1985. The ticks of Zim­ African Veterinary Association, 49,343-344. babwe. X. The genera Dermacentor and Rhipicentor. Zim­ LAWRENCE, J. A., NORVAL, R. A . l. & UILENBERG, G ., 1983. babwe Veterinary Journal, 16, 1-4. Rhipicephalus zambeziensis as a vector of bovine theileriae. NORVAL. R. A. 1., 1985. The ticks of Zimbabwe. XIV. The lesser Tropical Animal Health and Production, 15, 39-42. known Haemaphysalis species. Zimbabwe Veterinary Journal, LESSARD , P., L'EPLATTENIER, R., NORVAL , R. A. I., KUNDERT, 16, 54-59. K., DOLAN, T. T., CROZE, H., WALKER , J. B., IRVIN, A. D. & NORVAL , R. A. 1., SPICKETT, A . M. & CLIFFORD, C. M. , 1987. PERRY, B. D., 1990. Geographical information systems for stu­ The ticks of Zimbabwe. XVI. The genus Ixodes. Zimbabwe dying the epidemiology of cattle diseases caused by Theileria Veterinary Journal, 18, 1-10. parva. Veterinary Record, 126, 255-262. NUTTALL, G. H. F., WARBURTON , C., COOPER, W . F. & ROBIN­ LONDT, J. G. H. & ARTHUR, D. R ., 1975. The structure and SON, L. E ., 1908-1926. Ticks. A monograph of the Ixodoidea. parasitic life cycle ~f Booph!lus microplus (Canestrini, 1888) in Parts I-IV and Bibliographies [I] and II. London: Cambridge South Afnca (Acanna: Ixodtdae). Journal of the Entomological University Pr$!SS. Society of Southern Africa, 38, 321-340. NUTTALL. G. H. F. & WARBURTON, C., 1911. Ixodidae-Section LUCAS, J. M. S., 1954. Fatal anaemia in poulty caused by a heavy I: Classification; Section II: The genus Ixodes. Part II . In: NUT­ tick infestation. Veterinary Record, 66,573. TALL eta/. (1908-1926) . London: Cambridge University Press. NUTTALL. G. H. F., 1913. Rhipicephalus appendiculatus: Varia­ MACIVOR, K. M. DE F. & HORAK, I. G ., 1984. The internal and tion in size -and structure due to nutrition. Parasitology , 6, external parasites of Angora and Boer goats in Valley Bushveld near Uitenhage. Angora Goat and Mohair Journal26, 7-14. 195-203. PAINE, G. D., 1982. Ticks (Acari: Ixodoidea) in Botswana. Bulle­ MACIVOR, K. M. DE F., 1985. The distribution and hosts of tin of Entomological Research, 72, 1-16. Rhipicepahlus glabroscutatum. Onderstepoort Journal of Veteri­ nary Research, 52 , 43-46. PATTERSON, R . & BANNlSTER, A., 1987. Reptiles of Southern Africa. Cape Town: C. Struik (Pty} Ltd. MACIVOR, K. M. DE F. & HORAK, I. G. , 1987. Foot abscess in goats in relation to the seasonal abundance of adult Amb­ PEGRAM , R. G., CLIFFORD , C. M., WALKER , JAN E B. & KEI­ lyomma hebraeum and adult Rhipicephalus glabroscutatum RANS, J. E.,. 1987a. Clarification of the Rhipicephalus sangui­ (Acari: Ixodidae). Journal of the South African Veterinary As­ neus group (Acari, Ixodoidea, Ixodidae). I. R. sulcatus Neu­ sociation, 58, 113-118. mann, 1908 and R. turanicus Pomerantsev, 1936. Systematic Parasitology, 10, 3-26. I. & I., MACKENZIE, P. K. NORVAL, R. A. 1980. The transmis­ PEGRAM, R. G., KEIRANS , J. E .. CLIFFORD, C. M. & WALKER , sion of Cowdria ruminantium by Amblyomma tholloni. Veteri­ JANE B., 1987b. Clarification of the Rhipicephalus sanguineus nary Parasitology, 7, 265-268. group (Acari, Ixodoidea, Ixodidae). II . R. sanguineus (La­ MACLEAN, G. H., 1985. Roberts' birds of Southern Africa. Cape treille, 1806) and related species. Systematic Parasitology, 10, Town: John Voelcker Bird Book Fund. 27-44.

103 A REVIEW OF THE IXODID TICKS OCCURRING IN SOUTHERN AFRICA

PEGRAM , R. G., WALKER, JANE B., CLIFFORD , C. M. & KEI­ THEILER, GERTRUD, 1949a. Ticks in the South African Zoologi­ RANS , J. E ., 1987c. Comparison of populations of the Rhipice­ cal Survey Collection. Part VI. Little known African rhipice­ phalus simus group: R. simus, R. praetext{ltus and R. muhsamae phalids. Onderstepoort Journal of Veterinary Science and (Acari: Ixodidae). Journal of Medical Entomolgy, 24,666--682. Animal Industry, 21,253--300. PE1NEY, T. N., HORAK, I. G . & RECHAV, Y., 1987. The ecol­ THEILER, GERTRUD, 1949b. Zoological Survey of the Union of ogy of the African vectors of heartwater,' with particular refer­ South Africa. Part II. Distribution of Boophilus (Palpoboophi­ ence to Amblyomma hebraeum and Amblyomma variegatum. lus) decoloratus, the blue tick. Onderstepoort Journal of Onderstepoort Journal of Veterinary Research, 54, 381-395. Veterinary Science and Animal Industry, 22, 255-268 + 1 map. POTGIETER, F. T . & VAN RENSBURG , L., 1987. Tick transmis­ THEILER, GERTRUD, 1949c. Zoological Survey of the Union of sion of Anaplasma centrale. Onderstepoort Journal of Veteri­ South Africa: Tick Survey. Part III. Distribution of Rhipicepha­ nary Research, 54,5-7. lus appendiculatus, the brown tick. Onderstepoort Journal of RECHAV, Y ., 1986. Seasonal activity and hosts of the vectors of Veterinary Science and Animal Industry, 22,269--284 + 1 map. Crimean-Congo haemorrhagic fever in South Africa. South Af­ THEILER, GERTRUD & ZUMPT, F ., 1949. Description of new rican Medical Journal, 69,364-368. species. Rhipicephalus (s. str.) arnoldi Theiler and Zumpt, n. ROBINSON , L. E ., 1926. The genus Amblyomma. Part IV. In: sp., pp. 111-119. In: ZUMPT, F. (1949) Preliminary study to a NUTTALL et al. (1908-1926) . London: Cambridge University revision of the genus Rhipicephalus Koch. Key to the adult ticks Press. of the genus Rhipicephalus Koch and description of two new species. Mor,;ambique, No. 60,57-123. SALISBURY, LOIS E ., 1959. Ticks in the South African Zoological Survey Collection. Part X. Rhipicephalus miihlensi Zumpt. THEILER, GERTRUD, 1950a. Zoological Survey of the Union of Onderstepoort Journal of Veterinary Research, 28, 125-132. South Africa. Tick Survey-Part IV. Distribution of Rhipice­ phalus capensis, the Cape brown tick. Onderstepoort Journal of SANTOS DIAS , J. A . T ., 1955. Subsidios para o estudo da fauna Veterinary Science and Animal Industry, 24,7-32 + 1 map. ixodol6gica da BechuanaHindia. Mem6rias e Estudos do Museu Zool6gico da Universidade de Coimbra, No. 231,9 pp. THEILER, GERTRUD , 1950b. Zoological Survey of the Union of South Africa. Tick Survey-Part V. Distribution of Rhipice­ SANTOS , DIAS , J. A . T., 1958. Notas ixodol6gicas. V. Acerca de phalus evertsi, the red tick. Onderstepoort Journal of Veterinary alguns ixodideos do Museu de Hamburgo. Mem6rias e Estudos Science and Animal Industry, 24, 33--36 + 1 map. do Museu Zool6gico da UniveTSidade de Coimbra, No. 253, 32 pp. THEILER, GERTRUD , 1950c. Zoological Survey of the Union of South Africa. Tick Survey-Part IV. Distribution of the SANTOS , DIAS , J., A . T., 1963. Contribui~ao para o estudo da ixodeds: Ixodes pilosus and Ixodes rubicundus. Onderstepoort sistematica dos Acaros da subordem Ixodoidea Banks, 1894. 1. Journal of Veterinary Science and Animal Industry, 24, 37-51 + Farru1ia Ixodidae Murray, 1877. Mem6rias e Estudos do Museu 1map. Zool6gico da Universidade de Coimbra, No. 285 , 34 pp. THEILER, GERTRUD & ROBINSON, BRITHA N., 1953a. Zoologi­ SANTOS, DIAS, J. A. T., 1985. Previous note concerning the orga­ cal Survey of the Union of South Africa. Tick Survey: Part VII. nization of a catalogue for the genus Aponomma Neumann, Distribution of Haemaphysalis leachi, the yellow dog tick. 1899 (Acarina-Ixodoidea). Garcia de Orta, Sir. Zoologia, Lis­ Onderstepoort Journal of Veterinary Research, 26, 83--91 + 1 boa, 12, 31-42. map. SIGWART, H., 1915. Beitrag zur Zeckenkenntnis von Deutsch­ THEILER, GERTRUD & ROBINSON , BRITHA N., 1953b. Ticks in Siidwestafrika, unter Besonderer Beriicksichtigung der Funde the South African Zoological Survey Collection. Part VII. Six in den Bezirken Outjo und Waterberg. Zeitschrift fiir Infek­ lesser known African rhipicephalids. Onderstepoort Journal of tionskrankheiten, Parasitiire Krankheiten und Hygiene der Veterinary Research, 26,93--136 + 1 map. Haustiere, 16, 434-444. THEILER, GERTRUD, 1956. Zoological Survey of the Union of SOUSA DIAS, V. A. , 1950. Subsidios para o estudo dos ixodideos South Africa. Tick Survey, Part IX. The distribution of the de Angola. Pecwiria, Loanda (1947-1948), 2, 127-280. three South African Hyalommas or bontpoots. Onderstepoort SPICKETT, A. M. & HEYNE, HELOISE, 1988. A survey of Karoo Journal of Veterinary Research, 27, 239-269 + 3 maps. tick paralysis in South Africa. Onderstepoort Journal of Veteri­ THEILER, GERTRUD & SALISBURY, LOIS, E., 1958. Zoological nary Research, 55,89-92. Survey of the Union of South Africa. Tick Survey: Part X. SWANEPOEL, A., 1959. Tick paralysis: regional neurological Distribution of Margaropus winthemi, the winter horse tick. involvement caused by Hyalomma truncatum. South African Onderstepoort Journal of Veterinary Research, 27, 599--{)()4 + 1 Medica/Journal, 33 , 909--911. map. SWANEPOEL, R ., SHEPHERD, A . J. , LEMAN , P. A ., SHEPHERD, THEILER, GERTRUD & SALISBURY, LOIS, E ., 1959. Ticks in the S. P., MCGILLIVRAY, G. M. , ERASMUS , M. J., SEARLE, L.A. South African Zoological Survey Collection. Part IX. "The & GILL, D . E ., 1987. Epidemiologic and clinical features of Amblyomma marmoreum group". Onderstepoort Journal of Crimean-Congo hemorrhagic fever in southern Africa. Ameri­ Veterinary Research, 28, 47-124 + 1 map. can Journal of Tropical Medicine and Hygiene, 36, 120-132. THEILER, GERTRUD, 1961. A contribution to the knowledge of THEILER, A ., 1911. Diseases, ticks and their eradication. Agricul­ African Ixodidae. The genus Rhipicentor. Revue de Zoologie et tural Journal of the Union of South Africa, 1, 491-508. de Botanique Africaines, 66,297-308. THEILER, GERTRUD , 1941. Ticks in the South African Zoological THEILER, GERTRUD, 1962. The Ixodoidea parasites of verte­ Survey Collection, Part 1. Ixodes alluaudi Neumann, 1913, a brates in Africa south of the Sahara (Ethiopian region). Project primitive tick parasitic on shrews. Onderstepoort Journal of S 9958. Report to the Director of Veterinary Services, Veterinary Science and Animal Industry, 17, 51--{i(). Onderstepoort. 260 pp. Mimeographed. THEILER, GERTRUD, 1943a. Ticks in the South African Zoologi­ THEILER, GERTRUD, 1975. Past-workers on tick and tick-borne cal Survey Collection. Part II. Onderstepoort Journal of diseases in southern Africa. Journal of the South African Veteri­ Veterinary Science and Animal Industry, 18, 85-89. nary Association, 46,303-310. THEILER, GERTRUD, 1943b. Notes on the ticks off domestic stock TROMMSDORFF, -., 1914. Beitrag zur Kenntnis der in Deutsch from Portuguese East Africa. Esta~ao Anti-Malarica de Siidwestafrika vorkommenden Zeckenarten. Beihefte zum Louren~ Marques. Archiv fiir Schiffs- und Tropenhygiene, 18 (7), 731-747. THEILER, GERTRUD, 1945a. Ticks in the South African Zoologi­ VANDER BORGHT-ELBL, ALENA, 1977. Ixodid ticks (Acarina, cal Survey Collection. Part III. The ornate Aponommas. Ixodidae) of Central Africa. V. The larval and nymphal stages Onderstepoort Journal of Veterinary Science and Animal Indus­ of the more important species of the genus Amblyomma Koch, try, 20, 165-178. 1844. Annales du Musee Royal de /'Afrique Centrale, Serie in THEILER, GERTRUD , 1945b. Ticks in the South African Zoologi­ Bvo, Sciences Zoologiques, No. 222, xi + 158 pp. cal Survey Collection. Part IV. The inornate Aponommas. WALKER, JANE B. & LAURENCE, B. R ., 1973. Margaropus wileyi Onderstepoort Journal of Veterinary Science and Animal Indus­ sp. nov. (Ixodoidea, Ixodidae), a new species of tick from the try, 20, 179--190. reticulated giraffe. Onderstepoort Journal of Veterinary THEILER, GERTRUD, 1945c. Ticks in the South African Zoologi­ Research, 40, 13--22. cal Survey Collection. Part V. Three African haemaphysalids WALKER, JANE B., 1974. The ixodid ticks of Kenya. A review of parasitic on domestic stock. Onderstepoort Journal of Veterin­ present knowledge of their hosts and distribution. London: ary Science and Animal Industry, 20, 191-207. Commonwealth Institute of Entomology. THEILER, GERTRUD, 1948. Zoological Survey of the Union of WALKER, JANE B. , MEHLITZ, D . & JONES , G. E ., 1978. Notes South Africa. Tick Survey-Part 1. 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104 JANE B. WALKER

WALKER, JANE B., NORVAL, R. A. I. & CORWIN, M.D., 1981. physalis toxopei, Amblyomma robinsoni and A . dammermani, Rhipicephalus zambeziensis sp. nov., a new tick from eastern with a note on the ornate nymph of A. latum. Parasitology, 19, and southern Africa, together with a redescrip~ion of R~ipice­ 405-410 +plate XXVII. phalus appendiculatus Neumann, 1901 (Acanna, Ixodidae). WOUTERS, G., 1989. Hybridization model for R~ipicephal'!s ap­ Onderstepoort Journal of Veterinary Research, 48, 87-104. pendiculatus and R. zambeziensis by glucose-p-Isomerase Isoen­ WALKER, JANE B. & SCHULZ, K. C. A., 1984. Records of the zymes. Onderstepoort Journal of Veterinary Research, 56, boot tick, Amblyomma hebraeum, from the angulate tortoise, 235-238. Chersina angulata, and the leopard tortoise, Geochelone parda­ WOUTERS, G., BERKVENS, D. & GOMES, A. F. 1989. Genetic lis. Onderstepoort Journal of Veterinary Research, 51, 171-173. variation in the body weight of ticks of the Rhipicep~alus WALKER, JANE B. & 0LWAGE, A., 1987. The tick vectors of appendiculatus/zambeziensis complex. Medical and Vetermary Cowdria ruminantium (Ixodoidea, Ixodidae, genus Entomology, 3, 423-427. Amblyomma) and their distribution. Onderstepoon Journal of YEOMAN G . H . & WALKER, JANE B., 1%7. The ixodid ticks of Veterinary Research, 54,353-379. Tanzania. A study of the zoogeography of the Ixodidae of an WALKER, JANE B., KEIRANS, J. E., PEGRAM, R. G . & CLIF­ East African country. London: Commonwealth Institute of En­ FORD, C. M., 1988. Clarification of the status of Rhipicephalus tomology. tricuspis Donitz, 1906 and Rhipicephalus lunulatus Neumann, YOUNG, E., 1965. Aponomma exornatum (Koch) as a cause of 1907 (Ixodoidea, Ixodidae). Systematic Parasitology, 12, mortality among monitors. Journal of the South African Veteri­ 159-186. nary Medica/Association, 36,259. WALKER, JANE B., 1990. Two new species of ticks from southern YOUNG, E., ZUMPT, F., BOOMKER, J., PENZHORN, B. L. & Africa whose adults parasitize the feet of ungulates: Rhipice­ ERASMUS, B., 1973. Parasites and diseases of Cape mountain phalus lounsburyi n. sp. and Rhipicephalus neumanni n. sp. zebra, black wildebeest, mountain reedbuck and blesbok in the (Ixodoidea, Ixodidae). Onderstepoort Journal of Veterinary Mountain Zebra National Park. Koedoe, 16, 77-81. Research, 57,57-75. ZIVKOVIC, D., PEGRAM, R. G ., JONGEJAN , F. & MWASE , E . T ., WARBURTON, C., 1910. On two collections of Indian ticks. Para­ 1986. Biology of Rhipicephalus appendiculatus and R. zambe­ sitology, 3, 395-407. ziensis and production of a fertile hybrid under laboratory con­ WARBURTON, C., 1912. Notes on the genus Rhipicephalus, with ditions. Experimental and Applied Acarology, 2, 285-298. the description of new species, and the consideration of some ZUMPT, F., 1949. Preliminary study to a revision of the genus species hitherto described. Parasitology, 5, 1-20. Rhipicephalus Koch. Key to the adult ticks of the genus Rhipi­ WARBURTON, C., 1927. On five new species of ticks (Arachnida, cephauls and description of two new species. Mor;ambique, No. Ixodoidea), Ornithodorus nattereri, Ixodes theodori, Haema- 60, 57-123.

105