Horak_69-74.indd CONTEXT The most recently compiled world list of valid tick names contains the names of 692 ixodid and 186 argasid tick species as well as that of Nuttalliella namaqua, the only species in this genus (Nava, Guglielmone & Mangold 2008). Of these 879 spe- cies, 80 ixodid and 25 argasid species and N. na- maqua occur in South Africa. Twenty-one of the 80 ixodid species and two of the argasid species have distributions confined to this country (Theiler 1962; Walker 1991; Walker, Keirans & Horak 2000), and N. namaqua has only been recorded in South Africa and Namibia. Forty-six of the 80 ixodid and 16 of the 25 argasid species and N. namaqua have been de- scribed since 1908, and this review is confined to the latter 63 ticks. Twenty-four of these descriptions were written, or contributed to by people in South Africa, and the remainder by researchers in England, France, Germany, Israel, Russia, Mozambique and the USA. The South African authors and the South African ticks that they have described as being new to sci- ence since 1908 are summarized in tabular format. A similar format has been followed for tick species with South African distributions, but described by foreign researchers during the same period. The earliest description of a tick that has a strictly South African distribution is that by De Geer in 1778 for Amblyomma sylvaticum, a tick of tortoises and more particularly the angulate tortoise, Chersina angulata (Horak, McKay, Henen, Heyne, Hofmeyr & De Villiers 2006). The most recent description is that of Haemaphysalis colesbergensis, a tick of do- mestic and wild felids (Apanaskevich & Horak 2008c). South Africans have in recent times also been in- volved in some of the more controversial name changes of well-known ticks of veterinary impor- tance. The name changes of the common blue tick, Boophilus decoloratus and the Asiatic blue tick Boophilus microplus to Rhipicephalus (Boophilus) decoloratus and Rhipicephalus (Boophilus) micro- plus were first published by Horak, Camicas & Keirans (2002) in their world list of valid tick names. 69 Onderstepoort Journal of Veterinary Research, 76:69–74 (2009) A century of tick taxonomy in South Africa I.G. HORAK Department of Veterinary Tropical Diseases, Faculty of Veterinary Science, University of Pretoria Onderstepoort, 0110 South Africa ABSTRACT HORAK, I.G. 2009. A century of tick taxonomy in South Africa. Onderstepoort Journal of Veterinary Re search, 76:69–74 Eighty ixodid tick species, 25 argasid tick species and Nuttalliella namaqua occur in South Africa. Twenty-one of the 80 ixodid species and two of the argasid species occur only in this country, while N. namaqua is present only in South Africa and Namibia. Forty-six of the 80 ixodid species and 16 of the 25 argasid species as well as N. namaqua have been described as new species since 1908. People working in South Africa have written or contributed to the descriptions of 24 of these 63 new species, while foreign researchers have described the remainder. New species indigenous to South Africa are still being discovered, while the names of some species, well known because of their vet- erinary importance, have been altered. 70 A century of tick taxonomy in South Africa TABLE 1 Authors who have worked in South Africa and the South African tick species that they have described as new to science since 1908 South African author and species South African author and species Bedford, G.A.H. (1912–1938) Walker, Jane (1966–2000) Argas peringueyi (Bedford & Hewitt 1925) Rhipicephalus theileri Bedford & Hewitt, 1925 Haemaphysalis cooleyi Bedford, 1929 Ixodes elongatus Bedford, 1929 Nuttalliella namaqua Bedford, 1931 Argas striatus Bedford, 1932 Rhipicephalus distinctus Bedford, 1932 Haemaphysalis hyracophila Hoogstraal, Walker & Neitz, 1971 Argas africolumbae Hoogstraal, Kaiser, Walker, Ledger, Converse & Rice, 1975 Ixodes neitzi Clifford, Walker & Keirans, 1977 Rhipicephalus zambeziensis Walker, Norval & Corwin, 1981 Ixodes catherinei Keirans, Clifford & Walker, 1982 Ixodes corwini Keirans, Clifford & Walker, 1982 Rhipicephalus lounsburyi Walker, 1990 Rhipicephalus neumanni Walker, 1990 Rhipicephalus exophthalmos Keirans & Walker, 1993 Rhipicephalus oreotragi Walker & Horak, 2000 Rhipicephalus warburtoni Walker & Horak, 2000 Du Toit, R. (1929–1964) Rhipicephalus glabroscutatum Du Toit, 1941 Theiler, Gertrud (1940–1983) Argas eboris (Theiler, 1959) Ixodes drakensbergensis Clifford, Theiler & Baker, 1975 Rhipicephalus arnoldi Theiler & Zumpt, 1949 Horak I.G. (1974–present) Haemaphysalis colesbergensis Apanaskevich & Horak, 2008 Zumpt, F. (SA Institute of Medical Research) (1948–1980) Rhipicephalus muehlensi Zumpt, 1943* * Zumpt was working at the Bernard Nocht-Institut in Hamburg at the time he described this species TABLE 2 Tick species with South African distributions described as new to science by foreign researchers since 1908 Tick species and foreign authors* Tick species and foreign authors* Ixodid ticks Amblyomma nuttalli Dönitz, 1909 Haemaphysalis aciculifer Warburton, 1913 Haemaphysalis hoodi Warburton & Nuttall, 1909 Haemaphysalis pedetes Hoogstraal, 1972 Haemaphysalis silacea Robinson, 1912 Haemaphysalis zumpti Hoogstraal & El Kammah, 1974 Ixodes alluaudi Neumann, 1913 Ixodes aulacodi Arthur, 1956 Ixodes bakeri Arthur & Clifford, 1961 Ixodes bedfordi Arthur, 1959 Ixodes cavipalpus Nuttall & Warburton, 1908 Ixodes myotomys Clifford & Hoogstraal, 1970 Ixodes procaviae Arthur & Burrow, 1957 Ixodes rhabdomysae Arthur, 1959 Ixodes spiculae Arthur, 1956 Ixodes spinae Arthur, 1958 Ixodes theilerae Arthur, 1953 Ixodes transvaalensis Clifford & Hoogstraal, 1966 Rhipicentor bicornis Nuttall & Warburton, 1908 Rhipicentor nuttalli Cooper & Robinson, 1908 Rhipicephalus evertsi mimeticus Dönitz, 1910 Rhipicephalus follis Dönitz, 1910 Rhipicephalus gertrudae Feldman-Muhsam, 1960 Rhipicephalus simpsoni Nuttall, 1910 Rhipicephalus sulcatus Neumann, 1908 Rhipicephalus turanicus Pomerantsev, 1936 Rhipicephalus zumpti Santos Dias, 1950 Argasid ticks Argas arboreus Kaiser, Hoogstraal & Kohls, 1964 Argas streptopelia Kaiser, Hoogstraal & Horner, 1970 Argas theilerae Hoogstraal & Kaiser, 1970 Argas walkerae Kaiser & Hoogstraal, 1969 Argas zumpti Hoogstraal, Kaiser & Kohls, 1968 Carios boueti (Roubad, & Colas-Belcour, 1933) Carios confusus (Hoogstraal, 1955) Carios cordiformis (Hoogstraal & Kohls, 1967) Carios faini (Hoogstraal, 1960) Ornithodoros compactus Walton, 1962 Ornithodoros porcinus Walton, 1962 Ornithodoros zumpti Heisch & Guggisberg, 1953 * Not listed in references 71 I.G. HORAK TABLE 3 South African ticks, other than Rhipicephalus species, described by foreign researchers, for which additional descriptions have been produced by researchers in South Africa since 1908 Species and authors* Descriptions by researchers working in South Africa** Amblyomma hebraeum Koch, 1844 Amblyomma marmoreum Koch, 1844 Amblyomma nuttalli Dönitz, 1909 Amblyomma sylvaticum (De Geer, 1778) Amblyomma tholloni Neumann, 1899 Aponomma exornatum (Koch, 1844) Aponomma latum (Koch, 1844) Aponomma transversale (Lucas, 1844) Dermacentor rhinocerinus (Denny, 1843) Haemaphysalis aciculifer Warburton, 1913 Haemaphysalis elliptica (Koch, 1844) Haemaphysalis parmata Neumann, 1905 Haemaphysalis silacea Robinson, 1912 Hyalomma glabrum Delpy, 1949 Hyalomma rufipes Koch, 1844 Hyalomma truncatum Koch, 1844 Ixodes alluaudi Neumann, 1913 Ixodes bakeri Arthur & Clifford, 1961 Margaropus winthemi Karsch, 1879 Rhipicentor bicornis Nuttall & Warburton, 1908 Rhipicentor nuttalli Cooper & Robinson, 1908 Walker & Olwage (1987) Theiler & Salisbury (1959); Walker & Olwage (1987) Theiler & Salisbury (1959) Theiler (1943a) Walker & Olwage (1987) Theiler (1945a) Theiler (1945b) Theiler (1945b) Keirans (1993) Theiler (1945c) Apanaskevich, Horak & Camicas (2007) Theiler (1945c) Theiler (1945c) Apanaskevich & Horak (2006) Apanaskevich & Horak (2008a) Apanaskevich & Horak (2008b) Theiler (1941) Clifford, Walker & Keirans (1976) Walker & Laurence (1973); Gothe (1967) Theiler (1961) Theiler (1961) * Not listed in references ** Listed in references TABLE 4 South African Rhipicephalus species described by foreign researchers, for which additional descriptions have been pro- duced since 1908 by researchers in South Africa Rhipicephalus species and original authors* Descriptions by researchers in South Africa** Rhipicephalus (Boophilus) decoloratus (Koch, 1844) Rhipicephalus (Boophilus) microplus (Canestrini, 1887) Rhipicephalus appendiculatus Neumann, 1901 Rhipicephalus capensis Koch, 1844 Rhipicephalus evertsi evertsi Neumann, 1897 Rhipicephalus follis Dönitz, 1910 Rhipicephalus gertrudae Feldman-Muhsam, 1960 Rhipicephalus kochi Dönitz, 1905 Rhipicephalus lunulatus Neumann, 1907 Rhipicephalus maculatus Neumann, 1901 Rhipicephalus muehlensi Zumpt, 1943 Rhipicephalus nitens Neumann, 1904 Rhipicephalus oculatus Neumann, 1901 Rhipicephalus sanguineus (Latreille, 1806) Rhipicephalus simpsoni Nuttall, 1910 Rhipicephalus simus Koch, 1844 Rhipicephalus sulcatus Neumann, 1908 Rhipicephalus tricuspis Dönitz, 1906 Rhipicephalus turanicus Pomerantsev, 1936 Rhipicephalus zumpti Santos Dias, 1950 Gothe (1967); Arthur & Londt (1973); Heyne (1986) Gothe (1967); Londt & Arthur (1975); Heyne (1986) Walker, Norval & Corwin (1981) Walker, Keirans & Horak (2000) Walker et al. (2000) Walker et al. (2000) Walker et al. (2000) Walker et al. (2000) Walker, Keirans, Pegram & Clifford (1988); Walker et al. (2000) Theiler & Robinson (1953b); Walker et al. (2000) Salisbury (1959); Walker et al. (2000) Walker et al. (2000) Keirans, Walker, Horak & Heyne (1993); Walker et al. (2000) Pegram, Keirans, Clifford & Walker (1987b); Walker et al. (2000) Theiler 1949; Walker et al. 2000 Pegram, Walker, Clifford & Keirans (1987c); Walker et al. (2000) Pegram, Clifford, Walker & Keirans (1987a); Walker et al. (2000) Walker et al. (1988; 2000) Pegram et al. (1987a); Walker et al. (2000) Walker et al. (2000) * Not listed in references ** Listed in references 72 A century of tick taxonomy in South Africa This was done after molecular and morphological studies by Murrell, Campbell & Barker (2000) and Beati & Keirans (2001) had indicated that the genus Rhipicephalus was paraphyletic in respect of the genus Boophilus. Murrell & Barker (2003) later con- firmed these name changes. The yellow dog tick of South Africa, previously lumped with Haemaphysalis leachi of North and East Africa, has been reinstated as Haemaphysalis elliptica, an old taxon originally described from the Cape of Good Hope by Koch in 1844 (Apanaskevich, Horak & Camicas 2007). While the bont-legged ticks, to which sub-specific status within the Hyalomma marginatum species group had previously been assigned, have all been raised to specific status (Apanaskevich & Horak 2008a). Furthermore the tick previously known as Hyalomma marginatum turanicum in South Africa has been re- instated as Hyalomma glabrum, the only Hyalomma with a strictly southern hemisphere, and with that a strictly South African distribution (Apanaskevich & Horak 2006). Three of the abovementioned ticks are of consider- able veterinary importance in that R. (B.) decoloratus and R. (B.) microplus are the vectors of Babesia species, the cause of babesiosis or redwater in do- mestic cattle (De Vos, De Waal & Jackson 2004), while H. elliptica, and not H. leachi, is now the only proven vector of Babesia canis rossi the cause of the virulent form of babesiosis encountered in do- mestic dogs in South Africa (Lewis, Penzhorn, Lopez-Rebollar & De Waal 1996; Apanaskevich et al. 2007). A fourth member of this group, Hyalomma rufipes, is the principal vector in South Africa of the virus causing Crimean-Congo haemorrhagic fever in humans (Horak, Swanepoel & Gummow 2002). The name changes are thus not only of academic importance, but of scientific concern as they are oft used in the literature concerning the diseases that they transmit and in which the correct use of taxo- nomic epithets is essential. ACKNOWLEDGEMENTS I have been fortunate in having two monographs to consult, namely those by Theiler (1962) and Walker (1991), and I pay tribute to both authors and to their attention to detail. REFERENCES All authors mentioned in the descriptive text have been listed, but only South African authors or foreign authors that worked in South Africa at the time at which they de- scribed particular tick species have been included in the list of references. APANASKEVICH, D.A. & HORAK, I.G. 2006. The genus Hya- lomma Koch, 1844. I. Reinstatement of Hyalomma (Euhya- lomma) glabrum Delpy, 1949 (Acari, Ixodidae) as a valid species with a redescription of the adults, the first description of its immature stages and notes on its biology. Onderstepoort Journal of Veterinary Research, 73:1–12. APANASKEVICH, D.A., HORAK, I.G. & CAMICAS, J-L. 2007. Redescription of Haemaphysalis (Rhipistoma) elliptica (Koch, 1844), an old taxon of the Haemaphysalis (Rhipistoma) leachi group from East and southern Africa, and of Haem a- phy salis (Rhipistoma) leachi (Audouin, 1826) (Ixodida, Ixo- didae). Onderstepoort Journal of Veterinary Research, 74: 181–207. APANASKEVICH, D.A. & HORAK, I.G. 2008a. The genus Hy a- lomma Koch, 1844. V. Re-evaluation of the taxonomic rank of taxa comprising the H. (Euhyalomma) marginatum group of species (Acari: Ixodidae) with redescription of all parasitic stages and notes on biology. International Journal of Acarol- ogy, 34:13–42. APANASKEVICH, D.A. & HORAK, I.G. 2008b. The genus Hya- lomma. VI. Systematics of H. (Euhyalomma) truncatum and the closely related species, H. (E.) albiparmatum and H. (E.) nitidum (Acari, Ixodidae). Experimental and Applied Acarol- ogy, 44:115–136. APANASKEVICH, D.A. & HORAK, I.G. 2008c. Two new species of African Haemaphysalis ticks (Acari: Ixodidae), carnivore parasites of the H. (Rhipistoma) leachi group. Journal of Par- a sitology, 94:594–607. ARTHUR, D.R & LONDT, J.G.H. 1973. The parasitic life cycle of Boophilus decoloratus (Koch, 1844). Journal of the Ento mo- logical Society of Southern Africa, 36:87–116. BEATI, L. & KEIRANS, J.E. 2001. Analysis of the systematic re- lationships among ticks of the genera Rhipicephalus and Boophilus (Acari: Ixodidae) based on mitochondrial 12S ri- bosomal DNA gene sequences and morphological charac- ters. Journal of Parasitology, 87:32–48. BEDFORD, G.A.H. & HEWITT, J. 1925. Descriptions of two new species of ticks, with notes on rare and hitherto unrecorded species from South Africa. South African Journal of Natural History, 5:259–266. BEDFORD, G.A.H. 1929. Notes on some South African ticks, with descriptions of three new species. Annual Report of the Director of Veterinary Services, Union of South Africa, 15: 493–499. BEDFORD, G.A.H. 1931. Nuttalliella namaqua, a new genus and species of tick. Parasitology, 23:230–232. BEDFORD, G.A.H. 1932. Description of Argas striatus, a new species of tick. 18th Report of the Director of Veterinary Serv- ices and Animal Industry, Union of South Africa, 18:221–222. BEDFORD, G.A.H. 1932. A synoptic check-list and host-list of the ectoparasites found on South African Mammalia, Aves and Reptilia (2nd ed.). 18th Report of the Director of Veterinary Services and Animal Industry, Union of South Africa, 18:223– 523. CLIFFORD, C.M., THEILER, GERTRUD & BAKER, MAUREEN, 1975. Ixodes (Afrixodes) drakensbergensis n. sp. from do- mestic and wild animals in Natal, Republic of South Africa. Onderstepoort Journal of Veterinary Research, 42:33–44. CLIFFORD, C.M., WALKER, JANE B. & KEIRANS, J.E. 1976. Ixodes (Afrixodes) bakeri Arthur & Clifford, 1961 (Acarina: Ixodidae): descriptions of the male and immature stages from rodents and insectivores and notes on its biology in South Africa. Onderstepoort Journal of Veterinary Research, 43:105–112. 73 I.G. HORAK CLIFFORD, C.M., WALKER, JANE B. & KEIRANS, J.E. 1977. Ixodes (Afrixodes) neitzi, n. sp. (Acarina: Ixodidae) from the mountain reedbuck in South Africa. Onderstepoort Journal of Veterinary Research, 44:143–150. DE VOS, A.J., DE WAAL, D.T. & JACKSON, L.A. 2004. Bovine babesiosis, in Infectious diseases of livestock, edited by J.A.W. Coetzer & R.C. Tustin. Cape Town: Oxford University Press. DU TOIT, R. 1941. Description of a tick Rhipicephalus glabroscu- tatum, sp. nov. (Ixodidae), from the Karroo areas of the Union of South Africa. Onderstepoort Journal of Veterinary Science and Animal Industry, 16:115–118. GOTHE, R. 1967. Ticks in the South African Zoological Survey Collection. Part XII. The genera Boophilus Curtice, 1891 and Margaropus Karsch, 1879. Onderstepoort Journal of Veter- inary Research, 34:81–107. HEYNE, HELOISE 1986. Differentiation of Boophilus decoloratus and Boophilus microplus. Journal of the South African Veter- inary Association, 57:251–252. HOOGSTRAAL, H., WALKER, JANE B. & NEITZ, W.O. 1971. Notes on African Haemaphysalis ticks. VII. H. (Rhipistoma) hyracophila sp. nov. (Ixodoidea: Ixodidae), a parasite of hy- raxes in South Africa. Journal of Parasitology, 57:417–425. HOOGSTRAAL, H., KAISER, M.N., WALKER, JANE B., LEDG- ER, J.A., CONVERSE, J.D. & RICE, R.C.A. 1975. Obser va- tions on the subgenus Argas (Ixodoidea: Argasidae: Argas). 10. A. (A.) africolumbae, n. sp., a Pretoria virus-infected par- asite of birds in southern and eastern Africa. Journal of Med- ical Entomology, 12:194–201. HORAK, I.G., CAMICAS, J.-L. & KEIRANS, J.E. 2002. The Ixo- didae, Amblyommidae and Nuttalliellidae (Acari: Ixodida): a world list of valid tick names. Experimental and Applied Acar- ology, 28:7–54. HORAK I.G., SWANEPOEL R. & GUMMOW, B. 2002. The distri- bution of Hyalomma spp. and human cases of Crimean- Congo haemorrhagic fever in South Africa, in Proceedings of the 10th Conference of the Association of Institutions for Tropical Veterinary Medicine, Copenhagen, Denmark 20–23 August 2001: 501–509. HORAK, I.G., MCKAY, I.J., HENEN, B.T., HEYNE, HELOISE, HOFMEYR, MARGARETHA D. & DE VILLIERS, A.L. 2006. Parasites of domestic and wild animals in South Africa. XLVII. Ticks of tortoises and other reptiles. Onderstepoort Journal of Veterinary Research, 73:215–227. KEIRANS, J.E., CLIFFORD, C.M. & WALKER, JANE B. 1982. The Ixodes (Afrixodes) oldi group (Acari: Ixodidae) from sub- Saharan Africa with descriptions of five new species. Journal of Medical Entomology, 19:309–329. KEIRANS, J.E. 1993. Dermacentor rhinocerinus (Denny, 1843) (Acari: Ixodida: Ixodidae): rediscription of the male, female and nymph and first description of the larva. Onderstepoort Journal of Veterinary Research, 60:59–68. KEIRANS, J.E. WALKER, JANE B., HORAK, I.G. & HEYNE, HELOISE 1993. Rhipicephalus exophthalmos sp. nov., a new tick species from southern Africa, and redescription of Rhip- icephalus oculatus Neumann, 1901, with which it has hitherto been confused (Acari: Ixodida: Ixodidae). Onderstepoort Journal of Veterinary Research, 60:229–246. KEIRANS, J.E. & WALKER, JANE B. 1993. Rhipicephalus exoph- thalmos sp. nov., in KEIRANS, J.E., WALK ER, JANE B. & HORAK, I.G. 1993. Rhipicephalus exophthalmos sp. nov., a new tick species from southern Africa, and redescription of Rhipicephalus oculatus Neumann, 1901, with which it has hitherto been confused (Acari: Ixodida: Ixodidae). Onder ste- poort Journal of Veterinary Research, 60:229–246. LEWIS, B.D., PENZHORN, B.L., LOPEZ-REBOLLAR, L.M. & DE WAAL, D.T. 1996. Isolation of a South African vector-specific strain of Babesia canis. Veterinary Parasitology, 63:9–16. LONDT, J.G.H. & ARTHUR, D.R. 1975. The structure and para- sitic life cycle of Boophilus microplus (Canestrini, 1888) in South Africa (Acarina: Ixodidae). Journal of the Entomological Society of Southern Africa, 38:321–340. MURRELL, A.N., CAMPBELL, J.H. & BARKER, S.C. 2000. Phylogenetic analysis of the rhipicephaline ticks indicates that the genus Rhipicephalus is paraphyletic. Molecular Phylogenetics and Evolution, 16:1–7. MURRELL, A. & BARKER, S.C. 2003. Synonomy of Boophilus Curtice, 1981 with Rhipicephalus Koch, 1844 (Acari: Ixo di- dae). Systematic Parasitology, 56:169–172. NAVA, S., GUGLIELMONE, A.A. & MANGOLD, A.J. 2008. An overview of the systematics and evolution of ticks. Frontiers in Bioscience, in press. PEGRAM, R.G., CLIFFORD, C.M., WALKER, JANE B. & KEI- RANS, J.E. 1987a. Clarification of the Rhipicephalus san- guineus group (Acari, Ixodoidea, Ixodidae). I. R. sulcatus Neu mann 1908 and R. turanicus Pomarantsev, 1936. Sys- tematic Parasitology, 10:3–26. PEGRAM, R.G., KEIRANS, J.E., CLIFFORD, C.M. & WALKER, JANE B. 1987b. Clarification of the Rhipicephalus san- guineus group (Acari, Ixodoidea, Ixodidae). II. R. sanguineus (Latreille, 1806) and related species. Systematic Parasitology, 10:27–44. PEGRAM, R.G., WALKER, JANE B., CLIFFORD, C.M. & KEI- RANS, J.E. 1987c. Comparisons of populations of the Rhip- icephalus simus group: R. simus, R. praetextatus and R. muhsamae (Acari: Ixodidae). Journal of Medical Entomology, 24:666–682. SALISBURY, LOIS E. 1959. Ticks in the South African Zoological Survey Collection. Part X. Rhipicephalus mühlensi Zumpt. Onderstepoort Journal of Veterinary Research, 28:125–132. THEILER, GERTRUD 1941. Ticks in the South African Zoological Survey Collection. Part I. Ixodes alluaudi Neumann, 1913, a primitive tick parasitic on shrews. Onderstepoort Journal of Veterinary Science and Animal Industry, 17:51–60. THEILER, GERTRUD 1943. Ticks in the South African Zoological Survey Collection. Part II. Onderstepoort Journal of Veterinary Science and Animal Industry, 18:85–89. THEILER, GERTRUD 1945a. Ticks in the South African Zo o- logical Survey Collection. Part III. The ornate Aponommas. Onderstepoort Journal of Veterinary Science and Animal Industry, 20:165–178. THEILER, GERTRUD 1945b. Ticks in the South African Zoo- logical Survey Collection. Part IV. The inornate Aponommas. Onderstepoort Journal of Veterinary Science and Animal Industry, 20:179–190. THEILER, GERTRUD 1945c. Ticks in the South African Zoo log- ical Survey Collection. Part V. Three African haemaphysalids parasitic on domestic stock. Onderstepoort Journal of Vet- erinary Science and Animal Industry, 20:191–207. THEILER, GERTRUD 1949. Ticks in the South African Zoological Survey Collection. Part VI. Little known African rhipicepha- lids. Onderstepoort Journal of Veterinary Science and Animal Industry, 21:253–300. THEILER, GERTRUD & ZUMPT, F. 1949. Description of new spe cies. Rhipicephalus (s. str.) arnoldi Theiler & Zumpt, n. sp., in ZUMPT, F. 1949. Preliminary study to a revision of the genus Rhipicephalus Koch. Key to the adult ticks of the genus 74 A century of tick taxonomy in South Africa Rhipicephalus Koch and description of two new species. Mo- çambique, 60:57–123. THEILER, GERTRUD & ROBINSON, BRITHA N. 1953. Ticks in the South African Zoological Survey Collection. Part VII. Six lesser known African rhipicephalids. Onderstepoort Journal of Veterinary Research, 26:93–136. THEILER, GERTRUD 1959. Ticks in the South African zoologi- cal survey collection—Part XI—Ornithodoros eboris n. sp. Onderstepoort Journal of Veterinary Research, 28:145–153. THEILER, GERTRUD & SALISBURY, LOIS E. 1959. Ticks in the South African Zoological Survey Collection. Part IX. “the Amblyomma marmoreum group”. Onderstepoort Journal of Veterinary Research, 28:47–124. THEILER, GERTRUD 1961. A contribution to the knowledge of African Ixodidae. The genus Rhipicentor. Revue de Zoologie et de Botanique Africaines, 66:297–308. THEILER, GERTRUD 1962. The Ixodoidea parasites of verte- brates in Africa south of the Sahara (Ethiopian region). Project S 9958. Report to the Director of Veterinary Services, Onderstepoort. Mimeographed. WALKER, JANE B. & LAURENCE, B.R. 1973. Margaropus wi- leyi sp. nov. (Ixodoidea, Ixodidae), a new species of tick from the reticulated giraffe. Onderstepoort Journal of Veterinary Research, 40:13–22. WALKER, JANE B., NORVAL, R.A.I. & CORWIN, M.D. 1981. Rhipicephalus zambeziensis sp. nov., a new tick from east- ern and southern Africa, together with a rediscription of Rhip- icephalus appendiculatus Neumann, 1901 (Acarina, Ixo di- dae). Onderstepoort Journal of Veterinary Research, 48: 7–104. WALKER, JANE B. & OLWAGE, A. 1987. The tick vectors of Cowdria ruminantium (Ixodoidea, Ixodidae, genus Ambly om- ma) and their distribution. Onderstepoort Journal of Vet er in- ary Research, 54: 353–379. WALKER, JANE B., KEIRANS, J.E., PEGRAM, R.G. & CLIF- FORD, C.M. 1988. Clarification of the status of Rhipicephalus tricuspis Dönitz, 1906 and Rhipicephalus lunulatus Neu- mann, 1907 (Ixodoidea, Ixodidae). Systematic Parasitology, 12:159–186. WALKER, JANE B. 1990. Two new species of ticks from south- ern Africa whose adults parasitize the feet of ungulates: Rhip- i cephalus lounsburyi n. sp. and Rhipicephalus neumanni n. sp. (Ixodoidea, Ixodidae). Onderstepoort Journal of Vet er i- nary Research, 57:57–75. WALKER, JANE B. 1991. A review of the ixodid ticks (Acari: Ixodidae) occurring in southern Africa. Onderstepoort Journal of Veterinary Research, 58:81–105. WALKER, JANE B., KEIRANS, J.E. & HORAK, I.G. 2000. The genus Rhipicephalus (Acari, Ixodidae): a guide to the brown ticks of the world. Cambridge: Cambridge University Press. WALKER, JANE B. & HORAK, I.G. 2000. Rhipicephalus oreotra- gi, sp. nov., in WALKER, JANE B., KEIRANS, J.E. & HORAK, I.G. 2000. The genus Rhipicephalus (Acari, Ixodidae): a guide to the brown ticks of the world. Cambridge: Cambridge Uni - versity Press. WALKER, JANE B. & HORAK, I.G. 2000. Rhipicephalus warbur- toni sp. nov., in WALKER, JANE B., KEIRANS, J.E. & HORAK, I.G. 2000. The genus Rhipicephalus (Acari, Ixo di dae): a guide to the brown ticks of the world. Cambridge: Cambridge Uni- versity Press. ZUMPT, F. 1943. Rhipicephalus aurantiacus Neumann und ähn- liche Arten. VIII. Vorstudie zu einer Revision der Gattung Rhip icephalus Koch. Zeitschrift für Parasitenkunde, 13:102– 117. << /ASCII85EncodePages false /AllowTransparency false /AutoPositionEPSFiles true /AutoRotatePages /None /Binding /Left /CalGrayProfile (Dot Gain 20%) /CalRGBProfile (sRGB IEC61966-2.1) /CalCMYKProfile (Coated FOGRA27 \050ISO 12647-2:2004\051) /sRGBProfile (sRGB IEC61966-2.1) /CannotEmbedFontPolicy /Error /CompatibilityLevel 1.4 /CompressObjects /Tags /CompressPages true /ConvertImagesToIndexed true /PassThroughJPEGImages true /CreateJobTicket false /DefaultRenderingIntent /Default /DetectBlends true /DetectCurves 0.0000 /ColorConversionStrategy /CMYK /DoThumbnails false /EmbedAllFonts true /EmbedOpenType false /ParseICCProfilesInComments true /EmbedJobOptions true /DSCReportingLevel 0 /EmitDSCWarnings false /EndPage -1 /ImageMemory 1048576 /LockDistillerParams false /MaxSubsetPct 100 /Optimize true /OPM 1 /ParseDSCComments true /ParseDSCCommentsForDocInfo true /PreserveCopyPage true /PreserveDICMYKValues true /PreserveEPSInfo true /PreserveFlatness true /PreserveHalftoneInfo false /PreserveOPIComments true /PreserveOverprintSettings true /StartPage 1 /SubsetFonts false /TransferFunctionInfo /Apply /UCRandBGInfo /Preserve /UsePrologue false /ColorSettingsFile () /AlwaysEmbed [ true ] /NeverEmbed [ true ] /AntiAliasColorImages false /CropColorImages true /ColorImageMinResolution 300 /ColorImageMinResolutionPolicy /OK /DownsampleColorImages false /ColorImageDownsampleType /Bicubic /ColorImageResolution 2400 /ColorImageDepth -1 /ColorImageMinDownsampleDepth 1 /ColorImageDownsampleThreshold 1.50000 /EncodeColorImages true /ColorImageFilter /DCTEncode /AutoFilterColorImages true /ColorImageAutoFilterStrategy /JPEG /ColorACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /ColorImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000ColorACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000ColorImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasGrayImages false /CropGrayImages true /GrayImageMinResolution 300 /GrayImageMinResolutionPolicy /OK /DownsampleGrayImages false /GrayImageDownsampleType /Bicubic /GrayImageResolution 2400 /GrayImageDepth -1 /GrayImageMinDownsampleDepth 2 /GrayImageDownsampleThreshold 1.50000 /EncodeGrayImages true /GrayImageFilter /DCTEncode /AutoFilterGrayImages true /GrayImageAutoFilterStrategy /JPEG /GrayACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /GrayImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000GrayACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000GrayImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasMonoImages false /CropMonoImages true /MonoImageMinResolution 1200 /MonoImageMinResolutionPolicy /OK /DownsampleMonoImages false /MonoImageDownsampleType /Bicubic /MonoImageResolution 2400 /MonoImageDepth -1 /MonoImageDownsampleThreshold 1.50000 /EncodeMonoImages true /MonoImageFilter /CCITTFaxEncode /MonoImageDict << /K -1 >> /AllowPSXObjects false /CheckCompliance [ /None ] /PDFX1aCheck false /PDFX3Check false /PDFXCompliantPDFOnly false /PDFXNoTrimBoxError true /PDFXTrimBoxToMediaBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXSetBleedBoxToMediaBox true /PDFXBleedBoxToTrimBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXOutputIntentProfile (None) /PDFXOutputConditionIdentifier () /PDFXOutputCondition () /PDFXRegistryName () /PDFXTrapped /False /CreateJDFFile false /Description << /ARA /BGR /CHS /CHT /CZE /DAN /DEU /ESP /ETI /FRA /GRE /HEB /HRV (Za stvaranje Adobe PDF dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. Stvoreni PDF dokumenti mogu se otvoriti Acrobat i Adobe Reader 5.0 i kasnijim verzijama.) /HUN /ITA /JPN /KOR /LTH /LVI /NLD (Gebruik deze instellingen om Adobe PDF-documenten te maken die zijn geoptimaliseerd voor prepress-afdrukken van hoge kwaliteit. De gemaakte PDF-documenten kunnen worden geopend met Acrobat en Adobe Reader 5.0 en hoger.) /NOR /POL /PTB /RUM /RUS /SKY /SLV /SUO /SVE /TUR /UKR /ENU (Use these settings to create Adobe PDF documents best suited for high-quality prepress printing. Created PDF documents can be opened with Acrobat and Adobe Reader 5.0 and later.) >> /Namespace [ (Adobe) (Common) (1.0) ] /OtherNamespaces [ << /AsReaderSpreads false /CropImagesToFrames true /ErrorControl /WarnAndContinue /FlattenerIgnoreSpreadOverrides false /IncludeGuidesGrids false /IncludeNonPrinting false /IncludeSlug false /Namespace [ (Adobe) (InDesign) (4.0) ] /OmitPlacedBitmaps false /OmitPlacedEPS false /OmitPlacedPDF false /SimulateOverprint /Legacy >> << /AddBleedMarks false /AddColorBars false /AddCropMarks false /AddPageInfo false /AddRegMarks false /ConvertColors /ConvertToCMYK /DestinationProfileName () /DestinationProfileSelector /DocumentCMYK /Downsample16BitImages true /FlattenerPreset << /PresetSelector /MediumResolution >> /FormElements false /GenerateStructure false /IncludeBookmarks false /IncludeHyperlinks false /IncludeInteractive false /IncludeLayers false /IncludeProfiles false /MultimediaHandling /UseObjectSettings /Namespace [ (Adobe) (CreativeSuite) (2.0) ] /PDFXOutputIntentProfileSelector /DocumentCMYK /PreserveEditing true /UntaggedCMYKHandling /LeaveUntagged /UntaggedRGBHandling /UseDocumentProfile /UseDocumentBleed false >> ] >> setdistillerparams << /HWResolution [2400 2400] /PageSize [850.394 1133.858] >> setpagedevice