- Infectious Diseases of Livestock
- Part 1
- Non-pathogenic Theileria species in cattle
- Classification, epidemiology and control of arthropod-borne viruses
- Vectors: Ticks
- GENERAL INTRODUCTION: COCCIDIA
- Vectors: Tsetse flies
- Vectors: Muscidae
- Vectors: Tabanidae
- Vectors: Culicoides spp.
- Vectors: Mosquitoes
- Special factors affecting the control of livestock diseases in sub-Saharan Africa
- The control of infectious diseases of livestock: Making appropriate decisions in different epidemiological and socioeconomic conditions
- Infectious diseases of animals in sub-Saharan Africa: The wildlife⁄livestock interface
- Vaccination: An approach to the control of infectious diseases
- African animal trypanosomoses
- Amoebic infections
- Equine protozoal myeloencephalitis
- GENERAL INTRODUCTION: BABESIOSES
- Bovine babesiosis
- Equine piroplasmosis
- Porcine babesiosis
- Ovine babesiosis
- GENERAL INTRODUCTION: THEILERIOSES OF CATTLE
- East Coast fever
- Corridor disease
- Zimbabwe theileriosis
- Turning sickness
- Theileria taurotragi infection
- Theileria mutans infection
- Theileria annulata theileriosis
- Theileriosis of sheep and goats
- Theileria buffeli⁄orientalis infection
- Non-pathogenic Theileria species in cattle
- GENERAL INTRODUCTION: RICKETTSIAL, CHLAMYDIAL AND HAEMOTROPIC MYCOPLASMAL DISEASES
- Lesser-known rickettsias infecting livestock
- Q fever
- Bovine Haemobartonellosis
- Potomac horse fever
- GENERAL INTRODUCTION: ANAPLASMOSES
- Bovine anaplasmosis
- Ovine and caprine anaplasmosis
Non-pathogenic Theileria species in cattle
This content is distributed under the following licence: Attribution-NonCommercial CC BY-NC View Creative Commons Licence details here
Non-pathogenic Theileria species in cattle
J A LAWRENCE - Extraordinary Professor, DPhil, BSc, MRCVS (ret.), DTVM, Department of Paraclinical Veterinary Science, University of Zimbabwe, Harare, Zimbabwe
C BYARUHANGA - Post-doc Fellow, BVM, MVPM, PhD, Department of Veterinary Tropical Diseases, Para-clinical Building, Faculty of Veterinary Science, University of Pretoria, 100 Old Soutpan Road, Onderstepoort, Pretoria, Gauteng, 0110, South Africa
M OOSTHUIZEN - Parasitology, PhD, Department of Veterinary Tropical Diseases, Faculty of Veterinary Science, Private Bag X04, University of Pretoria, Pretoria, Gauteng, 0110, South Africa
B J MANS - Principal Researcher, BSc, BSc (Hons) Biochemistry, MSc (Biochemistry), PhD (Biochemistry), Agricultural Research Council, Onderstepoort Veterinary Research, 100 Old Soutpan Road, Pretoria, Gauteng, 0110, South Africa
A number of species of Theileria occur in cattle without causing disease. Theileria taurotragi, T. mutans and T. buffeli/orientalis most commonly cause inapparent to mild diseases and only occasionally result in more severe disease. Other species that have been identified without ever having been implicated in disease in cattle are:
- Theileria taurotragi clade - Theileria sp.(bougasvlei), Theileria sp. (buffalo)
- Theileria velifera clade - Theileria velifera, Theileria velifera A, Theileria velifera B, Theileria sp. RR-2012
- Antelope clade - Theileria sp. (sable), Theileria sp. (sable-like) – infectivity for cattle doubtful
- Theileria buffeli clade - Theileria sinensis, Theileria sinensis-like, Theileria sp. RMP-2013
The T. taurotragi clade is mostly composed of pathogenic Theileria, namely T. annulata, T. lestoquardi, T. parva and T. taurotragi (see Introduction Theilerioses of cattle). However, two more species are found in this clade that have to date not been associated with clinical disease, namely T. sp. (buffalo) and T. sp. (bougasvlei) 21, 22 These genotypes are unique species in their own right based on 18S, Cytochrome oxidase I (COI) and S5 ribosomal gene analysis 21, 34 but they have not been given appropriate species names as their tick vectors remain unknown and their complete life cycles have not been elucidated.
Theileria velifera was first described 40 in Madagascar, under the name Haematoxenus veliferus, as a benign parasite of cattle characterized by the presence of a veil-like structure associated with intra-erythrocytic piroplasms (Figure 1). The parasite is widespread in eastern and southern Africa and also occurs in the Caribbean. It has no practical significance other than its role in confusing the diagnosis of Theileria parva. The parasite also infects African buffalo (Syncerus caffer). 36 Ancestral relationships have shown that the T. velifera clade is composed of four genotypes, T. velifera, T. velifera A, T. velifera B and Theileria sp. RR-2012, a yet undescribed genotype from cattle in India. 22 Whereas T. velifera B has only been reported in African buffalo, T. velifera and T. velifera A have been found in cattle and African buffalo. 7, 8, 21, 22
Theileria sp. (sable) causes mortality in naïve roan antelope (Hippotragus equinus) and sable antelope (Hippotragus niger) and presented as a lymphoproliferative condition in these antelope. 28 Transformed schizonts of T. sp. (sable) can be maintained in cell culture similarly to T. parva and T. sp. (buffalo). 50 Sable and roan antelope are considered the natural carrier host but identical genotypes based on the 18S rRNA genes have been found in dogs. 24 Reverse line blot has also indicated the extensive presence of T. sp. (sable) in cattle, African buffalo, blesbuck (Damaliscus pygargus), blue wildebeest (Connochaetes taurinus), klipspringer (Oreotragus oreotragus), reedbuck (Redunca arundinum) and nyala (Tragelaphus angasii). 1, 27, 28, 32, 47 A closely related genotype, T. sp. (sable-like) that differs by 6 base pairs in the V4 hypervariable region of the 18S rRNA gene has been described in a single ox in South Africa that presented with clinical signs of theileriosis, with high schizont parasitaemia and lymphoid infiltration of organs. 21 However, subsequent screening of cattle from South Africa using sequencing of the 18S rRNA gene failed to detect either T. sp. (sable) or T. sp. (sable-like). 23 Potential cross-reactivity with T. velifera in the reverse line blot probe has been considered as a possible explanation for this discrepancy.21Theileria sinensis was first described in cattle in China based on infectivity studies, morphological observations and its inability to be transmitted by Hyalomma detritum and Haemaphysalis longicornis, the vectors of T. annulata and T. sergenti respectively. 4, 5 It was assigned to the T. buffeli/orientalis clade after molecular analysis and was named T. sinensis after classic taxonomic and molecular studies. 15 Molecular analysis based on the ITS1, ITS2 and 5.8S rRNA genes clearly differentiate it from T. sergenti and other members of the T. buffeli/orientalis clade, supporting its designation as a unique species in this clade. 18 A genotype and a possible new species, T. sinensis-like, that is closely related to T. sinensis, has been described in African buffalo in South Africa. 21 Screening of cattle and African buffalo using next-generation sequencing of the 18S rRNA gene indicated that this species is only found in buffalo. 23 A number of as yet undescribed genotypes in the T. buffeli/orientalis clade also occur in cattle, Nilgai (Boselaphus tragocamelus) and Asian buffalo in China and India, all designated as T. sp. RMP-2013. 22
Aetiology and life cycle
Theileria velifera has most likely a typical theilerial life cycle (see East Coast fever: Figure 29.2) but no detailed studies have yet been done. The parasite is transmitted by ticks of the genus Amblyomma, namely A. variegatum, 42 A. hebraeum, A. lepidum and A. astrion. 41The development of T. velifera in the gut and haemolymph of A. variegatum is typical of the Theileria genus 45 and the piroplasms are similar to those of Theileria mutans except that some are associated with the presence of a veil-like structure in the cytoplasm of the erythrocyte (Figure 1). This structure is believed to be composed of a crystalline derivative of haemoglobin. 43 Schizonts have not been described. The piroplasms are readily transmitted by inoculation of infected blood. 40The life cycle, including the tick vectors, of Theileria sp. (buffalo) and T. sp. (bougasvlei) remains obscure, beyond the ability to transform schizonts. Levels of parasitaemia in African buffalo are similar to those observed for T. parva. 33, 34 Rhipicephalus appendiculatus and R. evertsi evertsi have been implicated as vectors of T. sp. (sable). 38 Theileria sinensis is transmitted by Haemaphysalis quinghaiensis and infects cattle and yaks (Bos grunniens). 46 Transmission by infected blood inoculation indicated that the parasites multiply prolifically in the erythrocytic stage. 5 Piroplasms are polymorphic with shapes that range from pyriform or circular to rod-shaped and some may exhibit bars and veils similar to other members of this clade.5 It is expected that its general life cycle will resemble other members of the T. buffeli/orientalis clade.
Little is known of the epidemiology of T. velifera. It is assumed to be widespread throughout the range of its tick vectors. Theileria velifera infections were detected in cattle in Uganda, 3, 7, 25, 29, 31 Kenya, 27 Nigeria 20 and South Africa, 21 and in African buffalo in Uganda, 30, 31 Botswana 12 and South Africa. 8 21 The prevalence of T. velifera may reach 100 per cent, depending on location and ecology, 3, 30, 31 The protective effect of T. velifera co-infection against T. parva infection is discussed Theileria mutans infection.
Theileria sp. (buffalo) has been detected in African buffalo in Botswana, Kenya, Mozambique, Uganda, South Africa and Zimbabwe.8, 12, 21, 23, 34 It was first distinguished from T. parva due to differences observed in antigenicity using monoclonal antibody panels 11 and was shown to have a unique 18S rRNA sequence. 2 Theileria sp. (buffalo) was successfully cultivated in cell culture as schizonts from an African buffalo that showed patent Theileria schizont and a 10 per cent piroplasm parasitaemia.49 Most studies that screened cattle for Theileria parasites found no evidence of this parasite, while African buffalo showed a high prevalence (50-100 per cent). 21, 23, 30, 34, 47 However, recently the presence of T. sp. (buffalo) was detected in 67 per cent of cattle grazing with buffalo in Marula, a private ranching farm in Nakuru, Rift Valley in Kenya. 13 It was also showed that T. sp. (buffalo) could transform cattle-derived lymphocytes. 6 Whether this parasite plays a role in clinical theileriosis in cattle has not yet been established, but results thus far suggest that it is not transmissible between cattle. 6 Theileria sp. (bougasvlei) has been detected in Botswana, Mozambique, Namibia, South Africa and Zimbabwe.23 Studies of the distribution of T. sp. (buffalo) and T. sp. (bougasvlei) demonstrated an apparent antagonistic behaviour, in that buffalo herds or farms are almost exclusively infected by either one or the other, even in regions of sympatry such as the Kruger National Park in South Africa. 23, 34 This was interpreted as potential hybrid sterility that allowed speciation while still using the same tick vector and carrier host. 34
Theileria sp. (sable) has been detected in a variety of hosts using reverse line blot in South Africa as indicated above. It has also been detected in Nigeria 1and Kenya. 27 Its tick vectors, R. appendiculatus and R. evertsi occur from southern Africa to East Africa,44 and mortality in roan and sable antelope has been recorded in Swaziland and Zimbabwe.
Theileria sinensis has been detected mainly in China in cattle and yaks. 16, 35 A study that focused on piroplasms in dogs in the Gansu province of China found 6.4 per cent of animals sampled positive for T. sinensis. 26Theileria sinensis or a very closely related genotype was also detected in Asian buffalo in the Amazon region of Brazil. 37
No clinical signs have been recorded for T. velifera in intact infected animals. Anaemia has been described in infected splenectomized calves.42
Pathogenesis and pathology
No studies have been undertaken on the pathogenesis or pathology of T. velifera infection. There is currently no evidence that T. sp. (buffalo), T. sp. (bougasvlei), Theileria sp. (sable) or Theileria sinensis are pathogenic in cattle.
Diagnosis and differential diagnosis
Real-time PCR assays specific for T. sp. (buffalo) and T. sp. (bougasvlei) have been developed. 34 These are useful to differentiate and confirm mixed infection with T. parva, to estimate parasitaemia in carrier animals and to determine the geographic distribution of these parasites. A real-time PCR assay that can differentiate Theileria sp. (sable) and T. sp. (sable-like) has been developed and is being validated. 22
Theileria velifera infection can be differentiated from other benign theilerioses if piroplasms associated with veil-like structures are found in erythrocytes. However, as yet, no serological test or species-specific monoclonal antibodies are available. The RLB hybridisation assay was developed to detect T. velifera and other Theileria species, using species-specific probes. This was based on the assumption that the 18S hyper-variable region is conserved within species. 44 However, significant similarity exists between the RLB probe for Theileria sp. (sable) and T. velifera, resulting in the reporting of a relatively high prevalence (40-50 per cent) of Theileria sp. (sable) in cattle and African buffalo 12, 28, 47 that may be attributable to the presence of T. velifera. 21 Further, the RLB assay does not detect variations in the 18S rRNA gene sequences of T. velifera, 9 and therefore, the assay has to be validated when screening novel vertebrate species or animals from different geographical locations.
An ELISA has been developed based on the recombinant major piroplasm surface protein of T. sinensis. 48 Significant cross-reactivity with positive sera for T. annulata and T. orientalis were found, making the use of this assay for epidemiological surveys limited where these species share similar geographical distributions. Conventional PCR assays capable of distinguishing T. sergenti and T. sinensis in cattle and yaks have been developed based on the major piroplasm surface protein. 17 A LAMP PCR assay was also developed for the same purpose based on ITS sequences. 18, 19 A PCR-RFLP assay that targets the RPS8 gene can distinguish between T. annulata, T. sinensis, T. sergenti, Babesia ovata, B. bovis, B. bigemina,and B. major and Babesia species Kashi isolate, the major piroplasms of cattle in China. 39
As T. velifera is a benign parasite, there are no indications for its treatment or control. The same applies for T. sp. (buffalo) and T. sp. (bougasvlei). In the case of T. sp. (sable), buparvaquone was successful in treatment of clinical theileriosis in roan and sable antelope. 38
- ADAMU, M., TROSKIE, M., OSHADU, D.O., MALATJI, D.P., PENZHORN, B.L., MATJILA, P.T., 2014. Occurrence of tick-transmitted pathogens in dogs in Jos, Plateau State, Nigeria. Parasites & Vectors, 7, 119.
- ALLSOPP, B.A., BAYLIS, H.A., ALLSOPP, M.T., CAVALIER-SMITH, T., BISHOP, R.P., CARRINGTON, D.M., SOHANPAL, B., SPOONER, P., 1993. Discrimination between six species of Theileria using oligonucleotide probes which detect small subunit ribosomal RNA sequences. Parasitology, 107, 157-165.
- ASIIMWE, B.B., WEIR, W., TAIT, A., LUBEGA, G.W. & OURA, C.A.L., 2013. Haemoparasite infection kinetics and the population structure of Theileria parva on a single farm in Uganda. Veterinary Parasitology,193, 8-14.
- BAI, Q., LIU, G.Y., HAN, G.F., 1995. An unidentified Theileria sp. infective to cattle in China (in Chinese). Chinese Journal of Veterinary Science, 15, 16–21.
- BAI, Q., LIU, G., HEN, G., 1997. An unidentified species of Theileria infective for cattle discovered in China. Tropical Animal Health Production, 29, S43-S47.
- BISHOP, R.P., HEMMINK, J.D., MORRISON, W.I., WEIR, W., TOYE, P.G., SITT, T., SPOONER, P.R., MUSOKE, A.J., SKILTON, R.A., ODONGO, D.O., 2015. The African buffalo parasite Theileria sp. (buffalo) can infect and immortalize cattle leukocytes and encodes divergent orthologues of Theileria parva antigen genes. International Journal of Parasitology: Parasites and Wildlife, 4, 333-342.
- BYARUHANGA, C., COLLINS, N.E., KNOBEL, D., CHAISI, M.E., VORSTER, I., STEYN, H.C. & OOSTHUIZEN, M.C., 2016. Molecular investigation of tick-borne haemoparasite infections among transhumant zebu cattle in Karamoja Region, Uganda. Veterinary Parasitology: Regional Studies and Reports, 3-4, 27-35.
- CHAISI, M.E., SIBEKO, K.P., COLLINS, N.E., POTGIETER, F.T. & OOSTHUIZEN, M.C., 2011. Identification of Theileria parva and Theileria sp. (buffalo) 18S rRNA gene sequence variants in the African buffalo (Syncerus caffer) in southern Africa. Veterinary Parasitology, 182, 150-162.
- CHAISI, M.E., COLLINS, N.E., POTGIETER, F.T. & OOSTHUIZEN, M.C., 2013. Sequence variation identified in the 18S rRNA gene of Theileria mutans and Theileria velifera from the African buffalo (Syncerus caffer). Veterinary Parasitology, 191, 132-137.
- CHAISI, M.E., COLLINS, N.E., OOSTHUIZEN, M.C., 2014. Phylogeny of Theileria buffeli genotypes identified in the South African buffalo (Syncerus caffer) population. Veterinary Parasitology, 204, 87-95.
- CONRAD, P.A., STAGG, D.A., GROOTENHUIS, J.G., IRVIN, A.D., NEWSON, J., NJAMUNGGEH, R.E., ROSSITER, P.B., YOUNG, A.S., 1987. Isolation of Theileria parasites from African buffalo (Syncerus caffer) and characterization with anti-schizont monoclonal antibodies. Parasitology, 94, 413-423.
- EYGELAAR, D., JORI, F., MOKOPASETSO, M., SIBEKO, K.P., COLLINS, N.E., VORSTER, I., TROSKIE, M., OOSTHUIZEN, M.C., 2015. Tick-borne haemoparasites in African buffalo (Syncerus caffer) from two wildlife areas in Northern Botswana. Parasites & Vectors, 8, 26.
- GITHAKA, N., KONNAI, S., BISHOP, R., ODONGO, D., LEKOLOOL, I., KARIUKI, E., GAKUYA, F., KAMAU, L., ISEZAKI, M., MURATA, S., OHASHI, K., 2014. Identification and sequence characterization of novel Theileria genotypes from the waterbuck (Kobus defassa) in a Theileria parva-endemic area in Kenya. Veterinary Parasitology, 202, 180-193.
- GUBBELS, J.M., DE VOS, A.P., VAN DER WEIDE, M., VISERAS, J., SCHOULS, L.M., DE VRIES, E. & JONGEJAN, F., 1999. Simultaneous detection of bovine Theileria and Babesia species by reverse line blot hybridization. Journal of Clinical Microbiology, 37, 1782-1789.
- GUBBELS, M.J., YIN, H., BAI, Q., LIU, G., NIJMAN, I.J., JONGEJAN, F., 2002. The phylogenetic position of the Theileria buffeli group in relation to other Theileria species. Parasitology Research, 88, S28-S32.
- LI, S., LIU, J., LIU, A., LI, Y., WANG, S., WANG, S., YIN, H., LUO, J., GUAN, G., 2017. Molecular investigation of piroplasma infection in white yaks (Bos grunniens) in Gansu province, China. Acta Tropica, 171, 220-225.
- LIU, A., GUAN, G., LIU, Z., LIU, J., LEBLANC, N., LI, Y., GAO, J., MA, M., NIU, Q., REN, Q., BAI, Q., YIN, H., LUO, J., 2010. Detecting and differentiating Theileria sergenti and Theileria sinensis in cattle and yaks by PCR based on major piroplasm surface protein (MPSP). Experimental Parasitology, 126, 476-481.
- LIU, J., GUAN, G., LIU, Z., LIU, A., MA, M., BAI, Q., YIN, H., LUO, J., 2013a. Additional data for a new Theileria sp. from China based on the sequences of ribosomal RNA internal transcribed spacers. Experimental Parasitology, 133, 217-221.
- LIU, A., GUAN, G., DU, P., GOU, H., ZHANG, J., LIU, Z., MA, M., REN, Q., LIU, J., YANG, J., LI, Y., NIU, Q., BAI, Q., YIN, H., LUO, J., 2013b. Rapid identification and differentiation of Theileria sergenti and Theileria sinensis using a loop-mediated isothermal amplification (LAMP) assay. Veterinary Parasitology, 191, 15-22.
- LORUSSO, V., WIJNVELD, M., MAJEKODUNMI, A.O., DONGKUM, C., FAJINMI, A., DOGO, A.G., THRUSFIELD, M., MUGEYI, A., VAUMOURIN, E., IGWEH, A.C., JONGEJAN, F., WELBURN, S.C. & PICOZZI, K., 2016. Tick-borne pathogens of zoonotic and veterinary importance in Nigerian cattle. Parasites & Vectors, 9, 217.
- MANS, B.J., PIENAAR, R., LATIF, A.A. & POTGIETER, F.T., 2011. Diversity in the 18S SSU rRNA V4 hyper-variable region of Theileria in bovines and African buffalo (Syncerus caffer) from southern Africa. Parasitology, 138, 766–779.
- MANS, B.J., PIENAAR, R. & LATIF, A.A., 2015. A review of Theileria diagnostics and epidemiology. International Journal for Parasitology: Parasites and Wildlife, 4, 104-118.
- MANS, B.J., PIENAAR, R., RATABANE, J., PULE, B., LATIF, A.A., 2016. Investigating the diversity of the 18S SSU rRNA hyper-variable region of Theileria in cattle and Cape buffalo (Syncerus caffer) from southern Africa using a next generation sequencing approach. Ticks and Tick Borne Diseases, 7, 869-879.
- MATJILA, P.T., LEISEWITZ, A.L., OOSTHUIZEN, M.C., JONGEJAN, F., PENZHORN, B.L., 2008. Detection of a Theileria species in dogs in South Africa. Veterinary Parasitology, 157, 34-40.
- MUHANGUZI, D., MATOVU, E. & WAISWA, C., 2010. Prevalence and characterization of Theileria and Babesia species in cattle under different husbandry systems in western Uganda. International Journal of Animal and Veterinary Advances,2, 51-58.
- NIU, Q., YANG, J., LIU, Z., GAO, S., PAN, Y., GUAN, G., CHU, Y., LIU, G., LUO, J., YIN, H., 2017. First Molecular Detection of Piroplasm Infection in Pet Dogs from Gansu, China. Frontiers in Microbiology, 8, 1029.
- NJIIRI, N.E., BRONSVOORT, B.M., COLLINS, N.E., STEYN, H.C., TROSKIE, M., VORSTER, I., THUMBI, S.M., SIBEKO, K.P., JENNINGS, A., VAN WYK, I.C., MBOLE-KARIUKI, M., KIARA, H., POOLE, E.J., HANOTTE, O., COETZER, K., OOSTHUIZEN, M.C., WOOLHOUSE, M., TOYE, P., 2015. The epidemiology of tick-borne haemoparasites as determined by the reverse line blot hybridization assay in an intensively studied cohort of calves in western Kenya. Veterinary Parasitology, 210, 69-76.
- NIJHOF, A.M., PILLAY, V., STEYL, J., PROZESKY, L., STOLTSZ, W.H., LAWRENCE, J.A., PENZHORN, B.L., JONGEJAN, F., 2005. Molecular characterization of Theileria species associated with mortality in four species of African antelopes. Journal of Clinical Microbiology, 43, 5907-5911.
- OURA, C.A.L., BISHOP, R.P., WAMPANDE, E.M., LUBEGA, G.W. & TAIT, A., 2004. Application of a reverse line blot assay to the study of haemoparasites in cattle in Uganda. International Journal for Parasitology,34, 603-613.
- OURA, C.A.L., TAIT, A., ASIIMWE, B., LUBEGA, G.W. & WEIR, W., 2011a. Haemoparasite prevalence and Theileria parva strain diversity in Cape buffalo (Syncerus caffer) in Uganda. Veterinary Parasitology,175, 212-219.
- OURA, C.A., TAIT, A., ASIIMWE, B., LUBEGA, G.W., WEIR, W., 2011b. Theileria parva genetic diversity and haemoparasite prevalence in cattle and wildlife in and around Lake Mburo National Park in Uganda. Parasitology Research, 108, 1365-1374.
- PFITZER, S., OOSTHUIZEN, M.C., BOSMAN, A.M., VORSTER, I., PENZHORN, B.L., 2011. Tick-borne blood parasites in nyala (Tragelaphus angasii, Gray 1849) from KwaZulu-Natal, South Africa. Veterinary Parasitology, 176, 126-131.
- PIENAAR, R., POTGIETER, F.T., LATIF, A.A., THEKISOE, O.M.M. & MANS, B.J., 2011. Mixed Theileria infections in free-ranging buffalo herds: implications for diagnosing Theileria parva infections in Cape buffalo (Syncerus caffer). Parasitology, 138, 884-895.
- PIENAAR, R., LATIF, A.A., THEKISOE, O.M., MANS, B.J., 2014. Geographic distribution of Theileria sp. (buffalo) and Theileria sp. (bougasvlei) in Cape buffalo (Syncerus caffer) in southern Africa: implications for speciation. Parasitology, 141, 411-424.
- QIN, G., LI, Y., LIU, J., LIU, Z., YANG, J., ZHANG, L., LIU, G., GUAN, G., LUO, J., YIN, H., 2016. Molecular detection and characterization of Theileria infection in cattle and yaks from Tibet Plateau Region, China. Parasitology Research, 115, 2647-2652.
- SCHREUDER, B.E.C., UILENBERG, G. & TONDEUR, W., 1977. Studies on Theileriidae (Sporozoa) in Tanzania. VIII. Experiments with African buffalo (Syncerus caffer). Tropenmedizin und Parasitologie, 28, 367–371.
- SILVEIRA, J.A., DE OLIVEIRA, C.H., SILVESTRE, B.T., ALBERNAZ, T.T., LEITE, R.C., BARBOSA, J.D., OLIVEIRA, C.M., RIBEIRO, M., 2016. Molecular assays reveal the presence of Theileria spp. and Babesia spp. in Asian water buffaloes (Bubalus bubalis, Linnaeus, 1758) in the Amazon region of Brazil. Ticks and Tick Borne Diseases, 7, 1017-1023.
- STEYL, J.C., PROZESKY, L., STOLTSZ, W.H., LAWRENCE, J.A., 2012. Theileriosis (Cytauxzoonosis) in Roan antelope (Hippotragus equinus): field exposure to infection and identification of potential vectors. Onderstepoort Journal of Veterinary Research, 79, 1-8.
- TIAN, Z., DU, J., YANG, J., LIU, A., LIU, X., LIU, G., YIN, H., 2015. A PCR-RFLP Assay targeting RPS8 gene for the discrimination between bovine Babesia and Theileria species in China. Parasites and Vectors, 8, 475.
- UILENBERG, G., 1964. Haematoxenus veliferus, n.g., n. sp., parasite Incertae sedis du sang de bovins à Madagascar. Revue d’Élevage et de Médecine Vétérinaire des Pays Tropicaux, 17, 655–662.
- UILENBERG, G., 1983. Acquisitions nouvelles dans la connaissance du rôle vecteur de tiques du genre Amblyomma (Ixodidae). Revue d’Élevage et de Médecine Vétérinaire des Pays Tropicaux, 36, 61–66.
- UILENBERG, G. & SCHREUDER, B.E.C., 1976. Studies on Theileriidae (Sporozoa) in Tanzania. 1. Tick transmission of Haematoxenus veliferus. Tropenmedizin und Parasitologie, 27, 106–111.
- VAN VORSTENBOSCH, C.J.A.H.V., UILENBERG, G. & VAN DIJK, J.E., 1978. Erythrocytic forms of Theileria velifera. Research in Veterinary Science, 24, 214–221.
- WALKER J.B., KEIRANS, J.E., HORAK, I.G., 2000. The Genus Rhipicephalus (Acari, Ixodidae): A Guide to the Brown Ticks of the World. Cambridge University Press.
- WARNECKE, M., SCHEIN, E., VOIGT, W.P. & UILENBERG, G., 1979. On the lifecycle of Theileria velifera (Uilenberg, 1964) in the gut and the haemolymph of the tick vector Amblyomma variegatum (Fabricius, 1794). Tropenmedizin Parasitologie, 30, 318–322.
- YIN, H., LUO, J., SCHNITTGER, L., LU, B., BEYER, D., MA, M., GUAN, G., BAI, Q., LU, C., AHMED, J., 2004. Phylogenetic analysis of Theileria species transmitted by Haemaphysalis qinghaiensis. Parasitology Research, 92, 36-42.
- YUSUFMIA, S.B., COLLINS, N.E., NKUNA, R., TROSKIE, M., VAN DEN BOSSCHE, P., PENZHORN, B.L., 2010. Occurrence of Theileria parva and other haemoprotozoa in cattle at the edge of Hluhluwe-iMfolozi Park, KwaZulu-Natal, South Africa. Journal of the South African Veterinary Association, 81, 45-49.
- ZHAO, S., LIU, J., ZHAO, H., LI, Y., XIE, J., LIU, A., HASSAN, M.A., YIN, H., GUAN, G., LUO, J., 2017. Evaluating an indirect rMPSP enzyme-linked immunosorbent assay for the detection of bovine Theileria infection in China. Parasitology Research, 116, 667-676.
- ZWEYGARTH, E., KOEKEMOER, O., JOSEMANS, A.I., RAMBRITCH, N., PIENAAR, R., PUTTERILL, J., LATIF, A., POTGIETER, F.T., 2009a. Theileria-infected cell line from an African buffalo (Syncerus caffer). Parasitology Research, 105, 579-581.
- ZWEYGARTH, E., BENADE, J., STEYL, J., PROZESKY, L., KOEKEMOER, O., JOSEMANS, A.I., 2009b. In vitro cultivation of a Theileria species from a roan antelope (Hippotragus equinus). Parasitology Research, 105, 1755-1757.