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Barcoding in trypanosomes


Trypanosomes (genus Trypanosoma) are parasites of humans, and wild and domestic mammals, in which they cause several economically and socially important diseases, including sleeping sickness in Africa and Chagas disease in the Americas. Despite the development of numerous molecular diagnostics and increasing awareness of the importance of these neglected parasites, there is currently no universal genetic barcoding marker available for trypanosomes. In this review we provide an overview of the methods used for trypanosome detection and identification, discuss the potential application of different barcoding techniques and examine the requirements of the ‘ideal’ trypanosome genetic barcode. In addition, we explore potential alternative genetic markers for barcoding Trypanosoma species, including an analysis of phylogenetically informative nucleotide changes along the length of the 18S rRNA gene.

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*Corresponding author. Department of Biosciences, College of Life and Environmental Sciences, University of Exeter, Stocker Road, Exeter EX4 4QD, UK. E-mail:
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Abderrazak, S., Guerrini, F., Mathieu-Daudé, F., Truc, P., Neubauer, K., Lewicka, K., Barnabé, C. and Tibayrenc, M. (1993). Isoenzyme electrophoresis for parasite characterisation. In Hyde, J. (ed.). Methods in Molecular Biology, Vol 21. Protocols in Molecular Parasitology. Totowa, New Jersey: Humana Press Inc., pp. 361382.
Acup, C., Bardosh, K. L., Picozzi, K., Waiswa, C. and Welburn, S. C. (2016). Factors influencing passive surveillance for T. b. rhodesiense human African trypanosomiasis in Uganda. Acta Tropica 165, 230239.
Adams, E. R. and Hamilton, P. B. (2008). New molecular tools for the identification of trypanosome species. Future Microbiology 3, 167176.
Adams, E. R., Hamilton, P. B., Malele, I. I. and Gibson, W. C. (2008). The identification, diversity and prevalence of trypanosomes in field caught tsetse in Tanzania using ITS-1 primers and fluorescent fragment length barcoding. Infection, Genetics and Evolution 8, 439444.
Adams, E. R., Hamilton, P. B. and Gibson, W. C. (2010). African trypanosomes: celebrating diversity. Trends in Parasitology 26, 324328.
Adjé, C. A., Opperdoes, F. R. and Michels, P. A. M. (1998). Molecular analysis of phosphoglycerate kinase in Trypanosoma borrelia and the evolution of this enzyme in Kinetoplastida. Gene 217, 9199.
Adl, S. M., Simpson, A. G. B., Lane, C. E., Lukes, J., Bass, D., Bowser, S. S., Brown, M. W., Burki, F., Dunthorn, M., Hampl, V., Heiss, A., Hoppenrath, M., Lara, E., La Gall, L., Lynn, D. H., McManus, H., Mitchell, E. A. D., Mozley-Stanridge, S. E., Parfrey, L. W., Pawlowski, J., Rueckert, S., Shadwick, L., Schoch, C. L. and Smirnov, F. A. (2012). The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59, 429493.
Auty, H., Anderson, N. E., Picozzi, K., Lembo, T., Mubanga, J., Hoare, R., Fyumagwa, R. D., Mable, B., Hamill, L., Cleaveland, S. and Welburn, S. C. (2012a). Trypanosome diversity in wildlife species from the Serengeti and Luangwa Valley ecosystems. PLoS Neglected Tropical Diseases 6, e1828.
Auty, H. K., Picozzi, K., Malele, I., Torr, S. J., Cleaveland, S. and Welburn, S. (2012b). Using molecular data for epidemiological inference: assessing the prevalence of Trypanosoma brucei rhodesiense in tsetse in Serengeti, Tanzania. PLoS Neglected Tropical Diseases 6, e1501.
Averis, S., Thompson, R. C. A., Lymbery, A. J., Wayne, A. F., Morris, K. D. and Smith, A. (2009). The diversity, distribution and host-parasite associations of trypanosomes in Western Australian wildlife. Parasitology 136, 12691279.
Balmer, O., Beadell, J. S., Gibson, W. and Caccone, A. (2011). Phylogeography and taxonomy of Trypanosoma brucei. PLoS Neglected Tropical Diseases 5, e961.
Barbosa, A. D., Mackie, J. T., Stenner, R., Gillett, A., Irwin, P. and Ryan, U. (2016). Trypanosoma teixeirae: a new species belonging to the T. cruzi clade causing trypanosomiasis in an Australian little red flying fox (Pteropus scapulatus). Veterinary Parasitology 223, 214221.
Barrett, M. P. and Croft, S. L. (2012). Management of trypanosomiasis and leishmaniasis. British Medical Bulletin 104, 175196.
Basiye, F. L., Schoone, G. J., Beld, M., Minnaar, R., Ngeranwa, J. N., Wasunna, M. K. and Schallig, H. D. F. H. (2011). Comparison of short-term and long-term protocols for stabilization and preservation of RNA and DNA of Leishmania, Trypanosoma, and Plasmodium. Diagnostic Microbiology and Infectious Disease 69, 6673.
Besuschio, S. A., Murcia, M. L., Benatar, A. F., Monnerat, S., Cruz, I., Picado, A., Curto, M. D. L. A., Kubota, Y., Wehrendt, D. P., Pavia, P., Mori, Y., Puerta, C., Ndung'u, J. M. and Schijman, A. G. (2017). Analytical sensitivity and specificity of a loop-mediated isothermal amplification kit prototype for detection of Trypanosoma cruzi DNA in human blood samples. PLoS Neglected Tropical Diseases 11, e0005779.
Chappuis, F., Loutan, L., Simarro, P., Lejon, V. and Büscher, P. (2005). Options for field diagnosis of human African trypanosomiasis. Clinical Microbiology Reviews 18, 133146.
Clarivate Statistics (2017). Web of Science [v.5.24] – Web of Science Core Collection Home. Available at (Accessed September 2017).
Coissac, E., Hollingsworth, P. M., Lavergne, S. and Taberlet, P. (2016). From barcodes to genomes: extending the concept of DNA barcoding. Molecular Ecology 25, 14231428.
Dario, M. A., Moratelli, R., Schwabi, P., Jansen, A. M. and Llewellyn, M. S. (2017). Small subunit ribosomal metabarcoding reveals extraordinary trypanosomatid diversity in Brazilian bats. PLoS Neglected Tropical Diseases 11, e0005790.
D'Avila-Levy, C. M., Boucinha, C., Kostygov, A., Santos, H. L. C., Morelli, K. A., Grybchuk-Ieremenko, A., Duval, L., Votýpka, J., Yurchenko, V., Grellier, P. and Lukeš, J. (2015). Exploring the environmental diversity of kinetoplastid flagellates in the high-throughput DNA sequencing era. Memórias Do Instituto Oswaldo Cruz 110, 956965.
Del Campo, J., Sieracki, M. E., Molestina, R., Keeling, P., Massana, R. and Ruiz-Trillo, I. (2015). The others: our biased perspective of eukaryotic genomes. Trends in Evolution & Ecology 29, 252259.
De Oliveira Ramos Pereira, L. and Brandão, A. (2013). An analysis of trypanosomatids kDNA minicircle by absolute dinucleotide frequency. Parasitology International 62, 397403.
De Waal, T. (2012). Advances in diagnosis of protozoan diseases. Veterinary Parasitology 189, 6574.
Deborggraeve, S. and Büscher, P. (2010). Molecular diagnostics for sleeping sickness: what is the benefit for the patient? The Lancet Infectious Diseases 10, 433439.
Desquesnes, M. and Davila, A. (2002). Applications of PCR-based tools for detection and identification of animal trypanosomes: a review and perspectives. Veterinary Parasitology 109, 213231.
Desquesnes, M., Kamyingkird, K., Yangtara, S., Milocco, C., Ravel, S., Wang, M. H., Lun, Z. R., Morand, S. and Jittapalapong, S. (2011). Specific primers for PCR amplification of the ITS1 (ribosomal DNA) of Trypanosoma lewisi. Infection, Genetics and Evolution 11, 13611367.
Dunlop, J., Thompson, C. K., Godfrey, S. S. and Thompson, R. C. A. (2014). Sensitivity testing of trypanosome detection by PCR from whole blood samples using manual and automated DNA extraction methods. Experimental Parasitology 146, 2024.
Enyaru, J. C., Ouma, J. O., Malele, I. I., Matovu, E. and Masiga, D. K. (2010). Landmarks in the evolution of technologies for identifying trypanosomes in tsetse flies. Trends in Parasitology 26, 388394.
Fernandes, A. P., Nelson, K. and Beverley, S. M. (1993). Evolution of nuclear ribosomal RNAs in kinetoplastid protozoa: perspectives on the age and origins of parasitism. Proceedings of the National Academy of Sciences of the USA 90, 1160811612.
Ferri, G., Alù, M., Corradini, B., Licata, M. and Beduschi, G. (2009). Species identification through DNA ‘barcodes’. Genetic Testing and Molecular Biomarkers 13, 16.
Févre, E. M., Picozzi, K., Fyfe, J., Waiswa, C., Odiit, M., Coleman, P. G. and Welburn, S. C. (2005). A burgeoning epidemic of sleeping sickness in Uganda. Lancet 366, 745747.
Field, M. C., Horn, D., Fairlamb, A. H., Ferguson, M. A. J., Gray, D. W., Read, K. D., De Rycker, M., Torrie, L. S., Wyatt, P. G., Wyllie, S. and Gilbert, I. H. (2017). Anti-trypanosomatid drug discovery: an ongoing challenge and a continuing need. Nature Reviews Microbiology 15, 217231.
Flegontov, P., Votýpka, J., Skalický, T., Logacheva, M. D., Penin, A. A., Tanifuji, G., Onodera, N. T., Kondrashov, A. S., Volf, P., Archibald, J. M. and Lukeš, J. (2013). Paratrypanosoma is a novel early-branching Trypanosomatid. Current Biology 23, 17871793.
Franco, J. R., Simarro, P. P., Diarra, A., Ruiz-Postigo, J. A. and Jannin, J. G. (2014). The journey towards the elimination of gambiense human African trypanosomiasis: not far, nor easy. Parasitology 141, 748760.
Gashumba, J. K., Gibson, W. C. and Opiyo, E. A. (1986). A preliminary comparison of Trypanosoma simiae and T. congolense by isoenzyme electrophoresis. Acta Tropica 43, 1519.
Gibson, W. (2007). Resolution of the species problem in African trypanosomes. International Journal for Parasitology 37, 829838.
Gibson, W. (2009). Species-specific probes for the identification of the African tsetse-transmitted trypanosomes. Parasitology 136, 15011507.
Gibson, W. and Borst, P. (1986). Size-fractionation of the small chromosomes of Trypanozoon and Nannomonas trypanosomes by pulsed field gradient gel electrophoresis. Molecular and Biochemical Parasitology 18, 127140.
Gibson, W. C., Dukes, P. and Gashumba, J. K. (1988). Species-specific DNA probes for the identification of African trypanosomes in tsetse flies. Parasitology 97, 6373.
Gibson, W., Bingle, L., Blendeman, W., Brown, J., Wood, J. and Stevens, J. (2000). Structure and sequence variation of the trypanosome spliced leader transcript. Molecular and Biochemical Parasitology 107, 269277.
Godfrey, D. G. and Kilgour, V. (1976). Enzyme electrophoresis in characterizing the causative organism of Gambian trypanosomiasis. Transactions of the Royal Society of Tropical Medicine and Hygiene 70, 219224.
Godfrey, D. G., Baker, R. D., Rickman, L. R. and Mehlitz, D. (1990). The distribution, relationships and identification of enzymic variants within the subgenus Trypanozoon. Advances in Parasitology 29, 174.
Haag, J., O'hUigin, C. and Overath, P. (1998). The molecular phylogeny of trypanosomes: evidence for an early divergence of the Salivaria. Molecular and Biochemical Parasitology 91, 3749.
Hadziavdic, K., Lekang, K., Lanzen, A., Jonassen, I., Thompson, E. M. and Troedsson, C. (2014). Characterization of the 18S rRNA gene for designing universal eukaryote specific primers. PLoS ONE 9, eB7624.
Hamilton, P. B. and Stevens, J. R. (2011). Resolving relationships between Australian trypanosomes using DNA barcoding data. Trends in Parasitology 27, 99.
Hamilton, P. B., Stevens, J. R., Gaunt, M. W., Gidley, J. and Gibson, W. C. (2004). Trypanosomes are monophyletic: evidence from genes for glyceraldehyde phosphate dehydrogenase and small subunit ribosomal RNA. International Journal for Parasitology 34, 13931404.
Hamilton, P. B., Gibson, W. C. and Stevens, J. R. (2007). Patterns of co-evolution between trypanosomes and their hosts deduced from ribosomal RNA and protein-coding gene phylogenies. Molecular Phylogenetics and Evolution 44, 1525.
Hamilton, P. B., Adams, E. R., Malele, I. I. and Gibson, W. C. (2008). A novel, high-throughput technique for species identification reveals a new species of tsetse-transmitted trypanosome related to the Trypanosoma brucei subgenus, Trypanozoon. Infection, Genetics and Evolution 8, 2633.
Hamilton, P. B., Lewis, M. D., Cruickshank, C., Gaunt, M. W., Yeo, M., Llewellyn, M. S., Valente, S. A., Da Silva, F. M., Stevens, J. R., Miles, M. A. and Teixeira, M. M. G. (2011). Identification and lineage genotyping of South American trypanosomes using fluorescent fragment length barcoding. Infection, Genetics and Evolution 11, 4451.
Hashimoto, T., Nakamura, Y., Kamaishi, T., Adachi, J., Nakamura, F., Okamoto, K. and Hasegawa, M. (1995). Phylogenetic place of kinetoplastid protozoa inferred from a protein phylogeny of elongation factor 1α. Molecular and Biochemical Parasitology 70, 181185.
Hayden, E. (2014). The $1,000 genome. Nature 507, 294295.
Hebert, P. D. N., Cywinska, A., Ball, S. L. and deWaard, J. R. (2003). Biological identifications through DNA barcodes. Proceedings of the Royal Society of London B: Biological Sciences 270, 313321.
Hernández, C. and Ramírez, J. D. (2013). Molecular diagnosis of vector-borne parasitic diseases. Air & Water Borne Diseases 2, 110.
Hide, G., Cattand, P., Le Ray, D., Barry, J. D. and Tait, A. (1990). The identification of T. brucei subspecies using repetitive DNA sequences. Molecular and Biochemical Parasitology 39, 213226.
Hoare, C. (1972). The Trypanosomes of Mammals. Oxford: Blackwell Scientific Publications.
Hutchinson, R. and Gibson, W. (2015). Rediscovery of Trypanosoma (Pycnomonas) suis, a tsetse-transmitted trypanosome closely related to T. brucei. Infection, Genetics and Evolution 36, 381388.
Jensen, R. and Englund, P. (2012). Network news: the replication of kinetoplast DNA. Annual Review of Microbiology 66, 473491.
Kilgour, V. and Godfrey, D. G. (1973). Isoenzymes of alanine aminotransferase as possible specific characters of trypanosomes. Transactions of the Royal Society of Tropical Medicine and Hygiene 67, 11.
Koffi, M., De Meeûs, T., Séré, M., Bucheton, B., Simo, G., Njiokou, F., Salim, B., Kaboré, J., MacLeod, A., Camara, M., Solano, P., Belem, A. M. G. and Jamonneau, V. (2015). Population genetics and reproductive strategies of African trypanosomes: revisiting available published data. PLoS Neglected Tropical Diseases 9, e0003985.
Kukla, B. A., Majiwa, P. A. O., Young, J. R., Moloo, S. K. and Ole-Moiyoi, O. K. (1987). Use of species-specific DNA probes for detection and identification of trypanosome infection in tsetse flies. Parasitology 95, 116.
Lanham, S. M. and Godfrey, D. G. (1970). Isolation of salivarian trypanosomes from man and other mammals using DEAE-cellulose. Experimental Parasitology 28, 521534.
Laohasinnarong, D. (2011). Loop-mediated isothermal amplification (LAMP): an alternative molecular diagnosis. Journal of Applied Animal Science 4, 919.
Leonard, G., Soanes, D. M. and Stevens, J. R. (2011). Resolving the question of trypanosome monophyly: a comparative genomics approach using whole genome data sets with low taxon sampling. Infection, Genetics and Evolution 11, 955959.
Lima, L., Espinosa-Álvarez, O., Pinto, C. M., Cavazzana, M. Jr, Pavan, A. C., Carranza, J. C., Lim, B. K., Campaner, M., Takata, C. S. A., Camargo, E. P., Hamilton, P. B. and Teixeira, M. M. G. (2015). New insights into the evolution of the Trypanosoma cruzi clade provided by a new trypanosome species tightly linked to Neotropical Pteronotus bats and related to an Australian lineage of trypanosomes. Parasites & Vectors 8, 657.
Lukes, J., Jirku, M., Dolezel, D., Kral'ova, I., Hollar, L. and Maslov, D. A. (1997). Analysis of the ribosomal RNA gene suggests that trypanosomes are monophyletic. Journal of Molecular Evolution 44, 521527.
Lutumba, P., Matovu, E. and Boelaert, M. (2016). Human African Trypanosomiasis (HAT). In Gyapong, J and Boatin, B (eds). Neglected Tropical Diseases - Sub-Saharan Africa. Switzerland: Springer International Publishing, pp. 6385.
Mallo, D. and Posada, D. (2016). Multilocus inference of species trees and DNA barcoding. Philosophical Transactions of the Royal Society of London B 371, 20150335.
Maslov, D. A., Lukes, J., Jirku, M. and Simpson, L. (1996). Phylogeny of trypanosomes as inferred from the small and large subunit rRNAs: implications for the evolution of parasitism in the trypanosomatid protozoa. Molecular and Biochemical Parasitology 75, 197205.
Matovu, E., Mugasa, C. M., Ekangu, R. A., Deborggraeve, S., Lubega, G. W., Laurent, T., Schoone, G. J., Schallig, H. D. and Büscher, P. (2010). Phase II evaluation of sensitivity and specificity of PCR and NASBA followed by oligochromatography for diagnosis of human African trypanosomiasis in clinical samples from D.R. Congo and Uganda. PLoS Neglected Tropical Diseases 4, 38.
McNamara, J., Dukes, P., Snow, W. F. and Gibson, W. C. (1989). Use of DNA probes to identify Trypanosoma congolense and T. simiae in tsetse flies from The Gambia. Acta Tropica 46, 5561.
Messenger, L. A., Llewellyn, M. S., Bhattacharyya, T., Franzén, O., Lewis, M. D., Ramírez, J. D., Carrasco, H. J., Andersson, B. and Miles, M. A. (2012). Multiple mitochondrial introgression events and heteroplasmy in Trypanosoma cruzi revealed by maxicircle MLST and next generation sequencing. PLoS Neglected Tropical Diseases 6, e1584.
Miles, M. A., Toye, P. J., Oswald, S. C. and Godfrey, D. G. (1977). The identification by isoenzyme patterns of two distinct strain-groups of Trypanosoma cruzi circulating independently in a rural area of Brazil. Transactions of the Royal Society of Tropical Medicine and Hygiene 71, 217225.
Mpanya, A., Hendrickx, D., Vuna, M., Kanyinda, A., Lumbala, C., Tshilombo, V., Mitashi, P., Luboya, O., Kande, V., Boelaert, M., Lefèvre, P. and Lutumba, P. (2012). Should I get screened for sleeping sickness? A qualitative study in Kasai Province, Democratic Republic of Congo. PLoS Neglected Tropical Diseases 6, e1467.
Mugasa, C. M., Adams, E. R., Boer, K. R., Dyserinck, H. C., Büscher, P., Schallig, H. D. H. F. and Leeflang, M. M. G. (2012). Diagnostic accuracy of molecular amplification tests for human African trypanosomiasis-systematic review. PLoS Neglected Tropical Diseases 6, e1438.
Mugasa, C. M., Katiti, D., Boobo, A., Lubega, G. W., Schallig, H. D. F. H. and Matovu, E. (2014). Comparison of nucleic acid sequence-based amplification and loop-mediated isothermal amplification for diagnosis of human African trypanosomiasis. Diagnostic Microbiology and Infectious Disease 78, 144148.
National Center for Biotechnology Information (NCBI) (2017). Bethesda, MD: National Library of Medicine (US), National Center for Biotechnology Information website. Available at (Accessed April 2017).
Ouma, J. O., Masake, R. A., Masiga, D. K., Moloo, S. K., Njuguna, J. T. and Ndung'u, J. M. (2000). Comparative sensitivity of dot-ELISA, PCR and dissection method for the detection of trypanosome infections in tsetse flies (Diptera: Glossinidae). Acta Tropica 75, 315321.
Papadopoulos, M. C., Abel, P. M., Agranoff, D., Stich, A., Tarelli, E., Bell, B. A., Planche, T., Loosemore, A., Saadoun, S., Wilkins, P. and Krishna, S. (2004). A novel and accurate diagnostic test for human African trypanosomiasis. Lancet 363, 13581363.
Pawlowski, J., Audic, S., Adl, S., Bass, D., Belbahri, L., Berney, C., Bowser, S. S., Cepicka, I., Decelle, J., Dunthorn, M., Fiore-Donno, A. M., Gile, G. H., Holzmann, M., Jahn, R., Jirků, M., Keeling, P. J., Kostka, M., Kudryavtsev, A., Lara, E., Lukeš, J., Mann, D. G., Mitchell, E. A. D., Nitsche, F., Romeralo, M., Saunders, G. W., Simpson, A. G. B., Smirnov, A. V., Spouge, J. L., Stern, R. F., Stoeck, T., Zimmermann, J., Schindel, D. and De Vargas, C. (2012). CBOL protist Working Group: barcoding eukaryotic richness beyond the animal, plant, and fungal kingdoms. PLoS Biology 10, e1001419.
Pečnikar, Z. F. and Buzan, E. V. (2014). 20 years since the introduction of DNA barcoding: from theory to application. Journal of Applied Genetics 55, 4352.
Pompanon, F. and Samadi, S. (2015). Next generation sequencing for characterizing biodiversity: promises and challenges. Genetica 143, 133138.
Ratnasingham, S. and Hebert, P. D. N. (2007). BOLD: the barcode of life data system ( Molecular Ecology Notes 7, 355364.
Reeves, L. E., Holderman, C. J., Gillett-Kaufman, J. L., Kawahara, A. Y. and Kaufman, P. E. (2016). Maintenance of host DNA integrity in field-preserved mosquito (Diptera: Culicidae) blood meals for identification by DNA barcoding. Parasites & Vectors 9, 503.
Ricciardi, A. and Ndao, M. (2015). Diagnosis of parasitic infections: what's going on? Journal of Biomolecular Screening 20, 621.
Rivero, R., Bisio, M., Velázquez, E. B., Esteva, M. I., Scollo, K., González, N. L., Altcheh, J. and Ruiz, A. M. (2017). Rapid detection of Trypanosoma cruzi by colorimetric loop-mediated isothermal amplification (LAMP): A potential novel tool for the detection of congenital Chagas infection. Diagnostic Microbiology and Infectious Disease 89, 2628.
Rodrigues, A. C., Garcia, H. A., Batista, J. S., Minervino, A. H. H., Góes-Cavalcante, G., Da Silva, F. M., Ferreira, R. C., Campaner, M., Paiva, F. and Teixeira, M. M. G. (2010). Characterisation of spliced leader genes of Trypanosoma (Megatrypanum) theileri: phylogeographical analysis of Brazilian isolates from cattle supports spatial clustering of genotypes and parity with ribosomal markers. Parasitology 137, 111122.
Savolainen, V., Cowan, R. S., Vogler, A. P., Roderick, G. K. and Lane, R. (2005). Towards writing the encyclopaedia of life: an introduction to DNA barcoding. Philosophical Transactions of the Royal Society B 360, 18051811.
Silva-Iturriza, A., Nassar, J. M., Garcia-Rawlins, A. M., Rosales, R. and Mijares, A. (2013). Trypanosoma evansi kDNA minicircle found in the Venezuelan nectar-feeding bat Leptonycteris curasoae (Glossophaginae), supports the hypothesis of multiple origins of that parasite in South America. Parasitology International 62, 9599.
Simpson, A. G. B., Gill, E. E., Callahan, H. A., Litaker, R. W. and Roger, A. J. (2004). Early evolution within kinetoplastids (Euglenozoa), and the late emergence of trypanosomatids. Protist 155, 407422.
Smith, A., Clark, P., Averis, S., Lymbery, A. J., Wayne, A. F., Morris, K. D. and Thompson, R. C. A. (2008). Trypanosomes in a declining species of threatened Australian marsupial, the brush-tailed bettong Bettongia penicillata (Marsupialia: Potoroidae). Parasitology 135, 13291335.
Stevens, J. and Rambaut, A. (2001). Evolutionary rate differences in trypanosomes. Infection, Genetics and Evolution 1, 143150.
Stevens, J. and Wall, R. (2001). Genetic relationships between blowflies (Calliphoridae) of forensic importance. Forensic Science International 120, 116123.
Stevens, J., Noyes, H. and Gibson, W. (1998). The evolution of trypanosomes infecting humans and primates. Memórias Do Instituto Oswaldo Cruz 93, 669676.
Stevens, J. R. and Godfrey, D. G. (1992). Numerical taxonomy of Trypanozoon based on polymorphisms in a reduced range of enzymes. Parasitology 104, 7586.
Stevens, J. R., Noyes, H. A., Dover, G. A. and Gibson, W. C. (1999). The ancient and divergent origins of the human pathogenic trypanosomes, Trypanosoma brucei and T. cruzi. Parasitology 118, 107116.
Stockdale, L. and Newton, R. (2013). A review of preventative methods against human leishmaniasis infection. PLOS Neglected Tropical Disease 7, e2278.
Stuart, K., Brun, R., Croft, S., Fairlamb, A., Gürtler, R. E., McKerrow, J., Reed, S. and Tarleton, R. (2008). Kinetoplastids: related protozoan pathogens, different diseases. The Journal of Clinical Investigation 118, 13011310.
Swofford, D. L. (2002). PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4.0. Sunderland, Massachusetts: Sinauer Associates.
Taberlet, P., Coissac, E., Pompanon, F., Brochmann, C. and Willerslev, E. (2012). Towards next-generation biodiversity assessment using DNA metabarcoding. Molecular Ecology 21, 20452050.
Tihon, E., Imamura, H., Dujardin, J. C., Van Den Abbeele, J. and Van den Broeck, F. (2017). Discovery and genomic analysis of hybridization between divergent lineages of Trypanosoma congolense, causative agent of animal African trypanosomiasis. Molecular Ecology, 115, doi: 10.1111/mec.14271.
Uilenberg, G. and Boyt, W. P. (1998). A Field Guide for the Diagnosis, Treatment and Prevention of African Animal Trypanosomosis. Rome, Italy: Food and Agriculture Organisation of the United Nations.
Valentini, A., Pompanon, F. and Taberlet, P. (2009). DNA barcoding for ecologists. Trends in Ecology & Evolution 24, 110117.
Van Nimwegen, K. J. M., Van Soest, R. A., Veltman, J. A., Nelen, M. R., Van Der Wilt, G. J., Vissers, L. E. L. M. and Grutters, J. P. C. (2016). Is the $1000 genome as near as we think? A cost analysis of next-generation sequencing. Clinical Chemistry 62, 14581464.
Votýpka, J., Maslov, D. A., Yurchenko, V., Jirků, M., Kment, P., Lun, Z. and Lukeš, J. (2010). Probing into the diversity of trypanosomatid flagellates parasitizing insect hosts in South-West China reveals both endemism and global dispersal. Molecular Phylogenetics and Evolution 54, 243253.
Wastling, S. L. and Welburn, S. C. (2011). Diagnosis of human sleeping sickness: sense and sensitivity. Trends in Parasitology 27, 394402.
Westenberger, S. J., Sturm, N. R., Yanega, D., Podlipaev, S. A., Zeledon, R., Campbell, D. A. and Maslov, D. A. (2004). Trypanosomatid biodiversity in Costa Rica: genotyping of parasites from Heteroptera using the spliced leader RNA gene. Parasitology 129, 537547.
Woods, C. R. (2013). False-positive results for immunoglobulin M serologic results: explanations and examples. Journal of the Pediatric Infectious Diseases Society 2, 8790.
World Health Organisation (2013). Control and Surveillance of Human African Trypanosomiasis. WHO Technical Report Series. No 984. Geneva, Switzerland: World Health Organisation.
Yaro, M., Munyard, K. A., Stear, M. J. and Groth, D. M. (2016). Combatting African animal trypanosomiasis (AAT) in livestock: the potential role of trypanotolerance. Veterinary Parasitology 225, 4352.
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