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An appraisal of the taxonomy and nomenclature of trypanosomatids presently classified as Leishmania and Endotrypanum

  • O. A. ESPINOSA (a1), M. G. SERRANO (a2), E. P. CAMARGO (a1), M. M. G. TEIXEIRA (a1) and J. J. SHAW (a1)...


We propose a taxonomic revision of the dixenous trypanosomatids currently classified as Endotrypanum and Leishmania, including parasites that do not fall within the subgenera L. (Leishmania) and L. (Viannia) related to human leishmaniasis or L. (Sauroleishmania) formed by leishmanias of lizards: L. colombiensis, L. equatorensis, L. herreri, L. hertigi, L. deanei, L. enriettii and L. martiniquensis. The comparison of these species with newly characterized isolates from sloths, porcupines and phlebotomines from central and South America unveiled new genera and subgenera supported by past (RNA PolII gene) and present (V7V8 SSU rRNA, Hsp70 and gGAPDH) phylogenetic analyses of the organisms. The genus Endotrypanum is restricted to Central and South America, comprising isolates from sloths and transmitted by phlebotomines that sporadically infect humans. This genus is the closest to the new genus Porcisia proposed to accommodate the Neotropical porcupine parasites originally described as L. hertigi and L. deanei. A new subgenus Leishmania (Mundinia) is created for the L. enriettii complex that includes L. martiniquensis. The new genus Zelonia harbours trypanosomatids from Neotropical hemipterans placed at the edge of the Leishmania–Endotrypanum-Porcisia clade. Finally, attention is drawn to the status of L. siamensis and L. australiensis as nomem nudums.


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*Corresponding authors. Departamento de Parasitologia, Instituto de Ciências Biomédicas, Universidade de São Paulo, São Paulo, SP 05508-900 Brazil. E-mail:,


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Akhoundi, M., Kuhls, K., Cannet, A., Votypka, J., Marty, P., Delaunay, P. and Sereno, D. (2016). A historical overview of the classification, evolution, and dispersion of Leishmania Parasites and sandflies. PLoS Neglected Tropical Diseases 10, e0004349.
Arias, J. R., Miles, M. A., Naiff, R. D., Povoa, M. M., de Freitas, R. A., Biancardi, C. B. and Castellon, E. G. (1985). Flagellate infections of Brazilian sand flies (Diptera: Psychodidae): isolation in vitro and biochemical identification of Endotrypanum and Leishmania. American Journal of Tropical Medicine and Hygiene 34, 10981108.
Asato, Y., Oshiro, M., Myint, C. K., Yamamoto, Y., Kato, H., Marco, J. D., Mimori, T., Gomez, E. A., Hashiguchi, Y. and Uezato, H. (2009). Phylogenic analysis of the genus Leishmania by cytochrome b gene sequencing. Experimental Parasitology 121, 352361.
Bualert, L., Charungkiattikul, W., Thongsuksai, P., Mungthin, M., Siripattanapipong, S., Khositnithikul, R., Naaglor, T., Ravel, C., El Baidouri, F. and Leelayoova, S. (2012). Autochthonous disseminated dermal and visceral Leishmaniasis in an AIDS patient, Southern Thailand, caused by Leishmania siamensis. American Journal of Tropical Medicine and Hygiene 86, 821824.
Cassia-Pires, R., Boite, M. C., D'Andrea, P. S., Herrera, H. M., Cupolillo, E., Jansen, A. M. and Roque, A. L. (2014). Distinct Leishmania species infecting wild caviomorph rodents (Rodentia: Hystricognathi) from Brazil. PLoS Neglected Tropical Diseases 8, e3389.
Christensen, H. and Herrer, A. (1976). Neotropical sand flies (Diptera: Psychodidae), invertebrate hosts of Endotrypanum schaudinni (Kinetoplastidae). Journal of Medical Entomology 13, 299303.
Croan, D. G., Morrison, D. A. and Ellis, J. T. (1997). Evolution of the genus Leishmania revealed by comparison of DNA and RNA polymerase gene sequences. Molecular and Biochemical Parasitology 89, 149159.
Cupolillo, E., Pereira, L. O., Fernandes, O., Catanho, M. P., Pereira, J. C., Medina-Acosta, E. and Grimaldi, G. Jr. (1998). Genetic data showing evolutionary links between Leishmania and Endotrypanum. Memórias do Instituo Oswaldo Cruz 93, 677683.
Cupolillo, E., Medina-Acosta, E., Noyes, H., Momen, H. and Grimaldi, G. Jr. (2000). A revised classification for Leishmania and Endotrypanum. Parasitology Today 16, 142144.
Deane, L. M., da Silva, J. E. and de Figueiredo, P. Z. (1974). Leishmaniae in the viscera of porcupines from the state of Piaui, Brazil. Revista do Instituto de Medicina Tropical de São Paulo 16, 6869.
Dedet, J. P. and Pratlong, F. (2000). Leishmania, Trypanosoma and monoxenous trypanosomatids as emerging opportunistic agents. Journal of Eukaryotic Microbiology 47, 3739.
Dedet, J. P., Roche, B., Pratlong, F., Cales-Quist, D., Jouannelle, J., Benichou, J. C. and Huerre, M. (1995). Diffuse cutaneous infection caused by a presumed monoxenous trypanosomatid in a patient infected with HIV. Transactions of the Royal Society of Tropical Medicine and Hygiene 89, 644646.
Desbois, N., Pratlong, F., Quist, D. and Dedet, J. P. (2014). Leishmania (Leishmania) martiniquensis n. sp. (Kinetoplastida: Trypanosomatidae), description of the parasite responsible for cutaneous leishmaniasis in Martinique Island (French West Indies). Parasite 21, 12.
Dougall, A. M., Alexander, B., Holt, D. C., Harris, T., Sultan, A. H., Bates, P. A., Rose, K. and Walton, S. F. (2011). Evidence incriminating midges (Diptera: Ceratopogonidae) as potential vectors of Leishmania in Australia. International Journal for Parasitology 41, 571579.
Franco, A. M., Tesh, R. B., Guzman, H., Deane, M. P. and Grimaldi Junior, G. (1997). Development of Endotrypanum (Kinetoplastida:Trypanosomatidae) in experimentally infected phlebotomine sand flies (Diptera:Psychodidae). Journal of Medical Entomology 34, 189192.
Ghosh, S., Banerjee, P., Sarkar, A., Datta, S. and Chatterjee, M. (2012). Coinfection of Leptomonas seymouri and Leishmania donovani in Indian leishmaniasis. Journal of Clinical Microbiology 50, 27742778.
Grimaldi Junior, G., Kreutzer, R. D., Hashiguchi, Y., Gomez, E. A., Mimory, T. and Tesh, R. B. (1992). Description of Leishmania equatorensis sp. n (Kinetoplastida: Trypanosomatidae), a new parasite infecting arboreal mammals in Ecuador. Memórias do Instituo Oswaldo Cruz 87, 221228.
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.
Harkins, K. M., Schwartz, R. S., Cartwright, R. A. and Stone, A. C. (2016). Phylogenomic reconstruction supports supercontinent origins for Leishmania. Infection, Genetics and Evolution 38, 101109.
Herrer, A. (1971). Leishmania hertigi sp. n., from the tropical porcupine, Coendou rothschildi Thomas. Journal of Parasitology 57, 626629.
Jimenez, M. I., Lopez-Velez, R., Molina, R., Canavate, C. and Alvar, J. (1996). HIV co-infection with a currently non-pathogenic flagellate. Lancet 347, 264265.
Jirkú, M., Yurchenko, V. Y., Lukeš, J. and Maslov, D. A. (2012). New species of insect trypanosomatids from Costa Rica and the proposal for a new subfamily within the Trypanosomatidae. Journal of Eukaryotic Microbiology 59, 537547.
Kostygov, A. Y., Grybchuk-Ieremenko, A., Malysheva, M. N., Frolov, A. O. and Yurchenko, V. (2014). Molecular revision of the genus Wallaceina. Protist 165, 594604.
Kostygov, A. Y., Dobakova, E., Grybchuk-Ieremenko, A., Vahala, D., Maslov, D. A., Votypka, J., Lukeš, J. and Yurchenko, V. (2016). Novel Trypanosomatid-Bacterium Association: evolution of endosymbiosis in action. MBio 7, e01985.
Kreutzer, R. D., Corredor, A., Grimaldi, G., Grogl, M., Rowton, E. D., Young, D. G., Morales, A., McMahon-Pratt, D., Guzman, H. and Tesh, R. B. (1991). Characterization of Leishmania colombiensis sp. n (Kinetoplastida: Trypanosomatidae), a new parasite infecting humans, animals, and phlebotomine sand flies in Colombia and Panama. American Journal of Tropical Medicine and Hygiene 44, 662675.
Kwakye-Nuako, G., Mosore, M. T., Duplessis, C., Bates, M. D., Puplampu, N., Mensah-Attipoe, I., Desewu, K., Afegbe, G., Asmah, R. H., Jamjoom, M. B., Ayeh-Kumi, P. F., Boakye, D. A. and Bates, P. A. (2015). First isolation of a new species of Leishmania responsible for human cutaneous leishmaniasis in Ghana and classification in the Leishmania enriettii complex. International Journal for Parasitology 45, 679684.
Lainson, R. and Shaw, J. J. (1979). The Role of Animals in the Epidemiology of South American Leishmaniasis. In Biology of the Kinetoplastida, Vol. 2 (eds. Lumsden, W. H. R. and Evans, D. A.), pp. 1116. Academic Press, London, New York & San Francisco.
Lainson, R. and Shaw, J. J. (1977). Leishmanias of neotropical porcupines: Leishmania hertigi deanei nov.subsp. Acta Amazonica 7, 5157.
Lainson, R. and Shaw, J. J. (1987). Evolution, classification and geographical distribution. In The Leishmaniases in Biology and Medicine: Volume I Biology and Epidemiology, Vol. I (ed. Peters, W. and Killick-Kendrick, R.), pp. 1120. Academic Press Inc, London.
Lobsiger, L., Muller, N., Schweizer, T., Frey, C. F., Wiederkehr, D., Zumkehr, B. and Gottstein, B. (2010). An autochthonous case of cutaneous bovine leishmaniasis in Switzerland. Veterinary Parasitology 169, 408414.
Lopes, A. H., Iovannisci, D., Petrillo-Peixoto, M., McMahon-Pratt, D. and Beverley, S. M. (1990). Evolution of nuclear DNA and the occurrence of sequences related to new small chromosomal DNAs in the trypanosomatid genus Endotrypanum. Molecular and Biochemical Parasitology 40, 151161.
Lopes, A. H. C. S. and McMahon-Pratt, D. (1989). Monoclonal antibodies specific for members of the genus Endotrypanum. Journal of Protozoology 36, 354361.
Maslov, D. A., Yurchenko, V. Y., Jirku, M. and Lukes, J. (2010). Two new species of trypanosomatid parasites isolated from Heteroptera in Costa Rica. Journal of Eukaryotic Microbiology 57, 177188.
Mesnil, F. and Brimont, E. (1908). Sur un h‚matozaire nouveau Endotrypanum n.gen.) d'un ‚dent‚ de Guyane. Comptes Rendus Hebdomadaiarres des Séances de l´Académie des Sciences 65, 581583.
Morio, F., Reynes, J., Dollet, M., Pratlong, F., Dedet, J. P. and Ravel, C. (2008). Isolation of a protozoan parasite genetically related to the insect trypanosomatid Herpetomonas samuelpessoai from a human immunodeficiency virus-positive patient. Journal of Clinical Microbiology 46, 38453847.
Müller, N., Welle, M., Lobsiger, L., Stoffel, M. H., Boghenbor, K. K., Hilbe, M., Gottstein, B., Frey, C. F., Geyer, C. and von Bomhard, W. (2009). Occurrence of Leishmania sp. in cutaneous lesions of horses in Central Europe. Veterinary Parasitology 166, 346351.
Muniz, J. and Medina, H. (1948). Leishmaniose tegumentar do cobaio (Leishmania enriettii n.sp.). O Hospital. Rio de Janeiro 33, 725.
Nicolle, C. (1908). Sur trois cas d'infection splénique infantile corps de Leishman observés en Tunisia. Arch Inst Pasteur Tunis 3, 126.
Noyes, H. A., Arana, B. A., Chance, M. L. and Maingon, R. (1997). The Leishmania hertigi (Kinetoplastida; Trypanosomatidae) complex and the lizard Leishmania: their classification and evidence for a neotropical origin of the Leishmania-Endotrypanum clade. Journal of Eukaryotic Microbiology 44, 511517.
Noyes, H., Pratlong, F., Chance, M., Ellis, J., Lanotte, G. and Dedet, J. P. (2002). A previously unclassified trypanosomatid responsible for human cutaneous lesions in Martinique (French West Indies) is the most divergent member of the genus Leishmania ss. Parasitology 124, 1724.
Pacheco, R. S., Marzochi, M. C. A., Pires, M. Q., Brito, C. M. M., Madeira, M. D. and Barbosa-Santos, E. G. O. (1998). Parasite genotypically related to a monoxeneous trypanosomatid of dog's flea causing opportunistic infection in an HIV positive patient. Memórias do Instituto Oswaldo Cruz 93, 531537.
Podlipaev, S., Votypka, J., Jirku, M., Svobodova, M. and Lukes, J. (2004). Herpetomonas ztiplika n. sp. (Kinetoplastida: Trypanosomatidae): a parasite of the blood-sucking biting midge Culicoides kibunensis Tokunaga, 1937 (Diptera: Ceratopogonidae). Journal of Parasitology 90, 342347.
Poinar, G. Jr. (2008). Leptoconops nosopheris sp. n. (Diptera: Ceratopogonidae) and Paleotrypanosoma burmanicus gen. n., sp. n. (Kinetoplastida: Trypanosomatidae), a biting midge - trypanosome vector association from the Early Cretaceous. Memórias do Instituo Oswaldo Cruz 103, 468471.
Pothirat, T., Tantiworawit, A., Chaiwarith, R., Jariyapan, N., Wannasan, A., Siriyasatien, P., Supparatpinyo, K., Bates, M. D., Kwakye-Nuako, G. and Bates, P. A. (2014). First isolation of Leishmania from Northern Thailand: case report, identification as Leishmania martiniquensis and phylogenetic position within the Leishmania enriettii complex. PLoS Neglected Tropical Diseases 8, e3339.
Reuss, S. M., Dunbar, M. D., Calderwood Mays, M. B., Owen, J. L., Mallicote, M. F., Archer, L. L. and Wellehan, J. F. Jr. (2012). Autochthonous Leishmania siamensis in horse, Florida, USA. Emerging Infectious Diseases 18, 15451547.
Rodriguez-Bonfante, C., Bonfante-Garrido, R., Grimaldi, G. Jr., Momen, H. and Cupolillo, E. (2003). Genotypically distinct Leishmania colombiensis isolates from Venezuela cause both cutaneous and visceral leishmaniasis in humans. Infection, Genetics and Evolution 3, 119124.
Rose, K., Curtis, J., Baldwin, T., Mathis, A., Kumar, B., Sakthianandeswaren, A., Spurck, T., Low Choy, J. and Handman, E. (2004). Cutaneous leishmaniasis in red kangaroos: isolation and characterisation of the causative organisms. International Journal for Parasitology 34, 655664.
Seblova, V., Sadlova, J., Vojtkova, B., Votypka, J., Carpenter, S., Bates, P. A. and Volf, P. (2015). The Biting Midge Culicoides sonorensis (Diptera: Ceratopogonidae) Is Capable of Developing Late Stage Infections of Leishmania enriettii. PLoS Neglected Tropical Diseases 9, e0004060.
Shaw, J. J. (1963). A possible vector of Endotrypanum schaudinni of the Sloth, Choloepus hoffmanni, in Panama. Nature 201, 417418.
Shaw, J. J. (1969). The haemoflagellates of Sloths. London School of Hygiene and Tropical Medicine Memoir no. 13, pp. 132. London School of Hygiene and Tropical Medicine, London, England.
Shaw, J. J. (1981). The behaviour of Endotrypanum schaudinni (Kinetoplastidae:Trypanosomatidae) in three species of laboratory-bred neotropical sandflies (Diptera:Psychodidae) and its influence on the classification of the genus Leishmania. In Parasitological Topics A Presentation Volume to PCC Garnham FRS on the occasion of his 80th Birthday (ed. Canning, E. U.), pp. 232241. Society of Protozoologists Special Publication, Allen Press Inc, Lawrence Kansas.
Shaw, J. J. (1997). Ecological and evolutionary pressures on leishmanial parasites. Brazilian Journal of Genetics 20, 123128.
Silva, D. A., Madeira Mde, F., Barbosa Filho, C. J., Schubach, E. Y., Barros, J. H. and Figueiredo, F. B. (2013). Leishmania (Leishmania) hertigi in a porcupine (Coendou sp.) found in Brasilia, Federal District, Brazil. Revista Brasileira de Parasitologia Veterinária 22, 297299.
Stamatakis, A. (2006). RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22, 26882690.
Sukmee, T., Siripattanapipong, S., Mungthin, M., Worapong, J., Rangsin, R., Samung, Y., Kongkaew, W., Bumrungsana, K., Chanachai, K., Apiwathanasorn, C., Rujirojindakul, P., Wattanasri, S., Ungchusak, K. and Leelayoova, S. (2008). A suspected new species of Leishmania, the causative agent of visceral leishmaniasis in a Thai patient. International Journal for Parasitology 38, 617622.
Swofford, D. L. (2002). Phylogenetic Analysis using Parsimony (* and Other Methods). Version 4. pp. 197. Sinauer and Associates, Sunderland, MA, USA.
Teixeira, M. M., Borghesan, T. C., Ferreira, R. C., Santos, M. A., Takata, C. S., Campaner, M., Nunes, V. L., Milder, R. V., de Souza, W. and Camargo, E. P. (2011). Phylogenetic validation of the genera Angomonas and Strigomonas of trypanosomatids harboring bacterial endosymbionts with the description of new species of trypanosomatids and of proteobacterial symbionts. Protist 162, 503524.
Thompson, J. D., Gibson, T. J., Plewniak, F., Jeanmougin, F. and Higgins, D. G. (1997). The CLUSTAL_X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 25, 48764882.
Vianna, G. (1911). Söbre uma nova especie de Leishmania (Nota Preliminar). O Brazil Médico 25, 411.
Yakimoff, W. L. and Schokhor, N. I. (1914). Récherches sur les maladies tropicales humaines et animales au Turkestan - II. La leishmaniose cutannée spontanée du chien au Turkestan. Bulletin de la Société de Pathologie Exotique 7, 186187.
Yurchenko, V. Y., Lukes, J., Jirku, M., Zeledon, R. and Maslov, D. A. (2006). Leptomonas costaricensis sp. n. (Kinetoplastea: Trypanosomatidae), a member of the novel phylogenetic group of insect trypanosomatids closely related to the genus Leishmania. Parasitology 133, 537546.
Zeledon, R., Ponce, C. and de Ponce, E. (1977). Finding of Leishmania hertigi in the Costa Rican porcupine. Journal of Parasitology 63, 924925.
Zidkova, L., Cepicka, I., Votypka, J. and Svobodova, M. (2010). Herpetomonas trimorpha sp. nov. (Trypanosomatidae, Kinetoplastida), a parasite of the biting midge Culicoides truncorum (Ceratopogonidae, Diptera). International Journal of Systematic and Evolutionary Microbiology 60, 22362246.


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An appraisal of the taxonomy and nomenclature of trypanosomatids presently classified as Leishmania and Endotrypanum

  • O. A. ESPINOSA (a1), M. G. SERRANO (a2), E. P. CAMARGO (a1), M. M. G. TEIXEIRA (a1) and J. J. SHAW (a1)...


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