{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T11:09:38Z","timestamp":1785409778118,"version":"3.56.0"},"reference-count":85,"publisher":"MDPI AG","issue":"7","license":[{"start":{"date-parts":[[2026,6,25]],"date-time":"2026-06-25T00:00:00Z","timestamp":1782345600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100006769","name":"Russian Science Foundation","doi-asserted-by":"publisher","award":["25-74-10071"],"award-info":[{"award-number":["25-74-10071"]}],"id":[{"id":"10.13039\/501100006769","id-type":"DOI","asserted-by":"publisher"}]},{"award":["25-74-10071"],"award-info":[{"award-number":["25-74-10071"]}],"id":[{"id":"https:\/\/ror.org\/03y2gwe85","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Pathogens"],"abstract":"<jats:p>The subgenus Megatrypanum Hoare, 1964, with the type species Trypanosoma theileri Laveran, 1902, comprises stercorarian trypanosomes of mammals. A substantial portion of this subgenus consists of T. theileri-like trypanosomes parasitizing cervids and bovids worldwide. Similar to most other members of the genus Trypanosoma that lack obvious economic importance, the biology of T. theileri-like trypanosomes remains poorly understood. In particular, fundamental aspects such as their host specificity, host\u2013parasite interactions, and the morphology of developmental stages have been studied only to a limited extent. In this work, we provide a detailed description of the development and cellular organization of T. theileri-like trypanosomes in the horsefly Hybomitra montana using transmission electron microscopy (TEM). We show for the first time that T. theileri-like trypanosomes possess a well-developed cytostome\u2013cytopharyngeal complex, morphologically similar to those in other stercorarian trypanosomes. This complex is present in the studied trypanosomes at the epimastigote stage and degrades during metacyclogenesis. In the host ileum, epimastigotes and trypomastigotes at different stages of metacyclogenesis are embedded in a fibrillar matrix that isolates them from the gut lumen. This promotes their accumulation in the vector, thereby increasing the efficiency of future infection of the vertebrate host, which occurs via contamination of the oral mucosa.<\/jats:p>","DOI":"10.3390\/pathogens15070668","type":"journal-article","created":{"date-parts":[[2026,7,7]],"date-time":"2026-07-07T10:04:47Z","timestamp":1783418687000},"page":"668","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Developmental and Ultrastructural Characterization of Trypanosoma theileri-like Flagellates in a Horsefly Hybomitra montana"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1444-3104","authenticated-orcid":false,"given":"Alexander O.","family":"Frolov","sequence":"first","affiliation":[{"name":"Zoological Institute of the Russian Academy of Sciences, 199034 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1663-7706","authenticated-orcid":false,"given":"Anna I.","family":"Solovyeva","sequence":"additional","affiliation":[{"name":"Institute of Cytology of the Russian Academy of Sciences, 194064 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marina N.","family":"Malysheva","sequence":"additional","affiliation":[{"name":"Zoological Institute of the Russian Academy of Sciences, 199034 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0854-0621","authenticated-orcid":false,"given":"Maria E.","family":"Belokon","sequence":"additional","affiliation":[{"name":"Zoological Institute of the Russian Academy of Sciences, 199034 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Grigory N.","family":"Machakhtyrov","sequence":"additional","affiliation":[{"name":"Academy of Sciences of the Republic of Sakha (Yakutia), 677007 Yakutsk, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0988-0943","authenticated-orcid":false,"given":"Varvara A.","family":"Machakhtyrova","sequence":"additional","affiliation":[{"name":"Academy of Sciences of the Republic of Sakha (Yakutia), 677007 Yakutsk, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anatoly A.","family":"Bondarev","sequence":"additional","affiliation":[{"name":"Academy of Sciences of the Republic of Sakha (Yakutia), 677007 Yakutsk, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-9795-344X","authenticated-orcid":false,"given":"Maria S.","family":"Maximova","sequence":"additional","affiliation":[{"name":"Zoological Institute of the Russian Academy of Sciences, 199034 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anna I.","family":"Ganyukova","sequence":"additional","affiliation":[{"name":"Zoological Institute of the Russian Academy of Sciences, 199034 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,6,25]]},"reference":[{"key":"ref_1","unstructured":"Hoare, C.A. (1972). The Trypanosomes of Mammals. A Zoological Monograph, Blackwell Scientific Publications."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1111\/j.1550-7408.1987.tb03143.x","article-title":"Transmission of Megatiypanum trypanosomes to Cervus dama by Tabanidae","volume":"34","author":"Friedhoff","year":"1987","journal-title":"J. Protozool."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"308","DOI":"10.1186\/s13071-020-04169-0","article-title":"Pan-American Trypanosoma (Megatrypanum) trinaperronei n. sp. in the white-tailed deer Odocoileus virginianus Zimmermann and its deer ked Lipoptena mazamae Rondani, 1878: Morphological, developmental and phylogeographical characterization","volume":"13","author":"Garcia","year":"2020","journal-title":"Parasites Vectors"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1111\/j.1550-7408.2011.00599.x","article-title":"Trypanosoma (Megatrypanum) melophagium in the sheep ked Melophagus ovinus from organic farms in Croatia: Phylogenetic inferences support restriction to sheep and sheep keds and close relationship with trypanosomes from other ruminant species","volume":"59","author":"Rodrigues","year":"2012","journal-title":"J. Eukaryot. Microbiol."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Brot\u00e1nkov\u00e1, A., Fialov\u00e1, M., \u010cepi\u010dka, I., Brzo\u0148ov\u00e1, J., and Svobodov\u00e1, M. (2022). Trypanosomes of the Trypanosoma theileri group: Phylogeny and new potential vectors. Microorganisms, 10.","DOI":"10.3390\/microorganisms10020294"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1186\/s13071-022-05212-y","article-title":"Development of two species of the Trypanosoma theileri complex in tabanids","volume":"15","author":"Kostygov","year":"2022","journal-title":"Parasites Vectors"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"200407","DOI":"10.1098\/rsob.200407","article-title":"Euglenozoa: Taxonomy, diversity and ecology, symbioses and viruses","volume":"11","author":"Kostygov","year":"2021","journal-title":"Open Biol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1016\/j.pt.2023.11.009","article-title":"Phylogenetic framework to explore trait evolution in Trypanosomatidae","volume":"40","author":"Kostygov","year":"2024","journal-title":"Trends Parasitol."},{"key":"ref_9","first-page":"65","article-title":"Trypanosoma (Megatrypanum) melophagium: Modes of attachment of parasites to mid-gut, hindgut and rectum of the sheep ked, Meiophagus ovinus","volume":"32","author":"Molyneux","year":"1975","journal-title":"Acta Trop."},{"key":"ref_10","first-page":"319","article-title":"Trypanosoma (Megatrypanum) melophagium in the sheep ked, Melophagus ovinus. A scanning electron microscope (SEM) study of the parasites and the insect gut wall surfaces","volume":"35","author":"Molyneux","year":"1978","journal-title":"Acta Trop."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/0014-4894(65)90005-6","article-title":"Cytoplasmic inclusions and organelles of in vitro cultured Trypanosoma theileri and Trypanosoma melophagium and some speculations on their function","volume":"17","author":"Herbert","year":"1965","journal-title":"Exp. Parasitol."},{"key":"ref_12","first-page":"233","article-title":"Microscopic study of developmental stages of a Trypanosoma theileri-like trypanosome in axenic culture","volume":"17","author":"Malysheva","year":"2023","journal-title":"Protistology"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"223","DOI":"10.21685\/1680-0826-2018-12-4-6","article-title":"First record of Trypanosoma theileri-like flagellates in horseflies from Northwest Russia","volume":"12","author":"Ganyukova","year":"2018","journal-title":"Protistology"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"208","DOI":"10.1083\/jcb.17.1.208","article-title":"The use of lead citrate at high pH as an electron-opaque stain in electron microscopy","volume":"17","author":"Reynolds","year":"1963","journal-title":"J. Cell Biol."},{"key":"ref_15","first-page":"126","article-title":"Leptomonas jaculum (Leger, 1902) Woodcock 1914: A leptomonas or a blastocrithidia?","volume":"41","author":"Kostygov","year":"2007","journal-title":"Parazitologiia"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1393","DOI":"10.1016\/j.ijpara.2004.08.011","article-title":"Trypanosomes are monophyletic: Evidence from genes for glyceraldehyde phosphate dehydrogenase and small subunit ribosomal RNA","volume":"34","author":"Hamilton","year":"2004","journal-title":"Int. J. Parasitol."},{"key":"ref_17","first-page":"294","article-title":"DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates","volume":"3","author":"Folmer","year":"1994","journal-title":"Mol. Mar. Biol. Biotech."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"653","DOI":"10.1136\/vr.132.26.653","article-title":"Clinical disease associated with Trypanosoma theileri infection in a calf in Ireland","volume":"132","author":"Doherty","year":"1993","journal-title":"Vet. Rec."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1136\/vr.150.1.18","article-title":"Trypanosoma theileri in the cerebrospinal fluid and brain of a heifer with suppurative meningoencephalitis","volume":"150","author":"Braun","year":"2002","journal-title":"Vet. Rec."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"941","DOI":"10.12935\/jvma.64.941","article-title":"A case of a Japanese black cow developing trypanosomosis together with enzootic bovine leukosis","volume":"64","author":"Matsumoto","year":"2011","journal-title":"J. Jpn. Vet. Med. Assoc."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"82","DOI":"10.17221\/1580-VETMED","article-title":"Association of Trypanosoma theileri with peritonitis in a pregnant cross-bred cow: A case report","volume":"56","author":"Sood","year":"2011","journal-title":"Vet. Med."},{"key":"ref_22","first-page":"191","article-title":"Hemolytic anemia associated with Trypanosoma theileri in a cow from Kurdistan province, West of Iran","volume":"11","author":"Hajihassani","year":"2020","journal-title":"Vet. Res. Forum."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"102476","DOI":"10.1016\/j.parint.2021.102476","article-title":"Genetic and seasonal variations of Trypanosoma theileri and the association of Trypanosoma theileri infection with dairy cattle productivity in Northern Japan","volume":"86","author":"Suganuma","year":"2022","journal-title":"Parasitol. Int."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"788","DOI":"10.1111\/j.1550-7408.1993.tb04475.x","article-title":"Development of Trypanosoma (M.) theileri in Tabanids","volume":"40","author":"Heister","year":"1993","journal-title":"J. Euk. Mcrobrol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"365","DOI":"10.1017\/S0031182000014888","article-title":"An experimental study of the sheep trypanosome (Trypanosoma melophagium, Flu, 1908) and its transmission by sheep ked (Melophagus ovinus)","volume":"15","author":"Hoare","year":"1923","journal-title":"Parasitology"},{"key":"ref_26","first-page":"421","article-title":"High prevalence of Trypanosoma theileri-like trypanosomes and scarcity of monoxenous trypanosomatids in tabanids of Northwestern Russia","volume":"18","author":"Ganyukova","year":"2024","journal-title":"Protistology"},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Tur\u010dinavi\u010dien\u0117, J., Bernotien\u0117, R., and Petra\u0161i\u016bnas, A. (2024). Molecular detection and analysis of Trypanosoma (Megatrypanum) spp. diversity in Tabanidae (Diptera) collected in Lithuania. Insects, 15.","DOI":"10.3390\/insects15080581"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1155\/2018\/2597074","article-title":"Isolation of a trypanosome related to Trypanosoma theileri (Kinetoplastea: Trypanosomatidae) from Phlebotomus perfiliewi (Diptera: Psychodidae)","volume":"2018","author":"Calzolari","year":"2018","journal-title":"BioMed Res. Int."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1636","DOI":"10.1017\/S0031182021001360","article-title":"Genetic diversity of cervid Trypanosoma theileri in Honshu sika deer (Cervus nippon) in Japan","volume":"148","author":"Rosyadi","year":"2021","journal-title":"Parasitology"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1007\/BF00931269","article-title":"Host-parasite relationship of Trypanosoma corvi in Ornithomya avicularia","volume":"75","author":"Mungomba","year":"1989","journal-title":"Parasitol. Res."},{"key":"ref_31","first-page":"183","article-title":"Ultrastructure of the fibrous matrix surrounding cells of Trypanosoma melophagium in the hind-gut of the sheep ked, Melophagus ovinus","volume":"44","author":"Heywood","year":"1985","journal-title":"Cytobios"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1016\/S0171-9335(99)80036-3","article-title":"Filamentous proteophosphoglycan secreted by Leishmania promastigotes forms gel-like three dimensional networks that obstruct the digestive tract of infected sandfly vectors","volume":"78","author":"Stierhof","year":"1999","journal-title":"Eur. J. Cell Biol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"549","DOI":"10.1590\/S0074-02761997000400020","article-title":"Ultrastructure of the adhesion of Crithidia guilhermei (Kinetoplastida: Trypanosomatidae) to plastic coverslips: Formation of a fibrous matrix around the cells","volume":"92","author":"Oliveira","year":"1997","journal-title":"Mem. Inst. Oswaldo Cruz."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1016\/j.ijpara.2021.06.005","article-title":"First description of Lotmaria passim and Crithidia mellificae haptomonad stages in the honeybee hindgut","volume":"52","author":"Alunda","year":"2022","journal-title":"Int. J. Parasitol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"161","DOI":"10.21685\/1680-0826-2022-16-3-2","article-title":"Crithidia versiformis sp. n. (Kinetoplastea: Trypanosomatida), a parasite of the green lacewing Chrysoperla carnea (Stephens) (Neuroptera, Chrysopidae)","volume":"16","author":"Frolov","year":"2022","journal-title":"Protistology"},{"key":"ref_36","first-page":"318","article-title":"Ultrastructure of Crithidia bombi (Trypanosomatidae) choanomastigotes and haptomorphs with special reference to the flagellar pocket skeleton","volume":"19","author":"Karpov","year":"2025","journal-title":"Protistology"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"538","DOI":"10.1016\/j.pt.2021.02.004","article-title":"Development of monoxenous trypanosomatids and phytomonads in insects","volume":"37","author":"Frolov","year":"2021","journal-title":"Trends Parasitol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1007\/BF00538199","article-title":"T ransmission of Trypanosoma theileri to cattle by Tabanidae","volume":"73","author":"Friedhoff","year":"1987","journal-title":"Parasitol. Res."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"e84552","DOI":"10.7554\/eLife.84552","article-title":"Formation and three- dimensional architecture of Leishmania adhesion in the sand fly vector","volume":"12","author":"Yanase","year":"2023","journal-title":"eLife"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"699","DOI":"10.1038\/ncb0804-699","article-title":"Cytoplasmic intermediate filaments revealed as dynamic and multipurpose scaffolds","volume":"6","author":"Coulombe","year":"2004","journal-title":"Nat. Cell Biol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"2146","DOI":"10.1073\/pnas.1314138111","article-title":"Septin assemblies form by diffusion-driven annealing on membranes","volume":"111","author":"Bridges","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Gupta, C.M., and Thiyagarajan, S. (2025). Structure and functions of actin and actin-binding proteins in Leishmania. Pathogens, 14.","DOI":"10.3390\/pathogens14090948"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1016\/j.bbrc.2019.04.007","article-title":"Trypanosoma cruzi actins: Expression analysis of actin 2","volume":"513","author":"Hernandez","year":"2019","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"2837","DOI":"10.1016\/j.cub.2025.04.064","article-title":"Discovery of a novel flagellar filament system underpinning Leishmania adhesion to surfaces","volume":"35","author":"Owino","year":"2025","journal-title":"Curr. Biol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"6960","DOI":"10.1038\/s41467-024-51291-z","article-title":"Discovery of essential kinetoplastid-insect adhesion proteins and their function in Leishmania-sand flyinteractions","volume":"15","author":"Yanase","year":"2024","journal-title":"Nat. Commun."},{"key":"ref_46","first-page":"87","article-title":"Niche partitioning within an insect host: Trypanosomatids Wallacemonas raviniae and Trypanosoma (Megatrypanum) sp. in the horsefly Hybomitra solstitialis","volume":"16","author":"Malysheva","year":"2022","journal-title":"Protistology"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1017\/S0031182005008929","article-title":"Phylogeny of Trypanosoma (Megatrypanum) theileri and related trypanosomes reveals lineages of isolates associated with artiodactyl hosts diverging on SSU and ITS ribosomal sequences","volume":"132","author":"Rodrigues","year":"2006","journal-title":"Parasitology"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1385","DOI":"10.1016\/j.ijpara.2011.09.001","article-title":"Multilocus phylogeographical analysis of Trypanosoma (Megatrypanum) genotypes from sympatric cattle and water buffalo populations supports evolutionary host constraint and close phylogenetic relationships with genotypes found in other ruminants","volume":"41","author":"Garcia","year":"2011","journal-title":"Int. J. Parasitol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1016\/S0304-4017(03)00236-X","article-title":"Brazilian isolates of Trypanosoma (Megatrypanum) theileri: Diagnosis and differentiation of isolates from cattle and water buffalo based on biological characteristics and randomly amplified DNA sequences","volume":"116","author":"Rodrigues","year":"2003","journal-title":"Vet. Parasitol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1292\/jvms.09-0343","article-title":"Isolation of Trypanosoma (Megatrypanum) theileri from dairy cattle in Taiwan","volume":"72","author":"Lee","year":"2010","journal-title":"J. Vet. Med. Sci."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"560","DOI":"10.1016\/j.pt.2025.05.005","article-title":"Three Hungry Tryps: The efficient endocytic pathway of pathogenic trypanosomatids","volume":"41","author":"Pereira","year":"2025","journal-title":"Trends Parasitol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"320","DOI":"10.1111\/j.1550-7408.1969.tb02278.x","article-title":"The form and function of the cytostome-cytopharynx of the culture forms of the Elasmobranch haemoflagellate Trypanosoma raiae Laveran & Mesnil","volume":"16","author":"Preston","year":"1969","journal-title":"J. Protozool."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"563","DOI":"10.1083\/jcb.8.2.563","article-title":"The existence of a cytostome and the occurrence of pinocytosis in the trypanosome, Trypanosoma mega","volume":"8","author":"Steinert","year":"1960","journal-title":"J. Biophys. Biochem. Cytol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"730","DOI":"10.1111\/j.1550-7408.1969.tb02335.x","article-title":"The cytostome of Trypanosoma cruzi and T. conorhini","volume":"16","author":"Milder","year":"1969","journal-title":"J. Protozool."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1111\/j.1550-7408.1974.tb03646.x","article-title":"Fine structure of Trypanosoma cyclops in noncellular cultures","volume":"21","author":"Heywood","year":"1974","journal-title":"J. Protozool."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1712","DOI":"10.1139\/z76-198","article-title":"The ultrastructure of the epimastigote stages of Trypanosoma rotatorium in the leech Batracobdella picta","volume":"54","author":"Desser","year":"1976","journal-title":"Can. J. Zool."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1111\/j.1550-7408.1984.tb02990.x","article-title":"Trypanosoma lucknowi, a new species of trypanosome from Macaca mulatta with observations on its fine structure","volume":"31","author":"Weinman","year":"1984","journal-title":"J. Protozool."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1007\/BF00934379","article-title":"Development of Trypanosoma fallisi in the leech, Desserobdella picta, in toads (Bufo americanus), and in vitro. A light and electron microscopic study","volume":"77","author":"Martin","year":"1991","journal-title":"Parasitol. Res."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"594","DOI":"10.1111\/j.1550-7408.2009.00444.x","article-title":"Phylogenetic analyses based on small subunit rRNA and glycosomal glyceraldehyde-3-phosphate dehydrogenase genes and ultrastructural characterization of two snake trypanosomes: Trypanosoma serpentis n. sp. from Pseudoboa nigra and Trypanosoma cascavelli from Crotalus durissus terrificus","volume":"56","author":"Viola","year":"2009","journal-title":"J. Eukaryot. Microbiol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1186\/1756-3305-6-313","article-title":"The phylogeography of trypanosomes from South American alligatorids and African crocodilids is consistent with the geological history of South American river basins and the transoceanic dispersal of Crocodylus at the Miocene","volume":"6","author":"Fermino","year":"2013","journal-title":"Parasites Vectors"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1186\/1756-3305-6-221","article-title":"Trypanosoma livingstonei: A new species from African bats supports the bat seeding hypothesis for the Trypanosoma cruzi clade","volume":"6","author":"Lima","year":"2013","journal-title":"Parasites Vectors"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1186\/s13071-015-1193-7","article-title":"Phylogenetic and morphological characterization of trypanosomes from Brazilian armoured catfishes and leeches reveal high species diversity, mixed infections and a new fish trypanosome species","volume":"8","author":"Lemos","year":"2015","journal-title":"Parasites Vectors"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"610","DOI":"10.1111\/jeu.12310","article-title":"Developmental and ultrastructural characterization and phylogenetic analysis of Trypanosoma herthameyeri n. sp of brazilian Leptodactilydae frogs","volume":"63","author":"Attias","year":"2016","journal-title":"J. Eukaryot. Microbiol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1186\/s13071-019-3463-2","article-title":"Shared species of crocodilian trypanosomes carried by tabanid flies in Africa and South America, including the description of a new species from caimans, Trypanosoma kaiowa n. sp","volume":"12","author":"Fermino","year":"2019","journal-title":"Parasites Vectors"},{"key":"ref_65","first-page":"2227","article-title":"The three-dimensional structure of the cytostome-cytopharinx complex of Trypanosoma cruzi epimastigotes","volume":"127","author":"Alcantara","year":"2014","journal-title":"J. Cell Sci."},{"key":"ref_66","first-page":"164","article-title":"The cytostome\u2013cytopharynx complex of Trypanosoma cruzi epimastigotes disassembles during cell division","volume":"130","author":"Alcantara","year":"2017","journal-title":"J. Cell Sci."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"e13346","DOI":"10.1111\/cmi.13346","article-title":"The cytostome-cytopharynx complex of intracellular and extracellular amastigotes of Trypanosoma cruzi exhibit structural and functional differences","volume":"23","author":"Alcantara","year":"2021","journal-title":"Cell. Microbiol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1016\/j.jsb.2016.07.018","article-title":"Loss of the cytostome-cytopharynx and endocytic ability are late events in Trypanosoma cruzi metacyclogenesis","volume":"196","author":"Vidal","year":"2016","journal-title":"J. Struct. Biol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1111\/j.1550-7408.1989.tb05524.x","article-title":"Coated Vesicles from the Protozoan Parasite Trypanosoma brucei: Purification and Characterization","volume":"36","author":"Shapiro","year":"1989","journal-title":"J. Protozool."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"532","DOI":"10.1007\/BF00335057","article-title":"The fine structure of Crithidia fasciculata with special reference to the organelles involved in the ingestion and digestion of protein","volume":"116","author":"Brooker","year":"1971","journal-title":"Z. Zellforsch."},{"key":"ref_71","first-page":"1072","article-title":"Comparative morphology of kinetoplastids","volume":"37","author":"Frolov","year":"1995","journal-title":"Tsitologiia"},{"key":"ref_72","first-page":"159","article-title":"Ultrastructure of the parasitic flagellate Leptomonas nabiculae from the bug Nabicula flavomarginata","volume":"37","author":"Shaglina","year":"1995","journal-title":"Tsitologiia"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"11757","DOI":"10.1073\/pnas.1712311114","article-title":"Extensive flagellar remodeling during the complex life cycle of Paratrypanosoma, an early-branching trypanosomatid","volume":"114","author":"Wheeler","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"e2521233122","DOI":"10.1073\/pnas.2521233122","article-title":"The subcellular architecture of Paratrypanosoma confusum revealed by CryoET: A window into early trypanosome evolution","volume":"122","author":"Alcantara","year":"2025","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0065-308X(01)48003-1","article-title":"The Molecular Evolution of Trypanosomatidae","volume":"48","author":"Stevens","year":"2001","journal-title":"Adv. Parasitol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1016\/j.molbiopara.2015.10.003","article-title":"The origin and evolution of the acidocalcisome and its interactions with other organelles","volume":"209","author":"Docampo","year":"2016","journal-title":"Mol. Biochem. Parasitol."},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"de Souza, W. (2010). The glycosome of trypanosomatids. Structures and Organelles in Pathogenic Protists, Springer.","DOI":"10.1007\/978-3-642-12863-9"},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Qui\u00f1ones, W., Acosta, H., Gon\u00e7alves, C.S., Motta, M.C.M., Gualdr\u00f3n-L\u00f3pez, M., and Michels, P.A.M. (2020). Structure, properties, and function of glycosomes in Trypanosoma cruzi. Front. Cell. Infect. Microbiol., 10.","DOI":"10.3389\/fcimb.2020.00025"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1111\/j.1365-2818.1991.tb03118.x","article-title":"The glycosomes of trypanosomes: Number and distribution as revealed by electron spectroscopic imaging and 3-D reconstruction","volume":"162","author":"Tetley","year":"1991","journal-title":"J. Microsc."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1242\/jcs.198887","article-title":"Patterns of organelle ontogeny through a cell cycle revealed by whole cell reconstructions using 3D electron microscopy","volume":"130","author":"Hughes","year":"2017","journal-title":"J. Cell Sci."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"502","DOI":"10.1111\/j.1550-7408.1975.tb05219.x","article-title":"Fusion of host cell secondary lysosomes with the parasitiphorous vacuoles of Leishmania mexicana-infected macrophages","volume":"22","author":"Alexander","year":"1975","journal-title":"J. Protozool."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"246","DOI":"10.1007\/s00436-003-1027-y","article-title":"Ultrastructural and cytochemical identification of megasome in Leishmania (Leishmania) chagasi","volume":"92","author":"Alberio","year":"2004","journal-title":"Parasitol. Res."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"517","DOI":"10.1007\/BF00932700","article-title":"The reservosomes of epimastigote forms of Trypanosoma cruzi: Occurrence during in vitro cultivation","volume":"80","author":"Figueiredo","year":"1994","journal-title":"Parasitol. Res."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1017\/S1431927604040863","article-title":"Biogenesis of the reservosomes of Trypanosoma cruzi","volume":"10","year":"2004","journal-title":"Microsc. Microanal."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"346","DOI":"10.3390\/cells1030346","article-title":"Autophagy in Trypanosomatids","volume":"1","author":"Brennand","year":"2012","journal-title":"Cells"}],"container-title":["Pathogens"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-0817\/15\/7\/668\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,7]],"date-time":"2026-07-07T11:12:13Z","timestamp":1783422733000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-0817\/15\/7\/668"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,25]]},"references-count":85,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2026,7]]}},"alternative-id":["pathogens15070668"],"URL":"https:\/\/doi.org\/10.3390\/pathogens15070668","relation":{},"ISSN":["2076-0817"],"issn-type":[{"value":"2076-0817","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,25]]}}}