{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T11:46:26Z","timestamp":1791287186289,"version":"4.1.0"},"reference-count":45,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2021,4,14]],"date-time":"2021-04-14T00:00:00Z","timestamp":1618358400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2021,4,14]],"date-time":"2021-04-14T00:00:00Z","timestamp":1618358400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100007601","name":"Horizon 2020","doi-asserted-by":"publisher","award":["773830"],"award-info":[{"award-number":["773830"]}],"id":[{"id":"10.13039\/501100007601","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["DFG Research Training Group (GRK) 2046"],"award-info":[{"award-number":["DFG Research Training Group (GRK) 2046"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100011567","name":"Messerli-Stiftung","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100011567","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Friedrich-Loeffler-Institut, Bundesforschungsinstitut f\u00fcr Tiergesundheit"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Parasites Vectors"],"published-print":{"date-parts":[[2021,12]]},"abstract":"Abstract<\/jats:title>\n \n Background<\/jats:title>\n \n Besnoitia darlingi<\/jats:italic>\n ,\n B. neotomofelis<\/jats:italic>\n and\n B. oryctofelisi<\/jats:italic>\n are closely related coccidian parasites with felids as definitive hosts. These parasites use a variety of animal species as intermediate hosts. North American opossums (\n Didelphis virginiana<\/jats:italic>\n ), North American southern plains woodrats (\n Neotoma micropus<\/jats:italic>\n ) and South American domestic rabbits (\n Oryctolagus cuniculus<\/jats:italic>\n ) are intermediate hosts of\n B. darlingi<\/jats:italic>\n ,\n B. neotomofelis<\/jats:italic>\n and\n B. oryctofelisi<\/jats:italic>\n , respectively. Based on conserved regions in the internal transcribed spacer-1 (ITS1) sequence of the ribosomal DNA (rDNA), a real-time PCR for a sensitive detection of these\n Besnoitia<\/jats:italic>\n spp. in tissues of intermediate hosts and faeces of definitive hosts has recently been established. Available sequence data suggest that species such as\n B. akodoni<\/jats:italic>\n and\n B. jellisoni<\/jats:italic>\n are also covered by this real-time PCR. It has been hypothesised that additional\n Besnoitia<\/jats:italic>\n spp. exist worldwide that are closely related to\n B. darlingi<\/jats:italic>\n or\n B. darlingi<\/jats:italic>\n -like parasites (\n B. neotomofelis<\/jats:italic>\n ,\n B. oryctofelisi<\/jats:italic>\n ,\n B. akodoni<\/jats:italic>\n or\n B. jellisoni<\/jats:italic>\n ). Also\u00a0related, but not as\u00a0closely, is\n B. besnoiti<\/jats:italic>\n , the cause of bovine besnoitiosis.\n <\/jats:p>\n <\/jats:sec>\n \n Methods<\/jats:title>\n \n Faecal samples from two free-ranging cheetahs (\n Acinonyx jubatus<\/jats:italic>\n ) from Namibia that had previously tested positive for coccidian parasites by coproscopy were used for this study. A conventional PCR verified the presence of coccidian parasite DNA. To clarify the identity of these coccidia, the faecal DNA samples were further characterised by species-specific PCRs and Sanger sequencing.\n <\/jats:p>\n <\/jats:sec>\n \n Results<\/jats:title>\n \n One of the samples tested positive for\n B. darlingi<\/jats:italic>\n or\n B. darlingi<\/jats:italic>\n -like parasites by real-time PCR, while no other coccidian parasites, including\n Toxoplasma gondii<\/jats:italic>\n ,\n Hammondia hammondi<\/jats:italic>\n ,\n H. heydorni<\/jats:italic>\n ,\n B. besnoiti<\/jats:italic>\n and\n Neospora caninum<\/jats:italic>\n , were detected in the two samples. The rDNA of the\n B. darlingi<\/jats:italic>\n -like parasite was amplified and partially sequenced. Comparison with existing sequences in GenBank revealed a close relationship to other\n Besnoitia<\/jats:italic>\n spp., but also showed clear divergences.\n <\/jats:p>\n <\/jats:sec>\n \n Conclusions<\/jats:title>\n \n Our results suggest that a so far unknown\n Besnoitia<\/jats:italic>\n species exists in Namibian wildlife, which is closely related to\n B. darlingi<\/jats:italic>\n ,\n B. neotomofelis<\/jats:italic>\n ,\n B. oryctofelisi<\/jats:italic>\n ,\n B. akodoni<\/jats:italic>\n or\n B. jellisoni.<\/jats:italic>\n The cheetah appears to be the definitive host of this newly discovered parasite, while prey species of the cheetah may act as intermediate hosts.\n <\/jats:p>\n <\/jats:sec>\n \n Graphical Abstract<\/jats:title>\n <\/jats:sec>","DOI":"10.1186\/s13071-021-04697-3","type":"journal-article","created":{"date-parts":[[2021,4,14]],"date-time":"2021-04-14T16:34:50Z","timestamp":1618418090000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":7,"title":["Molecular analysis suggests that Namibian cheetahs (Acinonyx jubatus) are definitive hosts of a so far undescribed Besnoitia species"],"prefix":"10.1186","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3217-289X","authenticated-orcid":false,"given":"Gereon","family":"Schares","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7855-8579","authenticated-orcid":false,"given":"Maike","family":"Joeres","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6530-8645","authenticated-orcid":false,"given":"Franziska","family":"Rachel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mareen","family":"Tuschy","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9488-0069","authenticated-orcid":false,"given":"G\u00e1bor \u00c1.","family":"Czirj\u00e1k","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9457-0658","authenticated-orcid":false,"given":"Pavlo","family":"Maksimov","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7400-9409","authenticated-orcid":false,"given":"Franz J.","family":"Conraths","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0414-2298","authenticated-orcid":false,"given":"Bettina","family":"Wachter","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2021,4,14]]},"reference":[{"key":"4697_CR1","first-page":"521","volume":"126","author":"JP Dubey","year":"2003","unstructured":"Dubey JP, Sreekumar C, Lindsay DS, Hill D, Rosenthal BM, Venturini L, et al. Besnoitia oryctofelisi n. sp. (Protozoa: Apicomplexa) from domestic rabbits. Parasitology. 2003;126:521\u201339.","journal-title":"Parasitology"},{"key":"4697_CR2","doi-asserted-by":"publisher","first-page":"1731","DOI":"10.1017\/S0031182010000636","volume":"137","author":"JP Dubey","year":"2010","unstructured":"Dubey JP, Yabsley MJ. Besnoitia neotomofelis n. sp. (Protozoa: Apicomplexa) from the southern plains woodrat (Neotoma micropus). Parasitology. 2010;137:1731\u201347.","journal-title":"Parasitology"},{"key":"4697_CR3","doi-asserted-by":"publisher","first-page":"1066","DOI":"10.2307\/3279847","volume":"63","author":"DD Smith","year":"1977","unstructured":"Smith DD, Frenkel JK. Besnoitia darlingi (Protozoa: Toxoplasmatinae): cyclic transmission by cats. J Parasitol. 1977;63:1066\u201371.","journal-title":"J Parasitol"},{"key":"4697_CR4","doi-asserted-by":"publisher","first-page":"584","DOI":"10.1111\/j.1550-7408.1984.tb05510.x","volume":"31","author":"DD Smith","year":"1984","unstructured":"Smith DD, Frenkel JK. Besnoitia darlingi (Apicomplexa, Sarcocystidae, Toxoplasmatinae): transmission between opossums and cats. J Protozool. 1984;31:584\u20137.","journal-title":"J Protozool"},{"key":"4697_CR5","doi-asserted-by":"publisher","first-page":"84","DOI":"10.1016\/j.vetpar.2017.10.013","volume":"248","author":"SK Verma","year":"2017","unstructured":"Verma SK, Cerqueira-Cezar CK, Murata FHA, Lovallo MJ, Rosenthal BM, Dubey JP. Bobcats (Lynx rufus) are natural definitive host of Besnoitia darlingi. Vet Parasitol. 2017;248:84\u20139.","journal-title":"Vet Parasitol"},{"key":"4697_CR6","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1016\/S0304-4017(02)00156-5","volume":"107","author":"L Venturini","year":"2002","unstructured":"Venturini L, Petruccelli M, Piscopo M, Unzaga JM, Venturini MC, Bacigalupe D, et al. Natural Besnoitia sp. infection in domestic rabbits from Argentina. Vet Parasitol. 2002;107:273\u20138.","journal-title":"Vet Parasitol"},{"key":"4697_CR7","first-page":"61","volume":"45","author":"JP Dubey","year":"2003","unstructured":"Dubey JP, Sreekumar C, Rosenthal BM, Lindsay DS, Grisard EC, Vitor RW. Biological and molecular characterization of Besnoitia akodoni n. sp. (Protozoa: Apicomplexa) from the rodent Akodon montensis in Brazil. Parassitologia. 2003;45:61\u201370.","journal-title":"Parassitologia"},{"key":"4697_CR8","doi-asserted-by":"publisher","first-page":"545","DOI":"10.2307\/3277081","volume":"54","author":"JV Ernst","year":"1968","unstructured":"Ernst JV, Chobotar B, Oaks EC, Hammond DM. Besnoitia jellisoni (Sporozoa: Toxoplasmea) in rodents from Utah and California. J Parasitol. 1968;54:545\u20139.","journal-title":"J Parasitol"},{"key":"4697_CR9","first-page":"426","volume":"5","author":"JK Frenkel","year":"1953","unstructured":"Frenkel JK. Infections with organisms resembling Toxoplasma, together with the description of a new organism: Besnoitia jellisoni. Atti del VI Congr Intern Microbiol. 1953;5:426\u201337.","journal-title":"Atti del VI Congr Intern Microbiol"},{"key":"4697_CR10","doi-asserted-by":"publisher","first-page":"611","DOI":"10.2307\/3279560","volume":"63","author":"JK Frenkel","year":"1977","unstructured":"Frenkel JK. Besnoitia wallacei of cats and rodents: with a reclassification of other cyst-forming isosporoid coccidia. J Parasitol. 1977;63:611\u201328.","journal-title":"J Parasitol"},{"key":"4697_CR11","doi-asserted-by":"publisher","first-page":"369","DOI":"10.1126\/science.804183","volume":"188","author":"GD Wallace","year":"1975","unstructured":"Wallace GD, Frenkel JK. Besnoitia species (Protozoa, Sporozoa, Toxoplasmatidae): recognition of cyclic transmission by cats. Science. 1975;188:369\u201371.","journal-title":"Science"},{"key":"4697_CR12","first-page":"69","volume":"18","author":"S Ito","year":"1978","unstructured":"Ito S, Tsunoda K, Shimura K. Life cycle of the large type of Isospora bigemina of the cat. Natl Inst Anim Health Q (Tokyo). 1978;18:69\u201382.","journal-title":"Natl Inst Anim Health Q (Tokyo)"},{"key":"4697_CR13","doi-asserted-by":"publisher","first-page":"173","DOI":"10.1007\/BF00925462","volume":"61","author":"RW Mason","year":"1980","unstructured":"Mason RW. The discovery of Besnoitia wallacei in Australia and the identification of a free-living intermediate host. Z Parasitenkd. 1980;61:173\u20138.","journal-title":"Z Parasitenkd"},{"key":"4697_CR14","doi-asserted-by":"publisher","first-page":"120","DOI":"10.1080\/00480169.1980.34716","volume":"28","author":"PB McKenna","year":"1980","unstructured":"McKenna PB, Charleston WA. Coccidia (Protozoa: Sporozoasida) of cats and dogs. III. The occurrence of a species of Besnoitia in cats. N Z Vet J. 1980;28:120\u20132.","journal-title":"N Z Vet J"},{"key":"4697_CR15","doi-asserted-by":"publisher","first-page":"203","DOI":"10.1016\/0304-4017(94)90111-2","volume":"52","author":"CJ Ng'ang'a","year":"1994","unstructured":"Ng\u2019ang\u2019a CJ, Kanyari PW, Munyua WK. Isolation of Besnoitia wallacei in Kenya. Vet Parasitol. 1994;52:203\u20136.","journal-title":"Vet Parasitol"},{"key":"4697_CR16","doi-asserted-by":"publisher","first-page":"114","DOI":"10.1016\/j.ijppaw.2020.01.011","volume":"11","author":"G Schares","year":"2020","unstructured":"Schares G, Dubey JP, Rosenthal B, Tuschy M, B\u00e4rwald A, Conraths FJ. Sensitive, quantitative detection of Besnoitia darlingi and related parasites in intermediate hosts and to assess felids as definitive hosts for known and as-yet undescribed related parasite species. Int J Parasitol Parasites Wildl. 2020;11:114\u20139.","journal-title":"Int J Parasitol Parasites Wildl"},{"key":"4697_CR17","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1186\/s13071-019-3871-3","volume":"13","author":"A Seltmann","year":"2020","unstructured":"Seltmann A, Schares G, Aschenborn OHK, Heinrich SK, Thalwitzer S, Wachter B, et al. Species-specific differences in Toxoplasma gondii, Neospora caninum and Besnoitia besnoiti seroprevalence in Namibian wildlife. Parasites Vectors. 2020;13:7.","journal-title":"Parasites Vectors"},{"key":"4697_CR18","doi-asserted-by":"publisher","first-page":"407","DOI":"10.1016\/j.pt.2013.06.002","volume":"29","author":"G Alvarez-Garcia","year":"2013","unstructured":"Alvarez-Garcia G, Frey CF, Mora LM, Schares G. A century of bovine besnoitiosis: an unknown disease re-emerging in Europe. Trends Parasitol. 2013;29:407\u201315.","journal-title":"Trends Parasitol"},{"key":"4697_CR19","doi-asserted-by":"publisher","first-page":"851","DOI":"10.1007\/s00436-018-6190-2","volume":"118","author":"A Seltmann","year":"2019","unstructured":"Seltmann A, Webster F, Martins Ferreira SC, Czirj\u00e1k G\u00c1, Wachter B. Age-specific gastrointestinal parasite shedding in free-ranging cheetahs (Acinonyx jubatus) on Namibian farmland. Parasitol Res. 2019;118:851\u20139.","journal-title":"Parasitol Res"},{"key":"4697_CR20","doi-asserted-by":"publisher","first-page":"44837","DOI":"10.1038\/srep44837","volume":"7","author":"SK Heinrich","year":"2017","unstructured":"Heinrich SK, Hofer H, Courtiol A, Melzheimer J, Dehnhard M, Czirj\u00e1k G\u00c1, et al. Cheetahs have a stronger constitutive innate immunity than leopards. Sci Rep. 2017;7:44837.","journal-title":"Sci Rep"},{"key":"4697_CR21","doi-asserted-by":"publisher","first-page":"1203","DOI":"10.1128\/jcm.34.5.1203-1208.1996","volume":"34","author":"MSY Ho","year":"1996","unstructured":"Ho MSY, Barr BC, Marsh AE, Anderson ML, Rowe JD, Tarantal AF, et al. Identification of bovine Neospora parasites by PCR amplification and specific small-subunit rRNA sequence probe hybridization. J Clin Microbiol. 1996;34:1203\u20138.","journal-title":"J Clin Microbiol"},{"key":"4697_CR22","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1186\/s13071-020-04571-8","volume":"14","author":"G Schares","year":"2021","unstructured":"Schares G, Globokar Vrhovec M, Tuschy M, Joeres M, B\u00e4rwald A, Koudela B, et al. A real-time quantitative polymerase chain reaction for the specific detection of Hammondia hammondi and its differentiation from Toxoplasma gondii. Parasites Vectors. 2021;14:78.","journal-title":"Parasites Vectors"},{"key":"4697_CR23","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1053\/tvjl.2001.0599","volume":"163","author":"JR Slapeta","year":"2002","unstructured":"Slapeta JR, Koudela B, Votypka J, Modry D, Horejs R, Lukes J. Coprodiagnosis of Hammondia heydorni in dogs by PCR based amplification of ITS 1 rRNA: differentiation from morphologically indistinguishable oocysts of Neospora caninum. Vet J. 2002;163:147\u201354.","journal-title":"Vet J"},{"key":"4697_CR24","doi-asserted-by":"publisher","first-page":"1525","DOI":"10.1016\/j.ijpara.2005.08.008","volume":"35","author":"G Schares","year":"2005","unstructured":"Schares G, Pantchev N, Barutzki D, Heydorn AO, Bauer C, Conraths FJ. Oocysts of Neospora caninum, Hammondia heydorni, Toxoplasma gondii and Hammondia hammondi in faeces collected from dogs in Germany. Int J Parasitol. 2005;35:1525\u201337.","journal-title":"Int J Parasitol"},{"key":"4697_CR25","doi-asserted-by":"publisher","first-page":"187","DOI":"10.1017\/S0031182005008814","volume":"132","author":"J Abel","year":"2006","unstructured":"Abel J, Schares G, Orzeszko K, Gasser RB, Ellis JT. Hammondia isolated from dogs and foxes are genetically distinct. Parasitology. 2006;132:187\u201392.","journal-title":"Parasitology"},{"key":"4697_CR26","doi-asserted-by":"publisher","first-page":"808","DOI":"10.1007\/s004360100445","volume":"87","author":"G Schares","year":"2001","unstructured":"Schares G, Heydorn AO, C\u00fcppers A, Conraths FJ, Mehlhorn H. Hammondia heydorni-like oocysts shed by a naturally infected dog and Neospora caninum NC-1 cannot be distinguished. Parasitol Res. 2001;87:808\u201316.","journal-title":"Parasitol Res"},{"key":"4697_CR27","doi-asserted-by":"publisher","first-page":"303","DOI":"10.1017\/S0031182010001265","volume":"138","author":"B Gjerde","year":"2011","unstructured":"Gjerde B, Dahlgren SS. Hammondia triffittae n. comb. of foxes (Vulpes spp.): biological and molecular characteristics and differentiation from Hammondia heydorni of dogs. Parasitology. 2011;138:303\u201321.","journal-title":"Parasitology"},{"key":"4697_CR28","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1016\/j.vetpar.2009.04.033","volume":"163","author":"G Schares","year":"2009","unstructured":"Schares G, Basso W, Majzoub M, Cortes HC, Rostaher A, Selmair J, et al. First in vitro isolation of Besnoitia besnoiti from chronically infected cattle in Germany. Vet Parasitol. 2009;163:315\u201322.","journal-title":"Vet Parasitol"},{"key":"4697_CR29","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1016\/0020-7519(96)00030-6","volume":"26","author":"S Payne","year":"1996","unstructured":"Payne S, Ellis J. Detection of Neospora caninum DNA by the polymerase chain reaction. Int J Parasitol. 1996;26:347\u201351.","journal-title":"Int J Parasitol"},{"key":"4697_CR30","doi-asserted-by":"publisher","first-page":"2131","DOI":"10.1128\/JCM.00128-09","volume":"47","author":"H Talabani","year":"2009","unstructured":"Talabani H, Asseraf M, Yera H, Delair E, Ancelle T, Thulliez P, et al. Contributions of immunoblotting, real-time PCR, and the Goldmann-Witmer coefficient to diagnosis of atypical toxoplasmic retinochoroiditis. J Clin Microbiol. 2009;47:2131\u20135.","journal-title":"J Clin Microbiol"},{"key":"4697_CR31","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1111\/vde.12273","volume":"27","author":"S Legnani","year":"2016","unstructured":"Legnani S, Pantchev N, Forlani A, Zini E, Schares G, Balzer J, et al. Emergence of cutaneous neosporosis in a dog receiving immunosuppressive therapy: molecular identification and management. Vet Dermatol. 2016;27:49-e14.","journal-title":"Vet Dermatol"},{"key":"4697_CR32","doi-asserted-by":"publisher","first-page":"208","DOI":"10.1016\/j.vetpar.2011.01.038","volume":"178","author":"G Schares","year":"2011","unstructured":"Schares G, Maksimov A, Basso W, More G, Dubey JP, Rosenthal B, et al. Quantitative real time polymerase chain reaction assays for the sensitive detection of Besnoitia besnoiti infection in cattle. Vet Parasitol. 2011;178:208\u201316.","journal-title":"Vet Parasitol"},{"key":"4697_CR33","first-page":"148","volume":"124","author":"EM Constantin","year":"2011","unstructured":"Constantin EM, Schares G, Grossmann E, Sauter K, Romig T, Hartmann S. Studies on the role of the red fox (Vulpes vulpes) as a potential definitive host of Neospora caninum. Berl Munch Tierarztl Wochenschr. 2011;124:148\u201353.","journal-title":"Berl Munch Tierarztl Wochenschr."},{"key":"4697_CR34","doi-asserted-by":"publisher","first-page":"200","DOI":"10.1016\/j.jviromet.2006.05.020","volume":"136","author":"B Hoffmann","year":"2006","unstructured":"Hoffmann B, Depner K, Schirrmeier H, Beer M. A universal heterologous internal control system for duplex real-time RT-PCR assays used in a detection system for pestiviruses. J Virol Methods. 2006;136:200\u20139.","journal-title":"J Virol Methods"},{"key":"4697_CR35","doi-asserted-by":"crossref","DOI":"10.1093\/oso\/9780195135848.001.0001","volume-title":"Molecular evolution and phylogenetics","author":"M Nei","year":"2000","unstructured":"Nei M, Kumar S. Molecular evolution and phylogenetics. New York: Oxford University Press; 2000."},{"key":"4697_CR36","doi-asserted-by":"publisher","first-page":"1547","DOI":"10.1093\/molbev\/msy096","volume":"35","author":"S Kumar","year":"2018","unstructured":"Kumar S, Stecher G, Li M, Knyaz C, Tamura K. MEGA X: molecular evolutionary genetics analysis across computing platforms. Mol Biol Evol. 2018;35:1547\u20139.","journal-title":"Mol Biol Evol"},{"key":"4697_CR37","volume-title":"Toxoplasmosis of animals and humans","author":"JP Dubey","year":"2010","unstructured":"Dubey JP. Toxoplasmosis of animals and humans. 2nd ed. Boca Raton: CRC Press; 2010.","edition":"2"},{"key":"4697_CR38","first-page":"265","volume":"28","author":"JW Pols","year":"1960","unstructured":"Pols JW. Studies on bovine besnoitiosis with special reference to the aetiology. Onderstepoort J Vet Res. 1960;28:265\u2013356.","journal-title":"Onderstepoort J Vet Res"},{"key":"4697_CR39","first-page":"7","volume":"34","author":"RD Bigalke","year":"1967","unstructured":"Bigalke RD, van Niekerk JW, Basson PA, McCully RM. Studies on the relationship between Besnoitia of blue wildebeest and impala, and Besnoitia besnoiti of cattle. Onderstepoort J Vet Res. 1967;34:7\u201328.","journal-title":"Onderstepoort J Vet Res"},{"key":"4697_CR40","doi-asserted-by":"publisher","DOI":"10.1201\/9781315152561","volume-title":"Neosporosis in animals","author":"JP Dubey","year":"2017","unstructured":"Dubey JP, Hemphill A, Calero-Bernal R, Schares G. Neosporosis in animals. Boca Rotan: CRC Press; 2017."},{"key":"4697_CR41","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/0378-1119(92)90570-F","volume":"114","author":"JM Guay","year":"1992","unstructured":"Guay JM, Huot A, Gagnon S, Tremblay A, Levesque RC. Physical and genetic mapping of cloned ribosomal DNA from Toxoplasma gondii: primary and secondary structure of the 5S gene. Gene. 1992;114:165\u201371.","journal-title":"Gene"},{"key":"4697_CR42","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1017\/pao.2015.7","volume":"2","author":"ME Ogedengbe","year":"2016","unstructured":"Ogedengbe ME, Ogedengbe JD, Whale JCE, K., Juarez-Estrada MA, Barta JR, . Molecular phylogenetic analyses of tissue coccidia (sarcocystidae; apicomplexa) based on nuclear 18s RDNA and mitochondrial COI sequences confirms the paraphyly of the genus Hammondia. Parasitology Open. 2016;2:1\u201316.","journal-title":"Parasitology Open"},{"key":"4697_CR43","volume-title":"Cheetahs of the Serengeti plains: group living in an asocial species","author":"TM Caro","year":"1994","unstructured":"Caro TM. Cheetahs of the Serengeti plains: group living in an asocial species. Chicago: The University Press; 1994."},{"key":"4697_CR44","doi-asserted-by":"publisher","first-page":"1564","DOI":"10.1016\/j.meegid.2011.08.006","volume":"11","author":"P Olias","year":"2011","unstructured":"Olias P, Schade B, Mehlhorn H. Molecular pathology, taxonomy and epidemiology of Besnoitia species (Protozoa: Sarcocystidae). Infect Genet Evol. 2011;11:1564\u201376.","journal-title":"Infect Genet Evol"},{"key":"4697_CR45","doi-asserted-by":"publisher","first-page":"e1000436","DOI":"10.1371\/journal.pbio.1000436","volume":"8","author":"MA Nilsson","year":"2010","unstructured":"Nilsson MA, Churakov G, Sommer M, Tran NV, Zemann A, Brosius J, et al. Tracking marsupial evolution using archaic genomic retroposon insertions. PLoS Biol. 2010;8:e1000436.","journal-title":"PLoS Biol"}],"container-title":["Parasites & Vectors"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13071-021-04697-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s13071-021-04697-3\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13071-021-04697-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,8,28]],"date-time":"2024-08-28T05:43:16Z","timestamp":1724823796000},"score":1,"resource":{"primary":{"URL":"https:\/\/parasitesandvectors.biomedcentral.com\/articles\/10.1186\/s13071-021-04697-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,4,14]]},"references-count":45,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2021,12]]}},"alternative-id":["4697"],"URL":"https:\/\/doi.org\/10.1186\/s13071-021-04697-3","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.3.rs-235843\/v1","asserted-by":"object"}]},"ISSN":["1756-3305"],"issn-type":[{"value":"1756-3305","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,4,14]]},"assertion":[{"value":"12 February 2021","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"22 March 2021","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 April 2021","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"All experimental procedures, including animal immobilisation and sample collection, were approved by the Internal Ethics Committee of the Leibniz Institute for Zoo and Wildlife Research (Leibniz-IZW) (permit number 2002-04-01) and authorised by the Ministry of Environment, Forestry and Tourism of Namibia (permit numbers 1514\/2011 and 1689\/2012). Samples were transported to Germany in full compliance with the Convention on International Trade in Endangered Species (CITES). All experiments were carried out in compliance with the approved guidelines of the IZW and the laws of Germany and Namibia.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare that they have no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"201"}}