{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T09:38:15Z","timestamp":1788514695499,"version":"build-2803163510"},"reference-count":61,"publisher":"MDPI AG","issue":"10","license":[{"start":{"date-parts":[[2020,9,24]],"date-time":"2020-09-24T00:00:00Z","timestamp":1600905600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Pathogens"],"abstract":"<jats:p>Songbirds widely disperse ticks that carry a diversity of pathogens, some of which are pathogenic to humans. Among ticks commonly removed from songbirds, the blacklegged tick, Ixodes scapularis, can harbor any combination of nine zoonotic pathogens, including Babesia species. From May through September 2019, a total 157 ticks were collected from 93 songbirds of 29 species in the Canadian provinces of Ontario and Qu\u00e9bec. PCR testing for the 18S gene of Babesia species detected Babesia odocoilei in 12.63% of I. scapularis nymphs parasitizing songbirds in Ontario and Qu\u00e9bec; none of the relatively small numbers of Ixodes muris, Ixodes brunneus, or Haemaphysalis leporispalustris were PCR-positive. For ticks at each site, the prevalence of B. odocoilei was 16.67% in Ontario and 8.89% and 5.26% in Qu\u00e9bec. Of 31 live, engorged I. scapularis larvae and nymphs held to molt, 25 ticks completed the molt; five of these molted ticks were positive for B. odocoilei. PCR-positive ticks were collected from six bird species\u2014namely, Common Yellowthroat, Swainson\u2019s Thrush, Veery, House Wren, Baltimore Oriole, and American Robin. Phylogenetic analysis documented the close relationship of B. odocoilei to Babesia canis canis and Babesia divergens, the latter a known pathogen to humans. For the first time in Canada, we confirm the transstadial passage of B. odocoilei in I. scapularis molting from larvae to nymphs. A novel host record reveals I. scapularis on a Palm Warbler. Our findings show that B. odocoilei is present in all mobile life stages of I. scapularis, and it is widely dispersed by songbirds in Ontario and Qu\u00e9bec.<\/jats:p>","DOI":"10.3390\/pathogens9100781","type":"journal-article","created":{"date-parts":[[2020,9,24]],"date-time":"2020-09-24T08:33:03Z","timestamp":1600936383000},"page":"781","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":22,"title":["Detection of Babesia odocoilei in Ixodes scapularis Ticks Collected from Songbirds in Ontario and Quebec, Canada"],"prefix":"10.3390","volume":"9","author":[{"given":"John D.","family":"Scott","sequence":"first","affiliation":[{"name":"Department of Medicine and Epidemiology, School of Veterinary Medicine, University of California Davis, Davis, CA 95616, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3493-3485","authenticated-orcid":false,"given":"Emily L.","family":"Pascoe","sequence":"additional","affiliation":[{"name":"Department of Medicine and Epidemiology, School of Veterinary Medicine, University of California Davis, Davis, CA 95616, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Muhammad S.","family":"Sajid","sequence":"additional","affiliation":[{"name":"Department of Medicine and Epidemiology, School of Veterinary Medicine, University of California Davis, Davis, CA 95616, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Janet E.","family":"Foley","sequence":"additional","affiliation":[{"name":"Department of Medicine and Epidemiology, School of Veterinary Medicine, University of California Davis, Davis, CA 95616, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2020,9,24]]},"reference":[{"key":"ref_1","first-page":"692","article-title":"Sur l\u2019h\u00e9moglobinurie bact\u00e9rienne du boef","volume":"107","author":"Babes","year":"1888","journal-title":"CR Acad. Sci."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1788","DOI":"10.1016\/j.meegid.2012.07.004","article-title":"Babesia: A world emerging","volume":"12","author":"Schnittger","year":"2012","journal-title":"Infect. Genet. Evol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1016\/j.vetpar.2006.01.035","article-title":"Babesia\u2014A historical overview","volume":"138","author":"Uilenberg","year":"2006","journal-title":"Veter Parasitol."},{"key":"ref_4","first-page":"11","article-title":"Piroplasmosis in man: Report on a case","volume":"9","year":"1957","journal-title":"Doc. Med. Geogr. Trop."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Smith, T., and Kilbourne, F.L. (1893). Investigation into the Nature, Causation, and Prevention of Southern Cattle Fever.","DOI":"10.5962\/bhl.title.124068"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1517","DOI":"10.1111\/j.1537-2995.2011.03467.x","article-title":"The third described case of transfusion-transmitted Babesia duncani","volume":"52","author":"Bloch","year":"2011","journal-title":"Transfusion"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"130","DOI":"10.5858\/arpa.2017-0250-RS","article-title":"Transfusion-transmitted babesiosis","volume":"143","author":"Villatoro","year":"2019","journal-title":"Arch. Pathol. Lab. Med."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1869","DOI":"10.3201\/eid2211.151028","article-title":"Transmission of Babesia microti parasites by solid organ transplantation","volume":"22","author":"Brennan","year":"2016","journal-title":"Emerg. Infect. Dis."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1097\/01.inf.0000195438.09628.b0","article-title":"Neonatal babesiosis: Case report and review of the literature","volume":"25","author":"Fox","year":"2006","journal-title":"Pediatr. Infect. Dis. J."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"408","DOI":"10.1097\/IPC.0b013e31825b20c1","article-title":"Vertical transmission of babesiosis from a pregnant, splenectomized mother to her neonate","volume":"20","author":"Cornett","year":"2012","journal-title":"Infect. Dis. Clin. Pract."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1323","DOI":"10.1016\/S0020-7519(00)00137-5","article-title":"Human babesiosis: An emerging tick-borne disease","volume":"30","author":"Kjemtrup","year":"2000","journal-title":"Int. J. Parasitol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"214","DOI":"10.1016\/j.vetpar.2010.05.004","article-title":"Molecular and serologic evidence for Babesia bovis-like parasites in white-tailed deer (Odocoileus virginianus) in south Texas","volume":"172","author":"Ramos","year":"2010","journal-title":"Vet. Parasitol."},{"key":"ref_13","unstructured":"Australian Society for Parasitology (2020, July 27). Babesia\u00a0. Available online: http:\/\/parasite.org.au\/parasite\/text\/babesia-text.html#top."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"111","DOI":"10.2307\/3282486","article-title":"In vitro cultivation of a Babesia isolated from a white-tailed deer (Odocoileus virginianus)","volume":"74","author":"Holman","year":"1988","journal-title":"J. Parasitol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1111\/j.1550-7408.1993.tb04909.x","article-title":"The isolation and partial characterization of a Babesia sp. from desert bighorn sheep (Ovis canadensis nelsoni)","volume":"40","author":"Goff","year":"1993","journal-title":"J. Eukaryot. Microbiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"77","DOI":"10.2307\/3283281","article-title":"Isolation and in vitro cultivation of Babesia parasites from free-ranging desert bighorn sheep (Ovis canadensis nelsoni) and mule deer (Odocoileus hemionus) in California","volume":"79","author":"Thomford","year":"1993","journal-title":"J. Parasitol."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Scott, J.D. (2017). First record of locally acquired human babesiosis in Canada caused by Babesia duncani: A case report. Sage Open Med. Case Rep., 5.","DOI":"10.1177\/2050313X17725645"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"643","DOI":"10.7326\/0003-4819-124-7-199604010-00004","article-title":"A Fatal case of babesiosis in Missouri: Identification of another piroplasm that infects humans","volume":"124","author":"Herwaldt","year":"1996","journal-title":"Ann. Intern. Med."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1007\/s40588-015-0025-z","article-title":"Human babesiosis: Pathogens, prevalence, diagnosis, and treatment","volume":"2","author":"Ord","year":"2015","journal-title":"Curr. Clin. Microbiol. Rep."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1589","DOI":"10.3201\/eid1810.120511","article-title":"Wild birds and urban ecology of ticks and tick-borne pathogens, Chicago, Illinois, USA, 2005\u20132010","volume":"18","author":"Hamer","year":"2012","journal-title":"Emerg. Infect. Dis."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"5","DOI":"10.13188\/2325-4645.1000005","article-title":"The establishment of a blacklegged tick population by migratory songbirds in Ontario, Canada","volume":"1","author":"Scott","year":"2012","journal-title":"J. Vet. Sci. Med."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"133","DOI":"10.3201\/eid0602.000205","article-title":"Competence of American Robins as reservoir hosts for Lyme disease spirochetes","volume":"6","author":"Richter","year":"2000","journal-title":"Emerg. Infect. Dis."},{"key":"ref_23","unstructured":"Mullen, G.R., and Durden, L.A. (2019). Ticks (Ixodida). Medical and Veterinary Entomology, Academic Press\/Elsevier. [3rd ed.]."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Hersh, M.H., Osfeld, R.S., McHenry, D.J., Tibbetts, M., Brunner, J.L., Killilea, M.E., LoGiudice, K., Schmidt, K.A., and Keesing, F. (2014). Co-infestation of blacklegged ticks with Babesia microti and Borrelia burgdorferi is higher than expected and acquired from small mammal hosts. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0099348"},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Clifford, C.M., Anastos, G., and Elbl, A. (1961). The Larval Ixodid Ticks of the Eastern United States, Entomological Society of America Lanham.","DOI":"10.4182\/BHJB6050.2-1.3"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1093\/jmedent\/33.3.297","article-title":"Ixodes (Ixodes) scapularis (Acari: Ixodidae): Redescription of all active stages, distribution, hosts, geographical variation, and medical and veterinary importance","volume":"33","author":"Keirans","year":"1996","journal-title":"J. Med. Entomol."},{"key":"ref_27","unstructured":"Cooley, R.A. (1946). The Genera Boophilus, Rhipicephalus, and Haemaphysalis (Ixodidae) of the New World (National Institute of Health Bulletin No. 187)."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Durden, L.A., and Keirans, J.E. (1996). Nymphs of the Genus Ixodes (Acari: Ixodidae) of the United States: Taxonomy, Identification Key, Distribution, Hosts, and Medical\/Veterinary Importance, Thomas Say Publications in Entomology, Entomological Society of America.","DOI":"10.4182\/MAMR9602"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1093\/jmedent\/15.suppl2.1","article-title":"The genus Ixodes in the United States: A scanning electron microscope study and key to the adults","volume":"15","author":"Keirans","year":"1978","journal-title":"J. Med. Entomol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"869","DOI":"10.1093\/jmedent\/44.5.869","article-title":"Molecular identification of bloodmeal source in Ixodes ricinus ticks using 12 rDNA as a genetic marker","volume":"44","author":"Humair","year":"2007","journal-title":"J. Med. Entomol."},{"key":"ref_31","first-page":"65","article-title":"Presence of potentially pathogenic Babesia sp. for human in Ixodes ricinus in Switzerland","volume":"13","author":"Casati","year":"2016","journal-title":"Ann. Agric. Environ. Med."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1792","DOI":"10.1093\/nar\/gkh340","article-title":"MUSCLE: Multiple sequence alignment with high accuracy and high throughput","volume":"32","author":"Edgar","year":"2004","journal-title":"Nucleic Acids Res."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1547","DOI":"10.1093\/molbev\/msy096","article-title":"MEGA X: Molecular evolutionary genetics analysis across computing platforms","volume":"35","author":"Kumar","year":"2018","journal-title":"Mol. Biol. Evol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"772","DOI":"10.1038\/nmeth.2109","article-title":"jModelTest 2: More models, new heuristics and parallel computing","volume":"9","author":"Darriba","year":"2012","journal-title":"Nat. Methods"},{"key":"ref_35","unstructured":"US Fish and Wildlife Service (2020, July 30). Flyways, Available online: https:\/\/www.fws.gov\/birds\/management\/flyways.php."},{"key":"ref_36","first-page":"854","article-title":"Amblyomma dissimile Koch (Acari: Ixodidae) parasitizes bird captured in Canada","volume":"20","author":"Scott","year":"2015","journal-title":"Syst. Appl. Acarol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"780","DOI":"10.1645\/GE-3437.1","article-title":"Migratory songbirds disperse ticks across Canada, and first isolation of the Lyme disease spirochete, Borrelia burgdorferi, from the avian tick, Ixodes auritulus","volume":"91","author":"Morshed","year":"2005","journal-title":"J. Parasitol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1111\/j.1948-7134.2010.00068.x","article-title":"Detection of Lyme disease spirochete, Borrelia burgdorferi sensu lato, including three novel genotypes in ticks (Acari: Ixodidae) collected from songbirds (Passeriformes) across Canada","volume":"35","author":"Scott","year":"2010","journal-title":"J. Vector Ecol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1645\/GE-2874.1","article-title":"Widespread dispersal of Borrelia burgdorferi-infected ticks collected from songbirds across Canada","volume":"98","author":"Scott","year":"2012","journal-title":"J. Parasitol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1080\/01647954.2015.1038301","article-title":"New records of the Lyme disease bacterium in ticks collected from songbirds in central and eastern Canada","volume":"41","author":"Scott","year":"2015","journal-title":"Int. J. Acarol."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Scott, J.D., Clark, K.L., Foley, J.E., Bierman, B.C., and Durden, L.A. (2018). Far-reaching dispersal of Borrelia burgdorferi sensu lato-infected blacklegged ticks by migratory songbirds in Canada. Healthcare, 6.","DOI":"10.3390\/healthcare6030089"},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Scott, J.D., Clark, K.L., Foley, J.E., Anderson, J.F., Bierman, B.C., and Durden, L.A. (2018). Extensive distribution of the Lyme disease bacterium, Borrelia burgdorferi sensu lato, in multiple tick species parasitizing avian and mammalian hosts across Canada. Healthcare, 6.","DOI":"10.3390\/healthcare6040131"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Scott, J.D., Clark, K.L., Coble, N.M., and Ballantyne, T.R. (2019). Detection and transstadial passage of Babesia species and Borrelia burgdorferi sensu lato in ticks collected from avian and mammalian hosts in Canada. Healthcare, 7.","DOI":"10.3390\/healthcare7040155"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"149","DOI":"10.7589\/0090-3558-21.2.149","article-title":"Babesia odocoilei Emerson and Wright, 1970 in white-tailed deer, Odocoileus virginianus (Zimmermann), in Virginia","volume":"21","author":"Perry","year":"1985","journal-title":"J. Wildl. Dis."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"518","DOI":"10.7589\/0090-3558-36.3.518","article-title":"Antigenic, phenotypic and molecular characterization confirms Babesia odocoilei isolated from three cervids","volume":"36","author":"Holman","year":"2000","journal-title":"J. Wildl. Dis."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1177\/1040638713491746","article-title":"Babesia odocoilei infection in a Sakatchewan elk (Cervus elaphus canadensis) herd","volume":"25","author":"Pattullo","year":"2013","journal-title":"J. Vet. Diagn. Investig."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"942","DOI":"10.3201\/eid0908.020748","article-title":"Molecular characterization of a non-Babesia divergens organism causing zoonotic babesiosis in Europe","volume":"9","author":"Herwaldt","year":"2003","journal-title":"Emerg. Infect. Dis."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1093\/jmedent\/43.2.437","article-title":"Detection of Anaplasma phagocytophilum and Babesia odocoilei DNA in Ixodes scapularis (Acari: Ixodidae) collected in Indiana","volume":"43","author":"Steiner","year":"2006","journal-title":"J. Med. Entomol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1093\/jmedent\/45.2.289","article-title":"Infection and co-infection rates of Anaplasma phagocytophilum variant, Babesia spp., Borrelia burgdorferi, and richettsial endosymbiont in Ixodes scapularis (Acari: Ixodidae) from sites in Indiana, Maine, Pennsylvania, and Wisconsin","volume":"45","author":"Steiner","year":"2008","journal-title":"J. Med. Entomol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1131","DOI":"10.3201\/eid1307.070046","article-title":"Zoonotic pathogens in Ixodes scapularis, Michigan","volume":"13","author":"Hamer","year":"2007","journal-title":"Emerg. Infect. Dis."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"683","DOI":"10.7589\/0090-3558-41.4.683","article-title":"New ruminant hosts and wider geographic range identified for Babesia odocoilei (Emerson and Wright 1970)","volume":"41","author":"Schoelkopf","year":"2005","journal-title":"J. Wildl. Dis."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"406","DOI":"10.1089\/vbz.2019.2551","article-title":"Detection of Babesia, Borrelia, Anaplasma, and Rickettsia spp. in adult black-legged ticks (Ixodes scapularis) from Pennsylvania, United States, with a Luminex Bead Assay","volume":"20","author":"Livengood","year":"2020","journal-title":"Vector Borne Zoonotic Dis."},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Scott, J.D., Clark, K.L., Coble, N.M., and Ballantyne, T.R. (2019). Presence of Babesia odocoilei and Borrelia burgdorferi sensu stricto in a tick and dual parasitism of Amblyomma inornatum and Ixodes scapularis on a bird in Canada. Healthcare, 7.","DOI":"10.3390\/healthcare7010046"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"670","DOI":"10.1016\/j.ttbdis.2019.02.016","article-title":"Babesia odocoilei and zoonotic pathogens identified from Ixodes scapularis ticks in southern Ontario, Canada","volume":"10","author":"Milnes","year":"2019","journal-title":"Ticks Tick-Borne Dis."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1016\/j.ttbdis.2014.01.003","article-title":"Diversity of piroplasms detected in blood-fed and questing ticks from several states in the United States","volume":"5","author":"Shock","year":"2014","journal-title":"Ticks Tick-Borne Dis."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1017\/S0031182006001545","article-title":"Transstadial and transovarial persistence of Babesia divergens DNA in Ixodes ricinus ticks fed on infected blood in a new skin-feeding technique","volume":"134","author":"Bonnet","year":"2007","journal-title":"Parasitology"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1016\/S0065-308X(08)60285-7","article-title":"The piroplasms: Life cycle and sexual stages","volume":"23","author":"Mehlhorn","year":"1984","journal-title":"Adv. Parasitol."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Scott, J.D., Pascoe, E.L., Sajid, M.S., and Foley, J.E. (2020). Monitoring of nesting songbirds detects established population of blacklegged ticks and associated Lyme disease endemic area in Canada. Healthcare, 8.","DOI":"10.3390\/healthcare8010059"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1128\/CMR.13.3.451","article-title":"Babesiosis","volume":"13","author":"Homer","year":"2000","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"2779","DOI":"10.1128\/jcm.29.12.2779-2783.1991","article-title":"Babesia microti, human babesiosis, and Borrelia burgdorferi in Connecticut","volume":"29","author":"Anderson","year":"1991","journal-title":"J. Clin. Microbiol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1117","DOI":"10.1086\/319742","article-title":"Severe babesiosis in Long Island: Review of 34 cases and their complications","volume":"32","author":"Hatcher","year":"2001","journal-title":"Clin. Infect. Dis."}],"container-title":["Pathogens"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-0817\/9\/10\/781\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T10:13:09Z","timestamp":1760177589000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-0817\/9\/10\/781"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,9,24]]},"references-count":61,"journal-issue":{"issue":"10","published-online":{"date-parts":[[2020,10]]}},"alternative-id":["pathogens9100781"],"URL":"https:\/\/doi.org\/10.3390\/pathogens9100781","relation":{},"ISSN":["2076-0817"],"issn-type":[{"value":"2076-0817","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,9,24]]}}}