{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,3]],"date-time":"2026-11-03T17:42:11Z","timestamp":1793727731519,"version":"4.6.1"},"reference-count":57,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,1,20]],"date-time":"2026-01-20T00:00:00Z","timestamp":1768867200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,2,13]],"date-time":"2026-02-13T00:00:00Z","timestamp":1770940800000},"content-version":"vor","delay-in-days":24,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100002428","name":"FWF Austrian Science Fund","doi-asserted-by":"crossref","award":["P37076"],"award-info":[{"award-number":["P37076"]}],"id":[{"id":"10.13039\/501100002428","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000001","name":"US National Science Foundation","doi-asserted-by":"crossref","award":["NSF-DEB 2146653"],"award-info":[{"award-number":["NSF-DEB 2146653"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Parasites Vectors"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>\n                      Avian haemosporidians are single-celled eukaryotic parasites of vertebrates that require dipteran vectors for transmission. The genera\n                      <jats:italic>Plasmodium<\/jats:italic>\n                      ,\n                      <jats:italic>Haemoproteus<\/jats:italic>\n                      and\n                      <jats:italic>Leucocytozoon<\/jats:italic>\n                      currently comprise over 5000 parasite lineages based on a 478-bp section of the mitochondrial cytochrome\n                      <jats:italic>b<\/jats:italic>\n                      gene, which is the standard DNA barcode for avian haemosporidians. The mitochondrial genomes of apicomplexan parasites are highly condensed, with a length of approximately 6000\u00a0bp, containing three coding genes (cytochrome\n                      <jats:italic>c<\/jats:italic>\n                      oxidase subunit I, cytochrome\n                      <jats:italic>c<\/jats:italic>\n                      oxidase subunit III and cytochrome\n                      <jats:italic>b<\/jats:italic>\n                      ) and dispersed fragments of the small and large ribosomal RNA (rRNA) genes. Since the mitochondrial genomes are relatively conserved, they are valuable markers for studying the phylogenetic relationships between haemosporidian parasites. However, until recently, mitochondrial genomes were unavailable for parasites of the\n                      <jats:italic>Haemoproteus nisi<\/jats:italic>\n                      and\n                      <jats:italic>Leucocytozoon toddi<\/jats:italic>\n                      species groups, which are exclusive to accipitriform raptors and strongly diverged from other\n                      <jats:italic>Haemoproteus<\/jats:italic>\n                      and\n                      <jats:italic>Leucocytozoon<\/jats:italic>\n                      parasites.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      We screened 171 accipitriform raptors from Austria and Germany using new primers targeting the cytochrome\n                      <jats:italic>b<\/jats:italic>\n                      gene of a previously neglected\n                      <jats:italic>L. toddi<\/jats:italic>\n                      clade. We also developed a new primer assay that enables the amplification and sequencing of the complete mitochondrial genomes of haemosporidian parasites. This process involved long-range PCRs with lineage-specific primers placed within the cytochrome\n                      <jats:italic>b<\/jats:italic>\n                      gene, followed by five nested PCRs targeting conserved sequence regions.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      Screening the accipitriform raptors revealed 10 new\n                      <jats:italic>L. toddi<\/jats:italic>\n                      group lineages. We sequenced 18 mitochondrial genomes belonging to five\n                      <jats:italic>H. nisi<\/jats:italic>\n                      group, nine\n                      <jats:italic>L. toddi<\/jats:italic>\n                      group, and two other\n                      <jats:italic>Leucocytozoon<\/jats:italic>\n                      lineages. Phylogenetic analyses based on mt genome sequences placed the\n                      <jats:italic>L. toddi<\/jats:italic>\n                      lineages within the genus\n                      <jats:italic>Leucocytozoon<\/jats:italic>\n                      , but the results did not support a monophyly of the genus\n                      <jats:italic>Haemoproteus<\/jats:italic>\n                      .\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>\n                      The new nested PCR approach with lineage-specific primers used for the long-range PCRs described herein successfully enabled the sequencing of the complete mitochondrial genomes, even in samples with mixed infections. The mitochondrial genomes of the\n                      <jats:italic>H. nisi<\/jats:italic>\n                      and\n                      <jats:italic>L. toddi<\/jats:italic>\n                      group lineages are highly valuable for resolving the phylogenetic relationships of the order Haemosporida since these parasites belong to clades distinct from other\n                      <jats:italic>Haemoproteus<\/jats:italic>\n                      and\n                      <jats:italic>Leucocytozoon<\/jats:italic>\n                      parasites.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Graphical Abstract<\/jats:title>\n                  <\/jats:sec>","DOI":"10.1186\/s13071-026-07244-0","type":"journal-article","created":{"date-parts":[[2026,1,20]],"date-time":"2026-01-20T17:45:14Z","timestamp":1768931114000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["New mitochondrial genomes of parasites belonging to the Leucocytozoon toddi and Haemoproteus nisi groups (Haemosporida, Apicomplexa)"],"prefix":"10.1186","volume":"19","author":[{"given":"Josef","family":"Harl","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tanja","family":"Himmel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"M. Andre\u00edna","family":"Pacheco","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Herbert","family":"Weissenb\u00f6ck","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,1,20]]},"reference":[{"key":"7244_CR1","volume-title":"Avian malaria parasites and other Haemosporidia","author":"G Valki\u016bnas","year":"2005","unstructured":"Valki\u016bnas G. Avian malaria parasites and other Haemosporidia. Boca Raton: CRC Press; 2005."},{"key":"7244_CR2","doi-asserted-by":"publisher","first-page":"1353","DOI":"10.1111\/j.1755-0998.2009.02692.x","volume":"9","author":"S Bensch","year":"2009","unstructured":"Bensch S, Hellgren O, P\u00e9rez-Tris J. MalAvi: a public database of malaria parasites and related haemosporidians in avian hosts based on mitochondrial cytochrome b lineages. Mol Ecol Resour. 2009;9:1353\u20138. https:\/\/doi.org\/10.1111\/j.1755-0998.2009.02692.x.","journal-title":"Mol Ecol Resour"},{"key":"7244_CR3","doi-asserted-by":"publisher","first-page":"383","DOI":"10.1093\/molbev\/msx285","volume":"35","author":"MA Pacheco","year":"2018","unstructured":"Pacheco MA, Matta NE, Valki\u016bnas G, Parker PG, Mello B, Stanley CE, et al. Mode and rate of evolution of haemosporidian mitochondrial genomes: timing the radiation of avian parasites. Mol Biol Evol. 2018;35:383\u2013403. https:\/\/doi.org\/10.1093\/molbev\/msx285.","journal-title":"Mol Biol Evol"},{"key":"7244_CR4","doi-asserted-by":"publisher","DOI":"10.1016\/j.meegid.2020.104576","volume":"85","author":"MA Pacheco","year":"2020","unstructured":"Pacheco MA, Cer\u00edaco LMP, Matta NE, Vargas-Ram\u00edrez M, Bauer AM, Escalante AA. A phylogenetic study of Haemocystidium parasites and other haemosporida using complete mitochondrial genome sequences. Infect Genet Evol. 2020;85:104576. https:\/\/doi.org\/10.1016\/j.meegid.2020.104576.","journal-title":"Infect Genet Evol"},{"key":"7244_CR5","doi-asserted-by":"publisher","first-page":"471","DOI":"10.1080\/19401730802570926","volume":"19","author":"SL Perkins","year":"2008","unstructured":"Perkins SL. Molecular systematics of the three mitochondrial protein-coding genes of malaria parasites: corroborative and new evidence for the origins of human malaria. DNA Seq. 2008;19:471\u20138. https:\/\/doi.org\/10.1080\/19401730802570926.","journal-title":"DNA Seq"},{"key":"7244_CR6","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0038320","volume":"7","author":"JE Feagin","year":"2012","unstructured":"Feagin JE, Harrell MI, Lee JC, Coe KJ, Sands BH, Cannone JJ, et al. The fragmented mitochondrial ribosomal RNAs of Plasmodium falciparum. PLoS ONE. 2012;7:e38320. https:\/\/doi.org\/10.1371\/journal.pone.0038320.","journal-title":"PLoS ONE"},{"key":"7244_CR7","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1146\/annurev.genet.37.110801.142526","volume":"38","author":"MW Gray","year":"2004","unstructured":"Gray MW, Lang BF, Burger G. Mitochondria of protists. Annu Rev Genet. 2004;38:477\u2013524. https:\/\/doi.org\/10.1146\/annurev.genet.37.110801.142526.","journal-title":"Annu Rev Genet"},{"key":"7244_CR8","doi-asserted-by":"publisher","first-page":"9834","DOI":"10.1073\/pnas.1610643113","volume":"113","author":"G Karadjian","year":"2016","unstructured":"Karadjian G, Hassanin A, Saintpierre B, Gembu Tungaluna G-C, Ariey F, Ayala FJ, et al. Highly rearranged mitochondrial genome in Nycteria parasites (Haemosporidia) from bats. Proc Natl Acad Sci USA. 2016;113:9834\u20139. https:\/\/doi.org\/10.1073\/pnas.1610643113.","journal-title":"Proc Natl Acad Sci USA"},{"key":"7244_CR9","doi-asserted-by":"publisher","first-page":"498","DOI":"10.1038\/nature01097","volume":"419","author":"MJ Gardner","year":"2002","unstructured":"Gardner MJ, Hall N, Fung E, White O, Berriman M, Hyman RW, et al. Genome sequence of the human malaria parasite Plasmodium falciparum. Nature. 2002;419:498\u2013511. https:\/\/doi.org\/10.1038\/nature01097.","journal-title":"Nature"},{"key":"7244_CR10","doi-asserted-by":"publisher","first-page":"318","DOI":"10.1126\/science.1081449","volume":"300","author":"DA Joy","year":"2003","unstructured":"Joy DA, Feng X, Mu J, Furuya T, Chotivanich K, Krettli AU, et al. Early origin and recent expansion of Plasmodium falciparum. Science. 2003;300:318\u201321. https:\/\/doi.org\/10.1126\/science.1081449.","journal-title":"Science"},{"key":"7244_CR11","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1000765","volume":"6","author":"S Krief","year":"2010","unstructured":"Krief S, Escalante AA, Pacheco MA, Mugisha L, Andr\u00e9 C, Halbwax M, et al. On the diversity of malaria parasites in African apes and the origin of Plasmodium falciparum from bonobos. PLoS Pathog. 2010;6:e1000765. https:\/\/doi.org\/10.1371\/journal.ppat.1000765.","journal-title":"PLoS Pathog"},{"key":"7244_CR12","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1002015","volume":"7","author":"K-S Lee","year":"2011","unstructured":"Lee K-S, Divis PCS, Zakaria SK, Matusop A, Julin RA, Conway DJ, et al. Plasmodium knowlesi: reservoir hosts and tracking the emergence in humans and macaques. PLoS Pathog. 2011;7:e1002015. https:\/\/doi.org\/10.1371\/journal.ppat.1002015.","journal-title":"PLoS Pathog"},{"key":"7244_CR13","doi-asserted-by":"publisher","first-page":"1686","DOI":"10.1093\/molbev\/msi160","volume":"22","author":"J Mu","year":"2005","unstructured":"Mu J, Joy DA, Duan J, Huang Y, Carlton J, Walker J, et al. Host switch leads to emergence of Plasmodium vivax malaria in humans. Mol Biol Evol. 2005;22:1686\u201393. https:\/\/doi.org\/10.1093\/molbev\/msi160.","journal-title":"Mol Biol Evol"},{"key":"7244_CR14","doi-asserted-by":"publisher","first-page":"422","DOI":"10.1093\/molbev\/msu310","volume":"32","author":"MP Muehlenbein","year":"2015","unstructured":"Muehlenbein MP, Pacheco MA, Taylor JE, Prall SP, Ambu L, Nathan S, et al. Accelerated diversification of nonhuman primate malarias in Southeast Asia: adaptive radiation or geographic speciation? Mol Biol Evol. 2015;32:422\u201339. https:\/\/doi.org\/10.1093\/molbev\/msu310.","journal-title":"Mol Biol Evol"},{"key":"7244_CR15","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2148-11-299","volume":"11","author":"MA Pacheco","year":"2011","unstructured":"Pacheco MA, Battistuzzi FU, Junge RE, Cornejo OE, Williams CV, Landau I, et al. Timing the origin of human malarias: the lemur puzzle. BMC Evol Biol. 2011;11:299. https:\/\/doi.org\/10.1186\/1471-2148-11-299.","journal-title":"BMC Evol Biol"},{"key":"7244_CR16","doi-asserted-by":"publisher","first-page":"661","DOI":"10.1007\/s00436-006-0333-6","volume":"100","author":"S Omori","year":"2007","unstructured":"Omori S, Sato Y, Isobe T, Yukawa M, Murata K. Complete nucleotide sequences of the mitochondrial genomes of two avian malaria protozoa, Plasmodium gallinaceum and Plasmodium juxtanucleare. Parasitol Res. 2007;100:661\u20134. https:\/\/doi.org\/10.1007\/s00436-006-0333-6.","journal-title":"Parasitol Res"},{"key":"7244_CR17","doi-asserted-by":"publisher","first-page":"953","DOI":"10.1007\/s00436-008-1083-4","volume":"103","author":"S Omori","year":"2008","unstructured":"Omori S, Sato Y, Hirakawa S, Isobe T, Yukawa M, Murata K. Two extra chromosomal genomes of Leucocytozoon caulleryi; complete nucleotide sequences of the mitochondrial genome and existence of the apicoplast genome. Parasitol Res. 2008;103:953\u20137. https:\/\/doi.org\/10.1007\/s00436-008-1083-4.","journal-title":"Parasitol Res"},{"key":"7244_CR18","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2020.106947","volume":"153","author":"A Ciloglu","year":"2020","unstructured":"Ciloglu A, Ellis VA, Duc M, Downing PA, Inci A, Bensch S. Evolution of vector transmitted parasites by host switching revealed through sequencing of Haemoproteus parasite mitochondrial genomes. Mol Phylogenet Evol. 2020;153:106947. https:\/\/doi.org\/10.1016\/j.ympev.2020.106947.","journal-title":"Mol Phylogenet Evol"},{"key":"7244_CR19","doi-asserted-by":"publisher","first-page":"2967","DOI":"10.1007\/s00436-023-07986-1","volume":"122","author":"S Musa","year":"2023","unstructured":"Musa S. Mitochondrial genome amplification of avian haemosporidian parasites from single-infected wildlife samples using a novel nested PCR approach. Parasitol Res. 2023;122:2967\u201375. https:\/\/doi.org\/10.1007\/s00436-023-07986-1.","journal-title":"Parasitol Res"},{"key":"7244_CR20","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s12936-021-04019-z","volume":"21","author":"J Harl","year":"2022","unstructured":"Harl J, Himmel T, Valki\u016bnas G, Ilg\u016bnas M, Nedorost N, Matt J, et al. Avian haemosporidian parasites of accipitriform raptors. Malar J. 2022;21:1\u201338. https:\/\/doi.org\/10.1186\/s12936-021-04019-z.","journal-title":"Malar J"},{"key":"7244_CR21","doi-asserted-by":"publisher","DOI":"10.1051\/parasite\/2023066","volume":"31","author":"J Harl","year":"2024","unstructured":"Harl J, Fauchois A, Puech M-P, Gey D, Ariey F, Izac B, et al. Novel phylogenetic clade of avian Haemoproteus parasites (Haemosporida, Haemoproteidae) from Accipitridae raptors, with description of a new Haemoproteus species. Parasite. 2024;31:5. https:\/\/doi.org\/10.1051\/parasite\/2023066.","journal-title":"Parasite"},{"key":"7244_CR22","doi-asserted-by":"publisher","first-page":"797","DOI":"10.1645\/GE-184R1","volume":"90","author":"O Hellgren","year":"2004","unstructured":"Hellgren O, Waldenstr\u00f6m J, Bensch S. A new PCR assay for simultaneous studies of Leucocytozoon, Plasmodium, and Haemoproteus from avian blood. J Parasitol. 2004;90:797\u2013802. https:\/\/doi.org\/10.1645\/GE-184R1.","journal-title":"J Parasitol"},{"key":"7244_CR23","doi-asserted-by":"publisher","first-page":"375","DOI":"10.1645\/GE-656R.1","volume":"92","author":"RNM Sehgal","year":"2006","unstructured":"Sehgal RNM, Hull AC, Anderson NL, Valki\u016bnas G, Markovets MJ, Kawamura S, et al. Evidence for cryptic speciation of Leucocytozoon spp. (Haemosporida, Leucocytozoidae) in diurnal raptors. J Parasitol. 2006;92:375\u20139. https:\/\/doi.org\/10.1645\/GE-656R.1.","journal-title":"J Parasitol"},{"key":"7244_CR24","doi-asserted-by":"publisher","first-page":"767","DOI":"10.1007\/s10336-014-1064-6","volume":"155","author":"MA Jasper","year":"2014","unstructured":"Jasper MA, Hull JM, Hull AC, Sehgal RNM. Widespread lineage diversity of Leucocytozoon blood parasites in distinct populations of western red-tailed hawks. J Ornithol. 2014;155:767\u201375. https:\/\/doi.org\/10.1007\/s10336-014-1064-6.","journal-title":"J Ornithol"},{"key":"7244_CR25","doi-asserted-by":"publisher","first-page":"381","DOI":"10.1016\/j.ijpara.2012.12.007","volume":"43","author":"A P\u00e9rez-Rodr\u00edguez","year":"2013","unstructured":"P\u00e9rez-Rodr\u00edguez A, de la Puente J, Onrubia A, P\u00e9rez-Tris J. Molecular characterization of haemosporidian parasites from kites of the genus Milvus (Aves: Accipitridae). Int J Parasitol. 2013;43:381\u20137. https:\/\/doi.org\/10.1016\/j.ijpara.2012.12.007.","journal-title":"Int J Parasitol"},{"key":"7244_CR26","doi-asserted-by":"publisher","DOI":"10.1186\/s12936-024-04961-8","volume":"23","author":"MA Pacheco","year":"2024","unstructured":"Pacheco MA, Cepeda AS, Miller EA, Beckerman S, Oswald M, London E, et al. A new long-read mitochondrial-genome protocol (PacBio HiFi) for haemosporidian parasites: a tool for population and biodiversity studies. Malar J. 2024;23:134. https:\/\/doi.org\/10.1186\/s12936-024-04961-8.","journal-title":"Malar J"},{"key":"7244_CR27","doi-asserted-by":"publisher","DOI":"10.1186\/s12936-019-2940-6","volume":"18","author":"J Harl","year":"2019","unstructured":"Harl J, Himmel T, Valki\u016bnas G, Weissenb\u00f6ck H. The nuclear 18S ribosomal DNAs of avian haemosporidian parasites. Malar J. 2019;18:305. https:\/\/doi.org\/10.1186\/s12936-019-2940-6.","journal-title":"Malar J"},{"key":"7244_CR28","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s12936-020-03408-0","volume":"19","author":"J Harl","year":"2020","unstructured":"Harl J, Himmel T, Valki\u016bnas G, Ilg\u016bnas M, Bakonyi T, Weissenb\u00f6ck H. Geographic and host distribution of haemosporidian parasite lineages from birds of the family Turdidae. Malar J. 2020;19:1\u201335.","journal-title":"Malar J"},{"key":"7244_CR29","first-page":"95","volume":"41","author":"TA Hall","year":"1999","unstructured":"Hall TA. BioEdit: a user-friendly biological sequences alignment editor and analysis program for Windows 95\/98\/NT. Nucleic Acids Symp Ser. 1999;41:95\u20138.","journal-title":"Nucleic Acids Symp Ser"},{"key":"7244_CR30","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. https:\/\/doi.org\/10.1093\/molbev\/msy096.","journal-title":"Mol Biol Evol"},{"key":"7244_CR31","doi-asserted-by":"publisher","first-page":"772","DOI":"10.1093\/molbev\/mst010","volume":"30","author":"K Katoh","year":"2013","unstructured":"Katoh K, Standley DM. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Mol Biol Evol. 2013;30:772\u201380. https:\/\/doi.org\/10.1093\/molbev\/mst010.","journal-title":"Mol Biol Evol"},{"key":"7244_CR32","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1093\/molbev\/msu300","volume":"32","author":"L-T Nguyen","year":"2015","unstructured":"Nguyen L-T, Schmidt HA, von Haeseler A, Minh BQ. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Mol Biol Evol. 2015;32:268\u201374. https:\/\/doi.org\/10.1093\/molbev\/msu300.","journal-title":"Mol Biol Evol"},{"key":"7244_CR33","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1093\/sysbio\/sys029","volume":"61","author":"F Ronquist","year":"2012","unstructured":"Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, H\u00f6hna S, et al. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst Biol. 2012;61:539\u201342. https:\/\/doi.org\/10.1093\/sysbio\/sys029.","journal-title":"Syst Biol"},{"key":"7244_CR34","doi-asserted-by":"publisher","first-page":"979","DOI":"10.1111\/j.1420-9101.2008.01545.x","volume":"21","author":"A Marzal","year":"2008","unstructured":"Marzal A, Bensch S, Reviriego M, Balbontin J, De Lope F. Effects of malaria double infection in birds: one plus one is not two. J Evol Biol. 2008;21:979\u201387. https:\/\/doi.org\/10.1111\/j.1420-9101.2008.01545.x.","journal-title":"J Evol Biol"},{"key":"7244_CR35","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1016\/j.exppara.2016.01.009","volume":"163","author":"R Bernotien\u0117","year":"2016","unstructured":"Bernotien\u0117 R, Palinauskas V, Iezhova T, Murauskait\u0117 D, Valki\u016bnas G. Avian haemosporidian parasites (Haemosporida): a comparative analysis of different polymerase chain reaction assays in detection of mixed infections. Exp Parasitol. 2016;163:31\u20137. https:\/\/doi.org\/10.1016\/j.exppara.2016.01.009.","journal-title":"Exp Parasitol"},{"key":"7244_CR36","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1007\/s00436-015-4743-1","volume":"115","author":"J Hanel","year":"2016","unstructured":"Hanel J, Dole\u017ealov\u00e1 J, Stehl\u00edkov\u00e1 \u0160, Modr\u00fd D, Chudoba J, Synek P, et al. Blood parasites in northern goshawk (Accipiter gentilis) with an emphasis to Leucocytozoon toddi. Parasitol Res. 2016;115:263\u201370.","journal-title":"Parasitol Res"},{"key":"7244_CR37","doi-asserted-by":"publisher","first-page":"709","DOI":"10.1645\/GE-1357.1","volume":"94","author":"O Krone","year":"2008","unstructured":"Krone O, Waldenstr\u00f6m J, Valki\u016bnas G, Lessow O, M\u00fcller K, Iezhova TA, et al. Haemosporidian blood parasites in European birds of prey and owls. J Parasitol. 2008;94:709\u201315.","journal-title":"J Parasitol"},{"key":"7244_CR38","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1016\/j.ympev.2007.11.012","volume":"47","author":"ES Martinsen","year":"2008","unstructured":"Martinsen ES, Perkins SL, Schall JJ. A three-genome phylogeny of malaria parasites (Plasmodium and closely related genera): evolution of life-history traits and host switches. Mol Phylogenet Evol. 2008;47:261\u201373.","journal-title":"Mol Phylogenet Evol"},{"key":"7244_CR39","doi-asserted-by":"publisher","first-page":"199","DOI":"10.1016\/j.ijpara.2018.11.001","volume":"49","author":"S Musa","year":"2019","unstructured":"Musa S, Mackenstedt U, Woog F, Dinkel A. Avian malaria on Madagascar: prevalence, biodiversity and specialization of haemosporidian parasites. Int J Parasitol. 2019;49:199\u2013210. https:\/\/doi.org\/10.1016\/j.ijpara.2018.11.001.","journal-title":"Int J Parasitol"},{"key":"7244_CR40","doi-asserted-by":"publisher","first-page":"761","DOI":"10.1139\/z77-099","volume":"55","author":"EC Greiner","year":"1977","unstructured":"Greiner EC, Kocan AA. Leucocytozoon (Haemosporida; Leucocytozoidae) of the Falconiformes. Can J Zool. 1977;55:761\u201370.","journal-title":"Can J Zool"},{"key":"7244_CR41","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1645\/ge-2109.1","volume":"96","author":"G Valki\u016bnas","year":"2010","unstructured":"Valki\u016bnas G, Sehgal RNM, Iezhova TA, Hull AC. Identification of Leucocytozoon toddi group (Haemosporida: Leucocytozoidae), with remarks on the species taxonomy of leucocytozoids. J Parasitol. 2010;96:170\u20137. https:\/\/doi.org\/10.1645\/ge-2109.1.","journal-title":"J Parasitol"},{"key":"7244_CR42","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s13071-022-05623-x","volume":"16","author":"M Svobodov\u00e1","year":"2023","unstructured":"Svobodov\u00e1 M, \u010cepi\u010dka I, Z\u00eddkov\u00e1 L, Kassahun A, Vot\u00fdpka J, Pe\u0161ke L, et al. Blood parasites (Trypanosoma, Leucocytozoon, Haemoproteus) in the Eurasian sparrowhawk (Accipiter nisus): diversity, incidence and persistence of infection at the individual level. Parasit Vectors. 2023;16:1\u201316. https:\/\/doi.org\/10.1186\/s13071-022-05623-x.","journal-title":"Parasit Vectors"},{"key":"7244_CR43","doi-asserted-by":"publisher","first-page":"215","DOI":"10.3356\/JRR-08-34.1","volume":"44","author":"HD Ishak","year":"2010","unstructured":"Ishak HD, Loiseau C, Hull AC, Sehgal RNM. Prevalence of blood parasites in migrating and wintering California hawks. J Raptor Res. 2010;44:215\u201323. https:\/\/doi.org\/10.3356\/JRR-08-34.1.","journal-title":"J Raptor Res"},{"key":"7244_CR44","doi-asserted-by":"publisher","first-page":"1171","DOI":"10.1645\/GE-1982.1","volume":"95","author":"DC Outlaw","year":"2009","unstructured":"Outlaw DC, Ricklefs RE. On the phylogenetic relationships of haemosporidian parasites from raptorial birds (Falconiformes and Strigiformes). J Parasitol. 2009;95:1171\u20136. https:\/\/doi.org\/10.1645\/GE-1982.1.","journal-title":"J Parasitol"},{"key":"7244_CR45","unstructured":"Lemus, JA, Blanco G, Rodr\u00edguez-Estrella R. High infestation by blood parasites (Haematozoa) in nestlings of colonial ospreys (Pandion haliaetus) from Baja California, M\u00e9xico. In: Rodr\u00edguez-Estella R, editor. Current raptor studies in M\u00e9xico. Mexico DF: CIBNOR-CONABIO; 2006. p. 163\u2013180."},{"key":"7244_CR46","first-page":"281","volume":"40","author":"O Krone","year":"2001","unstructured":"Krone O, Priemer J, Streich J, S\u00f6mmer P, Langgemach T, Lessow O. Haemosporida of birds of prey and owls from Germany. Acta Protozool. 2001;40:281\u20139.","journal-title":"Acta Protozool"},{"key":"7244_CR47","doi-asserted-by":"publisher","first-page":"813","DOI":"10.1080\/00222938300770621","volume":"17","author":"MA Peirce","year":"1983","unstructured":"Peirce MA, Marquiss M. Haematozoa of British birds. VII. Haematozoa of raptors in Scotland with a description of Haemoproteus nisi sp. nov. from the sparrowhawk (Accipiter nisus). J Nat Hist. 1983;17:813\u201321. https:\/\/doi.org\/10.1080\/00222938300770621.","journal-title":"J Nat Hist"},{"key":"7244_CR48","unstructured":"Bennett GF, Whiteway M, Woodworth-Lynas C. A Host\u2013parasite catalogue of the avian Haematozoa. St. John\u2019s: Memorial University of Newfoundland; 1982."},{"key":"7244_CR49","doi-asserted-by":"publisher","first-page":"1762","DOI":"10.1139\/z75-211","volume":"53","author":"EC Greiner","year":"1975","unstructured":"Greiner EC, Bennett GF, White EM, Coombs RF. Distribution of the avian hematozoa of North America. Can J Zool. 1975;53:1762\u201387. https:\/\/doi.org\/10.1139\/z75-211.","journal-title":"Can J Zool"},{"key":"7244_CR50","doi-asserted-by":"publisher","first-page":"13183","DOI":"10.1073\/pnas.1109153108","volume":"108","author":"DC Outlaw","year":"2011","unstructured":"Outlaw DC, Ricklefs RE. Rerooting the evolutionary tree of malaria parasites. Proc Natl Acad Sci USA. 2011;108:13183\u20137.","journal-title":"Proc Natl Acad Sci USA"},{"key":"7244_CR51","doi-asserted-by":"publisher","first-page":"163","DOI":"10.1111\/j.1096-0031.2005.00059.x","volume":"21","author":"J Bergsten","year":"2005","unstructured":"Bergsten J. A review of long\u2010branch attraction. Cladistics. 2005;21:163\u201393. https:\/\/doi.org\/10.1111\/j.1096-0031.2005.00059.x.","journal-title":"Cladistics"},{"key":"7244_CR52","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1016\/j.pt.2023.04.004","volume":"39","author":"MA Pacheco","year":"2023","unstructured":"Pacheco MA, Escalante AA. Origin and diversity of malaria parasites and other Haemosporida. Trends Parasitol. 2023;39:501\u201316. https:\/\/doi.org\/10.1016\/j.pt.2023.04.004.","journal-title":"Trends Parasitol"},{"key":"7244_CR53","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1016\/j.ijppaw.2019.05.002","volume":"9","author":"IA Lotta","year":"2019","unstructured":"Lotta IA, Valki\u016bnas G, Pacheco MA, Escalante AA, Hern\u00e1ndez SR, Matta NE. Disentangling Leucocytozoon parasite diversity in the neotropics: descriptions of two new species and shortcomings of molecular diagnostics for leucocytozoids. Int J Parasitol Parasites Wildl. 2019;9:159\u201373.","journal-title":"Int J Parasitol Parasites Wildl"},{"key":"7244_CR54","doi-asserted-by":"publisher","first-page":"1031","DOI":"10.1007\/s00436-014-4269-y","volume":"114","author":"IA Lotta","year":"2015","unstructured":"Lotta IA, Gonzalez AD, Pacheco MA, Escalante AA, Valki\u016bnas G, Moncada LI, et al. Leucocytozoon pterotenuis sp. nov. (Haemosporida, Leucocytozoidae): description of the morphologically unique species from the Grallariidae birds, with remarks on the distribution of Leucocytozoon parasites in the Neotropics. Parasitol Res. 2015;114:1031\u201344. https:\/\/doi.org\/10.1007\/s00436-014-4269-y.","journal-title":"Parasitol Res"},{"key":"7244_CR55","doi-asserted-by":"publisher","first-page":"927","DOI":"10.1139\/z65-096","volume":"43","author":"GF Bennett","year":"1965","unstructured":"Bennett GF, Garnham PCC, Fallis AM. On the status of the genera Leucocytozoon Ziemann, 1898 and Haemoproteus Kruse, 1890 (Haemosporidiida: Leucocytozoidae and Haemoproteidae). Can J Zool. 1965;43:927\u201332. https:\/\/doi.org\/10.1139\/z65-096.","journal-title":"Can J Zool"},{"key":"7244_CR56","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s12936-022-04235-1","volume":"21","author":"G Valki\u016bnas","year":"2022","unstructured":"Valki\u016bnas G, Iezhova TA. Keys to the avian Haemoproteus parasites (Haemosporida, Haemoproteidae). Malar J. 2022;21:1\u201369.","journal-title":"Malar J"},{"key":"7244_CR57","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1016\/j.ympev.2015.09.003","volume":"94","author":"J Borner","year":"2016","unstructured":"Borner J, Pick C, Thiede J, Kolawole OM, Kingsley MT, Schulze J, et al. Phylogeny of haemosporidian blood parasites revealed by a multi-gene approach. Mol Phylogenet Evol. 2016;94:221\u201331. https:\/\/doi.org\/10.1016\/j.ympev.2015.09.003.","journal-title":"Mol Phylogenet Evol"}],"container-title":["Parasites &amp; Vectors"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s13071-026-07244-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13071-026-07244-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13071-026-07244-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,13]],"date-time":"2026-02-13T11:34:45Z","timestamp":1770982485000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s13071-026-07244-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,20]]},"references-count":57,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["7244"],"URL":"https:\/\/doi.org\/10.1186\/s13071-026-07244-0","relation":{},"ISSN":["1756-3305"],"issn-type":[{"value":"1756-3305","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,20]]},"assertion":[{"value":"24 July 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"3 January 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"20 January 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The collection of blood samples from living birds was approved by the institutional ethics and animal welfare committee and the national authority according to \u00a7\u00a7 26ff. of Animal Experiments Acts, Tierversuchsgesetz TVG 2012, Austria (BMWFW- 68.205\/0036-WF\/V\/3b\/2017). The samples collected by the Research Institute of Wildlife Ecology (University of Veterinary Medicine Vienna) were taken from carcasses submitted for routine pathological examinations. The samples from Germany originate from carcasses submitted to the Landeslabor Berlin-Brandenburg for routine diagnostics. No birds were euthanized for this study.","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 no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"80"}}