{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T15:23:50Z","timestamp":1788276230078,"version":"build-2803163510"},"reference-count":39,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,8,16]],"date-time":"2026-08-16T00:00:00Z","timestamp":1786838400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"vor","delay-in-days":16,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"DOI":"10.13039\/501100020912","name":"King Faisal University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100020912","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"DOI":"10.1038\/s41598-026-64985-9","type":"journal-article","created":{"date-parts":[[2026,8,16]],"date-time":"2026-08-16T11:34:01Z","timestamp":1786880041000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Molecular epidemiology of foot-and-mouth disease viruses collected from Khartoum, Blue Nile, and Northern States of Sudan (2019\u20132022)"],"prefix":"10.1038","volume":"16","author":[{"given":"Maysa","family":"Ragab","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yassir A.","family":"Almofti","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yazeed A.","family":"Rauouf","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nick J.","family":"Knowles","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Donald P.","family":"King","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jemma","family":"Wadsworth","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hayley M.","family":"Hicks","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haitham","family":"Saeed","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Omima Abd","family":"Algalil","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ibrahim","family":"Albokhadaim","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ahmed O.","family":"Alameen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mahmoud","family":"Kandeel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Samir","family":"Alhojaily","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mohammed Ali","family":"Al-Hammadi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jamal","family":"Hussen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mustafa","family":"Shukry","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bahaa","family":"Al-Trad","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,8,16]]},"reference":[{"issue":"2","key":"64985_CR1","doi-asserted-by":"publisher","first-page":"107","DOI":"10.3390\/vetsci12020107","volume":"12","author":"H Ma","year":"2025","unstructured":"Ma, H. et al. Comparative transcriptomics analysis of foot-and-mouth disease virus-infected cell model systems. Vet. Sci. 12 (2), 107 (2025).","journal-title":"Vet. Sci."},{"issue":"1","key":"64985_CR2","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1186\/s13567-025-01545-5","volume":"56","author":"C Stenfeldt","year":"2025","unstructured":"Stenfeldt, C. et al. The pathogenesis of foot-and-mouth disease virus: Current understandings and knowledge gaps. Vet. Res. 56 (1), 119 (2025).","journal-title":"Vet. Res."},{"issue":"1","key":"64985_CR3","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1016\/S0168-1702(02)00260-5","volume":"91","author":"NJ Knowles","year":"2003","unstructured":"Knowles, N. J. & Samuel, A. R. Molecular epidemiology of foot-and-mouth disease virus. Virus Res. 91 (1), 65\u201380 (2003).","journal-title":"Virus Res."},{"issue":"3","key":"64985_CR4","doi-asserted-by":"publisher","first-page":"437","DOI":"10.20506\/rst.21.3.1349","volume":"21","author":"W Vosloo","year":"2002","unstructured":"Vosloo, W. et al. Review of the status and control of foot and mouth disease in sub-Saharan Africa. Rev. Sci. Tech. 21 (3), 437\u2013449 (2002).","journal-title":"Rev. Sci. Tech."},{"issue":"3","key":"64985_CR5","doi-asserted-by":"publisher","first-page":"e0064322","DOI":"10.1128\/msphere.00643-22","volume":"8","author":"C Stenfeldt","year":"2023","unstructured":"Stenfeldt, C. et al. Heterogeneity and recombination of foot-and-mouth disease virus during multi-strain coinfection of cattle. mSphere 8 (3), e0064322 (2023).","journal-title":"mSphere"},{"issue":"1","key":"64985_CR6","doi-asserted-by":"publisher","first-page":"122","DOI":"10.1002\/vms3.208","volume":"6","author":"B Tadesse","year":"2020","unstructured":"Tadesse, B. et al. Foot and mouth disease outbreak investigation and estimation of its economic impact in selected districts in northwest Ethiopia. Vet. Med. Sci. 6 (1), 122\u2013132 (2020).","journal-title":"Vet. Med. Sci."},{"issue":"8","key":"64985_CR7","doi-asserted-by":"publisher","first-page":"1204","DOI":"10.1017\/S0950268809991427","volume":"138","author":"SN Balinda","year":"2010","unstructured":"Balinda, S. N. et al. Molecular characterization of SAT 2 foot-and-mouth disease virus from post-outbreak slaughtered animals: implications for disease control in Uganda. Epidemiol. Infect. 138 (8), 1204\u20131210 (2010).","journal-title":"Epidemiol. Infect."},{"issue":"5","key":"64985_CR8","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1111\/j.1865-1682.2010.01151.x","volume":"57","author":"M Habiela","year":"2010","unstructured":"Habiela, M. et al. Molecular characterization of foot-and-mouth disease viruses collected from Sudan. Transbound. Emerg. Dis. 57 (5), 305\u2013314 (2010).","journal-title":"Transbound. Emerg. Dis."},{"issue":"4","key":"64985_CR9","first-page":"443","volume":"64","author":"YA Raouf","year":"2018","unstructured":"Raouf, Y. A. et al. Sero-epidemiology of foot-and-mouth disease in Sudan. Bull. Anim. Health Prod. Afr. 64 (4), 443\u2013451 (2018).","journal-title":"Bull. Anim. Health Prod. Afr."},{"issue":"1","key":"64985_CR10","first-page":"145","volume":"65","author":"H Yousif","year":"2017","unstructured":"Yousif, H. et al. Role of small ruminants in the epidemiology of foot-and-mouth disease in Sudan. Bull. Anim. Health Prod. Afr. 65 (1), 145\u2013156 (2017).","journal-title":"Bull. Anim. Health Prod. Afr."},{"key":"64985_CR11","first-page":"443","volume":"64","author":"YA Raouf","year":"2016","unstructured":"Raouf, Y. A. et al. Sero-epidemiology of foot-and-mouth disease in Sudan. Anim. Health Prod. 64, 443\u2013451 (2016).","journal-title":"Anim. Health Prod."},{"issue":"2","key":"64985_CR12","doi-asserted-by":"publisher","first-page":"e404","DOI":"10.1111\/tbed.12774","volume":"65","author":"SS Hayer","year":"2018","unstructured":"Hayer, S. S. et al. Foot-and-mouth disease virus transmission dynamics and persistence in a herd of vaccinated dairy cattle in India. Transbound. Emerg. Dis. 65 (2), e404\u2013e415 (2018).","journal-title":"Transbound. Emerg. Dis."},{"issue":"3","key":"64985_CR13","doi-asserted-by":"publisher","first-page":"741","DOI":"10.20506\/rst.35.3.2565","volume":"35","author":"NJ Knowles","year":"2016","unstructured":"Knowles, N. J. et al. VP1 sequencing protocol for foot and mouth disease virus molecular epidemiology. Rev. Sci. Tech. 35 (3), 741\u2013755 (2016).","journal-title":"Rev. Sci. Tech."},{"issue":"1","key":"64985_CR14","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1186\/s42269-023-01004-1","volume":"47","author":"W Seyoum","year":"2023","unstructured":"Seyoum, W. & Tora, E. Foot and mouth disease in Ethiopia: A systematic review and meta-analysis in the last fifteen years (2007\u20132021). Bull. Natl. Res. Centre. 47 (1), 32 (2023).","journal-title":"Bull. Natl. Res. Centre"},{"issue":"5","key":"64985_CR15","doi-asserted-by":"publisher","first-page":"e1393","DOI":"10.1111\/tbed.14472","volume":"69","author":"YA Raouf","year":"2022","unstructured":"Raouf, Y. A. et al. Genotyping of foot-and-mouth disease viruses collected in Sudan between 2009 and 2018. Transbound. Emerg. Dis. 69 (5), e1393\u2013e1406 (2022).","journal-title":"Transbound. Emerg. Dis."},{"issue":"12","key":"64985_CR16","doi-asserted-by":"publisher","first-page":"435","DOI":"10.1007\/s00284-024-03944-x","volume":"81","author":"RT Hassanein","year":"2024","unstructured":"Hassanein, R. T. et al. Epidemiological and genetic insights of the circulating foot-and-mouth disease virus serotypes in Egypt. Curr. Microbiol. 81 (12), 435 (2024).","journal-title":"Curr. Microbiol."},{"issue":"1","key":"64985_CR17","doi-asserted-by":"publisher","first-page":"395","DOI":"10.1186\/s12917-025-04834-5","volume":"21","author":"GG Dagnaw","year":"2025","unstructured":"Dagnaw, G. G. & Dejene, H. Seroprevalence of foot and mouth disease in cattle in East Africa between 2014 and 2024: A systematic review and meta-analysis. BMC Vet. Res. 21 (1), 395 (2025).","journal-title":"BMC Vet. Res."},{"key":"64985_CR18","doi-asserted-by":"publisher","first-page":"100188","DOI":"10.1016\/j.vas.2021.100188","volume":"13","author":"NH Ahmed","year":"2021","unstructured":"Ahmed, N. H. et al. Serological detection and genetic characterization of foot-and-mouth disease virus from cattle in northern Sudan, 2016\u20132018. Vet. Anim. Sci. 13, 100188 (2021).","journal-title":"Vet. Anim. Sci."},{"key":"64985_CR19","first-page":"0NF","volume":"GU24","author":"K Bankowska","year":"2017","unstructured":"Bankowska, K. FAO world reference laboratory for foot-and-mouth disease (WRLFMD) genotyping report. Woking GU24, 0NF (2017).","journal-title":"Woking"},{"issue":"4","key":"64985_CR20","doi-asserted-by":"publisher","first-page":"343","DOI":"10.1007\/s11259-021-09815-1","volume":"45","author":"W Alfouz","year":"2021","unstructured":"Alfouz, W. et al. Sero-epidemiology of foot-and-mouth disease in Darfur area, western Sudan. Vet. Res. Commun. 45 (4), 343\u2013352 (2021).","journal-title":"Vet. Res. Commun."},{"issue":"33","key":"64985_CR21","doi-asserted-by":"publisher","first-page":"968","DOI":"10.1136\/vr.77.33.968","volume":"77","author":"P Sutmoller","year":"1965","unstructured":"Sutmoller, P. & Gaggero, A. Foot-and mouth diseases carriers. Vet. Rec. 77 (33), 968\u2013969 (1965).","journal-title":"Vet. Rec"},{"issue":"1","key":"64985_CR22","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1186\/1297-9716-44-12","volume":"44","author":"MJ Morelli","year":"2013","unstructured":"Morelli, M. J. et al. Evolution of foot-and-mouth disease virus intra-sample sequence diversity during serial transmission in bovine hosts. Vet. Res. 44 (1), 12 (2013).","journal-title":"Vet. Res."},{"key":"64985_CR23","unstructured":"World Organisation for Animal Health. Manual of Diagnostic Tests and Vaccines for Terrestrial Animals: Foot-and-Mouth Disease (World Organisation for Animal Health, 2023)."},{"key":"64985_CR24","doi-asserted-by":"crossref","unstructured":"Romey, A. et al. Field Evaluation of a Safe, Easy, and Low-Cost Protocol for Shipment of Samples from Suspected Cases of Foot-and-Mouth Disease to Diagnostic Laboratories. Vol. 2023(1). 9555213 (Transboundary and Emerging Diseases, 2023).","DOI":"10.1155\/2023\/9555213"},{"issue":"5040","key":"64985_CR25","doi-asserted-by":"publisher","first-page":"1079","DOI":"10.1038\/2101079a0","volume":"210","author":"WA Snowdon","year":"1966","unstructured":"Snowdon, W. A. Growth of foot-and mouth disease virus in monolayer cultures of calf thyroid cells. Nature 210 (5040), 1079\u20131080 (1966).","journal-title":"Nature"},{"issue":"3","key":"64985_CR26","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1016\/0378-1135(88)90024-7","volume":"16","author":"NP Ferris","year":"1988","unstructured":"Ferris, N. P. & Dawson, M. Routine application of enzyme-linked immunosorbent assay in comparison with complement fixation for the diagnosis of foot-and-mouth and swine vesicular diseases. Vet. Microbiol. 16 (3), 201\u2013209 (1988).","journal-title":"Vet. Microbiol."},{"issue":"3","key":"64985_CR27","doi-asserted-by":"publisher","first-page":"513","DOI":"10.1017\/S002217240002636X","volume":"83","author":"JR Crowther","year":"1979","unstructured":"Crowther, J. R. & Abu Elzein, E. M. Application of the enzyme linked immunosorbent assay to the detection and identification of foot-and-mouth disease viruses. J. Hyg. (Lond). 83 (3), 513\u2013519 (1979).","journal-title":"J. Hyg. (Lond)"},{"issue":"11","key":"64985_CR28","doi-asserted-by":"publisher","first-page":"1636","DOI":"10.2460\/javma.2002.220.1636","volume":"220","author":"JD Callahan","year":"2002","unstructured":"Callahan, J. D. et al. Use of a portable real-time reverse transcriptase-polymerase chain reaction assay for rapid detection of foot-and-mouth disease virus. J. Am. Vet. Med. Assoc. 220 (11), 1636\u20131642 (2002).","journal-title":"J. Am. Vet. Med. Assoc."},{"issue":"3","key":"64985_CR29","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1177\/104063870902100303","volume":"21","author":"SM Reid","year":"2009","unstructured":"Reid, S. M. et al. Performance of real-time reverse transcription polymerase chain reaction for the detection of foot-and-mouth disease virus during field outbreaks in the United Kingdom in 2007. J. Vet. Diagn. Invest. 21 (3), 321\u2013330 (2009).","journal-title":"J. Vet. Diagn. Invest."},{"issue":"3","key":"64985_CR30","doi-asserted-by":"publisher","first-page":"792","DOI":"10.1093\/molbev\/msi066","volume":"22","author":"BG Hall","year":"2004","unstructured":"Hall, B. G. Comparison of the accuracies of several phylogenetic methods using protein and DNA sequences. Mol. Biol. Evol. 22 (3), 792\u2013802 (2004).","journal-title":"Mol. Biol. Evol."},{"issue":"12","key":"64985_CR31","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1093\/molbev\/msae263","volume":"41","author":"S Kumar","year":"2024","unstructured":"Kumar, S. et al. MEGA12: Molecular evolutionary genetic analysis version 12 for adaptive and green computing. Mol. Biol. Evol. 41 (12), 1\u20139 (2024).","journal-title":"Mol. Biol. Evol."},{"key":"64985_CR32","unstructured":"Felsenstein, J. PHYLIP (Phylogeny Inference Package) Version 3.6. Seattle: Department of Genome Sciences (University of Washington, distributed by the author, 2005)."},{"issue":"1","key":"64985_CR33","doi-asserted-by":"publisher","first-page":"116","DOI":"10.1186\/1297-9716-44-116","volume":"44","author":"SM Jamal","year":"2013","unstructured":"Jamal, S. M. & Belsham, G. J. Foot-and-mouth disease: Past, present and future. Vet. Res. 44 (1), 116 (2013).","journal-title":"Vet. Res."},{"issue":"3\u20134","key":"64985_CR34","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1016\/j.prevetmed.2013.07.013","volume":"112","author":"TJ Knight-Jones","year":"2013","unstructured":"Knight-Jones, T. J. & Rushton, J. The economic impacts of foot and mouth disease - What are they, how big are they and where do they occur? Prev. Vet. Med. 112 (3\u20134), 161\u2013173 (2013).","journal-title":"Prev. Vet. Med."},{"key":"64985_CR35","doi-asserted-by":"crossref","unstructured":"Foglia, E. A. et al. Combining Multiple Assays Improves Detection and Serotyping of Foot-and-Mouth Disease Virus. A Practical Example with Field Samples from East Africa. Viruses 13(8) (2021).","DOI":"10.3390\/v13081583"},{"key":"64985_CR36","unstructured":"Saeed, H. Epidemiological study of foot and mouth disease in cattle in Northern State of the Sudan. Master Degree Thesis, Sudan Academy of Sciences (2019)."},{"issue":"6","key":"64985_CR37","doi-asserted-by":"publisher","first-page":"e246","DOI":"10.1111\/tbed.12338","volume":"63","author":"WT Jemberu","year":"2016","unstructured":"Jemberu, W. T. et al. Epidemiology of foot and mouth disease in Ethiopia: A retrospective analysis of district level outbreaks, 2007\u20132012. Transbound. Emerg. Dis. 63 (6), e246\u2013e259 (2016).","journal-title":"Transbound. Emerg. Dis."},{"issue":"5","key":"64985_CR38","doi-asserted-by":"publisher","first-page":"2108","DOI":"10.1128\/JCM.42.5.2108-2114.2004","volume":"42","author":"BC Bronsvoort","year":"2004","unstructured":"Bronsvoort, B. C. et al. Comparison of two 3ABC enzyme-linked immunosorbent assays for diagnosis of multiple-serotype foot-and-mouth disease in a cattle population in an area of endemicity. J. Clin. Microbiol. 42 (5), 2108\u20132114 (2004).","journal-title":"J. Clin. Microbiol."},{"key":"64985_CR39","doi-asserted-by":"crossref","unstructured":"Raouf, Y. et al. The prevalence of antibodies against SAT1 foot-and-mouth disease virus in cattle in Khartoum state: Epidemiological significance. In Bulletin of Animal Health and Production in Africa. Vol. 57(4) (2009).","DOI":"10.4314\/bahpa.v57i4.51680"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-64985-9","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-64985-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-64985-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T15:00:47Z","timestamp":1788274847000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-64985-9"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,16]]},"references-count":39,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["64985"],"URL":"https:\/\/doi.org\/10.1038\/s41598-026-64985-9","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,16]]},"assertion":[{"value":"5 March 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 July 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"16 August 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"27403"}}