{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,16]],"date-time":"2026-10-16T18:21:54Z","timestamp":1792174914192,"version":"4.4.0"},"reference-count":47,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,11,25]],"date-time":"2025-11-25T00:00:00Z","timestamp":1764028800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,11,25]],"date-time":"2025-11-25T00:00:00Z","timestamp":1764028800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    <jats:italic>Toxoplasma gondii<\/jats:italic>\n                    is a globally distributed apicomplexan protozoan of zoonotic importance. Domestic cats (\n                    <jats:italic>Felis catus<\/jats:italic>\n                    ) and related felids constitute its sole definitive hosts, while numerous warm-blooded species serve as intermediate hosts\n                    <jats:italic>.<\/jats:italic>\n                    Infection outcomes for intermediate hosts range from latent toxoplasmosis to behaviour alteration and fatal systemic disease. Oocysts, the lifecycle environmental stage, are often washed into watercourses, making river-dwelling species like Eurasian beavers (\n                    <jats:italic>Castor fiber<\/jats:italic>\n                    ) particularly at risk. To investigate the presence and impact of\n                    <jats:italic>T. gondii<\/jats:italic>\n                    on beavers in the United Kingdom, archived brain, heart, and liver from 23 beavers necropsied between 2021 and 2023 were screened for\n                    <jats:italic>T. gondii<\/jats:italic>\n                    DNA by nested- and quantitative PCR. Positive samples were genotyped using a panel of six published PCR-Restriction Fragment Length Polymorphism (PCR-RFLP) assays and amplicon sequencing. Sequences were aligned with those reported previously from UK and other wildlife species to assess the occurrence of shared genotypes.\n                    <jats:italic>Toxoplasma gondii<\/jats:italic>\n                    DNA was detected in 19\/23 (83%) Eurasian beavers; all consistent with PCR-RFLP ToxoDB#3 with limited genetic diversity, concurring with findings from previous surveillance of UK wildlife species. Following histopathological screening non-suppurative toxoplasmosis was characterised in 3\/6 tissues selected from two animals. These results demonstrate a high occurrence of\n                    <jats:italic>T. gondii<\/jats:italic>\n                    infection in UK beavers\n                  <\/jats:p>","DOI":"10.1038\/s41598-025-26504-0","type":"journal-article","created":{"date-parts":[[2025,11,25]],"date-time":"2025-11-25T07:35:11Z","timestamp":1764056111000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Occurrence and characterisation of Toxoplasma gondii infection in free-living Eurasian beavers, Castor fiber, from southern England"],"prefix":"10.1038","volume":"15","author":[{"given":"Ambre","family":"Jacquart","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Damer P.","family":"Blake","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Simon","family":"Spiro","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"J. M. Jaramillo","family":"Ortiz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rebecca","family":"Foskett","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sophie M.","family":"Common","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,11,25]]},"reference":[{"key":"26504_CR1","unstructured":"Howe, C. & Dalton, L. Beavering away: The reintroduction of Beavers in England. In Pract. (2023)."},{"key":"26504_CR2","unstructured":"Common, S., Donald, H. & Sainsbury, A. Revised disease risk analysis for the conservation translocation of the Eurasian Beaver (Castor fiber) to England [NECR540]. Nat. Engl. (2024)."},{"key":"26504_CR3","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1017\/S0031182097008949","volume":"115","author":"JP Dubey","year":"1997","unstructured":"Dubey, J. P. Tissue cyst tropism in Toxoplasma gondii: A comparison of tissue cyst formation in organs of cats, and rodents fed oocysts. Parasitology 115, 15\u201320 (1997).","journal-title":"Parasitology"},{"key":"26504_CR4","doi-asserted-by":"publisher","first-page":"264","DOI":"10.1128\/CMR.05013-11","volume":"25","author":"F Robert-Gangneux","year":"2012","unstructured":"Robert-Gangneux, F. & Dard\u00e9, M. L. Epidemiology of and diagnostic strategies for toxoplasmosis. Clin. Microbiol. Rev. 25, 264\u2013296 (2012).","journal-title":"Clin. Microbiol. Rev."},{"key":"26504_CR5","doi-asserted-by":"publisher","first-page":"1","DOI":"10.7589\/JWD-D-23-00077","volume":"60","author":"P Scherrer","year":"2024","unstructured":"Scherrer, P., Frey, C. F. & Basso, W. Toxoplasma Gondii infection in the Eurasian Beaver (Castor fiber) in Switzerland: Seroprevalence genetic characterization, and clinicopathologic relevance. J. Wildl. Dis. 60, 1\u201313 (2024).","journal-title":"J. Wildl. Dis."},{"key":"26504_CR6","doi-asserted-by":"publisher","first-page":"404","DOI":"10.7589\/JWD-D-21-00097","volume":"58","author":"M Maas","year":"2022","unstructured":"Maas, M. et al. Zoonotic pathogens in Eurasian Beavers (Castor fiber) in the Netherlands. J. Wildl. Dis. 58, 404\u2013408 (2022).","journal-title":"J. Wildl. Dis."},{"key":"26504_CR7","doi-asserted-by":"publisher","first-page":"108","DOI":"10.1016\/j.vetpar.2012.08.026","volume":"191","author":"DC Herrmann","year":"2013","unstructured":"Herrmann, D. C., Wibbelt, G., G\u00f6tz, M., Conraths, F. J. & Schares, G. Genetic characterisation of Toxoplasma gondii isolates from European beavers (Castor fiber) and European wildcats (Felis silvestris silvestris). Vet. Parasitol. 191, 108\u2013111 (2013).","journal-title":"Vet. Parasitol."},{"key":"26504_CR8","doi-asserted-by":"publisher","first-page":"110","DOI":"10.1016\/j.ijppaw.2013.02.002","volume":"2","author":"T Hollings","year":"2013","unstructured":"Hollings, T., Jones, M., Mooney, N. & McCallum, H. Wildlife disease ecology in changing landscapes: Mesopredator release and toxoplasmosis. Int. J. Parasitol. Parasites Wildl. 2, 110\u2013118 (2013).","journal-title":"Int. J. Parasitol. Parasites Wildl."},{"key":"26504_CR9","doi-asserted-by":"publisher","first-page":"1065","DOI":"10.1080\/09273948.2023.2203215","volume":"32","author":"S Inchauspe","year":"2023","unstructured":"Inchauspe, S. et al. Association between ocular trauma and activation of ocular toxoplasmosis. Ocul. Immunol. Inflamm. 32, 1065\u20131069 (2023).","journal-title":"Ocul. Immunol. Inflamm."},{"key":"26504_CR10","doi-asserted-by":"publisher","first-page":"87351","DOI":"10.18632\/oncotarget.13568","volume":"7","author":"B Shen","year":"2016","unstructured":"Shen, B. et al. Activation of chronic toxoplasmosis by transportation stress in a mouse model. Oncotarget 7, 87351\u201387360 (2016).","journal-title":"Oncotarget"},{"key":"26504_CR11","doi-asserted-by":"publisher","first-page":"515","DOI":"10.1017\/S0031182001007624","volume":"122","author":"J Havl\u00edcek","year":"2001","unstructured":"Havl\u00edcek, J., Ga\u0161ov\u00e1, Z., Smith, A. P., Zv\u00e1ra, K. & Flegr, J. Decrease of psychomotor performance in subjects with latent \u2018asymptomatic\u2019 toxoplasmosis. Parasitology 122, 515\u2013520 (2001).","journal-title":"Parasitology"},{"key":"26504_CR12","doi-asserted-by":"crossref","unstructured":"Shapiro, K. et al. Type X strains of Toxoplasma gondii are virulent for southern sea otters (Enhydra lutris nereis) and present in felids from nearby watersheds (2019).","DOI":"10.1098\/rspb.2019.1334"},{"key":"26504_CR13","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/s13071-018-3256-z","volume":"12","author":"N Bachand","year":"2019","unstructured":"Bachand, N. et al. Serological and molecular detection of Toxoplasma gondii in terrestrial and marine wildlife harvested for food in Nunavik, Canada. Paras. Vectors 12, 1\u201310 (2019).","journal-title":"Paras. Vectors"},{"key":"26504_CR14","doi-asserted-by":"crossref","unstructured":"VanWormer, E. et al. Using molecular epidemiology to track Toxoplasma gondii from terrestrial carnivores to marine hosts: Implications for Public Health and Conservation. PLoS Negl. Trop. Dis. 8 (2014).","DOI":"10.1371\/journal.pntd.0002852"},{"key":"26504_CR15","doi-asserted-by":"publisher","first-page":"997","DOI":"10.1016\/S0020-7519(02)00069-3","volume":"32","author":"MA Miller","year":"2002","unstructured":"Miller, M. A. et al. Coastal freshwater runoff is a risk factor for Toxoplasma gondii infection of southern sea otters (Enhydra lutris nereis). Int. J. Parasitol. 32, 997\u20131006 (2002).","journal-title":"Int. J. Parasitol."},{"key":"26504_CR16","doi-asserted-by":"publisher","first-page":"4460","DOI":"10.1128\/AEM.64.11.4460-4466.1998","volume":"64","author":"GJ Medema","year":"1998","unstructured":"Medema, G. J., Schets, F. M., Teunis, P. F. M. & Havelaar, A. H. Sedimentation of free and attached Cryptosporidium oocysts and Giardia cysts in water. Appl. Environ. Microbiol. 64, 4460\u20134466 (1998).","journal-title":"Appl. Environ. Microbiol."},{"key":"26504_CR17","doi-asserted-by":"publisher","first-page":"674","DOI":"10.1644\/07-MAMM-A-168R.1","volume":"89","author":"CR Pavey","year":"2008","unstructured":"Pavey, C. R., Eldridge, S. R. & Heywood, M. Population dynamics and prey selection of native and introduced predators during a rodent outbreak in arid Australia. J. Mammal. 89, 674\u2013683 (2008).","journal-title":"J. Mammal."},{"key":"26504_CR18","doi-asserted-by":"publisher","first-page":"1768","DOI":"10.1017\/S0031182013001169","volume":"140","author":"A Burrells","year":"2013","unstructured":"Burrells, A. et al. Evidence of the three main clonal Toxoplasma gondii lineages from wild mammalian carnivores in the UK. Parasitology 140, 1768\u20131776 (2013).","journal-title":"Parasitology"},{"key":"26504_CR19","doi-asserted-by":"publisher","first-page":"828","DOI":"10.1111\/apha.12458","volume":"213","author":"J Xiao","year":"2015","unstructured":"Xiao, J. & Yolken, R. H. Strain hypothesis of Toxoplasma gondii infection on the outcome of human diseases. Acta Physiol. 213, 828\u2013845 (2015).","journal-title":"Acta Physiol."},{"key":"26504_CR20","unstructured":"Homan, W. L., Vercammen, M., De Braekeleer, J. & Verschueren, H. Identification of a 200-to 300-fold repetitive 529 bp DNA fragment in Toxoplasma gondii, and its use for diagnostic and quantitative PCR (2000)."},{"key":"26504_CR21","doi-asserted-by":"publisher","first-page":"5881","DOI":"10.1128\/JCM.43.12.5881-5887.2005","volume":"43","author":"A Khan","year":"2005","unstructured":"Khan, A. et al. Genotyping of Toxoplasma gondii strains from immunocompromised patients reveals high prevalence of type I strains. J. Clin. Microbiol. 43, 5881\u20135887 (2005).","journal-title":"J. Clin. Microbiol."},{"key":"26504_CR22","doi-asserted-by":"crossref","unstructured":"Dini, F. M. et al. Sentinels in the shadows: Exploring Toxoplasma gondii and other Sarcocystidae parasites in synanthropic rodents and their public health implications. Int. J. Parasitol. Parasites Wildl. 24 (2024).","DOI":"10.1016\/j.ijppaw.2024.100939"},{"key":"26504_CR23","first-page":"463","volume":"72","author":"DJ Ferguson","year":"1991","unstructured":"Ferguson, D. J., Graham, D. I. & Hutchison, W. M. Pathological changes in the brains of mice infected with Toxoplasma gondii: A histological, immunocytochemical and ultrastructural study. Int. J. Exp. Pathol. 72, 463\u2013474 (1991).","journal-title":"Int. J. Exp. Pathol."},{"key":"26504_CR24","doi-asserted-by":"publisher","first-page":"2749","DOI":"10.1098\/rstb.2009.0087","volume":"364","author":"DL Sibley","year":"2009","unstructured":"Sibley, D. L., Khan, A., Ajioka, J. W. & Rosenthal, B. M. Genetic diversity of Toxoplasma gondii in animals and humans. Philos. Trans. R. Soc. B Biol. Sci. 364, 2749\u20132761 (2009).","journal-title":"Philos. Trans. R. Soc. B Biol. Sci."},{"key":"26504_CR25","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/s13071-014-0608-1","volume":"8","author":"Q Liu","year":"2015","unstructured":"Liu, Q., Wang, Z. D., Huang, S. Y. & Zhu, X. Q. Diagnosis of toxoplasmosis and typing of Toxoplasma gondii. Paras. Vectors 8, 1\u201314 (2015).","journal-title":"Paras. Vectors"},{"key":"26504_CR26","doi-asserted-by":"crossref","unstructured":"Deagle, B. E., Eveson, J. P. & Jarman, S. N. Quantification of damage in DNA recovered from highly degraded samples\u2014A case study on DNA in faeces. Front. Zool. 3 (2006).","DOI":"10.1186\/1742-9994-3-11"},{"key":"26504_CR27","first-page":"1","volume":"4","author":"MM Krassner","year":"2023","unstructured":"Krassner, M. M. et al. Postmortem changes in brain cell structure: A review. Free Neuropathol. 4, 1\u201353 (2023).","journal-title":"Free Neuropathol."},{"key":"26504_CR28","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/1297-9716-42-121","volume":"42","author":"F Katzer","year":"2011","unstructured":"Katzer, F. et al. Increased Toxoplasma gondii positivity relative to age in 125 Scottish sheep flocks; Evidence of frequent acquired infection. Vet. Res. 42, 1\u20139 (2011).","journal-title":"Vet. Res."},{"key":"26504_CR29","doi-asserted-by":"publisher","first-page":"52","DOI":"10.2193\/2007-317","volume":"72","author":"SM Nusser","year":"2008","unstructured":"Nusser, S. M., Clark, W. R., Otis, D. L. & Huang, L. Sampling considerations for disease surveillance in wildlife populations. J. Wildl. Manage. 72, 52\u201360 (2008).","journal-title":"J. Wildl. Manage."},{"key":"26504_CR30","doi-asserted-by":"publisher","first-page":"4527","DOI":"10.1111\/1462-2920.12927","volume":"17","author":"C Krusor","year":"2015","unstructured":"Krusor, C. et al. Concentration and retention of Toxoplasma gondii oocysts by marine snails demonstrate a novel mechanism for transmission of terrestrial zoonotic pathogens in coastal ecosystems. Environ. Microbiol. 17, 4527\u20134537 (2015).","journal-title":"Environ. Microbiol."},{"key":"26504_CR31","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3389\/fmars.2020.00336","volume":"7","author":"J D\u00edaz-Delgado","year":"2020","unstructured":"D\u00edaz-Delgado, J. et al. Fatal systemic toxoplasmosis by a novel non-archetypal Toxoplasma gondii in a Bryde\u2019s Whale (Balaenoptera edeni). Front. Mar. Sci. 7, 1\u20139 (2020).","journal-title":"Front. Mar. Sci."},{"key":"26504_CR32","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1017\/S0031182009991065","volume":"137","author":"C Su","year":"2010","unstructured":"Su, C., Shwab, E. K., Zhou, P., Zhu, X. Q. & Dubey, J. P. Moving towards an integrated approach to molecular detection and identification of Toxoplasma gondii. Parasitology 137, 1\u201311 (2010).","journal-title":"Parasitology"},{"key":"26504_CR33","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/s13071-019-3862-4","volume":"13","author":"M Fern\u00e1ndez-Escobar","year":"2020","unstructured":"Fern\u00e1ndez-Escobar, M. et al. Isolation and genetic characterization of Toxoplasma gondii in Spanish sheep flocks. Paras. Vectors 13, 1\u201314 (2020).","journal-title":"Paras. Vectors"},{"key":"26504_CR34","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.ijppaw.2023.03.005","volume":"21","author":"P Scherrer","year":"2023","unstructured":"Scherrer, P. et al. Exploring the epidemiological role of the Eurasian lynx (Lynx lynx) in the life cycle of Toxoplasma gondii. Int. J. Parasitol. Parasites Wildl. 21, 1\u201310 (2023).","journal-title":"Int. J. Parasitol. Parasites Wildl."},{"key":"26504_CR35","doi-asserted-by":"publisher","first-page":"1365","DOI":"10.1645\/GE-1409.1","volume":"93","author":"JP Dubey","year":"2007","unstructured":"Dubey, J. P. et al. Mouse-virulent Toxoplasma gondii isolated from feral cats on Mona island. Puerto Rico. J. Parasitol. 93, 1365\u20131369 (2007).","journal-title":"Puerto Rico. J. Parasitol."},{"key":"26504_CR36","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1371\/journal.pone.0156255","volume":"11","author":"JP Dubey","year":"2016","unstructured":"Dubey, J. P. et al. Experimental toxoplasmosis in rats induced orally with eleven strains of Toxoplasma gondii of seven genotypes: Tissue tropism, tissue cyst size, neural lesions, tissue cyst rupture without reactivation, and ocular lesions. PLoS ONE 11, 1\u201326 (2016).","journal-title":"PLoS ONE"},{"key":"26504_CR37","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1007\/s13364-018-0405-x","volume":"64","author":"C Stefen","year":"2019","unstructured":"Stefen, C. Causes of death of beavers (Castor fiber) from eastern Germany and observations on parasites, skeletal diseases and tooth anomalies\u2014A long-term analysis. Mammal Res. 64, 279\u2013288 (2019).","journal-title":"Mammal Res."},{"key":"26504_CR38","doi-asserted-by":"publisher","first-page":"4177","DOI":"10.1111\/1462-2920.12685","volume":"17","author":"K Shapiro","year":"2015","unstructured":"Shapiro, K., Vanwormer, E., Aguilar, B. & Conrad, P. A. Surveillance for Toxoplasma gondii in California mussels (Mytilus californianus) reveals transmission of atypical genotypes from land to sea. Environ. Microbiol. 17, 4177\u20134188 (2015).","journal-title":"Environ. Microbiol."},{"key":"26504_CR39","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1016\/S0166-6851(01)00297-3","volume":"116","author":"A Jacquet","year":"2001","unstructured":"Jacquet, A. et al. The surface antigen SAG3 mediates the attachment of Toxoplasma gondii to cell-surface proteoglycans. Mol. Biochem. Parasitol. 116, 35\u201344 (2001).","journal-title":"Mol. Biochem. Parasitol."},{"key":"26504_CR40","doi-asserted-by":"publisher","first-page":"3","DOI":"10.25518\/1780-4507.13461","volume":"21","author":"F Debode","year":"2017","unstructured":"Debode, F., Marien, A., Janssen, \u00c9., Bragard, C. & Berben, G. The influence of amplicon length on real-time PCR results. Base 21, 3\u201311 (2017).","journal-title":"Base"},{"key":"26504_CR41","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1371\/journal.pntd.0009458","volume":"15","author":"IN Nzelu","year":"2021","unstructured":"Nzelu, I. N. et al. Detection and genetic characterisation of Toxoplasma gondii circulating in free-range chickens, pigs and seropositive pregnant women in benue state, Nigeria. PLoS Negl. Trop. Dis. 15, 1\u201319 (2021).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"26504_CR42","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1371\/journal.pntd.0001142","volume":"5","author":"AK Gibson","year":"2011","unstructured":"Gibson, A. K. et al. Polyparasitism is associated with increased disease severity in Toxoplasma gondii-infected marine sentinel species. PLoS Negl. Trop. Dis. 5, 14\u201316 (2011).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"26504_CR43","doi-asserted-by":"crossref","unstructured":"Gonzales-Viera, O. A. et al. Postmortem findings in free-ranging North American beavers (Castor canadensis) reveal potential threats to California\u2019s freshwater ecosystems. Animals 15 (2025).","DOI":"10.3390\/ani15030338"},{"key":"26504_CR44","doi-asserted-by":"crossref","unstructured":"Coombes, J. L. et al. Motile invaded neutrophils in the small intestine of Toxoplasma gondii-infected mice reveal a potential mechanism for parasite spread. Proc. Natl. Acad. Sci. USA. 110 (2013).","DOI":"10.1073\/pnas.1220272110"},{"key":"26504_CR45","doi-asserted-by":"crossref","unstructured":"Romito, G., Fracassi, F. & Cipone, M. Transient myocardial thickening associated with acute myocardial injury and congestive heart failure in two Toxoplasma gondii-positive cats. J. Feline Med. Surg. Open Reports 8 (2022).","DOI":"10.1177\/20551169221131266"},{"key":"26504_CR46","doi-asserted-by":"crossref","unstructured":"Campbell-Palmer, R. et al. Eurasian beaver (Castor fiber) health surveillance in Britain: Assessing a disjunctive reintroduced population. Vet. Rec. 188 (2021).","DOI":"10.1002\/vetr.84"},{"key":"26504_CR47","doi-asserted-by":"crossref","unstructured":"Zhang, Q. et al. Genomic expansion of Aldh1a1 protects beavers against high metabolic aldehydes from lipid oxidation. Cell Rep. 37 (2021).","DOI":"10.1016\/j.celrep.2021.109965"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-26504-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-26504-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-26504-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,11,27]],"date-time":"2025-11-27T13:27:16Z","timestamp":1764250036000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-26504-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,11,25]]},"references-count":47,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["26504"],"URL":"https:\/\/doi.org\/10.1038\/s41598-025-26504-0","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,11,25]]},"assertion":[{"value":"19 June 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 October 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"25 November 2025","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 authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}},{"value":"No live animals were used in these studies. All animal materials were collected from animals found dead and submitted to the DRAHS programme for routine surveillance\n                      .","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical review"}}],"article-number":"42516"}}