{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,12]],"date-time":"2026-02-12T09:02:20Z","timestamp":1770886940359,"version":"3.50.1"},"reference-count":96,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,11,7]],"date-time":"2025-11-07T00:00:00Z","timestamp":1762473600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2025,11,7]],"date-time":"2025-11-07T00:00:00Z","timestamp":1762473600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Commun"],"DOI":"10.1038\/s41467-025-64908-8","type":"journal-article","created":{"date-parts":[[2025,11,7]],"date-time":"2025-11-07T10:52:55Z","timestamp":1762512775000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["The contribution of mycetoma grains to suboptimal disease management"],"prefix":"10.1038","volume":"16","author":[{"given":"Ahmed","family":"Hassan Fahal","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Abdalla Osaman","family":"Ahmed","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3550-3428","authenticated-orcid":false,"given":"Lamyaa","family":"El Hassan","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3524-4825","authenticated-orcid":false,"given":"Ali Awadallah","family":"Saeed","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2025,11,7]]},"reference":[{"key":"64908_CR1","doi-asserted-by":"publisher","first-page":"e0003423","DOI":"10.1128\/cmr.00034-23","volume":"37","author":"WWJ Van de Sande","year":"2024","unstructured":"Van de Sande, W. W. J. & Fahal, A. H. An updated list of eumycetoma causative agents and their differences in grain formation and treatment response. Clin. Microbiol. Rev. 37, e0003423 (2024).","journal-title":"Clin. Microbiol. Rev."},{"key":"64908_CR2","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1016\/S0035-9203(03)00009-9","volume":"98","author":"AH Fahal","year":"2004","unstructured":"Fahal, A. H. Mycetoma: a thorn in the flesh. Trans. R. Soc. Trop. Med. Hyg. 98, 3\u201311 (2004).","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"64908_CR3","doi-asserted-by":"publisher","first-page":"e0004244","DOI":"10.1371\/journal.pntd.0004244","volume":"10","author":"EE Zijlstra","year":"2016","unstructured":"Zijlstra, E. E., van de Sande, W. W. & Fahal, A. H. Mycetoma: a long journey from neglect. PLoS Negl. Trop. Dis. 10, e0004244 (2016).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR4","doi-asserted-by":"publisher","first-page":"100","DOI":"10.1016\/S1473-3099(15)00359-X","volume":"16","author":"EE Zijlstra","year":"2016","unstructured":"Zijlstra, E. E. et al. Mycetoma: a unique neglected tropical disease. Lancet Infect. Dis. 16, 100\u2013112 (2016).","journal-title":"Lancet Infect. Dis."},{"key":"64908_CR5","doi-asserted-by":"publisher","first-page":"e0007019","DOI":"10.1371\/journal.pntd.0007019","volume":"12","author":"M Abbas","year":"2018","unstructured":"Abbas, M. et al. The disabling consequences of Mycetoma. PLoS Negl. Trop. Dis. 12, e0007019 (2018).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR6","doi-asserted-by":"publisher","first-page":"146","DOI":"10.1097\/QCO.0b013e32803d38fe","volume":"20","author":"AA Ahmed","year":"2007","unstructured":"Ahmed, A. A. et al. Management of mycetoma: major challenge in tropical mycoses with limited international recognition. Curr. Opin. Infect. Dis. 20, 146\u2013151 (2007).","journal-title":"Curr. Opin. Infect. Dis."},{"key":"64908_CR7","doi-asserted-by":"publisher","first-page":"639","DOI":"10.1016\/j.trstmh.2012.07.003","volume":"106","author":"HA Zein","year":"2012","unstructured":"Zein, H. A., Fahal, A. H., Mahgoub, S. E., El Hassan, T. A. & Abdel-Rahman, M. E. Predictors of cure, amputation and follow-up dropout among patients with mycetoma seen at the Mycetoma Research Centre, University of Khartoum, Sudan. Trans. R. Soc. Trop. Med. Hyg. 106, 639\u2013644 (2012).","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"64908_CR8","doi-asserted-by":"publisher","first-page":"e0004945","DOI":"10.1371\/journal.pntd.0004945","volume":"10","author":"NA Mohamed","year":"2016","unstructured":"Mohamed, N. A. & Fahal, A. H. Mycetoma pulmonary secondaries from a gluteal eumycetoma: an unusual presentation. PLoS Negl. Trop. Dis. 10, e0004945 (2016).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR9","doi-asserted-by":"publisher","first-page":"e0006391","DOI":"10.1371\/journal.pntd.0006391","volume":"12","author":"SM Bakhiet","year":"2018","unstructured":"Bakhiet, S. M. et al. A holistic approach to the mycetoma management. PLoS Negl. Trop. Dis. 12, e0006391 (2018).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR10","doi-asserted-by":"publisher","first-page":"e3271","DOI":"10.1371\/journal.pntd.0003271","volume":"8","author":"A Fahal","year":"2014","unstructured":"Fahal, A. et al. A new model for management of mycetoma in the Sudan. PLoS Negl. Trop. Dis. 8, e3271 (2014).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR11","doi-asserted-by":"publisher","first-page":"e0005638","DOI":"10.1371\/journal.pntd.0005638","volume":"11","author":"AA Ahmed","year":"2017","unstructured":"Ahmed, A. A., van de Sande, W. & Fahal, A. H. Mycetoma laboratory diagnosis: review article. PLoS Negl. Trop. Dis. 11, e0005638 (2017).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR12","doi-asserted-by":"publisher","first-page":"e2667","DOI":"10.1371\/journal.pntd.0002667","volume":"8","author":"WW Van de Sande","year":"2014","unstructured":"Van de Sande, W. W. et al. The mycetoma knowledge gap: identification of research priorities. PLoS Negl. Trop. Dis. 8, e2667 (2014).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR13","doi-asserted-by":"crossref","unstructured":"Fahal, A. H., Suliman, S. H. & Hay, R. Mycetoma: the spectrum of clinical presentation. Trop. Med. Infect. Dis. 3, 97 (2018).","DOI":"10.3390\/tropicalmed3030097"},{"key":"64908_CR14","doi-asserted-by":"crossref","unstructured":"Ahmed, A. et al. Environmental occurrence of Madurella mycetomatis, the major agent of human eumycetoma in Sudan. J. Clin. Microbiol. 40, 1031\u20131036 (2002).","DOI":"10.1128\/JCM.40.3.1031-1036.2002"},{"key":"64908_CR15","doi-asserted-by":"publisher","first-page":"12","DOI":"10.11599\/germs.2015.1066","volume":"5","author":"MF Bafghi","year":"2015","unstructured":"Bafghi, M. F. et al. Nocardia isolation from clinical samples with the paraffin baiting technique. Germs 5, 12\u201316 (2015).","journal-title":"Germs"},{"key":"64908_CR16","doi-asserted-by":"publisher","first-page":"175","DOI":"10.3114\/sim.2008.61.18","volume":"61","author":"H Badali","year":"2008","unstructured":"Badali, H. et al. Biodiversity of the genus Cladophialophora. Stud. Mycol. 61, 175\u2013191 (2008).","journal-title":"Stud. Mycol."},{"key":"64908_CR17","first-page":"26","volume":"36","author":"P Boiron","year":"1998","unstructured":"Boiron, P. et al. Nocardia, nocardiosis and mycetoma. Med. Mycol. 36, 26\u201337 (1998).","journal-title":"Med. Mycol."},{"key":"64908_CR18","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1093\/trstmh\/trab037","volume":"115","author":"A Bonifaz","year":"2021","unstructured":"Bonifaz, A. et al. Thoracic actinomycetoma: a retrospective clinical-epidemiological study of 64 cases. Trans. R. Soc. Trop. Med. Hyg. 115, 337\u2013339 (2021).","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"64908_CR19","doi-asserted-by":"publisher","first-page":"285","DOI":"10.4103\/idoj.IDOJ_474_20","volume":"12","author":"A Bonifaz","year":"2021","unstructured":"Bonifaz, A., Tirado-Sanchez, A., Vazquez-Gonzalez, D., Araiza, J. & Hernandez-Castro, R. Actinomycetoma by Actinomadura madurae: clinical characteristics and treatment of 47 cases. Indian Dermatol. Online J. 12, 285\u2013289 (2021).","journal-title":"Indian Dermatol. Online J."},{"key":"64908_CR20","doi-asserted-by":"publisher","first-page":"582","DOI":"10.1016\/S0035-9203(03)80036-6","volume":"97","author":"ME Abd Bagi","year":"2003","unstructured":"Abd Bagi, M. E. et al. Pathological fractures in mycetoma. Trans. R. Soc. Trop. Med. Hyg. 97, 582\u2013584 (2003).","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"64908_CR21","doi-asserted-by":"publisher","first-page":"665","DOI":"10.2214\/ajr.180.3.1800665","volume":"180","author":"ME Abd El Bagi","year":"2003","unstructured":"Abd El Bagi, M. E. New radiographic classification of bone involvement in pedal mycetoma. Am. J. Roentgenol. 180, 665\u2013668 (2003).","journal-title":"Am. J. Roentgenol."},{"key":"64908_CR22","first-page":"529","volume":"30","author":"ME Abd El-Bagi","year":"2009","unstructured":"Abd El-Bagi, M. E. & Fahal, A. H. Mycetoma revisited. Incidence of various radiographic signs. Saudi Med. J. 30, 529\u2013533 (2009).","journal-title":"Saudi Med. J."},{"key":"64908_CR23","doi-asserted-by":"crossref","unstructured":"Hay, R. J., Mahgoub, E. S., Leon, G., al-Sogair, S. & Welsh, O. Mycetoma. J. Med. Vet. Mycol. 30 (Suppl 1), 41\u201349 (1992).","DOI":"10.1080\/02681219280000751"},{"key":"64908_CR24","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1016\/j.clindermatol.2006.05.011","volume":"25","author":"O Welsh","year":"2007","unstructured":"Welsh, O., Vera-Cabrera, L. & Salinas-Carmona, M. C. Mycetoma. Clin. Dermatol. 25, 195\u2013202 (2007).","journal-title":"Clin. Dermatol."},{"key":"64908_CR25","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/0035-9203(72)90065-X","volume":"66","author":"AM El Hassan","year":"1972","unstructured":"El Hassan, A. M. & Mahgoub, E. S. Lymph node involvement in mycetoma. Trans. R. Soc. Trop. Med. Hyg. 66, 165\u2013169 (1972).","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"64908_CR26","doi-asserted-by":"publisher","first-page":"420","DOI":"10.1093\/trstmh\/traa181","volume":"115","author":"EA Ezaldeen","year":"2021","unstructured":"Ezaldeen, E. A., Ahmed, E. S. & Fahal, A. H. Massive complicated secondary inguinal mycetoma: a case series. Trans. R. Soc. Trop. Med. Hyg. 115, 420\u2013425 (2021).","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"64908_CR27","doi-asserted-by":"publisher","first-page":"e0010867","DOI":"10.1371\/journal.pntd.0010867","volume":"16","author":"SAY Adam","year":"2022","unstructured":"Adam, S. A. Y. et al. Eumycetoma with pulmonary dissemination an unusual complication: case series and literature review. PLoS Negl. Trop. Dis. 16, e0010867 (2022).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR28","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2334-11-290","volume":"11","author":"JA Almaguer-Chavez","year":"2011","unstructured":"Almaguer-Chavez, J. A. et al. Decrease of virulence for BALB\/c mice produced by continuous subculturing of Nocardia brasiliensis. BMC Infect. Dis. 11, 290 (2011).","journal-title":"BMC Infect. Dis."},{"key":"64908_CR29","doi-asserted-by":"publisher","first-page":"382","DOI":"10.1016\/j.clindermatol.2011.09.008","volume":"30","author":"L Vera-Cabrera","year":"2012","unstructured":"Vera-Cabrera, L. et al. Host defenses in subcutaneous mycoses. Clin. Dermatol. 30, 382\u2013388 (2012).","journal-title":"Clin. Dermatol."},{"key":"64908_CR30","doi-asserted-by":"publisher","first-page":"e0011736","DOI":"10.1371\/journal.pntd.0011736","volume":"17","author":"AH Fahal","year":"2023","unstructured":"Fahal, A. H. & Bakhiet, S. M. Mycetoma and the environment. PLoS Negl. Trop. Dis. 17, e0011736 (2023).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR31","doi-asserted-by":"publisher","first-page":"e3250","DOI":"10.1371\/journal.pntd.0003250","volume":"8","author":"AM Samy","year":"2014","unstructured":"Samy, A. M., van de Sande, W. W., Fahal, A. H. & Peterson, A. T. Mapping the potential risk of mycetoma infection in Sudan and South Sudan using ecological niche modeling. PLoS Negl. Trop. Dis. 8, e3250 (2014).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR32","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1093\/trstmh\/traa177","volume":"115","author":"A Santona","year":"2021","unstructured":"Santona, A. et al. Metagenomics of black grains: new highlights in the understanding of eumycetoma. Trans. R. Soc. Trop. Med. Hyg. 115, 307\u2013314 (2021).","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"64908_CR33","doi-asserted-by":"publisher","first-page":"e0010385","DOI":"10.1371\/journal.pntd.0010385","volume":"16","author":"A Santona","year":"2022","unstructured":"Santona, A. et al. Metagenomic detection of eumycetoma causative agents from households of patients residing in two Sudanese endemic villages in White Nile State. PLoS Negl. Trop. Dis. 16, e0010385 (2022).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR34","doi-asserted-by":"crossref","unstructured":"Hashizume, H. et al. Environmental detection of eumycetoma pathogens using multiplex real-time PCR for soil DNA in Sennar State, Sudan. Trop. Med. Health 51, 71 (2023).","DOI":"10.1186\/s41182-023-00563-3"},{"key":"64908_CR35","doi-asserted-by":"publisher","first-page":"e0010274","DOI":"10.1371\/journal.pntd.0010274","volume":"16","author":"H Hashizume","year":"2022","unstructured":"Hashizume, H. et al. Detection of multiple mycetoma pathogens using fungal metabarcoding analysis of soil DNA in an endemic area of Sudan. PLoS Negl. Trop. Dis. 16, e0010274 (2022).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR36","doi-asserted-by":"crossref","unstructured":"Kapinusova, G., Lopez Marin, M. A. & Uhlik, O. Reaching unreachables: Obstacles and successes of microbial cultivation and their reasons. Front. Microbiol. 14, 1089630 (2023).","DOI":"10.3389\/fmicb.2023.1089630"},{"key":"64908_CR37","doi-asserted-by":"publisher","first-page":"e57774","DOI":"10.1371\/journal.pone.0057774","volume":"8","author":"AI Ibrahim","year":"2013","unstructured":"Ibrahim, A. I., El Hassan, A. M., Fahal, A. & van de Sande, W. W. A histopathological exploration of the Madurella mycetomatis grain. PLoS ONE 8, e57774 (2013).","journal-title":"PLoS ONE"},{"key":"64908_CR38","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1080\/02681219580000041","volume":"33","author":"AH Fahal","year":"1995","unstructured":"Fahal, A. H., el Toum, E. A., El Hassan, A. M., Mahgoub, E. S. & Gumaa, S. A. The host tissue reaction to Madurella mycetomatis: new classification. J. Med. Vet. Mycol. 33, 15\u201317 (1995).","journal-title":"J. Med. Vet. Mycol."},{"key":"64908_CR39","doi-asserted-by":"publisher","first-page":"e0008053","DOI":"10.1371\/journal.pntd.0008053","volume":"14","author":"RS Ali","year":"2020","unstructured":"Ali, R. S., Newport, M. J., Bakhiet, S. M., Ibrahim, M. E. & Fahal, A. H. Host genetic susceptibility to mycetoma. PLoS Negl. Trop. Dis. 14, e0008053 (2020).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR40","doi-asserted-by":"crossref","unstructured":"Nyuykonge, B. et al. Madurella mycetomatis grains within a eumycetoma lesion are clonal. Med. Mycol. 60, myac051 (2022).","DOI":"10.1093\/mmy\/myac051"},{"key":"64908_CR41","doi-asserted-by":"publisher","first-page":"420","DOI":"10.1302\/0301-620X.96B3.31421","volume":"96-B","author":"AH Fahal","year":"2014","unstructured":"Fahal, A. H., Shaheen, S. & Jones, D. H. The orthopaedic aspects of mycetoma. Bone Jt. J. 96-B, 420\u2013425 (2014).","journal-title":"Bone Jt. J."},{"key":"64908_CR42","doi-asserted-by":"crossref","unstructured":"Mhmoud, N. A. et al. Pleurostomophora ochracea, a novel agent of human eumycetoma with yellow grains. J. Clin. Microbiol. 50, 2987\u20132994 (2012).","DOI":"10.1128\/JCM.01470-12"},{"key":"64908_CR43","doi-asserted-by":"publisher","first-page":"324","DOI":"10.1093\/trstmh\/traa168","volume":"115","author":"W Lim","year":"2021","unstructured":"Lim, W., Parel, F., de Hoog, S., Verbon, A. & van de Sande, W. W. J. Melanin production in coelomycetous agents of black grain eumycetoma. Trans. R. Soc. Trop. Med. Hyg. 115, 324\u2013327 (2021).","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"64908_CR44","doi-asserted-by":"crossref","unstructured":"Lim, W. et al. Inhibiting DHN- and DOPA-melanin biosynthesis pathway increased the therapeutic value of itraconazole in Madurella mycetomatis infected Galleria mellonella. Med. Mycol. 60, myac003 (2022).","DOI":"10.1093\/mmy\/myac003"},{"key":"64908_CR45","doi-asserted-by":"crossref","unstructured":"Nyuykonge, B., Croughs, P. D., Fahal, A. H., Verbon, A. & van de Sande, W. W. J. Correction for Nyuykonge et al., \u201cPyomelanin Secretion in Madurella mycetomatis Interferes with Spectrophotometric Endpoint Reading Using the Sensititre YeastOne alamarBlue Assay but Not with Visual Endpoint Reading\u201d. Antimicrob. Agents Chemother. 64, e01848-20 (2020).","DOI":"10.1128\/AAC.01848-20"},{"key":"64908_CR46","doi-asserted-by":"publisher","first-page":"1114","DOI":"10.1016\/j.micinf.2007.05.015","volume":"9","author":"WW Van de Sande","year":"2007","unstructured":"Van de Sande, W. W. et al. Melanin biosynthesis in Madurella mycetomatis and its effect on susceptibility to itraconazole and ketoconazole. Microbes Infect. 9, 1114\u20131123 (2007).","journal-title":"Microbes Infect."},{"key":"64908_CR47","first-page":"71","volume":"57","author":"SS Chouhan","year":"1969","unstructured":"Chouhan, S. S. & Agarwal, S. Histological diagnosis of mycetoma: a clinicopathological study of 24 cases. Indian J. Med. Res. 57, 71\u201377 (1969).","journal-title":"Indian J. Med. Res."},{"key":"64908_CR48","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1080\/00362177385190361","volume":"11","author":"ES Mahgoub","year":"1973","unstructured":"Mahgoub, E. S. Mycetomas caused by Curvularia lunata, Madurella grisea, Aspergillus nidulans, and Nocardia brasiliensis in Sudan. Sabouraudia 11, 179\u2013182 (1973).","journal-title":"Sabouraudia"},{"key":"64908_CR49","doi-asserted-by":"publisher","first-page":"415","DOI":"10.1007\/s11046-004-2844-y","volume":"158","author":"R Vilela","year":"2004","unstructured":"Vilela, R. et al. A case of eumycetoma due to Madurella grisea in northern Brazil. Mycopathologia 158, 415\u2013418 (2004).","journal-title":"Mycopathologia"},{"key":"64908_CR50","doi-asserted-by":"publisher","first-page":"617","DOI":"10.1016\/j.ijantimicag.2009.08.004","volume":"34","author":"D Cerar","year":"2009","unstructured":"Cerar, D., Malallah, Y. M., Howard, S. J., Bowyer, P. & Denning, D. W. Isolation, identification and susceptibility of Pyrenochaeta romeroi in a case of eumycetoma of the foot in the UK. Int. J. Antimicrob. Agents 34, 617\u2013618 (2009).","journal-title":"Int. J. Antimicrob. Agents"},{"key":"64908_CR51","doi-asserted-by":"crossref","unstructured":"Ma, J., Konings, M., Verbon, A. & van de Sande, W. W. J. A Falciformispora senegalensis grain model in Galleria mellonella larvae. Med. Mycol. 61, myad070 (2023).","DOI":"10.1093\/mmy\/myad070"},{"key":"64908_CR52","doi-asserted-by":"crossref","unstructured":"Ahmed, S. A. et al. Nigrograna mackinnonii, Not Trematosphaeria grisea (syn., Madurella grisea), Is the Main Agent of Black Grain Eumycetoma in Latin America. J. Clin. Microbiol. 56, e01723-17 (2018).","DOI":"10.1128\/JCM.01723-17"},{"key":"64908_CR53","doi-asserted-by":"publisher","first-page":"4537","DOI":"10.1128\/JCM.41.10.4537-4541.2003","volume":"41","author":"A Ahmed","year":"2003","unstructured":"Ahmed, A., van de Sande, W., Verbrugh, H., Fahal, A. & van Belkum, A. Madurella mycetomatis strains from mycetoma lesions in Sudanese patients are clonal. J. Clin. Microbiol. 41, 4537\u20134541 (2003).","journal-title":"J. Clin. Microbiol."},{"key":"64908_CR54","doi-asserted-by":"publisher","first-page":"e0003926","DOI":"10.1371\/journal.pntd.0003926","volume":"9","author":"W Kloezen","year":"2015","unstructured":"Kloezen, W., van Helvert-van Poppel, M., Fahal, A. H. & van de Sande, W. W. A Madurella mycetomatis Grain Model in Galleria mellonella Larvae. PLoS Negl. Trop. Dis. 9, e0003926 (2015).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR55","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1111\/j.1439-0507.2004.00964.x","volume":"47","author":"GS De Hoog","year":"2004","unstructured":"De Hoog, G. S., Adelmann, D., Ahmed, A. O. & van Belkum, A. Phylogeny and typification of Madurella mycetomatis, with a comparison of other agents of eumycetoma. Mycoses 47, 121\u2013130 (2004).","journal-title":"Mycoses"},{"key":"64908_CR56","doi-asserted-by":"crossref","unstructured":"Siddig, E. E. et al. The first case of Fusarium falciforme Eumycetoma in Sudan and an extensive literature review about treatment worldwide. J. Fungi 9, 730 (2023).","DOI":"10.3390\/jof9070730"},{"key":"64908_CR57","doi-asserted-by":"crossref","unstructured":"Badarol Hisham, N., Abdul Rhani, S., Mohd Ali, R. & Masri, S. N. Fusarium falciforme eumycetoma: a diagnostic challenge of a neglected tropical disease. BMJ Case Rep. 17, e258657 (2024).","DOI":"10.1136\/bcr-2023-258657"},{"key":"64908_CR58","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1128\/CMR.00039-07","volume":"21","author":"KJ Cortez","year":"2008","unstructured":"Cortez, K. J. et al. Infections caused by Scedosporium spp. Clin. Microbiol. Rev. 21, 157\u2013197 (2008).","journal-title":"Clin. Microbiol. Rev."},{"key":"64908_CR59","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1111\/j.1365-2133.1974.tb12899.x","volume":"91","author":"GH Findlay","year":"1974","unstructured":"Findlay, G. H. & Vismer, H. F. Black grain mycetoma. A study of the chemistry, formation and significance of the tissue grain in Madurella mycetomi infection. Br. J. Dermatol. 91, 297\u2013303 (1974).","journal-title":"Br. J. Dermatol."},{"key":"64908_CR60","doi-asserted-by":"publisher","first-page":"497","DOI":"10.1111\/j.1365-2133.1977.tb14125.x","volume":"97","author":"GH Findlay","year":"1977","unstructured":"Findlay, G. H. & Vismer, H. F. Black grain mycetoma. Atomic absorption and spark source mass spectrophotometry of the tissue grain in Madurella mycetomi infection. Br. J. Dermatol. 97, 497\u2013499 (1977).","journal-title":"Br. J. Dermatol."},{"key":"64908_CR61","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1007\/BF00436241","volume":"67","author":"GH Findlay","year":"1979","unstructured":"Findlay, G. H., Vismer, H. F. & Kiebenberg, N. W. Black grain mycetoma: the ultrastructure of Madurella mycetomi. Mycopathologia 67, 51\u201354 (1979).","journal-title":"Mycopathologia"},{"key":"64908_CR62","doi-asserted-by":"publisher","first-page":"e0008190","DOI":"10.1371\/journal.pntd.0008190","volume":"14","author":"G Sheehan","year":"2020","unstructured":"Sheehan, G. et al. Proteomic analysis of the processes leading to Madurella mycetomatis grain formation in Galleria mellonella larvae. PLoS Negl. Trop. Dis. 14, e0008190 (2020).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR63","doi-asserted-by":"publisher","first-page":"343","DOI":"10.1080\/02681219480000461","volume":"32","author":"AH Fahal","year":"1994","unstructured":"Fahal, A. H., el Toum, E. A., el Hassan, A. M., Mahgoub, E. S. & Gumaa, S. A. A preliminary study on the ultrastructure of Actinomadura pelletieri and its host tissue reaction. J. Med. Vet. Mycol. 32, 343\u2013348 (1994).","journal-title":"J. Med. Vet. Mycol."},{"key":"64908_CR64","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1016\/S0928-8244(03)00096-8","volume":"37","author":"AO Ahmed","year":"2003","unstructured":"Ahmed, A. O. et al. A murine model of Madurella mycetomatis eumycetoma. FEMS Immunol. Med. Microbiol. 37, 29\u201336 (2003).","journal-title":"FEMS Immunol. Med. Microbiol."},{"key":"64908_CR65","doi-asserted-by":"publisher","first-page":"341","DOI":"10.1016\/0021-9975(81)90003-7","volume":"91","author":"SA Gumaa","year":"1981","unstructured":"Gumaa, S. A. & Abu-Samra, M. T. Experimental mycetoma infection in the goat. J. Comp. Pathol. 91, 341\u2013346 (1981).","journal-title":"J. Comp. Pathol."},{"key":"64908_CR66","first-page":"773","volume":"99","author":"A Gonzalez-Ochoa","year":"1969","unstructured":"Gonzalez-Ochoa, A. [Experimental production of mycetoma by Nocardia braziliensis in the mouse]. Gac. Med. Mex. 99, 773\u2013781 (1969).","journal-title":"Gac. Med. Mex."},{"key":"64908_CR67","doi-asserted-by":"publisher","first-page":"S45","DOI":"10.3109\/13693786.2010.517224","volume":"48","author":"A Harun","year":"2010","unstructured":"Harun, A., Serena, C., Gilgado, F., Chen, S. C. & Meyer, W. Scedosporium aurantiacum is as virulent as S. prolificans, and shows strain-specific virulence differences, in a mouse model. Med. Mycol. 48, S45\u2013S51 (2010).","journal-title":"Med. Mycol."},{"key":"64908_CR68","doi-asserted-by":"publisher","first-page":"233","DOI":"10.3390\/pathogens13030233","volume":"13","author":"A Giammarino","year":"2024","unstructured":"Giammarino, A., Bellucci, N. & Angiolella, L. Galleria mellonella as a model for the study of fungal pathogens: advantages and disadvantages. Pathogens 13, 233 (2024).","journal-title":"Pathogens"},{"key":"64908_CR69","doi-asserted-by":"publisher","first-page":"696","DOI":"10.1111\/tmi.12871","volume":"22","author":"K Eadie","year":"2017","unstructured":"Eadie, K., Parel, F., Helvert-van Poppel, M., Fahal, A. & van de Sande, W. Combining two antifungal agents does not enhance survival of Galleria mellonella larvae infected with Madurella mycetomatis. Trop. Med. Int. Health 22, 696\u2013702 (2017).","journal-title":"Trop. Med. Int. Health"},{"key":"64908_CR70","doi-asserted-by":"publisher","first-page":"641258","DOI":"10.3389\/fmicb.2021.641258","volume":"12","author":"MQ Granato","year":"2021","unstructured":"Granato, M. Q. et al. Silver(I) and copper(II) complexes of 1,10-phenanthroline-5,6-dione against Phialophora verrucosa: a focus on the interaction with human macrophages and Galleria mellonella Larvae. Front. Microbiol. 12, 641258 (2021).","journal-title":"Front. Microbiol."},{"key":"64908_CR71","doi-asserted-by":"publisher","first-page":"469","DOI":"10.1093\/mmy\/myx064","volume":"56","author":"W Kloezen","year":"2018","unstructured":"Kloezen, W. et al. Amphotericin B and terbinafine but not the azoles prolong survival in Galleria mellonella larvae infected with Madurella mycetomatis. Med. Mycol. 56, 469\u2013478 (2018).","journal-title":"Med. Mycol."},{"key":"64908_CR72","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1016\/j.pdpdt.2018.12.010","volume":"25","author":"MH Paziani","year":"2019","unstructured":"Paziani, M. H. et al. Antimicrobial photodynamic therapy with phenothiazinium photosensitizers in non-vertebrate model Galleria mellonella infected with Fusarium keratoplasticum and Fusarium moniliforme. Photodiagnosis Photodyn. Ther. 25, 197\u2013203 (2019).","journal-title":"Photodiagnosis Photodyn. Ther."},{"key":"64908_CR73","doi-asserted-by":"publisher","first-page":"174","DOI":"10.4269\/ajtmh.1987.37.174","volume":"37","author":"M Nasher","year":"1987","unstructured":"Nasher, M., Wethered, D., Hay, R. J., Mahgoub, E. S. & Gumaa, S. A. The ultrastructure of actinomycetoma grains caused by Streptomyces somaliensis. Am. J. Trop. Med. Hyg. 37, 174\u2013179 (1987).","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"64908_CR74","doi-asserted-by":"publisher","first-page":"406","DOI":"10.1093\/trstmh\/traa145","volume":"115","author":"EE Siddig","year":"2021","unstructured":"Siddig, E. E. et al. Human actinomycetoma caused by Actinomadura mexicana in Sudan: the first report. Trans. R. Soc. Trop. Med. Hyg. 115, 406\u2013410 (2021).","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"64908_CR75","doi-asserted-by":"publisher","first-page":"1332","DOI":"10.1001\/archderm.1985.01660100112030","volume":"121","author":"W Sindhuphak","year":"1985","unstructured":"Sindhuphak, W., Macdonald, E. & Head, E. Actinomycetoma caused by Nocardiopsis dassonvillei. Arch. Dermatol. 121, 1332\u20131334 (1985).","journal-title":"Arch. Dermatol."},{"key":"64908_CR76","doi-asserted-by":"crossref","unstructured":"Bonifaz, A. et al. Actinomycetoma by Actinomadura madurae. Clinical and therapeutic characteristics of 18 cases with two treatment modalities. J. Dermatolog. Treat. 33, 954\u2013958 (2022).","DOI":"10.1080\/09546634.2020.1793887"},{"key":"64908_CR77","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1016\/S0035-9203(01)90346-3","volume":"95","author":"AM El Hassan","year":"2001","unstructured":"El Hassan, A. M., Fahal, A. H., Ahmed, A. O., Ismail, A. & Veress, B. The immunopathology of actinomycetoma lesions caused by Streptomyces somaliensis. Trans. R. Soc. Trop. Med. Hyg. 95, 89\u201392 (2001).","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"64908_CR78","first-page":"1","volume":"137","author":"MC Salinas-Carmona","year":"2001","unstructured":"Salinas-Carmona, M. C. Anti-Nocardia brasiliensis antibodies in patients with actinomycetoma and their clinical usefulness. Gac. Med. Mex. 137, 1\u20138 (2001).","journal-title":"Gac. Med. Mex."},{"key":"64908_CR79","doi-asserted-by":"publisher","first-page":"e0008775","DOI":"10.1371\/journal.pntd.0008775","volume":"14","author":"MC Salinas-Carmona","year":"2020","unstructured":"Salinas-Carmona, M. C. et al. Inducible nitric oxide synthase blockade with aminoguanidine, protects mice infected with Nocardia brasiliensis from actinomycetoma development. PLoS Negl. Trop. Dis. 14, e0008775 (2020).","journal-title":"PLoS Negl. Trop. Dis."},{"key":"64908_CR80","doi-asserted-by":"crossref","unstructured":"Salinas-Carmona, M. C., Torres-Lopez, E., Ramos, A. I., Licon-Trillo, A. & Gonzalez-Spencer, D. Immune response to Nocardia brasiliensis antigens in an experimental model of actinomycetoma in BALB\/c mice. Infect. Immun. 67, 2428\u20132432 (1999).","DOI":"10.1128\/IAI.67.5.2428-2432.1999"},{"key":"64908_CR81","doi-asserted-by":"publisher","first-page":"2239","DOI":"10.1099\/mic.0.037036-0","volume":"156","author":"P Tielen","year":"2010","unstructured":"Tielen, P. et al. Extracellular enzymes affect biofilm formation of mucoid Pseudomonas aeruginosa. Microbiology 156, 2239\u20132252 (2010).","journal-title":"Microbiology"},{"key":"64908_CR82","first-page":"91","volume":"15","author":"C Rodriguez Mdel","year":"1996","unstructured":"Rodriguez Mdel, C., Torres, J. A. & Zlotnik, H. Investigation of the temporal humoral immune response in a murine model of actinomycetoma. P R. Health Sci. J. 15, 91\u201395 (1996).","journal-title":"P R. Health Sci. J."},{"key":"64908_CR83","doi-asserted-by":"publisher","first-page":"204173141455711","DOI":"10.1177\/2041731414557112","volume":"5","author":"JK Kular","year":"2014","unstructured":"Kular, J. K., Basu, S. & Sharma, R. I. The extracellular matrix: Structure, composition, age-related differences, tools for analysis and applications for tissue engineering. J. Tissue Eng. 5, 2041731414557112 (2014).","journal-title":"J. Tissue Eng."},{"key":"64908_CR84","doi-asserted-by":"publisher","first-page":"127","DOI":"10.1007\/s11046-008-9093-4","volume":"165","author":"B Castro-Matteotti","year":"2008","unstructured":"Castro-Matteotti, B. et al. Immune response to Nocardia brasiliensis extracellular antigens in patients with mycetoma. Mycopathologia 165, 127\u2013134 (2008).","journal-title":"Mycopathologia"},{"key":"64908_CR85","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1111\/j.1574-695X.2009.00575.x","volume":"57","author":"ML Gonzalez-Suarez","year":"2009","unstructured":"Gonzalez-Suarez, M. L., Salinas-Carmona, M. C. & Perez-Rivera, I. IgM but not IgG monoclonal anti-Nocardia brasiliensis antibodies confer protection against experimental actinomycetoma in BALB\/c mice. FEMS Immunol. Med. Microbiol. 57, 17\u201324 (2009).","journal-title":"FEMS Immunol. Med. Microbiol."},{"key":"64908_CR86","doi-asserted-by":"publisher","first-page":"e100064","DOI":"10.1371\/journal.pone.0100064","volume":"9","author":"I Meester","year":"2014","unstructured":"Meester, I., Rosas-Taraco, A. G. & Salinas-Carmona, M. C. Nocardia brasiliensis induces formation of foamy macrophages and dendritic cells in vitro and in vivo. PLoS ONE 9, e100064 (2014).","journal-title":"PLoS ONE"},{"key":"64908_CR87","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1111\/j.1749-6632.1996.tb52972.x","volume":"797","author":"MC Salinas-Carmona","year":"1996","unstructured":"Salinas-Carmona, M. C. & Torres-Lopez, E. Role of passive humoral immunity in experimental mycetoma by Nocardia brasiliensis. Ann. N. Y. Acad. Sci. 797, 263\u2013265 (1996).","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"64908_CR88","doi-asserted-by":"crossref","unstructured":"Vazquez-Marmolejo, A. V. et al. Nitric oxide determines the development of actinomycetoma by Nocardia brasiliensis in eNOS knockout C57BL\/6 mice. FEMS Microbiol. Lett. 368, fnab048 (2021).","DOI":"10.1093\/femsle\/fnab048"},{"key":"64908_CR89","doi-asserted-by":"publisher","first-page":"918","DOI":"10.1016\/0035-9203(88)90042-9","volume":"82","author":"DB Wethered","year":"1988","unstructured":"Wethered, D. B., Markey, M. A., Hay, R. J., Mahgoub, E. S. & Gumaa, S. A. Humoral immune responses to mycetoma organisms: characterization of specific antibodies by the use of enzyme-linked immunosorbent assay and immunoblotting. Trans. R. Soc. Trop. Med. Hyg. 82, 918\u2013923 (1988).","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"64908_CR90","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1111\/j.1365-4362.1968.tb05624.x","volume":"7","author":"L Ajello","year":"1968","unstructured":"Ajello, L. & Basom, W. C. A Mexican case of mycetoma caused by Streptomyces somaliensis. Dermatol. Int. 7, 17\u201322 (1968).","journal-title":"Dermatol. Int."},{"key":"64908_CR91","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1007\/BF00437238","volume":"97","author":"A Kamalam","year":"1987","unstructured":"Kamalam, A. & Thambiah, A. S. A clinico-pathological study of actinomycotic mycetomas caused by Actinomadura madurae and Actinomadura pelletierii. Mycopathologia 97, 151\u2013163 (1987).","journal-title":"Mycopathologia"},{"key":"64908_CR92","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1111\/j.1440-0960.1990.tb00648.x","volume":"31","author":"PV Venugopal","year":"1990","unstructured":"Venugopal, P. V. & Venugopal, T. V. Actinomadura madurae mycetomas. Australas. J. Dermatol. 31, 33\u201336 (1990).","journal-title":"Australas. J. Dermatol."},{"key":"64908_CR93","first-page":"301","volume":"28","author":"P Dockx","year":"1972","unstructured":"Dockx, P. The ultrastructure of the elementary lesion of actinomycotic mycetoma by Nocardia asteroides. Arch. Belg. Dermatol. Syphiligr. 28, 301\u2013313 (1972).","journal-title":"Arch. Belg. Dermatol. Syphiligr."},{"key":"64908_CR94","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1016\/S0928-8244(03)00102-0","volume":"37","author":"A Licon-Trillo","year":"2003","unstructured":"Licon-Trillo, A., Angeles Castro-Corona, M. & Salinas-Carmona, M. C. Immunogenicity and biophysical properties of a Nocardia brasiliensis protease involved in pathogenesis of mycetoma. FEMS Immunol. Med. Microbiol. 37, 37\u201344 (2003).","journal-title":"FEMS Immunol. Med. Microbiol."},{"key":"64908_CR95","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1080\/00362176785190171","volume":"5","author":"E Macotela-Ruiz","year":"1966","unstructured":"Macotela-Ruiz, E. & Gonzalez-Angulo, A. Electron microscopic studies on granules of Nocardia brasiliensis in man. Sabouraudia 5, 92\u201398 (1966).","journal-title":"Sabouraudia"},{"key":"64908_CR96","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1007\/BF00444033","volume":"105","author":"A Van Gelderen de Komaid","year":"1989","unstructured":"Van Gelderen de Komaid, A. Physiological and pathogenic characteristics of Nocardia brasiliensis isolated from human mycetomas. Mycopathologia 105, 111\u2013116 (1989).","journal-title":"Mycopathologia"}],"updated-by":[{"DOI":"10.1038\/s41467-026-69568-w","type":"correction","label":"Correction","source":"publisher","updated":{"date-parts":[[2026,2,11]],"date-time":"2026-02-11T00:00:00Z","timestamp":1770768000000}}],"container-title":["Nature Communications"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41467-025-64908-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41467-025-64908-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41467-025-64908-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,12]],"date-time":"2026-02-12T08:02:48Z","timestamp":1770883368000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41467-025-64908-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,11,7]]},"references-count":96,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["64908"],"URL":"https:\/\/doi.org\/10.1038\/s41467-025-64908-8","relation":{},"ISSN":["2041-1723"],"issn-type":[{"value":"2041-1723","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,11,7]]},"assertion":[{"value":"28 June 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 September 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"7 November 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 February 2026","order":5,"name":"change_date","label":"Change Date","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Correction","order":6,"name":"change_type","label":"Change Type","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"A Correction to this paper has been published:","order":7,"name":"change_details","label":"Change Details","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"https:\/\/doi.org\/10.1038\/s41467-026-69568-w","URL":"https:\/\/doi.org\/10.1038\/s41467-026-69568-w","order":8,"name":"change_details","label":"Change Details","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"9855"}}