{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T05:14:41Z","timestamp":1788498881329,"version":"build-2803163510"},"reference-count":89,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T00:00:00Z","timestamp":1788480000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T00:00:00Z","timestamp":1788480000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"German Federal Ministry of Agriculture, Food and Regional Identity (BMLEH) through the Federal Office for Agriculture and Food"},{"name":"Leibniz-Zentrum f\u00fcr Agrarlandschaftsforschung (ZALF) e.V."}],"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                      Despite its importance for\n                      <jats:italic>Culicoides<\/jats:italic>\n                      -borne disease risk assessment, the winter activity of species of this biting midge genus remains poorly characterized. Moreover, information on virus circulation in winter-active\n                      <jats:italic>Culicoides<\/jats:italic>\n                      vectors, both under endemic conditions and during outbreaks, is almost non-existent.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      UV-light traps to collect\n                      <jats:italic>Culicoides<\/jats:italic>\n                      biting midges were operated once a week for 24\u00a0h in cattle, sheep, goat and horse stables at 12 to 55 locations during four consecutive winter periods (November 1 to March 31, 2021 to 2025) throughout Germany. Collected females of the genus\n                      <jats:italic>Culicoides<\/jats:italic>\n                      were morphologically assigned to Obsoletus Group or Pulicaris Complex. Temperature data were recorded to relate biting midge activity to temperatures. Screening for bluetongue virus (BTV), Schmallenberg virus and epizootic hemorrhagic disease virus was done by real-time RT-PCR on 1,581 pools (46,358 females) of both groups collected before (winter 2022\/2023) and during the BTV serotype 3 (BTV-3) outbreak (winters 2023\/2024 and 2024\/2025).\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      <jats:italic>Culicoides<\/jats:italic>\n                      collections displayed notable variation between sites and months. November and March accounted for 95% of captures, while mid-winter months (December to February) yielded only 5%, with January and February each contributing 1% of total captures. The March 2024 and 2025 activity peaks coincided with mild temperatures and the inclusion of new monitoring sites, suggesting that one or both factors contributed. Although blood-fed females were very rare during mid-winter, their abundance often exceeded the EU threshold for parous\n                      <jats:italic>Culicoides<\/jats:italic>\n                      to define a vector-free period of\u2009&lt;\u20095 individuals per trap and night. However, only a single\n                      <jats:italic>Culicoides<\/jats:italic>\n                      pool from November 2024 tested positive for BTV-3-RNA. All other pools were negative for all three viruses.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>\n                      Potential\n                      <jats:italic>Culicoides<\/jats:italic>\n                      vectors are active and feed on blood in livestock housings during wintertime. The presence of blood-fed females, exceeding the EU threshold, challenges the concept of a \u2018vector-free period\u2019. However, despite extensive monitoring over four years including outbreak conditions, only one virus-positive\n                      <jats:italic>Culicoides<\/jats:italic>\n                      pool was detected during winter. Combined with the very low number of biting\u00a0midges and the temperatures not allowing virus replication in arthropods, virus circulation during wintertime appears to be hardly existent even under endemic or outbreak conditions. Considering both\n                      <jats:italic>Culicoides<\/jats:italic>\n                      activity and virus circulation, the findings suggest that the risk of BTV infection in ruminants via biting midges during the central European winter (December to March) is negligible. BTV overwintering in central Europe likely depends on either rare infection events in ruminants and vectors during winter that sustain viremia until spring, or on persistence mechanisms that are currently unknown.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1186\/s13071-026-07620-w","type":"journal-article","created":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T02:34:21Z","timestamp":1788489261000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Winter activity and virus infection of Culicoides biting midges (Diptera: Ceratopogonidae) in German livestock housing before and during the 2024 BTV-3 outbreak"],"prefix":"10.1186","volume":"19","author":[{"given":"Anja","family":"Voigt","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Helge","family":"Kampen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Martin","family":"Beer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kerstin","family":"Wernike","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dorothee","family":"Scheuch","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Franziska","family":"Sick","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sophie","family":"Zeiske","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Doreen","family":"Werner","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,9,4]]},"reference":[{"key":"7620_CR1","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1146\/annurev.ento.45.1.307","volume":"45","author":"PS Mellor","year":"2000","unstructured":"Mellor PS, Boorman J, Baylis M. Culicoides biting midges: their role as arbovirus vectors. Annu Rev Entomol. 2000;45:307\u201340.","journal-title":"Annu Rev Entomol"},{"key":"7620_CR2","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1136\/vr.161.8.253","volume":"161","author":"KE Darpel","year":"2007","unstructured":"Darpel KE, Batten CA, Veronesi E, Shaw AE, Anthony S, Bachanek-Bankowska K, et al. Clinical signs and pathology shown by British sheep and cattle infected with bluetongue virus serotype 8 derived from the 2006 outbreak in northern Europe. Vet Rec. 2007;161:253\u201361.","journal-title":"Vet Rec"},{"key":"7620_CR3","doi-asserted-by":"publisher","first-page":"142","DOI":"10.1016\/j.tvjl.2009.05.004","volume":"182","author":"F Dal Pozzo","year":"2009","unstructured":"Dal Pozzo F, Saegerman C, Thiry E. Bovine infection with bluetongue virus with special emphasis on European serotype 8. Vet J. 2009;182:142\u201351.","journal-title":"Vet J"},{"key":"7620_CR4","unstructured":"European Commission. Regulation (EU) No. 2016\/429 of 9 March 2016 on transmissible animal diseases and amending and repealing certain acts in the area of animal health (\u2018Animal Health Law\u2019). Off J EU. 2016; L84:1\u2013208."},{"key":"7620_CR5","unstructured":"European Commission. Regulation (EU) No. 2020\/689 of 17 December 2019 supplementing Regulation (EU) 2016\/429 of the European Parliament and of the Council as regards rules for surveillance, eradication programs, and disease-free status for certain listed and emerging diseases. Off J EU. 2020; L174:211\u2013340."},{"key":"7620_CR6","doi-asserted-by":"publisher","first-page":"65","DOI":"10.3389\/fvets.2020.00065","volume":"7","author":"J Gethmann","year":"2020","unstructured":"Gethmann J, Probst C, Conraths FJ. Economic impact of a bluetongue serotype 8 epidemic in Germany. Front Vet Sci. 2020;7:65.","journal-title":"Front Vet Sci"},{"key":"7620_CR7","doi-asserted-by":"publisher","first-page":"294","DOI":"10.1016\/j.prevetmed.2009.11.007","volume":"93","author":"AG Velthuis","year":"2010","unstructured":"Velthuis AG, Saatkamp HW, Mourits MC, de Koeijer AA, Elbers AR. Financial consequences of the Dutch bluetongue serotype 8 epidemics of 2006 and 2007. Prev Vet Med. 2010;93:294\u2013304.","journal-title":"Prev Vet Med"},{"key":"7620_CR8","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1016\/j.prevetmed.2014.10.010","volume":"117","author":"D Tago","year":"2014","unstructured":"Tago D, Hammitt JK, Thomas A, Raboisson D. Cost assessment of the movement restriction policy in France during the 2006 bluetongue virus episode (BTV-8). Prev Vet Med. 2014;117:577\u201389.","journal-title":"Prev Vet Med"},{"key":"7620_CR9","doi-asserted-by":"publisher","first-page":"1552","DOI":"10.3201\/eid3008.231331","volume":"30","author":"M Holwerda","year":"2024","unstructured":"Holwerda M, Santman-Berends IMGA, Harders F, Engelsma M, Vloet RPM, Dijkstra E, et al. Emergence of bluetongue virus serotype 3, the Netherlands, September 2023. Emerg Infect Dis. 2024;30:1552\u201361.","journal-title":"Emerg Infect Dis"},{"key":"7620_CR10","doi-asserted-by":"publisher","first-page":"1438","DOI":"10.3201\/eid3007.240275","volume":"30","author":"A Voigt","year":"2024","unstructured":"Voigt A, Kampen H, Heuser E, Zeiske S, Hoffmann B, H\u00f6per D, et al. Bluetongue virus serotype 3 and Schmallenberg virus in Culicoides biting midges, western Germany, 2023. Emerg Infect Dis. 2024;30:1438\u201341.","journal-title":"Emerg Infect Dis"},{"key":"7620_CR11","doi-asserted-by":"publisher","DOI":"10.1002\/vetr.4910","volume":"196","author":"K Newbrook","year":"2025","unstructured":"Newbrook K, Obishakin E, Jones LA, Waters R, Ashby M, Batten C, et al. Clinical disease in British sheep infected with an emerging strain of bluetongue virus serotype 3. Vet Rec. 2025;196:e4910.","journal-title":"Vet Rec"},{"key":"7620_CR12","doi-asserted-by":"publisher","first-page":"521","DOI":"10.3390\/v17040521","volume":"17","author":"V Van Leeuw","year":"2025","unstructured":"Van Leeuw V, De Leeuw I, Degives N, Depoorter P, Dewulf J, Hanon JB, et al. Impact of BTV-3 circulation in Belgium in 2024 and current knowledge gaps hindering an evidence-based control program. Viruses. 2025;17:521.","journal-title":"Viruses"},{"key":"7620_CR13","unstructured":"DEFRA (Department for Environment, Food and Rural Affairs). Updated Outbreak Assessment #11: Bluetongue virus in Europe: 4 September 2024. London: DEFRA; 2024. https:\/\/assets.publishing.service.gov.uk\/media\/66e151e0865c0eef0bc42dfd\/Updated_Outbreak_Assessment_11_Bluetongue_Virus_in_Europe.pdf. Accessed 31 Mar 2026."},{"key":"7620_CR14","doi-asserted-by":"publisher","first-page":"1845","DOI":"10.3390\/v16121845","volume":"16","author":"SC Barros","year":"2024","unstructured":"Barros SC, Henriques AM, Ramos F, Lu\u00eds T, Fagulha T, Magalh\u00e3es A, et al. Emergence of bluetongue virus serotype 3 in Portugal (2024). Viruses. 2024;16:1845.","journal-title":"Viruses"},{"key":"7620_CR15","unstructured":"DEFRA (Department for Environment, Food and Rural Affairs). Updated Outbreak Assessment #12: Bluetongue virus in Europe: 17 December 2024. London: DEFRA; 2024. https:\/\/assets.publishing.service.gov.uk\/media\/67693d924e2d5e9c0bde9d0f\/BTV_Europe_12.1.pdf. Accessed 31 Mar 2026."},{"key":"7620_CR16","doi-asserted-by":"publisher","first-page":"377","DOI":"10.3390\/pathogens14040377","volume":"14","author":"M Larska","year":"2025","unstructured":"Larska M, Or\u0142owska A, \u0141opuszy\u0144ski W, Skurka \u0141, Nowakowska A, Tr\u0119bas P, et al. First detection of bluetongue virus type 3 in Poland in 2024: a case study in European bison (Bison bonasus). Pathogens. 2025;14:377.","journal-title":"Pathogens"},{"key":"7620_CR17","doi-asserted-by":"publisher","DOI":"10.1016\/j.prevetmed.2024.106289","volume":"231","author":"IMGA Santman-Berends","year":"2024","unstructured":"Santman-Berends IMGA, van den Brink KMJA, Dijkstra E, van Schaik G, Spierenburg MAH, van den Brom R. The impact of the bluetongue serotype 3 outbreak on sheep and goat mortality in the Netherlands in 2023. Prev Vet Med. 2024;231:106289.","journal-title":"Prev Vet Med"},{"key":"7620_CR18","doi-asserted-by":"publisher","first-page":"469","DOI":"10.3201\/eid1803.111905","volume":"18","author":"B Hoffmann","year":"2012","unstructured":"Hoffmann B, Scheuch M, H\u00f6per D, Jungblut R, Holsteg M, Schirrmeier H, et al. Novel orthobunyavirus in cattle, Europe, 2011. Emerg Infect Dis. 2012;18:469\u201372.","journal-title":"Emerg Infect Dis"},{"key":"7620_CR19","doi-asserted-by":"publisher","first-page":"391","DOI":"10.1016\/j.prevetmed.2014.02.012","volume":"116","author":"A Afonso","year":"2014","unstructured":"Afonso A, Abrahantes JC, Conraths F, Veldhuis A, Elbers A, Roberts H, et al. The Schmallenberg virus epidemic in Europe-2011-2013. Prev Vet Med. 2014;116:391\u2013403.","journal-title":"Prev Vet Med"},{"key":"7620_CR20","first-page":"1005","volume":"18","author":"MM Garigliany","year":"2012","unstructured":"Garigliany MM, Hoffmann B, Dive M, Sartelet A, Bayrou C, Cassart D, et al. Schmallenberg virus in calf born at term with porencephaly. Belgium Emerg Infect Dis. 2012;18:1005\u20136.","journal-title":"Belgium Emerg Infect Dis"},{"key":"7620_CR21","first-page":"106","volume":"137","author":"R Van den Brom","year":"2012","unstructured":"Van den Brom R, Luttikholt SJ, Lievaart-Peterson K, Peperkamp NH, Mars MH, van der Poel WH, et al. Epizootic of ovine congenital malformations associated with Schmallenberg virus infection. Tijdschr Diergeneeskd. 2012;137:106\u201311.","journal-title":"Tijdschr Diergeneeskd"},{"key":"7620_CR22","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1017\/S0950268812002245","volume":"141","author":"M Beer","year":"2013","unstructured":"Beer M, Conraths FJ, van der Poel WH. \u201cSchmallenberg virus\u201d \u2013 a novel orthobunyavirus emerging in Europe. Epidemiol Infect. 2013;141:1\u20138.","journal-title":"Epidemiol Infect"},{"key":"7620_CR23","doi-asserted-by":"publisher","first-page":"1063","DOI":"10.3201\/eid2905.221773","volume":"29","author":"A Lorusso","year":"2023","unstructured":"Lorusso A, Cappai S, Loi F, Pinna L, Ruiu A, Puggioni G, et al. Epizootic hemorrhagic disease virus serotype 8, Italy, 2022. Emerg Infect Dis. 2023;29:1063\u20135.","journal-title":"Emerg Infect Dis"},{"key":"7620_CR24","doi-asserted-by":"publisher","DOI":"10.1016\/j.virusres.2024.199489","volume":"350","author":"M Gondard","year":"2024","unstructured":"Gondard M, Postic L, Garin E, Turpaud M, Vorimore F, Ngwa-Mbot D, et al. Exceptional bluetongue virus (BTV) and epizootic hemorrhagic disease virus (EHDV) circulation in France in 2023. Virus Res. 2024;350:199489.","journal-title":"Virus Res"},{"key":"7620_CR25","doi-asserted-by":"publisher","first-page":"1339","DOI":"10.3390\/microorganisms11051339","volume":"11","author":"L Jim\u00e9nez-Cabello","year":"2023","unstructured":"Jim\u00e9nez-Cabello L, Utrilla-Trigo S, Lorenzo G, Ortego J, Calvo-Pinilla E. Epizootic hemorrhagic disease virus: current knowledge and emerging perspectives. Microorganisms. 2023;11:1339.","journal-title":"Microorganisms"},{"key":"7620_CR26","unstructured":"DEFRA (Department for Environment, Food and Rural Affairs). Epizootic haemorrhagic disease in Europe: updated outbreak assessment: 14 August 2024. London: DEFRA; 2024. https:\/\/assets.publishing.service.gov.uk\/media\/66cdc8160671fa213911b341\/14_August_2024_Epizootic_Haemorrhagic_Disease_in_Europe.pdf. Accessed 31 Mar 2026."},{"key":"7620_CR27","doi-asserted-by":"publisher","first-page":"901","DOI":"10.7589\/2016-12-284","volume":"53","author":"MG Ruder","year":"2017","unstructured":"Ruder MG, Johnson D, Ostlund E, Allison AB, Kienzle C, Phillips JE, et al. The first 10 years (2006\u201315) of epizootic hemorrhagic disease virus serotype 6 in the USA. J Wildl Dis. 2017;53:901\u20135.","journal-title":"J Wildl Dis"},{"key":"7620_CR28","doi-asserted-by":"publisher","first-page":"171","DOI":"10.1038\/nrmicro1090","volume":"3","author":"BV Purse","year":"2005","unstructured":"Purse BV, Mellor PS, Rogers DJ, Samuel AR, Mertens PP, Baylis M. Climate change and the recent emergence of bluetongue in Europe. Nat Rev Microbiol. 2005;3:171\u201381.","journal-title":"Nat Rev Microbiol"},{"key":"7620_CR29","doi-asserted-by":"publisher","first-page":"1421","DOI":"10.3201\/eid1409.080417","volume":"14","author":"B Hoffmann","year":"2008","unstructured":"Hoffmann B, Sasserath M, Thalheim S, Bunzenthal C, Strebelow G, Beer M. Bluetongue virus serotype 8 reemergence in Germany, 2007 and 2008. Emerg Infect Dis. 2008;14:1421\u20133.","journal-title":"Emerg Infect Dis"},{"key":"7620_CR30","doi-asserted-by":"publisher","first-page":"1701","DOI":"10.3201\/eid1910.130622","volume":"19","author":"K Wernike","year":"2013","unstructured":"Wernike K, Kohn M, Conraths FJ, Werner D, Kameke D, Hechinger S, et al. Transmission of Schmallenberg virus during winter, Germany. Emerg Infect Dis. 2013;19:1701\u20133.","journal-title":"Germany Emerg Infect Dis"},{"key":"7620_CR31","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1016\/S0378-1135(02)00106-2","volume":"88","author":"KR Bonneau","year":"2002","unstructured":"Bonneau KR, DeMaula CD, Mullens BA, MacLachlan NJ. Duration of viraemia infectious to Culicoides sonorensis in bluetongue virus-infected cattle and sheep. Vet Microbiol. 2002;88:115\u201325.","journal-title":"Vet Microbiol"},{"key":"7620_CR32","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1099\/vir.0.18705-0","volume":"84","author":"H Takamatsu","year":"2003","unstructured":"Takamatsu H, Mellor PS, Mertens PPC, Kirkham PA, Burroughs JN, Parkhouse RME. A possible overwintering mechanism for bluetongue virus in the absence of the insect vector. J Gen Virol. 2003;84:227\u201335.","journal-title":"J Gen Virol"},{"key":"7620_CR33","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.0060210","volume":"6","author":"A Wilson","year":"2008","unstructured":"Wilson A, Darpel K, Mellor PS. Where does bluetongue virus sleep in the winter? PLoS Biol. 2008;6:e210.","journal-title":"PLoS Biol"},{"key":"7620_CR34","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1007\/s00436-009-1411-3","volume":"105","author":"B Vorsprach","year":"2009","unstructured":"Vorsprach B, Meiser CK, Werner D, Balczun C, Schaub GA. Monitoring of Ceratopogonidae in Southwest Germany. Parasitol Res. 2009;105:337\u201344.","journal-title":"Parasitol Res"},{"key":"7620_CR35","doi-asserted-by":"publisher","first-page":"881","DOI":"10.1007\/s00436-016-5361-2","volume":"116","author":"D Kameke","year":"2017","unstructured":"Kameke D, Kampen H, Walther D. Activity of Culicoides spp (Diptera: Ceratopogonidae) inside and outside of livestock stables in late winter and spring. Parasitol Res. 2017;116:881\u20139.","journal-title":"Parasitol Res"},{"key":"7620_CR36","doi-asserted-by":"publisher","first-page":"552","DOI":"10.1111\/mve.12756","volume":"38","author":"S Groschupp","year":"2024","unstructured":"Groschupp S, Kampen H, Werner D. Winter activity of Culicoides (Diptera: Ceratopogonidae) inside and outside stables in Germany. Med Vet Entomol. 2024;38:552\u201365.","journal-title":"Med Vet Entomol"},{"key":"7620_CR37","doi-asserted-by":"publisher","first-page":"246","DOI":"10.1186\/1756-3305-5-246","volume":"5","author":"C Kaufmann","year":"2012","unstructured":"Kaufmann C, Steinmann IC, Hegglin D, Schaffner F, Mathis A. Spatio-temporal occurrence of Culicoides biting midges in the climatic regions of Switzerland, along with large scale species identification by MALDI-TOF mass spectrometry. Parasit Vectors. 2012;5:246.","journal-title":"Parasit Vectors"},{"key":"7620_CR38","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1111\/mve.12260","volume":"32","author":"A Magliano","year":"2018","unstructured":"Magliano A, Scaramozzino P, Ravagnan S, Montarsi F, Rold DA. Indoor and outdoor winter activity of Culicoides biting midges, vectors of bluetongue virus, in Italy. Med Vet Entomol. 2018;32:70\u20137.","journal-title":"Med Vet Entomol"},{"key":"7620_CR39","doi-asserted-by":"publisher","first-page":"356","DOI":"10.1016\/j.vetpar.2008.11.016","volume":"160","author":"Y Braverman","year":"2009","unstructured":"Braverman Y, Mumcuoglu K. Newly emerged nulliparous Culicoides imicola Kieffer (Diptera: Ceratopogonidae) with pigmented abdomen. Vet Parasitol. 2009;160:356\u20138.","journal-title":"Vet Parasitol"},{"key":"7620_CR40","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1111\/j.1440-6055.1969.tb00727.x","volume":"8","author":"AL Dyce","year":"1969","unstructured":"Dyce AL. The recognition of nulliparous and parous Culicoides (Diptera: Ceratopogonidae) without dissection. J Aust Entomol Soc. 1969;8:11\u20135.","journal-title":"J Aust Entomol Soc"},{"key":"7620_CR41","unstructured":"Del\u00e9colle JC. Nouvelle contribution \u00e0 l\u02bc\u00e9tude syst\u00e9matique et iconographique des esp\u00e8ces du genre Culicoides (Dipt\u00e9ra: C\u00e9ratopogonidae) du Nord-Est de la France [Ph.D. thesis]. Strasbourg: Universit\u00e9 Louis Pasteur de Strasbourg; 1985."},{"key":"7620_CR42","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1186\/1756-3305-5-137","volume":"5","author":"B Mathieu","year":"2012","unstructured":"Mathieu B, C\u00eatre-Sossah C, Garros C, Chavernac D, Balenghien T, Carpenter S, et al. Development and validation of IIKC: an interactive identification key for Culicoides (Diptera: Ceratopogonidae) females from the western Palaearctic region. Parasit Vectors. 2012;5:137.","journal-title":"Parasit Vectors"},{"key":"7620_CR43","doi-asserted-by":"publisher","first-page":"1133","DOI":"10.1007\/s00436-011-2357-9","volume":"109","author":"SA Nielsen","year":"2011","unstructured":"Nielsen SA, Kristensen M. Morphological and molecular identification of species of the Obsoletus Group (Diptera: Ceratopogonidae) in Scandinavia. Parasitol Res. 2011;109:1133\u201341.","journal-title":"Parasitol Res"},{"key":"7620_CR44","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1016\/j.vetpar.2009.07.020","volume":"165","author":"N Pag\u00e8s","year":"2009","unstructured":"Pag\u00e8s N, Mu\u00f1oz-Mu\u00f1oz F, Talavera S, Sarto V, Lorca C, N\u00fa\u00f1ez JI. Identification of cryptic species of Culicoides (Diptera: Ceratopogonidae) in the subgenus Culicoides and development of species-specific PCR assays based on barcode regions. Vet Parasitol. 2009;165:298\u2013310.","journal-title":"Vet Parasitol"},{"key":"7620_CR45","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1016\/j.jviromet.2015.02.023","volume":"217","author":"K Wernike","year":"2015","unstructured":"Wernike K, Hoffmann B, Beer M. Simultaneous detection of five notifiable viral diseases of cattle by single-tube multiplex real-time RT-PCR. J Virol Methods. 2015;217:28\u201335.","journal-title":"J Virol Methods"},{"key":"7620_CR46","doi-asserted-by":"publisher","first-page":"236","DOI":"10.1016\/j.vetmic.2012.03.035","volume":"159","author":"S Bilk","year":"2012","unstructured":"Bilk S, Schulze C, Fischer M, Beer M, Hlinak A, Hoffmann B. Organ distribution of Schmallenberg virus RNA in malformed newborns. Vet Microbiol. 2012;159:236\u20138.","journal-title":"Vet Microbiol"},{"key":"7620_CR47","doi-asserted-by":"publisher","first-page":"699","DOI":"10.3390\/d15060699","volume":"15","author":"O D\u00e4hn","year":"2023","unstructured":"D\u00e4hn O, Werner D, Mathieu B, Kampen H. Development of conventional multiplex PCR assays for the identification of 21 West Palaearctic biting midge taxa (Diptera: Ceratopogonidae) belonging to the Culicoides subgenus Culicoides, including recently discovered species and genetic variants. Diversity. 2023;15:699.","journal-title":"Diversity"},{"key":"7620_CR48","doi-asserted-by":"publisher","first-page":"323","DOI":"10.3390\/genes15030323","volume":"15","author":"O D\u00e4hn","year":"2024","unstructured":"D\u00e4hn O, Werner D, Mathieu B, Kampen H. Large-scale cytochrome c oxidase subunit I gene data analysis for the development of a multiplex polymerase chain reaction test capable of identifying biting midge vector species and haplotypes (Diptera: Ceratopogonidae) of the Culicoides subgenus Avaritia Fox, 1955. Genes. 2024;15:323.","journal-title":"Genes"},{"key":"7620_CR49","doi-asserted-by":"publisher","first-page":"381","DOI":"10.1007\/s00436-009-1417-x","volume":"105","author":"PH Clausen","year":"2009","unstructured":"Clausen PH, Stephan A, Bartsch S, Jandowsky A, Hoffmann-K\u00f6hler P, Schein E, et al. Seasonal dynamics of biting midges (Diptera: Ceratopogonidae, Culicoides spp.) on dairy farms of central Germany during the 2007\/2008 epidemic of bluetongue. Parasitol Res. 2009;105:381\u20136.","journal-title":"Parasitol Res"},{"key":"7620_CR50","doi-asserted-by":"publisher","first-page":"1481","DOI":"10.3201\/eid1509.090562","volume":"15","author":"B Hoffmann","year":"2009","unstructured":"Hoffmann B, Bauer B, Bauer C, B\u00e4tza HJ, Beer M, Clausen PH, et al. Monitoring of putative vectors of bluetongue virus serotype 8, Germany. Emerg Infect Dis. 2009;15:1481\u20134.","journal-title":"Germany Emerg Infect Dis"},{"key":"7620_CR51","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1007\/s00436-009-1413-1","volume":"105","author":"H Mehlhorn","year":"2009","unstructured":"Mehlhorn H, Walldorf V, Klimpel S, Schmahl G, Al-Quraishy S, Walldorf U, et al. Entomological survey on vectors of bluetongue virus in North Rhine-Westphalia (Germany) during 2007 and 2008. Parasitol Res. 2009;105:321\u20139.","journal-title":"Parasitol Res"},{"key":"7620_CR52","doi-asserted-by":"publisher","first-page":"219","DOI":"10.1007\/s00436-007-0519-6","volume":"101","author":"H Mehlhorn","year":"2007","unstructured":"Mehlhorn H, Walldorf V, Klimpel S, Jahn B, Jaeger F, Eschweiler J, et al. First occurrence of Culicoides obsoletus-transmitted bluetongue virus epidemic in central Europe. Parasitol Res. 2007;101:219\u201328.","journal-title":"Parasitol Res"},{"key":"7620_CR53","doi-asserted-by":"publisher","first-page":"98","DOI":"10.1016\/j.prevetmed.2008.06.013","volume":"87","author":"R Meiswinkel","year":"2008","unstructured":"Meiswinkel R, Goffredo M, Leijs P, Conte A. The Culicoides \u201csnapshot\u201d: a novel approach used to assess vector densities widely and rapidly during the 2006 outbreak of bluetongue (BT) in The Netherlands. Prev Vet Med. 2008;87:98\u2013118.","journal-title":"Prev Vet Med"},{"key":"7620_CR54","doi-asserted-by":"publisher","first-page":"512","DOI":"10.1186\/s13071-018-3082-3","volume":"11","author":"C Sohier","year":"2018","unstructured":"Sohier C, Deblauwe I, De Deken R, Madder M, Fassotte C, Losson B, et al. Longitudinal monitoring of Culicoides in Belgium between 2007 and 2011: local variation in population dynamics parameters warrant cautious use of monitoring data. Parasit Vectors. 2018;11:512.","journal-title":"Parasit Vectors"},{"key":"7620_CR55","doi-asserted-by":"publisher","first-page":"112","DOI":"10.1186\/s13071-018-2706-y","volume":"11","author":"AC Cu\u00e9llar","year":"2018","unstructured":"Cu\u00e9llar AC, Kj\u00e6r LJ, Kirkeby C, Skovgard H, Nielsen SA, Stockmarr A, et al. Spatial and temporal variation in the abundance of Culicoides biting midges (Diptera: Ceratopogonidae) in nine European countries. Parasit Vectors. 2018;11:112.","journal-title":"Parasit Vectors"},{"key":"7620_CR56","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1016\/j.vetpar.2014.08.029","volume":"205","author":"JY Zimmer","year":"2014","unstructured":"Zimmer JY, Brostaux Y, Haubruge E, Francis F. Larval development sites of the main Culicoides species (Diptera: Ceratopogonidae) in northern Europe and distribution of coprophilic species larvae in Belgian pastures. Vet Parasitol. 2014;205:676\u201386.","journal-title":"Vet Parasitol"},{"key":"7620_CR57","doi-asserted-by":"publisher","DOI":"10.1016\/j.vetpar.2025.110423","volume":"335","author":"S Groschupp","year":"2025","unstructured":"Groschupp S, Kampen H, Werner D. Deep litter as a production substrate of Culicoides biting midges (Diptera: Ceratopogonidae). Vet Parasitol. 2025;335:110423.","journal-title":"Vet Parasitol"},{"key":"7620_CR58","doi-asserted-by":"publisher","first-page":"5000","DOI":"10.3390\/ijerph17145000","volume":"17","author":"D Werner","year":"2020","unstructured":"Werner D, Groschupp S, Bauer C, Kampen H. Breeding habitat preferences of major Culicoides species (Diptera: Ceratopogonidae) in Germany. Int J Environ Res Public Health. 2020;17:5000.","journal-title":"Int J Environ Res Public Health"},{"key":"7620_CR59","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1016\/j.prevetmed.2008.06.008","volume":"87","author":"R Meiswinkel","year":"2008","unstructured":"Meiswinkel R, Goffredo M, Dijkstra EG, van der Ven IJ, Baldet T, Elbers A. Endophily in Culicoides associated with BTV-infected cattle in the province of Limburg, south-eastern Netherlands, 2006. Prev Vet Med. 2008;87:182\u201395.","journal-title":"Prev Vet Med"},{"key":"7620_CR60","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1111\/j.1365-2915.2009.00842.x","volume":"24","author":"M Baylis","year":"2010","unstructured":"Baylis M, Parkin H, Kreppel K, Carpenter S, Mellor PS, McIntyre KM. Evaluation of housing as a means to protect cattle from Culicoides biting midges, the vectors of bluetongue virus. Med Vet Entomol. 2010;24:38\u201345.","journal-title":"Med Vet Entomol"},{"key":"7620_CR61","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1186\/1756-3305-6-43","volume":"6","author":"C Kirkeby","year":"2013","unstructured":"Kirkeby C, B\u00f8dker R, Stockmarr A, Lind P. Spatial abundance and clustering of Culicoides (Diptera: Ceratopogonidae) on a local scale. Parasit Vectors. 2013;6:43.","journal-title":"Parasit Vectors"},{"key":"7620_CR62","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0027987","volume":"6","author":"S Carpenter","year":"2011","unstructured":"Carpenter S, Wilson A, Barber J, Veronesi E, Mellor P, Venter G, et al. Temperature dependence of the extrinsic incubation period of orbiviruses in Culicoides biting midges. PLoS ONE. 2011;6:e27987.","journal-title":"PLoS ONE"},{"key":"7620_CR63","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1146\/annurev-ento-011613-162023","volume":"59","author":"DL Denlinger","year":"2014","unstructured":"Denlinger DL, Armbruster PA. Mosquito diapause. Annu Rev Entomol. 2014;59:73\u201393.","journal-title":"Annu Rev Entomol"},{"key":"7620_CR64","first-page":"243","volume-title":"Aquatic Insects of North Europe \u2013 A Taxonomic Handbook","author":"R Szadziewski","year":"2004","unstructured":"Szadziewski R, Krzywi\u0144ski J, Gi\u0142ka W. Diptera Ceratopogonidae, biting midges. In: Nilsson AN, editor. Aquatic Insects of North Europe \u2013 A Taxonomic Handbook, vol. 2. Stenstrup (Denmark): Apollo Books; 2004. p. 243\u201363."},{"key":"7620_CR65","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1186\/s13071-022-05370-z","volume":"15","author":"JE Stokes","year":"2022","unstructured":"Stokes JE, Carpenter S, Sanders C, Gubbins S. Emergence dynamics of adult Culicoides biting midges at two farms in South-East England. Parasit Vectors. 2022;15:251.","journal-title":"Parasit Vectors"},{"key":"7620_CR66","doi-asserted-by":"publisher","DOI":"10.1016\/j.jtherbio.2024.103783","volume":"119","author":"AL Hochstrasser","year":"2024","unstructured":"Hochstrasser AL, Mathis A, Verhulst NO. Thermal preference of Culicoides biting midges in laboratory and semi-field settings. J Therm Biol. 2024;119:103783.","journal-title":"J Therm Biol"},{"key":"7620_CR67","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1016\/j.vetmic.2013.07.015","volume":"166","author":"M Larska","year":"2013","unstructured":"Larska M, Lechowski L, Grochowska M, \u017bmudzi\u0144ski JF. Detection of the Schmallenberg virus in nulliparous Culicoides obsoletus\/scoticus complex and C. punctatus \u2013 the possibility of transovarial virus transmission in the midge population and of a new vector. Vet Microbiol. 2013;166:467\u201373.","journal-title":"Vet Microbiol"},{"key":"7620_CR68","doi-asserted-by":"publisher","first-page":"16078","DOI":"10.1038\/s41598-025-00849-y","volume":"15","author":"D Erram","year":"2025","unstructured":"Erram D, McGregor B, Acevedo C, Alto BW, Burkett-Cadena N. Epizootic hemorrhagic disease virus oral infection affects midge reproduction and is vertically transmitted to offspring in Culicoides sonorensis. Sci Rep. 2025;15:16078.","journal-title":"Sci Rep"},{"key":"7620_CR69","doi-asserted-by":"publisher","first-page":"1571","DOI":"10.3390\/pathogens10121571","volume":"10","author":"M Goffredo","year":"2021","unstructured":"Goffredo M, Quaglia M, De Ascentis M, d\u2019Alessio SG, Federici V, Conte A, et al. The absence of abdominal pigmentation in livestock associated Culicoides following artificial blood feeding and the epidemiological implication for arbovirus surveillance. Pathogens. 2021;10:1571.","journal-title":"Pathogens"},{"key":"7620_CR70","doi-asserted-by":"publisher","first-page":"228","DOI":"10.1111\/j.1365-2915.2008.00732.x","volume":"22","author":"M Baylis","year":"2008","unstructured":"Baylis M, O\u2019Connell L, Mellor PS. Rates of bluetongue virus transmission between Culicoides sonorensis and sheep. Med Vet Entomol. 2008;22:228\u201337.","journal-title":"Med Vet Entomol"},{"key":"7620_CR71","doi-asserted-by":"publisher","first-page":"851","DOI":"10.3390\/microorganisms8060851","volume":"8","author":"E Veronesi","year":"2020","unstructured":"Veronesi E, Darpel K, Gubbins S, Batten C, Nomikou K, Mertens P, et al. Diversity of transmission outcomes following co-infection of sheep with strains of bluetongue virus serotype 1 and 8. Microorganisms. 2020;8:851.","journal-title":"Microorganisms"},{"key":"7620_CR72","doi-asserted-by":"publisher","first-page":"262","DOI":"10.1186\/s13071-025-06913-w","volume":"18","author":"S Zeiske","year":"2025","unstructured":"Zeiske S, Kampen H, Sick F, D\u00e4hn O, Voigt A, Heuser A, et al. Monitoring of Schmallenberg virus, bluetongue virus and epizootic haemorrhagic disease virus in biting midges in Germany 2019\u20132023. Parasit Vectors. 2025;18:262.","journal-title":"Parasit Vectors"},{"key":"7620_CR73","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1111\/mve.70007","volume":"40","author":"M Uiterwijk","year":"2026","unstructured":"Uiterwijk M, Jacobs F, de Boer K, Feddema A-J, Vloet RPM, Dik M, et al. Culicoides species involved in the BTV-3 epidemic, the Netherlands, 2023\u20132024. Med Vet Entomol. 2026;40:111\u201322.","journal-title":"Med Vet Entomol"},{"key":"7620_CR74","first-page":"160","volume":"40","author":"BA Mullens","year":"2004","unstructured":"Mullens BA, Gerry AC, Lysyk TJ, Schmidtmann ET. Environmental effects on vector competence and virogenesis of bluetongue virus in Culicoides: interpreting laboratory data in a field context. Vet Ital. 2004;40:160\u20136.","journal-title":"Vet Ital"},{"key":"7620_CR75","doi-asserted-by":"publisher","first-page":"230","DOI":"10.1016\/j.prevetmed.2013.05.014","volume":"111","author":"M Goffredo","year":"2013","unstructured":"Goffredo M, Monaco F, Capelli G, Quaglia M, Federici V, Catalani M, et al. Schmallenberg virus in Italy: a retrospective survey in Culicoides stored during the bluetongue Italian surveillance program. Prev Vet Med. 2013;111:230\u20136.","journal-title":"Prev Vet Med"},{"key":"7620_CR76","doi-asserted-by":"publisher","first-page":"489","DOI":"10.2478\/jvetres-2025-0066","volume":"69","author":"M Kwa\u015bnik","year":"2025","unstructured":"Kwa\u015bnik M, Or\u0142owska A, Ro\u017cek W, Larska M, Rola J. Entomological surveillance for bluetongue virus in Poland: late-season activity and abundance of Culicoides vectors in 2024. J Vet Res. 2025;69:489\u2013503.","journal-title":"J Vet Res"},{"key":"7620_CR77","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1111\/mve.12109","volume":"29","author":"N De Regge","year":"2015","unstructured":"De Regge N, De Deken R, Fassotte C, Losson B, Deblauwe I, Madder M, et al. Culicoides monitoring in Belgium in 2011: analysis of spatiotemporal abundance, species diversity and Schmallenberg virus detection. Med Vet Entomol. 2015;29:263\u201375.","journal-title":"Med Vet Entomol"},{"key":"7620_CR78","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1016\/0147-9571(94)90043-4","volume":"17","author":"NJ MacLachlan","year":"1994","unstructured":"MacLachlan NJ. The pathogenesis and immunology of bluetongue virus infection of ruminants. Comp Immunol Microbiol Infect Dis. 1994;17:197\u2013206.","journal-title":"Comp Immunol Microbiol Infect Dis"},{"key":"7620_CR79","unstructured":"WOAH (World Organisation of Animal Health). Terrestrial Animal Health Code: Infection with bluetongue virus. Chapter 8.3. Paris: WOAH; 2014. Available from: https:\/\/www.woah.org\/fileadmin\/Home\/eng\/Health_standards\/tahc\/2016\/en_chapitre_bluetongue.htm. Accessed 31 Mar 2026."},{"key":"7620_CR80","first-page":"20241105","volume":"291","author":"GA Tung","year":"2024","unstructured":"Tung GA, Fonseca DM. Internal and external drivers interact to create highly dynamic mosquito blood-feeding behaviour. Proc Biol Sci. 2024;291:20241105.","journal-title":"Proc Biol Sci"},{"key":"7620_CR81","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1186\/s13071-020-04552-x","volume":"14","author":"LA Tugwell","year":"2021","unstructured":"Tugwell LA, England ME, Gubbins S, Sanders CJ, Stokes JE, Stoner J, et al. Thermal limits for flight activity of field-collected Culicoides in the United Kingdom defined under laboratory conditions. Parasit Vectors. 2021;14:55.","journal-title":"Parasit Vectors"},{"key":"7620_CR82","doi-asserted-by":"crossref","unstructured":"Brugger K, K\u00f6fer J, Rubel F. Outdoor and indoor monitoring of livestock-associated Culicoides spp. to assess vector-free periods and disease risks. BMC Vet Res. 2016;12:88.","DOI":"10.1186\/s12917-016-0710-z"},{"key":"7620_CR83","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1016\/j.prevetmed.2008.06.005","volume":"87","author":"R Meiswinkel","year":"2008","unstructured":"Meiswinkel R, Baldet T, de Deken R, Takken W, Del\u00e9colle JC, Mellor PS. The 2006 outbreak of bluetongue in northern Europe \u2013 the entomological perspective. Prev Vet Med. 2008;87:55\u201363.","journal-title":"Prev Vet Med"},{"key":"7620_CR84","doi-asserted-by":"publisher","first-page":"451","DOI":"10.1136\/vr.160.13.451-b","volume":"160","author":"B Losson","year":"2007","unstructured":"Losson B, Mignon B, Paternostre J, Madder M, De Deken R, De Deken G, et al. Biting midges overwintering in Belgium. Vet Rec. 2007;160:451\u20132.","journal-title":"Vet Rec"},{"key":"7620_CR85","unstructured":"ECDC (European Centre for Disease Prevention and Control). Culicoides imicola \u2013 current known distribution: October 2023.Stockholm: ECDC; 2023. https:\/\/www.ecdc.europa.eu\/en\/publications-data\/culicoides-imicola-current-known-distribution-october-2023. Accessed 31 Mar 2026."},{"key":"7620_CR86","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1186\/s13071-016-1426-4","volume":"9","author":"S Jacquet","year":"2016","unstructured":"Jacquet S, Huber K, Guis H, Setier-Rio ML, Goffredo M, All\u00e8ne X, et al. Spatio-temporal genetic variation of the biting midge vector species Culicoides imicola (Ceratopogonidae) Kieffer in France. Parasit Vectors. 2016;9:141.","journal-title":"Parasit Vectors"},{"key":"7620_CR87","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1111\/j.1365-2915.2006.00621.x","volume":"20","author":"A Cagienard","year":"2006","unstructured":"Cagienard A, Griot C, Mellor PS, Denison E, St\u00e4rk KD. Bluetongue vector species of Culicoides in Switzerland. Med Vet Entomol. 2006;20:239\u201347.","journal-title":"Med Vet Entomol"},{"key":"7620_CR88","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1186\/1297-9716-42-4","volume":"42","author":"S Napp","year":"2011","unstructured":"Napp S, Gubbins S, Calistri P, Allepuz A, Alba A, Garc\u00eda-Bocanegra I, et al. Quantitative assessment of the probability of bluetongue virus overwintering by horizontal transmission: application to Germany. Vet Res. 2011;42:4.","journal-title":"Vet Res"},{"key":"7620_CR89","unstructured":"IPCC (Intergovernmental Panel on Climate Change) AR6 Climate change 2021: Regional fact sheet \u2013 Europe. Climate change 2021: The physical science basis. Working group 1 contribution to the sixth assessment report of the Intergovernmental Panel on Climate Change. Geneva: IPCC; 2021. https:\/\/www.ipcc.ch\/report\/ar6\/wg1\/downloads\/factsheets\/IPCC_AR6_WGI_Regional_Fact_Sheet_Europe.pdf. Accessed 31 Mar 2026."}],"container-title":["Parasites &amp; Vectors"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s13071-026-07620-w","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13071-026-07620-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13071-026-07620-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T04:23:26Z","timestamp":1788495806000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s13071-026-07620-w"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,4]]},"references-count":89,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["7620"],"URL":"https:\/\/doi.org\/10.1186\/s13071-026-07620-w","relation":{},"ISSN":["1756-3305"],"issn-type":[{"value":"1756-3305","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,4]]},"assertion":[{"value":"6 April 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 August 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"4 September 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Not applicable.","order":1,"name":"Ethics","label":"Ethics approval and consent to participate","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","label":"Consent for publication","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors state that there are no conflicts of interest.","order":3,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"382"}}