{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T21:31:46Z","timestamp":1775079106251,"version":"3.50.1"},"reference-count":96,"publisher":"American Society of Tropical Medicine and Hygiene","issue":"4","license":[{"start":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T00:00:00Z","timestamp":1775001600000},"content-version":"unspecified","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,4,1]]},"abstract":"<jats:title>ABSTRACT.<\/jats:title>\n                  <jats:p>\n                    West Nile virus (WNV) is an endemic arboviral infection in the United States that has undergone phylogenetic evolution since its introduction 25 years ago. An integrated vector\u2013human\u2013pathogen study was conducted in the summer of 2023 to unearth contemporary\n                    <jats:italic>Culex quinquefasciatus<\/jats:italic>\n                    habitat patterns and human transmission spillover foci in South Carolina, a state with historically little WNV data. A serosurvey revealed WNV seroprevalence 10 times the national average (22% versus 2%, respectively), with unusual epidemiologic risk factors. Female\n                    <jats:italic>Culex quinquefasciatus<\/jats:italic>\n                    WNV positivity was low (2.7%), with viral phylogenetics 100% homologous to the WN02 clade. Mosquito vectors clustered in affluent urban neighborhoods with greater tree canopy cover and abundant waterbodies.\n                    <jats:italic>Culex quinquefasciatus<\/jats:italic>\n                    abundance was greatest when climate variance was nominal in the 72 hours preceding collection. An unusual bimodal mosquito temporal pattern was observed, reflecting changing climate patterns. The present comprehensive WNV study reveals emerging transmission factors as WNV continues to evolve and persist in the southeastern United States.\n                  <\/jats:p>","DOI":"10.4269\/ajtmh.25-0305","type":"journal-article","created":{"date-parts":[[2026,2,5]],"date-time":"2026-02-05T15:17:56Z","timestamp":1770304676000},"page":"742-750","source":"Crossref","is-referenced-by-count":0,"title":["Ecologic Factors Contributing to West Nile Virus Hyperendemicity in Central South Carolina: An Integrated Vector\u2013Human\u2013Environmental Study"],"prefix":"10.4269","volume":"114","author":[{"given":"Elba S.","family":"Fridriksson","sequence":"first","affiliation":[{"name":"Arnold School of Public Health, University of South Carolina, Columbia, South Carolina;"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ahalya","family":"Muraleedharan","sequence":"additional","affiliation":[{"name":"Arnold School of Public Health, University of South Carolina, Columbia, South Carolina;"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kyndall C.","family":"Dye-Braumuller","sequence":"additional","affiliation":[{"name":"Arnold School of Public Health, University of South Carolina, Columbia, South Carolina;"},{"name":"Institute for Infectious Disease Translational Research, University of South Carolina, Columbia, South Carolina"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Madeleine M.","family":"Meyer","sequence":"additional","affiliation":[{"name":"Arnold School of Public Health, University of South Carolina, Columbia, South Carolina;"},{"name":"Institute for Infectious Disease Translational Research, University of South Carolina, Columbia, South Carolina"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kia","family":"Zellars","sequence":"additional","affiliation":[{"name":"Arnold School of Public Health, University of South Carolina, Columbia, South Carolina;"},{"name":"Institute for Infectious Disease Translational Research, University of South Carolina, Columbia, South Carolina"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Huixuan","family":"Li","sequence":"additional","affiliation":[{"name":"Arnold School of Public Health, University of South Carolina, Columbia, South Carolina;"},{"name":"Institute for Infectious Disease Translational Research, University of South Carolina, Columbia, South Carolina"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Melissa S.","family":"Nolan","sequence":"additional","affiliation":[{"name":"Arnold School of Public Health, University of South Carolina, Columbia, South Carolina;"},{"name":"Institute for Infectious Disease Translational Research, University of South Carolina, Columbia, South Carolina"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"2795","reference":[{"key":"b1","series-title":"Am J Trop Med Hyg","first-page":"471","article-title":"A neurotropic virus isolated from the blood of a native of Uganda","volume":"s1-20","author":"Smithburn","year":"1940"},{"key":"b2","series-title":"J Virol","first-page":"2964","article-title":"Phylogeography of West Nile virus: From the cradle of evolution in Africa to Eurasia, Australia, and the Americas","volume":"85","author":"May","year":"2011"},{"key":"b3","series-title":"Lancet","first-page":"261","article-title":"Epidemic West Nile encephalitis, New York, 1999: Results of a household-based seroepidemiological survey","volume":"358","author":"Mostashari","year":"2001"},{"key":"b4","series-title":"Emerg Infect Dis","first-page":"684","article-title":"Neuroinvasive disease and West Nile virus infection, North Dakota, USA, 1999-2008","volume":"18","author":"Carson","year":"2012"},{"key":"b5","series-title":"PLoS One","first-page":"e102953","article-title":"Survival analysis, long-term outcomes, and percentage of recovery up to 8 years post-infection among the Houston West Nile virus cohort","volume":"9","author":"Murray","year":"2014"},{"key":"b6","series-title":"Am J Trop Med Hyg","first-page":"1006","article-title":"Long-term neurological outcomes in West Nile virus-infected patients: An observational study","volume":"92","author":"Weatherhead","year":"2015"},{"key":"b7","series-title":"PLoS One","first-page":"e0148898","article-title":"West Nile virus retinopathy and associations with long term neurological and neurocognitive sequelae","volume":"11","author":"Hasbun","year":"2016"},{"key":"b8","series-title":"PLoS One","first-page":"e40374","article-title":"Prevalence of chronic kidney disease and progression of disease over time among patients enrolled in the Houston West Nile virus cohort","volume":"7","author":"Nolan","year":"2012"},{"key":"b9","series-title":"Viral Immunol","first-page":"327","article-title":"Evaluation of prolonged fatigue post-West Nile virus infection and association of fatigue with elevated antiviral and proinflammatory cytokines","volume":"27","author":"Garcia","year":"2014"},{"key":"b10","series-title":"PLoS Negl Trop Dis","first-page":"e0009190","article-title":"A 20-year historical review of West Nile virus since its initial emergence in North America: Has West Nile virus become a neglected tropical disease?","volume":"15","author":"Ronca","year":"2021"},{"key":"b11","series-title":"Emerg Infect Dis","first-page":"325","article-title":"Cumulative incidence of West Nile virus infection, continental United States, 1999\u20132016","volume":"25","author":"Ronca","year":"2019"},{"key":"b12","article-title":"Data and Maps for West Nile.","year":"2024"},{"key":"b13","series-title":"Ann Neurol","first-page":"93","article-title":"Clinical, prognostic, and longitudinal functional and neuropsychological features of West Nile virus neuroinvasive disease in the United States: A systematic review and meta-analysis","volume":"98","author":"Roberts","year":"2025"},{"key":"b14","series-title":"J Med Entomol","first-page":"401","article-title":"Riding the wave: Reactive vector-borne disease policy renders the United States vulnerable to outbreaks and insecticide resistance","volume":"59","author":"Dye-Braumuller","year":"2022"},{"key":"b15","series-title":"Trop Med Infect Dis","first-page":"73","article-title":"Needs assessment of southeastern United States vector control agencies: Capacity improvement is greatly needed to prevent the next vector-borne disease outbreak","volume":"7","author":"Dye-Braumuller","year":"2022"},{"key":"b16","series-title":"Emerg Infect Dis J","first-page":"1587","article-title":"Use of testing for West Nile virus and other arboviruses","volume":"22","author":"Vanichanan","year":"2016"},{"key":"b17","series-title":"MMWR Morb Mortal Wkly Rep","first-page":"646","article-title":"West Nile virus activity\u2013United States, July 3-9, 2003","volume":"52","year":"2003"},{"key":"b18","series-title":"Am J Forensic Med Pathol","first-page":"192","article-title":"West Nile virus encephalitis","volume":"26","author":"Batalis","year":"2005"},{"key":"b19","series-title":"Vector Borne Zoonotic Dis","first-page":"1","article-title":"Prediction of equine risk of West Nile virus infection based on dead bird surveillance","volume":"6","author":"Roberts","year":"2006"},{"key":"b20","series-title":"J Wildl Dis","first-page":"863","article-title":"Hematology, parasitology, and serology of free-ranging coyotes (Canis latrans) from South Carolina","volume":"45","author":"Miller","year":"2009"},{"key":"b21","series-title":"Geospat Health","first-page":"15","article-title":"Application of a degree-day model of West Nile virus transmission risk to the East Coast of the United States of America","volume":"7","author":"Konrad","year":"2012"},{"key":"b22","series-title":"J Wildl Dis","first-page":"354","article-title":"Survey for West Nile virus antibodies in wild ducks, 2004-06, USA","volume":"52","author":"Hofmeister","year":"2016"},{"key":"b23","article-title":"County steps up efforts to prevent West Nile virus spread","year":"2022"},{"key":"b24","series-title":"Identification and Geographical Distribution of the Mosquitoes of North America, North of Mexico","author":"Darsie","year":"2005"},{"key":"b25","series-title":"Acta Trop","first-page":"106972","article-title":"Employing oxford nanopore technologies (ONT) for understanding the ecology and transmission dynamics of flaviviruses in mosquitoes (Diptera: Culicidae) from Eastern Colombia","volume":"245","author":"Mart\u00ednez","year":"2023"},{"key":"b26","series-title":"Mol Biol Evol","first-page":"3022","article-title":"MEGA11: Molecular evolutionary genetics analysis version 11","volume":"38","author":"Tamura","year":"2021"},{"key":"b27","series-title":"Nucleic Acids Res","first-page":"1792","article-title":"MUSCLE: Multiple sequence alignment with high accuracy and high 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2005\u20132007","volume":"365","author":"Amore","year":"2010"},{"key":"b32","series-title":"Virology","first-page":"381","article-title":"Fine-scale genetic variation and evolution of West Nile Virus in a transmission \u201chot spot\u201d in suburban Chicago, USA","volume":"374","author":"Bertolotti","year":"2008"},{"key":"b33","series-title":"PLoS One","first-page":"e0227160","article-title":"The drivers of West Nile virus human illness in the Chicago, Illinois, USA area: Fine scale dynamic effects of weather, mosquito infection, social, and biological conditions","volume":"15","author":"Karki","year":"2020"},{"key":"b34","series-title":"Vector Borne Zoonotic Dis","first-page":"73","article-title":"Factors affecting the geographic distribution of West Nile virus in Georgia, USA: 2002\u20132004","volume":"6","author":"Gibbs","year":"2006"},{"key":"b35","series-title":"PLoS One","first-page":"e15437","article-title":"Economic conditions predict prevalence of West Nile virus","volume":"5","author":"Harrigan","year":"2010"},{"key":"b36","series-title":"Clin Infect Dis","first-page":"1565","article-title":"Beyond viruses: Clinical profiles and etiologies associated with encephalitis","volume":"43","author":"Glaser","year":"2006"},{"key":"b37","series-title":"Parasit Vectors","first-page":"547","article-title":"Updated distribution maps of predominant Culex mosquitoes across the Americas","volume":"14","author":"Gorris","year":"2021"},{"key":"b38"},{"key":"b39","series-title":"Parasit Vectors","first-page":"116","article-title":"Geographic variation in the response of Culex pipiens life history traits to temperature","volume":"9","author":"Ruybal","year":"2016"},{"key":"b40","series-title":"Glob Chang Biol","first-page":"2373","article-title":"Temperature variation makes ectotherms more sensitive to climate change","volume":"19","author":"Paaijmans","year":"2013"},{"key":"b41","series-title":"Proc Biol Sci","first-page":"20201065","article-title":"Thermal thresholds heighten sensitivity of West Nile virus transmission to changing temperatures in coastal California","volume":"287","author":"Skaff","year":"2020"},{"key":"b42","series-title":"Acta Trop","first-page":"1","article-title":"Effect of temperature on life history traits during immature development of Aedes aegypti and Culex quinquefasciatus (Diptera: Culicidae) from C\u00f3rdoba city, Argentina","volume":"146","author":"Grech","year":"2015"},{"key":"b43"},{"key":"b44","series-title":"Integr Comp Biol","first-page":"45","article-title":"Fine-scale microclimatic variation can shape the responses of organisms to global change in both natural and urban environments","volume":"56","author":"Pincebourde","year":"2016"},{"key":"b45","series-title":"PLoS Negl Trop Dis","first-page":"e0005640","article-title":"Fine-scale variation in microclimate across an urban landscape shapes variation in mosquito population dynamics and the potential of Aedes albopictus to transmit arboviral disease","volume":"11","author":"Murdock","year":"2017"},{"key":"b46","series-title":"Int J Health Geogr","first-page":"8","article-title":"Environmental and social determinants of human risk during a West Nile virus outbreak in the greater Chicago area, 2002","volume":"3","author":"Ruiz","year":"2004"},{"key":"b47","series-title":"Int J Health Geogr","first-page":"12","article-title":"Locating suitable habitats for West Nile Virus-infected mosquitoes through association of environmental characteristics with infected mosquito locations: A case study in Shelby County, Tennessee","volume":"7","author":"Ozdenerol","year":"2008"},{"key":"b48","series-title":"Int J Rad Oncol Biol Phys","first-page":"S71","article-title":"Centering Black Voices: Factors Influencing a Cancer Patients Decision to Join a Clinical Trial","volume":"120","author":"Gomez","year":"2024"},{"key":"b1","series-title":"Am J Trop Med Hyg","first-page":"471","article-title":"A neurotropic virus isolated from the blood of a native of Uganda","volume":"s1-20","author":"Smithburn","year":"1940"},{"key":"b2","series-title":"J Virol","first-page":"2964","article-title":"Phylogeography of West Nile virus: From the cradle of evolution in Africa to Eurasia, Australia, and the Americas","volume":"85","author":"May","year":"2011"},{"key":"b3","series-title":"Lancet","first-page":"261","article-title":"Epidemic West Nile encephalitis, New York, 1999: Results of a household-based seroepidemiological survey","volume":"358","author":"Mostashari","year":"2001"},{"key":"b4","series-title":"Emerg Infect Dis","first-page":"684","article-title":"Neuroinvasive disease and West Nile virus infection, North Dakota, USA, 1999-2008","volume":"18","author":"Carson","year":"2012"},{"key":"b5","series-title":"PLoS One","first-page":"e102953","article-title":"Survival analysis, long-term outcomes, and percentage of recovery up to 8 years post-infection among the Houston West Nile virus cohort","volume":"9","author":"Murray","year":"2014"},{"key":"b6","series-title":"Am J Trop Med Hyg","first-page":"1006","article-title":"Long-term neurological outcomes in West Nile virus-infected patients: An observational study","volume":"92","author":"Weatherhead","year":"2015"},{"key":"b7","series-title":"PLoS One","first-page":"e0148898","article-title":"West Nile virus retinopathy and associations with long term neurological and neurocognitive sequelae","volume":"11","author":"Hasbun","year":"2016"},{"key":"b8","series-title":"PLoS One","first-page":"e40374","article-title":"Prevalence of chronic kidney disease and progression of disease over time among patients enrolled in the Houston West Nile virus cohort","volume":"7","author":"Nolan","year":"2012"},{"key":"b9","series-title":"Viral Immunol","first-page":"327","article-title":"Evaluation of prolonged fatigue post-West Nile virus infection and association of fatigue with elevated antiviral and proinflammatory cytokines","volume":"27","author":"Garcia","year":"2014"},{"key":"b10","series-title":"PLoS Negl Trop Dis","first-page":"e0009190","article-title":"A 20-year historical review of West Nile virus since its initial emergence in North America: Has West Nile virus become a neglected tropical disease?","volume":"15","author":"Ronca","year":"2021"},{"key":"b11","series-title":"Emerg Infect Dis","first-page":"325","article-title":"Cumulative incidence of West Nile virus infection, continental United States, 1999\u20132016","volume":"25","author":"Ronca","year":"2019"},{"key":"b12","article-title":"Data and Maps for West Nile.","year":"2024"},{"key":"b13","series-title":"Ann Neurol","first-page":"93","article-title":"Clinical, prognostic, and longitudinal functional and neuropsychological features of West Nile virus neuroinvasive disease in the United States: A systematic review and meta-analysis","volume":"98","author":"Roberts","year":"2025"},{"key":"b14","series-title":"J Med Entomol","first-page":"401","article-title":"Riding the wave: Reactive vector-borne disease policy renders the United States vulnerable to outbreaks and insecticide resistance","volume":"59","author":"Dye-Braumuller","year":"2022"},{"key":"b15","series-title":"Trop Med Infect Dis","first-page":"73","article-title":"Needs assessment of southeastern United States vector control agencies: Capacity improvement is greatly needed to prevent the next vector-borne disease outbreak","volume":"7","author":"Dye-Braumuller","year":"2022"},{"key":"b16","series-title":"Emerg Infect Dis J","first-page":"1587","article-title":"Use of testing for West Nile virus and other arboviruses","volume":"22","author":"Vanichanan","year":"2016"},{"key":"b17","series-title":"MMWR Morb Mortal Wkly Rep","first-page":"646","article-title":"West Nile virus activity\u2013United States, July 3-9, 2003","volume":"52","year":"2003"},{"key":"b18","series-title":"Am J Forensic Med Pathol","first-page":"192","article-title":"West Nile virus encephalitis","volume":"26","author":"Batalis","year":"2005"},{"key":"b19","series-title":"Vector Borne Zoonotic Dis","first-page":"1","article-title":"Prediction of equine risk of West Nile virus infection based on dead bird surveillance","volume":"6","author":"Roberts","year":"2006"},{"key":"b20","series-title":"J Wildl Dis","first-page":"863","article-title":"Hematology, parasitology, and serology of free-ranging coyotes (Canis latrans) from South Carolina","volume":"45","author":"Miller","year":"2009"},{"key":"b21","series-title":"Geospat Health","first-page":"15","article-title":"Application of a degree-day model of West Nile virus transmission risk to the East Coast of the United States of America","volume":"7","author":"Konrad","year":"2012"},{"key":"b22","series-title":"J Wildl Dis","first-page":"354","article-title":"Survey for West Nile virus antibodies in wild ducks, 2004-06, USA","volume":"52","author":"Hofmeister","year":"2016"},{"key":"b23","article-title":"County steps up efforts to prevent West Nile virus spread","year":"2022"},{"key":"b24","series-title":"Identification and Geographical Distribution of the Mosquitoes of North America, North of Mexico","author":"Darsie","year":"2005"},{"key":"b25","series-title":"Acta Trop","first-page":"106972","article-title":"Employing oxford nanopore technologies (ONT) for understanding the ecology and transmission dynamics of flaviviruses in mosquitoes (Diptera: Culicidae) from Eastern Colombia","volume":"245","author":"Mart\u00ednez","year":"2023"},{"key":"b26","series-title":"Mol Biol Evol","first-page":"3022","article-title":"MEGA11: Molecular evolutionary genetics analysis version 11","volume":"38","author":"Tamura","year":"2021"},{"key":"b27","series-title":"Nucleic Acids Res","first-page":"1792","article-title":"MUSCLE: Multiple sequence alignment with high accuracy and high throughput","volume":"32","author":"Edgar","year":"2004"},{"key":"b28","series-title":"Mol Biol Evol","first-page":"512","article-title":"Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees","volume":"10","author":"Tamura","year":"1993"},{"key":"b29","series-title":"Epidemiol Infect","first-page":"591","article-title":"Estimated cumulative incidence of West Nile virus infection in US adults, 1999\u20132010","volume":"141","author":"Petersen","year":"2013"},{"key":"b30","series-title":"Am J Transplant","first-page":"1959","article-title":"Surveillance for West Nile virus disease \u2013 United States, 2009\u20132018","volume":"21","author":"McDonald","year":"2021"},{"key":"b31","series-title":"Philos Trans R Soc Lond B Biol Sci","first-page":"1871","article-title":"Multi-year evolutionary dynamics of West Nile virus in suburban Chicago, USA, 2005\u20132007","volume":"365","author":"Amore","year":"2010"},{"key":"b32","series-title":"Virology","first-page":"381","article-title":"Fine-scale genetic variation and evolution of West Nile Virus in a transmission \u201chot spot\u201d in suburban Chicago, USA","volume":"374","author":"Bertolotti","year":"2008"},{"key":"b33","series-title":"PLoS One","first-page":"e0227160","article-title":"The drivers of West Nile virus human illness in the Chicago, Illinois, USA area: Fine scale dynamic effects of weather, mosquito infection, social, and biological conditions","volume":"15","author":"Karki","year":"2020"},{"key":"b34","series-title":"Vector Borne Zoonotic Dis","first-page":"73","article-title":"Factors affecting the geographic distribution of West Nile virus in Georgia, USA: 2002\u20132004","volume":"6","author":"Gibbs","year":"2006"},{"key":"b35","series-title":"PLoS One","first-page":"e15437","article-title":"Economic conditions predict prevalence of West Nile virus","volume":"5","author":"Harrigan","year":"2010"},{"key":"b36","series-title":"Clin Infect Dis","first-page":"1565","article-title":"Beyond viruses: Clinical profiles and etiologies associated with encephalitis","volume":"43","author":"Glaser","year":"2006"},{"key":"b37","series-title":"Parasit Vectors","first-page":"547","article-title":"Updated distribution maps of predominant Culex mosquitoes across the Americas","volume":"14","author":"Gorris","year":"2021"},{"key":"b38"},{"key":"b39","series-title":"Parasit Vectors","first-page":"116","article-title":"Geographic variation in the response of Culex pipiens life history traits to temperature","volume":"9","author":"Ruybal","year":"2016"},{"key":"b40","series-title":"Glob Chang Biol","first-page":"2373","article-title":"Temperature variation makes ectotherms more sensitive to climate change","volume":"19","author":"Paaijmans","year":"2013"},{"key":"b41","series-title":"Proc Biol Sci","first-page":"20201065","article-title":"Thermal thresholds heighten sensitivity of West Nile virus transmission to changing temperatures in coastal California","volume":"287","author":"Skaff","year":"2020"},{"key":"b42","series-title":"Acta Trop","first-page":"1","article-title":"Effect of temperature on life history traits during immature development of Aedes aegypti and Culex quinquefasciatus (Diptera: Culicidae) from C\u00f3rdoba city, Argentina","volume":"146","author":"Grech","year":"2015"},{"key":"b43"},{"key":"b44","series-title":"Integr Comp Biol","first-page":"45","article-title":"Fine-scale microclimatic variation can shape the responses of organisms to global change in both natural and urban environments","volume":"56","author":"Pincebourde","year":"2016"},{"key":"b45","series-title":"PLoS Negl Trop Dis","first-page":"e0005640","article-title":"Fine-scale variation in microclimate across an urban landscape shapes variation in mosquito population dynamics and the 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