{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T11:47:58Z","timestamp":1793533678865,"version":"4.5.0"},"reference-count":59,"publisher":"Public Library of Science (PLoS)","issue":"8","license":[{"start":{"date-parts":[[2025,8,28]],"date-time":"2025-08-28T00:00:00Z","timestamp":1756339200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/"}],"funder":[{"DOI":"10.13039\/100018190","name":"Cornell Atkinson Center for Sustainability, Cornell University","doi-asserted-by":"publisher","award":["Academic Venture Fund"],"award-info":[{"award-number":["Academic Venture Fund"]}],"id":[{"id":"10.13039\/100018190","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100018190","name":"Cornell Atkinson Center for Sustainability, Cornell University","doi-asserted-by":"publisher","award":["Rapid Response Fund"],"award-info":[{"award-number":["Rapid Response Fund"]}],"id":[{"id":"10.13039\/100018190","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plosntds.org"],"crossmark-restriction":false},"short-container-title":["PLoS Negl Trop Dis"],"abstract":"<jats:p><jats:italic>Aedes aegypti,<\/jats:italic> the primary vector of dengue virus, is predominantly considered an urban mosquito, especially in the Americas, where its reemergence began in cities after the end of continent-wide eradication campaigns. The results of our study diverge from this narrative, demonstrating the recent and widespread rural invasion of <jats:italic>Ae. aegypti<\/jats:italic> along major shipping routes in the northern Peruvian Amazon between the major cities of Iquitos, Pucallpa, and Yurimaguas. Using prokopack aspirators to conduct indoor mosquito collections, we identified <jats:italic>Ae. aegypti<\/jats:italic> populations in 29 of 30 sites surveyed across a rural to urban gradient and quantified <jats:italic>Ae. aegypti<\/jats:italic> adult metrics. In multiple instances, adult <jats:italic>Ae. aegypti<\/jats:italic> indices in rural villages were equal to or greater than indices in dengue-endemic cities, suggesting the entomological risk level in some rural areas is sufficient to support dengue transmission. Fourteen rural sites were sampled in transects from the community river port into town. In seven of these sites, houses closer to the port were significantly more likely to be infested with <jats:italic>Ae. aegypti<\/jats:italic> adults than houses further from the ports, and four additional sites showed a similar trend. This pattern suggests that <jats:italic>Ae. aegypti<\/jats:italic> is still actively invading many rural sites by adult <jats:italic>Ae. aegypti<\/jats:italic> disembarking from boats at the port, finding nearby oviposition sites, and advancing stepwise towards the interior, with sections of towns still <jats:italic>Ae. aegypti-<\/jats:italic>free. Only one site showed a strong signal of invasion via the egg or larval stage, with a focus of <jats:italic>Ae. aegypti<\/jats:italic> far removed from the port. The widespread infestation of <jats:italic>Ae. aegypti<\/jats:italic> in rural areas is a major public health threat given the far distance of communities to hospital care. It is important to implement control measures now before the mosquito gains a stronger foothold in zones of active invasion.<\/jats:p>","DOI":"10.1371\/journal.pntd.0012506","type":"journal-article","created":{"date-parts":[[2025,8,28]],"date-time":"2025-08-28T17:40:27Z","timestamp":1756402827000},"page":"e0012506","update-policy":"https:\/\/doi.org\/10.1371\/journal.pntd.corrections_policy","source":"Crossref","is-referenced-by-count":8,"title":["Changing dynamics of Aedes aegypti invasion and vector-borne disease risk for rural communities in the Peruvian Amazon"],"prefix":"10.1371","volume":"19","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5850-9948","authenticated-orcid":true,"given":"Kara","family":"Fikrig","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Arnold O.","family":"Noriega","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rosa A.","family":"Rodriguez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"John","family":"Bardales","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jos\u00e9","family":"Rivas","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Becker","family":"Reyna","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guido","family":"Izquierdo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gissella M.","family":"Vasquez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryan T.","family":"Larson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amy C.","family":"Morrison","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Laura C.","family":"Harrington","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2025,8,28]]},"reference":[{"issue":"10427","key":"pntd.0012506.ref001","doi-asserted-by":"crossref","first-page":"667","DOI":"10.1016\/S0140-6736(23)02576-X","article-title":"Dengue","volume":"403","author":"G Paz-Bailey","year":"2024","journal-title":"The Lancet"},{"issue":"7446","key":"pntd.0012506.ref002","doi-asserted-by":"crossref","first-page":"504","DOI":"10.1038\/nature12060","article-title":"The global distribution and burden of dengue","volume":"496","author":"S Bhatt","year":"2013","journal-title":"Nature"},{"issue":"8","key":"pntd.0012506.ref003","doi-asserted-by":"crossref","first-page":"935","DOI":"10.1016\/S1473-3099(16)00146-8","article-title":"The global economic burden of dengue: a systematic analysis","volume":"16","author":"DS Shepard","year":"2016","journal-title":"Lancet Infect Dis"},{"issue":"1","key":"pntd.0012506.ref004","doi-asserted-by":"crossref","first-page":"7","DOI":"10.2105\/AJPH.53.1.7","article-title":"The elimination of urban yellow fever in the Americas through the eradication of Aedes aegypti","volume":"53","author":"FL Soper","year":"1963","journal-title":"Am J Public Health Nations Health"},{"issue":"3","key":"pntd.0012506.ref005","first-page":"201","article-title":"First documented outbreak of dengue in the Peruvian Amazon region","volume":"26","author":"I Phillips","year":"1992","journal-title":"Bull Pan Am Health Organ"},{"issue":"4","key":"pntd.0012506.ref006","doi-asserted-by":"crossref","first-page":"584","DOI":"10.4269\/ajtmh.2012.11-0770","article-title":"The history of dengue outbreaks in the Americas","volume":"87","author":"O Brathwaite Dick","year":"2012","journal-title":"Am J Trop Med Hyg"},{"issue":"9","key":"pntd.0012506.ref007","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0009631","article-title":"The role of urbanisation in the spread of Aedes mosquitoes and the diseases they transmit-a systematic review","volume":"15","author":"A Kolimenakis","year":"2021","journal-title":"PLoS Negl Trop Dis"},{"issue":"6","key":"pntd.0012506.ref008","article-title":"Characterizing dengue transmission in rural areas: a systematic review","volume":"17","author":"O Man","year":"2023","journal-title":"PLoS Negl Trop Dis"},{"issue":"2","key":"pntd.0012506.ref009","first-page":"169","article-title":"Aedes aegypti in rural areas: public health implications","volume":"36","author":"VA Olano","year":"2016","journal-title":"Biomedica"},{"issue":"1","key":"pntd.0012506.ref010","doi-asserted-by":"crossref","first-page":"356","DOI":"10.1186\/s13071-017-2295-1","article-title":"A cross-sectional survey of Aedes aegypti immature abundance in urban and rural household containers in central Colombia","volume":"10","author":"HJ Overgaard","year":"2017","journal-title":"Parasit Vectors"},{"issue":"1","key":"pntd.0012506.ref011","article-title":"Comportamiento estacional y temporal de Aedes aegypti y Aedes albopictus en La Habana, Cuba","volume":"60","author":"MDC Marquetti","year":"2008","journal-title":"Revista Cubana de Medicina Tropical"},{"issue":"1","key":"pntd.0012506.ref012","first-page":"89","article-title":"Primer registro de Aedes (Stegomyia) aegypti (L.) en \u00e1reas remotas del estado Bol\u00edvar","volume":"51","author":"Y Rubio-Palis","year":"2011","journal-title":"Bolet\u00edn de Malariolog\u00eda y Salud Ambiental"},{"issue":"3","key":"pntd.0012506.ref013","first-page":"163","article-title":"Expansi\u00f3n del Aedes aegypti a localidades rurales de Cajamarca","volume":"23","author":"L Troyes","year":"2006","journal-title":"Revista Peruana de Medicina Experimental y Salud Publica"},{"issue":"4","key":"pntd.0012506.ref014","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1590\/0074-02760150363","article-title":"Detection of all four dengue serotypes in Aedes aegypti female mosquitoes collected in a rural area in Colombia","volume":"111","author":"R P\u00e9rez-Castro","year":"2016","journal-title":"Mem Inst Oswaldo Cruz"},{"issue":"0","key":"pntd.0012506.ref015","doi-asserted-by":"crossref","first-page":"193","DOI":"10.7705\/biomedica.v37i0.3584","article-title":"Dengue virus detection in Aedes aegypti larvae and pupae collected in rural areas of Anapoima, Cundinamarca, Colombia","volume":"37","author":"ML Velandia-Romero","year":"2017","journal-title":"Biomedica"},{"issue":"9","key":"pntd.0012506.ref016","article-title":"A dengue outbreak in a rural community in Northern Coastal Ecuador: An analysis using unmanned aerial vehicle mapping","volume":"15","author":"GO Lee","year":"2021","journal-title":"PLoS Negl Trop Dis"},{"issue":"1","key":"pntd.0012506.ref017","article-title":"Dengue serotype differences in urban and semi-rural communities in Ecuador","volume":"10","author":"S M\u00e1rquez","year":"2018","journal-title":"ACI Avances en Ciencias e Ingenierias"},{"key":"pntd.0012506.ref018","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1016\/j.socscimed.2016.12.010","article-title":"What caused the 2012 dengue outbreak in Pucallpa, Peru? A socio-ecological autopsy","volume":"174","author":"M Charette","year":"2017","journal-title":"Soc Sci Med"},{"issue":"3","key":"pntd.0012506.ref019","first-page":"175","article-title":"Comportamiento poblacional de larvas de aedes egypti para estimar los casos de dengue en Yurimaguas, Per\u00fa, 2000-2004","volume":"22","author":"W Fern\u00e1ndez","year":"2005","journal-title":"Revista peruana de medicina experimental y Salud P\u00fablica"},{"issue":"5","key":"pntd.0012506.ref020","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0000670","article-title":"Epidemiology of dengue virus in Iquitos, Peru 1999 to 2005: interepidemic and epidemic patterns of transmission","volume":"4","author":"AC Morrison","year":"2010","journal-title":"PLoS Negl Trop Dis"},{"issue":"8","key":"pntd.0012506.ref021","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0003033","article-title":"Patterns of geographic expansion of Aedes aegypti in the Peruvian Amazon","volume":"8","author":"SA Guagliardo","year":"2014","journal-title":"PLoS Negl Trop Dis"},{"issue":"9","key":"pntd.0012506.ref022","article-title":"The genetic structure of Aedes aegypti populations is driven by boat traffic in the Peruvian Amazon","volume":"13","author":"SAJ Guagliardo","year":"2019","journal-title":"PLoS Negl Trop Dis"},{"issue":"4","key":"pntd.0012506.ref023","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0003648","article-title":"River boats contribute to the regional spread of the dengue vector Aedes aegypti in the Peruvian Amazon","volume":"9","author":"SA Guagliardo","year":"2015","journal-title":"PLoS Negl Trop Dis"},{"issue":"4","key":"pntd.0012506.ref024","doi-asserted-by":"crossref","first-page":"726","DOI":"10.1093\/jme\/tjv048","article-title":"Evidence for Aedes aegypti (Diptera: Culicidae) Oviposition on Boats in the Peruvian Amazon","volume":"52","author":"SA Guagliardo","year":"2015","journal-title":"J Med Entomol"},{"issue":"6","key":"pntd.0012506.ref025","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0196988","article-title":"The Amazon Hope: a qualitative and quantitative assessment of a mobile clinic ship in the Peruvian Amazon","volume":"13","author":"NP Limaye","year":"2018","journal-title":"PLoS One"},{"issue":"1","key":"pntd.0012506.ref026","doi-asserted-by":"crossref","first-page":"180","DOI":"10.4269\/ajtmh.13-0547","article-title":"Healthcare access and health beliefs of the indigenous peoples in remote Amazonian Peru","volume":"90","author":"CK Brierley","year":"2014","journal-title":"Am J Trop Med Hyg"},{"key":"pntd.0012506.ref027","unstructured":"Nacional IG. L\u00edmites Departamentales. 2023."},{"key":"pntd.0012506.ref028","unstructured":"(ROLAC) OLAatC. Hidrograf\u00eda de Per\u00fa. 2015."},{"issue":"6","key":"pntd.0012506.ref029","doi-asserted-by":"crossref","first-page":"1256","DOI":"10.1603\/033.046.0602","article-title":"A new, cost-effective, battery-powered aspirator for adult mosquito collections","volume":"46","author":"GM Vazquez-Prokopec","year":"2009","journal-title":"J Med Entomol"},{"key":"pntd.0012506.ref030","volume-title":"Identification and geographical distribution of the mosquitoes of North America, north of Mexico","author":"RF Darsie","year":"2005"},{"key":"pntd.0012506.ref031","volume-title":"Loreto Censos Nacionales de Poblaci\u00f3n y Vivienda 2017","year":"2017"},{"key":"pntd.0012506.ref032","volume-title":"Ucayali Censos Nacionales de Poblaci\u00f3n y Vivienda 2017","year":"2017"},{"key":"pntd.0012506.ref033","volume-title":"R: A Language and Environment for Statistical Computing","author":"RCoreTeam","year":"2019"},{"key":"pntd.0012506.ref034","volume-title":"emmeans: estimated marginal means, aka least-squares means. R package v. 1.3. 4","author":"R Lenth","year":"2019"},{"key":"pntd.0012506.ref035","article-title":"Fitting linear mixed-effects models using lme4","author":"D Bates","year":"2014","journal-title":"arXiv preprint"},{"key":"pntd.0012506.ref036","volume-title":"Package \u2018glmmtmb\u2019. R Package Version 02 0","author":"A Magnusson","year":"2017"},{"key":"pntd.0012506.ref037","volume-title":"Changing dynamics of Aedes aegypti invasion and vector-borne disease risk for rural communities in the Peruvian Amazon","author":"K Fikrig","year":"2025"},{"key":"pntd.0012506.ref038","volume-title":"Population of Cities in Peru 2024","author":"Review WP","year":"2024"},{"key":"pntd.0012506.ref039","volume-title":"Ucayali Resultados Definitivos","year":"2018"},{"key":"pntd.0012506.ref040","volume-title":"Loreto Resultados Definitivos","year":"2018"},{"issue":"3","key":"pntd.0012506.ref041","doi-asserted-by":"crossref","first-page":"502","DOI":"10.1603\/0022-2585-41.3.502","article-title":"Evaluation of a sampling methodology for rapid assessment of Aedes aegypti infestation levels in Iquitos, Peru","volume":"41","author":"AC Morrison","year":"2004","journal-title":"J Med Entomol"},{"issue":"3","key":"pntd.0012506.ref042","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0005429","article-title":"The relationship between entomological indicators of Aedes aegypti abundance and dengue virus infection","volume":"11","author":"EA Cromwell","year":"2017","journal-title":"PLoS Negl Trop Dis"},{"issue":"4","key":"pntd.0012506.ref043","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0006378","article-title":"Efficacy of Aedes aegypti control by indoor Ultra Low Volume (ULV) insecticide spraying in Iquitos, Peru","volume":"12","author":"CE Gunning","year":"2018","journal-title":"PLoS Negl Trop Dis"},{"issue":"26","key":"pntd.0012506.ref044","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2118283119","article-title":"Efficacy of a spatial repellent for control of Aedes-borne virus transmission: a cluster-randomized trial in Iquitos, Peru","volume":"119","author":"AC Morrison","year":"2022","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"6","key":"pntd.0012506.ref045","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0003829","article-title":"Human-mediated marine dispersal influences the population structure of Aedes aegypti in the Philippine Archipelago","volume":"9","author":"E Fonzi","year":"2015","journal-title":"PLoS Negl Trop Dis"},{"issue":"9","key":"pntd.0012506.ref046","article-title":"Genetic evidence of expansion by passive transport of Aedes (Stegomyia) aegypti in Eastern Argentina","volume":"10","author":"LM D\u00edaz-Nieto","year":"2016","journal-title":"PLoS Negl Trop Dis"},{"issue":"4805","key":"pntd.0012506.ref047","doi-asserted-by":"crossref","first-page":"1114","DOI":"10.1126\/science.3576225","article-title":"Aedes albopictus in North America: probable introduction in used tires from northern Asia","volume":"236","author":"WA Hawley","year":"1987","journal-title":"Science"},{"issue":"3","key":"pntd.0012506.ref048","first-page":"494","article-title":"The used tire trade: a mechanism for the worldwide dispersal of container breeding mosquitoes","volume":"3","author":"P Reiter","year":"1987","journal-title":"J Am Mosq Control Assoc"},{"key":"pntd.0012506.ref049","article-title":"Pathways for introduction and dispersal of invasive Aedes mosquito species in Europe: a review","volume":"38","author":"A Ib\u00e1\u00f1ez-Justicia","year":"2020","journal-title":"J Eur Mosq Control Assoc"},{"key":"pntd.0012506.ref050","volume-title":"Madre de Dios Resultados Definitivos","year":"2018"},{"key":"pntd.0012506.ref051","volume-title":"San Mart\u00edn Resultados Definitivos","year":"2018"},{"key":"pntd.0012506.ref052","volume-title":"Hu\u00e1nuco Resultados Definitivos","year":"2018"},{"issue":"1","key":"pntd.0012506.ref053","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/1742-7622-2-1","article-title":"Dengue fever: new paradigms for a changing epidemiology","volume":"2","author":"D Guha-Sapir","year":"2005","journal-title":"Emerg Themes Epidemiol"},{"issue":"14","key":"pntd.0012506.ref054","doi-asserted-by":"crossref","first-page":"1157","DOI":"10.1001\/jama.1996.03540140045025","article-title":"Yellow fever: a decade of reemergence","volume":"276","author":"SE Robertson","year":"1996","journal-title":"JAMA"},{"issue":"8","key":"pntd.0012506.ref055","doi-asserted-by":"crossref","first-page":"926","DOI":"10.3201\/eid0908.030075","article-title":"Enzootic transmission of yellow fever virus in Peru","volume":"9","author":"J Bryant","year":"2003","journal-title":"Emerg Infect Dis"},{"issue":"9","key":"pntd.0012506.ref056","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0005897","article-title":"Geographic patterns and environmental factors associated with human yellow fever presence in the Americas","volume":"11","author":"PN Hamrick","year":"2017","journal-title":"PLoS Negl Trop Dis"},{"issue":"6686","key":"pntd.0012506.ref057","doi-asserted-by":"crossref","first-page":"957","DOI":"10.1126\/science.ado0050","article-title":"Peru\u2019s zoning amendment endangers forests","volume":"383","author":"C Martel","year":"2024","journal-title":"Science"},{"issue":"4","key":"pntd.0012506.ref058","doi-asserted-by":"crossref","DOI":"10.1002\/rmv.2333","article-title":"Aedes mosquitoes in the emerging threat of urban yellow fever transmission","volume":"32","author":"G Gabiane","year":"2022","journal-title":"Rev Med Virol"},{"issue":"12","key":"pntd.0012506.ref059","doi-asserted-by":"crossref","first-page":"724","DOI":"10.1016\/j.micinf.2016.10.007","article-title":"Mayaro virus: a forest virus primed for a trip to the city?","volume":"18","author":"IM Mackay","year":"2016","journal-title":"Microbes Infect"}],"container-title":["PLOS Neglected Tropical Diseases"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pntd.0012506","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,8,28]],"date-time":"2025-08-28T17:40:39Z","timestamp":1756402839000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pntd.0012506"}},"subtitle":[],"editor":[{"given":"Clarence","family":"Mang'era","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2025,8,28]]},"references-count":59,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2025,8,28]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pntd.0012506","relation":{},"ISSN":["1935-2735"],"issn-type":[{"value":"1935-2735","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,28]]}}}