{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,3]],"date-time":"2026-06-03T15:09:59Z","timestamp":1780499399365,"version":"3.54.1"},"reference-count":94,"publisher":"American Society of Tropical Medicine and Hygiene","issue":"6","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,6,3]]},"abstract":"<jats:title>ABSTRACT.<\/jats:title>\n                  <jats:p>\n                    <jats:italic>Anopheles stephensi<\/jats:italic>\n                    is rapidly expanding across Africa, posing new challenges for malaria control. Its biting time patterns, however, remain poorly characterized, raising uncertainty about the effectiveness of bed nets against this invasive vector. To address this gap, we investigated diel biting activity, feeding propensity, and flight behavior using complementary behavioral assays on females reared from wild-caught larvae in Hawassa City, Ethiopia. Flight monitoring assays revealed that\n                    <jats:italic>An. stephensi<\/jats:italic>\n                    exhibited pronounced activity at dusk, beginning between 18:00 and 19:00 with scotophase onset, and little to no activity during photophase. Blood-feeding propensity, defined as the proportion of mosquitoes taking a blood meal when offered, peaked during early scotophase (18:00\u201322:00) at 33.3\u201351.7% but was markedly reduced during daylight hours (0\u201316.7%). Human landing catches in large-cage enclosures confirmed this early-evening activity: 83.5% of total landings occurred between 18:00 and 22:00, with a sharp peak at 18:00\u201319:00, corresponding to a mean biting rate of 17.8 bites per person per hour. These findings demonstrate that invasive\n                    <jats:italic>An. stephensi<\/jats:italic>\n                    primarily seek hosts and bite during the early evening, when people are often unprotected by bed nets. This behavior reduces the protective impact of conventional net-based interventions and underscores the need for African National Malaria Control Programs to deploy complementary measures such as spatial repellents and larval source management to mitigate early-evening transmission. Moreover, this study highlights the utility of integrated behavioral assays for estimating biting time, offering approaches that can be extended to other vector species across Africa.\n                  <\/jats:p>","DOI":"10.4269\/ajtmh.25-0589","type":"journal-article","created":{"date-parts":[[2026,4,14]],"date-time":"2026-04-14T13:32:12Z","timestamp":1776173532000},"page":"1114-1119","source":"Crossref","is-referenced-by-count":0,"title":["Early-Evening Biting by Anopheles stephensi in Southern Ethiopia: A Challenge for Bed Net Use as Vector Control in Africa"],"prefix":"10.4269","volume":"114","author":[{"given":"Maxwell G.","family":"Machani","sequence":"first","affiliation":[{"name":"Centre for Global Health Research, Kenya Medical Research Institute, Kisumu, Kenya;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Samuel S. C.","family":"Rund","sequence":"additional","affiliation":[{"name":"Center for Research Computing, University of Notre Dame, Notre Dame, Indiana, USA;"},{"name":"Department of Biological Sciences, University of Notre Dame, Notre Dame, Indiana, USA;"},{"name":"Eck Institute of Global Health, University of Notre Dame, Notre Dame, Indiana, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dawit","family":"Hawaria","sequence":"additional","affiliation":[{"name":"Malaria and Other Vector-Borne Diseases Research Center, College of Medicine and Health Science, Hawassa University, Ethiopia;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guiyun","family":"Yan","sequence":"additional","affiliation":[{"name":"School of Population and Public Health, University of California, Irvine, California, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"2795","reference":[{"key":"b1","series-title":"Acta Trop","first-page":"180","article-title":"First detection of Anopheles stephensi Liston, 1901 (Diptera: Culicidae) in Ethiopia using molecular and morphological approaches","volume":"188","author":"Carter","year":"2018"},{"issue":"12","key":"b2","series-title":"Emerg Infect Dis","first-page":"2498","article-title":"Detection of Anopheles stephensi mosquitoes by molecular surveillance, Kenya","volume":"29","author":"Ochomo","year":"2023"},{"key":"b3","series-title":"Acta Trop","first-page":"39","article-title":"First record of the Asian malaria vector Anopheles stephensi and its possible role in the resurgence of malaria in Djibouti, Horn of Africa","volume":"139","author":"Faulde","year":"2014"},{"key":"b4","article-title":"Vector Alert: Anopheles Stephensi Invasion and Spread in Africa and Sri Lanka.","year":"2023"},{"key":"b5","series-title":"Proc Natl Acad Sci USA","first-page":"24900","article-title":"A new malaria vector in Africa: Predicting the expansion range of Anopheles stephensi and identifying the urban populations at risk","volume":"117","author":"Sinka","year":"2020"},{"issue":"1","key":"b6","series-title":"BMC Med","first-page":"135","article-title":"The potential impact of Anopheles stephensi establishment on the transmission of Plasmodium falciparum in Ethiopia and prospective control measures","volume":"20","author":"Hamlet","year":"2022"},{"issue":"1","key":"b7","series-title":"Sci Rep","first-page":"19878","article-title":"Spatial distribution and population structure of the invasive Anopheles stephensi in Kenya from 2022 to 2024","volume":"15","author":"Samake","year":"2025"},{"issue":"1","key":"b8","series-title":"Malar J","first-page":"373","article-title":"First report of Anopheles stephensi from southern Ethiopia","volume":"22","author":"Hawaria","year":"2023"},{"issue":"1918","key":"b9","series-title":"Philos Trans R Soc Lond B Biol Sci","first-page":"20230343","article-title":"Seasonal plasticity in daily timing of flight activity in Anopheles stephensi is driven by temperature modulation of dawn entrainment","volume":"380","author":"Rund","year":"2025"},{"issue":"2","key":"b10","series-title":"Insects","first-page":"14","article-title":"Daily rhythms in mosquitoes and their consequences for malaria transmission","volume":"7","author":"Rund","year":"2016"},{"key":"b11","series-title":"Annu Rev Entomol","first-page":"53","article-title":"Mosquito behavior and vector control","volume":"50","author":"Pates","year":"2005"},{"issue":"6","key":"b12","series-title":"Trans R Soc Trop Med Hyg","first-page":"647","article-title":"Four years\u2019 entomological study of the transmission of seasonal malaria in Senegal and the bionomics of Anopheles gambiae and A. arabiensis.","volume":"91","author":"Fontenille","year":"1997"},{"issue":"3","key":"b13","series-title":"Exp Parasitol","first-page":"306","article-title":"Some observations on the biting behavior of Anopheles gambiae s.s., Anopheles arabiensis, and Anopheles funestus and their implications for malaria control","volume":"82","author":"Githeko","year":"1996"},{"issue":"1","key":"b14","series-title":"Sci Rep","first-page":"17569","article-title":"Insecticide resistance and behavioural adaptation as a response to long-lasting insecticidal net deployment in malaria vectors in the Cascades region of Burkina Faso","volume":"11","author":"Sanou","year":"2021"},{"issue":"30","key":"b15","series-title":"Proc Natl Acad Sci USA","first-page":"15086","article-title":"Mosquito feeding behavior and how it influences residual malaria transmission across Africa","volume":"116","author":"Sherrard-Smith","year":"2019"},{"issue":"4","key":"b16","series-title":"Evolution","first-page":"1218","article-title":"The importance of mosquito behavioural adaptations to malaria control in Africa","volume":"67","author":"Gatton","year":"2013"},{"key":"b17","article-title":"Partners Convening: A Regional Response to the Invasion of Anopheles Stephensi in Africa.","year":"2023"},{"issue":"1","key":"b18","series-title":"Indian J Med Res","first-page":"120","article-title":"Mosquito biting activity on humans & detection of Plasmodium falciparum infection in Anopheles stephensi in Goa, India","volume":"135","author":"Korgaonkar","year":"2012"},{"issue":"2","key":"b19","series-title":"Iran J Arthropod-Borne Dis","first-page":"35","article-title":"Mosquito vector biting and community protection in a malarious area, Siahoo District, Hormozgan, Iran","volume":"4","author":"Shahandeh","year":"2010"},{"issue":"1","key":"b20","series-title":"Malar J","first-page":"70","article-title":"Key to the females of Afrotropical Anopheles mosquitoes (Diptera: Culicidae)","volume":"19","author":"Coetzee","year":"2020"},{"issue":"1","key":"b21","series-title":"Parasit Vectors","first-page":"35","article-title":"Geographical distribution of Anopheles stephensi in eastern Ethiopia","volume":"13","author":"Balkew","year":"2020"},{"key":"b22","series-title":"Sci 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Ethiopia","volume":"216","author":"Degefa","year":"2021"},{"issue":"1","key":"b31","series-title":"Parasit Vectors","first-page":"520","article-title":"Impact of nighttime human behavior on exposure to malaria vectors and effectiveness of using long-lasting insecticidal nets in the Ethiopian lowlands and highlands","volume":"17","author":"Esayas","year":"2024"},{"issue":"8","key":"b32","series-title":"Trends Parasitol","first-page":"731","article-title":"Invasive Anopheles stephensi in Africa: Insights from Asia","volume":"40","author":"Taylor","year":"2024"},{"key":"b33","series-title":"Malar J","first-page":"10","article-title":"Colonization of malaria vectors under semi-field conditions as a strategy for maintaining genetic and phenotypic similarity with wild populations","volume":"14","author":"Ng\u2019habi","year":"2015"},{"key":"b34","series-title":"Malar J","first-page":"19","article-title":"Differential effects of inbreeding and selection on male reproductive phenotype associated with the colonization and laboratory maintenance of Anopheles gambiae.","volume":"13","author":"Baeshen","year":"2014"},{"issue":"12","key":"b35","series-title":"Heliyon","first-page":"e12178","article-title":"Malaria vector feeding, peak biting time and resting place preference behaviors in line with Indoor based intervention tools and its implication: Scenario from selected sentinel sites of Ethiopia","volume":"8","author":"Bedasso","year":"2022"},{"issue":"1","key":"b36","series-title":"Malar J","first-page":"66","article-title":"Early morning anopheline mosquito biting, a potential driver of malaria transmission in Busia County, western Kenya","volume":"23","author":"Odero","year":"2024"},{"issue":"1","key":"b37","series-title":"Parasit Vectors","first-page":"376","article-title":"Differences in malaria vector biting behavior and changing vulnerability to malaria transmission in contrasting ecosystems of western Kenya","volume":"16","author":"Nzioki","year":"2023"},{"key":"b38","series-title":"Parasit Vectors","first-page":"48","article-title":"Diel flight activity of wild-caught Anopheles farauti (s.s.) and An. hinesorum malaria mosquitoes from northern Queensland, Australia","volume":"12","author":"Duffield","year":"2019"},{"key":"b39","series-title":"Front Physiol","first-page":"199","article-title":"Why use of interventions targeting outdoor biting mosquitoes will be necessary to achieve malaria elimination","volume":"3","author":"Govella","year":"2012"},{"key":"b40","series-title":"Malar J","first-page":"97","article-title":"Potential causes and consequences of behavioural resilience and resistance in malaria vector populations: A mathematical modelling analysis","volume":"13","author":"Killeen","year":"2014"},{"key":"b41","series-title":"Malar J","first-page":"259","article-title":"\u201cA bite before bed\u201d: Exposure to malaria vectors outside the times of net use in the highlands of western Kenya","volume":"14","author":"Cooke","year":"2015"},{"issue":"2","key":"b42","series-title":"Emerg Infect Dis","first-page":"603","article-title":"Anopheles stephensi mosquitoes as vectors of Plasmodium vivax and falciparum, Horn of Africa, 2019","volume":"27","author":"Tadesse","year":"2021"},{"issue":"12","key":"b43","series-title":"Nat Med","first-page":"3203","article-title":"Evidence for a role of Anopheles stephensi in the spread of drug- and diagnosis-resistant malaria in Africa","volume":"29","author":"Emiru","year":"2023"},{"issue":"2","key":"b44","series-title":"Trends Parasitol","first-page":"102","article-title":"Anopheles stephensi ecology and control in Africa","volume":"40","author":"Zhou","year":"2024"},{"issue":"10473","key":"b45","series-title":"Lancet","first-page":"147","article-title":"Effect of a spatial repellent on malaria incidence in an area of western Kenya characterised by high malaria transmission, insecticide resistance, and universal coverage of insecticide treated nets (part of the AEGIS Consortium): A cluster-randomised, controlled trial","volume":"405","author":"Ochomo","year":"2025"},{"key":"b46","series-title":"Malar J","first-page":"19","article-title":"MalariaSphere: A greenhouse-enclosed simulation of a natural Anopheles gambiae (Diptera: Culicidae) ecosystem in western Kenya","volume":"1","author":"Knols","year":"2002"},{"key":"b47","series-title":"Malar J","first-page":"158","article-title":"Establishment of a large semi-field system for experimental study of African malaria vector ecology and control in Tanzania","volume":"7","author":"Ferguson","year":"2008"},{"key":"b1","series-title":"Acta Trop","first-page":"180","article-title":"First detection of Anopheles stephensi Liston, 1901 (Diptera: Culicidae) in Ethiopia using molecular and morphological approaches","volume":"188","author":"Carter","year":"2018"},{"issue":"12","key":"b2","series-title":"Emerg Infect Dis","first-page":"2498","article-title":"Detection of Anopheles stephensi mosquitoes by molecular surveillance, Kenya","volume":"29","author":"Ochomo","year":"2023"},{"key":"b3","series-title":"Acta Trop","first-page":"39","article-title":"First record of the Asian malaria vector Anopheles stephensi and its possible role in the resurgence of malaria in Djibouti, Horn of Africa","volume":"139","author":"Faulde","year":"2014"},{"key":"b4","article-title":"Vector Alert: Anopheles Stephensi Invasion and Spread in Africa and Sri Lanka.","year":"2023"},{"key":"b5","series-title":"Proc Natl Acad Sci USA","first-page":"24900","article-title":"A new malaria vector in Africa: Predicting the expansion range of Anopheles stephensi and identifying the urban populations at risk","volume":"117","author":"Sinka","year":"2020"},{"issue":"1","key":"b6","series-title":"BMC Med","first-page":"135","article-title":"The potential impact of Anopheles stephensi establishment on the transmission of Plasmodium falciparum in Ethiopia and prospective control measures","volume":"20","author":"Hamlet","year":"2022"},{"issue":"1","key":"b7","series-title":"Sci Rep","first-page":"19878","article-title":"Spatial distribution and population structure of the invasive Anopheles stephensi in Kenya from 2022 to 2024","volume":"15","author":"Samake","year":"2025"},{"issue":"1","key":"b8","series-title":"Malar J","first-page":"373","article-title":"First report of Anopheles stephensi from southern Ethiopia","volume":"22","author":"Hawaria","year":"2023"},{"issue":"1918","key":"b9","series-title":"Philos Trans R Soc Lond B Biol Sci","first-page":"20230343","article-title":"Seasonal plasticity in daily timing of flight activity in Anopheles stephensi is driven by temperature modulation of dawn entrainment","volume":"380","author":"Rund","year":"2025"},{"issue":"2","key":"b10","series-title":"Insects","first-page":"14","article-title":"Daily rhythms in mosquitoes and their consequences for malaria transmission","volume":"7","author":"Rund","year":"2016"},{"key":"b11","series-title":"Annu Rev Entomol","first-page":"53","article-title":"Mosquito behavior and vector control","volume":"50","author":"Pates","year":"2005"},{"issue":"6","key":"b12","series-title":"Trans R Soc Trop Med Hyg","first-page":"647","article-title":"Four years\u2019 entomological study of the transmission of seasonal malaria in Senegal and the bionomics of Anopheles gambiae and A. arabiensis.","volume":"91","author":"Fontenille","year":"1997"},{"issue":"3","key":"b13","series-title":"Exp Parasitol","first-page":"306","article-title":"Some observations on the biting behavior of Anopheles gambiae s.s., Anopheles arabiensis, and Anopheles funestus and their implications for malaria control","volume":"82","author":"Githeko","year":"1996"},{"issue":"1","key":"b14","series-title":"Sci Rep","first-page":"17569","article-title":"Insecticide resistance and behavioural adaptation as a response to long-lasting insecticidal net deployment in malaria vectors in the Cascades region of Burkina Faso","volume":"11","author":"Sanou","year":"2021"},{"issue":"30","key":"b15","series-title":"Proc Natl Acad Sci USA","first-page":"15086","article-title":"Mosquito feeding behavior and how it influences residual malaria transmission across Africa","volume":"116","author":"Sherrard-Smith","year":"2019"},{"issue":"4","key":"b16","series-title":"Evolution","first-page":"1218","article-title":"The importance of mosquito behavioural adaptations to malaria control in Africa","volume":"67","author":"Gatton","year":"2013"},{"key":"b17","article-title":"Partners Convening: A Regional Response to the Invasion of Anopheles Stephensi in Africa.","year":"2023"},{"issue":"1","key":"b18","series-title":"Indian J Med Res","first-page":"120","article-title":"Mosquito biting activity on humans & detection of Plasmodium falciparum infection in Anopheles stephensi in Goa, India","volume":"135","author":"Korgaonkar","year":"2012"},{"issue":"2","key":"b19","series-title":"Iran J Arthropod-Borne Dis","first-page":"35","article-title":"Mosquito vector biting and community protection in a malarious area, Siahoo District, Hormozgan, Iran","volume":"4","author":"Shahandeh","year":"2010"},{"issue":"1","key":"b20","series-title":"Malar J","first-page":"70","article-title":"Key to the females of Afrotropical Anopheles mosquitoes (Diptera: Culicidae)","volume":"19","author":"Coetzee","year":"2020"},{"issue":"1","key":"b21","series-title":"Parasit Vectors","first-page":"35","article-title":"Geographical distribution of Anopheles stephensi in eastern Ethiopia","volume":"13","author":"Balkew","year":"2020"},{"key":"b22","series-title":"Sci Rep","first-page":"2494","article-title":"Daily rhythms in antennal protein and olfactory sensitivity in the malaria mosquito Anopheles gambiae.","volume":"3","author":"Rund","year":"2013"},{"issue":"12","key":"b23","series-title":"J Insect Physiol","first-page":"1609","article-title":"Strain- and sex-specific differences in daily flight activity and the circadian clock of Anopheles gambiae mosquitoes","volume":"58","author":"Rund","year":"2012"},{"key":"b24","series-title":"Parasit Vectors","first-page":"187","article-title":"Monitoring population and environmental parameters of invasive mosquito species in Europe","volume":"7","author":"Petri\u0107","year":"2014"},{"issue":"5","key":"b25","series-title":"Ecology","first-page":"985","article-title":"Do invasive species perform better in their new ranges?","volume":"94","author":"Parker","year":"2013"},{"issue":"1","key":"b26","series-title":"PLoS Negl Trop Dis","first-page":"e0007831","article-title":"The importance of vector control for the control and elimination of vector-borne diseases","volume":"14","author":"Wilson","year":"2020"},{"issue":"21","key":"b27","series-title":"Proc Natl Acad Sci USA","first-page":"e2104282119","article-title":"Diurnal biting of malaria mosquitoes in the Central African Republic indicates residual transmission may be \u201cout of control.\u201d","volume":"119","author":"Sangbakembi-Ngounou","year":"2022"},{"key":"b28","series-title":"Curr Opin Insect Sci","first-page":"101179","article-title":"Circadian and daily rhythms of disease vector mosquitoes","volume":"63","author":"Duffield","year":"2024"},{"key":"b29","series-title":"BMC Physiol","first-page":"23","article-title":"Molecular analysis of photic inhibition of blood-feeding in Anopheles gambiae.","volume":"8","author":"Das","year":"2008"},{"key":"b30","series-title":"Acta Trop","first-page":"105837","article-title":"Patterns of human exposure to early evening and outdoor biting mosquitoes and residual malaria transmission in 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associated with the colonization and laboratory maintenance of Anopheles gambiae.","volume":"13","author":"Baeshen","year":"2014"},{"issue":"12","key":"b35","series-title":"Heliyon","first-page":"e12178","article-title":"Malaria vector feeding, peak biting time and resting place preference behaviors in line with Indoor based intervention tools and its implication: Scenario from selected sentinel sites of Ethiopia","volume":"8","author":"Bedasso","year":"2022"},{"issue":"1","key":"b36","series-title":"Malar J","first-page":"66","article-title":"Early morning anopheline mosquito biting, a potential driver of malaria transmission in Busia County, western Kenya","volume":"23","author":"Odero","year":"2024"},{"issue":"1","key":"b37","series-title":"Parasit Vectors","first-page":"376","article-title":"Differences in malaria vector biting behavior and changing vulnerability to malaria transmission in contrasting ecosystems of western 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western Kenya","volume":"14","author":"Cooke","year":"2015"},{"issue":"2","key":"b42","series-title":"Emerg Infect Dis","first-page":"603","article-title":"Anopheles stephensi mosquitoes as vectors of Plasmodium vivax and falciparum, Horn of Africa, 2019","volume":"27","author":"Tadesse","year":"2021"},{"issue":"12","key":"b43","series-title":"Nat Med","first-page":"3203","article-title":"Evidence for a role of Anopheles stephensi in the spread of drug- and diagnosis-resistant malaria in Africa","volume":"29","author":"Emiru","year":"2023"},{"issue":"2","key":"b44","series-title":"Trends Parasitol","first-page":"102","article-title":"Anopheles stephensi ecology and control in Africa","volume":"40","author":"Zhou","year":"2024"},{"issue":"10473","key":"b45","series-title":"Lancet","first-page":"147","article-title":"Effect of a spatial repellent on malaria incidence in an area of western Kenya characterised by high malaria transmission, insecticide resistance, and universal coverage of 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