J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 1 http://jad.tums.ac.ir Published Online: March 31, 2023 Review article Worldwide Status of Insecticide Resistance of Aedes aegypti and Ae. albopictus, Vectors of Arboviruses of Chikungunya, Dengue, Zika and Yellow Fever Tahereh Sadat Asgarian1, *Hassan Vatandoost1,2, Ahmad Ali Hanafi-Bojd1,2, Fatemeh Nikpoor2 1Department of Vector Biology and Control of Diseases, School of Public Health, Tehran University of Medical Sciences, Tehran, Iran 2Department of Chemical Pollutants and Pesticides, Institute for Environmental Research, Tehran University of Medical Sciences, Tehran, Iran *Corresponding author: Prof Hassan Vatandoost, E-mail: hvatandoost1@yahoo.com, vatando@tums.ac.ir (Received 19 Aug 2022; accepted 24 Dec 2022) Abstract Background: Controlling of Aedes aegypti and Ae. albopictus, vectors of five important mosquito-borne diseases, is known as the most effective method to prevent the transmission of arboviruses to humans, but the emergence of insecti - cide resistance is threat for control and prevention of vector borne diseases. A better understanding of mosquito re- sistance to insecticides will help to develop more effective methods to control insecticide resistance in mosquito vectors. Methods: Worldwide geographical distribution of insecticide resistance in Ae. aegypti and Ae. albopictus by the availa- ble papers and map of the data for carbamates, organochlorines, organophosphates, pyrethroids, microbial and insect growth regulator insecticides were reviewed. Article data published up to December 2022 were investigated by search- ing the following databases: "Google Scholar", "PubMed", "Scopus", "SID" and "Web of Knowledge". Results: The results showed that the susceptibility and resistance status of Ae. aegypti and Ae. albopictus to insecticides in the world is very diverse. Conclusion: Due to the importance of Ae. aegypti and Ae. albopictus in the transmission of mosquito-borne arbovirus- es, resistance management should be given more attention worldwide to prevent insecticide resistance in the arbovirus vector and replace the new approach for vector control. Keywords: Insecticide; Resistant; Aedes; Arboviruses; World Introduction Among arboviruses that are widespread worldwide, chikungunya, dengue, Zika and yel- low fever are widely transmitted by Aedes spp. (1, 2). Aedes aegypti mosquitoes originated from Africa, but they are also found in tropical, sub- tropical, and temperate parts of the world (3, 4). Aedes albopictus has adapted to urban, sub- urban, and rural regions. This species is a for- est species that has spread from Asia to Afri- ca, America, and Europe through the trade of used tires (5). Dengue is the fastest mosquito transmitted disease, it has increased more than 15 -fold since 2000 and has affected more than 129 countries. Because vaccines or drug treatments exist for only a small number of vector-borne pathogens, the primary method for controlling many vector-borne diseases is direct vector control (6). Chemical control by organic or inorganic insecticides is one of the methods of mosquito control and as part of integrated management. Insecticides used for mosquitoes, that are neu- rotoxic: organochlorines, organophosphates, car- bamates, and pyrethroids (7). Insecticide re- sistance is a growing topic around the world that limits the effectiveness of control methods against vector mosquitoes. The data on insec- ticide resistance in Aedes is varying. Insecti- cides have an important role in the control of dengue fever, but the resistance of mosquitoes Copyright © 2023 The Authors. Published by Tehran University of Medical Sciences. This work is licensed under a Creative Commons Attribution-NonCommercial 4.0 International license (https://creativecommons.org/licenses/by- nc/4.0/). Non-commercial uses of the work are permitted, provided the original work is properly cited. http://jad.tums.ac.ir/ mailto:vatando@tums.ac.ir https://creativecommons.org/licenses/by-nc/4.0/ https://creativecommons.org/licenses/by-nc/4.0/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 2 http://jad.tums.ac.ir Published Online: March 31, 2023 reduces the effectiveness of these interventions in some countries (8). According to the World Health Organization (WHO), resistance to in- secticides is a major risk in interventions to con- trol vector-borne diseases. It is essential to iden- tify the geographical regions where vector re- sistance to insecticides exists and can make it difficult to control the vector, and to improve the induction of innovative tools for vector con- trol. A better identifying of resistance of insecti- cides helps to formulate a global measure to con- trol insecticide resistance in disease vectors. Materials and Methods Published articles were searched by terms “resistance”, “Aedes aegypti”, “Aedes albopic- tus”, “pyrethroid”, “organochlorines”, “organ- ophosphate”, “carbamate”, “IGR”, and “kdr”, “P450”, “monooxygenase”, “glutathione”, or “esterase”. Data were extracted from articles published up to December 2022. The terms in the following databases: PubMed, Web of Knowledge, Scopus, Google Scholar, and SID were reviewed. The WHO guideline was con- sidered for insecticide resistant level. Results Resistant status of Aedes aegypti and Ae. al- bopictus to Organochlorines insecticides In Africa Resistance of two Aedes species to dichlo- rodiphenyltrichloroethane (DDT) has been re- ported from each country where susceptibility testing has been performed. In Ghana (9, 10), Cape Verde, and Senegal (11) high Ae. ae- gypti resistance to DDT was reported. Experi- ments showed one Ae. aegypti population from Gabon and two populations of Ae. albopictus from Cameroon were resistant to DDT (12, 13). Also, in another study adult bioassays of three field populations of Ae. albopictus and Ae. ae- gypti from Cameroon showed different sus- ceptibility levels from 68.75% to 100% against 4% DDT (14). In Central African Republic Ae. aegypti was resistant to DDT and Ae. albopic- tus had a sensitive population and the rest of the populations were tolerant (15). In Lagos State of Nigeria all populations of Ae. aegypti showed resistance to DDT (16), while in Kwara state, Nigeria, Ae. aegypti was completely sus- ceptible to this insecticide (17). Highest re- sistance to 4% of DDT was observed in pop- ulations of Ae. aegypti from northern Nigeria, while the knockdown rate in dieldrin was very high (18). Two population types of Ae. ae- gypti (white and brown populations) examined in Cote D’ivoire, both populations were re- sistant to DDT (19, 20). Nine populations test- ed of Ae. aegypti from Senegal were resistant to DDT (21) (Fig. 1). In Asia In Thailand, resistance, or increased toler- ance to dieldrin has been reported, and there is resistance to DDT, although DDT is not used in Aedes control programs. DDT has been used for agriculture in Thailand since 1934 was banned in agriculture in 1983. In health and for indoor spraying, it has been used against malaria vectors since 1949, and since 2000, its use in health has been ceased due to its de- structive effects on the environment and the development of physiological resistance in oth- er mosquitoes (22–27). In Malaysia, Ae. ae- gypti and Ae. albopictus showed resistance to DDT. Aedes aegypti was susceptible to diel- drin except in one area. Dieldrin resistance was reported in Ae. albopictus except in Kuala Lum- pur and two other areas. The higher resistance of Ae. albopictus to dieldrin was due to the ecology of this species and its breeding sites, which was close to plants and in agricultural areas, and therefore exposed to dieldrin be- cause it was still used in agriculture to control soil pests (28, 29). The adult Ae. aegypti from Jharia, Bihar were the first case of DDT re- sistance in India (30). In India DDT resistance in field collected Ae. aegypti has reported from http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 3 http://jad.tums.ac.ir Published Online: March 31, 2023 Goa (31) and Jharkhand (32). DDT and diel- drin resistance have been reported in all Indi- an Ae. aegypti. Aedes albopictus from western Bengal has shown resistance to DDT. Aedes aegypti females’ exposure with 0.4% dieldrin resulted in 48% and 100% mortality, while 4% dieldrin for same exposure period showed 88% and 100% mortality (33–38). In China all Ae. albopictus adults tested populations showed resistance to DDT. Urban Ae. albopictus had a higher level of resistance to DDT than strains collected from rural areas (39–42). In Laos Ae. albopictus was resistant to DDT (43). There was high DDT resistance in adult of Ae. aegypti in Myanmar (44). In Philippines Ae. aegypti and Ae. albopictus larvae were sensitive to lindane, dieldrin, and DDT (45). Also, there was DDT resistance in Vietnam (46–48). In Japan all strains of Ae. albopictus were highly resistant to DDT (49). In Phnom Penh, Cambodia pa- pers impregnated with DDT at 4% and diel- drin at 0.4 and 4% were used for the bioas- says of adult Ae. aegypti. The mortality rates were reported 0% for 4% DDT and 87.6± 2.1% and 97.8±4.3, for 0.4% and 4% diel- drin%, respectively (50) (Fig. 1). In America The development of resistance to DDT in Mexico and many parts of the Americas was due to the widespread use of this insecticide during the 1950s and 1960s (51). In the Amer- icas Ae. aegypti resistance to DDT was initial- ly found in Trinidad in 1955, and it was sus- ceptible to dieldrin. Now, all parts of the Car- ibbean have Ae. aegypti populations that are resistant to DDT and dieldrin. From French Gui- ana to the Bahamas, the situation is uniform re- garding this resistance (52). DDT resistance has been confirmed in Ae. aegypti Caribbean pop- ulations (53) and in two Florida populations and one New Jersey population (54). There were 45 dengue epidemics in Cuba in 1977, 1981, 1997 and 2002 and Ae. aegypti resistance to DDT has been seen in this country (55, 56). In Colombia resistance to DDT was seen in all mosquito populations tested, although this in- secticide did not use for vector control in any of the study areas (57–59). In Venezuela low mortality in larvae Ae. aegypti was exhibited by field strains toward DDT and confirmed the presence of high resistance to this insecticide (60). After exposure of adults of Ae. aegypti from Santo Domingo, Dominican Republic to discriminating concentrations of DDT, it was found that wild populations were resistant to this insecticide (61). In Brazil eradication pro- grams from 1950 to 1970 might have led to spread of DDT resistance (62). All of popula- tions of Ae. aegypti from Peru were resistant to DDT (4%), after 1 hour of exposure with papers treated (63) (Fig. 1). In Australia and Oceania Adult bioassays detected significant re- sistance to dieldrin and DDT in Ae. aegypti population from Townsville City, despite DDT has not been used for adult control in this city for minimum 12 years, and dieldrin is not an adulticide (64). Susceptibility to DDT of Ae. aegypti and Ae. albopictus populations seems to have decreased in Papua New Guinea (65) (Fig. 1). Resistant status of Aedes aegypti and Ae. albopictus to Organophosphates insecticides In Africa Complete susceptibility to temephos, fenitro- thion and malathion has been reported from both vectors in Cameroun, Gabon (12, 13), and Ma- yotte (66). In Central African Republic larvae of both species were sensitive to temephos and fenitrothion and only one population of resistant to fenitrothion was reported (15). In Cape Verde Ae. aegypti was sensitive to fenitrothion (11), but it shows resistant to temephos and sensi- tivity to malathion (67). In Dakar, Senegal al- so its sensibility to fenitrothion was reported (11). Organophosphates (0.8% and 5% mala- thion, 0.05% pirimiphos-methyl and 1% fenitro- thion) were used for bioassays of Ae. aegypti http://jad.tums.ac.ir/ https://www.google.com/search?sa=X&bih=588&biw=1213&hl=en&q=Oceania&stick=H4sIAAAAAAAAAONgVuLQz9U3MM0rSn_EaMwt8PLHPWEprUlrTl5jVOHiCs7IL3fNK8ksqRQS42KDsnikuLjgmngWsbL7J6cm5mUmAgBnMPSDTAAAAA J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 4 http://jad.tums.ac.ir Published Online: March 31, 2023 from Senegal. All the Ae. aegypti populations tested were susceptible to 5% malathion, while only two populations showed susceptibility to 0.8% malathion. Resistance in all populations were reported for 0.05% pirimiphos-methyl. Louga, Mbour and Barke´dji strains showed resistance to 1% fenitrothion (21). The Ae. ae- gypti populations in northern Nigeria were sus- ceptible to fenitrothion in 2018 and 2019, with mortality of 98.6% and 100%, respectively. But, in 2020, mortalities of 93% observed and moderate resistance were registered (18) (Fig. 2). In Asia In Thailand Ae. aegypti and Ae. albopictus resistance to temephos was demonstrated in some areas. Aedes aegypti showed susceptibil- ity to malathion. All strains of Ae. aegypti were resistant to temephos except strains of Nakhon Ratchasima. Aedes albopictus was susceptible to both insecticides (68). In other study the lar- vae of both species were resistant to temephos and resistance or tolerance to Malathion and fenitrothion has been reported in different parts of north and south of Thailand, while in cen- tral and eastern parts of Thailand, were still susceptible to fenitrothion (24). Temephos has been used against Ae. aegypti larvae in Thai- land since 1950, and malathion, fenitrothion, and pirimiphos methyl have been used for IRS and fogging (22–24, 26, 68, 69). In Sri Lanka Ae. aegypti and Ae. albopictus were sensitive to malathion. Although this insecticide has been widely used in the control of malaria and fil- ariasis in endemic areas (70). Also, in 2020 re- ported that adult Ae. aegypti mosquitoes from Sri Lanka were susceptible to malathion 5%. For temephos the resistance ratios (RR50) var- ied between 0.69 and 3.93 (71). In one study in Malaysia, Ae. aegypti was sensitive to mal- athion and fenitrothion (29), in 2015 was sen- sitive to this insecticide, except in Kuala Lum- pur and Ae. albopictus was resistant to mala- thion except in Kuala Lumpur. No resistance was observed in larvae to temephos, and vari- ous populations were susceptible or tolerant, although it has been widely used since the 1970s and during the 1998 global pandemic (28). Chen et al. reported greater resistance of Ae. aegypti to temephos than Ae. albopictus (72). In India the susceptibility of two species was reported to malathion in 1993 (73). In other studies, Ae. aegypti adults were tolerant to fenitrothion and sensitive to malathion, also the larvae were sensitive to three tested insec- ticides including temephos, fenitrothion and mal- athion (32–34). Resistance of this species to di- chlorvos was also reported (74). Results from the studies in 2015 showed that only one Ae. aegypti population had high levels of resistance to temephos and the rest also Ae. albopictus were susceptible (35, 74). In 2017 Ae. aegypti was reported sensitive to temephos except one population. In this study, exposure of Ae. ae- gypti with 1% fenitrothion for 30 and 60 minutes caused 40% and 100% mortality, re- spectively (36). In 2018, larvae of Ae. aegypti was susceptible to temephos (37). In a recent study in West Bengal all populations Ae. ae- gypti were reported sensitive to malathion (75). In China, resistance to malathion was clear in Ae. aegypti in Hainan Province but Ae. al- bopictus was susceptible (76). Aedes albopic- tus larvae were resistant to temephos and adult was sensitive to dichlorvos, with 100% mor- tality (41), also adult populations of Ae. al- bopictus in the four districts were all sensitive to malathion, but high resistance to temephos was registered (42). In Cambodia there is lar- val resistance to temephos in endemic areas. In the first study on the sensitivity of two Ae. aegypti populations to temephos, in one popu- lation, the larvae showed some degree of re- sistance to temephos and the other population was sensitive (77). In 2022, moderate Ae. ae- gypti resistance was observed for temephos in Phnom Penh, Cambodia. Also, papers impreg- nated with fenitrothion at 1%, malathion at 0.8% and pirimiphos-methyl at 0.2%, were used for the bioassays of adult Ae. aegypti. Organophos- phate insecticides showed high mortality rates (50). Development of mild resistance of Ae. ae- http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 5 http://jad.tums.ac.ir Published Online: March 31, 2023 gypti to malathion was found in Jakarta, Indo- nesia. South Denpasar mosquitoes had the high- est resistance to malathion, which was signifi- cantly different from the resistance reported in North Denpasar (78–80) and populations of Ae. aegypti in Banjarmasin, Kalimantan, Indonesia were susceptible to malathion 5% (81). In Vien- tiane Laos, based on larval bioassay, the wild population of Ae. aegypti strain was relatively resistant to temphos (82). In other study in Laos all seven population of Ae. albopictus but one was resistant to malathion and three of popu- lations showed resistance to temephos and the rest were tolerant (43). In Philippines Ae. ae- gypti and Ae. albopictus larvae were reported sensitive to malathion (45). In Singapore, both species were reported to be sensitive to pirimi- phos methyl (83). In another study, all Ae. ae- gypti populations were sensitive to pirimiphos methyl, and all populations of Ae. albupictus were resistant to pirimiphos methyl but south- eastern populations were tolerant (84, 85). In another study, Ae. aegypti larvae were report- ed to be sensitive to temephos, although re- sistance was possible (86). Aedes aegypti from Dili, Timor-Leste was sensitive to malathion (87), also in Vietnam was sensitive to mala- thion (46–48). In Bangladesh Ae. aegypti mos- quitoes varied in susceptibility to malathion (50μg/bottle) (88) (Fig. 2). In America High levels of resistance to chlorpyrifos in Ae. albopictus has been found in Alabama and Florida (89). None of the eight popula- tions of US Ae. albopictus larvae were resistant to temephos, also Ae. albopictus resistance to malathion was reported in Florida and New Jersey (54). Levels of malathion and temephos resistance were observed in Ae. aegypti Car- ibbean populations. Guyana, Jamaica, and Su- riname Ae. aegypti populations showed only slight resistance to the larvicide. In most Car- ibbean countries, where malathion has been used sporadically for 20 to 30 years to control adults, there is little resistance to this insecti- cide (90) and low levels of temephos resistance were confirmed in Grand Cayman (53). In Unit- ed States Ae. albopictus from Florida, Califor- nia, three different North Carolina populations and Ae. aegypti from Texas were susceptible to malathion. Conversely, Ae. albopictus from North Carolina and Texas were resistant to the same malathion doses (91). Mortality of Ae. ae- gypti mosquitoes tested in Jamaica with mala- thion, was from 84 to 90% at 30 minutes of ex- posure and 100% with increasing exposure time to 45 minutes (92). In 2019, Ae. aegypti strains from New Mexico were susceptible to the chlorpyrifos (93). In Mexico, the Ae. aegypti populations showed less resistant to chlorpyri- fos than to pyrethroids (94). Resistance to fen- thion, fenitrothion, and temephos, has been found in Cuba (55, 56). On the Island of Mar- tinique (Caribbean), Ae. aegypti populations were resistant to naled (organophosphate) that showed insecticide resistance reduced its ef- fect when applied by ultra-low volume (ULV) thermal fogging (95). There is evident that the organophosphate insecticide resistance (mala- thion, fenthion and temephos) is prevalent in Trinidad and Tobago larval strains of Ae. ae- gypti (96). In Cuba, Venezuela, Costa Rica and Jamaica, Ae. aegypti was sensitive to malathi- on despite its widespread use in vector control programs in these countries (97). Resistance to organophosphates is found throughout Latin America due to its intense use to control lar- vae, including Colombia, Cuba, Martinique, Costa Rica, Havana, Brazil, Bolivia, French Guiana, Argentina, French Polynesia, and the Caribbean. But resistance to temephos is low- er in West Africa. The susceptibility of adults and larvae of Ae. aegypti to insecticides from Santo Domingo, Dominican Republic, was eval- uated. Hatched larvae from eggs collected from ovitraps were resistant to temephos. Adults were resistant to malathion (61). larvae of Ae. aegypti from Cuba and other Latin-American countries were investigated for organophosphate insec- ticide resistance, including malathion, pirimi- phos methyl, temephos, fenthion, fenitrothion http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 6 http://jad.tums.ac.ir Published Online: March 31, 2023 and chlorpyriphos. All the strains showed re- sistance to temephos except Nicaragua strain with moderate resistance to temephos. The high- est resistant ratio (RR) value to temephos was in the Havana city strain followed by Panama, Costa Rica, Peru, Jamaica, and Venezuela. Ae- des aegypti larvae were susceptible to mala- thion in all of the strains, and the same results obtained for fenthion and fenitrothion, but Peru strain showed moderate resistance to fenthion, and the Havana city strain had high resistance to fenthion and moderate to fenitrothion. High resistance to pirimiphos methyl has been re- ported in Panama, Costa Rica and Venezuela; however, Santiago de Cuba, Jamaica and Peru showed moderate resistance to this chemical. The Santiago de Cuba strain had the highest chlorpyriphos, resistant ration of 50 followed by Costa Rica and Jamaica, with Havana city, panama, Nicaragua, and Peru showing suscep- tibility to chlorpyriphos. The Venezuela strain showed moderate resistance to chlorpyriphos (98). Aedes aegypti mosquitoes from Peru were exposed for 1 hour with malathion (5%) and pirimiphos-methyl (0.25%) papers, and for 2 hours with fenitrothion (1%) papers. The Chosi- ca population showed resistance to fenitrothi- on and pirimiphos-methyl and developing re- sistance to malathion. While the Punchana and Piura populations were susceptible to malathion and showed resistance to other evaluated insec- ticides (63). In southern Ecuador Ae. aegypti was resistant to malathion and had a mortality rate below 80% (99). In Brazil from 1999–2000 until 2010–2011, the effectiveness of temeph- os on Ae. aegypti was high, resulting in larval mortality of more than 80% throughout Bra- zil. The frequency of temephos-resistant indi- viduals in the insect population increased stead- ily during each biennial survey, indicating a sig- nificant reduction in the effectiveness of temeph- os. This trend reached high levels (less than 50% mortality) in about half of the country in early 2004–2005 (100). In Colombia mosquitoes were susceptible to malathion. A high frequency of fenitrothion resistance was reported in all Ae. aegypti mosquitoes (58), also in 2019 no evi- dence of malathion resistance was found in Ae. aegypti (59). In Venezuela high mortality was obtained with fenitrothion and fenthion against adults and larvae Ae. aegypti in field strains, which suggested the absence of any apprecia- ble amount of resistance to these insecticides, but resistance to malathion, temephos, chlorpyr- ifos, and pirimiphos-methyl was observed (60). Larvae of Ae. aegypti mosquito from the Gua- deloupe and Saint Martin islands were resistant to temephos and malathion compared with the susceptible Bora Bora strain. For the first time in Guadeloupe, mosquito populations of Gua- deloupe and Saint Martin were resistant (weak- ly) to malathion (101). Aedes aegypti resistance to temephos has been observed in Colombia (102), and the Caribbean (90). Result of re- sistance of Ae. aegypti in Brazil show resistance to at least one of the organophosphates (temeph- os, fenitrothion, malathion) tested in all popu- lations in the states of Rio de Janeiro and Es- pírito Santo (103). A study in the city of Curi- tiba, State of Paraná in Brazil reported that temephos could be used for control larvae of Ae. aegypti (104). In Sao Paulo, Ae. aegypti was resistant to temephos and susceptible to mala- thion. It was suggested that malathion be used in ULV instead of cypermethrin in the dengue fever control program in Sao Peteo, and fenitro- thion be used in residual spraying (105). The larvae of all studied Ae. aegypti populations from different Brazilian regions were resistant to temephos (106, 107). Aedes aegypti larvae from northeastern Brazil showed the highest levels of resistance to temephos, also adults from northeastern showed the lowest levels of susceptibility to malathion (108), and adults from Recife Brazil were susceptible to mala- thion (109). In Tocantins state in Brazil all eval- uated populations of Ae. aegypti were resistant to temephos (110). Resistance to fenitrothion for the four populations of Ae. aegypti distrib- uted along the French Guiana exhibited re- sistance to fenitrothion (111). A high level of resistance to temephos was reported from Ae. http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 7 http://jad.tums.ac.ir Published Online: March 31, 2023 aegypti populations in Martinique (French West Indies) (112). In this study the resistance level of Juazeiro do Norte population was much less (RR=7.2), which is only slightly lower than the 2003 level (RR=10.2), because temephos has not been used for at least seven years to con- trol Aedes and has been replaced by Bacillus thuriginensis (Bti). The increased resistance lev- els to temephos in Crato and Barbalha popula- tions, indicate that the resistance management programs in Juazeiro do Norte should be done in neighboring cities. These results also show that even if temephos was replaced in Juazeiro do Norte, the recovery of susceptibility was slow (113). The results of bioassays with temeph- os, that has been used for decades against Ae. aegypti larvae in Brazil caused all the evalu- ated populations to be resistant to temephos (110) (Fig. 2). In Europe Exposure of Ae. aegypti from Pau´l do Mar (Madeira) to fenitrothion given 100% mor- tality that indicate mosquito susceptibility to this insecticide but Ae. aegypti from Funchal (Madeira) was resistant to malathion and fenitro- thion (114). Aedes albopictus adults from the Swiss-Italian border region were susceptible malathion (109) (Fig. 2). In Australia and Oceania Investigating the state of sensitivity to in- secticides in Ae. aegypti from Townsville showed that mosquitoes were resistant to mal- athion and fenthion and susceptible to other organophosphates, temephos, fenitrothion and chlorpyrifos (64). Aedes aegypti and Ae. al- bopictus were susceptible to malathion in Pa- pua New Guinea (65) (Fig. 2). Resistant status of Aedes aegypti and Ae. al- bopictus to Carbamates insecticides In Africa In Cameroon, resistance of Ae. aegypti to bandiocarb and tolerance of Ae. albopictus to this insecticide were reported (13). In 2021 it was reported this species was susceptible to bendiocarb (14). In Central African Republic both vectors were sensitive to propoxur and only one resistant population to propoxur was reported (15). In Kwara state, Nigeria, mortal- ities from exposure of Ae. aegypti to bendio- carb showed that all the mosquito samples test- ed were resistant (17). In northern Nigeria, Ae. aegypti mosquitoes were resistant to 0.1% propoxur (18). In Cape Verde low sensitivity Ae. aegypti to propoxur was reported and in Dakar, Senegal mortality of Ae. aegypti with propoxur was 87.2% (11). In Cote D’ivoire white and brown populations of Ae. aegypti were resistant to propoxur (19, 20). In Senegal, apart from the Matam populations of Ae. ae- gypti were susceptible to 0.1% bendiocarb, and all other populations were resistant to bendio- carb and propoxur (21) (Fig. 3). In Asia Various levels of tolerance/ resistance to propoxur and resistant to bendiocarb and propoxur in Ae. aegypti has been reported in parts of central and southern in Thailand (24, 26, 69). Aedes aegypti resistance to bendiocarb and propoxur was also reported from Malaysia as well as Ae. albopictus resistance (28, 29). In India, Ae. aegypti females were exposed to 0.1% propoxur for 30 and 60min and mortali- ty after 24h was 96% and 100%, but in other study Ae. aegypti was resistant to propoxur, alt- hough this insecticide has not been used to con- trol vectors in India. Resistance has been due to accidental exposure or to intersection re- sistance (36, 37), also Ae. albopictus from sub- Himalayan areas of West Bengal exhibit se- vere resistance against propoxur (38). In China, Ae. albopictus adult was sensitive to propoxur (41), and resistant to bendiocarb (42). In In- donesia Ae. aegypti was resistant to bendio- carb. Developed resistance, with about 80–90% mortality, was observed against bendiocarb 0.1% (80, 81). Papers impregnated with bendiocarb at 0.1% and propoxur at 0.1% were used for the bioassays of adult Ae. aegypt in Phnom Penh, Cambodia, results showed high mortality rates http://jad.tums.ac.ir/ https://www.google.com/search?sa=X&bih=588&biw=1213&hl=en&q=Oceania&stick=H4sIAAAAAAAAAONgVuLQz9U3MM0rSn_EaMwt8PLHPWEprUlrTl5jVOHiCs7IL3fNK8ksqRQS42KDsnikuLjgmngWsbL7J6cm5mUmAgBnMPSDTAAAAA J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 8 http://jad.tums.ac.ir Published Online: March 31, 2023 with carbamate (50). Complete susceptibility to bendiocarb was reported in populations of Ae. aegypti from Bangladesh (88) (Fig. 3). In America In the states of Alabama and Florida, propoxur cross-resistance levels were evalu- ated in Ae. albopictus. Propoxur showed the least toxicity to Ae. albopictus. The results showed that propoxur tolerance levels were absent or very low in all strains (89). In New Jersey, Penn- sylvania, and Florida Ae. albopictus was sensi- tive to propoxur (54). In Colombia all of Ae. aegypti field-collected strains were suscepti- ble to propoxur (59). In Venezuela Ae. aegypti resistance to propoxur was reported (60). Ae- des aegypti adults from Santo Domingo, Do- minican Republic, were resistant to propoxur (61) (Fig. 3). In Europe Aedes aegypti from two localities in Ma- deira (Funchal and Pau´l do Mar) was resistant to bendiocarb (114). After 1h exposure and 24 h holding period to bendiocarb, Ae. albopictus field populations from the Swiss-Italian border region were susceptible (109) (Fig. 3). In Australia and Oceania In Australian 1995 strain of Ae. aegypti was significantly less susceptible to bendiocarb than the 1989 strain, and both strains showed high resistance to propoxur (64). Aedes aegypti from the Madang population, Papua New Guinea was susceptible to bendiocarb and Ae. albopictus also were found to be susceptible to bendio- carb (65) (Fig. 3). Resistant status of Aedes aegypti and Ae. al- bopictus to Pyrethroids insecticides In Africa Both the Aedes species are present in Cen- tral Africa, but Ae. albopictus is more influen- tial and was first reported in 2009. A study in this area found that both vectors were resistant to deltamethrin, and their mortality was less than 90% (15). In Ghana, Ae. aegypti was re- sistant to deltamethrin, and lambda-cyhalothrin but was sensitive to permethrin (9). In other study some populations were resistant to per- methrin and some tolerant (10). In Mayotte both species had highly sensitive to deltamethrin. No resistance to deltamethrin has been record- ed on the island, although deltamethrin has been used on the island since 1984 (66). In Came- roun complete susceptibility of both species was reported to deltamethrin except in Yaounde, where Ae. albopictus populations showed a mor- tality rate of about 80% (12), but in 2017 both species were resistant to deltamethrin and de- creased sensitivity to permethrin was observed (13). In 2021 Ae. aegypti and Ae. albopictus mosquitoes from Cameroun were exposed to 0.05%, and 0.75% permethrin. Aedes albopic- tus populations showed full susceptibility to permethrin and deltamethrin insecticides, but Ae. aegypti was resistant to these insecticides (14). In a study conducted in agricultural and non-agricultural areas of Lagos State, Nigeria, all mosquitoes from both sites were suscepti- ble to permethrin, but about deltamethrin the most populations in non-agricultural areas were resistant and mosquitoes collected from agri- cultural areas were reported sensitive (16). In another study, Ae. aegypti was completely sen- sitive to permethrin (17). Adult bioassays of Ae. aegypti in northern Nigeria were conduct- ed by pyrethroids: 0.25% and 0.75% perme- thrin, 0.03% and 0.05% of deltamethrin, Lamb- da-cyhalothrin, and α-cypermethrin, and 0.15% cyfluthrin. Populations were susceptible to cyfluthrin but highly resistant to other type I and II pyrethroids (18). In Dakar, Senegal, Ae. aegypti populations was susceptible permethrin. About deltamethrin mortality was 94.5% and for lambda-cyhalothrin 81.6%. In Cape Verde Ae. aegypti was sensitive to deltamethrin, per- methrin, and lambda-cyhalothrin (11). In an- other study, Ae. aegypti was resistant to del- tamethrin, and cypermethrin (67). Bioassays of pyrethroids (0.25% and 0.75% permethrin), http://jad.tums.ac.ir/ https://www.google.com/search?sa=X&bih=588&biw=1213&hl=en&q=Oceania&stick=H4sIAAAAAAAAAONgVuLQz9U3MM0rSn_EaMwt8PLHPWEprUlrTl5jVOHiCs7IL3fNK8ksqRQS42KDsnikuLjgmngWsbL7J6cm5mUmAgBnMPSDTAAAAA J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 9 http://jad.tums.ac.ir Published Online: March 31, 2023 0.05% deltamethrin, 0.03% and 0.5% lambda- cyhalothrin, 0.05% alpha-cypermethrin), by standard WHO test kits for adults of Ae. ae- gypti from Senegal showed that susceptibility to 0.75% permethrin was observed in all pop- ulations except in three, and one population had suspected resistance to 0.25% permethrin. On- ly populations of the southern regions were susceptible to type II pyrethroids (21). In Dar es Salaam, Tanzania all sites showed lower sus- ceptibility to deltamethrin, permethrin and re- sistance to lambda-cyhalothrin (115). In Cote D’ivoire white populations of Ae. aegypti were resistant to deltamethrin and brown populations were susceptible (19, 20) (Fig. 4). In Asia In Japan Ae. albopictus in Nagasaki showed high tolerance to pyrethroids. The cause for this may be due to the widespread use of DDT in the 1950s (116). Most studies on pyrethroid re- sistance in Ae. aegypti in Southeast Asia are from Thailand. The first outbreak of dengue fe- ver in Thailand was in 1958 (117). Since 1950, carbamates, organophosphates, and organochlo- rines have been used for control. Synthetic py- rethroids have been used since 1992, low price of pyrethroids, their quick knockdown effect and relative safety for humans due to low tox- icity to mammals has increased their use (22). Increased tolerance or resistance to pyrethroids has been reported from Ae. aegypti and Ae. al- bopictus in Thailand, such as deltamethrin and permethrin, which are widespread throughout Thailand, resistance to cypermethrin in parts of northern and western Thailand, and resistance to bioallethrin, bioresmethrin, and alpha-cyper- methrin in larvae and adults. Mortality of Ae. aegypti with lambda cyhalothrin was 100% and had tolerated to mosquito coils. Aedes aegypti has shown behavioral resistance and avoidance to alpha-cypermethrin, deltamethrin, permethrin, tetramethrin, cyphenothrin in Thailand (26, 68, 118–120). In Malaysia, many studies have been conducted on the resistance of the two species to insecticides. The first study on pyrethroid resistance was in 2001. The Ae. aegypti urban strain had the highest resistance to permethrin, cyphenothrin, lambda-cyhalothrin, and deltame- thrin. Aedes albopictus was sensitive to perme- thrin and deltamethrin (121–123). The first study on mosquito coils against Ae. aegypti was in 1996, that mortality was minimal (124). In 2017, in a study was reported that mosquito coils had low insecticidal activity against Ae. aegypti and therefore may have little protection for mos- quito bites (125). In Malaysia was reported Ae. aegypti resistance to permethrin (29). In the oth- er study across Malaysia, Ae. aegypti was re- sistant to permethrin and deltamethrin (28). In India, Ae. aegypti and Ae. albopictus resistance to pyrethroids such as permethrin has been re- ported in Delhi and Kerala (73). In 2001 Ae. aegypti adults was sensitive to deltamethrin, permethrin, and lambda cyhalothrin in Delhi (33). In Another study a total of five different pyrethroid compounds were tested against Ae. aegypti females with exposure period of 5 and 15min in Madurai West city in Tamil Nadu State, they were found invariably susceptible to all tested pyrethroid compounds. Based on the results, the efficacy in descending order of pyrethroids was cyfluthrin > permethrin > del- tamethrin > lambda cyhalothrin and > etofenprox (36). In recent studies in West Bengal, all pop- ulations Ae. aegypti except one site were sen- sitive to deltamethrin and alpha-cypermethrin (75), and two Ae. albopictus populations, from sub-Himalayan districts were severely resistant to permethrin (38). In China, Ae. albopictus is resistant to deltamethrin, permethrin, cyperme- thrin, and beta-cypermethrin, and resistance to pyrethroids is greater than resistance to other common insecticide groups (40–42). In Cam- bodia, there is Ae. aegypti adult resistance to the two main insecticides including permethrin (nets) and deltamethrin (fumigation) in all re- gions (126). The pyrethroids alpha-cyperme- thrin at 0.03%, bifenthrin at 0.2%, cyfluthrin at 0.15%, deltamethrin at 0.03%, etofenprox at 0.5%, lambda-cyhalothrin at 0.03% and perme- thrin at 0.25% were used for the bioassays of http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 10 http://jad.tums.ac.ir Published Online: March 31, 2023 adult Ae. aegypti. The adult mortalities were 0 % for etofenprox, 1±1.9% for permethrin, 3.1± 4.1% for bifenthrin, 4.2±8.3% for lambda- cyhalothrin, 10.2±7.1% for deltamethrin, 11± 4.5% for alpha-cypermethrin and 35±11% for cyfluthrin and there were low mortality rates with all the tested pyrethroid insecticides (50). In Indonesia, Ae. aegypti larvae were sensi- tive to pyrethroids (78, 127). Aedes aegypti adult resistance was reported to more than one pyrethroids of cypermethrin, permethrin and D-allethrin (78). In other study, adult Ae. ae- gypti resistance to permethrin and deltame- thrin and high levels of resistance to alpha- cypermethrin and lambda cyhalothrin has been reported (128). In 2017, all Ae. aegypti adults tested in Jakarta were resistant to deltamethrin, permethrin, and lambda cyhalothrin, with a mor- tality rate of less than 90%. Resistance to per- methrin was higher than the others (79). Also, resistance of Ae. aegypti to deltamethrin, permethrin, lambda cyhalothrin, cyfluthrin in Denpasar, Bali has been reported (80). In Ban- jarmasin, Kalimantan, Indonesia, all Ae. ae- gypti populations showed different degree of resistance to 0.15% cyfluthrin 0.05% deltame- thrin, 0.05% Lambda-cyhalothrin, and 0.75% permethrin, with mortalities less than 90% (81). In Laos high level of resistance against per- methrin, and high susceptibility to deltamethrin was reported in Ae. aegypti (129). In the other study seven populations of Ae. albopictus were sensitive to permethrin and deltamethrin (43). In Myanmar high pyrethroid resistance (perme- thrin and deltamethrin) has been reported in the larvae and adult of Ae. aegypti (44). In Sin- gapore, Ae. aegypti is the primary vector of Dengue Virus. The first case was reported in the 1960s (130). Aedes aegypti resistance to per- methrin, cypermethrin, and deltamethrin has been demonstrated in Singapore (83-85). In other two studies, Ae. aegypti larvae were resistant to permethrin and etofenprox (86), and Ae. ae- gypti adult with PermaNet® had less than 80 % mortality (131). Aedes aegypti from Dili, Timor-Leste, has become resistant to perme- thrin, lambda-cyhalothrin and resmethrin (87). In the first insecticide resistance investigation of Ae. aegypti in several locations of Central Highlands and Nam Bo in Vietnam, the mos- quitoes were susceptible to pyrethroids such as deltamethrin in the central and northern regions. In the southern and central mountainous regions Ae. aegypti was resistant to deltamethrin, per- methrin, lambda-cyhalothrin and alpha-cyper- methrin. This is due to the long use of pyre- throids in the control of malaria and dengue fever and its use in agriculture, especially in the southern and central regions (46). In the next study, Ae. aegypti's sensitivity to permethrin, lambda-cyhalothrin, deltamethrin, and alpha- cypermethrin was reported in many parts of the north and center but was resistant to these insecticides in southern and central Vietnam (47). In 2009, it was reported that, Ae. aegypti sensitivity to D-allethrin has decreased. Mos- quitoes were sensitive to pyrethroids in north- ern and central Vietnam but were resistant in southern and central Vietnam (132). In 2016 and 2003 also, Ae. aegypti resistance was re- ported to permethrin and lambda-cyhalothrin (48, 133). Also, populations of Ae. albopictus tested from Vietnam exhibited high-level re- sistance to permethrin (134). In 2010, study of pyrethroid susceptibility of larvae Ae. albopic- tus collected from Nagasaki City, Japan, indi- cated populations of Ae. albopictus tolerant to pyrethroids spread widely in Nagasaki (116). In other study in this city insecticide suscepti- bility tests were performed on Ae. albopictus adults and larvae of F1 colonies collected from Nagasaki City, Japan. The results were com- pared with those of several such colonies col- lected from other locations in Japan. The lar- vae collected from Nagasaki City, also from other locations in Japan were resistant to d- T80-allethrin and more than half of the adults of the Nagasaki and Fukuoka colonies showed resistance to permethrin (135). Susceptibility of adult Ae. aegypti mosquitoes from Sri Lanka to deltamethrin 0.05%, permethrin 0.75% were evaluated by the standard WHO mosquito bi- http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 11 http://jad.tums.ac.ir Published Online: March 31, 2023 oassay protocol. Resalts revealed resistance in all Ae. aegypti populations for both permethrin (10–89%) and deltamethrin (40–92%) (71). In Bangladesh Ae. aegypti mosquitoes were ex- posed to diagnostic dose of 15μg/ bottle for permethrin, and 10μg/bottle for deltamethrin. High levels of resistance to permethrin were reported in Ae. aegypti, while sensitivity to del- tamethrin was different between populations (88) (Fig. 4). In America High resistance to pyrethroids were observed in Ae. aegypti. All Grand Cayman Ae. aegypti populations survived after one hour of expo- sure to pyrethroid-impregnated papers (53). In the USA adult mortality after a 24h exposure to the pyrethroid insecticides (deltamethrin, prallethrin, and phenothrin) at discriminating doses showed that, all the field populations test- ed were susceptible (99–100% mortality) (54). In Florida, California, North Carolina, and Texas, vectors were resistant to low doses of etofenprox. Ae. aegypti was also resistant to high doses in Texas. Six populations of Ae. al- bopictus showed susceptibility to bifenthrin and two populations Ae. aegypti and Ae. albopic- tus were resistant and possible resistance was shown in one Ae. albopictus population. Seven Aedes spp. populations were susceptible (100%) to permethrin and one population was resistant. Possible resistance was shown in one Ae. al- bopictus population. All tested Ae. albopictus were susceptible (100%) to phenothrin and one population of Ae. aegypti was resistant. Six Ae. albopictus populations were susceptible to the low dose of deltamethrin. At the high deltame- thrin dose, seven Ae. albopictus populations were susceptible and two Ae. albopictus and Ae. aegypti populations showed possibility of resistance (91). Aedes aegypti from New Mex- ico showed high pyrethroid resistance and kdr mutation F1534C in the para gene (93). In Tex- as/Mexico border cities all populations of Ae. aegypti showed resistance to permethrin, del- tamethrin and sumithrin, although none of these insecticides are commonly used for vector con- trol activities in this region (136). All Ae. ae- gypti mosquitoes tested in Jamaica were re- sistant to permethrin (92). Laboratory bioassay on the Island of Martinique (Caribbean) showed that Ae. aegypti populations were strongly re- sistant to pyrethrins and deltamethrin therefore insecticide resistance decreased the efficacy of space sprays for adult mosquito and dengue con- trol (95, 137). There is evidence of resistance to permethrin in populations in Baja California North, South and in Quintana Roo, south of Mexico and some states of northeast Mexico due to use for more than 10 years in Mexico for control of Ae. aegypti (138-140). The results obtained from Veracruz State Mexico showed that the field strains of Ae. aegypti were resistant to most of the pyrethroids analyzed including cypermethrin, deltamethrin, δ-phenothrin, α-cy- permethrin, d-phenothrin, z-cypermethrin, λ- cyhalothrin, bifenthrin, and permethrin and sug- gested that populations in the state of Veracruz were under the strong selection pressure, caus- ing from the continuous use of permethrin for more than a decade (141). In Cuba, Ae. aegypti resistance was reported to deltamethrin, lamb- da cyhalothrin, beta-cypermethrin, cypermethrin, and cyfluthrin (55, 56). Aedes aegypti from Ja- maica was resistant to permethrin with types of the mode of mechanism (92). One study on mosquitoes collected from eight Latin Ameri- can countries showed that larvae from the Ha- vana city strain were resistant to all the pyre- throids, the highest RR50 was to deltamethrin, as well as lambda cyhalothrin, betacypermethrin and cyfluthrin showed high resistance. Larvae from Santiago De Cuba, Peru and Venezuela showed high deltamethrin RR50 values as well as Havana City (98). Populations of Ae. aegypti from Peru were exposed for 1h to papers treated with alpha-cypermethrin (0.05%), cypermethrin (0.05%), deltamethrin (0.05%), etofenprox (0.5 %), lambda-cyhalothrin (0.05%), and permethrin (0.75%). Results showed Ae. aegypti from Chosi- ca was resistant to alpha-cypermethrin, cyper- methrin, etofenprox, also developing resistance http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 12 http://jad.tums.ac.ir Published Online: March 31, 2023 to lambda-cyhalothrin, and susceptibility to del- tamethrin were reported. But the Punchana and Piura populations were resistant to all insecti- cides (63). In Tocantins state in Brazil the da- ta showed different resistance to deltamethrin among the samples of Ae. aegypti (110). In southern Ecuador Ae. aegypti was resistant to deltamethrin and alpha-cypermethrin (99). In study of insecticide resistance status of Ae. ae- gypti from Colombia, susceptibility to deltame- thrin and cyfluthrin was reported in all mos- quito populations studied. Six populations were resistant to lambda-cyhalothrin. Four field pop- ulations showed resistance to permethrin, two populations were also resistant to lambda- cyhalothrin. Pyrethroid resistance was not found in populations from the state of Antioquia. Re- sistance to etofenprox was reported in all mos- quito populations, however, there is no record of using this insecticide for dengue control pro- grams at any of the study sites (58). In other study the most of field populations were re- sistant to permethrin. The resistance to deltame- thrin was observed only in one site and lamb- da cyhalothrin resistance in three sites (59). The lambda cyhalothrin resistance has been com- monly detected in Colombia (142). In Venezue- la Ae. aegypti moderate levels of resistance to permethrin and lambda cyhalothrin was report- ed (60). Aedes aegypti in different Brazilian re- gions were resistant to deltamethrin, cyperme- thrin, permethrin (104–108, 113, 143). In Bra- zil Ae. albopictus adult mortality rates with per- methrin and λ-cyhalothrin were close to 100% (109). Evaluation of insecticide resistance in Ae. aegypti populations the case of Tocantins state in Brazil showed different state of resistance to deltamethrin among the samples (110). In one study, severe resistance to deltamethrin with weak knockdown effect and low mortality was observed in French Guiana Ae. aegypti popu- lations (111). Aedes aegypti adults from Santo Domingo, Dominican Republic, were reported resistant to permethrin and deltamethrin (61). Tests done with adult mosquitoes of Ae. aegypti reported moderate resistance status to deltame- thrin from Guadeloupe and Saint Martin islands (101) (Fig. 4). In Europe In Italy resistance to α-cypermethrin and per- methrin in adult Ae. albopictus has been report- ed. Resistance to permethrin was seen from Fer- rara Province in Emilia-Romagna and from Bari Province in Puglia while the field-population from Athens (Greece) showed possibility of re- sistance. Resistance to α-cypermethrin was re- ported for the mosquitoes in Ferrara Province Venezia Province and Rome but four Italian pop- ulations were susceptible. Eight Italian popula- tions, and the Albanian one, showed suscepti- bility to deltamethrin but the Greek laboratory colony was resistant (144). In the Swiss-Ital- ian border region Ae. albopictus adult mortali- ty rates with permethrin and λ-cyhalothrin were close to 100% (109). In Spain, 5 populations of Ae. albopictus were studied, four populations were sensitive to deltamethrin and permethrin. In the case of cypermethrin, two populations of Ae. albopictus were sensitive to this toxin and two populations were tolerant and one popu- lation was reported to be resistant (145). Ae- des aegypti from Funchal was reported to be resistant to cyfluthrin and permethrin. Resistance to pyrethroids was also seen in the Pau´l do Mar population. Pyrethroids mortality rates were be- tween 2% (permethrin), and 88% (cyfluthrin) (114). Populations of Ae. albopictus tested from Italy exhibited high-level resistance to perme- thrin (134) (Fig. 4). In Australia and Oceania Results a study in 1999 showed that there was no evidence of resistance to synthetic py- rethroids (deltamethrin and permethrin) (64). Ex- periments on Queensland mosquitoes with py- rethroids (bifenthrin, deltamethrin and lamb- da-cyhalothrin) did not show Ae. aegypti re- sistant, and bioassays showed only weak toler- ance. There was no evidence of kdr mutations in Queensland Ae. aegypti. In Cairns South, a low resistance to lambda-cyhalothrin was re- http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 13 http://jad.tums.ac.ir Published Online: March 31, 2023 ported in a sample of Ae. aegypti at 24h after exposure. Obvious resistance to bifenthrin has not been detected in field populations of Ae. ae- gypti from northern Queensland (146). Study of resistance of Ae. aegypti and Ae. albopictus in Papua New Guinea clearly show high pyrethroid resistance: deltamethrin and lambda cyhalothrin in both Madang and Port Moresby Ae. aegypti populations. The Ae. aegypti population in Port Moresby show a higher level of deltamethrin resistance than Madang where susceptibility to lambda-cyhalothrin has been greatly reduced (65) (Fig. 4). Resistant status of Aedes aegypti and Ae. al- bopictus to Insect growth regulator (IGRs) and microbial insecticides In Africa In Cameroon, Gabon (12) and Central Af- rican Republic (15) larvae of both species were susceptible to Bacillus thuringiensis israelensis (Bti). In Mayotte complete susceptibility to Bti, and IGRs including spinosad, diflubenzuron, pyriproxyfen, and methoprene was reported (66). In Cape Verde also was observed sensitivity to diflubenzuron and Bti (67) (Fig. 5). In Asia In China, Ae. albopictus larvae from all studied areas were sensitive to Bti and hex- afluoromoron, but in one population showed high resistance to pyriproxyfen and moderate resistance to pyriproxyfen (42). In Singapore, larval mortality due to Bti was 100% (86). In Malaysia susceptibility of field Ae. aegypti lar- vae to Bti was reported (147), but Ae. aegypti and Ae. albopictus populations from 12 states in Malaysia showed a different state of resistance to five IGRs (pyriproxyfen, methoprene, cyrom- azine, novaluron and diflubenzuron). Aedes al- bopictus was susceptible against cyromazine, pyriproxyfen, and methoprene. Field-collect- ed Ae. aegypti showed high susceptibility to diflubenzuron, cyromazine, and novaluron (148). Susceptibility of a wild larval population of Ae. aegypti against Bti, diflubenzuron, pyriproxyfen and spinosad was tested in Vientiane Laos. With bioassays of larvae was found that the wild Ae. aegypti strain was moderately resistant to spinosad and susceptible to the other insecti- cides. Based on field bioassays, all tested in- secticides remained above the WHO accepta- ble larvicidal threshold after 28 weeks (82). In Cambodia for larvae, moderate Ae. aegypti re- sistance was reported for spinosad (RR90 < 5.6) but, there was no resistance against Bti (RR90 < 1.6) (50) (Fig. 5). In America The susceptibility Ae. albopitus to imidaclo- prid, spinosad, and Bti was investigated in Al- abama and Florida. These results reported that mosquitoes collected from field were much more susceptible to spinosad than to the other insecticides tested in the study (89). Of the eight USA populations of Ae. albopictus, none were resistant to the larvicides tested (Bti, spinosad, methoprene, and pyriproxyfen) but reduced susceptibility Ae. albopictus to the IGRs pyriproxyfen and methoprene was observed in Florida and New Jersey (54). The larvae of all studied Ae. aegypti populations from different Brazilian regions were susceptible to difluben- zuron and Bti (106, 107, 109). A tolerance to IGRs and full susceptibility to spinosad and Bti was reported in Martinique. In experiments pyriproxyfen and Bti had 28- and 37-weeks ac- tivities in permanent breeding containers, where- as under field conditions they could not con- trol Ae. Aegypti populations after four weeks. But diflubenzuron and spinosad were effective for 16 weeks, therefore these chemicals can be alternatives to Bti and temephos to control re- sistant Ae. aegypti (112, 149). In French Guiana, ULV spraying with BT was caused 100% mor- tality in Ae. aegypti populations (111) (Fig. 5). In Europe The Ae. albopictus populations from the Canton of Ticino in southern Switzerland, and the Como area in northern Italy were all sus- http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 14 http://jad.tums.ac.ir Published Online: March 31, 2023 ceptible to Bti. The efficacy of Lysinibacillus sphaericus, another entomopathogenic bacte- rium, was tested against Ae. albopictus and showed good activity. The activity a mixture of Bti and L. sphaericus crystals “Vectomax®”, was also studied to determine if these insecti- cidal components are effective for controlling Ae. albopictus. In this study Ae. albopictus was also susceptible to diflubenzuron (109) (Fig. 5). Mechanisms of insecticide resistance in Ae- des aegypti and Ae. albopictus The mode of action of insecticides is mostly on the nervous system of insects. Target-site mu- tations and metabolic resistance are two main resistance mechanisms in Ae. aegypti and Ae. albopictus. In the mutation at the target site, which is one of the worst resistance mecha- nisms in insects, the insect makes the insecti- cides ineffective by changing the structure of the targets of the insecticides. Pyrethroids and DDT act on voltage-gated sodium channels, and the mutation in the amino acid sequence of this gene causes the sensitivity of the channels to prevent the binding of pyrethroids and DDT to decrease. The use of pyrethroids with cross-re- sistance mechanisms (especially kdr mutations) existing in DDT caused to the rapid rise of py- rethroid resistance. Also, resistance to acetyl- cholinesterase (AChE) is current in Aedes and is related to organophosphate and carbamate in- secticides. Mutations in the gene coding for the neurotransmitter acetylcholinesterase number one (ace-1) decrease the inhibition effect of the insecticide on the enzyme. Metabolic resistance is very current in mosquitoes and has been ob- served against all insecticides used for vector control. Mosquitoes have enzyme systems that protect them from foreign compounds, and some of these enzymes can break down insecticides before they reach their site of action. In meta- bolic resistance, the enzymes that detoxify the insecticide are overexpressed or the enzyme's affinity for the insecticide is altered by amino acid substitution. Three of the most important of these enzymes are cytochrome P450 monoox- ygenases, glutathione S-transferases (GSTs) and esterases (62, 150, 151). Bti bacterial larvicide due to the presence of an intact endotoxin com- plex and synergy among individual toxins, es- pecially the presence of Cyt1A, it has the low- est risk of developing resistance. Bti plays an important role in reducing the development of resistance and maintaining sensitivity in other biolarvicids. Binary toxins from Bacillus sphaer- icus have many advantages in controlling mos- quito larvae. Laboratory data and field studies worldwide showed that a combination of Bti with B. sphaericus, the best way to increase larvicidal activity is to prevent resistance and restore susceptibility to B. sphaericus. In the case of IGR insecticides, the risk of develop- ing resistance is low. Late 4th instar larvae and pupae with lower internal juvenile hormone ti- ters are more susceptible to juvenile hormone analogs such as methoprene and pyriproxyfen in the transfer from late 4th instar larvae to pu- pae and adults. In aquatic habitats, different ag- es of mosquito larvae coexist, with very differ- ent levels of internal rejuvenation hormones. This phenomenon leads to sublethal exposures, tolerance and even the development of resistance (152). http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 15 http://jad.tums.ac.ir Published Online: March 31, 2023 Fig. 1. World mapping of organochlorine insecticide resistance in Aedes aegypti and Aedes albopictus up to December 2022 Fig. 2. World mapping of organophosphates insecticide resistance in Aedes aegypti and Aedes albopictus up to Decem- ber 2022 http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 16 http://jad.tums.ac.ir Published Online: March 31, 2023 Fig. 3. World mapping of carbamates insecticide resistance in Aedes aegypti and Aedes albopictus up to December 2022 Fig. 4. World mapping of pyrethroids insecticide resistance in Aedes aegypti and Aedes albopictus up to December 2022 http://jad.tums.ac.ir/ J Arthropod-Borne Dis, March 2023, 17(1): 1–27 TS Asgarian et al.: Worldwide Status of … 17 http://jad.tums.ac.ir Published Online: March 31, 2023 Fig. 5. World mapping of Insect growth regulator (IGRs) and microbial insecticides insecticide resistance in Aedes aegypti and Aedes albopictus up to December 2022 Conclusions WHO recommended bendiocarb, propoxur, DDT, fenitrothion, malathion, pirimiphos-me- thyl, á-cypermethrin, bifenthrin, cyfluthrin, del- tamethrin, etofenprox, lambda cyhalothrin has been recommend as indoor residual spraying for mosquito control. Alpha-cypermethrin, cyfluth- rin, deltamethrin, deltamethrin, etofenprox, lambda cyhalothrin, permethrin are being used for impregnation of bednets. Fenitrothion, mal- athion, pirimiphos-methyl, bioresmethrin, cyfluthrin, cypermethrin, cyphenothrin, d,d- trans-cyphenothrin, deltamethrin, d-phenothrin, etofenprox, lambda cyhalothrin, permethrin, resmethrin for thermal fog and space spraying. Fuel oil, B. thurigiensis, chlorpyrifos, difluben- zuron, fenthion, methoprene, novaluron, pyriproxyfen, pirimiphos-methyl, and temephos are recommended as larvicides. Because of importance of Ae. aegypti and Ae. albopictus in transmission mosquito-borne arboviruses, resistance management programs and strategies should be further considered in worldwide to prevention from insecticide re- sistance in arbovirus vectors. Acknowledgements This research is supported by Ministry of Health and Medical Education under code num- ber of NIMAD 982984. Conflict of interest statement The authors declare there is no conflict of interests. References 1. Vega-Rúa A, Lourenço-de-Oliveira R, Mous- son L, Vazeille M, Fuchs S, Yébakima A (2015) Chikungunya virus transmission potential by local Aedes mosquitoes in the Americas and Europe. PLoS Negl Trop Dis. 9(5): 1–5. 2. Ferreira-de-Brito A, Ribeiro IP, Miranda RMd, Fernandes RS, Campos SS, Silva KABd (2016) First detection of natural infection of Aedes aegypti with Zika virus in Brazil and throughout South America. 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