Developing effective and eco-friendly antiparasitic drugs and insecticides is an issue of high importance nowadays. In this study, we evaluated the anthelminthic and insecticidal potential of the leaf essential oil obtained from Origanum syriacum against the L3 larvae of the parasitic nematode Anisakis simplex and larvae and adults of the mosquito Culex quinquefasciatus. Tests on A. simplex were performed by standard larvicidal and penetration assays, while mosquito toxicity was assessed relying on larvicidal, tarsal contact, and fumigation tests. To shed light on the possible mode of action, we analyzed the oil impact as acetylcholinesterase (AChE) inhibitor. This oil was particularly active on L3 larvae of A. simplex, showing a LC50 of 0.087 and 0.067 mg mL-1 after 24 and 48 h treatment, respectively. O. syriacum essential oil was highly effective on both larvae and adults of C. quinquefasciatus, showing LC50 values of 32.4 mg L-1 and 28.1 µg cm-2, respectively. Its main constituent, carvacrol, achieved larvicidal LC50(90) of 29.5 and 39.2 mg L-1, while contact toxicity assays on adults had an LC50(90) of 25.5 and 35.8 µg cm-2, respectively. In fumigation assays, the LC50 was 12.1 µL L-1 after 1 h and decreased to 1.3 µL L-1 in 24 h of exposure. Similarly, the fumigation LC50 of carvacrol was 8.2 µL L-1 after 1 h of exposure, strongly decreasing to 0.8 µL L-1 after 24 h of exposure. These results support the folk usage of Lebanese oregano as an antiparasitic agent, providing new insights about its utilization for developing new effective and eco-friendly nematocidal and insecticidal products.

Crop Research Institute Drnovska 507 110 00 119 99 Prague Czech Republic

Department of Agriculture Food and Environment University of Pisa via del Borghetto 80 56124 Pisa Italy

Department of Agronomy Food Natural Resources Animals and Environment University of Padova 35020 Legnaro Italy

Department of Pharmaceutical and Pharmacological Sciences University of Padova 35128 Padova Italy

Department of Pharmacy Faculty of Health Sciences Universidad San Jorge Villanueva de Gállego 50830 Zaragoza Spain

Department of Veterinary Medicine University of Bari Aldo Moro 70100 Bari Italy

Instituto Agroalimentario de Aragón IA2 50013 Zaragoza Spain

School of Bioscience and Veterinary Medicine University of Camerino Centro Ricerche Floristiche dell'Appennino San Colombo 67021 Barisciano Italy

School of Pharmacy University of Camerino via Sant'Agostino 1 62032 Camerino Italy

See more in PubMed

Guardone L., Armani A., Nucera D., Costanzo F., Mattiucci S., Bruschi F. Human anisakiasis in Italy: A retrospective epidemiological study over two decades. Parasite. 2018;25:41. doi: 10.1051/parasite/2018034. PubMed DOI PMC

Herrador Z., Daschner Á., Perteguer M.J., Benito A. Epidemiological scenario of anisakidosis in Spain based on associated hospitalizations: The tipping point of the iceberg. Clin. Infect. Dis. 2018 doi: 10.1093/cid/ciy853. PubMed DOI PMC

Kassai T., Del Campillo M.C., Euzeby J., Gaafar S., Hiepe T., Himonas C.A. Standardized nomenclature of animal parasitic diseases (SNOAPAD) Vet. Parasitol. 1988;29:299–326. doi: 10.1016/0304-4017(88)90148-3. PubMed DOI

Shimamura Y., Muwanwella N., Chandran S., Kandel G., Marcon N. Common symptoms from an uncommon infection: Gastrointestinal anisakiasis. Can. J. Gastroenterol. Hepatol. 2016;2016:5176502. doi: 10.1155/2016/5176502. PubMed DOI PMC

Kilpatrick A.M., Randolph S.E. Drivers, dynamics, and control of emerging vector-borne zoonotic diseases. Lancet. 2012;380:1946–1955. doi: 10.1016/S0140-6736(12)61151-9. PubMed DOI PMC

Otranto D., Dantas-Torres F. Vector-Borne Parasitic Zoonotic Infections in Humans. In: Singh S.K., editor. Human Emerging and Re-emerging Infections. John Wiley & Sons, Inc.; Chichester, UK: 2015. pp. 505–516. DOI

Waltner-Toews D. Zoonoses, One Health and complexity: Wicked problems and constructive conflict. Phil. Trans. R. Soc. B Biol. Sci. 2017;372:20160171. doi: 10.1098/rstb.2016.0171. PubMed DOI PMC

Petersen E., Wilson M.E., Touch S., McCloskey B., Mwaba P., Bates M., Dar O., Mattes F., Kidd M., Ippolito G., et al. Unexpected and rapid spread of Zika virus in the Americas—Implications for public health preparedness for mass gatherings at the 2016 Brazil Olympic Games. Int. J. Infect. Dis. 2015;44:11–15. doi: 10.1016/j.ijid.2016.02.001. PubMed DOI

Benelli G., Duggan M.F. Management of arthropod vector data—Social and ecological dynamics facing the One Health perspective. Acta Trop. 2018;182:80–91. doi: 10.1016/j.actatropica.2018.02.015. PubMed DOI

WHO . Lymphatic Filariasis: Fact Sheet N°102. World Health Organization; Geneva, Switzerland: 2014.

Wilke A.B.B., Beier J.C., Benelli G. Filariasis vector control down-played due to the belief the drugs will be enough—Not true! Entomol. Gen. 2019 doi: 10.1127/entomologia/2019/0776. DOI

Benelli G., Romano D. Mosquito vectors of Zika virus. Entomol. Gen. 2017;36:309–318. doi: 10.1127/entomologia/2017/0496. DOI

Guedes D.R., Paiva M.H., Donato M.M., Barbosa P.P., Krokovsky L., dos SRocha S.W., La Saraiva K., Crespo M.M., Rezende T.M.T., Wallau G.L., et al. Zika virus replication in the mosquito Culex quinquefasciatus in Brazil. Emerg. Microbes Infect. 2017;6:e69. doi: 10.1038/emi.2017.59. PubMed DOI PMC

Van den Hurk A.F., Hall-Mendelin S., Jansen C.C., Higgs S. Zika virus and Culex quinquefasciatus mosquitoes: A tenuous link. Lancet Infect. Dis. 2017;17:1014–1016. doi: 10.1016/S1473-3099(17)30518-2. PubMed DOI

Corbel V., N’guessan R., Brengues C., Chandre F., Djogbenou L., Martin T., Akogbéto M., Hougard J.M., Rowland M. Multiple insecticide resistance mechanisms in Anopheles gambiae and Culex quinquefasciatus from Benin, West Africa. Acta Trop. 2007;101:207–216. doi: 10.1016/j.actatropica.2007.01.005. PubMed DOI

Benelli G., Beier J.C. Current vector control challenges in the fight against malaria. Acta Trop. 2017;174:91–96. doi: 10.1016/j.actatropica.2017.06.028. PubMed DOI

Dantas-Torres F., Chomel B.B., Otranto D. Ticks and tick-borne diseases: A One Health perspective. Trends Parasitol. 2012;28:437–446. doi: 10.1016/j.pt.2012.07.003. PubMed DOI

Paternoster G., Tomassone L., Favretto A., Balduzzi G., Tamba M., Chiari M., Lavazza A., Vogler B. Evaluation of One Health practices to tackle zoonoses: The example of the integrated WNV surveillance in Northern Italy; Proceedings of the 8th International Conference on Emerging Zoonoses focusing on Emerging and Transboundary Infectious Diseases; Manhattan, KS, USA. 7–10 May 2017; p. 28.

Isman M.B. Botanical insecticides, deterrents, and repellents in modern agriculture and an increasingly regulated world. Annu. Rev. Entomol. 2006;51:45–66. doi: 10.1146/annurev.ento.51.110104.151146. PubMed DOI

Stevenson P.C., Isman M.B., Belmain S.R. Pesticidal plants in Africa: A global vision of new biological control products from local uses. Ind. Crop. Prod. 2017;110:2–9. doi: 10.1016/j.indcrop.2017.08.034. DOI

Benelli G., Pavela R. Beyond mosquitoes—Essential oil toxicity and repellency against bloodsucking insects. Ind. Crops Prod. 2018;117:382–392. doi: 10.1016/j.indcrop.2018.02.072. DOI

Benelli G., Pavela R. Repellence of essential oils and selected compounds against ticks—A systematic review. Acta Trop. 2018;179:47–54. doi: 10.1016/j.actatropica.2017.12.025. PubMed DOI

Miresmailli S., Isman M.B. Botanical insecticides inspired by plant—Herbivore chemical interactions. Trends Plant. Sci. 2014;19:29–35. doi: 10.1016/j.tplants.2013.10.002. PubMed DOI

Benelli G. Managing mosquitoes and ticks in a rapidly changing world—Facts and trends. Saudi J. Biol. Sci. 2019 doi: 10.1016/j.sjbs.2018.06.007. PubMed DOI PMC

AlShebly M.M., AlQahtani F.S., Govindarajan M., Gopinath K., Vijayan P., Benelli G. Toxicity of ar-curcumene and epi-β-bisabolol from Hedychium larsenii (Zingiberaceae) essential oil on malaria, chikungunya and St. Louis encephalitis mosquito vectors. Ecotoxicol. Environ. Saf. 2017;137:149–157. doi: 10.1016/j.ecoenv.2016.11.028. PubMed DOI

Giarratana F., Muscolino D., Beninati C., Giuffrida A., Panebianco A. Activity of Thymus vulgaris essential oil against Anisakis larvae. Exp. Parasitol. 2014;142:7–10. doi: 10.1016/j.exppara.2014.03.028. PubMed DOI

Gómez-Rincón C., Langa E., Murillo P., Valero M.S., Berzosa C., López V. Activity of tea tree (Melaleuca alternifolia) essential oil against L3 larvae of Anisakis simplex. BioMed Res. Int. 2014;2014 doi: 10.1155/2014/549510. PubMed DOI PMC

Romero M.C., Navarro M.C., Martín-Sánchez J., Valero A. Peppermint (Mentha piperita) and albendazole against anisakiasis in an animal model. Trop. Med. Int. Health. 2014;19:1430–1436. doi: 10.1111/tmi.12399. PubMed DOI

Gómez-Mateos Pérez M., Navarro Moll C., Merino Espinosa G., Valero López A. Evaluation of different Mediterranean essential oils as prophylactic agents in anisakidosis. Pharm Biol. 2017;55:456–461. doi: 10.1080/13880209.2016.1247880. PubMed DOI PMC

Giarratana F., Muscolino D., Ziino G., Giuffrida A., Marotta S.M., Lo Presti V., Chiofalo V., Panebianco A. Activity of Tagetes minuta Linnaeus (Asteraceae) essential oil against L3 Anisakis larvae type 1. Asian Pac. J. Trop. Med. 2017;10:461–465. doi: 10.1016/j.apjtm.2017.05.005. PubMed DOI

López V., Cascella M., Benelli G., Maggi F., Gómez-Rincón C. Green drugs in the fight against Anisakis simplex—Larvicidal activity and acetylcholinesterase inhibition of Origanum compactum essential oil. Parasitol. Res. 2018;117:861–867. doi: 10.1007/s00436-018-5764-3. PubMed DOI PMC

Ietswaart J.H. A Taxonomic Revision of the Genus Origanum (Labiatae) Leiden University Press; Leiden, The Netherlands: 1980. (Leiden Botanical Series).

Carlström A. New species of Alyssum, Consolida, Origanum and Umblicus from the SE Aegean Sea. Willdenowia. 1984;14:15–26.

Danin A. Two new species of Origanum (Labiatae) from Jordan. Willdenowia. 1990;19:401–404.

Danin A., Künne I. Origanum jordanicum (Labiatae), a New Species from Jordan, and Notes on the Other Species of O. sect. Campanulaticalyx. Willdenowia. 1996;25:601–611.

Duman H., Aytaç Z., Ekici M., Karavelioğulları F.A., Dönmez A.A., Duran A. Three New Species (Labiatae) From Turkey. Flora Mediter. 1996;5:221–228.

Duman H., Başer K.H.C., Aytaç Z. Two New Species and a New Hybrid from Anatolia. Turk. J. Bot. 1998;22:51–55.

Dirmenci T., Yazıcı T., Özcan T., Çelenk Ç., Martin E. A new species and a new natural hybrid of Origanum, L. (Lamiaceae) from the west of Turkey. Turk. J. Bot. 2018;42:73–90. doi: 10.3906/bot-1704-35. DOI

Greuter W., Burdet H.M., Long G. Med-Checklist. Conserv. Jard. Bot. Ville Genkre. 1986;3:308.

Salah S.M., Jäger A.K. Screening of traditionally used Lebanese herbs for neurological activities. J. Ethnopharmacol. 2005;97:145–149. doi: 10.1016/j.jep.2004.10.023. PubMed DOI

El Beyrouthy M., Arnold N.A., Annick D.D., Frederic D. Plants used as remedies antirheumatic and antineuralgic in the traditional medicine of Lebanon. J. Ethnopharmacol. 2008;120:315–334. PubMed

Khoury M., Stien D., Eparvier V., Ouaini N., El Beyrouthy M. Report on the medicinal use of eleven Lamiaceae species in Lebanon and rationalization of their antimicrobial potential by examination of the chemical composition and antimicrobial activity of their essential oils. Evid. Based Complement. Alternat. Med. 2016;2016 doi: 10.1155/2016/2547169. PubMed DOI PMC

Yaniv Z., Dafni A., Friedman J., Palevitch D. Plants used for the treatment of diabetes in Israel. J. Ethnopharmacol. 1987;19:145–151. doi: 10.1016/0378-8741(87)90038-9. PubMed DOI

Darwish R.M., Aburjai T.A. Effect of ethnomedicinal plants used in folklore medicine in Jordan as antibiotic resistant inhibitors on Escherichia coli. BMC Complement. Altern. Med. 2010;10:9. PubMed PMC

Loizzo M.R., Menichini F., Conforti F., Tundis R., Bonesi M., Saab A.M., Statti G.A., de Cindio B., Houghton P.J., Menichini F., et al. Chemical analysis, antioxidant, antiinflammatory and anticholinesterase activities of Origanum ehrenbergii Boiss and Origanum syriacum L. essential oils. Food Chem. 2009;117:174–180. doi: 10.1016/j.foodchem.2009.03.095. DOI

El Gendy A.N., Leonardi M., Mugnaini L., Bertelloni F., Ebani V.V., Nardoni S., Mancianti F., Hendawy S., Omer E., Pistelli L. Chemical composition and antimicrobial activity of essential oil of wild and cultivated Origanum syriacum plants grown in Sinai, Egypt. Ind. Crops Prod. 2015;67:201–207. doi: 10.1016/j.indcrop.2015.01.038. DOI

Viuda-Martos M., El Gendy A.E.N.G., Sendra E., Fernandez-Lopez J., Abd El Razik K.A., Omer E.A., Pérez-Alvarez J.A. Chemical composition and antioxidant and anti-Listeria activities of essential oils obtained from some Egyptian plants. J. Agric. Food Chem. 2010;58:9063–9070. doi: 10.1021/jf101620c. PubMed DOI

Lukas B., Samuel R., Novak J. Oregano or marjoram? The enzyme γ-terpinene synthase affects chemotype formation in the genus Origanum. Isr. J. Plant. Sci. 2010;58:211–220. doi: 10.1560/IJPS.58.3-4.211. DOI

Baser K.H.C., Kurkcuoglu M., Demirci B., Ozek T. The essential oil of Origanum syriacum L. var. sinaicum (Boiss.) Ietswaart. Flavour. Fragr. J. 2003;18:98–99.

Zein S., Awada S., Rachidi S., Hajj A., Krivoruschko E., Kanaan H. Chemical analysis of essential oil from Lebanese wild and cultivated Origanum syriacum L. (Lamiaceae) before and after flowering. J. Med. Plants Res. 2011;5:379–387.

Al Hafi M., El Beyrouthy M., Ouaini N., Stien D., Rutledge D., Chaillou S. Chemical composition and antimicrobial activity of Origanum libanoticum, Origanum ehrenbergii, and Origanum syriacum growing wild in Lebanon. Chem. Biodivers. 2016;13:555–560. doi: 10.1002/cbdv.201500178. PubMed DOI

Traboulsi A.F., Taoubi K., El-Haj S., Bessiere J.M., Rammal S. Insecticidal properties of essential plant oils against the mosquito Culex pipiens molestus (Diptera: Culicidae) Pest. Manag. Sci. 2002;58:491–495. doi: 10.1002/ps.486. PubMed DOI

Kordali S., Emsen B., Yıldırım E. Fumigant toxicity of essential oils from fifteen plant species against Sitophilus zeamais Motschulsky (Coleoptera: Curculionidae) Egypt. J. Biol. Pest. Co. 2013;23:241–246.

Sener O., Arslan M., Demirel N., Uremis I. Insecticidal effects of some essential oils against the confused flour beetle (Tribolium confusum du Val) (Col.: Tenebrinoidea) in stored wheat. Asian J. Chem. 2009;21:3995.

Tunc I., Berger B.M., Erler F., Dağlı F. Ovicidal activity of essential oils from five plants against two stored-product insects. J. Stored Prod. Res. 2000;36:161–168. doi: 10.1016/S0022-474X(99)00036-3. DOI

Oka Y., Nacar S., Putievsky E., Ravid U., Yaniv Z., Spiegel Y. Nematicidal activity of essential oils and their components against the root-knot nematode. Phytopathology. 2000;90:710–715. doi: 10.1094/PHYTO.2000.90.7.710. PubMed DOI

Dudai N., Poljakoff-Mayber A., Mayer A.M., Putievsky E., Lerner H.R. Essential oils as allelochemicals and their potential use as bioherbicides. J. Chem. Ecol. 1999;25:1079–1089. doi: 10.1023/A:1020881825669. DOI

Benelli G., Pavela R., Iannarelli R., Petrelli R., Cappellacci L., Cianfaglione K., Afshar F., Nicoletti M., Canale A., Maggi F. Synergized mixtures of Apiaceae essential oils and related plant-borne compounds: Larvicidal effectiveness on the filariasis vector Culex quinquefasciatus Say. Ind. Crops Prod. 2017;96:186–195. doi: 10.1016/j.indcrop.2016.11.059. DOI

Benelli G., Pavela R., Petrelli R., Cappellacci L., Canale A., Senthil-Nathan S., Maggi F. Not just popular spices! Essential oils from Cuminum cyminum and Pimpinella anisum are toxic to insect pests and vectors without affecting non-target invertebrates. Ind. Crops Prod. 2018;124:236–243. doi: 10.1016/j.indcrop.2018.07.048. DOI

Benelli G., Pavela R., Giordani C., Casettari L., Curzi G., Cappellacci L., Petrelli R., Maggi F. Acute and sub-lethal toxicity of eight essential oils of commercial interest against the filariasis mosquito Culex quinquefasciatus and the housefly Musca domestica. Ind. Crop. Prod. 2018;112:668–680. doi: 10.1016/j.indcrop.2017.12.062. DOI

Benelli G., Pavela R., Petrelli R., Cappellacci L., Bartolucci F., Canale A., Maggi F. Origanum syriacum subsp. syriacum: From an ingredient of Lebanese ‘manoushe’to a source of effective and eco-friendly botanical insecticides. Ind. Crop. Prod. 2019;134:26–32.

Romero M.C., Valero A., Martín-Sánchez J., Navarro-Moll M.C. Activity of Matricaria chamomilla essential oil against anisakiasis. Phytomedicine. 2012;19:520–523. doi: 10.1016/j.phymed.2012.02.005. PubMed DOI

Audicana M.T., Kennedy M.W. Anisakis simplex: From Obscure Infectious Worm to Inducer of Immune Hypersensitivity. Clin. Microbiol. Rev. 2008;21:360–379. doi: 10.1128/CMR.00012-07. PubMed DOI PMC

Benelli G., Pavela R., Canale A., Cianfaglione K., Ciaschetti G., Conti F., Nicoletti M., Senthil-Nathan S., Mehlhorn H., Maggi F. Acute larvicidal toxicity of five essential oils (Pinus nigra, Hyssopus officinalis, Satureja montana, Aloysia citrodora and Pelargonium graveolens) against the filariasis vector Culex quinquefasciatus: Synergistic and antagonistic effects. Parasitol. Int. 2017;66:166–171. doi: 10.1016/j.parint.2017.01.012. PubMed DOI

Marchese A., Orhan I.E., Daglia M., Barbieri R., Di Lorenzo A., Nabavi S.F., Gortzi O., Izadi M., Nabavi S.M. Antibacterial and antifungal activities of thymol: A brief review of the literature. Food Chem. 2016;210:402–414. doi: 10.1016/j.foodchem.2016.04.111. PubMed DOI

Sharifi-Rad M., Varoni E.M., Iriti M., Martorell M., Setzer W.N., del Mar Contreras M., Salehi B., Soltani-Nejad A., Rajabi S., Tajbakhsh M., et al. Carvacrol and human health: A comprehensive review. Phytother. Res. 2018;32:1675–1687. doi: 10.1002/ptr.6103. PubMed DOI

Jukic M., Politeo O., Maksimovic M., Milos M., Milos M. In vitro acetylcholinesterase inhibitory properties of thymol, carvacrol and their derivatives thymoquinone and thymohydroquinone. Phytother. Res. 2007;21:259–261. doi: 10.1002/ptr.2063. PubMed DOI

Aazza S., Lyoussi B., Miguel M.G. Antioxidant and antiacetylcholinesterase activities of some commercial essential oils and their major compounds. Molecules. 2011;16:7672–7690. doi: 10.3390/molecules16097672. PubMed DOI PMC

Pavela R. Essential oils for the development of eco-friendly mosquito larvicides: A review. Ind. Crop. Prod. 2015;76:174–187. doi: 10.1016/j.indcrop.2015.06.050. DOI

Pavela R., Zabka M., Bednar J., Tříska J., Vrchotová N. New knowledge for yield, composition and insecticidal activity of essential oils obtained from the aerial parts or seeds of fennel (Foeniculum vulgare Mill.) Ind. Crops Prod. 2016;83:275–282. doi: 10.1016/j.indcrop.2015.11.090. DOI

Cheng S.S., Huang C.G., Chen Y.J., Yu J.J., Chen W.J., Chang S.T. Chemical compositions and larvicidal activities of leaf essential oils from two Eucalyptus species. Bioresour. Technol. 2009;100:452–456. doi: 10.1016/j.biortech.2008.02.038. PubMed DOI

Pavela R. Insecticidal properties of Pimpinella anisum essential oils against the Culex quinquefasciatus and the non-target organism Daphnia magna. J. Asia Pac. Entomol. 2014;17:287–293. doi: 10.1016/j.aspen.2014.02.001. DOI

Pavela R. Encapsulation—A convenient way to extend the persistence of the effect of eco-friendly mosquito larvicides. Curr. Org. Chem. 2016;20:2674–2680. doi: 10.2174/1385272820666151026231851. DOI

Pavela R., Benelli G., Pavoni L., Bonacucina G., Cespi G., Cianfaglione K., Bajalan I., Morshedloo M.R., Lupidi G., Romano D., et al. Microemulsions for delivery of Apiaceae essential oils—Towards highly effective and eco-friendly mosquito larvicides? Ind. Crops Prod. 2019;129:631–640. doi: 10.1016/j.indcrop.2018.11.073. DOI

Tabari M.A., Youssefi M.R., Maggi F., Benelli G. Toxic and repellent activity of selected monoterpenoids (thymol, carvacrol and linalool) against the castor bean tick, Ixodes ricinus (Acari: Ixodidae) Vet. Parasitol. 2017;245:86–91. doi: 10.1016/j.vetpar.2017.08.012. PubMed DOI

Pavela R., Benelli G. Essential oils as eco-friendly biopesticides? Challenges and constraints. Trends Plant Sci. 2016;21:1000–1007. doi: 10.1016/j.tplants.2016.10.005. PubMed DOI

Lee K.W., Everts H., Kappert H.J., Yeom K.H., Beynen A.C. Dietary carvacrol lowers body weight gain but improves feed conversion in female broiler chickens. J. Appl. Poult. Res. 2003;12:394–399. doi: 10.1093/japr/12.4.394. DOI

Mattila H.R., Otis G.W., Daley J., Schultz T. Trials of apiguard, a thymol-based miticide part 2. Non-target effects on honey bees. Am. Bee J. 2000;140:68–70.

George D.R., Sparagano O.A.E., Port G., Okello E., Shiei R.S., Guy J.H. Repellence of plant essential oils to Dermanyssus gallinae and toxicity to the non-target invertebrate Tenebrio molitor. Vet. Parasitol. 2009;162:129–134. doi: 10.1016/j.vetpar.2009.02.009. PubMed DOI

Lahlou M. Potential of Origanum compactum as a cercaricide in Morocco. Ann. Trop. Med. Parasitol. 2002;96:587–593. doi: 10.1179/000349802125001447. PubMed DOI

Tong F., Coats J.R. Effects of monoterpenoid insecticides on [3H]-TBOB binding in house fly GABA receptor and 36Cl—Uptake in American cockroach ventral nerve cord. Pestic. Biochem. Physiol. 2010;98:317–324. doi: 10.1016/j.pestbp.2010.07.003. DOI

Enan E. Insecticidal activity of essential oils: Octopaminergic sites of action. Comp. Biochem. Physiol. C Toxicol. Pharmacol. 2001;130:325–337. doi: 10.1016/S1532-0456(01)00255-1. PubMed DOI

Govindarajan M., Rajeswary M., Hoti S.L., Benelli G. Larvicidal potential of carvacrol and terpinen-4-ol from the essential oil of Origanum vulgare (Lamiaceae) against Anopheles stephensi, Anopheles subpictus, Culex quinquefasciatus and Culex tritaeniorhynchus (Diptera: Culicidae) Res. Vet. Sci. 2016;104:77–82. doi: 10.1016/j.rvsc.2015.11.011. PubMed DOI

Mouterde P. Nouvelle Flore du Liban et de la Syrie. Volume 3 Dar El-Machreq; Beyrouth, Lebanon: 1984.

Benelli G., Pavela R., Drenaggi E., Maggi F. Insecticidal efficacy of the essential oil of jambú (Acmella oleracea (L.) R.K. Jansen) cultivated in central Italy against filariasis mosquito vectors, houseflies and moth pests. J. Ethnopharmacol. 2019;229:272–279. doi: 10.1016/j.jep.2018.08.030. PubMed DOI

WHO . Report of the Who Informal Consultation on the Evaluation and Testing of Insecticides. WHO; Geneva, Switzerland: 1996. CTD/WHOPES/IC/96.1.

Ellman G.L., Courtney K.D., Andres V., Jr., Featherstone R.M. A new and rapid colorimetric determination of acetylcholinesterase activity. Biochem. Pharmacol. 1961;7:88–90. doi: 10.1016/0006-2952(61)90145-9. PubMed DOI

Abbott W.S. A method of computing the effectiveness of an insecticide. J. Econ. Entomol. 1925;18:265–267. doi: 10.1093/jee/18.2.265a. DOI

Finney D.J. Probit Analysis. Cambridge University; London, UK: 1971. pp. 68–78.

Pavela R., Pavoni L., Bonacucina G., Cespi M., Kavallieratos N.G., Cappellacci L., Petrelli R., Maggi F., Benelli G. Rationale for developing novel mosquito larvicides based on isofuranodiene microemulsions. J. Pest Sci. 2019;92:909–921. doi: 10.1007/s10340-018-01076-3. DOI