2 Disc

Acta Tropica 193 (2019) 236–271
Contents lists available at ScienceDirect
Acta Tropica
journal homepage: www.elsevier.com/locate/actatropica
Plant extracts for developing mosquito larvicides: From laboratory to the T

field, with insights on the modes of action
Roman Pavelaa,1, Filippo Maggib,1, , Romilde Iannarellib, Giovanni Benellic,
⁎ ⁎
a
Crop Research Institute, Drnovska 507, 161 06, Prague 6, Ruzyne, Czech Republic
b
School of Pharmacy, University of Camerino, via Sant’Agostino, 62032 Camerino, Italy
c
Department of Agriculture, Food and Environment, University of Pisa, via del Borghetto 80, 56124 Pisa, Italy
ARTICLE INFO ABSTRACT
Keywords: In the last decades, major research efforts have been done to investigate the insecticidal activity of plant-based products
Biopesticides against mosquitoes. This is a modern and timely challenge in parasitology, aimed to reduce the frequent overuse of
Botanical insecticides synthetic pesticides boosting resistance development in mosquitoes and causing serious threats to human health and
Chemical characterization environment. This review covers the huge amount of literature available on plant extracts tested as mosquito larvicides,
Culicidae
particularly aqueous and alcoholic ones, due to their easy formulation in water without using surfactants. We analysed
Dengue
Insecticide resistance
results obtained on more than 400 plant species, outlining that 29 of them have outstanding larvicidal activity (i.e., LC50
Filariasis values below 10 ppm) against major vectors belonging to the genera Anopheles, Aedes and Culex, among others.
Malaria Furthermore, synergistic and antagonistic effects between plant extracts and conventional pesticides, as well as among
Mosquito control selected plant extracts are discussed. The efficacy of pure compounds isolated from the most effective plant extracts and
Microemulsions – when available – their mechanism of action, as well as the impact on non-target species, is also covered. These belong
Nanoparticles to the following class of secondary metabolites: alkaloids, alkamides, sesquiterpenes, triterpenes, sterols, flavonoids,
Vector-borne disease coumarins, anthraquinones, xanthones, acetogenonins and aliphatics. Their mode of action on mosquito larvae ranges
West Nile
from neurotoxic effects to inhibition of detoxificant enzymes and larval development and/or midugut damages. In the
Zika virus
final section, current drawbacks as well as key challenges for future research, including technologies to synergize
efficacy and improve stability - thus field performances - of the selected plant extracts, are outlined. Unfortunately,
despite the huge amount of laboratory evidences about their efficacy, only a limited number of studies was aimed to
validate their efficacy in the field, nor the epidemiological impact potentially arising from these vector control op-
erations has been assessed. This strongly limits the development of commercial mosquito larvicides of botanical origin,
at variance with plant-borne products developed in the latest decades to kill or repel other key arthropod species of
medical and veterinary importance (e.g., ticks and lice), as well as mosquito adults. Further research on these issues is
urgently needed.
1. Introduction Despite decades of intensive research, the effective and sustainable

management of mosquito vector populations is still a hard challenge to deal
Arthropods act as vectors of many diseases, caused by a huge with (Baldacchino et al., 2015; Benelli and Mehlhorn, 2016; Hemingway
number of pathogens and parasites (Kilpatrick and Randolph, 2012; et al., 2016). Indeed, even if synthetic pesticides played a major role in
Dantas-Torres and Otranto, 2016; Leitner et al., 2015; Fernandes et al., reducing the number of several diseases worldwide, with special reference
2018). Among them, a prominent role is played by mosquitoes (Diptera: to malaria in African countries (Benelli and Beier, 2017), their massive
Culicidae), responsible for the spread of malaria, dengue, yellow fever, overuse has been found responsible for resistance development in targeted
chikungunya, West Nile, lymphatic filariasis, Rift Valley fever, St. Louis vectors (Hemingway et al., 2004; Liu, 2015; Ranson and Lissenden, 2016),
encephalitis, and – as recently pointed out by outbreaks in South along with serious non-target effects on human health and the environment
America and the Pacific – Zika virus, among others (Brady et al., 2012; (Barata et al., 2004, 2006; Benelli, 2018a).
Bird and McElroy, 2016; Kindhauser et al., 2016; Kotsakiozi et al., Therefore, several routes have been explored to reduce/eliminate the
2017; Mayer et al., 2017; Benelli et al., 2018a). use of synthetic insecticides, in the framework of Integrated Vector
⁎
Corresponding authors.
E-mail addresses: filippo.maggi@unicam.it (F. Maggi), giovanni.benelli@unipi.it (G. Benelli).
1
These authors contributed equally to this work.
https://doi.org/10.1016/j.actatropica.2019.01.019
Received 14 December 2018; Received in revised form 23 January 2019; Accepted 23 January 2019
Available online 31 January 2019
0001-706X/ © 2019 Elsevier B.V. All rights reserved.
R. Pavela, et al. Acta Tropica 193 (2019) 236–271
Management (IVM) and One Health (OH) approaches (Dantas-Torres et al., structures with outstanding activities against ectoparasites and vectors
2012; Benelli and Duggan, 2018). Biotechnology helped with several solu- (Banumathi et al., 2017a; Benelli et al., 2017a, b; Benelli, 2018b, c). In
tions, ranging from the Sterile Insect Technique (SIT, currently available this scenario, the present work is aimed to provide a systematic review
also as “boosted SIT”) (Bouyer and Lefrançois, 2014; Lees et al., 2015; Azrag on the effectiveness of plant extracts as mosquito larvicides, particu-
et al., 2016; Chung et al., 2018) to World Mosquito Program’s Wolbachia larly aqueous and alcoholic ones, due to their easy formulation in water
method (Bourtzis et al., 2016; Puggioli et al., 2016) and the employ of without using surfactants, focusing on the most effective botanical fa-
genetically-modified mosquito strains for population suppression or popu- milies and the chemical characterization of species used for extract
lation replacement purposes (Wilke et al., 2018). While the efficacy of SIT- preparation leading to a LC50 lower than 50 ppm, according to the
based approach has been validated for several species of economic im- criteria earlier established by Pavela (2015a). Furthermore, synergistic
portance, including agricultural pests, its potential to control mosquitoes and antagonistic effects between plant extracts and conventional pes-
remains patchy, due to the low competitiveness of most mass-reared ster- ticides, as well as among selected plant extracts are discussed. The ef-
ilized males, when interacting with potential mates (Benelli, 2015a, see also ficacy of pure compounds isolated from the most effective plant extracts
Bourtzis et al., 2016). No evidences about the epidemiological impact of and – when available – their mode of action, is covered. A further focus
GM-based population suppression or replacement strategies are currently is provided on field studies evaluating the efficacy of plant extracts and
available. In addition, 100% refractoriness of a given pathogen or parasite in the subsequent development of commercial products for real-world
GM mosquitoes is a difficult challenge, still unaddressed (Wilke et al., applications. In the final section, major limitations and challenges for
2018). Developing proper GM strains for multiple vectors of a given mos- future research, including technologies to improve stability and field
quito-borne disease is often not suitable for a rather wide number of de- performances of the selected plant extracts, are discussed.
veloping countries registering high numbers of deaths and morbidity yearly
(Wilke et al., 2018). 2. Methods and criteria for literature selection
From a biological control perspective, several natural enemies have
been considered, mostly against mosquito young instars, such as fishes, Considering the non-uniform methods used by various authors, we
anurans (particularly tadpoles), waterbugs and predatory copepods have determined criteria to select studies suitable for the aim of this
(Bowatte et al., 2013; Lazaro et al., 2015; Cuthbert et al., 2018). However, review. We selected researches focusing on the larvicidal activity of
non-target effects have been reported for most of them (Mischke et al., plant extracts against mosquitoes as listed in the Web of Science data-
2016), with special reference to fishes (e.g., the mosquitofish, Gambusia base, using all combinations of “plant extract”, “mosquito” and “larvi-
affinis Baird & Girard predating a wide range of aquatic invertebrates) (see cide”, as well as “plant extract”, “mosquito” and “larvicidal activity”, as
Benelli et al., 2016 for a dedicated review). Biocontrol also includes the keywords; “botanical extract” was also used as keyword. Similar re-
employ of entomopathogenic species, mainly Lagenidium, Coelomomyces, searches we carried out on Scopus database (accessed: October 2017).
Entomophthora, Culicinomyces, Beauveria (e.g., Beauveria bassiana (Bals.- Therefore, here all LC50(90) are given in ppm to compare the results
Criv.) Vuill.), and Metarhizium (Metarhizium anisopliae (Metchnikoff) Sor- between themselves. Moreover, due to the large amount of literature
okin) (Scholte et al., 2004; Kanzok and Jacobs-Lorena, 2006; Amerasan available on this topic, for a single plant species, we selected only the
et al., 2016), the symbiotic control of mosquitoes by recombinant bacteria most efficient extracts. In other words, if the authors tested various
and yeasts (e.g., Asaia sp., Wickerhamomyces anomalus) (Ricci et al., 2011, solvents to prepare a given plant extract, only the extract with the
2012; Jiménez-Cortés et al., 2018), as well as the widely-used bacterial lowest LC50 was presented in Table 1. Lastly, we omitted doubtful ar-
toxins from Bacillus thuringiensis Berliner var. israelensis (Bti) (Bravo et al., ticles (e.g., showing an LC50 value of < 0.1 ppm if other authors re-
2007) currently available on the marked since 30 years – even as slow- ported an LC50 of > 100 ppm) for the same plant extract.
release formulations (Zhang et al., 2016) – showing a little environmental
impact (Lawler, 2017). On the other hand, evidences about mosquito re- 3. Mosquito larvicidal activity of 429 plant extracts
sistance to Bti toxins have been documented (Bonin et al., 2015; Wirth et al.,
2015; Melo et al., 2016). Based on the analysis carried out in the present review, 429 plant
Therefore, one may argue: which cheap and effective tools can be species were assessed for their larvicidal efficacy against various mos-
exploited by marginalized poor populations worldwide to reduce the in- quito vectors (Table 1). They belong to 101 botanical families. How-
cidence of mosquito bites during their lifespan? In this framework, plants ever, most the plant extracts tested on mosquitoes were obtained from a
offer an extraordinary diversity of secondary metabolites with proven effi- few families, namely Asteraceae (49 species), Lamiaceae (41 species),
cacy against mosquito species of medical and veterinary importance as well Fabaceae (40 species) and Rutaceae (18 species) (Fig. 1).
as against other arthropod noxious pests and vectors (Isman, 2006; Isman These families belong to the groups of Rosidae (e.g., Fabaceae and
and Grieneisen, 2014; Benelli and Pavela, 2018a, b; Beck et al., 2013). Rutaceae) and Asteridae (e.g., Asteraceae and Lamiaceae) which abandoned
Essential oils extracted from many plant species have been recognized as the woody habitus in favor of shrub and herbaceous forms (Hufford, 1992).
extremely interesting mosquito ovicides (Benelli, 2015b), larvicides (Amer Also, their flowers acquired more specialization towards pollinators con-
and Mehlhorn, 2006; Pavela, 2015a; Maggi and Benelli, 2018), pupicides, sisting in zygomorphic, complex corollas (Donoghue et al., 1998). Rosidae
adulticides (Chellappandian et al., 2018), insect growth regulators, ovipo- explored the biosynthetic pathway of coumarins, tannins and flavonoids
sition deterrents (Shaalan and Canyon, 2018) and adult repellents (Pavela with a residual production of alkaloids, whereas limited was the production
and Benelli, 2016a). The multiple mechanisms of action of plant-based in- of terpenoids (Gardner, 1977). The latter were over synthesized in the As-
secticides, inhibiting the neurotransmitter acetylcholinesterase and acting teridae group where these compounds reached their maximum expression
both on GABA and octopamine receptors (Enan, 2001, 2005; Price and on a qualitative and quantitative point of view. This specialization in the
Berry, 2006), make the development of resistance in targeted species un- production of secondary metabolites relied on the presence of secretory
likely (Rattan, 2010; Jankowska et al., 2017). In addition, they are safe to tissues such as epidermal glandular trichomes of Lamiaceae and Asteraceae,
vertebrates (Pavela and Benelli, 2016b), and low toxicity levels have been or internal oil glands and channels of Rutaceae and Fabaceae, respectively
reported also on non-target aquatic invertebrates, even if with some con- (Gardner, 1977).
flicting evidences (e.g., Conti et al., 2014). Considering the overall toxicity of these plant extracts, only 29 out of
However, while reviews currently published to date mostly focused 429 owned notable larvicidal activity showing LC50 values below 10 ppm
on essential oils, it should be pointed out that there is an enormous (Table 1). Plants with such efficiency are promising for development of new
amount of literature on the effectiveness of plant extracts, including insecticidal formulations that can replace the synthetic ones without giving
algal ones, against mosquitoes. In addition, some of them have been the same disadvantages (e.g. induction of insect resistance, impact on en-
used with success to fabricate nanosized metal, metal oxide and carbon vironment and human health), thus having a significant potential for future
237
Table 1
Current knowledge on 429 plant extracts evaluated as mosquito larvicides.
Plant Family Part of the Solvent used LC50 (ppm) LC90 (ppm) Target organisms Larval Reference
plant instar
Abrus precatorius L. Fabaceae Seed Ethyl acetate 19 72 An. vagus 4 Bagavan and
Seed Methanol 89; 137 663; 1248 Ar. subalbatus, Cx. Rahuman, 2011
vishnui resp.
Seed Acetone 99 486 Ar. subalbatus
Leaf Petroleum ether 359 n.d. Cx. quinquefasciatus 4 Karmegam et al.,
1997
Abutilon indicum (L.) Malvaceae Leaf Methanol 333; 373 904; 1001 Ae. aegypti, An. 3 and 4 Mohankumar et al.,
Sweet stephensi resp. 2016
Whole plant Methanol 112 405 Cx. quinquefasciatus 3 Kovendan et al.,
2012d
Stem Hexane 184 470 Ae. aegypti 4 Kumar et al., 2012c
Acacia caesia (L.) Willd. Fabaceae Leaf Distilled water 125; 136; 242; 257; 274 An. subpictus, Ae. 3 Benelli et al., 2018b
147 albopictus, Cx.
tritaeniorhynchus
Acacia pennata (L.) Fabaceae Shoot tips Ethanol 244.50 n.d. Ae. aegypti 3 Thongwat et al., 2017
Willd.
Acacia nilotica (L.) Willd. Fabaceae Leaf Petroleum ether 70; 56; 58 338; 195; 211 Ae. aegypti, An. 4 Sakthivadivel and
ex Delile stephensi, Cx. Daniel, 2008
quinquefasciatus resp.
Acalypha alnifolia Klein Euphorbiaceae Leaf Ethanol 68844 172104 An. stephensi 3 Murugan et al., 2012
ex Willd. Euphorbiaceae Leaf Methanol 129; 126; 382; 396; 386 Ae. aegypti, An. 4 Kovendan et al.,
128 stephensi, Cx. 2012f
Acalypha indica L. Euphorbiaceae Leaf Methanol 292.5, n.d. Ae. aegypti 1, 2, 3 Kamalakannan et al.,
327.9, and 4 2011
365.3,
420.6 and
467.6
Acanthophora muscoides Rhodomelaceae Whole plant Methanol - 63 n.d. Cx. quinquefasciatus 3 and 4 Devi et al., 1997
(Linnaeus) Bory Petroleum ether
fraction
Acer campestre L. Sapindaceae Leaf Methanol 52 86 Cx. quinquefasciatus 4 Pavela, 2008
Acer cissifolium (Siebold Sapindaceae Leaf Methanol 70 123 Cx. quinquefasciatus 4 Pavela, 2008
& Zucc.) K. Koch
Acer negundo L. Sapindaceae Leaf Methanol 51 89 Cx. quinquefasciatus 4 Pavela, 2008
Acer platanoides L. Sapindaceae Leaf Methanol 28 62 Cx. quinquefasciatus 4 Pavela, 2008
Acer pseudoplatanus L. Sapindaceae Leaf Methanol 23 76 Cx. quinquefasciatus 4 Pavela, 2008
Achillea millefolium L. Asteraceae Stem Methanol 120 159 Cx. quinquefasciatus 4 Pavela, 2008
Achyranthes aspera L. Amaranthaceae Stem Hexane 58 91 Ae. aegypti 4 Kumar et al., 2012c
Leaf Methanol 283.1, n.d. Ae. aegypti 1-4 Kamalakannan et al.,
322.2, 2011
355.3 and
409.1
Adansonia digitata L. Malvaceae Leaf Methanol 78 155 An. stephensi 3 Krishnappa et al.,
2012
Adenanthera pavonina L. Fabaceae Seed Methanol 420; 475 744; 835 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Adhatoda vasica Nees Acanthaceae Leaf Acetone 18; 27 96; 70 An. stephensi, Cx. 4 Kamaraj et al., 2010b
Adiantum raddianum K. Pteridaceae Leaf Distilled water 156; 143; 306; 286; 322 Ae. aegypti, An. 3 Govindarajan et al.,
Presl 170 stephensi, Cx. 2017a
Aegle marmelos (L.) Corr. Rutaceae Leaf Ethyl acetate 167; 99 588; 479 An. subpictus, Cx. 4 Elango et al., 2009
Serr. tritaeniorhynchus resp.
Aganosma cymosa Apocynaceae Leaf Distilled water 183; 204; 364; 400; 424 An. stephensi, Ae. 3 Benelli and
(Roxb.) G.Don 225 aegypti, Cx. Govindarajan, 2017
quinquefasciatus
Ageratina adenophora Asteraceae Leaf Petroleum ether 187 514 Cx. quinquefasciatus 3 Samuel et al., 2014
(Spreng.) King & Leaf Acetone 357; 227 n.d. Ae. aegypti, Cx. 4 Mohan and
H.E. Robins. quinquefasciatus resp. Ramaswamy, 2007
Ageratum conyzoides L. Asteraceae Leaf Methanol 406; 233 n.d.; n.d. An. arabiensis, An. 3 Muema et al., 2016
gambiae sensu stricto
resp.
Leaf Petroleum ether 426 n.d. Cx. quinquefasciatus 3 Sharma et al., 2009b
Aglaia elaeagnoidea Meliaceae Leaf Distilled water 207; 230; 408; 443; 462 An. stephensi, Ae. 3 Benelli et al., 2018c
(A.Juss.) Benth. 246 aegypti, Cx.
quinquefasciatus
Albizia polyantha Fabaceae Stem Ethanol 548 n.d. Ae. aegypti 3 Garcez et al., 2009
(A.Spreng.)
G.P.Lewis
Allium sativum L. Amaryllidaceae Stem Hexane 218 435 Ae. aegypti 4 Kumar et al., 2012c
Bulb Ethanol 166 n.d. Cx. quinquefasciatus 3 Kalu et al., 2010
(continued on next page)
238
Table 1 (continued)
plant instar
Aloe fibrosa Lavranos & Xanthorrhoeaceae Leaf Hexane 50 90 Ae. aegypti 3 Chore et al., 2014
L.E. Newton Leaf Hexane 1760 n.d. An. gambiae 3 Matasyoh et al., 2008
Aloe ngongensis Christian Xanthorrhoeaceae Leaf Hexane 110 480 Ae. aegypti 3 Chore et al., 2014
Ethyl acetate 150 320 Ae. aegypti
Leaf Hexane 840 n.d. An. gambiae 3 Matasyoh et al., 2008
Aloe turkanensis Xanthorrhoeaceae Leaf Ethyl acetate 110 190 Ae. aegypti 3 Chore et al., 2014
Christian Leaf Ethyl acetate 110 n.d. An. gambiae 3 Matasyoh et al., 2008
Aloe vera (L.) Burm. f. Xanthorrhoeaceae Leaf Petroleum ether 253 563 Ae. aegypti 3 Subramaniam et al.,
2012
Leaf Ethanol 5500; 5900; 12400; 12900; Ae. aegypti, An. 3 Arjunan et al., 2012
5100 11800 stephensi, Cx.
Alstonia scholaris (L.) R. Apocynaceae Leaf Isopropanol 394 n.d. Ae. albopictus 3 Yadav et al., 2014
Br.
Alternanthera sessilis (L.) Amaranthaceae Leaf Distilled water 10300 n.d. Cx. quinquefasciatus 3 Rawani et al., 2014b
R. Br. ex DC.
Amaranthus oleraceus L. Amaranthaceae Leaf Petroleum ether 849 n.d. An. stephensi 3 Sharma et al., 2009a
Ammi visnaga (L.) Lam. Apiaceae Seed Methanol 9 45 Cx. quinquefasciatus 4 Pavela, 2008
Anacardium occidentale Anacardiaceae Leaf Methanol 912; 57; 11 n.d. Ae. aegypti, An. 4 Tripathy et al., 2011
L. stephensi, Cx.
Andrographis paniculata Acanthaceae Leaf Ethanol 12 n.d. Ae. aegypti 4 Edwin et al., 2016
(Burm.f.) Nees Leaf Distilled water 129; 109 235; 198 Ae. aegypti, Cx. 4 Renugadevi et al.,
quinquefasciatus resp. 2013
Leaf Methanol 80 155 Anophleles stephensi 3 Govindarajan, 2011b
Leaf Chloroform 100; 92 190; 176 Ae. aegypti, Cx. 3 Govindarajan, 2011c
Leaf and root Ethanol 36 51 An. stephensi 3 Chenniappan et al.,
2011
Leaf Hexane 67; 89 372; 416 An. subpictus, Cx. 4 Elango et al., 2009
tritaeniorhynchus resp.
Whole plant Ethanol 35 47 An. stephensi 4 Chenniappan and
Kadarkarai, 2008
Andrographis lineata Nees Acanthaceae Leaf Distilled water 152; 147 241; 239 Ae. aegypti, Cx. 4 Renugadevi et al.,
Leaf Methanol 2310 7450 An. subpictus 4 Elango et al., 2011
Angelica archangelica L. Apiaceae Fruit Methanol 60 146 Cx. quinquefasciatus 4 Pavela, 2009
Anisomeles indica (L.) Lamiaceae Leaf Distilled water 121; 109; 236; 222; 250 Ae. albopictus, An. 3 Govindarajan et al.,
Kuntze 131 subpictus, Cx. 2016e
Anisomeles malabarica Lamiaceae Leaf Methanol 59 118 An. stephensi 3 Unpaprom et al.,
(L.) R. Br. ex Sims Flower Methanol 71 124 An. stephensi 2015
Leaf Ethyl acetate 2530 6400 An. subpictus 4 Elango et al., 2011
Annona glabra L. Annonaceae Leaf Methanol 114; 449 321; 900 Ae. aegypti, An. 3 and 4 Mohankumar et al.,
stephensi resp. 2016
Annona muricata L. Annonaceae Leaf Distilled water 618; 547; 1241; 937; 937 Ae. aegypti, An. 4 Santhosh et al.,
516 stephensi, Cx. 2015a
Leaf Distilled water 51; 61; 89 82; 157; 200 Ae. aegypti, An. 3 Santhosh et al.,
stephensi, Cx. 2015b
Seed Ethanol 93 442 Ae. aegypti 3 Grzybowski et al.,
2012
Leaf Ethanol 56 n.d. Cx. quinquefasciatus 3 Magadula et al., 2009
Annona reticulata L. Annonaceae Leaf Methanol 95; 263 263; 637 Ae. aegypti, An. 3 and 4 Mohankumar et al.,
Annona senegalensis Pers. Annonaceae Leaf Methanol - N- 299; 2808 573; 16241 An. gambiae, Cx. 4 Lame et al., 2015
hexane fraction quinquefasciatus resp.
Annona squamosa L. Annonaceae Bark Methanol 94; 105 525; 444 An. subpictus, Cx. 4 Kamaraj et al., 2011b
Seed Ethanol - Acetone 3.6 19 Ae. albopictus 3 Kempraj and Bhat,
fraction 2011
Bark Ethyl acetate 25; 43 94; 172 An. stephensi, Cx. 4 Kamaraj et al., 2010b
Leaf Ethyl acetate 19; 60 75; 224 An. subpictus, Cx. 4 Bagavan et al., 2009
Leaf Acetone 17 196 An. subpictus
Leaf Chloroform 64 192 Cx. tritaeniorhynchus
Stem bark Chloroform 17 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Root bark Chloroform 13 n.d. An. gambiae
Anthemis tinctoria L. Asteraceae Flower Methanol 396 628 Cx. quinquefasciatus 4 Pavela, 2009
239
Table 1 (continued)
plant instar
Apium graveolens L. Apiaceae Seed Ethanol 81 177=LC95 Ae. aegypti 4 Choochote et al.,
2004
Argemone mexicana L. Papaveraceae Root Hexane 91 157 Ae. aegypti 4 Warikoo and Kumar,
Leaf Hexane 169 377 2013
Stem Hexane 218 453
Leaf Petroleum ether 140 n.d. Cx. quinquefasciatus 3 Sharma et al., 2009b
Leaf Petroleum ether 49; 48; 30 189; 158; 246 Ae. aegypti, An. 4 Sakthivadivel and
stephensi, Cx. Daniel, 2008
Seed Petroleum ether 33; 20; 24 299; 81; 185 Ae. aegypti, An. 4 Sakthivadivel and
Leaf Benzene 30 n.d. Cx. quinquefasciatus 4 Karmegam et al.,
1997
Aristolochia bracteata Aristolochiaceae Leaf Petroleum ether 362 n.d. Cx. quinquefasciatus 4 Karmegam et al.,
Retz. 1997
Aristolochia indica L. Aristolochiaceae Leaf Methanol 12; 26 62; 106 Cx. gelidus, Cx. 4 Kamaraj et al., 2010a
Leaf Ethanol 262.66, 692.47, 733.08, An. stephensi 1, 2, 3 Murugan et al., 2015f
292.59, 837.49 and and 4
385.71 and 974.64
490.31
Artemisia abrotanum L. Asteraceae Stem Methanol 609 836 Cx. quinquefasciatus 4 Pavela, 2009
Artemisia afra Jacq. ex Asteraceae Leaf Methanol 960 4010 An. gambiae 3 Wachira et al., 2014
Willd.
Artemisia annua L. Asteraceae Leaf Chloroform 0.7; 0.8 5.5; 4.9 Ae. aegypti, An. 3 and 4 Sharma et al., 2014
stephensi resp.
Asteraceae Callus culture Hexane 276; 245; 887; 751; 1018 Ae. aegypti, Anopheles 3 and 4 Cheah et al., 2013
375 sinensis, Cx.
Artemisia campestris L. Asteraceae Stem Methanol 23 36 Cx. quinquefasciatus 4 Pavela, 2009
Artemisia nilagirica Asteraceae Leaf Methanol 395; 362 805; 789 Ae. aegypti, An. 3 Panneerselvam et al.,
(C.B.Clarke) Pamp. stephensi resp. 2012
1997
Artocarpus lakoocha Moraceae Leaf Petroleum ether 1163; 241; 1549; 2714; Ae. aegypti, An. 3 and 4 Prashanthi et al.,
Roxb. 625 2995 stephensi, Cx. 2016
Leaf Ethanol 1022 1549 An. stephensi
Fruit Petroleum ether 1399; 1181; 1659; 1359; Ae. aegypti, An.
Asparagus racemosus Asparagaceae Root Methanol 98; 91; 115 180; 169; 211 Ae. aegypti, An. 3 Govindarajan and
Willd. stephensi, Cx. Sivakumar, 2014a
Asteranthe asterias Annonaceae Leaf Chloroform 267 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
(S.Moore) Engl. & Stem bark Petroleum ether 238 n.d. An. gambiae
Diels
Asteranthe lutea Vollesen Annonaceae Stem bark Chloroform 212 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Root bark Petroleum ether 59 n.d. An. gambiae
Astrodaucus persicus Apiaceae Fruit Methanol - Ethyl 34490 108610 An. stephensi 3 Goodarzi et al., 2017
(Boiss.) Drude acetate fraction
Apiaceae Root Methanol - 101210 478940 An. stephensi
Petroleum ether
fraction
Atalantia monophylla DC. Rutaceae Leaf Methanol 0.1; 2; 0.1 0.3; 6.1; 0.3 Ae. aegypti, An. 4 Sivagnaname and
stephensi, Cx. Kalyanasundaram,
Azadirachta indica A. Meliaceae Leaf Methanol 440 1410 An. gambiae 3 Wachira et al., 2014
Juss. Seed Distilled water 159 441 Ae. aegypti 3 Murugan et al., 2011
Fruit Methanol 22 n.d. Cx. quinquefasciatus 3 Sharma et al., 2009b
neem cake Distilled water 106, 122, 233, 267, 318, Ae. aegypti 1, 2, 3, Chandramohan et al.,
148, 189 414 4 2016
Balanites aegyptiaca (L.) Zygophyllaceae Root Methanol 290; 102 678; 276 Ae. aegypti, An. 4 Patil et al., 2010
Del. stephensi resp.
Balsamita major Desf. Asteraceae Stem Methanol 732 891 Cx. quinquefasciatus 4 Pavela, 2009
Barleria cristata L. Acanthaceae Leaf Distilled water 135; 124; 261; 247; 276 Ae. albopictus, An. 3 Govindarajan and
146 subpictus, Cx. Benelli, 2016b
Basella rubra L. Basellaceae Leaf Benzene 54 86 Ae. aegypti 3 Krishnappa and
Elumalai, 2013
240
Table 1 (continued)
plant instar
Berberis tinctoria Lesch. Berberidaceae Leaf Ethanol 34 n.d. Ae. aegypti 3 Banumathi et al.,
2017b
Distilled water 183; 231; 448; 543; 613; Ae. albopictus 1, 2, 3, Kumar et al., 2016
270; 322 695 4
Bidens pilosa L. Asteraceae Leaf Methanol 620 6540 An. gambiae 3 Wachira et al., 2014
Biophytum sensitivum (L.) Oxalidaceae Leaf Acetone 13 138=LC99 Ae. aegypti 4 Shivakumar et al.,
DC. 2013
Boenninghausenia Rutaceae Leaf Petroleum ether 125; 105; 190; 180; 200 Ae. aegypti, An. 4 Alam et al., 2011
albiflora (Hook.) 115 stephensi, Cx.
Rchb. ex Meisn. quinquefasciatus resp.
Bombax malabaricum DC. Malvaceae Leaf Methanol 49 264 Cx. quinquefasciatus 3 Hossain et al., 2011
Borago officinalis L. Boraginaceae Stem Methanol 371 915 Cx. quinquefasciatus 4 Pavela, 2009
Bougainvillea glabra Nyctaginaceae Leaf Distilled water 199; 235; 477; 564; 309; Cx. tritaeniorhynchus 1, 2, 3, Vincent et al., 2017
Choisy 309; 372 372 4
Bryonia dioica Jacq. Cucurbitaceae Stem Methanol 112 320 Cx. quinquefasciatus 4 Pavela, 2009
Bryopsis pennata J.V. Bryopsidaceae Whole plant Methanol - 83 n.d. Ae. aegypti Yu et al., 2015
Lamouroux Chloroform fraction
Bupleurum exaltatum Apiaceae Stem Methanol 120 210 Cx. quinquefasciatus 4 Pavela, 2008
M.Bieb.
Caesalpinia pulcherrima Fabaceae Leaf Methanol 113; 165 208; 299 An. subpictus, Cx. 3 Govindarajan et al.,
(L.) Sw. tritaeniorhynchus resp. 2012
Leaf Benzene 136; 127; 272; 258; 288 Ae. aegypti, An. 3 Govindarajan et al.,
Leaf Ethyl acetate 131 254 An. stephensi
Callistemon rigidus R. Br. Myrtaceae Leaf Methanol 447 767 Cx. quinquefasciatus 4 Pierre et al., 2014
Leaf Methanol - Hexane 56; 17 174; 83 Ae. aegypti, An. gambiae
fraction resp.
Leaf Methanol 1025; 115 4358; 295 Ae. aegypti, An. gambiae
resp.
Callistemon viminalis Myrtaceae Leaf Isopropanol 71 n.d. Ae. albopictus 3 Yadav et al., 2014
(Soland. ex Gaertn.)
Cheel
Calotropis gigantea (L.) Apocynaceae Leaf Ethanol 136; 122; 328; 374; 432 Ae. aegypti, An. 3 Kovendan et al.,
Ait. f. 174 stephensi, Cx. 2012c
Leaf Methanol 467; 629 963; 1092 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Calotropis procera (Ait.) Apocynaceae Seed Methanol 195; 127; n.d. Ae. aegypti, An. 3 and 4 Bansal et al., 2012
Ait. f. 361 stephensi, Cx.
Seed Acetone 193 n.d. Cx. quinquefasciatus
Leaves Methanol 171; 89; n.d. Ae. aegypti, An.
370 stephensi, Cx.
Flower Methanol 617; 95; n.d. Ae. aegypti, An.
1384 stephensi, Cx.
Root Ethanol 215 300 An. labranchiae 3 and 4 Markouk et al., 2000
Campanula longistyla Campanulaceae Stem Methanol 480 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Fomin
Canna indica L. Cannaceae Leaf Petroleum ether 56 248 Cx. quinquefasciatus 4 Rahuman et al.,
2009a
Capsicum annuum L. Solanaceae Fruit Methanol 208; 241 321; 315 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Cardiospermum Sapindaceae Leaf Methanol 157; 150 300; 296 Ae. aegypti, Cx. 3 Govindarajan, 2011a
halicacabum L. quinquefasciatus resp.
Leaf Chloroform 155 295 Cx. quinquefasciatus
Leaf Methanol 133 271 Anophleles stephensi 3 Govindarajan, 2011b
Carica papaya L. Caricaceae Leaf Methanol 336 680 Ae. aegypti 3 and 4 Kovendan et al.,
2012g
1997
Carissa carandas Apocynaceae Leaf Distilled water 177, 164, 353, 330, 370 Ae. albopictus, An. 3 Govindarajan and
194 stephensi, Cx. Benelli, 2017
Carissa spinarum Apocynaceae Leaf Distilled water 104, 112, 204, 216, 227 Ae. albopictus, An. Govindarajan et al.,
120 subpictus, Cx. 2016g
Cassia angustifolia Vahl Fabaceae Leaf Methanol 18 94 Cx. gelidus 4 Kamaraj et al., 2010a
Leaf Ethyl acetate 35 173 Cx. quinquefasciatus
241
Table 1 (continued)
plant instar
Cassia auriculata L. Fabaceae Flower Methanol 34; 51 133; 205 An. stephensi, Cx. 4 Kamaraj et al., 2010b
Leaf Petroleum ether 44; 70 187; 335 An. subpictus, Cx. 4 Kamaraj et al., 2009
Flower Methanol 45; 51 188; 246 An. subpictus, Cx.
Cassia mimosoides L. Fabaceae Leaf and pod Petroleum ether 280 1360 An. gambiae 3 Alayo et al., 2015
Cassia obtusifolia L. Fabaceae Leaf Ethanol 52 109 An. stephensi 3 Rajkumar and
Jebanesan, 2009
Cassia occidentalis L. Fabaceae Leaf Hexane 75 202 Ae. aegypti 4 Kumar et al., 2012c
Cassia roxburghii DC. Fabaceae Leaf Distilled water 150; 134; 275; 244; 293 Ae. aegypti, An. 3 Muthukumaran et al.,
Catharanthus roseus (L.) Apocynaceae Leaf Petroleum ether 5900 21060 An. stephensi 3 and 4 Panneerselvam et al.,
G. Don 2013a
Leaf Distilled water 69000 185000 An. stephensi 4 Subarani et al., 2013
Leaf Petroleum ether 145; 150; 255; 280; 270 Ae. aegypti, An. 4 Alam et al., 2011
160 stephensi, Cx.
Seed Petroleum ether 53; 38; 47 153; 125; 143 Ae. aegypti, An. 4 Sakthivadivel and
Cedrus deodara (Roxb. ex Pinaceae Stem bark Distilled water (hot) 134 583 Cx. quinquefasciatus 4 Rahuman et al.,
D. Don) G. Don Stem bark Methanol 95 640 Cx. quinquefasciatus 2009b
Ceiba pentandra (L.) Malvaceae Leaf Methanol 119 171=LC95 Ae. aegypti 3 Deepa et al., 2015
Gaertn.
Centella asiatica (L.) Apiaceae Leaf Methanol 27 111 An. subpictus 4 Bagavan et al., 2009
Urban
Cestrum diurnum L. Solanaceae Leaf Distilled water 5700 122700=LC95 Cx. quinquefasciatus 3 Ghosh and Chandra,
2006
Cestrum nocturnum L. Solanaceae Leaf Dichloromethane 30 89 Ae. aegypti 3 Patil et al., 2011
Chloroxylon swietenia DC. Rutaceae Bark Methanol 125; 138; 226; 246; 235 Ae. aegypti, An. 4 Balasubramanian
131 stephensi, Cx. et al., 2015
Chomelia asiatica (L.) Rubiaceae Leaf Distilled water 97; 90; 103 174; 165; 183 Ae. aegypti, An. 3 Muthukumaran et al.,
Kuntze stephensi, Cx. 2015c
Chrysanthemum Asteraceae Leaf Methanol 227; 219 324; 399 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
cinerariifolium quinquefasciatus resp.
(Trevir.) Vis.
Chrysanthemum indicum Asteraceae Leaf Methanol 2560 6980 An. subpictus 4 Elango et al., 2011
L. Leaf Ethyl acetate 40; 42 146; 172 An. subpictus, Cx. 4 Kamaraj et al., 2011b
Cichorium intybus L. Asteraceae Whole plant Petroleum ether 15 n.d. An. pharoensis 4 Mansour et al., 2014
Citrulluscolocynthis (L.) Cucurbitaceae Leaf Petroleum ether 75; 88 538; 338 Ae. aegypti, Cx. 4 Rahuman and
Schrad. quinquefasciatus resp. Venkatesan, 2008
Citrus aurantifolia (L.) Rutaceae Peel Methanol 80; 96 117; 128 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Swingle quinquefasciatus resp.
Citrus sinensis (L.) Osbeck Rutaceae Peel Methanol 96 303 An. stephensi 4 Kamaraj et al., 2008
Citrus grandis (L.) Osbeck Rutaceae Fruit - peel N-hexane 1.1 3.3 Ae. aegypti 3 and 4 Torres et al., 2016
Clausena anisata (Willd.) Rutaceae Leaf Hexane 68 n.d. Ae. aegypti 3 Mukandiwa et al.,
Hook.f. ex Benth. 2015
Clausena dentata (Willd.) Rutaceae Leaf Acetone 169; 46; 1100; 1124; Ae. aegypti, An. 4 Manjari et al., 2014
M. Roemer 150 7303 stephensi, Cx.
Leaf Petroleum benzene 242 2059 Cx. quinquefasciatus
Clausena lansium (Lour.) Rutaceae Seed Petroleum ether 23 89 Ae. albopictus 4 Han et al., 2013
Skeels
Cleistanthus collinus Phyllanthaceae Leaf Ethyl acetate 226 1025 Cx. quinquefasciatus 3 Kovendan et al.,
(Roxb.) Benth. ex 2012b
Hook.f.
Cleome rupicola Vicary Cleomaceae Leaf Methanol 2.2 4.1 Ae. caspius 4 Al-Mekhlafi et al.,
Stem Methanol 1; 3.6 3.9; 7.5 Ae. caspius, Cx. pipiens 2013
Fruit Methanol 2.3; 4.6 4.4; 8.7 resp.
Cleome viscosa L. Cleomaceae Leaf Benzene 82 149 Ae. aegypti 3 Krishnappa and
Elumalai, 2013
Seed Methanol 100; 127 354; 475 Ae. aegypti, Cx. 3 and 4 Bansal et al., 2014
Seed Acetone 106; 119 439; 548 An. stephensi, Cx. 3 and 4
Clerodendrum chinense Lamiaceae Leaf Distilled water 67; 62; 72 130; 123; 136 Ae. albopictus, An. 3 Govindarajan et al.,
(Osb.) Mabberley subpictus, Cx. 2016d
242
Table 1 (continued)
plant instar
Clerodendrum inerme (L.) Lamiaceae Leaf Methanol - 850 n.d. Cx. quinquefasciatus 3 and 4 Devi et al., 1997
Gaertn. Petroleum ether
fraction
Clerodendrum phlomidis Lamiaceae Leaf Chloroform - Ethyl 33; 5 82; 62 Ae. aegypti, Cx. 4 Muthu et al., 2012
L.f. acetate fraction quinquefasciatus resp.
Clitoria ternatea L. Fabaceae Leaf Methanol 556 1190 An. stephensi 4 Mathew et al., 2009
Seed Methanol 195; 117; 599; 298; 681 Ae. aegypti, An.
148 stephensi, Cx.
Root Methanol 376 793 Cx. quinquefasciatus
Flower Methanol 631 1079 Cx. quinquefasciatus
Cocciniaindica Wight & Cucurbitaceae Leaf Methanol 309; 378 1330; 1336 Ae. aegypti, Cx. 4 Rahuman and
Arn. quinquefasciatus resp. Venkatesan, 2008
Coccoloba mollis Casar. Polygonaceae Stem bark Ethanol - Hexane 0.1 n.d. Ae. aegypti 4 Oliveira et al., 2010
fraction
Cocculus hirsutus (L.) Menispermaceae Leaf Methanol 143; 105 830; 508 An. subpictus, Cx. 4 Elango et al., 2009
W.Theob. tritaeniorhynchus resp.
Coix lacryma-joby L. Poaceae Leaf Methanol 93 114=LC95 Ae. aegypti 3 Deepa et al., 2015
Coldenia procumbens L. Coldeniaceae Leaf Petroleum ether 334 n.d. Cx. quinquefasciatus 4 Karmegam et al.,
1997
Coleus amboinicus Lour. Lamiaceae Leaf Ethanol 6200; 6800; 13300; 13500; Ae. aegypti, An. 3 Arjunan et al., 2012
Coleus aromaticus Benth. Lamiaceae Leaf Ethyl acetate 214; 216 33990; 165291 Ae. aegypti, Cx. 4 Ramkumar et al.,
Commiphora berryi (Arn.) Burseraceae Leaf Methanol 97 121=LC95 Ae. aegypti 3 Deepa et al., 2015
Engl.
Convolvulus cantabrica L. Convolvulaceae Stem Methanol 390 490 Cx. quinquefasciatus 4 Pavela, 2008
Cosmos bipinnatus Cav. Asteraceae Leaf Ethanol 1180 n.d. Cx. quinquefasciatus 4 Modise and Ashafa,
2016
Leaf Methanol 443; 258 1226; 1048 Ae. aegypti, An. 3 and 4 Mohankumar et al.,
Cotula cinerea Delile Asteraceae Whole plant Ethyl ether 310 505 An. labranchiae 3 and 4 Markouk et al., 2000
Couroupita guianensis Lecythidaceae Leaf Ethyl acetate 45 318 Ae. aegypti 4 Vimala et al., 2015
Aubl. Fruit Ethyl acetate 50 569 Ae. aegypti
Flower Distilled water 199, 226, 488, 581, 633, An. stephensi 1, 2, 3, Subramaniam et al.,
257, 308 718 4 2015
Croton bonplandianum Euphorbiaceae Leaf Methanol 132 215=LC95 Ae. aegypti 3 Deepa et al., 2015
Baill. Leaf Ethyl acetate 40 218 An. vagus 4 Bagavan and
Rahuman, 2011
Croton macrostachyus Euphorbiaceae Leaf Methanol 89 225 An. arabiensis 3 Karunamoorthi and
Hochst. ex Delile Ilango, 2010
Croton sylvaticus Hochst. Euphorbiaceae Stem bark Chloroform 232 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Root bark Petroleum ether 110 n.d. An. gambiae
Cryptomeria japonica Cupressaceae Sapwood Methanol 12; 16 23; 62 Ae. aegypti, Ae. 4 Cheng et al., 2008
(Thunb. ex L. f.) D. albopictus resp.
Don
Cucumis sativus L. Cucurbitaceae Leaf Methanol 493; 624 1824; 2407 Ae. aegypti, Cx. 4 Rahuman and
quinquefasciatus resp. Venkatesan, 2008
Cunninghamia konishii Cupressaceae Wood Ethanol 240; > 400 > 400; > 400 Aedes eagypti, Ae. 4 Cheng et al., 2013
(Lamb.) Hook. Leaf Ethanol > 400; > 400; > 400 albopictus resp.
> 400
Curcuma amada Roxb. Zingiberaceae Rhizome Methanol 481; 395 783; 607 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Curcuma xanthorrhiza Zingiberaceae Root Hexane 26 n.d. Ae. aegypti 3 Sukari et al., 2010
Roxb. orth. var.
Curcuma heyneana Zingiberaceae Root Petroleum ether 35 n.d. Ae. aegypti 3
Valeton & Zijp
Curcuma mangga Valeton Zingiberaceae Root Ethanol 134 n.d. Ae. aegypti 3
& Zijp
Cymbopogon citratus (DC. Poaceae Leaf Distilled water 349; 298 667; 595 Ae. aegypti, An. 3 Murugan et al.,
ex Nees) Stapf stephensi resp. 2015d
Leaf Methanol 74 158 An. arabiensis 3 Karunamoorthi and
Ilango, 2010
Cynodon dactylon (L.) Poaceae Leaf Ethanol 4000; 4400; 10400; 10900; Ae. aegypti, An. 3 Arjunan et al., 2012
Pers. 3300 9100 stephensi, Cx.
Leaf Methanol 41 184 An. vagus 4 Bagavan and
Leaf Ethyl acetate 22 98 Ar. subalbatus Rahuman, 2011
Dalbergia sissoo Roxb. ex Fabaceae Flower Hexane 55 180 An. stephensi 3 Haq et al., 2016
DC. Branche Acetone 115 342 An. stephensi
Seed Methanol 122 301 An. stephensi
243
Table 1 (continued)
plant instar
Dalbergia oliveri Gamble Fabaceae Hearthwood N-hexane - 289 n.d. Ae. aegypti 3 Pluempanupat et al.,
ex Prain Dichloromethane 2013
fraction
Datura metel L. Solanaceae Leaf Distilled water 47 38 An. stephensi 3 Murugan et al.,
2015a
Delonix elata (L.) Gamble Fabaceae Leaf Methanol 112; 93 203; 164 Ae. aegypti, An. 3 Marimuthu et al.,
Fabaceae Seed Methanol 139; 115 273; 225 Ae. aegypti, An.
stephensi resp.
Derris sp. Lour. Fabaceae Root Hexane 61 94 Ae. aegypti 3 Zaridah et al., 2006
Derris heterophylla Fabaceae Leaf Methanol - 800 n.d. Cx. quinquefasciatus 3 and 4 Devi et al., 1997
(Willd.) K.Heyne Petroleum ether
fraction
Derrisurucu (Killip & Fabaceae Root bark Methanol 18 55 Ae. aegypti 4 Gusmao et al., 2002
A.C.Sm.) J.F.Macbr. Root medulla Methanol 33 84 Ae. aegypti
Dicranopteris linearis Gleicheniaceae Leaf Ethanol 165; 181; 113; 142; 189; Ae. aegypti 1-4 Rajaganesh et al.,
(Burm. f.) Underw. 202; 219 205 2016
Dictamnus albus L. Rutaceae Rhizome Methanol 982 > 1000 Cx. quinquefasciatus 4 Pavela, 2009
Dolichos biflorus L. Fabaceae Leaf Ethyl acetate 33; 37 129; 177 Cx. gelidus, Cx. 4 Kamaraj et al., 2010a
Dracocephalum moldavica Lamiaceae Stem Methanol 789 > 1000 Cx. quinquefasciatus 4 Pavela, 2009
L.
Dregea volubilis (L. f.) Apocynaceae Leaf Methanol 57 218 Cx. quinquefasciatus 3 Hossain et al., 2011
Benth. ex Hook. f.
Dysphania ambrosioides Chenopodiaceae Leaf Distilled water 124.55, 335.30, 398.83, Ae. albopictus 1, 2, 3 Subramaniam et al.,
(L.) Mosyakin & 143.24, 453.72 and and 4 2017
Clemants 172.10 and 488.86
199.55
Echinacea purpurea (L.) Asteraceae Root Methanol 278 503 Cx. quinquefasciatus 4 Pavela, 2009
Moench
Eclipta alba (L.) Hassk. Asteraceae Leaf Methanol 113 221 An. stephensi 3 Govindarajan, 2011b
Leaf Methanol 128 246 Ae. aegypti 3 Govindarajan and
Karuppannan, 2011
Eclipta paniculata Hassk. Asteraceae Aerial part Ethanol 3.3 17 Ae. fluviatilis 4 Macedo et al., 1997
Eclipta prostrata (L.) L. Asteraceae Leaf Acetone 2820 8060 An. subpictus 4 Elango et al., 2011
Leaf Distilled water 28; 27 71; 70 An. subpictus, Cx. 4 Rajakumar and
quinquefasciatus resp. Rahuman, 2011
Leaf Ethyl acetate 78; 120 361; 565 An. subpictus, Cx. 4 Elango et al., 2009
Eichhornia crassipes Pontederiaceae Leaf and Petroleum ether - 120 235 Cx. quinquefasciatus 3 Jayanthi et al., 2012
(Mart.) Solms shoot Ethanol fraction
Ervatamia coronaria (L.) Apocynaceae Leaf Methanol 86; 132 160; 245 An. subpictus, Cx. 3 Govindarajan et al.,
Stapf tritaeniorhynchus resp. 2012
Leaf Benzene 90; 79; 96 166; 150; 174 Ae. aegypti, An. 3 Govindarajan et al.,
stephensi, Cx. 2011a
Erythrina indica Lam. Fabaceae Leaf Methanol 75; 69; 91 134; 125; 167 Ae. aegypti, An. 3 Govindarajan and
stephensi, Cx. Sivakumar, 2014b
Eugenia jambolana Lam. Myrtaceae Leaf Petroleum ether 41 83 Ae. aegypti 4 Raghavendra et al.,
2013
Leaf Petroleum ether 41; 96; 54 83; 156; 127 Ae. aegypti, An. 4 Prathibha et al., 2014
stephensi, Cx.
Euodia ridleyi Hochr. Rutaceae Leaf Ethyl acetate 149; 120; 196; 185; 98 Ae. aegypti, An. 4 Prathibha et al., 2014
75 stephensi, Cx.
Eupatorium adenophorum Asteraceae Leaf Ethanol 41 n.d. Ae. aegypti 3 Banumathi et al.,
Spreng. 2017b
Eupatorium odoratum (L.) Asteraceae Leaf Distilled water 397 717 Cx. quinquefasciatus 3 Elemike et al., 2017
King & H.E. Robins. Leaf Petroleum ether 155; 135; 290; 250; 300 Ae. aegypti, An. 4 Alam et al., 2011
145 stephensi, Cx.
Euphorbia hirta L. Euphorbiaceae Leaf Methanol 218 591 An. stephensi 3 Panneerselvam et al.,
2013b
Leaf Methanol 169 331 An. stephensi 3 Priyadarshini et al.,
2012
Leaf Petroleum ether 9694 n.d. An. stephensi 3 Sharma et al., 2009a
Euphorbia rothiana Euphorbiaceae Leaf Acetone 36.64 n.d. Ae. aegypti 3 Banumathi et al.,
Spreng. 2017b
Leaf Distilled water 8.28 40.19 Ae. aegypti 3 Banumathi et al.,
2017c
Evodia rutaecarpa (A. Rutaceae Unripe fruit Ethanol 43 106=LC95 Ae. albopictus 4 Liu et al., 2012
Juss.) Benth.
244
Table 1 (continued)
plant instar
Feronia elephantum Rutaceae Leaf Distilled water 62; 55; 67 111; 97; 118 Ae. aegypti, An. 3 Veerakumar et al.,
Corrêa stephensi, Cx. 2014b
Ferula lancerottensis Parl. Apiaceae Stem Methanol 435 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Ficus benghalensis L. Moraceae Leaf Methanol 57; 86 101; 160 An. subpictus, Cx. 3 Govindarajan et al.,
tritaeniorhynchus resp. 2011b
Leaf Methanol 70; 76; 58 137; 153; 127 Ae. aegypti, An. 3 Govindarajan, 2010
stephensi, Cx.
Ficus microcarpa L. f. Moraceae Leaf Methanol 92 111=LC95 Ae. aegypti 3 Deepa et al., 2015
Ficus racemosa L. Moraceae Bark Distilled water 64; 68 n.a. Cx. gelidus, Cx. 4 Velayutham et al.,
Ficus religiosa L. Moraceae Leaf Methanol 111 155=LC95 Ae. aegypti 3 Deepa et al., 2015
Foeniculum vulgare Mill. Apiaceae Leaf Ethanol 100 n.d. Cx. quinquefasciatus 4 Modise and Ashafa,
2016
Fumaria indica Papaveraceae Leaf Water 158; 144; 304; 289; 320 Ae. albopictus, An. 3 Benelli et al., 2017c
(Hausskn.) Pugsley 172 subpictus, Cx.
Galatella villosa (L.) Asteraceae Stem Methanol 395 480 Cx. quinquefasciatus 4 Pavela, 2008
Rchb. f.
Garcinia mangostana L. Clusiaceae Fruit - Ethanol 5.5 22 Ae. aegypti 3 and 4 Torres et al., 2015
pericarp
Fruit - crown Hexane 25 64 Ae. aegypti 3 and 4 Torres et al., 2015
Gardenia carinata Griff. Rubiaceae Leaf Distilled water 40700 n.d. Cx. quinquefasciatus 3 Rawani et al., 2014b
Gilia capitata Sims Polemoniaceae Stem Methanol 687 > 1000 Cx. quinquefasciatus 4 Pavela, 2009
Glebionis coronaria (L.) Asteraceae Flower Methanol 53 110 Cx. quinquefasciatus 4 Pavela, 2008
Cass. ex Spach
Gloriosa superba L. Colchicaceae Leaf Acetone 18; 87 63; 313 An. subpictus, Cx. 4 Bagavan et al., 2009
Gluta renghas L. Anacardiaceae Leaf Acetone 2854 3980=LC95 Ae. aegypti 3 and 4 Zuharah et al., 2014
Gmelina asiatica L. Lamiaceae Leaf Distilled water 130; 114; 230; 202; 244 Ae. aegypti, An. 3 Muthukumaran et al.,
139 stephensi, Cx. 2015b
Gossypium herbaceum L. Malvaceae Leaf Methanol 216; 299 620; 1222 Ae. aegypti, An. 3 and 4 Mohankumar et al.,
Gossypium hirsutum L. Malvaceae Leaf Water 212; 233 529; 625 Ae. aegypti, An. 4 Patil et al., 2014
stephensi resp.
Leaf Ethanol 299 604 An. stephensi
Guarea kunthiana A. Meliaceae Seed Ethanol 169.93 496.11 Ae. aegypti 3 Sarmento et al., 2016
Juss.
Guettarda grazielae Rubiaceae Stem Ethanol - Ethyl 52 n.d. Ae. aegypti 4 Oliveira et al., 2010
M.R.Barbosa acetate fraction
Stem bark Ethanol - Ethyl 86 n.d. Ae. aegypti
acetate fraction
Leaf Ethanol - Hexane 132 n.d. Ae. aegypti
fraction
Gymnema sylvestre Apocynaceae Leaf Petroleum ether 166; 187 n.d. An. subpictus, Cx. 4 Khanna et al., 2011
(Retz.) Schultes quinquefasciatus resp.
Habenaria plantaginea Orchidaceae Leaf Distilled water 103; 112; 202; 216; 231; An. stephensi, Ae. 3 Aarthi et al., 2018
Lindl. 123; 124; 243; 266; 280 aegypti, Cx.
137; 150 Quinquefasciatus, An.
subpictus, Ae.
albopictus, Cx.
tritaeniorhynchus
Halodule uninervis Cymodoceaceae Leaf Distilled water 296 663 Ae. aegypti 4 Mahyoub et al., 2017
(Forssk.) Boiss.
Halophila ovalis (R. Br.) Hydrocharitaceae Whole plant Methanol - 1650 n.d. Cx. quinquefasciatus 3 and 4 Devi et al., 1997
Hook. f. Petroleum ether
fraction
Hedychium coronarium Zingiberaceae Rhizome Distilled water 0.7; 0.8; 24; 40; 53; 73 Ae. aegypti 1-4 Kalimuthu et al.,
J.Koenig 1.0; 1.1, 2017
LC50 in %
Hedyotis puberula Rubiaceae Leaf Distilled water 183; 199; 370; 385; 404 An. stephensi, Ae. 3 Azarudeen et al.,
(G.Don) R.Br. ex 218 aegypti, Cx. 2016
Arn. quinquefasciatus
Heliotropium indicum L. Boraginaceae Leaf Distilled water 73; 69; 79 127; 121; 134 Ae. aegypti, An. 3 Veerakumar et al.,
stephensi, Cx. 2014a
Hemidesmus indicus (L.) Apocynaceae Leaf Ethanol 4800; 5100; 11300; 11700; Ae. aegypti, An. 3 Arjunan et al., 2012
R. Br. ex Schult. 4300 10500 stephensi, Cx.
Leaf Methanol 155 608 An. stephensi 4 Kamaraj et al., 2010b
Leaf Acetone 109 646 Cx. quinquefasciatus
245
Table 1 (continued)
plant instar
Hibiscus rosa-sinensis L. Malvaceae Leaf Methanol 295; 390 502; 613 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Hiptage benghalensis (L.) Malpighiaceae Root bark Acetone 13; 15; 17 n.d. Ae. albopictus, 3 Lalrotluanga et al.,
Kurz Anopheles barbirostris, 2012
Cx. quinquefasciatus
resp.
Holarrhena Apocynaceae Bark Methanol 715 2010 Cx. quinquefasciatus 3 Kumar et al., 2018
antidysenterica Bark Hexane 739 1708 Ae. aegypti
(Roth) Wall. ex
A.DC.
Holostemma ada-kodien Apocynaceae Leaf Distilled water 186; 204; 374; 395; 417 An. stephensi, Ae. 3 Alyahya et al., 2018
Schult. 222 aegypti, Cx.
quinquefasciatus
Hoslundia opposita Vahl Lamiaceae Leaf Petroleum ether 171 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Root bark Methanol 368 n.d. An. gambiae
Hugonia mystax Cav. Linaceae Leaf Distilled water 162; 182; 326; 356; 375 An. stephensi, Ae. 3 Govindarajan et al.,
199 aegypti, Cx. 2017b
quinquefasciatus
Humulus japonicus Sieb. Cannabaceae Leaf Methanol 25 87 Cx. quinquefasciatus 4 Pavela, 2008
& Zucc.
Hymenodictyon orixense Rubiaceae Leaf Distilled water 105, 114, 206, 219, 234 An. subpictus, Ae. 3 Govindarajan and
(Roxb.) Mabb. 124 albopictus, Cx. Benelli, 2016c
Hydrocotyle javanica Araliaceae Leaf Ethyl acetate 88; 97 561; 303 An. stephensi, Cx. 4 Kamaraj et al., 2010b
Thunb. quinquefasciatus resp.
Hypericum perforatum L. Hypericaceae Flower Methanol 496 980 Cx. quinquefasciatus 4 Pavela, 2009
Hypericum polyanthemum Hypericaceae Aerial part N-hexane - 121 n.d. Ae. aegypti 4 da Silva et al., 2013
Klotzsch ex H.Reich. Benzopyran
enriched fraction
Hyptis suaveolens (L.) Lamiaceae Leaf Acetone 258 n.d. Ae. albopictus 3 Yadav et al., 2014
Poit. Leaf Methanol 327; 392 720; 844 Ae. aegypti, An. 3 and 4 Mohankumar et al.,
Leaf Acetone 820 3030 Ae. albopictus 3 Yadav et al., 2015
Whole plant Methanol 87 415 Cx. quinquefasciatus 3 Kovendan et al.,
2012d
Hyssopus officinalis L. Lamiaceae Stem Methanol 150 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Ichnocarpus frutescens Apocynaceae Leaf Distilled water 208; 186; 396; 365; 418 Ae. albopictus, An. 3 Govindarajan et al.,
(L.) R. Br. 226 subpictus, Cx. 2016a
Imperatoria ostruthium L. Apiaceae Root Methanol 49 83 Cx. quinquefasciatus 4 Pavela, 2009
Indonesiella echioides (L.) Acanthaceae Leaf Petroleum ether 244 n.d. Cx. quinquefasciatus 4 Karmegam et al.,
Streem 1997
Inula britannica L. Asteraceae Stem Methanol 382 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Inula helenium L. Asteraceae Stem Methanol 420 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Inula racemosa Hook.f. Asteraceae Root Ethanol 25 95=LC95 Ae. albopictus 4 He et al., 2014
Ipomoea cairica (L.) Convolvulaceae Leaf Acetone 102; 106 n.d.; n.d. Ae. aegypti, Ae. 3 Zuharah et al., 2016
Sweet albopictus resp.
Leaf Methanol 884 1769 Cx. quinquefasciatus 3 Samuel et al., 2014
Flower Methanol 76 216 Cx. quinquefasciatus
Petal Methanol 18; 32 180; 174 Ae. aegypti, Ae. 3 Ishak et al., 2014
albopictus resp.
Root Methanol 38; 121 38; 115 Ae. aegypti, Ae.
albopictus resp.
Leaf Methanol 12; 22 83; 118 Ae. aegypti, Ae.
albopictus resp.
Leaf Methanol 115; 122 320; 316=LC95 Ae. aegypti, Ae. 3 AhbiRami et al., 2014
albopictus resp.
Leaf Acetone 102; 106 448; 322=LC95 Ae. aegypti, Ae.
albopictus resp.
Flower Acetone 106; 136 279; 219=LC95 Ae. aegypti, Ae.
albopictus resp.
Flower Methanol 138 307=LC95 Ae. albopictus
Stem Acetone 133; 146 375; 357=LC95 Ae. aegypti, Ae.
albopictus resp.
Ipomoea carnea Jacq. Convolvulaceae Leaf Petroleum ether 42 163 Cx. quinquefasciatus 4 Rahuman et al.,
2009a
Jacobaea maritima (L.) Asteraceae Stem Methanol 270 428 Cx. quinquefasciatus 4 Pavela, 2009
Pelser & Meijden
Jasminum nervosum Lour. Oleaceae Leaf Distilled water 196 585=LC95 Cx. quinquefasciatus 3 Lallawmawma et al.,
2015
246
Table 1 (continued)
plant instar
Jatropha curcas L. Euphorbiaceae Leaf Methanol 113 481 Cx. quinquefasciatus 3 Kovendan et al.,
2012d
Leaf Methanol 68234 126771 Cx. quinquefasciatus 3 Kovendan et al., 2011
1997
Justicia procumbens L. Acanthaceae Leaf Ethyl acetate 12; 21 63; 88 Cx. gelidus, Cx. 4 Kamaraj et al., 2010a
Kaempferia angustifolia Zingiberaceae Root Methanol 88 n.d. Ae. aegypti 3 Sukari et al., 2010
Roscoe
Kaempferia rotunda L. Zingiberaceae Root Chloroform 68 n.d. Ae. aegypti 3 Sukari et al., 2010
Knema attenuata (Hook.f. Myristicaceae Fruit - kernel Ethanol 159; 162 342; 458 Ae. albopictus, An. 3 and 4 Vinayachandra et al.,
& Th.) Warb. (seed) stephensi resp. 2011
Fruit - aril Chloroform 141; 160 290; 445 Ae. albopictus, An.
stephensi resp.
Laburnum anagyroides Fabaceae Leaf Methanol 111 198 Cx. quinquefasciatus 4 Pavela, 2008
Medik.
Lansium domesticum Meliaceae Seed Ethanol 9368 n.d. Ae. aegypti 3 Ni’mah et al., 2015
Corrêa
Lantana camara L. Verbenaceae Leaf Hexane 31 86 Ae. aegypti 4 Kumar et al., 2012c
Root Methanol 28; 133; 12 n.d. Ae. aegypti, An. 4 Tripathy et al., 2011
stephensi, Cx.
Laurencia papillosa (C. Rhodomelaceae Whole plant Diethyl ether 64 246 Cx. pipiens 3 Abou-Elnaga et al.,
Agardh) Greville 2011
Laurus nobilis L. Lauraceae Leaf Methanol 303; 337 814; 1198 Ae. aegypti, An. 3 and 4 Mohankumar et al.,
Lavandula officinalis Lamiaceae Flower Methanol 59 123 Cx. quinquefasciatus 4 Pavela, 2008
Chaix
Lettowianthus stellatus Annonaceae Stem bark Petroleum ether 93 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Diels
Leucas aspera (Willd.) Lamiaceae Leaf Methanol 10 93 Ae. aegypti 4 Suganya et al., 2014
Link Leaf Ethyl acetate 17 31 Ae. aegypti
Leaf and Methanol 108 411 Cx. quinquefasciatus 3 Kovendan et al.,
flower 2012d
Whole plant Ethanol 108230 166770 An. stephensi 3 Kovendan et al.,
2012h
Flower Methanol 53; 81 233; 300 An. subpictus, Cx. 4 Kamaraj et al., 2009
Leucas linifolia (Roth.) Lamiaceae Leaf Methanol 233; 250 306; 315 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Sprengel quinquefasciatus resp.
Linaria genistifolia (L.) Plantaginaceae Stem Methanol 490 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Mill.
Lippia nodiflora (L.) Verbenaceae Leaf Methanol 190; 199 268; 302 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Michx. quinquefasciatus resp.
Lobelia leschenaultiana Campanulaceae Leaf Acetone 22.83 n.d. Ae. aegypti 3 Banumathi et al.,
(C.Presl) Skottsb. 2017b
Lobelia siphilitica L. Campanulaceae Stem Methanol 633 929 Cx. quinquefasciatus 4 Pavela, 2009
Malva sylvestris L. Malvaceae Leaf Distilled water 158; 144; 305; 282; 318 Ae. aegypti, An. 3 Govindarajan et al.,
171 stephensi, Cx. 2016c
Malvastrum Malvaceae Leaf Methanol 337 n.d. Ae. albopictus 3 Yadav et al., 2014
coromandelianum Leaf Acetone 620 1760 Ae. albopictus 3 Yadav et al., 2015
(L.) Garcke
Mangifera indica L. Anacardiaceae Leaf Acetone 630 779=LC95 Ae. aegypti 3 and 4 Zuharah et al., 2014
Marrubium vulgare L. Lamiaceae Leaf Methanol 1893 2924 Cx. pipiens 4 Amel and Sélima,
2015
Matricaria maritima L. Asteraceae Flower Methanol 72 139 Cx. quinquefasciatus 4 Pavela, 2008
Medicago romanica Fabaceae Stem Methanol 482 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Prodan
Melanochyla fasciculiflora Anacardiaceae Leaf Acetone 2338 6481=LC95 Ae. aegypti 3 and 4 Zuharah et al., 2014
Kochummen
Melia dubia Cav. Meliaceae Leaf Methanol 100 127=LC95 Ae. aegypti 3 Deepa et al., 2015
Millettia ferruginea Fabaceae Seed Methanol 15 43 An. arabiensis 3 and 4 Andemo et al., 2014
(Hochst.) Baker
Mentha arvensis L. Lamiaceae Leaf Methanol 90; 99 112; 194 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Merremia aegyptia (L.) Convolvulaceae Leaf Acetone 121 n.d. Ae. aegypti 4 Oliveira et al., 2010
Urban
247
Table 1 (continued)
plant instar
Merremia emarginata Convolvulaceae Leaf Distilled water 158; 147; 302; 286; 316 Ae. aegypti, An. 3 Azarudeen et al.,
(Burm. f.) Hallier f. 169 stephensi, Cx. 2017a
Microdictyon Anadyomenaceae Whole plant Methanol - 50 n.d. Cx. quinquefasciatus 3 and 4 Devi et al., 1997
pseudohapteron Petroleum ether
A.Gepp et E.S.Gepp fraction
Mimosa pudica L. Fabaceae Leaf Distilled water 46; 36 n.d. An. subpictus, Cx. 4 Marimuthu et al.,
Leaf Ethyl acetate 54; 57 271; 284 Cx. gelidus, Cx. 4 Kamaraj et al., 2010a
Mimusops elengi L. Sapotaceae Leaf Distilled water 169-302; 418-683; 394- An. stephensi, Ae. 1-4 Subramaniam et al.,
155-278 696 albopictus 2015
Momordica charantia L. Cucurbitaceae Fruit Hexane 260 663 Ae. aegypti 4 Kumar et al., 2012c
Fruit Petroleum ether 28; 41 155; 270 An. stephensi, Cx. 3 Maurya et al., 2009
Leaf Methanol 199; 208 780; 787 Ae. aegypti, Cx. 4 Rahuman and
Morinda citrifolia L. Rubiaceae Leaf Methanol 278; 262; 568; 505; 620 Ae. aegypti, An. 3 Kovendan et al.,
Rubiaceae Leaf Distilled water 325 644 Cx. quinquefasciatus
Morinda tinctoria Roxb. Rubiaceae Leaf Acetone 8 n.d. Cx. quinquefasciatus 3 Kumar et al., 2015
Moringa oleifera Lam. Moringaceae Flower Distilled water 16700 n.d. Ae. aegypti 3 Pontual et al., 2012
Root Methanol 283; 318 498; 487 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Moschosma polystachyon Lamiaceae Leaf Methanol 153 n.d. Cx. quinquefasciatus 3 Rajkumar and
(L.) Benth. Jebanesan, 2004
Murraya koenigii (L.) Rutaceae Leaf Methanol 461; 407 1022; 908 Ae. aegypti, An. 3 Suganya et al., 2013
Jack. stephensi resp.
Rutaceae Leaf Ethyl acetate 858 1564 Cx. quinquefasciatus 3 Kovendan et al.,
2012b
Musa paradisiaca L. Musaceae Flower Chloroform 35 312 An. vagus 4 Bagavan and
Flower Methanol 49; 103 188; 340 Ar. subalbatus, Cx. Rahuman, 2011
vishnui resp.
Stem juice Distilled water 117; 137; 279; 330; 379; An. stephensis 1-4 Anbazhagan et al.,
163; 190 414 2017
Mussaenda glabra Vahl Rubiaceae Leaf Distilled water 81, 88, 95 161, 170, 177 An. subpictus, Ae. 3 Govindarajan et al.,
albopictus, Cx. 2016h
Myristica fragrans Houtt. Myristicaceae Mace Ethanol 75 124=LC95 Ae. aegypti 4 Intirach et al., 2016
Leaf Methanol 162, 194, 502, 543, 605, Ae. aegypti 1-4 Ashokan et al., 2017
240, 274 661
Naregamia alata Wight & Meliaceae Leaf Distilled water 165; 179; 322; 342; 372 An. stephensis, Ae. 3 Azarudeen et al.,
Arn. 196 aegypti, Cx. 2017b
quinquefasciatus
Nelumbo nucifera Gaertn. Nelumbonaceae Leaf Ethyl acetate 35; 37 173; 177 An. stephensi, Cx. 4 Kamaraj et al., 2011a
Leaf Methanol 8.9; 9.5 29; 28 An. subpictus, Cx. 4 Santhoshkumar et al.,
Nerium oleander L. Apocynaceae Leaf Distilled water 319 640 An. stephensi 3 Roni et al., 2013
Nicandra physalodes (L.) Solanaceae Leaf Distilled water 203, 223, 405, 430, 449 An. stephensi, Ae. 3 Govindarajan et al.,
Gaertn. 242 aegypti, Cx. 2016f
Nicotiana tabacum L. Solanaceae Leaf Distilled water (hot) 76 335 Cx. quinquefasciatus 4 Rahuman et al.,
2009b
Nyctanthes arbortistis L. Oleaceae Leaf Petroleum ether 180; 185; 340; 320; 335 Ae. aegypti, An. 4 Alam et al., 2011
160 stephensi, Cx.
Leaf Dichloromethane 261; 115 588; 367 Ae. aegypti, An. 4 Patil et al., 2010
stephensi resp.
Leaf Chloroform 526; 781 1670; 2022 Ae. aegypti, An. 4 Mathew et al., 2009
stephensi resp.
Flower Methanol 679 1218 An. stephensi
Ocimum basilicum L. Lamiaceae Leaf Ethanol 46 105 An. stephensi 3 Murugan et al., 2015c
Stem Methanol 32 69 Cx. quinquefasciatus 4 Pavela, 2008
Leaf Methanol 5.785, n.d. An. stephensi 2, 3 Aarthi and Murugan,
5.967 and and 4 2010
6.369%
Ocimum canum Sims Lamiaceae Leaf Methanol 43; 30; 28 61; 48; 44 Ae. aegypti, An. 4 Murugan et al., 2016
stephensi, Cx.
Leaf Ethyl acetate 102 492 An. stephensi 4 Kamaraj et al., 2008
Flower Ethyl acetate 29 168 An. stephensi
248
Table 1 (continued)
plant instar
Ocimum sanctum L. Lamiaceae Leaf Acetone 156 799 An. stephensi 4 Kamaraj et al., 2008
Ocotea velloziana Lauraceae Trunk bark Ethanol 214 n.d. Ae. aegypti 3 Garcez et al., 2009
(Meisn.) Mez
Origanum vulgare L. Lamiaceae Stem Ethanol 256 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Ormocarpum Fabaceae Leaf Distilled water 178; 165; 343; 324; 364 Ae. aegypti, An. 3 Govindarajan and
cochinchinense 193 stephensi, Cx. Benelli, 2016a
(Lour.) Merr. quinquefasciatus resp.
Ormosia arborea (Vell.) Fabaceae Leaf Ethanol 238 347 Ae. aegypti 3 Porto et al., 2017
Harms Seed Ethanol 111 194 Ae. aegypti
Orthosiphon thymiflorus Lamiaceae Leaf Methanol 150; 119; 426; 373; 411 Ae. aegypti, An. 3 Kovendan et al.,
(Roth) Sleesen 137 stephensi, Cx. 2012e
Otanthus maritimus (L.) Asteraceae Stem Methanol 7 15 Cx. quinquefasciatus 4 Pavela, 2008
Hoffmanns. & Link
Padina australis Hauck Dictyotaceae Whole plant Methanol - 341 n.d. Ae. aegypti 4 Yu et al., 2015
Chloroform fraction
Parthenium hysterophorus Asteraceae Leaf Methanol 1880 15210 An. gambiae 3 Wachira et al., 2014
L. Leaf Distilled water 74; 59 n.d. Ae. aegypti, Cx. 3 Thandapani et al.,
quinquefasciatus 2018
Passiflora foetida L. Passifloraceae Leaf Methanol 172; 441 301; 922 Ae. aegypti, An. 3 and 4 Mohankumar et al.,
Pavonia zeylanica (L.) Malvaceae Leaf Methanol 143; 193 496; 565 An. stephensi, Cx. 4 Kamaraj et al., 2010b
Cav. quinquefasciatus resp.
Pedalium murex L. Pedaliaceae Seed Distilled water 125 234 Ae. aegypti 4 Ishwarya et al., 2017
Pedilanthus tithymaloides Euphorbiaceae Leaf Distilled water 17720 n.d. Ae. aegypti 3 Sundaravadivelan
L. et al., 2013
Leaf Ethanol 260 450 Cx. quinquefasciatus 3 Kamalakannan et al.,
2010
Pelargonium graveolens Geraniaceae Leaf Methanol 96 118=LC95 Ae. Aegypti 3 Deepa et al., 2015
(Thunb.) L'Hér.
Pereskia bleo (Kunth) DC. Cactaceae Fruit - Ethanol 1095 n.d. Ae. aegypti 3 Thongwat et al., 2014
endocarp
Pergularia daemia Apocynaceae Leaf Hexane 14 57 An. subpictus 4 Bagavan et al., 2009
(Forsskal) Chiov. Leaf Ethyl acetate 32 115 Cx. tritaeniorhynchus
Persea americana P. Mill. Lauraceae Seed Hexane 9.8 22 Ae. aegypti 3 Torres et al., 2014
Fruit - peel Ethanol 10 26 Ae. aegypti
Fruit - pulp Ethanol 21 59 Ae. aegypti
Pergulariaextensa (Jacq.) Asclepiadaceae Root Ethanol 63 143 An. stephensi 3 Abirami et al., 2017
N.E. Br.
Phyllanthus emblica L. Phyllanthaceae Fruit Hexane 299 454 Ae. aegypti 4 Kumar et al., 2012c
Leaf Methanol 55 200 Cx. tritaeniorhynchus 4 Bagavan et al., 2009
Physalis alkekengi L. Solanaceae Stem Methanol 899 > 1000 Cx. quinquefasciatus 4 Pavela, 2009
Piper aduncum L. Piperaceae Leaf Chloroform 192 346 Ae. aegypti 3 Porto et al., 2017
Piper hispidum Sw. Piperaceae Leaf Ethanol 169 474 Ae. aegypti 3 Porto et al., 2017
Piper longum L. Piperaceae Seed Methanol 55; 70 92; 114 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Fruit Ethanol 2.2 4.8=LC95 Ae. aegypti 4 Chaithong et al.,
2006
Piper nigrum L. Piperaceae Unripe fruit Ethanol 1.8 2.8 Ae. aegypti 3 Grzybowski et al.,
2012
Seed Ethyl acetate 25; 44 108; 217 An. stephensi, Cx. 4 Kamaraj et al., 2011a
Seed Methanol 0.6; 0.7 6.8; 8.4 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Piper ribesioides Wall. Piperaceae Wood Ethanol 8.1 14=LC95 Ae. aegypti 4 Chaithong et al.,
2006
Piper sarmentosum Roxb. Piperaceae Whole plant Ethanol 4.1 12.1=LC95 Ae. aegypti 4 Chaithong et al.,
2006
Pithecellobium dulce Fabaceae Leaf Methanol 156; 145 280; 251 Ae. aegypti, An. 3 Govindarajan et al.,
(Roxb.) Benth. Seed Methanol 194; 168 377; 315 stephensi resp. 2013
Plectranthus amboinicus Lamiaceae Leaf Methanol 47 123 Ae. aegypti 3 Murugan et al., 2013
(Lour.) Spreng.
Plumbago zeylanica L. Plumbaginaceae Root Ethanol 16 50 Ae. aegypti 3 Patil et al., 2011
Root Methanol 170; 222 415; 499 Ae. aegypti, An. 4 Patil et al., 2010
stephensi resp.
Polyalthia tanganyikensis Annonaceae Root bark Methanol 70 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Vollesen
Prosopis juliflora (Sw.) Fabaceae Leaf Methanol 80 n.d. Ae. albopictus 3 Yadav et al., 2014
DC. Leaf Methanol 440 1850 Ae. albopictus 3 Yadav et al., 2015
Leaf Methanol 128; 96; n.d. Ae. aegypti, An. 3 and 4 Bansal et al., 2012
119 stephensi, Cx.
249
Table 1 (continued)
plant instar
Psychotria nilgiriensis Deb Rubiaceae Leaf Distilled water 320 664 Ae. aegypti 3 Kovendan et al., 2016
& M.G.Gangop.
Pteridium aquilinum (L.) Dennstaedtiaceae Leaf Ethanol 302 582 An. stephensi 3 and 4 Panneerselvam et al.,
Kuhn 2016
Quassia africana (Baill.) Simaroubaceae Root Methanol 18 n.d. An. gambiae 4 Sama et al., 2014
Baill.
Quercus infectoria Oliv. Fagaceae Gall Methanol - Ethyl 117 145 An. stephensi 4 Aivazi and Vijayan,
acetate fraction 2009
Gall Acetone 556 n.d. Cx. pipiens 4 Redwane et al., 2002
Quisqualis indica L. Combretaceae Leaf Distilled water 186; 203; 367; 388; 413 An. stephensi, Ae. 3 Govindarajan et al.,
220 aegypti, Cx. 2016i
quinquefasciatus
Reseda odorata L. Resedaceae Stem Methanol 381 915 Cx. quinquefasciatus 4 Pavela, 2009
Rhinacanthus nasutus (L.) Acanthaceae Flower Methanol 41; 72 143; 362 An. subpictus, Cx. 4 Kamaraj et al., 2009
Kurz tritaeniorhynchus resp.
Leaf Hexane 173 630 An. stephensi 4 Kamaraj et al., 2008
Rhizophora apiculata Rhizophoraceae Stilt root Petroleum ether 26 n.d. Cx. quinquefasciatus 4 Thangam and
Blume Kathiresan, 1997
Rhizophora mucronata Rhizophoraceae Stilt root Petroleum ether 69 n.d. Cx. quinquefasciatus 4 Thangam and
Lamarck ex Poiret Kathiresan, 1997
Ricinus communis L. Euphorbiaceae Leaf Methanol 400 1130 An. gambiae 3 Wachira et al., 2014
Leaf Hexane 64 140 Ae. aegypti 4 Kumar et al., 2012c
Seed Acetone 17; 12; 7.1 n.d. Ae. albopictus, An. 4 Mandal, 2010
stephensi, Cx.
Rourea doniana Baker Connaraceae Leaf Ethanol - 170 n.d. Ae. aegypti 4 Oliveira et al., 2010
Chloroform fraction
Stem Ethanol - Hexane 12 n.d. Ae. aegypti
fraction
Rubia tinctorum L. Rubiaceae Stem Methanol 725 943 Cx. quinquefasciatus 4 Pavela, 2009
Rubus ellipticus Sm. Rosaceae Leaf Distilled water 184; 167; 354; 329; 377 Ae. aegypti, An. 3 AlQahtani et al.,
202 stephensi, Cx. 2017
Ruellia tuberosa L. Acanthaceae Leaf Distilled water 45200 n.d. Cx. quinquefasciatus 3 Rawani et al., 2014b
Salicornia fruticosa (L.) Chenopodiaceae Leaf Distilled water 2.4 6.4 Cx. pipiens 4 Mahyoub et al., 2016
A.J. Scott
Salvadora persica Wall. Salvadoraceae Leaf Methanol - 850 n.d. Cx. quinquefasciatus 3 and 4 Devi et al., 1997
Chloroform fraction
Salvia farinacea Benth. Lamiaceae Stem Methanol 195 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Salvia officinalis L. Lamiaceae Stem Methanol 159 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Salvia verbenaca L. Lamiaceae Stem Methanol 311 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Salvia verticillata L. Lamiaceae Stem Methanol 410 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Salvia viridis L. Lamiaceae Flower Methanol 110 183 Cx. quinquefasciatus 4 Pavela, 2008
Sargassum binderi Sonder Sargassaceae Whole Methanol - 192 n.d. Ae. aegypti 4 Yu et al., 2015
ex J.Agardh seaweed Chloroform fraction
Sargassum muticum Sargassaceae Whole Distilled water 12-22, 17- 39-55, 43-58, Ae. aegypti, An. 1-4 Madhiyazhagan
(Yendo) Fensholt seaweed 24, 23-30 52-69, stephensi, Cx. et al., 2015
Sargassum wightii Sargassaceae Whole Ethanol 57; 62; 68; 117; 122; 129; An. stephensi 1-4 Murugan et al., 2018
Greville ex J.Agardh seaweed 79 145
Sapindus emarginatus Sapindaceae Seed Aqueous 2500; 2380; n.d. Ae. aegypti, An. 3 Koodalingam et al.,
Vahl (physiological 2810 stephensi, Cx. 2009
saline) quinquefasciatus resp.
Saponaria officinalis L. Caryophyllaceae Stem Methanol 516 928 Cx. quinquefasciatus 4 Pavela, 2009
Satureja hortensis L. Lamiaceae Stem Methanol 28 56 Cx. quinquefasciatus 4 Pavela, 2008
Schisandra chinensis Schisandraceae Leaf Methanol 15 33 Cx. quinquefasciatus 4 Pavela, 2009
(Turcz.) Baill.
Scindapsus officinalis Araceae Fruit Petroleum ether 3.2 8.6 Ae. aegypti 4 Sreedev et al., 2014
(Roxb.) Schott
Scrophularia nodosa L. Scrophulariaceae Stem Methanol 982 > 1000 Cx. quinquefasciatus 4 Pavela, 2009
Senecio laetus Edgew. Asteraceae Leaf Petroleum ether 67; 58; 83 257; 246; 281 Ae. aegypti, An. 4 Ali et al., 2018
stephensi, Cx.
Leaf Methanol 83 253 Ae. aegypti
Root Methanol 22; 17; 31 145; 109; 184 Ae. aegypti, An.
stephensi, Cx.
Senna didymobotrya Fabaceae Leaf Methanol 530 1370 An. gambiae 3 Wachira et al., 2014
(Fresen.) Irwin &
Barneby
Senna occidentalis (L.) Fabaceae Leaf Ethanol 52 123 An. stephensi 3 Murugan et al., 2015c
Link
Sesamum indicum L. Pedaliaceae Leaf Methanol 106 139=LC95 Ae. aegypti 3 Deepa et al., 2015
250
Table 1 (continued)
plant instar
Sesbania grandiflora (L.) Fabaceae Leaf Methanol 2080 6200 An. stephensi 4 Elango et al., 2011
Poir.
Seseli diffusum (Roxb. ex Apiaceae Seed Ethanol 126 n.d. Ae. aegypti 4 Kabir et al., 2013
Sm.) Santapau &
Wagh
Seseli pallasii Besser Apiaceae Stem Methanol 6 10 Cx. quinquefasciatus 4 Pavela, 2009
Seseli tortuosum L. Apiaceae Stem Methanol 428 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Sida acuta Burm. f. Malvaceae Leaf Distilled water 119; 110; 214; 202; 228 Ae. aegypti, An. 3 Veerakumar et al.,
130 stephensi, Cx. 2013
Sideritis euxina Juz. Lamiaceae Stem Methanol 250 425 Cx. quinquefasciatus 4 Pavela, 2008
Solanum nigrum L. Solanaceae Leaf Benzene 16 59 Cx. vishnui 3 Rawani et al., 2014a
Fruit Hexane 18; 16; 9; 6; 65; 95; 49; 43; Ae. aegypti, An. 3 and 4 Raghavendra et al.,
12 56 culicifacies species A, 2009
An. culicifacies species
C, An. stephensi, Cx.
Solanum torvum Sw. Solanaceae Leaf Methanol 30; 21 133; 83 An. stephensi, Cx. 4 Kamaraj et al., 2010b
Seed Methanol 31 131 An. stephensi
Leaf Methanol 35; 44 152; 185 An. subpictus, Cx. 4 Kamaraj et al., 2009
Seed Methanol 29; 84 121; 351 An. subpictus, Cx.
Solanum variabile Mart. Solanaceae Leaf Ethanol 188 284 Ae. aegypti 3 Porto et al., 2017
Solanum villosum Mill. Solanaceae Berry Chloroform : 12 n.d. Ae. aegypti 3 Chowdhury et al.,
Methanol (1:1) 2008
Solanum violaceum Solanaceae Leaf Petroleum ether 36.92 n.d. Ae. aegypti 3 Banumathi et al.,
Ortega 2017b
Solanum xanthocarpum L. Solanaceae Fruit Methanol 122; 78; 1262; 466; 1004 Ae. aegypti, An. 3 and 4 Bansal et al., 2015
143 stephensi, Cx.
Leaf Ethanol 271 1012 Cx. quinquefasciatus 3 Kumar et al., 2012a
Fruit Methanol 221 410 Ae. aegypti 3 Kumar et al., 2012b
Fruit Ethanol 788; 573 1289; 1067 Ae. aegypti, Cx. 3 and 4 Changbunjong et al.,
Fruit Ethanol 378; 132; 1633; 420; 787; Ae. aegypti, Anopheles 3 and 4 Bansal et al., 2009a
196; 520 2860 culicifacies, An.
stephensi, Cx.
Fruit without Ethanol 273; 80; 1063; 254; 403; Ae. aegypti, Anopheles
seed 131; 385 1936 culicifacies, An.
stephensi, Cx.
Seed Ethanol 291; 92; 866; 379; 490; Ae. aegypti, Anopheles
187; 451 1881 culicifacies, An.
stephensi, Cx.
Seed Methanol 124; 67; 74; 364; 194; 241; Ae. aegypti, Anopheles 3 and 4 Bansal et al., 2009b
155 501 culicifacies, An.
stephensi,Cx.
Fruit (green Distilled water 498; 113; 1414; 432; 2228 Ae. aegypti, An.
unripe) 846 stephensi,Cx.
Fruit (yellow Methanol 118; 52; 52; 233; 159; 177; Ae. aegypti, Anopheles
ripe) 157 529 culicifacies, An.
stephensi,Cx.
Fruit Carbon 5.1 n.d. An. stephensi 3 Mohan et al., 2005
tetrachloride
Fruit Petroleum ether 63 n.d. Cx. quinquefasciatus
Solidago canadensis L. Asteraceae Leaf Petroleum ether 16; 35; 28 61; 101; 96 Ae. aegypti, An. 4 Prathibha et al., 2014
stephensi, Cx.
Leaf Petroleum ether 22 44 Ae. aegypti 4 Raghavendra et al.,
2013
Sonchus arvensis L. Asteraceae Stem Methanol 68 118 Cx. quinquefasciatus 4 Pavela, 2008
Sonneratia alba Sm. Lythraceae Leaf Distilled water 192; 222; 443; 491; 524; Ae. aegypti 1-4 Murugan et al., 2017
264; 325 666
Sonneratia caseolaris (L.) Lythraceae Leaf Methanol - 1350 n.d. Cx. quinquefasciatus 3 and 4 Devi et al., 1997
Engl. Petroleum ether
fraction
Spermacoce hispida L. Rubiaceae Root Ethanol 37 125 An. stephensi 3 Abirami et al., 2017
251
Table 1 (continued)
plant instar
Spermacoce latifolia Aubl. Rubiaceae Leaf Methanol 625 1122 Ae. aegypti 3 Porto et al., 2017
Leaf Methanol - Hexane 415 901 Ae. aegypti
fraction
Spermacoce verticillata L. Rubiaceae Stem Ethanol - Hexane 116 n.d. Ae. aegypti 4 Oliveira et al., 2010
fraction
Aerial part Ethanol - Hexane 84 n.d. Ae. aegypti
fraction
Root Ethanol - Hexane 92 n.d. Ae. aegypti
fraction
Sphaeranthus indicus L. Asteraceae Leaf Ethyl acetate 275 1081 Cx. quinquefasciatus 3 Kovendan et al.,
2012b
Spilanthes acmella (L.) L. Asteraceae Flower Hexane - Ethyl 0.9; 4.6; 3.1 1.9; 7.8; 8.9 Anopheles culicifacies, 3 and 4 Pandey et al., 2013
acetate An. stephensi, Cx.
Spilanthes acmella var. Asteraceae Flower Hexane 0.9; 4.6; 3.1 1.9; 7.8; 8.9 Anopheles culicifacies, 3 and 4 Pandey et al., 2007
oleracea (L.) An. stephensi, Cx.
C.B.Clarke quinquefasciatus resp.
Spilanthes calva DC. Asteraceae Flower Hexane 0.9; 5.1; 3.5 2; 8.5; 9.9 Anopheles culicifacies, 3 and 4 Pandey et al., 2007
An. stephensi, Cx.
Spilanthes mauritiana Asteraceae Flower Ethyl acetate 14; 12; 5 33; 28; 11 Ae. aegypti, An. 4 Prathibha et al., 2014
(A.Rich. ex Pers.) stephensi, Cx.
DC. quinquefasciatus resp.
Spilanthes paniculata Asteraceae Flower Hexane 3.2; 5.1; 3.4 7.1; 13.6; 6.3 Anopheles culicifacies, 3 and 4 Pandey et al., 2007
Wall. ex DC. An. stephensi, Cx.
Spondias mombin L. Anacardiaceae Leaf MEthanol - Hexane 23; 92; 327 96; 245; 795 Ae. aegypti, An. 4 Eze et al., 2014
fraction gambiae, Cx.
Stachys byzantina K. Lamiaceae Stem Methanol 65 133 Cx. quinquefasciatus 4 Pavela, 2009
Koch ex Scheele
Stachys cretica L. Lamiaceae Stem Methanol 292 430 Cx. quinquefasciatus 4 Pavela, 2008
Suaeda maritima (L.) Amaranthaceae Leaf Ethanol 135.034, 329.083, Ae. aegypti 1, 2, 3 Suresh et al., 2018
Dumort. 162.454, 380.995, and 4
183.604 399.344 and
and 436.988
210.809
Syringodium isoetifolium Cymodoceaceae Whole plant Methanol - 450 n.d. Cx. quinquefasciatus 3 and 4 Devi et al., 1997
(Aschers.) Dandy Petroleum ether
fraction
Syzygium aromaticum (L.) Myrtaceae Flower Hexane 86; 150 409; 996 An. vagus, Cx. vishnui 4 Bagavan and
Merr. & Perry resp. Rahuman, 2011
Flower Ethyl acetate 49 188 Ar. subalbatus
Tabebuia avellanedae Bignoniaceae Trunk bark Ethanol 442 n.d. Ae. aegypti 3 Garcez et al., 2009
Lorentz ex Griseb
Tagetes minuta L. Asteraceae Leaf Hexane 1010 n.d. Cx. quinquefasciatus 4 Modise and Ashafa,
2016
Aerial part Ethanol 1 1.5 Ae. fluviatilis 4 Macedo et al., 1997
Tagetes patula L. Asteraceae Flower Methanol 85; 118 185; 261 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Tanacetum vulgare L. Asteraceae Flower Methanol 178 198 Cx. quinquefasciatus 4 Pavela, 2008
Tephrosia purpurea (L.) Fabaceae Seed Methanol 63; 75; 47 n.d. Ae. aegypti, An. 3 and 4 Bansal et al., 2012
Pers. gambiae, Cx.
Tephrosia vogelii Hook. f. Fabaceae Leaf Distilled water 1.2 n.d. Ae. albopictus 4 Li et al., 2015
Terminalia chebula Retz. Combretaceae Seed Methanol 138 220 Ae. aegypti 4 Thanigaivel et al.,
2017
Terminalia fagifolia Mart. Combretaceae Trunk Ethanol 373 n.d. Ae. aegypti 3 Garcez et al., 2009
Tessmannia densiflora Fabaceae Stem bark Chloroform 104 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Harms Root bark Chloroform 162 n.d. An. gambiae
Tessmannia martiniana Fabaceae Stem bark Chloroform 83 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
var. martiniana Root bark Methanol 114 n.d. An. gambiae
Harms
Tessmannia martiniana Fabaceae Stem bark Chloroform 256 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
var pauloi Harms Root bark Methanol 148 n.d. An. gambiae
Teucrium chamaedrys L. Lamiaceae Stem Methanol 710 957 Cx. quinquefasciatus 4 Pavela, 2009
Stem Methanol 320 > 500 Cx. quinquefasciatus 4 Pavela, 2008
Teucrium hircanicum L. Lamiaceae Stem Methanol 316 688 Cx. quinquefasciatus 4 Pavela, 2009
Thespesia populnea (L.) Malvaceae Leaf Methanol 353; 841 969; 1344 Ae. aegypti, An. 3 and 4 Mohankumar et al.,
Soland. ex Correa stephensi resp. 2016
Thevetia peruviana (Pers.) Apocynaceae Leaf Methanol 101 131=LC95 Ae. aegypti 3 Deepa et al., 2015
K. Schum.
252
Table 1 (continued)
plant instar
Thymus serphyllum L. Lamiaceae Stem Methanol 711 > 1000 Cx. quinquefasciatus 4 Pavela, 2009
Thymus vulgaris L. Lamiaceae Stem Methanol 48 98 Cx. quinquefasciatus 4 Pavela, 2008
Tinospora cordifolia Menispermaceae Leaf Distilled water 54; 59 n.d. An. subpictus, Cx. 4 Jayaseelan et al.,
(Willd.) Hook. f. & quinquefasciatus resp. 2011
Thomson
Tithonia diversifolia Asteraceae Leaf Methanol 740 2270 An. gambiae 3 Wachira et al., 2014
(Hemsl.) Gray
Trachyspermum ammi Apiaceae Fruit Hexane 66 109 Ae. aegypti 4 Kumar et al., 2012c
(L.) Sprague ex
Turrill
Toddalia asiatica (L.) Rutaceae Leaf Petroleum ether 115 n.d. Cx. quinquefasciatus 4 Karmegam et al.,
Lam. 1997
Trema orientalis (L.) Cannabaceae Leaf Distilled water 17900 n.d. Cx. quinquefasciatus 3 Rawani et al., 2014b
Blume
Tribulus terrestris L. Zygophyllaceae Leaf Petroleum ether 65 n.d. Ae. aegypti 3 El-Sheikh et al., 2016
Leaf Methanol 197; 124 721; 545 Ae. aegypti, Cx. 3 and 4 Bansal et al., 2014
Leaf Petroleum ether 176; 186 739; 597 Ae. aegypti, An.
stephensi resp.
Fruit Methanol 103; 71 346; 240 Ae. aegypti, An.
stephensi resp.
Fruit Acetone 132 638 Cx. quinquefasciatus
Trichodesma africanum Boraginaceae Flower Methanol 4.1 8.1 Ae. caspius 4 Al-Mekhlafi et al.,
(L.) Sm. Leaf Methanol 2.6 4.5 2013
Stem Methanol 5.4; 1.2 8.1; 2.1 Cx. pipiens, Ae. caspius
resp.
Trichosanthes anguina L. Cucurbitaceae Leaf Acetone 554; 842 2235; 3399 Ae. aegypti, Cx. 4 Rahuman and
Tridax procumbens L. Asteraceae Leaf Acetone 52; 69 227; 287 An. subpictus, Cx. 4 Kamaraj et al., 2011b
Triplaris americana L Polygonaceae Stem Ethanol - Hexane 118 n.d. Ae. aegypti 4 Oliveira et al., 2010
fraction
Root Ethanol - Hexane 98 n.d. Ae. aegypti
fraction
Fruit Ethyl acetate - 135 n.d. Ae. aegypti
Chloroform fraction
Tropaeolum majus L. Tropaeloceae Leaf Methanol 780 > 1000 Cx. quinquefasciatus 4 Pavela, 2009
Turnera ulmifolia L. Passifloraceae Leaf Ethanol 242 899 Ae. aegypti 3 Porto et al., 2017
Ulva lactuca L. Ulvaceae Whole Ethanol 30 76 An. stephensi 3 Murugan et al.,
seaweed 2015b
Whole Distilled water 155 284 Ae. aegypti 4 Ishwarya et al., 2018
seaweed
Umbilicaria aprina Nyl. Umbilicariaceae Whole plant Methanol - 1150 n.d. Cx. quinquefasciatus 3 and 4 Devi et al., 1997
Petroleum ether
fraction
Uvaria faulknerae Verdc. Annonaceae Stem bark Chloroform 33 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Uvaria kirkii Oliv. ex Annonaceae Stem bark Petroleum ether 48 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Hook. f. Root bark Petroleum ether 76 n.d. An. gambiae
Uvaria leptocladon Oliv. Annonaceae Stem bark Chloroform 88 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Uvaria lungonyana Annonaceae Stem bark Chloroform 245 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Vollesen Root bark Chloroform 93 n.d. An. gambiae
Uvaria scheffleri Diels Annonaceae Stem bark Petroleum ether 130 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
Root bark Methanol 164 n.d. An. gambiae
Uvariodendron Annonaceae Stem bark Chloroform 56 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
pycnophyllum (Diels) Root bark Chloroform 56 n.d. An. gambiae
R.E. Fr. Root bark Methanol 56 n.d. An. gambiae
Uvariodendron Annonaceae Stem bark Chloroform 188 n.d. An. gambiae 3 and 4 Kihampa et al., 2009
usambarense R.E. Fr. Stem bark Petroleum ether 188 n.d. An. gambiae
Valeriana hardwickii Caprifoliaceae Root Petroleum ether 235; 225; 415; 390; 385 Ae. aegypti, An. 4 Alam et al., 2011
Wall. 180 stephensi, Cx.
Ventilago maderaspatana Rhamnaceae Leaf Distilled water 245; 267; 482; 508; 539 An. stephensi, Ae. 3 Azarudeen et al.,
Gaertn. 290 aegypti, Cx. 2017c
quinquefasciatus
Vernonia ammophila Asteraceae Aerial part Ethanol 40 88 Ae. fluviatilis 4 Macedo et al., 1997
Gardn.
Vernonia cinerea (L.) Asteraceae Leaf Acetone 65 n.d. Ae. albopictus 3 Yadav et al., 2014
Less. Leaf Acetone 220 960 Ae. albopictus 3 Yadav et al., 2015
Verbascum pinnatifidum Scrophulariaceae Methanol 53 110 Cx. quinquefasciatus 4 Pavela, 2008
Vahl
Verbena officinalis L. Verbenaceae Stem Methanol 38 115 Cx. quinquefasciatus 4 Pavela, 2009
253
Table 1 (continued)
plant instar
Vetiveria zizanoides (L.) Poaceae Root Ethanol 1374; 483; n.d. Ae. aegypti, An. 3 and 4 Kadarohman et al.,
Nash 7095 sundaicus, Culex sp. 2013
resp.
Methanol 0.276, n.d. An. stephensi 2, 3 Aarthi and Murugan,
0.285, and 4 2010
0.305%
Vincetoxicum Apocynaceae Stem Methanol 717 957 Cx. quinquefasciatus 4 Pavela, 2009
hirundinaria Medik.
Vitex cymosa Bertero ex Lamiaceae Stem Ethanol 875 n.d. Ae. aegypti 3 Garcez et al., 2009
Spreng.
Vitex negundo L. Lamiaceae Leaf Distilled water 29.32; n.d. An. subpictus, Cx. 4 Gandhi et al., 2016
33.65 quinquefasciatus resp.
Leaf Hexane 34; 57 127; 211 An. stephensi, Cx. 4 Kamaraj et al., 2010b
Leaf Hexane 65 302 Cx. tritaeniorhynchus 4 Kamaraj et al., 2009
Vitex payos (Lour.) Merr. Lamiaceae Root bark Acetone 16 n.d. An. gambiae 3 and 4 Nyamoita et al., 2013
Stem bark Acetone 512 n.d.
Vitex trifolia L. Lamiaceae Leaf Methanol 77 n.d. An. gambiae 3 and 4 Nyamoita et al., 2013
Stem bark Acetone 926 n.d.
Withania somnifera (L.) Solanaceae Fruit Methanol 135; 108; 711; 523; 2159 Ae. aegypti, An. 3 and 4 Bansal et al., 2015
Dunal 1092 stephensi, Cx.
Green fruit Methanol 197; 115; n.d. Ae. aegypti, An. 3 and 4 Bansal et al., 2011
555 stephensi, Cx.
Red fruit Methanol 95; 95; 269; 282; n.d. Ae. aegypti, An.
1013 stephensi resp.
Red fruit Distilled water 445 n.d. Cx. quinquefasciatus
Red fruit Petroleum ether 45; 30; 54 225; 128; 198 Ae. aegypti, An.
(without stephensi, Cx.
seed) quinquefasciatus resp.
Seed Petroleum ether 140; 778 n.d. An. stephensi, Cx.
Seed Methanol 769 n.d. Ae. aegypti
Leaf Benzene 342 n.d. Cx. quinquefasciatus 4 Karmegam et al.,
1997
Wrightia tinctoria (Roxb.) Apocynaceae Leaf Petroleum ether 421 n.d. Cx. quinquefasciatus 4 Karmegam et al.,
R.Br. 1997
Xanthium strumarium L. Asteraceae Seed Methanol 502; 531 867; 906 Cx. pipiens, Ae. caspius 4 Al-Mekhlafi et al.,
resp. 2017
Zanthoxylum spp. Rutaceae Stem Ethanol 538 n.d. Ae. aegypti 3 Garcez et al., 2009
Leaf Ethanol 435 n.d. Ae. aegypti
Zingiber officinale Roscoe Zingiberaceae Stem Hexane 55 129 Ae. aegypti 4 Kumar et al., 2012c
Rhizome Methanol 521; 390 524; 628 Ae. albopictus, Cx. 3 and 4 Nath et al., 2006
Zingiber zerumbet (L.) Zingiberaceae Root Hexane 26 128 Cx. gelidus 4 Kamaraj et al., 2010a
Roscoe ex Sm. Root Ethyl acetate 19 87 Cx. quinquefasciatus
Zornia diphylla (Benth.) Fabaceae Leaf Distilled water 65; 61; 71 127; 121; 134 Ae. albopictus, An. 3 Govindarajan et al.,
Pittier subpictus, Cx. 2016b
n.d.=not determined.
resp.=respectively.
IVM programs. alkaloids, alkamides, anthraquinones, coumarins, flavonoids, limonoids,

polyacetylenes, sesquiterpene lactones, sterols, thiophenes, triterpenoids
4. Phytochemical constituents of extracts with noteworthy and xanthones (Fig. 2). Most of them are known to play an important role in
larvicidal activity the constitutive or induced plant chemical defense against microorganisms
(e.g., fungi, bacteria, virus) and harmful insects.
Notably, for 19 out of 29 plant extracts we found in literature sev-
eral phytochemical studies highlighting the marker compounds that are 4.1. Alkaloids and alkamides
likely to be responsible for the larvicidal properties of extracts. These
constituents are summarized in Table 2 where they are listed according Alkaloids are nitrogen-containing heterocycles, with basic reactivity,
to the plant species from which they are obtained and their chemical biosynthetically derived from amino acids and involved in the plant defense
group. Other informations, such as the possible mode of action (ex- against herbivores and pathogens (Mithöfer and Boland, 2012). Their
pected for a certain group of substances) and the calculated LC50 values classification is based on the biogenetic origin, which influences the type of
on some mosquito species, whenever available, are also reported. functional group of the molecule. Alkaloids were found in the extracts from
The marker compounds of the bioactive plant extracts are divided into Nelumbo nucifera Gaertn. (nuciferine and neferine), Piper nigrum L. and P.
the following main chemical classes: i.e. (in alphabetical order) acetogenins, longum L. (piperine, piperlongumine, pipericide, retrofractamide A),
254
Fig. 1. Contribution of botanical families in terms of plant extracts tested as botanical insecticides against mosquito larvae; the analysis includes 429 plant species
belonging to 101 botanical families.
Scindapsus officinalis (Roxb.) Schott (scinamide A, scinamide B), Solanum on third instar larvae of Culex, Anopheles and Aedes species, respectively. Its
xanthocarpum Schrad. & H. Wendl. (solasodine) and Trichodesma africanum larvicidal effect has been correlated with the alteration of muscular and
(L.) Sm. (trichodesmine). Only the pepper alkaloids, also known as piper- locomotor activity (Saraf and Dixit, 2002), which in turn is supposed to
amides for the presence of an amine group, were successfully assayed depend on the interaction with the GABAergic system (Boonen et al., 2012).
against mosquito larvae confirming the insecticidal efficacy of Piper extracts Spilanthol efficacy proved to be superior to that of some commercial in-
as source of effective, safe and scalable insecticides (Scott et al., 2008). secticides (Kadir et al., 1989).
Notably, pipericide and retrofractamide A (Fig. 2) were found as the most
potent larvicides, showing LC50 values of 0.004, 0.1 and 0.26 ppm, and
4.2. Coumarins
0.028, 0.039 and 0.01 ppm, on Cx. pipiens pallens, Ae. aegypti and Ae. togoi,
respectively, (Park et al., 2002) (Table 2). These values were consistent with
Coumarins are bicyclic aromatic compounds endowed with the 5,6-
those of many synthetic insecticides. On the same target mosquitoes, pi-
benzo-2-pyrone (α-cromone) moiety and biosynthetically derived from the
perine, the most abundant alkaloid in pepper extracts, was found as sig-
shikimate pathway (de Souza et al., 2016). They occur in plants under free
nificantly less toxic, with LC50 values of 3.2, 5.1 and 4.6 ppm, respectively
or glycosylated form and are abundant in several families such as Apiaceae,
(Park et al., 2002).
Rutaceae, Fabaceae and Poaceae. They are classified into four groups,
Piperamides showed multiple modes of action, for instance neu-
namely hydroxycoumarins, furanocoumarins, pyranocoumarins and glyco-
tralizing the detoxifying system of insect by interacting with the oxi-
sylated coumarins. Among them, furanocoumarins (also known as fur-
doreductase, monooxygenase and glutathione S-transferase activity, or
anochromones) contained in Ammi visnaga (L.) Lam. extract, namely khellin
giving neurotoxicity by activating the nicotinic acetylcholine receptors
and visnagin (Fig. 2), are characterized by a furan ring condensed with the
(Godara et al., 2018; Samuel et al., 2016; Scott et al., 2008). Further-
benzopyrone moiety and exhibited important insecticidal activity through
more, they showed antifeedant activity and disruption of embryonic
induction of photosensitization (Pavela, 2015b; Maleck et al., 2013). The
development (Scott et al., 2008). Also, synergism of piperamides with
latter is particularly important due to the presence of extended conjugated
conventional insecticides such as pyrethrum was observed (Jensen
groups. Another possible mechanism of action of khellin and visnagin is
et al., 2006). This may allow to reduce the dose of the latter producing
their interaction with detoxification enzymes such as transaminase and
harmful effects to non-target organisms.
cytochrome P450 monooxygenase (Wen et al., 2009). Similar mechanism
Solasodine is a steroidal alkaloid produced by the Solanaceae, such
may be ascribed to visnadin, a pyranocoumarin formed by a pyran ring
as S. xanthocarpum, and exerting distortions in the development of
condensed to the benzopyrone group (Winderl et al., 2016; Wen et al.,
pupae and adults (Patel et al., 2012; Lingampally et al., 2014). How-
2009). Khellin LC50 values on Cx. quinquefasciatus and Ae. aegypti were 10
ever, their LC50 values have not been provided yet.
and 50 ppm, respectively, whereas visnagin showed an LC50 of 26 ppm on
Trichodesmine is a pyrrolizidine alkaloid isolated from T. africanum
Cx. quinquefasciatus (Pavela et al., 2016; Maleck et al., 2013).
that exerts antifeedant activity on insects (Macel, 2011; Omar et al.,
1983), but expected toxicity on humans, i.e. hepatotoxicity, could limit
its use in insecticidal formulations (Molyneux et al., 2011). 4.3. Anthraquinones
Other alkaloids contained in larvicidal extracts, are nuciferine and
neferine from N. nucifera and scinamides from S. officinalis that are Anthraquinones are aromatic compounds bearing two ketonic functions
worthy of investigation to determine their LC50 values and mechanism derived from the oxidation of anthracene. They are tricyclic compounds
of action on mosquitoes (Fang et al., 2017; Yu et al., 2018). endowed with high lipophilicity under free forms (not glycosilated). They
N-alkylamides are secondary metabolites playing an important role in are derived from polymerization of eight units of acetyl-CoA forming a liner
the plant defense (Boonen et al., 2012). They are made up of a poly-un- chain undergoing cyclization leading to oxygenated functional groups such
saturated fatty chain linked to a variously substituted amine group. Both can as phenols and ketons (Sendelbach, 1989). These compounds are typical of
present cyclic systems and heteroatoms (e.g. S, O, N) (Boonen et al., 2012). several families belonging to dicotyledons such as Polygonaceae, Fabaceae,
N-alkylamides are distributed in several families such as Asteraceae, Ruta- Rubiaceae and Rhamnaceae.
ceae, Piperaceae, Rutaceae and Menispermiaceae. Among them, spilanthol, Emodin is one of the main antraquinones obtained from the stem of
the main alkylamide produced by Spilanthes acmella DC. (Asteraceae, syn. Coccoloba mollis Casar. and responsible for its larvicidal efficacy.
Acmella oleracea (L.) R.K.Jansen), is a promising insecticide (Benelli et al., Notably, it exerted a pronounced toxicity, with LC50 values of 1.4, 1.9
2019). This compound is characterized by an isobutyl side chain which is and 2.2 ppm on Cx. pipiens pallens, Ae. aegypti and Ae. togoi, respectively
responsible for the tingling effect given by the plant consumption (Borges (Yang et al., 2003). In insects, emodin exerts antifeedant activity and
et al., 2012). Spilanthol is particularly effective on ticks and mosquitoes post-ingestive damages in the midgut (Georges et al., 2008; De Liberto
(Benelli et al., 2019). Notably, in the study by Ramsewak et al. (1999) and Werner, 2016). Similar mechanism might be ascribed to physcion,
spilanthol showed an LC50 of 6.8 ppm on Ae. aegypti larvae. Saraf and Dixit another marker components of C. mollis extracts, although studies on its
(2002) confirmed this result reporting LC50 values of 4.5, 5.1 and 4.5 ppm insecticidal capacity are missing.
255
Table 2
Selected marker compounds of plant extracts endowed with relevant mosquito larvicidal activity (LC50 < 10 ppm).
Compound Chemical group Molecular Chemical Plant extract Possible mechanism of action for the LC50 (ppm) (target species) References
R. Pavela, et al.
weight formula chemical group
visnagin furanochromones 230.22 C13H10O4 Ammi visnaga phototoxicity; interaction with transaminase 26 (Cx. quinquefasciatus) Pavela, 2015a,b,c; Pavela et al., 2016;
khellin 260.07 C14H12O5 and cytochrome P450 monooxygenase 10 (Cx. quinquefasciatus); 50 Wen et al., 2009; Maleck et al., 2013
(Ae.aegypti)
visnadin pyranocoumarins 388.15 C21H24O7 interaction with transaminase and not found Winderl et al., 2016; Wen et al., 2009
cytochrome P450 monooxygenase
squamocin G acetogenins 622.48 C37H66O7 Annona squamosa inhibition of NADH ubiquinone oxydo- 0.01 (Ae. aegypti) Chen et al., 2012; Bonneau et al., 2017;
reductase; neuronal toxicity; reduction of Boreddy et al., 2000; Rongnoparut
protein level; midgut damage; antifeedant et al., 2016; Costa et al., 2017; da Silva
activity Costa et al., 2016; Begum and Pandey,
2017
artemisinin sesquiterpene 282.15 C15H22O5 Artemisia annua generation of ROS 1.9 (An. stephensi); 14.3 (Cx. Efferth, 2017; Sharma et al., 2012,
lactones quinquefasciatus); 4.1 (Cx. 2014
tritaeniorhynchus)
dihydroartemisinin 284.16 C15H24O5 not found
pyropheophorbide a chlorin derivatives 534.26 C33H34N4O3 Atalantia phototoxicity 8.4 (Culex sp.) Chansakaow et al., 1996; Singh et al.,
monophylla 2017
atalantin limonoids 500.54 C27H32O9 antifeedant activity not found Dreyer et al., 1976; Wu et al., 1997
β-sitosterol phytosterols 414.39 C29H50O Citrus grandis; antifeedant activity 11.5 (Ae. aegypt); 3.6 (An. stephensi); Mokbel and Hashinaga, 2006;
Persea americana 26.7 (Cx. quinquefasciatus) Chandramu et al., 2003; Rahuman
et al., 2008; Leite et al., 2009
limonin limonoids 470.19 C26H30O8 Citrus grandis inhibition of glutathione S-transferase; 71.6 (Ae. aegypti) Xiang et al., 2014; Mendel et al., 1991;
antifeedant activity 32.4 (Ae.albopictus) Kiprop et al., 2005; Liu et al., 2012
cleomdiolic acid secodammarane 490.37 C30H50O5 Cleome rupicola not reported not found Al-Rehaily et al., 2017
triterpenes
256
pectolinaringenin flavones 314.29 C17H14O6 Clerodendrum glutathione S-transferase and esterase 10.2 (An. stephensi); 0.6 (Cx. Baskar et al., 2014;
phlomidis activities; reduction of protein level; quinquefasciatus); 0.8 (Ae. aegypti) Duraipandiyan et al., 2015; Muthu
antifeedant activity et al., 2012, 2015;
emodin anthraquinones 270.24 C15H10O5 Coccoloba mollis antifeedant activity; post-ingestive damage 1.4 (Cx. pipiens pallens); Barros et al., 2011;
1.9 (Ae. aegypti); De Liberto and Werner, 2016; Ahn
2.2 (Ae. togoi) et al., 2013; Yang et al., 2003; Georges
< 25 (An. gambiae) et al., 2008
physcion 284.27 C16H12O5 not found
4-hydroxy-3-methoxybenzoic benzoic acids 168.15 C8H8O4 not reported not found Oliveira et al., 2008
acid
α-mangostin prenylated xanthones 410.47 C24H26O6 Garcinia blocking of sterol carrying proteins; 2.2 (Ae. aegypti); 2.9 (An. gambiae); Mohan et al., 2018; Kumar et al., 2010;
mangostana phototoxicity; interaction with cytochrome 1.7 (An. quadrimaculatus); 1.9 (An. Larson et al., 2010
P450, glutathione S-transferase and esterase stephensi); 0.8 (Cx. pipiens); 1.6 (Cx.
activities quinquefasciatus)
nuciferine alkaloids 295.38 C19H21NO2 Nelumbo nucifera not reported not found Fang et al., 2017
neferine 624.78 C36H44N2O6 Nelumbo nucifera
persin acetogenins 380.57 C23H40O4 Persea americana antifeedant activity not found Rodríguez-López et al., 2015;
Rodriguez-Saona et al., 1997
1,2,4-trihydroxynonadecane alkane derivatives 316.53 C19H40O3 not reported Leite et al., 2009; Oberlies et al., 1998
1,2,4-trihydroxyheptadec-16- alkene derivatives 286.25 C17H34O3
ene
1,2,4-trihydroxyheptadec-16- alkyne derivatives 284.24 C17H32O3
yne
piperine alkaloid amides 285.34 C17H19NO3 Piper longum, Piper antifeedant activity, nicotinic acetylcholine 3.2. (Cx. pipiens); 5.1 (Ae. aegypti); Godara et al., 2018: Scott et al., 2008;
nigrum activated cation selective channel activity, 4.6 (Ae. togoi) Samuel et al., 2016; Corbel et al., 2002;
piperlongumine 317.13 C17H19NO5 oxidoreductase, monooxygenase and not found Park et al., 2002
pipericide 355.48 C22H29NO3 glutathione S-transferase activity, inhibition 0.004 (Cx. pipiens pallens); 0.1 (Ae.
of embryonic development aegypti); 0.26 (Ae. togoi)
retrofractamide A 327.42 C20H25NO3 0.028 (Cx. pipiens pallens);
0.039 (Ae. aegypti); 0.01 (Ae. togoi)
Acta Tropica 193 (2019) 236–271

4.4. Xanthones
2012; Vats, 2018; Santiago et al., 2012;
2009; Lu et al., 2017; Liang et al., 2015

Stevenson et al., 2012; Belmain et al.,
2002; Ramsewak et al., 1999; Boonen
Patel et al., 2012; Lingampally et al.,
Yenesew et al., 2003; Yenesew et al.,

Benelli et al., 2019; Saraf and Dixit,
Ibrahim et al., 2018; Marles et al.,

Anthraquinones-related compounds are xanthones which differ for the
Macel, 2011; Omar et al., 1983

presence of only one ketonic function into the tricyclic backbone (Fotie and
Bohle, 2006). Among them, α-mangostin is a prenylated xanthone isolated
1991; Perich et al., 1995

from mango (Garcinia mangostana L.) pericarp with high potential to be
used as an ingredient of botanical insecticides. Notably, it displayed several
Yu et al., 2018
et al., 2012 modes of action in insects, such as the inhibition of sterol carrying proteins
and interaction with cytochrome P450, glutathione S-transferase and es-
References
terase activities (Larson et al., 2010; Kumar et al., 2010). A photo-induced

2014
toxicity of α-mangostin in water was also recorded (Ibrahim et al., 2016).

Therefore, this xanthone can be a promising compound to combat in-
secticide resistance. α-Mangostin showed significant toxicity on mosquito
larvae, with LC50 values of 2.2, 2.9, 1.7, 1.9, 0.8 and 1.6 ppm on Ae. agypti,
An. gambiae, An. quadrimaculatus, An. stephensi, Cx. pipiens and Cx. quin-
LC50 (ppm) (target species)
quefasciatus, respectively (Larson et al., 2010).

1.9 (Cx. pipiens pallens)
5.1 (Anopheles sp.)
3.2 (An. stephensi)
4.5. Flavonoids
6.8 (Ae. aegypti)
1.6 (Ae. aegypti)

1.4 (Ae. aegypti)
0.5 (Ae.aegypti)
4.5 (Aedes sp.)
4.5 (Culex sp.)
not found
not found
not found
not found
Flavonoids represent a large class of plant secondary metabolites de-

rived from two distinct metabolic pathways, namely shikimate and acetyl-
coenzyme A (Treutter, 2006). They consist in a tricyclic system made up of
an aromatic ring (A) showing one or two hydroxylic groups, a central
inhibition of pupation; formation of mosaics;
activity; interaction with GABAergic systme
pyranic ring (C) and a second aromatic ring (B), variously substituted,
linked to the former at position 2. When the link between these rings occurs
alteration of movement and muscular
antifeedant activity; genotoxicity and

Possible mechanism of action for the
oxidoreductase; inhibition of protein
at position 3, isoflavones are formed. The variability in functionalization of

inhibition of NADH:ubiquinone
the pyranic ring gives rise to different subtypes (e.g., flavones, flavonols,
synthesis in the integument
anthocyanins, flavan-3-ols and flavanones, among others) (Treutter, 2006).

deformed pupae and adults
In the plant cell, flavonoids are accumulated into the vacuole under gly-
cosylated forms. They are endowed with several biological activities which
depend on their interaction with numerous enzymes. Flavonoids play an
chemical group
phototoxicity
mutagenicity
important role as ecological mediators, giving an important contribution

not reported
also in the plant defense against microorganisms and insects (Treutter,

2006). Among flavones, one of the most promising compounds is pectoli-
naringenin (5,7-dihydroxy-4′,6-dimethoxy-flavone) (Fig. 2) obtained from
the leaves of Clerodendrum phlomidis L.f. (Table 2). This compound was
Spilanthes acmella
Tephrosia vogelii
proved to affect the detoxification system of insects through reduction of

Tagetes minuta
xanthocarpum
Plant extract
Trichodesma
glutathione S-transferase (GST) and esterase activities (Baskar et al., 2014).

Scindapsus
africanum
officinalis
Solanum
It also reduces the total protein content and exerted antifeedant activity
(Muthu et al., 2015). When assayed on mosquito larvae, pectolinaringenin
exhibited noteworthy toxicity showing LC50 values of 10.2, 0.6 and 0.8 ppm
on An. stephensi, Cx. quinquefasciatus and Ae. agypti, respectively (Muthu
C21H21ClOS2
et al., 2012, 2015).

C27H2402S5
C18H21NO5
C19H19NO6
C18H27NO6
C27H45NO
C14H23NO
C23H22O7
C23H22O6
C23H22O6
Chemical
formula
Rotenoids are particular forms of isoflavonoids produced by plants to

defend themselves against insects (Fig. 2) (Fukami and Nakajima, 1971).
Rotenone is the parent molecule of this class of secondary metabolites,
firstly isolated from the roots of the genus Derris and also obtained from
Molecular
Lonchocarpus and Tephrosia, all belonging to the Fabaceae family. It is

331.37
357.36
399.66
221.34
540.79
388.97
394.42
410.42
394.42
353.42
weight
marketed as a natural insecticide in several countries (Pesticide Action

Network, 2004), however, its relatively high toxicity to mammals (LD50 in
rats of 132 mg/kg) limited its use (El-Wakeil, 2013). Rotenone is a mi-
tochondrial activity disruptor, being able to inhibit the NADH:ubiquinone
Chemical group
oxidoreductase activity making insect unable of getting energy and causing

pyrrolizidine
alkylamides
arrest of feeding (El-Wakeil, 2013). Tested on mosquito larvae, rotenone

thiophenes
rotenoids
alkaloids
alkaloids
alkaloids
showed very high toxicity with LC50 values of 0.5, 3.2 and 1.9 ppm on Ae.
agypti, An. stephensi and Cx. pipiens pallens, respectively (Vats, 2018). Rote-
noids are the marker compounds of Tephrosia vogelii Hook.f. (Fabaceae), a
tropical plant traditionally used as a fish poison and insecticide in Africa
(Stevenson et al., 2012). From the leaves of T. vogelii different rotenoids are
obtained. Besides rotenone, deguelin and tephrosin (Fig. 2) are the major
Table 2 (continued)
rotenoids of leaf extracts. The latter showed similar insecticidal power when
compared to rotenone. Their LC50 on Ae. aegypti was 1.6 and 1.4 ppm, re-
trichodesmine
thiotagetin A
thiotagetin B
spectively (Belmain et al., 2012). Thus, these two new-characterized rote-

scinamide A
scinamide B
Compound
solasodine
spilanthol
tephrosin
rotenone
noids from T. vogelii are promising ingredients to replace rotenone in in-

deguelin
secticidal formulations. However, furthers studies supporting their lower

toxicity to mammals, when compared to rotenone, are needed.
257
4.6. Acetogenins Marles et al., 1991). On the above, the possibility to induce resistance in
insects by these compounds is relatively unlikely. Squamocin G showed
Acetogenins are derivatives of fatty acids bearing an odd-carbon a very low LC50 value (0.01 ppm) on Ae. aegypti larvae (Costa et al.,
aliphatic chain composed of 35–37 carbon atoms ending with an α-β- 2017). However, further studies are needed to prove the safety of these
unsaturated γ-lactone ring. This group of natural products is typical of compounds on humans since some acetogenins are suspected to cause
the Annonaceae family (Annonaceous acetogenins) and are responsible neurodegenerative diseases (Bonneau et al., 2017).
for several biological activities of these plants (Rupprecht et al., 1990). Acetogenins are also contained in the seeds of avocado (Persea
Among them, squamocin G (Fig. 2) is the major acetogenin contained in Americana Mill.), notably persin which is structurally similar to fatty
the seeds of Annona squamosa L. bearing the bistetrahydrofuran ring acids (Fig. 2). They may contribute to the possible antifeedant activity
into the backbone skeleton (Chen et al., 2012). Annonaceous acet- displayed by the avocado extracts on insects (Rodríguez-López et al.,
ogenins display multiple effects on insects, such as the inhibition of 2015; Rodriguez-Saona et al., 1997). However, further investigation
NADH ubiquinone oxidoreductase and protein synthesis, and midgut using the pure compound is needed.
damages (Bonneau et al., 2017; da Silva Costa et al., 2016). Antifeedant
activity and phototoxicity are also possible (Begum and Pandey, 2017;
Fig. 2. Selected chemical constituents of plant extracts reported in Table 2 being promising for the development of mosquito larvicidal formulations.
258
Fig. 2. (continued)
4.7. Thiophenes they are mainly accumulated in the roots. They play an important
ecological role, being able to interact with plant pathogens, insects and
Thiophenes are a class of natural compounds composed of a poly- parasites (Ibrahim et al., 2018; Perich et al., 1995). Thiophenes are
acetylene chain and one or more sulfur-containing heterocycles (Sperry particularly abundant in the genus Tagetes which showed potential as
and Wright, 2005). They are typical of the Asteraceae family where source of larvicidal extracts. Notably, Tagetes minuta L. is a plant spread
259
in temperate to tropical regions with a long history of use as an in- insecticides at lower doses thus minimizing their potential impact on
secticidal agent (Ibrahim et al., 2018). From this species, thiotagetin A the environment, including reduced risks for non-target organisms and
and thiotagetin B (Fig. 2) have been characterized as two of the main human health as well. In this section, the most important cases of sy-
thiophenes which are probably involved in the larvicidal effects dis- nergistic blends made up of plant extracts with synthethic insecticides
played by the plant extract (Table 1). Overall, thiophenes are light- assayed as larvicides against mosquitoes are summarized.
activated toxins producing notable toxicity in insects (Marles et al., About four decades ago, Kalyanasundaram and Babu (1982) assayed
1991). However, they are still to be assayed for determining their LC50 five petroleum ether extracts, namely Ocimum basilicum L., O. sanctum L.,
values and safety to humans. Azadirachta indica A. Juss., Lantana camara L. and Vitex negundo L., alone or
in combination with organophosphates like phenthoate and fenthion,
4.8. Sesquiterpene lactones against fourth instar larvae of Cx. quinquefasciatus. When mixed with syn-
thetic insecticides at 1 and 5 ppm concentrations, the plant extracts reduced
Sesquiterpene lactones are an important group of terpenoids (more significantly the LC50 values (up to 0.001174 and 0.000495 ppm with
than 3000 compounds identified so far) characterizing the Asteraceae phenthoate and fenthion, respectively) compared with single treatments.
family where they are secreted in the glandular trichomes and involved Notably, the extracts from O. basilicum and A. indica at 5 ppm were the most
in the plant defense, displaying repellents and allelopathic activities effective in reducing the LC50 values of phenthoate and fenthion, respec-
(Sut et al., 2018; Brunetti et al., 2018). These compounds own a C15 tively. Moreover, Kalyanasundaram and Das (1985) evaluated the syner-
carbon skeleton encompassing a lactone ring formed from the cyclic gism of several petroleum ether extracts, including Vinca rosea L., Leucas
condensation between the alcoholic and acid functional groups. The aspera (Willd.) Link, Pedalium murex L., Clerodendron inerme (L.) Gaertn.,
presence of a lactone conjugated with a double bound (α-methylene-γ- Rauvolfia canescens L., Turnera ulmifolia L., Parthenium hysterophorus L., Ca-
lactone group) makes these compounds extremely reactive with cy- lotropis sp. and Adathoda sp., with phenthoate and fenthion against larvae of
steine thiol groups of proteins by alkylation reactions (Sut et al., 2018). Cx. quinquefasciatus, An. stephensi Liston and Ae. aegypti L. They highlighted
Artemisinin is the major sesquiterpene lactone isolated from the leaves that most of them, at 5 ppm, displayed significant degrees of synergism.
of Artemisia annua L., which is known as qinghao in the traditional Later, a fraction from the leaf petroleum ether extract of O. basilicum was
Chinese medicine. This compound, having an endoperoxide ring, is effective in combination 1:1 with the neonicotinoid imidacloprid on An.
responsible for the antimalarial activity of qinghao extracts (Efferth, stephensi larvae, showing LC50 values of 0.010 amd 0.007 ppm at 24 and
2017). For its discovery, the scholar Tu Youyou received the Nobel 48 h, respectively (Maurya et al., 2012). This combination was evaluated as
Prize for Medicine in 2015 (Benelli and Mehlhorn, 2016; Efferth, 2017). one of the most effective in the management of anopheline mosquitoes
Concerning the mechanism of action, the endoperoxide ring of artemi- (Maurya et al., 2012). Harve and Kamath (2004) found out that acetone and
sinin is easily broken into radicals which exert relevant toxicity to many petroleum ether extracts from Murraya koenigii (L.) Spreng. leaves, Corian-
parasites (Pavela et al., 2018). Furthermore, this reactivity of the molecule drum sativum L. seeds, Ferula assa-foetida L. powder and Trigonella foenum-
has been found crucial also for the larvicidal activity. Indeed, artemisinin graecum L. seeds (in the concentration range of 25–100 ppm) in combina-
showed noteworthy toxicity to An. stephensi, Cx. quinquefasciatus and Cx. tion with fenthion and temephos (at 0.05 ppm) produced a synergistic effect
tritaeniorhynchus with LC50 values of 1.9, 14.3 and 4.1 ppm, respectively against Ae. aegypti.
(Sharma et al., 2012, 2014). Another possible mode of action of artemisinin Furthermore, Raghavendra et al. (2013) showed that the extract
that is worthy of further investigation in insects can be the inhibition of from leaves of the weed Solidago canadensis L. increased the efficacy of
glutathione S-transferase (Efferth, 2017). deltamethrin on Ae. aegypti when used in a mixture at 1:1 ratio. Madhu
and Vijayan (2010) reported on the synergistic effect of P. longum
4.9. Sterols petroleum ether extract with propoxur at 1:1 ratio against Cx. quin-
quefasciatus larvae. Mohan et al. (2010) studied the effect of combining
Sterols are substances endowed with a steroid backbone including a the S. xanthocarpum petroleum ether extract with some synthetic in-
hydroxylic function in beta-configuration at position 3 (Bouic and secticides (temephos and fenthion) on Cx. quinquefasciatus. Whereas the
Lamprecht, 1999). They are ubiquitous in plant organisms in which are combination with temephos gave antagonistic effects, the mixture 1:1
considered structurally and functionally important for cells. β-Sitosterol with fenthion produced synergistc effects, showing LC50 of 0.0144 and
(Fig. 2) is one of the most abundant sterols in plants where it is con- 0.056 ppm at 24 and 48 h, respectively. The same extract resulted sy-
sidered essential to maintain the membrane fluidity (Hennessey, 1992). nergic with cypermethrin in a 1:1 blend, giving LC50 values on Cx.
The lipophilic extracts obtained from the seeds of P. americana and peel quinquefasciatus of 0.089 and 0.087 ppm at 24 and 48 h, respectively
of Citrus grandis (L.) Osbeck are particularly rich in this compound. β- (Mohan et al., 2006). Besides, Aivazi and Vijayan (2010) found out that
Sitosterol was assayed as larvicidal on Ae. aegypti, An. stephensi and Cx. the leaf petroleum ether extract of Ruta graveolens L. synergized the
quinquefasciatus, showing LC50 values of 11.5, 3.6 and 26.7 ppm, reactivity of cypemethrin on An. stephensi when assayed at 1:1 ratio.
spectively (Rahuman et al., 2008). The mechanism of action of this Thangam and Kathiresan (1997) screened a pool of mangrove species
sterol on mosquitoes has not been investigated in deep, though the for larvicidal activity and synergism with pyrethrum against Cx. quin-
antifeedant activity is believed to be involved in the effect displayed quefasciatus. They found that the petroleum ether extract from roots of
(Chandramu et al., 2003). Rhizophora apiculata Blume gave the best results in terms of overall
toxicity (LC50 of 25.7 ppm) and synergistic effects with pyrethrum at
5. Larvicidal plant extracts synergizing synthetic larvicides 5 ppm, reducing its LC50 to 0.107 ppm.
On the above it is evident that plant extracts have the potential to be
As outlined in the paragraphs above, a plant extract is a complex incorporated in conventional insecticidal formulations together with syn-
mixture made up of hundreds of secondary metabolites among which thetic compounds to minimize the development of resistance. However,
several of them are credited with the most powerful biological effects, further studies should be performed, especially on their effects on non-target
oftenly acting on insects at different levels (see paragraph 3). organisms. At the same time, only those plant extracts assuring scalability of
Frequently, an effective strategy to boost the insecticidal efficacy of a production on an industrial level should be selected.
conventional (synthetic) insecticide is to prepare synergistic blends
with one or more plant extracts, that can act on different targets 6. Synergized blends of larvicidal plant extracts
compared with the active ingredient of the commercial product, thus
improving the overall toxicity coupled with the reduction of probability An alternative to mix synthetic larvicides with plant extracts sy-
of inducing resistance. This strategy allows to use the conventional nergizing their action, can be exploiting the synergism between
260
different plant extracts and/or selected ingredients (see also Pavela, the leaf aqueous extract of Berberis tinctoria Lesch. achieved LC50 values
2015c). To the best of our knowledge, this has been carried out in of 552 and 481 ppm against the non-target copepod Mesocyclops ther-
several studies. Recently, Yankanchi et al. (2014) evaluated the sy- mocyclopoides and the mosquito predator Toxorhynchites splendens, re-
nergistic effects of different plant extracts against Ae. aegypti and spectively, while against 1st-4th instar larvae of Ae. albopictus LC50 va-
showed that the most promising were those from leaves of V. negundo lues ranged from 183 to 360 ppm, with no overlapping 95% CI when
and C. inerme in combination with the seed extract of Pongamia glabra comparing their values to those estimated for the two non-target species
Vent. Furthermore, the mixture of Andrographis echioides (L.) Nees and (Kumar et al., 2016). The flower aqueous extract of Couroupita guia-
Cadaba trifoliata Wight & Arn. leaf extracts was tested by Rajkumar nensis Aubl. at 102 ppm significantly improved predation efficacy of
et al. (2012) on Ae. agypti 4th instar larvae. Authors found that this Aplocheilus lineatus fishes against An. stephensi larvae from 53% (stan-
combination reduced significantly the LC50 values compared with dard laboratory conditions) to 84% (aquatic area where fishes were
single treatments, i.e. from 123.4 to 68.3 ppm. Singha et al. (2011) exposed to the flower extract) (Subramaniam et al., 2016). Simarly, low
reported a synergistic effect between the extracts (chloroform: me- concentration of the leaf extract of Chenopodium ambrosoides L. (syn.
thanol (1:1) and ethyl acetate) from fruits of Croton caudatus Geiseler Dysphania ambrosioides(L.) Mosyakin & Clemants) significantly boosted
and flowers of Tiliacora acuminata Miers. against larvae of Cx. quin- the predation rates of Oryzias melastigma fishes against both 2nd and 3rd
quefasciatus. Annona muricata L. seed and P. nigrum fruit ethanolic ex- instar larvae of Ae. albopictus (Subramaniam et al., 2017).
tracts, containing acetogenins and piperamides, were synergistically It is also worthy to note that the exposure to sub-lethal concentra-
used in combination against third instar larvae of Ae. aegypti from 90:10 tions of various plant extracts (i.e., generally 1/3 of the LC50 calculated
to 10:90 ratio (Grzybowski et al., 2013). The most active combination on larval instars of selected mosquito species, otherwise 1–3 ppm) used
was 9:1 of A. muricata-P. nigrum extracts, respectively; this mixture to fabricate metal and metal oxide larvicidal nanoparticles did not led
caused pronounced alterations in the body of mosquito larvae. Shaalan to detrimental effects on various non-target organisms, such as odonate
et al. (2005a,Shaalan et al., 2005b reported on the occurrence of sy- nymphs (Murugan et al., 2015a), copepods (Murugan et al., 2015d) and
nergistic effects between different extracts of Callitris glaucophylla tadpoles (Murugan et al., 2015e), even if genotoxicity on Carassius
J.Thomps. & L.A.S.Johnson and Khaya senegalensis (Desv.) A.Juss., as auratus fishes exposed to some of these green-synthesized nanomater-
well as between these extracts and three commercial insecticides such ials (e.g., neem-cake synthesized Ag nanoparticles, 12 ppm) has been
as fenitrothion, lambda-cyhalothrin and methoprene against both Ae. detected (Chandramohan et al., 2016; see Benelli et al., 2017a for a
aegypti and Cx. annulirostris Skuse. dedicated review on non-target effects of green-fabricated nanosized
In this framework, further research studying the potential sy- insecticides).
nergistic and antagonistic effects among the most effective extracts Besides, literature reported toxicity on non-target organisms is
outlined in this review for their LC50 < 10 ppm, as well as among se- really limited for some of the marker compounds responsible for the
lected pure constituents (see Fig. 2) is needed, along with proper field mosquitocidal effects of extracts detailed in Table 2. Thus, further in-
formulation to improve stability and efficacy and the assessment of vestigations on this important aspect should be assured in the future.
non-target toxicity in the aquatic environment. Considering potential toxicity to mammals, it is worth noting that
when a plant extract comes from a species traditionally used as a food,
7. Non-target toxicity of plant extracts effective as larvicides scarce toxicity to humans and other vertebrates can be expected.
Indeed, several species including A. squamosa, Citrus spp., G. mangos-
Due to the large amount of data reviewed in the earlier paragraphs tana, P. americana, Piper spp. and S. acmella, which have been used as
on mosquito larvicides, this review cannot cover also the impact of food for long time, may be considered as relatively safe. In this regard,
botanical larvicides on non-target species with a quantitative analysis of some of them have been classified as generally recognized as safe
literature. For detailed non-target issues the reader is referred to other (GRAS) by the FDA and/or EFSA (Pavela and Benelli, 2016a,b; Benelli
comprehensive reviews and research studies (e.g., Shaalan et al., et al., 2019). On the other hand, for other species for which a docu-
2005a,b; Promsiri et al., 2006; Chowdhury et al., 2008; Ghosh et al., mented use as poison and insecticide has been reported (e.g., C. phlo-
2012). However, general concepts on this topic are highlighted below. midis, T. minuta, T. africanum, T. vogelii), a serious evaluation of their
As complex mixtures of active constituents, which are selective, biotoxicity is needed through determination of LD50 in rats after oral ad-
degradable and owning a wide range of activities, plant-borne extracts ministration and related genotoxicity and mutagenicity. This point re-
are expected to be scarcely toxic (at least little acute toxicity) to various presents the future challenge to pursue before attaining the regulatory
non-target organisms, including fish, odonates, waterbugs and crusta- approval and marketing of a botanical insecticide.
ceans (including copepods), with extremely high LC50 values. For in-
stance, Govindarajan and Benelli (2016b) outlined that the aqueous leaf 8. Field experiments testing plant extracts as larvicides
extract of Barleria cristata L. was scarcely toxic to three non-target
aquatic species, i.e., Anisops bouvieri, Diplonychus indicus and Gambusia In the paragraphs above, we have outlined how this review sum-
affinis, with LC50 values ranging from 6231 to 8596 μg/mL, while their marized knowledge arising from several hundreds of researches carried
toxicity against 3rd instar larvae of An. subpictus, Ae. albopictus and Cx. out on plant extracts as larvicides in laboratory conditions. The findings
tritaeniorhynchus ranged from 124 to 146 μg/mL. Comparable results are promising, therefore one may argue: did researchers validated the
were also obtained testing the aqueous leaf extracts of Clerodendrum bioactivity of these plant extracts in the field? To the best of our
chinense (Osbeck) Mabb. (Govindarajan et al., 2016d) and Carissa spi- knowledge, only in a limited number of papers the larvicidal activity of
narum L. (Govindarajan et al., 2016g) against the same three non-target plant extracts has been evaluated in the field, mostly relying on ex-
species, with LC50 values in the ranges 3111–6877 μg/mL and periments against Anopheles, Aedes and Culex species in storage water
4146–6403 μg/mL, respectively. A further good example is the aqueous tanks and sewage water bodies.
leaf extract of Nicandra physalodes (L.) Gaertn., which achieved a very Earlier, Aarthi and Murugan (2010) showed that after 72 h from a
high LC50 value, 19.057 μg/mL, on adults of the non-target waterbug D. treatment with Vetiveria zizanioides (L.) Nash and O. basilicum metha-
indicus (Govindarajan et al., 2016f), while the LC50 of the leaf aqueous nolic extracts, a > 90% reduction of An. stephensi larval populations in
extract of Hymenodictyon orixense (Roxb.) Mabb. on the same waterbug overhead water tanks, cement tanks and cement containers was ob-
species was 6233 μg/mL (Govindarajan and Benelli, 2016c) tained. Then, Kamalakannan et al. (2011) outlined that the treatment of
In some cases, the toxicity towards non-target species is closer to water storage tanks (about 0.5 m3) in the field with Acalypha indica L.
that observed on targeted mosquito young instars. However, still with and Achyranthes aspera L. combined methanolic extracts (i.e., 50 + 50 g
significant differences among them. Kumar et al. (2016) outlined that of extracts plus 0.5 g of 0.05% Teepol® emulsifying agent per 1 L)
261
reduced Ae. aegypti larval (3rd instar and 4th instar) and pupal popula- borne terpenes active against a rather wide number of crop pests and
tions by 97% and 81%, respectively, after 5 days. characterized by 0-day pre-harvest interval). Besides, garlic-based in-
Arjunan et al. (2012) tested Cynodon dactylon (L.) Pers., Aloe vera secticidal formulations have been developed in US and UK by SME and
(L.) Burm.f., Hemidesmus indicus (L.) R. Br. ex Schult. and Coleus am- used to control agricultural pests (Isman and Machial, 2006). More
boinicus Lour. ethanolic extracts at a high concentration (51 g/L) recently, an extract made with garlic oil and capsicum oleoresin has
against An. stephensi in overhead water tanks, as well as on Ae. aegypti in been marketed and approved by EPA in the US (Isman, 2015). In India
storage water tanks and against Cx. quinquefasciatus in sewage water and China, which are considered leader countries for development and
systems. Concerning trials on An. stephensi, none of the tested extracts marketing of botanical insecticides, several plant extracts (e.g. Milletia
achieved 90% larval mortality 48 h post-treatment, while mortality pinnata (Roxb.) Steud., Apocynum venetum L., Tripterygium wilfordii
values higher than 90% were obtained testing C. dactylon and H. indicus Hook. f., A. squamosa, Sophora flavescens Aiton, Veratrum nigrum L.,
extracts on Ae. aegypti, as well as C. dactylon, H. indicus and A. vera Celastrus angulatus Maxim.) have been recently commercialised (Isman,
extracts on Cx. quinquefasciatus (Arjunan et al., 2012). Moreover, 2015). However, comparable plant extract-based products currently
Kovendan et al. (2012d) also tested extracts of Jatropha curcas L., Hyptis registered against mosquito larvae are sadly lacking.
suaveolens (L.) Poit., Abutilon indicum (L.) Sweet and Leucas aspera Looking at the botanical products developed to manage arthropods
(Willd.) Link against this mosquito species, using the same field method of medical and veterinary importance, a major effort has been done
described above. L. aspera, A. indicum and H. suaveolens achieved 99.7, about repellents. An important example is the efficacy of p-menthane-
92 and 90.4% larval mortality after 72 h, respectively. Complete elim- 3,8-diol, which is related to that of essential oils obtained from the
ination of An. stephensi and Ae. aegypti larval populations in field water lemon-scented leaves of eucalyptus tree [e.g., Corymbia citriodora
bodies (2.0 × 1.8 × 1.4 m) was achieved within 72 h testing the leaf (Hook.) K.D.Hill & L.A.S.Johnson] (Carroll and Loye, 2006). In United
extract of Acalypha alnifolia Klein ex Willd. plus 0.05% Teepol® States and Europe, there are repellent products on the market con-
(Kovendan et al., 2013). In these two studies, the solvent used to pre- taining oils of lemon eucalyptus (Semmler et al., 2014). Besides, neem
pare the extracts tested in the field was not specified. However, it looks seed constituents have been widely exploited to develop a number of
conceivable that the authors used methanol, since LC50 of methanolic products (e.g., Tre-san® and MiteStop® developed by the university spin-
plant extracts were the lowest ones in laboratory assays presented in off AlphaBioCare, Düsseldorf) highly effective against various pests and
both articles. vectors, including house dust mites, poultry mites, harvest mites, Ixodes
Field experiments aimed to control Cx. quinquefasciatus larvae in and Rhipicephalus ticks, cat fleas, bed bugs, head lice and Mallophaga,
sewage water bodies (2.0 × 1.8 × 1.4 m) pointed out that combining cockroaches (Blatta, Blattella, Gomphadorhina) and raptor bugs (Tria-
the S. xanthocarpum leaf ethanolic extract with B. thuringiensis israelensis toma), among others (Schmahl et al., 2010). On the other hand, plant
(both at their LC90 estimated in laboratory conditions plus 0.05% extract-based commercial larvicides to be used against mosquitoes are
Teepol®) achieved 100% mortality after 72 h, while the extract tested not available. This is a major issue deserving further attention by ap-
alone achieved 94% larval reduction after 72 h from the treatment plied entomologists worldwide.
(Kumar et al., 2012a). Also, the fruit methanol extract of the same plant One of the possible reasons behind the limited expansion of bota-
species at a sub-lethal concentration increased the predation of Meso- nical insecticides is certainly the expensive and time-consuming pro-
cyclops thermocyclopoides copepods against Ae. aegypti (particularly cedures to get the regulatory approval. To date only the large agro-
against first and third instar larvae), probably through reduction of chemical companies (multinational) can afford it. Consequently, they
larval motility (Kumar et al., 2012b). preferred to develop a new generation of synthetic products (e.g.,
Later, 3rd and 4th instar larval populations of various mosquito neonicotinoids) with lower toxicity respect to the previous ones (e.g.,
vectors were eliminated from external water storage reservoirs (250 L) nicotine and pyrethroids) and that displayed a favourable cost/benefit
after 72 h from the treatment with the extracts of Mimusops elengi L. ratio compared with botanical insecticides (Isman, 2008). However, the
(target: An. stephensi and Ae. albopictus, aqueous extract, Subramaniam overuse of these products revealed to be deleterious to many non-target
et al., 2015), Phyllanthus niruri L. (target: Ae. aegypti, aqueous extract, organisms accompanied by environment pollution and human health
Suresh et al., 2015), Pteridium aquilinum (L.) Kuhn (target: An. stephensi, concerns and caused the development of many forms of resistance in
ethanolic extract, Panneerselvam et al., 2016), C. ambrosioides (target: insects.
Ae. albopictus, aqueous extract, Subramaniam et al., 2017) and Suaeda
maritima (L.) Dumort. leaves (target: Ae. aegypti, aqueous extract, 10. Conclusions, research perspectives and challenges for real-
Suresh et al., 2018). However, in these studies, the tested concentration world applications
was rather high (10xLC50). Larval elimination was also obtained within
72 h from the treatement testing the Couroupita guianensis Aubl. flower This comprehensive survey allowed to identify a batch of extremely
aqueous extract (Subramaniam et al., 2016) and the Aristolochia indica effective plant extracts and their chemical markers with a relevant
L. leaf aqueous extract (Murugan et al., 2015e) against An. stephensi 3rd potential in mosquito control programs. The botanical families en-
and 4th instar larvae in similar field experiments. compassing the highest number of larvicidal plant extracts are re-
Overall, it can be observed that the field studies validating the ef- presented by Asteraceae, Fabaceae and Lamiaceae, which display a long
ficacy of plant extracts and their selected constituents as mosquito history as either food or medicine all around the world. In developing
larvicides remain scarce, if compared to the overall scientific produc- countries where the mosquito-borne diseases are endemic, the usage of
tion on the same botanicals as mosquito larvicides in laboratory con- these plant extracts and essential oils should be facilitated – pending
ditions; this strongly limits the development of products for real-world formulation in stable and synergized blends and proper field validation
use (Semmler et al., 2009; Pavela and Benelli, 2016a,b). of their epidemiological impact (see Benelli and Beier, 2017; Fernandes
et al., 2018; Wilke et al., 2018) – in the years to come by the availability
9. Does this knowledge help to develop commercial products? of low-cost extraction instruments and appropriate formulation tech-
nology, including microencapsulation, microemulsions and nano-
Starting from the field efficacy evidences reported above, a further formulations (Benelli, 2016, 2018c; Pavela et al., 2019a,b). On the
question could be: which larvicidal products have been developed for other hand, in industrial nations the renaissance of plant-borne in-
real-world commercial purposes? Unfortunately, little has been done in secticides may be the natural consequence of the failure of the strate-
this framework. Botanical products have been widely exploited to de- gies adopted by multinational agrochemical companiens in the latest
velop commercial insecticides to be used against important agricultural years.
pests (e.g., Requiem®, recently developed by Bayer based on plant- The compounds characterizing the most active extracts belong to
262
different chemical groups and possess different mechanisms of action in Anopheles stephensi Liston (Insecta: diptera: culicidae). J. Biopesticides 3, 199–204.
insects, although several extracts still deserve phytochemical in- Aarthi, C., Govindarajan, M., Rajaraman, P., Alharbi, N.S., Kadaikunnan, S., Khaled, J.M.,
et al., 2018. Eco-friendly and cost-effective Ag nanocrystals fabricated using the leaf
vestigation. These compounds can be valuable as alternative to con- extract of Habenaria plantaginea: toxicity on six mosquito vectors and four non-target
ventional products as well as in combination with them. Comparing species. Environ. Sci. Pollut. Res. 25, 10317–10327.
with synthetic products that exert commonly neurotoxic effects and Abirami, D., Kovendan, K., Chandramohan, B., 2017. Ovicidal and larvicidal properties of
Pergularia extensa and Spermacoce hispida ethanol root extracts on Anopheles stephensi
promote the development of cross-resistance in insects, the constituents Liston. Southeast Asian J. Trop. Med. Public Health 48, 1188–1199.
of extracts reported in this survey may also have other modes of action, Abou-Elnaga, Z.S., Alarif, W.M., Al-Iihaibi, S.S., 2011. New larvicidal acetogenin from the
including the inhibition of mitochondrial respiration and detoxification red alga Laurencia papillosa. Clean (Weinh) 39, 787–794.
AhbiRami, R., Zuharah, W.F., Thiagaletchumi, M., Subramaniam, S., Sundarasekar, J.,
enzymes (e.g., transferases, esterases and monooxygenase), antifeedant 2014. Larvicidal efficacy of different plant parts of railway creeper, Ipomoea carica
activity, phototoxicity, generation of ROS, midgut damages and dis- extract against dengue vector mosquitoes, Aedes albopictus (Diptera: culicidae) and
ruption of embryonic development. Within this batch of compounds, Aedes aegypti (Diptera: culicidae). J. Insect Sci. 14, 80.
Ahn, J.Y., Lee, J.Y., Yang, E.J., Lee, Y.J., Koo, K.B., Song, K.S., Lee, K.Y., 2013.
only those obtained from plants with a long history of use as food and
Mosquitocidal activity of anthraquinones isolated from symbiotic bacteria
flavoring and having high LD50 values after oral administration in rats Photorhabdus of entomopathogenic nematode. J. Asia Pac. Entomol. 16, 317–320.
should be selected due to safety concern and facilitation in regulatory Aivazi, A.A., Vijayan, V.A., 2009. Larvicidal activity of oak Quercus infectoria Oliv.
approval. Further investigations are encouraged on these compounds (Fagaceae) gall extracts against Anopheles stephensi Liston. Parasitol. Res. 104,
1289–1293.
with the aim to develop effective, safe and eco-friendly micro- and Aivazi, A.A., Vijayan, V.A., 2010. Efficacy of Ruta graveolens extract and its synergistic
nanoemulsions for the control of mosquito vector of medical relevance. effect with cypermethrin against Anopheles stephensi Liston larvae. Toxicol. Environ.
Finally, the regulatory framework is expected to change in the years to Chem. 92, 893–901.
Alam, M.F., Safhi, M.M., Chopra, A.K., Dua, V.K., 2011. Toxicological properties of sev-
come allowing to facilitate the marketing of botanical larvicides (Pavela eral medicinal plants from the Himalayas (India) against vectors of malaria, filariasis
and Benelli, 2016a,b). and dengue. Trop. Biomed. 28, 343–350.
Unfortunately, despite the abundance of laboratory data stressing Alayo, M.A., Femi-Oyewo, M.N., Bakre, L.G., Fashina, A.O., 2015. Larvicidal potential
and mosquito repellent activity of Cassia mimosoides extracts. Southeast Asian J. Trop.
the efficacy of plant extracts against mosquito young instars, less than Med. Public Health 46, 596–601.
twenty research items shed light on their effectiveness in the field (as Ali, S.I., Gopalakrishnan, B., Venkatesalu, V., 2018. Evaluation of larvicidal activity of
detailed in paragraph 8). In addition, the epidemiological impact on Senecio laetus Edgew. Against the malarial vector, Anopheles stephensi, dengue vector,
Aedes aegypti and Bancroftian filariasis vector, Culex quinquefasciatus. S. Afr. J. Bot.
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extracts as mosquito larvicides has not been evaluated. This really Al-Mekhlafi, F.A., Abu Taha, N., Mashaly, A.M.A., Wadaan, M.A., 2013. Larvicidal ac-
limits the development of commercial mosquito larvicides of botanical tivity of selected xerophytic plants against Culex pipiens and Aedes caspius (Diptera:
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Al-Mekhlafi, F.A., Abutaha, N., Mashaly, A.M.A., Nasr, F.A., Ibrahim, K.E., Wadaan, M.A.,
decades to kill or repell other key arthropod species of medical and 2017. Biological activity of Xanthium strumarium seed extracts on different cancer cell
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AlQahtani, F.S., AlShebly, M.M., Govindarajan, M., Senthilmurugan, S., Vijayan, P.,
2019b). Further research on these issues is urgently needed. Benelli, G., 2017. Green and facile biosynthesis of silver nanocomposites using the
Overall, we believe that the possibility to develop effective, eco-friendly aqueous extract of Rubus ellipticus leaves: toxicity and oviposition deterrent activity
and safe botanical insecticides starting from active ingredients found in against Zika virus, malaria and filariasis mosquito vectors. J. Asia Pac. Entomol. 20,
157–164.
these plant extracts is highly likely. Moreover, these prospects are enhanced Al-Rehaily, A.J., Ahmad, M.S., Yousaf, M., Ahmed, S., Nur-e-Alam, M., Al-Dosari, M.S.,
by the fact that some from the most effective extracts come from plants Parvez, M.K., 2017. Chemical constituents of Cleome rupicola growing in Saudi
which could be grown as a commercial crop (e.g., A. visnaga, Atalantia Arabia. Chem. Nat. Comp. 53, 670–673.
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monophyla DC., T. africanum, T. minuta, and S. acmella), and provided that a R.A., et al., 2018. Swift fabrication of Ag nanostructures using a colloidal solution of
suitable growing technology is used, a relatively large amount of dry mass Holostemma ada-kodien (Apocynaceae)–Antibiofilm potential, insecticidal activity
can be obtained from one hectare. These crops may provide a well available against mosquitoes and non-target impact on water bugs. J. Photochem. Photobiol. B,
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and relatively inexpensive source of active substances for potential botanical
Amel, A., Sélima, B., 2015. Larvicidal effect of Marrubium vulgare on Culex pipiens in
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Author contributions
Amerasan, D., Nataraj, T., Murugan, K., Panneerselvam, C., Madhiyazhagan, P., Nicoletti,
M., Benelli, G., 2016. Myco-synthesis of silver nanoparticles using Metarhizium ani-
All authors conceived and designed the research. RP, FM and GB sopliae against the rural malaria vector Anopheles culicifacies Giles (Diptera: culi-
collected and analysed the literature data. All authors wrote the cidae). J. Pest Sci. 89, 249–256.
Anbazhagan, P., Murugan, K., Jaganathan, A., Sujitha, V., Samidoss, C.M., Jayashanthani,
manuscript. S., et al., 2017. Mosquitocidal, antimalarial and antidiabetic potential of Musa
paradisiaca-synthesized silver nanoparticles: in vivo and in vitro approaches. J. Clust.
Conflict of interest Sci. 28, 91–107.
Andemo, A., Yewhalaw, D., Alemayehu, B., Ambelu, A., 2014. Evaluation of the mos-
quitocidal effect of Birbira (Mellitia ferruginea) seed extract against Anopheles ara-
The authors declare no competing interests. biensis (Diptera: culicidae) from Ethiopia. Acta Trop. 136, 68–73.
Arjunan, N., Murugan, K., Madhiyazhagan, P., Kovendan, K., Prasannakumar, K.,
Thangamani, S., Barnard, D.R., 2012. Mosquitocidal and water purification proper-
Acknowledgements ties of Cynodon dactylon, Aloe vera, Hemidesmus indicus and Coleus amboinicus leaf
extracts against the mosquito vectors. Parasitol. Res. 110, 1435–1443.
The authors are grateful to the Prof. John C. Beier for inviting this Ashokan, A.P., Paulpandi, M., Dinesh, D., Murugan, K., Vadivalagan, C., et al., 2017.
Toxicity on dengue mosquito vectors through Myristica fragrans-synthesized zinc
Review on Acta Tropica. Dr. Roman Pavela would like to thank the
oxide nanorods, and their cytotoxic effects on liver cancer cells (HepG2). J. Clust. Sci.
Ministry of Agriculture of the Czech Republic for its financial support 28, 205–226.
concerning botanical pesticide and basic substances research (Project Azarudeen, R.M.S.T., Govindarajan, M., Amsath, A., Kadaikunnan, S., Alharbi, N.S.,
Vijayan, P., et al., 2016. Size-controlled fabrication of silver nanoparticles using the
No. RO0418). The Cx. quinquefasciatus photo used in the graphical
Hedyotis puberula leaf extract: toxicity on mosquito vectors and impact on biological
abstract was kindly provided by Dr. Harry Weinburgh (PHIL-CDC). control agents. RSC Adv. 6, 96573–96583.
Azarudeen, R.M.S.T., Govindarajan, M., AlShebly, M.M., AlQahtani, F.S., Amsath, A.,
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Acta Tropica 193 (2019) 236–271

Contents lists available at ScienceDirect

Plant extracts for developing mosquito larvicides: From laboratory to the T

ARTICLE INFO ABSTRACT

1. Introduction Despite decades of intensive research, the effective and sustainable

IVM programs. alkaloids, alkamides, anthraquinones, coumarins, flavonoids, limonoids,

weight formula chemical group

(continued on next page)

2012; Vats, 2018; Santiago et al., 2012;

2009; Lu et al., 2017; Liang et al., 2015

Yenesew et al., 2003; Yenesew et al.,

Ibrahim et al., 2018; Marles et al.,

Macel, 2011; Omar et al., 1983

1991; Perich et al., 1995

terase activities (Larson et al., 2010; Kumar et al., 2010). A photo-induced

toxicity of α-mangostin in water was also recorded (Ibrahim et al., 2016).

quefasciatus, respectively (Larson et al., 2010).

3.2 (An. stephensi)

1.6 (Ae. aegypti)

Flavonoids represent a large class of plant secondary metabolites de-

activity; interaction with GABAergic systme

antifeedant activity; genotoxicity and

oxidoreductase; inhibition of protein

at position 3, isoflavones are formed. The variability in functionalization of

anthocyanins, flavan-3-ols and flavanones, among others) (Treutter, 2006).

important role as ecological mediators, giving an important contribution

also in the plant defense against microorganisms and insects (Treutter,

proved to affect the detoxification system of insects through reduction of

glutathione S-transferase (GST) and esterase activities (Baskar et al., 2014).

et al., 2012, 2015).

Rotenoids are particular forms of isoflavonoids produced by plants to

Lonchocarpus and Tephrosia, all belonging to the Fabaceae family. It is

marketed as a natural insecticide in several countries (Pesticide Action

oxidoreductase activity making insect unable of getting energy and causing

arrest of feeding (El-Wakeil, 2013). Tested on mosquito larvae, rotenone

spectively (Belmain et al., 2012). Thus, these two new-characterized rote-

noids from T. vogelii are promising ingredients to replace rotenone in in-

secticidal formulations. However, furthers studies supporting their lower

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