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Two-dimensional chromatogram of GC×GC/MS analysis of Bactrocera dorsalis and B. carambolae cuticular profiles. Cuticular profiles of B. carambolae female (A) and male (B) and of B. dorsalis female (C), male (D). Intensity of the signals is colour coded from blue (zero) to red (maximum). The compounds are assigned according to S1 Table.
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Bactrocera invadens Drew, Tsuruta & White, Bactrocera papayae Drew & Hancock, and Bactrocera philippinensis Drew & Hancock, key pest species within the Bactrocera dorsalis species complex, have been recently synonymized under the name Bactrocera dorsalis (Hendel). The closely related Bactrocera carambolae Drew & Hancock remains as a discrete taxono...
Citations
... EL, EM, and EP have been found in Bactrocera correcta (33), Bactrocera oleae (34), Bactrocera frauenfeldi (35), Bactrocera musae (36), Bactrocera bryoniae (37), and Bactrocera tryoni (38) female extracts. In addition, EL, EM, and EP were found in the cuticle of mature female B. tryoni, Bactrocera carambolae, Bactrocera papaya, Bactrocera philippinensis and Bactrocera invadens, with varying relative abundances among different species (39,40), suggesting that these compounds might have roles in chemical rec ognition. Notably, EL, EM, and EP elicited electrophysiological responses in male and female B. bryoniae (37) and B. oleae (34), but only EL elicited responses in male B. musae (36), which reveals the functional divergence of these fatty acid esters in different tephritid flies. ...
Chemical communication plays a vital role in mate attraction and discrimination among many insect species. Here, we document a unique example of semiochemical parsimony, where four chemicals act as both aphrodisiacs and anti-aphrodisiacs in different contexts in Bactrocera dorsalis . Specifically, we identified four female-specific semiochemicals, ethyl laurate, ethyl myristate, ethyl cis -9-hexadecenoate, and ethyl palmitate, which serve as aphrodisiacs to attract male flies and arouse male courtship. Interestingly, these semiochemicals, when sexually transferred to males during mating, can function as anti-aphrodisiacs, inhibiting the receptivity of subsequent female mates. We further showed that the expression of elongase11, a key enzyme involved in the biosynthesis of these semiochemicals, is under the control of doublesex, facilitating the exclusive biosynthesis of these four semiochemicals in females and guaranteeing effective chemical communication. The dual roles of these semiochemicals not only ensure the attractiveness of mature females but also provide a simple yet reliable mechanism for female mate discrimination. These findings provide insights into chemical communication in B. dorsalis and add elements for the design of pest control programs.
... These non-polar compounds serve primarily to protect insects from desiccation (Blomquist and Bagnères 2010). They can also play a major role in inter-and intra-specific recognition, especially mate recognition and territorial displays (Blomquist and Bagnères 2010;Everaerts et al. 2010;Benelli 2015b;Vaníčková et al. 2017). In tephritid flies CHs have been used for species differentiation in cryptic species complexes, such as the Anastrepha fraterculus complex or the so-called Ceratitis FAR complex (Vaníčková 2012;Vaníčková et al. 2014Vaníčková et al. , 2015a (Vaníčková et al. 2017). ...
... They can also play a major role in inter-and intra-specific recognition, especially mate recognition and territorial displays (Blomquist and Bagnères 2010;Everaerts et al. 2010;Benelli 2015b;Vaníčková et al. 2017). In tephritid flies CHs have been used for species differentiation in cryptic species complexes, such as the Anastrepha fraterculus complex or the so-called Ceratitis FAR complex (Vaníčková 2012;Vaníčková et al. 2014Vaníčková et al. , 2015a (Vaníčková et al. 2017). In addition to CHs, abundant complex mixtures of sex-specific oxygenated lipids (3 fatty acids and 22 fatty acid esters) with unknown functions have been identified in epicuticular extracts from B. dorsalis and B. carambolae females. ...
... However, the reproductive isolation between species may be linked to rapid changes in the olfactory pathways involved in pheromone detection (Dekker et al. 2015). Among Dacinae species, many cuticular compounds are shared or derived (Baker and Bacon 1985;Carlson and Yocom 1986;Goh et al. 1993;Fletcher and Kitching 1995;Benelli et al. 2014a;Vaníčková et al. 2017;Noushini et al. 2020), and the olfactory system is largely conserved at the molecular and functional levels (Jacob et al. 2017). This also suggests that pheromone emission and detection may evolve gradually. ...
The melon fly, Zeugodacus cucurbitae (Coquillett) (Diptera: Tephritidae), is considered to be the most destructive pest of melons and other related cucurbit crops worldwide. Despite the potential of behaviour-based control strategies, little is known about the mechanisms involved in female mate choice. Herein, we investigated the production and chemoreception of cuticular hydrocarbons in both sexes of Z. cucurbitae, and the behavioural responses they induce. We studied the epicuticular composition of virgin males and females, using two-dimensional gas chromatography coupled with mass spectrometric detection. Data were interpreted using multivariate factorial analysis. The differentiation of chemical profiles was consistently observed over time. In young individuals, the chemical profiles did not differ between sexes, while sex-specific differences were noted in mature flies. The fly olfactory sensitivity to these compounds was explored using gas chromatography combined with chopped triple electroantennography and electropalpography detectors. This extensive exploration of the pest olfactory sensitivity highlighted three compounds produced by the male. When blended, they induced a robust positive response in unmated naive females in a six-choice olfactometer. The responsiveness of other Tephritidae species (a polyphagous species Bactrocera dorsalis (Hendel) and the cucurbit specialist Dacus demmerezi (Bezzi)) to whole body extracts of Z. cucurbitae was also investigated. Our findings showed that Z. cucurbitae uses species-specific olfactory receptors to detect male produced compounds. In addition, the palps were sensitive to a female-specific component, 1,7-dioxaspiro[5.5]undecane, which the males produce in minute quantities. Overall, this study provides a starting point for a pheromone-based tephritid lure that targets unmated females. The potential implications for pest management are discussed.
... ranging from 0.55% -to 5.82%). Vaničkova et al., (2017) stated that the chemical constituents of Bactrocera dorsalis epicuticle comprising of fatty acid esters, aliphatic hydrocarbons, aldehydes, and fatty acids were qualitatively and quantitatively different in both sexes. The present investigation adds aromatic hydrocarbons to the recorded compounds in the cuticle. ...
Cuticular hydrocarbons are present on the surface of all insects and play an important role in the life of insects. Although primarily anti-desiccation agents, cuticular hydrocarbons are emerging as important chemicals in insect communication. Using Cuticular hydrocarbons of insect cuticles is important in taxonomy because they have a wide variety of chemical compounds e.g., hydrocarbons, monoester waxes, triglycerides, and free fatty acids. So cuticular chemical components are a precise tool for chemotaxonomy, and they can be used as an accompaniment to morphology and genetic characters in phylogenetic studies. Solitary bees (leaf-cutting) of Anthidium Fabricius, 1804 are economically important in natural and agroecosystems and play an important role in pollinating many domesticated and wild plant species. This study aimed to describe the cuticular chemical profile of males of two species of the genus Anthidium to be applied as a chemotaxonomic tool. The investigation used gas chromatography coupled to mass spectrometry (GC-MS). Investigated specie revealed 40 compounds in total, with nine compounds in common. The chemical diversity was higher in, A. tessellatum (30 compounds) while, in A. pulchellum (19 compounds). Males of A. tessellatum were distinguished by twenty-one exclusive compounds, and males of A. pulchellum were characterized by ten compounds. Most abundant compounds and that represented by scarce quantity were recorded for each species.
... The retention time and the Kovats retention index of extracts were used to identify alkanes by comparing to the retention time of C 7 -C 40 n-alkanes standards. The peak areas of these alkanes were used for quantification by comparing them to the internal standard peak area (Vanickova et al., 2017). ...
... They can also be chemotaxonomic markers for insect species, biotypes and population identification. In B. dorsalis, this difference can be seen in the different contents of CHCs in females and males, as well in the results of the present study(Vanickova et al., 2017).As commonly found in Diptera, B. dorsalis has two types of CYP4G genes, as does D. melanogaster. CYP4G100 (CYP4G1 type) also has a similar expression pattern as CYP4G1. ...
The oriental fruit fly Bactrocera dorsalis (Hendel) is expanding its distribution to higher latitudes. Our goal in this study was to understand how B. dorsalis adapts to higher latitude environments that are more arid than tropical regions. Cuticular hydrocarbons (CHCs) on the surface of the epicuticle in insects act as a hydrophobic barrier against water loss. The essential decarbonylation reaction in CHC synthesis is catalyzed by CYP4G, a cytochrome P450 subfamily protein. Hence, in B. dorsalis it is necessary to clarify the function of the CYP4G gene and its role in desiccation resistance. CYP4G100 was identified in the B. dorsalis genome. The complete open reading frame (ORF) encodes a CYP4 family protein (552 amino acid residues) that has the CYP4G‐specific insertion. This CYP4G gene was highly expressed in adults, especially in the oenocyte‐rich peripheral fat body. The gene can be induced by desiccation treatment, suggesting its role in CHC synthesis and waterproofing. Silencing of CYP4G100 resulted in a decrease of CHC levels and the accumulation of triglycerides. It also increased water loss and resulted in higher desiccation susceptibility. CYP4G100 is involved in hydrocarbon synthesis and contributes to cuticle waterproofing to help B. dorsalis resist desiccation in arid environments.
... Cuticular hydrocarbons have been studied principally in insects from Diptera, Hymenoptera, and Coleoptera (Blomquist & Bagnères 2010). For instance, studies have identified the cuticular hydrocarbons of larvae and adults from tephritid fruit flies, including species of the genera Ceratitis, Bactrocera, and Anastrepha (Carlson & Yocom 1986;Sutton & Carlson 1993;Vaníčková et al. 2014Vaníčková et al. , 2015Vaníčková et al. , 2017Bosa et al. 2018). The principal objective of these studies was to identify the cuticular hydrocarbons as a tool for chemotaxonomical discrimination against the cryptic species (Carlson & Yocom 1986;Vaníčková et al. 2014Vaníčková et al. , 2015Vaníčková et al. , 2017. ...
... For instance, studies have identified the cuticular hydrocarbons of larvae and adults from tephritid fruit flies, including species of the genera Ceratitis, Bactrocera, and Anastrepha (Carlson & Yocom 1986;Sutton & Carlson 1993;Vaníčková et al. 2014Vaníčková et al. , 2015Vaníčková et al. , 2017Bosa et al. 2018). The principal objective of these studies was to identify the cuticular hydrocarbons as a tool for chemotaxonomical discrimination against the cryptic species (Carlson & Yocom 1986;Vaníčková et al. 2014Vaníčková et al. , 2015Vaníčková et al. , 2017. In contrast, relatively few studies have investigated the dynamic of these compounds in relation to abiotic and biotic factors (Sutton & Carlson 1993;Vaníčková et al. 2012;Bosa et al. 2018). ...
The primary function of cuticular hydrocarbons is to keep insects from losing water. However, cuticular hydrocarbons also may mediate chemical communication in a number of species. In this study, we investigated the effect of the extraction method, natal host, and age (maturation) on the cuticular hydrocarbon profiles of the West Indian fruit fly, Anastrepha obliqua (Macquart) (Diptera: Tephritidae). Cuticular hydrocarbons from female and male adults of different natal hosts (Mangifera indica L. or Spondias mombin L. [both Anacardiaceae]) and age were extracted by solvent extraction and direct contact solid-phase microextraction. Cuticular hydrocarbons were identified by gas chromatography coupled to mass spectrometry. In total, we recorded 12 compounds, but only 9 of them were identified. The identified cuticular hydrocarbons were linear alkanes (n-heneicosane, n-nonacosane, and n-hentriacontane), alkenes (n-heneicosene, n-tricosene, n-nonacosene, and n-hentriacontene), and branched alkanes (2-methyl-octacosane and 2-methyl-triacontane). There were no qualitative differences between sampling techniques. The solvent extraction method extracted more cuticular hydrocarbons from flies reared on mango compared to those extracted from flies reared on hog plum. In contrast, solid-phase microextraction extracted a higher concentration of cuticular hydrocarbons from flies reared on hog plum than to those extracted from flies reared on mango. Higher levels of 2-methyl-octacosane and 2-methyl-triacontane were detected with solvent extraction than with solid-phase microextraction; the opposite occurred with hentriacontane and unknown compound 2. The compounds n-heneicosene, n-heneicosane, and n-tricosene were present in mature males but not in mature females; n-nonacosene was found only in the mature flies of both sexes.
... In tephritids, sex-and species-specificity of CHs have been described in species of the genera Anastrepha, Bactrocera and Ceratitis (Table 4; Table S4). [89,272,412,[414][415][416]418,419]. Nevertheless, studies focused on the elucidation of CHs behavioural function in fruit fly mating system are still missing, except for those performed on B. oleae [79,84,286]. ...
... In species of the B. dorsalis complex, abundant complex mixtures of sex-specific oxygenated lipids (i.e., three fatty acids and 22 fatty acid esters) with so far unknown biological function were identified in epicuticle extracts from females [419]. Such sexspecificity may be driven by sexual selection if the chemical composition of the cuticle is used as a pheromone signal in mate choice. ...
... Preparation of samples for gas chromatographic analysis of tephritid CHs is usually made by solvent extraction of whole insect bodies with solvents such as pentane, hexane, dichlormethane, and chloroform [89,425]. The most widely used method for recovering tephritid CHs is hexane washing [272,419,426]. This process may contaminate the cuticular sample with other materials, such as those from the exocrine glands. ...
The Dipteran family Tephritidae (true fruit flies) comprises more than 5000 species classified in 500 genera distributed worldwide. Tephritidae include devastating agricultural pests and highly invasive species whose spread is currently facilitated by globalization, international trade and human mobility. The ability to identify and exploit a wide range of host plants for oviposition, as well as effective and diversified reproductive strategies, are among the key features supporting tephritid biological success. Intraspecific communication involves the exchange of a complex set of sensory cues that are species- and sex-specific. Chemical signals, which are standing out in tephritid communication, comprise long-distance pheromones emitted by one or both sexes, cuticular hydrocarbons with limited volatility deposited on the surrounding substrate or on the insect body regulating medium- to short-distance communication, and host-marking compounds deposited on the fruit after oviposition. In this review, the current knowledge on tephritid chemical communication was analysed with a special emphasis on fruit fly pest species belonging to the Anastrepha, Bactrocera, Ceratitis, and Rhagoletis genera. The multidisciplinary approaches adopted for characterising tephritid semiochemicals, and the real-world applications and challenges for Integrated Pest Management (IPM) and biological control strategies are critically discussed. Future perspectives for targeted research on fruit fly chemical communication are highlighted.
... Some aspects of tephritid fruit fly semiochemistry have received significant attention, particularly the pheromones [34,35] they use to attract mates and for aggregation and the particular compounds found in fruit, food and certain flowers to which they are attracted [36][37][38][39][40][41][42][43][44][45][46][47]. Some work has also been performed on fruit fly cuticular chemistry because their cuticular chemical profiles tend to be highly species-specific [48,49] and have been used to resolve species, cryptic species and geographic variation in larvae [50][51][52] and adults [53][54][55][56][57][58]. Beyond their use as chemotaxonomic tools, however, relatively little work has been performed on tephritid cuticular compounds. ...
... The solvent choice of n-hexane was based on anticipated polarities of insect cuticular compounds that are generally less or non-polar [99]. The method used in the present study is more similar to methods widely used in studies of the cuticular hydrocarbons of other tephritids [53][54][55][56][57][58][59]100]. ...
... The C 12 to C 20 esters we found in cuticle of mature female B. tryoni have also been reported in the cuticle of females in five B. dorsalis complex species, but at different relative abundances between the species, suggesting that they may have a role in species recognition [58,59]. On the other hand, the methyl branched fatty acid esters found in the present study have not been previously reported in either the emission profile or rectal gland extracts of B. tryoni [67], or other fruit flies. ...
The cuticular layer of the insect exoskeleton contains diverse compounds that serve important biological functions, including the maintenance of homeostasis by protecting against water loss, protection from injury, pathogens and insecticides, and communication. Bactrocera tryoni (Froggatt) is the most destructive pest of fruit production in Australia, yet there are no published accounts of this species' cuticular chemistry. We here provide a comprehensive description of B. tryoni cuticular chemistry. We used gas chromatography-mass spectrometry to identify and characterize compounds in hexane extracts of B. tryoni adults reared from larvae in naturally infested fruits. The compounds found included spiroacetals, aliphatic amides, saturated/unsaturated and methyl branched C12 to C20 chain esters and C29 to C33 normal and methyl-branched alkanes. The spiroacetals and esters were found to be specific to mature females, while the amides were found in both sexes. Normal and methyl-branched alkanes were qualitatively the same in all age and sex groups but some of the alkanes differed in amounts (as estimated from internal standard-normalized peak areas) between mature males and females, as well as between mature and immature flies. This study provides essential foundations for studies investigating the functions of cuticular chemistry in this economically important species.
... Furthermore, under conditions of high competition for mates, males of C. capitata increase courtship behavior in comparison with those males that were under low competition (Leftwich et al., 2012). On the other hand, differences in the CHC profiles among populations and also between sexes have been observed in A. fraterculus, Ceratitis rosa Karsch, and also among populations of Bactrocera dorsalis Hendel ( Van ı ckov a et al., 2012Van ı ckov a et al., , 2014Van ı ckov a et al., , 2015Van ı ckov a et al., , 2017. Differences in CHC profiles between sexes and among flies of different ages have been reported in A. fraterculus and in A. ludens (Van ı ckov a et al., 2012; Bosa et al., 2018). ...
... In D. serrata, female presence in groups of males induces changes in CHC expression, pheromones, and locomotive activity, which are conditions that favor male mating success (Krupp et al., 2008). CHC profiles vary between populations of C. rosa, and also among populations of the B. dorsalis complex ( Van ı ckov a et al., 2014Van ı ckov a et al., , 2015Van ı ckov a et al., , 2017. CHC profiles of A. fraterculus vary among populations and also between sexes; however, the profile between males and females does not differ at early ages (5 days old), indicating that CHC profile changing is a dynamic process similar to the one observed in Drosophila (Van ı ckov a et al., 2012). ...
The sterile insect technique (SIT), used for the control of many tephritid fly pests, is based on the rearing and release of large numbers of sexually competitive sterile insects into a wild population. In the interest of reducing expenses and increasing SIT effectiveness, genetic sexing strains (GSS) have been developed. These strains allow the production and release of only males. The objective of our study was to assess the effects of pre‐release adult exposure to methoprene and to females on the mating propensity and mating competitiveness of GSS sterile males of Anastrepha ludens (Loew) (Diptera: Tephritidae). GSS sterile males were kept on a protein‐sugar (protein‐fed) or a protein‐sugar‐methoprene diet and were exposed to different proportions of females for the normal pre‐release period of 5 days. Using laboratory and field‐cage bioassays, we examined the influence of methoprene and female presence on the mating success of sterile males of 3‐9 days old, in competition for wild females with untreated males and with wild males. Methoprene and female exposure had no significant effects on male mating success in the laboratory, whereas age had a positive relationship with the number of copulations observed. However, in field‐cage bioassays, males exposed to females obtained a higher number of copulations than unexposed control males. Possible implications of these findings for programs that use GSS and especially for the campaign against Mexican fruit flies are discussed.
... invadens') as B. dorsalis. The latter taxonomic revision was supported by multidisciplinary evidence from morphological, morphometric, molecular/genetic, cytogenetic, behavioral/sexual compatibility and chemo-ecological studies [8][9][10][11][12][13][14][15][16][17][18][19][20][21]. However, the synonymization by Schutze et al. [8] was criticized [22] and debated [23]. ...
... The species status of B. carambolae in relation to B. dorsalis has also been explored by many researchers using multiple approaches. Data on morphology/morphometrics [1,24], certain genetic markers [12,14,21,[25][26][27][28][29], mating behavior [11] and chemoecology [20,[30][31][32] supported the identity of B. carambolae as a separate biological species and provided some diagnostic features for species discrimination. On the other hand, identification of morphological hybrids [33] and data from nuclear protein coding genes [14] and microsatellite analysis [34] suggest naturally occurring hybridization and gene flow between the two taxa. ...
... invadens'). Their status has been verified by taxonomists and the insect materials have been used in several research projects [11,[17][18][19][20]69]. ...
Bactrocera carambolae is one of the approximately 100 sibling species of the Bactrocera dorsalis complex and considered to be very closely related to B. dorsalis. Due to their high morphological similarity and overlapping distribution, as well as to their economic impact and quarantine status, the development of reliable markers for species delimitation between the two taxa is of great importance. Here we present the complete mitochondrial genome of B. carambolae sourced from its native range in Malaysia and its invaded territory in Suriname. The mitogenome of B. carambolae presents the typical organization of an insect mitochondrion. Comparisons of the analyzed B. carambolae sequences to all available complete mitochondrial sequences of B. dorsalis revealed several species-specific polymorphic sites. Phylogenetic analysis based on Bactrocera mitogenomes supports that B. carambolae is a differentiated taxon though closely related to B. dorsalis. The present complete mitochondrial sequences of B. carambolae could be used, in the frame of Integrative Taxonomy, for species discrimination and resolution of the phylogenetic relationships within this taxonomically challenging complex, which would facilitate the application of species-specific population suppression strategies, such as the sterile insect technique.
... A heat map was used for visualizing the complex data sets organized as matrices as described in Vaníčková et al. (2017). A heat map performed two actions on a matrix. ...
