The Role of Chemical Cues in Host and Mate Location in the Pear Psylla Cacopsylla bidens (Homoptera: Psyllidae)

Chemical communication was shown to play a role in the pear psylla, Cacopsylla bidens. Electrophysiological (EAG) and behavioral responses were investigated in males and females pear psylla . Males were found to be attracted to females, and especially to those on host plants, but not to males, uninfested host plants, or plants infested with conspecific larvae. On the other hand, females were not attracted to males or females but displayed some attraction to host plants. Furthermore, females showed a preference for uninfested pear versus plants infested with conspecific larvae. The antennae of males gave highest electroantenographic response to volatiles from pears infested with females but not males, while females, responded also toward the volatiles of pear alone. These results indicate that females of C. bidens emit sex pheromones that are attractive to the males and suggest that, host volatiles may play a role in host selection by pear psylla females.     

Males of Hylamorpha elegans burmeister (Coleoptera: Scarabaeidae) are attracted to odors released from conspecific females

The behavioral responses of Hylamorpha elegans L. (Coleoptera: Scarabaeidae, Rutelinae) to the semiochemicals released from conspecific individual adults were studied, with particular attention paid to female attraction of males. Odors released from virgin females significantly attracted male conspecifics in both the field and laboratory olfactometer and wind tunnel bioassays. However, females did not attract other females, and males attracted no one. The response of male H. elegans to (1) compounds (1,4-hydroquinone and 1,4-benzoquinone) released only by unmated females; (2) the essential oil of the secondary host (Nothofagus obliqua); and (3) the blend of 1,4-hydroquinone and 1,4-benzoquinone with N. obliqua essential oil was studied. The blend of 1,4-benzoquinone mixed with essential oil at the trial concentration was attractive with males. The same response was found with 1,4-hydroquinone alone. The essential oil did not have the expected attractant effect on conspecific males. These results suggest that, when combined with essential oil, 1,4-benzoquinone may function in the sexual behavior of males and females. These findings are discussed in terms of the ecological role of this putative sexual pheromone and its potential use in a strategy of control of this pest.     

Sex‐specific responses of Asian citrus psyllid to volatiles of conspecific and host‐plant origin

Diaphorina citri Kuwayama (Hemiptera: Psyllidae) is the primary vector of Candidatus Liberibacter spp. bacteria that cause citrus greening, a disease of worldwide importance. Olfactometry was employed to test responses of D. citri to odours from intact citrus plants (Mexican lime, Citrus aurantifolia, sour orange, Citrus aurantium, Marsh grapefruit, Citrus paradisi and Valencia orange, Citrus sinensis), citrus plants previously infested with D. citri, and odours of conspecifics including nymphs, adult insects of same and opposite sex, and their products (honeydew), both alone and in combination. In contrast to other studies, psyllids of both sexes were attracted to volatiles of undamaged Mexican lime leaves, whereas undamaged grapefruit attracted only females, and leaves of Valencia and sour orange did not attract either sex. All four plant species attracted female psyllids when previously infested, but only Mexican lime and sour orange‐attracted males. Thus, Citrus species appear to vary in the production of both constituitive and induced volatiles that attract adult psyllids. Volatiles emitted by nymphs did not attract either sex, but psyllid honeydew was attractive to males, likely due to female pheromone residues. Males oriented to the odour of females, whereas the reverse was not true, and neither males nor females oriented to same‐sex volatiles. The addition of conspecific cues (adults, nymphs or honeydew) did not increase female attraction to previously infested leaves, but male response was increased by the presence of adults and honeydew, regardless of plant species. Thus, female psyllids appear to orient more strongly to volatiles of plant origin, whereas males respond more strongly to cues emanating from females and conspecific excretions. These results suggest that female psyllids drive the initial colonization of host plants, whereas males orient to females and infested plants. Identification of the specific volatiles involved may permit their use in monitoring and management of this pest.     

Sex attraction in Bactericera cockerelli (Hemiptera: Triozidae)

The potato psyllid, Bactericera (=Paratrioza) cockerelli (?ulc) (Hemiptera: Triozidae), is a major pest of potato. We examined the role of chemical signals in sex attraction, assessing male and female response to male- and female-produced volatile chemicals. In laboratory olfactometer assays, potato psyllid males were attracted to odorants emitted from live females and from solvent extract of females. These results indicate that the female-produced chemicals responsible for attracting males may be isolated by means of insect extractions. Males were also attracted to volatile chemicals from males and extracts of males, providing the first example of male-male attraction in the Psylloidea. Males exposed simultaneously to odorants from conspecific females and males were preferentially attracted to female odorants, suggesting the presence of a female-specific sex attractant for males. Potato psyllid females avoided volatile chemicals emitted by females and extracts of females and by volatile chemicals emitted by males and extracts of males. Possible explanations for avoidance of conspecifics by females are discussed. This study is the first report of male attraction to volatile chemicals emitted by females and female extracts in the Triozidae and more specifically in the potato psyllid.     

Role of the plant–conspecific complex in host location and intra-specific communication of Lygus rugulipennis

Abstract.  Wind tunnel and vertical open Y-shaped olfactometer studies are used to test whether volatile cues from the host plant ( Vicia faba ), from conspecific bugs, and from a plant–conspecifics combination, would elicit behavioural responses in mated males and females of Lygus rugulipennis . In the olfactometer, females move towards volatiles from healthy plants but they do not respond to volatiles released by oviposition- and/or feeding-damaged plants without conspecifics, nor to conspecifics alone. Both in the wind tunnel and olfactometer, females respond to volatiles emitted by the plant–insect complex. By contrast, in the wind tunnel, both sexes move significantly towards damaged host plants, even if the presence of conspecifics on these plants enhances only the female response. However, the presence of eggs from conspecifics on host plants reduces the responses of both sexes in the wind tunnel. Finally, males, as well as females, are less responsive to conspecifics alone compared with damaged plants, especially when conspecifics are present on the host plants (host plant– Lygus complex). The results suggest that volatiles emitted by plants and conspecifics influence L.   rugulipennis behaviour, giving information to both sexes on the presence of suitable host plants that have been colonized by other conspecifics acting as pioneers, or providing information on the presence of an already exploited host plant (presence of eggs), thus preventing competition. Males also can use this information to increase the probability of encountering mature females.     

Female Attraction to Conspecific Chemical Cues in the Palmate Newt Triturus helveticus

Although chemosignals are largely used in sexual communication in urodeles, olfactometer studies in newts provided contrasting results about the sex specificity of female behavioural responses. Because long‐range sexual advertisement is believed to be costly, some species might restrain this activity to close interactions with conspecifics. We tested chemical‐mediated sexual attraction in female palmate newt (Triturus helveticus) by measuring the attraction to male and female odours in a linear water olfactometer. Unexpectedly, females were attracted towards conspecifics regardless of sex. They did not show attraction towards Limnaea stagnalis, a common sympatric aquatic gastropod. These results do not support the use of long‐range male sexual signalling in the palmate newt. Instead, conspecific attraction is likely to promote aggregation of males and females in breeding ponds. Observations in the field and in the laboratory tend to support the aggregative behaviour of this species. We discuss the possible function of conspecific attraction in this context. Heading towards any conspecific would increase the probability of finding potential mates. Chemical cues do not need to be sex‐specific at that stage so that long‐range sexual advertisement might be unnecessary. This work emphasizes the need for studies investigating the evolutionary relationships between sexual signalling systems and population‐distribution patterns.     

Effects of Carrion-Independent Pheromone Emission by Male Burying Beetles (Silphidae: Necrophorus)

All former publications on the reproductive biology of burying beetles (genus Necrophorus) state that in these species copulations take place on carcasses only, this being the only place for the sexes to meet. Our laboratory investigations have shown that males emit pheromones if no carrion is around. We now present data from a field investigation showing the effects of pheromone emission by males that have not secured a carcass. Such males are successful in attracting conspecific females. But conspecific males and individuals of other species are also attracted by the scent. Possible benefits for attracted individuals of different species and sex are discussed. Pheromone emission and attraction of conspecifics are restricted to a distinct species-specific period of the day in at least two of the common species.     

Influence of host fruit and conspecifics on the release of the sex pheromone By Toxotrypana curvicauda males (Diptera: Tephritidae)

The release of tephritid sex pheromones depends significantly on the age of the male, the social context (presence of conspecifics) and chemical context (host volatiles). In this study, the influence of host fruit and conspecific (males and females) on the emission of the pheromonal compound 2-methyl-6-vinylpyrazine (2,6 mvp) by Toxotrypana curvicauda (Gerstaecker) males was investigated under laboratory conditions. Males were divided into one control group (nonexposed to treatment) and five experimental groups were placed 1 hr before volatile collection with: 1) host fruit (unripe papaya), 2) two females, 3) two males, 4) host fruit plus two females, and 5) host fruit plus two males. The volatiles were sampled by means of solid phase microextraction and analyzed with gas chromatography-mass spectrometry, with a focus on 2,6 mvp. Males emitted volatiles from the first day after adult emergence. The maximum peak and the temporal pattern of 2,6 mvp release were modified by the presence of host fruit and conspecific males or females. On day 5, males in the presence of fruit maintained a constant release of 2,6 mvp while the presence of conspecific caused a decrease of pheromone release. The release of 2,6 mvp was increased significantly in males exposed to both types of stimuli simultaneously. The stimuli (fruit and conspecifics) modified the release of 2,6 mvp, however the effect depended on male age.     

Prey Habitat Location by the Cassava Mealybug Predator Exochomus flaviventris: Olfactory Responses to Odor of Plant, Mealybug, Plant–Mealybug Complex, and Plant–Mealybug–Natural Enemy Complex

Exochomus flaviventris Mader is considered to be the most active predator of the cassava mealybug Phenacoccus manihoti Matile–Ferrero in Central Africa. The response of experienced gravid female coccinellids to the odor of cassava plant (var. Zanaga), unparasitized mealybugs, plant–mealybug complex with or without feeding prey (parasitized or not), and plant–mealybug complex with or without conspecific coccinellids was investigated in a Y-tube olfactometer. The odor of uninfested cassava plants was not more attractive than clean air. Dual-choice tests revealed that mealybug-infested plants were preferred to mealybugs alone and mealybug-damaged plants and were the major sources of volatiles that attract females coccinellids to the microhabitat of its prey. The emission of volatile chemicals did not appear to be limited to the infested parts of the plant but did occur systemically throughout the plant. The presence of conspecific coccinellid larvae or adult males did not modify the attractiveness of the mealybug-infested plants. However, when an infested plant with conspecific predator females (alone or with conspecific males) was compared to an infested plant or infested plant with conspecific males, E. flaviventris females showed a preference for the last two sources of odor. The uninfested plant with conspecific males was also preferred to the uninfested plant with conspecific females. In addition, the odor of conspecific males was preferred over that of conspecific females. Female predators preferred the plant infested with unparasitized mealybugs over the plant infested with mealybugs previously parasitized. These results showed that E. flaviventris females use herbivore-induced plant volatiles during foraging and can detect via olfaction the presence of conspecific gravid females and parasitized prey, thus assessing patch suitability from a distance.     

Attraction of male winterform pear psylla to female-produced volatiles and to female extracts and evidence of male–male repellency

Pear psylla, Cacopsylla pyricola (Förster) (Homoptera: Psyllidae), is a major pest of commercial pears in North America and Europe. Olfactometer trials have shown that males of both the summer and winter morphotype are attracted to female-infested host material. Additional work with the summer morphotype has shown that males are attracted to females even in the absence of the host plant, which is evidence that female C. pyricola produce a volatile sex attractant. Here, we describe similar results with the winterform, confirming for this morphotype that the female psylla rather than the infested host material is the source of the attractant. Male winterforms displayed attraction to odors from live females in the absence of the host plant, freshly killed females, and female whole body extracts. The female whole body extracts were at least as attractive as a comparable number of live females, suggesting that we were successful at extracting the components of the attractant with this procedure. All previous olfactometer trials with C. pyricola used the insect as the attractant source; the current study is the first to demonstrate that volatile chemicals isolated from the female insect were attractive to male conspecifics. Winterform males were also assayed to odors produced by conspecific males. We found that male psylla avoided volatile odors from live males, freshly killed males, or whole body extracts of males. To our knowledge, these results are the first indication that males of any member of the Psyllidae avoid odors associated with conspecific males.     

Attraction of pepper weevil to volatiles from damaged pepper plants

Pioneer herbivorous insects may find their host plants through a combination of visual and constitutive host‐plant volatile cues, but once a site has been colonized, feeding damage changes the quantity and quality of plant volatiles released, potentially altering the behavior of conspecifics who detect them. Previous work on the pepper weevil, Anthonomus eugenii Cano (Coleoptera: Curculionidae), demonstrated that this insect can detect and orient to constitutive host plant volatiles released from pepper [Capsicum annuum L. (Solanaceae)]. Here we investigated the response of the weevil to whole plants and headspace collections of plants damaged by conspecifics. Mated weevils preferred damaged flowering as well as damaged fruiting plants over undamaged plants in a Y‐tube olfactometer. They also preferred volatiles from flowering and fruiting plants with actively feeding weevils over plants with old feeding damage. Both sexes preferred volatiles from fruiting plants with actively feeding weevils over flowering plants with actively feeding weevils. Females preferred plants with 48 h of prior feeding damage over plants subjected to weevil feeding for only 1 h, whereas males showed no preference. When attraction to male‐ and female‐inflicted feeding damage was compared in the Y‐tube, males and females showed no significant preference. Wind tunnel plant assays and four‐choice olfactometer assays using headspace volatiles confirmed the attraction of weevils to active feeding damage on fruiting plants. In a final four‐choice olfactometer assay using headspace collections, we tested the attraction of mated males and virgin and mated females to male and female feeding damage. In these headspace volatile assays, mated females again showed no preference for male feeding; however, virgin females and males preferred the headspace volatiles of plants fed on by males, which contained the male aggregation pheromone in addition to plant volatiles. The potential for using plant volatile lures to improve pepper weevil monitoring and management is discussed.     

Mate finding in the forest cockchafer, Melolontha hippocastani, mediated by volatiles from plants and females

The response of the forest cockchafer, Melolontha hippocastani F. (Coleoptera, Scarabaeidae), towards volatiles emitted by different host plants and conspecifics was tested in field experiments during the flight period at dusk. Funnel traps containing artificially damaged leaves from the host plants Carpinus betulus L. and Quercus rubra L., as well as from the non‐host plant Prunus serotina Ehrh. caught significantly more beetles than empty control traps. On the other hand, traps baited with undamaged leaves from Q. rubra did not catch significantly more beetles than empty controls. Leaves from C. betulus damaged by beetle feeding did not attract more beetles than artificially damaged leaves. By use of gas chromatography coupled with electroantennographic detection (GC‐EAD) electrophysiological responses of males and females were shown for 18 typical plant volatiles. A synthetic mixture of selected typical green plant volatiles was also highly attractive in the field. A total of 9982 beetles was caught during the field experiments, among them only 33 females. This suggests that attraction to damaged foliage during flight period at dusk is male‐specific. Field experiments testing the attractiveness of female M. hippocastani towards conspecific males by employing caged beetles and beetle extracts indicated that males of M. hippocastani use a female‐derived sex pheromone for mate location. On wired cages containing either unmated feeding females, or unmated females without access to foliage, or feeding males in combination with extracts from unmated females, significantly more males landed during the flight period than on comparable control cages containing feeding males or male extracts. A possible scenario of mate location in M. hippocastani involving feeding‐induced plant volatiles and a female‐derived sex pheromone is discussed.     

The role of biotic and abiotic cues in stimulating aggregation by larval cane toads (Rhinella marina)

Tadpoles of the cane toad (Rhinella marina) form dense aggregations in the field, but the proximate cues eliciting this behavior are not well understood. We sampled water‐bodies in the Northern Territory of Australia, finding that the density of cane toad tadpoles increased with increasing temperature. Furthermore, we conducted laboratory experiments to explore the roles of biotic factors (attraction to conspecifics; chemical cues from an injured conspecific; food) and spatially heterogeneous abiotic factors (light levels, water depth, physical structure) to identify the cues that induce tadpole aggregation. Annulus and binary choice trials demonstrated weak but significant attraction between conspecifics. Tadpoles decreased swimming speeds, but did not increase grouping in response to cues from an injured conspecific. Larvae aggregated in response to abiotic cues (high levels of illumination and proximity to physical structures) and were strongly attracted to feeding conspecifics. Overall, aggregation by cane toad tadpoles is likely driven by weak social attraction coupled with a shared preference for specific abiotic features, creating loose aggregations that are then reinforced by movement toward feeding conspecifics.     

Aggregation pheromone in Sternechus subsignatus (Coleoptera: Curculionidae): olfactory behaviour and temporal pattern of emission

The Brazilian soybean stalk weevil, Sternechus subsignatus Boehman (Coleoptera: Curculionidae), is a pest of economic importance in many regions of Brazil. Volatiles from both sexes of S. subsignatus were collected by aeration and the behavioural response of males and females was evaluated using a Y‐olfactometer. The results obtained demonstrate that the communication in S. subsignatus is mediated by aggregation pheromone as both sexes were attracted to host plant (HP) volatiles, and this attraction was increased by the addition of male volatiles. At least five male‐specific compounds (1–5) were detected in the chromatographic analysis, providing chemical support to the behavioural data. Release of these volatiles is dependent on the presence of the HP as the amount of compounds differs significantly when volatiles are collected from weevils with or without access to food. The release takes place mainly during photophase, showing a peak between 4 and 6 h after its beginning, which is also a peak of the insect activity in the field. Studies are underway to elucidate the structures of these putative aggregation pheromone components, in order to evaluate their biological activity under laboratory and field conditions.     

An omnivorous arthropod, <Emphasis Type="Italic">Nesidiocoris tenuis</Emphasis>, induces gender-specific plant volatiles to which conspecific males and females respond differently

Nesidiocoris tenuis Reuter (Heteroptera: Miridae) is an omnivorous mirid bug that preys on diverse generalist herbivorous arthropods. N. tenuis adults are attracted to volatiles from plants induced by their prey, such as tobacco cutworms (CCW) (Spodoptera litura larvae) and two-spotted spider mites (Tetranychus urticae). N. tenuis adults also induce volatiles when they infest plants. In this study, we focused on olfactory responses of N. tenuis males and females to volatiles from eggplants and sesame plants induced by conspecifics of the same or different gender by using a Y-tube olfactometer. Males were attracted to volatiles from plants of both species induced by either males or females. The male preference was biased to volatiles from plants of both species induced by females, probably because the biased response would facilitate their mate-finding. Females were attracted only to volatiles from plants of both species induced by females. Mating occurs multiple times in this species. Thus, the responses would indirectly affect mating of males and females. Slight but significant qualitative and quantitative differences were detected between the volatiles of plants of both species induced by N. tenuis females and those of the plants induced by conspecific males. N. tenuis might use such differences in their gender-specific responses.     

Concurrent modulation of neuronal and behavioural olfactory responses to sex and host plant cues in a male moth

Mating has profound effects on animal physiology and behaviour, not only in females but also in males, which we show here for olfactory responses. In cotton leafworm moths, Spodoptera littoralis, odour-mediated attraction to sex pheromone and plant volatiles are modulated after mating, producing a behavioural response that matches the physiological condition of the male insect. Unmated males are attracted by upwind flight to sex pheromone released by calling females, as well as to volatiles of lilac flowers and green leaves of the host plant cotton, signalling adult food and mating sites, respectively. Mating temporarily abolishes male attraction to females and host plant odour, but does not diminish attraction to flowers. This behavioural modulation is correlated with a response modulation in the olfactory system, as shown by electro-physiological recordings from antennae and by functional imaging of the antennal lobe, using natural odours and synthetic compounds. An effect of mating on the olfactory responses to pheromone and cotton plant volatiles but not to lilac flowers indicates the presence of functionally independent neural circuits within the olfactory system. Our results indicate that these circuits interconnect and weigh perception of social and habitat odour signals to generate appropriate behavioural responses according to mating state.     

Alternative models of conspecific attraction in flies and crabs

Animals are often attracted to one another when selecting habitats,but little is known about the rules governing conspecific attraction.We use Akaike Information Criterion to evaluate alternativemodels of the effects of conspecifics on individual choice inthe context of habitat selection. One set of models was testedusing data collected on virgin female flies, Drosophila melanogaster,selecting habitats in the laboratory; a second set of modelswas tested using data collected on crabs, Ocypode rotundata,selecting foraging patches in the field. Patterns of space usein the flies were most consistent with models indicating thatindividuals were attracted to other females that selected trapsduring the same hour, as well as to cues left by females thathad entered traps during the previous hour. Results for thecrabs were most consistent with a model which assumes that individualspreferred to join the patch with the most crabs but that theirability to assess the number of crabs in alternative patcheswas constrained by basic psychophysical principles (Weber'slaw). These results provide support for hypotheses about thefunctional significance of conspecific attraction in the fliesand the crabs and illustrate the richness of information aboutconspecific attraction that can be obtained when the same dataset is confronted with alternative models of the ways that animalsrespond to one another when selecting habitats.     

Attraction of clover head weevil, Hypera meles, to flower bud volatiles of several species of Trifolium

Steam distilled volatiles from flower buds of white clover, Trifolium repens, attracted significantly greater numbers of female clover head weevils, Hypera meles, than males, in laboratory bioassays. Persian clover, T. resupinatum, bud extract, which attracted significantly more females than either small hop clover, T. dubium or red clover, T. pratense, also displayed qualitative and quantitative differences in the gas chromatographic profile of its bud volatiles. The strong attraction of normal females to Persian clover in laboratory bioassays was abolished by antennectomy, suggesting that attraction is regulated by olfactory receptors located on weevil antennal clubs. Weevils demonstrated a similar attraction to Persian clover in the field, indicating the possibility of developing a lure made from the flower bud extract of this clover to trap weevils for control or population monitoring purposes.     

Intended and unintended receivers of the male pheromones of the burying beetles Nicrophorus humator and Nicrophorus vespilloides

Male Nicrophorus beetles (Coleoptera: Silphidae) attract females through volatiles that are emitted at species‐specific times of day. Not only beetles of the opposite sex but also conspecific males are attracted. Another observation is the co‐attraction of congeners, a phenomenon that was shown in particular for Nicrophorus vespilloides Herbst, the smallest Nicrophorus species in Central Europe. In the current study, we identified the Nicrophorus humator Gleditsch male pheromone as methyl 4‐methyloctanoate through gas chromatography/mass spectrometry analysis. In field experiments, we tested and compared the attractiveness of synthetic analogs of the male pheromones of N. humator and N. vespilloides in baited pitfall traps. An asymmetric cross‐attraction to the synthetic male pheromones was observed, which is best explained by the skewed competitive relationship of the two species, with regard to the restricted availability of breeding resources. Nicrophorus humator is attracted by both its own male pheromone and by the pheromone of the smaller N. vespilloides, whereas N. vespilloides is almost exclusively attracted by its own male pheromone. The observed attraction of conspecific males of either species to male pheromone baits can be explained by both competition for females and competition for breeding resources.     

Response of the pollen beetle Meligethes aeneus to volatiles emitted by intact plants and conspecifics

The response of the pollen beetle Meligethes aeneus Fab. (Coleoptera, Nitidulidae) to the volatiles of undamaged plants and conspecifics was tested in a Y-tube-olfactometer-bioassay. Beetles that had hibernated preferred significantly the volatiles emitted by their most important host plant, oilseed-rape (Brassica napus L. ‘Lorar’) in the early bud-stage. However, the odour emitted by rye (Secale cereale L.), tomato plants (Solanum lycopersicum L.), and yarrow (Achillea millefolium L.) were also attractive when tested against the corresponding growing-medium. Dock plants (Rumex obtusifolius L.) and touch-me-not (Impatiens parviflora L.) possessed no attractive properties. When tested against each other, oilseed-rape was preferred significantly by M. aeneus above all other plants, with the exception of tomato. The results indicate that M. aeneus is able to locate its host plant by olfactory stimuli in the early bud-stage, i.e. in that stage in which the infestation begins in the field and when the typical yellow colour and floral scent of oilseed-rape are absent. Female pollen beetles avoided significantly the volatiles emitted by female conspecifics and an ether extract of conspecifics of mixed sex, while volatiles from males had no significant effect on females. Furthermore, males showed no preferences when responding to conspecific odour in the olfactometer. These results suggest the existence of an epideictic pheromone for M. aeneus.