Orientation of flying male Anobium punctatum (Coleoptera: Anobiidae) to sex pheromone: separating effects of visual stimuli and physical barriers to wind

Abstract. Flying male woodworm, Anobium punctatum , are able to locate a source of female pheromone offered as a point source but land more quickly if the same odour source is presented on a model 'tree' (cylinder). We tested the response to a two-dimensional black 'tree' marked as a silhouette on the upwind screen of the wind tunnel, with or without a hidden upwind barrier to the wind. The beetles responded equally to pheromone presented on a visual silhouette with wind barrier and to the hidden upwind wind-barrier alone, suggesting that the wind-barrier itself has a significant effect on insect orientation and landing: visual stimuli do not add to the landing response if the barrier is present. However, without the barrier upwind, landing was faster when the visual stimulus was present than without. This is the third forest-living species to be shown to respond in this way, which suggests this phenomenon may be more widespread. The possibility that this phenomenon is present in other, non-forest, phytophagous insects, their predators and parasitoids is discussed.     

Positive and negative impacts of insect frass quality on soil nitrogen availability and plant growth

Frass deposition to soil is an important pathway by which herbivorous insects impact decomposition and soil nutrient availability. However, little is known about how frass quality influences ecosystem properties. Here, we examined the effects of frass quality on the decomposition process, soil nitrogen (N) availability, and plant growth, using frass of Mamestra brassicae (L.) that fed on fertilized or unfertilized Brassica rapa L. var. perviridis Bailey. The frass quality was largely dependent on the host plant quality. Frass excreted by larvae that fed on the fertilized plants had higher N than that of larvae that fed on the unfertilized plants. The decomposition rate of the frass did not differ between N-rich and N-poor frass, except during the early decomposition period. The inorganic N concentration decreased during decomposition in both frass types. However, difference in the initial inorganic N concentration led to different consequences regarding soil N availability. Furthermore, addition of frass to the soil differently influenced the growth of B. rapa plants depending on the frass quality: plant biomass was increased by N-rich frass addition but decreased by N-poor frass addition, compared to the biomass without frass addition. These results indicate that frass quality is an important factor in determining the impact of herbivorous insects on nutrient dynamics, and that frass positively or negatively influences soil N availability and plant growth, depending on its quality.     

Kairomones from scarabaeid grubs and their frass as cues in below-ground host location by the parasitoids Tiphia vernalis and Tiphia pygidialis

Cues used in below-ground host-searching behaviour and host discrimination were examined for Tiphia vernalis Rohwer and Tiphia pygidialis Allen (Hymenoptera: Tiphiidae), ecto-parasitoids of root-feeding larvae of the Japanese beetle, Popillia japonica Newman, and masked chafers, Cyclocephala spp. (Coleoptera: Scarabaeidae), respectively. Response to potential stimuli was compared in dual choice tests in an observation chamber filled with soil. Each wasp showed species-specific, directed movement along residual body odor trails made by dragging its respective host through the soil. Presence of a grub was not necessary for wasps to follow such trails. Frass from either host- or non-host grubs elicited trail-following, but each Tiphia species followed frass trails from its respective host when a choice was presented. Frass trails elicited stronger responses than body odor trails. The combination of host frass and body odor elicited the strongest trail-following responses. Our results suggest that once in the soil, Tiphia spp. locate their hosts using contact kairomones present in grub body odor trails and frass.     

Parasitoids and dipteran predators exploit volatiles from microbial symbionts to locate bark beetles

Host location by parasitoids and dipteran predators of bark beetles is poorly understood. Unlike coleopteran predators that locate prey by orienting to prey pheromones, wasps and flies often attack life stages not present until after pheromone production ceases. Bark beetles have important microbial symbionts, which could provide sources of cues. We tested host trees, trees colonized by beetles and symbionts, and trees colonized by symbionts alone for attractiveness to hymenopteran parasitoids and dipteran predators. Field studies were conducted with Ips pini in Montana. Three pteromalid wasps were predominant. All were associated with the second and third instars of I. pini. Heydenia unica was more attracted to logs colonized by either I. pini or the fungus Ophiostoma ips than logs alone or blank controls (screen with no log). Rhopalicus pulchripennis was more attracted to logs colonized by I. pini than logs alone or blank controls. Dibrachys cavus was attracted to logs but did not distinguish whether or not they were colonized. Two dolichopodid predators were predominant. A Medetera species was more attracted to colonized than uncolonized logs and more attracted to logs than blank controls. It was also more attracted to logs colonized with the yeast Pichia scolyti than uncolonized logs, but attraction was less consistent. An unidentified dolichopodid was more attracted to logs colonized with I. pini, O. ips, and the bacteria Burkholderia sp., than to uncolonized logs. It was also attracted to uncolonized logs. Its responses were less consistent and pronounced than H. unica. These results suggest some parasitoids and dipteran predators exploit microbial symbionts of bark beetles to locate hosts. Overall, specialists showed strong attraction to fungal cues, whereas generalists were more attracted by plant volatiles. These results also show how microbial symbionts can have conflicting effects on host fitness.     

Sequence matters – selective adaptation in electroantennographic response to binary odour mixtures by the Colorado potato beetle

Chemically mediated behaviour of insects is often strongly affected by mixtures of odour stimuli and their temporal characteristics. Both sensory transduction and central processing of odour mixtures can give rise to several different kinds of interaction, which can influence how individual components are perceived and processed. In particular, odour mixtures have been examined in model experiments as premixed binary mixtures in comparison with pure odour stimuli. Only in few experiments, the influence of the temporal structure of odour mixtures on odour perception has been taken into account. Natural odour stimuli often have a pulsed structure and may in general be superimposed on a background of irrelevant or interfering compounds, which can fluctuate with different frequencies, depending on their source. To achieve a better representation of these natural conditions, our odour mixing experiments apply a new kind of stimulation protocol: odours were not premixed but superimposed with a specific time pattern; one odour stimulus was presented as a longer persisting background and the second stimulus was a superimposed short test signal. To gain an overview of odour interaction patterns in the Colorado potato beetle by causing adaptation of one receptor population at naturally occurring levels of concentration and time intervals, electroantennographic recordings were made on excised antennae. A matrix of 12 stimulus compounds led to 132 pairs of compounds tested, each in the role of background and test stimulus. In 64 cases, the interaction was significantly different, when the role of background and stimulus was exchanged. Interaction patterns ranging from no interference (independence) to suppression were found and assigned to four clearly distinguishable types. We discuss that the observed effects of the presentation sequence in odour mixtures may contribute to the mechanisms of olfactory pattern recognition and olfactory contrast perception by insects.     

The role of frass and cocoon volatiles in host location by Monodontomerus aeneus, a parasitoid of Megachilid solitary bees

Monodontomerus aeneus (Fonscolombe) is a parasitic wasp that oviposits on the prepupae and pupae of Osmia cornuta (Latreille) and other solitary bee species. A two-armed olfactometer was used to test the olfactory attractiveness of O. cornuta prepupae, cocoon, and larval frass to female M. aeneus. Both cocoon and frass attracted the female parasitoids, but frass alone was more attractive than the cocoon and the cocoon with frass was more attractive than frass alone. Female parasitoids were not attracted by the host prepupa. M33 (methanol) was the organic volatile most emitted by cocoons and m61 (acetic acid) was the compound most emitted by frass. However, cocoons showed higher emission for almost all compounds, including m61 (acetic acid). Although acetic acid alone attracted M. aeneus, a complex volatile signal is probably involved in the attraction process because the ratio of acetic acid and acetaldehyde characteristic of the frass was more attractive than other ratios.     

Visual and olfactory location of biotopes, prey patches, and individual prey by the ladybeetleChilocorus nigritus

Foraging behaviour of the predatorChilocorus nigritus (Fabricius) (Coleoptera: Coccinellidae) at the three spatial levels of biotope, prey patch and individual prey, was studied in the laboratory, and related to behaviour in the field. Vertically oriented parallel lines were more attractive than the same shapes in a horizontal position. A simulated horizon with a tree line was preferred to a simulated flat horizon. They were attracted to a tree image for the first 2 h of exposure, but were less attracted after longer exposure, possibly due to habituation. Leaf shape was recognised, and simple ovate leaves were preferred to compound bipinnate leaves and to squares. These responses were associated with biotope selection for feeding and aggregation at aestivation sites. The location of prey patches by adults involved prey odour but the location of such sites by larvae did not. Adults detected individual prey visually and olfactorily over short distances but physical contact with prey was required for detection by larvae. Location of individual prey and prey patches by adults and larvae was facilitated by alternation between intensive and extensive search. The differences in the ability of larvae and adults to locate prey, stem from the adults being the active locators of biotope and patch, whereas the comparatively immobile larvae depend on their parents’ ability for long-range location of prey. Two hypotheses concerning coccinellid foraging behaviour are proposed. Firstly, the duration of response to a visual cue is related to the distance over which such a cue may be perceived. It follows that habituation to closer range cues occurs more rapidly than to longer range cues. Secondly, visual cues used by adults at the different spatial levels of prey location, and the location of mates and aggregation sites, have the same or similar shape. These results also provide guidelines for orchard management to maximise the biocontrol value of this species.     

Divergent responses of two cereal aphids to previous infestation of their host plant

The role of pollen odour in resource location by the pollen beetle, Meligethes aeneus (Fabricius) (Coleoptera: Nitidulidae), a pollen-feeding insect regarded as a pest of oilseed rape, Brassica napus L., (Brassicaceae) crops, was investigated in a linear track olfactometer. Both male and female beetles were attracted to the odour of whole oilseed rape flowers, indicating that these insects can locate their host plants using floral odours as cues. The attractive odour of flowers was found to emanate from all floral parts tested: the petals/sepals, the anthers, and from pollen itself. Therefore, at least part of the attractive odour of oilseed rape flowers emanates from pollen. Beetles were more attracted to floral samples containing anthers than those without anthers when these odours were directly compared in a choice-test, and this indicates that there were detectable differences between them. Anthers and pollen may therefore release distinctive odours that are quantitatively and/or qualitatively different from the odour of the rest of the flower. These experiments support the hypothesis that pollen-seeking insects use pollen odour cues to locate this food source.     

Learning of Multi-Modal Stimuli in Hawkmoths

The hawkmoth, Manduca sexta, uses both colour and odour to find flowers when foraging for nectar. In the present study we investigated how vision and olfaction interact during learning. Manduca sexta were equally attracted to a scented blue coloured feeding target (multimodal stimulus) as to one that does not carry any scent (unimodal stimulus; visual) or to an invisible scented target (unimodal stimulus; odour). This naive attraction to multimodal as well as to unimodal stimuli could be manipulated through training. Moths trained to feed from a blue, scented multimodal feeding target will, when tested in a set-up containing all three feeding targets, select the multimodal target as well as the scented, unimodal target, but ignore the visual target. Interestingly, moths trained to feed from a blue, unimodal visual feeding target will select the visual target as well as the scented, multimodal target, but ignore the unimodal odour target. Our results indicate that a multimodal target is perceived as two separate modalities, colour and odour, rather than as a unique fused target. These findings differ from earlier studies of desert ants that perceive combined visual and odour signals as a unique fused stimulus following learning trials.     

Attraction of the predator,Harmonia axyridis (Coleoptera: Coccinellidae), to its prey,Myzus persicae (Hemiptera: Aphididae), feeding on Chinese cabbage

The predatory multicolored Asian lady beetle, Harmonia axyridis, was attracted to volatiles released from Chinese cabbage infested by the green peach aphid, Myzus persicae, in T-tube olfactometer choice tests. However, lady beetle adults and larvae did not respond to clean air, Chinese cabbage alone or green peach aphid alone. Of different prey densities, H. axyridis adults were most attracted to Chinese cabbage infested by 60 M. persicae adults after 24 h. However, H. axyridis larvae were not attracted to Chinese cabbage infested by M. persicae. Mechanically damaged Chinese cabbage attracted neither lady beetle adults nor larvae. Predatory adults were attracted to 60 M. persicae adults after 24 and 48 h, and to 90 M. persicae adults after 12 h, suggesting that the predatory response depends on the prey density. Lady adult beetles did not prefer the volatiles induced by Diamondback moth, Plutella xylostella, indicating that specific host insect specificity attracts respective natural enemies. It can be explained that the volatile compounds emitted from the host plant as a result of herbivore attack preferred by the specific insect species.     

Modelling mating success of saproxylic beetles in relation to search behaviour, population density and substrate abundance

We compared the efficiency of two mate-finding strategies exploited by representatives of the beetle families Cisidae and Anobiidae (genus Dorcatoma) that live inside fruiting bodies of wood-decaying fungi. In the Cisidae both sexes are attracted to host odour, but no pheromones seem to be present (nonpheromone strategy). In the Dorcatoma species only the females are attracted to host odour, but having found a host they attract males with a sexual pheromone (pheromone strategy). With a simulation model, we compared the efficiency of the two strategies at four densities of trees hosting fungal fruiting bodies and at three relative densities of insects. We found only small differences in efficiency between the two strategies at high relative densities of conspecific individuals, regardless of host tree density. The pheromone strategy was relatively more efficient when the relative density of insects or the density of host trees decreased. Thus, species adopting the nonpheromone strategy are probably more sensitive to habitat fragmentation and more likely to decline and go extinct at low population densities (because of Allee effects) than species using the pheromone strategy. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

The culturable bacterial community of frass produced by larvae of Rhynchophorus ferrugineus Olivier (Coleoptera: Curculionidae) in the Canary island date palm

Aims: Larvae of the red palm weevil (RPW) Rhynchophorus ferrugineus Olivier (Coleoptera: Curculionidae) feed inside palm stem tissues, making galleries and producing a wet fermenting frass. We characterized the culturable bacteria associated with frass produced by tunnelling larvae inside the Canary island date palms and investigated the role of frass and gut bacteria in plant polymers breakdown. Methods and Results: A culture‐dependent method was used to isolate bacteria from frass and noninfested palm tissues. Bacterial isolates were grouped into operational taxonomic units based on polymorphisms in the ITS‐PCR profiles, and representative isolates were identified by partial sequencing of the 16S rRNA gene. Frass bacteria were dominated by 2,3‐butanediol fermenter Enterobacteriaceae. None of the bacterial isolates was able to degrade cellulose; however, cellulolytic and hemicellulolytic bacteria were isolated from the larval gut enrichment cultures. Conclusions: Frass bacteria are specifically associated with the RPW larvae and might play beneficial roles for RPW, other than nutritional, that deserve further investigations. Breakdown of plant polymers probably occurs inside the larvae digestive system. Significance and Impact of the Study: Frass and gut micro‐organisms of R. ferrugineus should be included in studies of the interactions between RPW, its plant hosts, and its enemies.     

Medetera aldrichii larval feeding behavior and prey consumption [Dipt.: Dolichopodidae]

Predatory behavior ofMedetera aldrichii Wheeler (Dolichopodidae) larvae was observed in bark-Lucite sandwiches infested with immature broods of the bark beetle preyDendroctonus pseudotsugae Hopkins (Scolytidae). Predator larvae could not penetrate unmined phloem or prey galleries tightly packed with frass, but moved through loose frass and bark-Lucite interspaces formed when the mined inner bark dried. At high prey densities interspace formation and gallerly contiguity enabled the predator to move freely and attack several prey in rapid succession. At low prey densities the predator was frequently isolated from prey by impenetrable strips of phloem. Predator larvae maintained singly in small plastic arenas consumed an average of 14.7 immature prey, but required only 6.2 prey when reared exclusively on large prey.     

Zigzagging and casting as a programmed response to wind-borne odour: a review

ABSTRACT. 'Counterturning' (meaning here the execution of a succession of alternating left and right turns) is the common feature in upwind zigzagging and cross-wind casting by flying insects manoeuvring towards a small source of wind-borne odour. Recent progress in understanding its control and function is discussed. Counterturning is internally controlled ('self-steered') in the limited sense that, once initiated by a chemical stimulus, it continues without further changes in the chemical input both in clean air and in a homogeneous cloud of odour. As a reaction it appears to be the kind of chemotaxis distinguished as longitudinal klinotaxis, for which the stimulus is a difference of chemical concentration detected over time along the insect's path, not across it. The new directions taken in response to the stimulus, being self-steered in the above sense, have no necessary relation to the direction of the chemical gradient that provided the stimulus but are influenced by the visual cues generated by wind drift. In wind, the counterturning programme is modulated by changes in the chemical input and simultaneously integrated with anemotaxis, but it can then continue in similar form after the wind has ceased. Unambiguous evidence for these conclusions is so far available only for certain flying male moths responding to sex pheromone. The primary function of counterturning, of all amplitudes and in both zigzagging and casting, appears to be the regaining of contact with an elusive scent.     

The attraction of virgin female hide beetles (Dermestes maculatus) to cadavers by a combination of decomposition odour and male sex pheromones

ABSTRACT: INTRODUCTION: The hide beetle Dermestes maculatus (Coleoptera: Dermestidae) feeds as an adult and larva on decomposing animal remains and can also be found on human corpses. Therefore, forensic entomological questions with regard to when and how the first receptive females appear on carcasses are important, as the developmental stages of their larvae can be used to calculate the post-mortem interval. To date, we know that freshly emerged males respond to the cadaver odour of post-bloated carcasses (approximately 9 days after death at Tmean = 27degrees), being attracted by benzyl butyrate. This component occurs at its highest concentration at this stage of decay. The aim of our study was to determine the principle of attraction of virgin females to the feeding and breeding substrate. For this purpose, we tested the response of these females to headspace samples of piglet cadavers and male sex pheromones [(Z9)-unsaturated fatty acid isopropyl esters] in a Y-olfactometer. Because we expected that such an odour combination is of importance for virgin female attraction, we tested the following two questions: 1) Are virgin female hide beetles attracted by a combination of cadaver odour and male sex pheromones? 2) During which decomposition stage do the first virgin females respond to cadaver odour when combined with male sex pheromones? RESULTS: We found that young virgin females were attracted to the cadaver by a combination of cadaver odour and male sex pheromones. Neither cadaver odour alone nor male sex pheromones alone was significantly more attractive than a solvent control. Our results also gave a weak indication that the first young virgin females respond as early as the post-bloating stage to its associated decomposition odour when combined with male sex pheromones. CONCLUSIONS: Our results indicate that freshly emerged males possibly respond to cadaver odour and visit carcasses before virgin females. Being attracted to cadavers when male sex pheromone is perceived as well, virgin females can optimise their reproductive possibilities.     

Behavioural responses of the vine weevil, Otiorhynchus sulcatus, to semiochemicals from conspecifics, Otiorhynchus salicicola, and host plants

The vine weevil Otiorhynchus sulcatus is a parthenogenetic reproducing species which forages for suitable host plants at night, but is found congregated in dark places during the day. Frass of this weevil species is suspected to contain attractive compounds that are host‐plant related. Using a still‐air olfactometer, we tested adult vine weevils at night for their behavioural response to odours from conspecifics, feeding on a mixture of spindle tree (Euonymus fortunei) and yew (Taxus baccata), and to a sexually reproducing related species (Otiorhynchus salicicola), feeding on a mixture of ivy (Hedera helix) and cherry laurel (Prunus laurocerasus). Their attraction to conspecifics and O. salicicola appeared to be related to frass production. Freshly collected frass from O. sulcatus and from O. salicicola males and females was attractive. Prunus laurocerasus and H. helix have not been observed to be hosts of the vine weevil in the field. However, our tests showed that the vine weevil was attracted to mechanically damaged leaves of both plant species, whereas undamaged leaves were not attractive. Only undamaged young unfolding leaves of H. helix were also attractive. The attraction to odours from mechanically damaged host and non‐host plants suggested the involvement of compounds that are commonly found in many plant species. The involvement of plant compounds and/or aggregation pheromones in attraction to frass of the vine weevil and frass of the related weevil species O. salicicola is discussed.     

Static antennae act as locomotory guides that compensate for visual motion blur in a diurnal,keen-eyed predator

High visual acuity allows parallel processing of distant environmental features, but only when photons are abundant enough. Diurnal tiger beetles (Carabidae: Cicindelinae) have acute vision for insects and visually pursue prey in open, flat habitats. Their fast running speed causes motion blur that degrades visual contrast, forces stop-and-go pursuit and potentially impairs obstacle detection. We demonstrate here that vision is insufficient for obstacle detection during running, and show instead that antennal touch is both necessary and sufficient for obstacle detection. While running, tiger beetle vision appears to be photon-limited in a way reminiscent of animals in low-light habitats. Such animals often acquire wide-field spatial information through mechanosensation mediated by longer, more mobile appendages. We show that a nocturnal tiger beetle species waves its antennae in elliptical patterns typical of poorly sighted insects. While antennae of diurnal species are also used for mechanosensation, they are rigidly held forward with the tips close to the substrate. This enables timely detection of path obstructions followed by an increase in body pitch to avoid collision. Our results demonstrate adaptive mechanosensory augmentation of blurred visual information during fast locomotion, and suggest that future studies may reveal non-visual sensory compensation in other fast-moving animals.     

Masquerading predators deceive prey by aggressively mimicking bird droppings in a crab spider

In aggressive mimicry, a predator accesses prey by mimicking the appearance and/or behavior of a harmless or beneficial model in order to avoid being correctly identified by its prey. The crab spider genus Phrynarachne is often cited as a textbook example of masquerading as bird droppings (BDs) in order to avoid predation. However, Phrynarachne spiders may also aggressively mimic BDs in order to deceive potential prey. To date, there is no experimental evidence to support aggressive mimicry in masquerading crab spiders; therefore, we performed a field survey, a manipulative field experiment, and visual modeling to test this hypothesis using Phrynarachne ceylonica. We compared prey-attraction rates among BDs, spiders, and control empty leaves in the field. We found that although all prey combined and agromyzid dipterans, in particular, were attracted to BDs at a higher rate than to spiders, other dipterans and hymenopterans were attracted to BDs at a similar rate as to spiders. Both spiders and BDs attracted insects at a significantly higher rate than did control leaves. As predicted, prey was attracted to experimentally blackened or whitened spiders significantly less frequently than to unmanipulated spiders. Finally, visual modeling suggested that spiders and BDs can be detected by dipterans and hymenopterans against background leaves, but they are indistinguishable from each other. Taken together, our results suggest that insects lured by spiders may misidentify them as BDs, and bird-dropping masquerading may serve as aggressive mimicry in addition to predator avoidance in P. ceylonica.     

Olfactory learning of plant genotypes by a polyphagous insect predator

Olfactory learning may allow insects to forage optimally by more efficiently finding and using favourable food sources. Although olfactory learning has been shown in bees, insect herbivores and parasitoids, there are fewer examples from polyphagous predators. In this study, olfactory learning by a predatory coccinellid beetle is reported for the first time. In laboratory trials, adults of the aphidophagous ladybird Coccinella septempunctata did not prefer the odour of one aphid-infested barley cultivar over another. However, after feeding on aphids for 24 h on a cultivar, they preferred the odour of that particular cultivar. The mechanism appeared to be associative learning rather than sensitisation. Although inexperienced ladybirds preferred the odour of an aphid-infested barley cultivar over uninfested plants of the same cultivar, after feeding experience on a different cultivar this preference disappeared. This may indicate the acquisition and replacement of olfactory templates. The odour blends of the different aphid-infested barley cultivars varied qualitatively and quantitatively, providing a potential basis for olfactory discrimination by the ladybird. The results show that predatory coccinellids can learn to associate the odour of aphid-infested plants with the presence of prey, and that this olfactory learning ability is sensitive enough to discriminate variability between different genotypes of the same plant.