Plant scents modify innate colour preference in foraging swallowtail butterflies

Flower-visiting insects exhibit innate preferences for particular colours. A previous study demonstrated that naive Papilio xuthus females prefer yellow and red, whereas males are more attracted to blue. Here, we demonstrate that the innate colour preference can be modified by olfactory stimuli in a sexually dimorphic manner. Naive P. xuthus were presented with four coloured discs: blue, green, yellow and red. The innate colour preference (i.e. the colour first landed on) of the majority of individuals was blue. When scent from essential oils of either orange flower or lily was introduced to the room, females’ tendency to select the red disc increased. Scents of lavender and flowering potted Hibiscus rosa-sinensis, however, were less effective. Interestingly, the odour of the non-flowering larval host plant, Citrus unshiu, shifted the preference to green in females. In males, however, all plant scents were less effective than in females, such that blue was always the most favoured colour. These observations indicate that interactions between visual and olfactory cues play a more prominent role in females.     

Sex recognition in zebrafish (<Emphasis Type="Italic">Danio rerio</Emphasis>)

We investigated sex recognition in female zebrafish (Danio rerio) to better understand the underlying sensory mechanisms and identify male secondary sexual traits. Females were simultaneously presented with two fish, a male and a female, in a flow-chamber apparatus, and females’ relative attraction towards males was observed under different conditions. With domesticated fish, females were more attracted to males when presented with both visual and chemosensory cues from stimulus fish. They still discriminated the sexes when only visual cues were provided, but not when white ambient light was changed to yellow, indicating that colour plays a role. Sex discrimination under yellow light was improved when chemosensory cues were also provided. Surprisingly, females’ attraction to males was not more pronounced in the morning when mating occurs. Domesticated females discriminated the sexes when presented with wild-derived, as well as domesticated fish, whereas wild-derived females did not show any biases for domesticated or wild-derived males. Behavioural observations indicated that the wild-derived females were distressed, which explains their lack of attraction to males. In summary, domesticated female zebrafish discriminated the sexes using both visual (body colour) and olfactory cues; however, wild-derived zebrafish were too distressed for behavioural experiments under these laboratory conditions.     

Oviposition preference in the oligophagous tomato fruit fly, Neoceratitis cyanescens

We assessed the role of visual and olfactory cues on oviposition preference in the oligophagous tomato fruit fly, Neoceratitis cyanescens (Bezzi) (Diptera: Tephritidae). In a field survey, we evaluated the stage of susceptibility of field‐grown tomatoes by monitoring N. cyanescens infestations from fruit‐setting up to harvest, in relation to post‐flowering time, size, and visual properties of fruit. In two‐choice laboratory experiments, we tested the degree to which females use visual and olfactory cues to select their host plant for oviposition. In addition, we investigated the ability of flies to avoid fruit already infested by conspecific eggs or larvae, and the influence of natal host fruit on oviposition preference. Neoceratitis cyanescens females preferentially lay their eggs in small yellow‐green unripe fruit (2–3.5 cm diameter, 10–21 days post‐flowering). Damage to fruit was significantly affected by brightness and size properties. In laboratory experiments, females chose to lay their eggs in bright orange rather than yellow domes. On the sole basis of olfactory stimuli, females showed a significant preference for unripe vs. ripe host fruit, for unripe fruit vs. flowers or leaves, and for host vs. non‐host fruit (or control). However, colour interacted with odour as females dispatched their eggs equally between the yellow dome and the bright orange dome when unripe fruit of tomato was placed under the yellow dome vs. ripe fruit under the bright orange dome. When offered real ripe and unripe tomatoes, females preferred unripe tomatoes. Females significantly chose to lay eggs in non‐infested fruit when they were given the choice between these or fruit infested with larvae. In contrast, recent stings containing eggs did not deter females from laying eggs. Rather, they could have an attractive effect when deposited within <1 h. Regardless of their natal host plant, tomato or bugweed, N. cyanescens females laid significantly more eggs in a dome containing bugweed fruit. However, 15% of females originating from tomato laid eggs exclusively in the dome with tomato, against 3% of females originating from bugweed.     

Olfactory host‐finding behaviour of Oulema melanopus (Coleoptera: Chrysomelidae) and its parasitoid,Tetrastichus julis (Hymenoptera: Eulophidae)

Behavioural responses to the host‐associated olfactory cues have not been completely understood for the cereal leaf beetle, Oulema melanopus, and its principal parasitoid, Tetrastichus julis. We, therefore, investigated the role of olfactory cues in the host‐finding behaviour of these species using olfactory bioassays. Behavioural responses of O. melanopus to odours emanating from intact host plants (wheat, oat, barley) vs. a clean‐air control were tested using multichoice and two‐choice bioassays. For T. julis, responses of naïve and experienced adult female wasps to odours associated with the faecal coat of O. melanopus larvae were measured under multichoice and two‐choice conditions. Our results indicate that olfactory cues are involved in the host‐finding behaviour of both O. melanopus and T. julis. Olfactory responses of O. melanopus were influenced by the sex of the beetle and the physiological stage of adults (reproductively active vs. in reproductive diapause). Females respond to olfactory cues in greater proportions than males, and reproductively active, overwintered adults show greater responsiveness than teneral adults in reproductive diapause. Behavioural responses to cues emanating from different crop species were different in multichoice bioassays but not in two‐choice bioassays. Further, we report for the first time that the olfactory cues associated with the faecal coat of O. melanopus evoke host‐finding behaviour of its parasitoid, T. julis. Naïve female wasps are more likely to use these cues to locate the potential host than experienced females. The results of this investigation provide insights into host finding by both the species and the nature of behavioural response brought about by olfactory stimuli, and the results can help to design strategies to improve parasitoid activity by enhancing the crop environment to generate cues for host finding and to manage O. melanopus populations.     

Mate Choice by Both Sexes Maintains Reproductive Isolation in a Species Flock of Pupfish (Cyprinodon spp) in the Bahamas

Female and male mate choices can reinforce reproductive isolation after sympatric speciation. Using a binary choice design, we examine the importance of visual cues in female mate choice in all three sympatric species of pupfish on San Salvador Island. We also examine the importance of olfactory cues in female choice of the hard‐shelled invertebrate specialist (Cyprinodon brontotheroides). We examine male mate choice in two of the three species, the scale eater (C. desquamator) and the detritivore (C. variegatus). Females of all three species use visual cues and prefer conspecific males. C. brontotheroides females do not use olfactory cues to discriminate between conspecific and heterospecific males. Males of C. desquamator and C. variegatus also preferentially court conspecific females. Thus, mutual mate choice, where both females and males exhibit mate choice, acts as a strong behavioral pre‐mating isolation mechanism in these sympatrically speciated pupfish.     

The parasitoid fly Exorista japonica uses visual and olfactory cues to locate herbivore‐infested plants

Some parasitoid flies exploit odors derived from plants as olfactory cues for locating the food plants of host insects, but the role of visual cues associated with plants remains largely unknown. The generalist tachinid Exorista japonica Townsend (Diptera: Tachinidae) is attracted to odors derived from maize plants [Zea mays L. (Poaceae)] infested by the larvae of Mythimna separata (Walker) (Lepidoptera: Noctuidae). In this study, we examined the effects of visual parameters on the olfactory attraction of female flies to host‐infested plants. A paper plant model of one of four colors (blue, green, yellow, or red) was placed in front of a host‐infested plant, which was hidden behind a mesh screen in a wind tunnel. The landing rate of females was significantly higher on the green plant model than on the other three models. When an achromatic plant model of one of four gray scales (white, light gray, dark gray, or black) was tested, the response rate of females was significantly higher towards the white model and decreased as the brightness of models decreased. Few female flies responded to the green plant model without odors of the host‐infested plants. When the four color plant models were placed together in a cage filled with odors of host‐infested plants, females remained significantly longer on the green model than on the other three models. These results showed that E. japonica females preferred the color green when odors of the host‐infested plants were present and suggest that E. japonica uses visual as well as olfactory cues to locate the host habitat.     

颜色对梨小食心虫产卵选择性的影响

梨小食心虫(Grapholitha molesta Busck)是一种重要的果树害虫,早春产卵喜在桃嫩梢叶上,为探明寄主颜色在其产卵选择中的作用,利用彩色卡纸模拟寄主颜色,室内比较了红、粉红、浅粉、橙黄、深黄、浅黄、青绿、深绿、浅绿、蓝、紫、褐色等12种不同颜色基质对其成虫产卵选择性的影响。结果发现:基质颜色对梨小食心虫的产卵选择性有显著影响,其产卵偏嗜浅黄和浅绿色,白色和黑色参比时其产卵选择率分别依次为68.9%、63.8%和64.1%、65.5%,蓝和浅粉色则表现一定的拒避作用,白色和黑色参比时其产卵选择率分别依次为47.7%、40.4%和47.2%、42.7%,且参比色对其产卵选择性影响差异显著。基质颜色对其产卵量亦有显著性影响,无论黑或白色参比,黄、绿颜色上的产卵量均较多,尤其是深黄、深绿和青绿色。基质颜色对1—7日龄梨小食心虫成虫的产卵选择性均有显著影响。白色参比时,2、3日龄蛾对浅绿和浅黄色的产卵选择率显著高于其他颜色;黑色参比时,2日龄时明显偏嗜浅绿色(79.7%),6日龄时明显偏嗜浅黄色(74.8%)。表明寄主颜色在梨小食心虫产卵场所选择中具有重要作用,为其卵期监测和防控中颜色应用乃至进一步揭示其产卵寄主选择机理提供了依据和参考。     

Elevated anthocyanins protect young Eucalyptus leaves from high irradiance but also indicate foliar nutritional quality to visually attuned psyllids

1. Foliar colour changes with age and, as a consequence, reflects the internal physiology of leaves. Anthocyanins are ‘red’ pigments known for their photoprotective role in young leaves and have been suggested to influence the host‐finding behaviour of insect herbivores. The existence of colour vision in some species of Eucalyptus‐feeding psyllid provides evidence for the possibility of them being able to locate and select leaves based on their age. 2. The preferences of three psyllid species, namely Anoeconeossa bundoorensis, Glycaspis brimblecombei, and Ctenarytaina bipartita, for leaf colours were tested using live leaves of different age, presented without olfactory cues. Changes in foliar pigment concentrations and relationships with amino acid composition in these psyllid's hosts, namely Eucalyptus camaldulensis and Eucalyptus kitsoniana, were studied to consider the adaptive significance of selecting leaves based on their age. 3. The preference for and attraction to young, anthocyanic leaves of two red‐sensitive psyllid species (A. bundoorensis and G. brimblecombei) were demonstrated, whilst the green‐yellow‐sensitive species (C. bipartita) was shown to discriminate between young ‘yellow’ and older ‘green’ leaves. Age‐related variation in leaf colour was positively correlated with greater availability of essential amino acids. 4. This study presents a unique example of herbivore attraction to ‘red’ leaves and strong evidence for reliance on colour vision in insect orientation at the within‐host level.     

How do sex and sexual maturation influence the response of Monochamus galloprovincialis to host odours?

Finding the right host plant for feeding and reproduction is crucial for herbivorous insects. Usually they need both visual and olfactory cues to locate and colonize their host plants, but the respective role of the two types of stimuli is difficult to disentangle. Model plants mimicking the shape and colour of natural hosts, combined with odours dispensers, are relevant tools for this purpose. We used dummy pines and cut branches of natural pines to investigate host attraction in Monochamus galloprovincialis, the insect vector of the pine wood nematode in Europe. We compared the behavioural response of males and females, sexually mature or immature, in a large climatic chamber and in an olfactometer. Immature females were significantly attracted by the sole odours released by pines. Immature males and mature males and females needed a combination of visual and olfactory cues to find the host tree. These findings suggest that beetles use both visual and chemical cues to locate pine trees but that relevant stimuli likely differ between beetles of different sex and sexual maturation status. Our results question the implementation of precautionary clear‐cuts around infested trees for limiting the spread of PWN.     

Pheromones and body coloration affect mate recognition in the Japanese nine-spotted moth Amata fortunei (Lepidoptera: Arctiidae)

Adults of the Japanese nine-spotted moth, Amata fortunei, are diurnal and have white-spotted black wings and a black-and-yellow striped body pattern. We evaluated whether this species uses sex pheromones and whether visual cues from the female body are used in mate recognition. We introduced extracts of potentially scent-bearing abdominal tips of females to males. Males responded more to the extracts than to dried female specimens, suggesting the presence of sex pheromones in the extracts. Indeed, no males responded to the dried female specimens. To evaluate the importance of visual cues, we conducted the experiments with crude extract and an additional model stimulant. Males responded more to model females with the same number and similar area of yellow bands to the original conspecific females than to those with more bands and greater total band area, suggesting that dissimilarity in band number and area to conspecific females could interrupt male mating behavior. Males of A. fortunei likely find mating partners using olfactory cues over relatively long distances, while using both olfactory and visual cues over short distances. These results suggest that olfactory cues play a major role in mate recognition, whereas visual cues play a supplementary role.     

Oviposition preferences in Grapholita molesta: the relative importance of visual and olfactory cues

The oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae), is a major pest of fruit trees worldwide. Females oviposit on the young leaves of peach [Prunus persica (L.) Batsch (Rosaceae)] shoots at twilight, when light intensity changes markedly. Previous researches have shown that the oviposition response of this moth is guided particularly by visual and olfactory cues from host plants. However, the relative importance of visual and olfactory cues in their oviposition preference is largely unknown. As a crepuscular moth, how do choices change as light intensity drops rapidly from day to night? In the present study, through two‐choice behavioural experiments, the oviposition responses of G. molesta to visual and olfactory cues (alone or in combination) from peach shoots were tested at four light intensities ranging from daylight to starlight. Grapholita molesta showed similar oviposition responses to plant cues under 1 000, 100, 1 and 0.01 mW m^?2 illumination. Olfactory cues from peach shoots attracted significantly more oviposition than the negative control, whereas visual cues alone did not. Furthermore, olfactory cues were more attractive than visual cues, and no significant interaction was observed between the responses to the two cues. Our findings indicate that G. molesta females mainly rely on olfactory cues to recognize oviposition sites, regardless of differences in light intensity. These results do not provide evidence for attraction to visual cues, but the possibility that the brightness of leaves might be used to guide oviposition is discussed.     

Visual cues override olfactory cues in the host-finding process of the monophagous leaf beetle Altica engstroemi

It is generally assumed that specialist insect herbivores utilize plant odours to find their particular host plants and that visual cues are of minor importance in the host‐finding process. We performed Y‐tube olfactometer bioassays and small‐scale field experiments to determine whether, under laboratory and field conditions, the monophagous herbivore Altica engstroemi J. Sahlberg (Coleoptera: Chrysomelidae: Alticinae) is guided to its host plant Filipendula ulmaria (L.) Maxim. (Rosaceae) by visual or olfactory cues. The olfactometer tests showed that A. engstroemi was never attracted to odours, either from undamaged or from damaged plants. Even starvation for 24 h did not change this behaviour. However, the field experiment showed that visual cues alone were sufficient to attract a significant number of starved beetles when offered a choice between bagged host plants and bagged green plastic control ‘plants’. Our findings contrast with the general view that plant odours constitute the major cue in the host‐finding process among specialized phytophagous insects. A review of the literature for the period 1986–2006 inclusive, relating to host‐plant finding in Chrysomelidae, identified studies of 19 chrysomelid species, all of which were guided by olfactory cues. No species were guided to their host by visual cues. Although some studies demonstrated that chrysomelids may exhibit orientation responses to colour or contrast, our study on A. engstroemi is the only one demonstrating that visual cues affect host‐plant selection in a chrysomelid species. We suggest that the use of visual cues in host‐finding may evolve among chrysomelids with limited dispersal ability in persistent habitats and may be found among species monophagous on abundant host plants that dominate the structure of the plant community, that is, where the host plant's presence is predictable in time and space.     

Female mate choice and species recognition between two closely related cichlid fish of Lake Malawi,Metriaclima estherae and M. callainos

Cichlids are one of the most diverse and colourful groups of freshwater fishes in the world. Despite much investigation, the factors that promote speciation in these fishes are still uncertain. However, previous studies suggest that sexual selection on male colour is one of the main drivers of speciation among these fishes. Metriaclima estherae is a polymorphic cichlid species from Lake Malawi, and thus provides an ideal model for the investigation of the importance of colour as a species recognition cue. M. callainos is a closely related and morphologically similar species, with male colour pattern very similar to that of M. estherae. We tested female choice by giving females of the two species a choice between conspecific and heterospecific males in the presence and absence of visual (colour) and chemical cues. The results show that females of M. callainos were able to reliably recognize conspecific males, even when colour was eliminated as a cue. However, females of M. estherae did not prefer conspecific males, although they were able to discriminate between red and blue conspecific colour morphs by using chemical cues. These results suggest that species recognition cues may differ even among closely related species of cichlid fish, and that female preferences for male coloration may be weak in certain species.     

Nonvisual mate choice in the Pyrenean mountain newt (<Emphasis Type="Italic">Euproctus asper</Emphasis>): females prefer small males

In many species, females prefer large males as mates. In this paper, we examined whether female Pyrenean mountain newts (Euproctus asper) would show a preference for large males in a simultaneous choice situation. When we tested for a visual preference relative to male body size using visual cues, females did not prefer large males. When chemical cues were presented, females did show a preference, but contrary to our prediction, they preferred the smaller of two males. E. asper males exhibit coercive mating behavior (amplexus), keeping females from feeding for several hours or even days, which may explain why females avoid large, physically stronger, and thus, more coercive males. The predominantly nocturnal lifestyle may explain why visual cues are less important for mate discrimination in this species.     

Behavioural plasticity and sex differences in host finding of a specialized bee species

Many animals feed on flowers, and visual as well as olfactory cues are considered as most important mediators in animal–plant interactions. However, the relative importance of these cues is not well understood. Bees are the most important animal pollinators worldwide and here, we determined the importance of decoupled and combined visual and olfactory cues of Lysimachia punctata (Primulaceae) for host plant location in both sexes of the specialized, solitary bee, Macropis fulvipes (Melittidae). Lysimachia-inexperienced female bees preferred olfactory over visual cues though visual cues increased the attractiveness of olfactory ones. In experienced females, the importance of visual cues was increased. Both Lysimachia-naive and -experienced males relied more on visual cues as compared to females. This study demonstrates that the relative weighting of cues used for host plant finding depends on the sex and experience of M. fulvipes. The latter finding reveals the presence of learning-induced behavioural plasticity in host plant finding for a bee species. It may allow the bee to forage highly efficient. Visually guided female detection on flowers by males is a likely functional explanation for the differences in the weighting of visual and olfactory cues between the sexes.     

Ultraviolet reflection and female mate choice in the pied flycatcher, Ficedula hypoleuca

In pied flycatchers females seem to prefer male territory quality rather than male characteristics, and the results of female mate choice experiments are divergent. In this outdoor aviary study, we examined how altering the ultraviolet reflection of males affects female mate choice behaviour. We chose pairs of males with similar human-visible dorsal colour and morphological traits. We then reduced the proportional ultraviolet reflectance in one male with sunscreen chemicals. The other male was treated with a chemical that slightly increased the ultraviolet reflectance of the plumage. In the experiment females clearly preferred males with slightly increased ultraviolet reflection. Our results indicate that pied flycatcher females use ultraviolet cues for mate choice when the effect of territory quality is controlled for. The results give us new information about a possible mechanism of mate assessment in this species, and indicate the importance of colour cues in avian mate choice behaviour.     

Sugar in moderation: sugar diets affect short‐term parasitoid behaviour

The biological control potential of parasitic wasps in the field is expected to increase with the provisioning of sugar sources, which increase longevity and replenish carbohydrate reserves. Apanteles aristoteliae is an important parasitoid of Argyrotaenia franciscana, the orange tortrix, an economic pest in fruit crops. In the present study, the effect of sugar diet on the physiological status of A. aristoteliae is investigated in the laboratory, as well as the effects of nutritional status on short‐term olfactory orientation and parasitism behaviour, and the association between olfactory orientation and immediate parasitism activity. Levels of glycogen, fructose, total sugars, proportional weight gain and volume consumed are higher among females fed 25% sucrose solution than 10% sucrose solution, and lowest for those fed water. Sugar feeding also affects behaviour: wasps with higher levels of fructose or weight gain have a lower probability of making a choice in the olfactometer. Wasps with intermediate fructose levels or weight gain have a higher probability of orienting towards hosts than wasps with low or high levels. Among wasps that make a choice in the olfactometer, wasps fed 10% or 25% sucrose significantly prefer host versus food cues, whereas starved wasps are just as likely to select food cues as host cues.     

Influence of visual cues on host‐searching and learning behaviour of the egg parasitoids Telenomus podisi and Trissolcus basalis

Insect parasitoids use a variety of chemical and physical cues when foraging for hosts and food. Parasitoids can learn cues that lead them to the hosts, thus contributing to better foraging. One of the cues that influence host‐searching behaviour could be colour. In this study, we investigated the ability of females of the parasitoid wasps Telenomus podisi Ashmead and Trissolcus basalis Wollaston (both Hymenoptera: Scelionidae) to respond to colours and to associate the presence of hosts – eggs of Euschistus heros (Fabricius) (Hemiptera: Pentatomidae) – with coloured substrates after training (associative learning). Two sets of experiments were conducted: in one the innate preference for substrate colours was examined, in the other associative learning of substrate colour and host presence was tested in multiple‐choice and dual‐choice experiments. In the associative learning experiments, Te. podisi and Tr. basalis were trained to respond to differently coloured substrates containing hosts in two sessions of 2 h each, with 1‐h intervals. In multiple‐choice experiments, the wasps displayed innate preference for yellow substrates over green, brown, black, or white ones. Even after being trained on substrates of different colours, both parasitoids continued to show preference for yellow substrates. The response to the colours of substrates of both parasitoids was related with the orientation to the plant foliage during the search for hosts.     

Lack of evolution in a leaf beetle that lives on two contrasting host plants

The interactions between plant‐eating insects and their hosts have shaped both the insects and the plants, driving evolution of plant defenses and insect specialization. The leaf beetle Trirhabda eriodictyonis (Chrysomelidae) lives on two shrubs with differing defenses: Eriodictyon crassifolium has hairy leaves, whereas E. trichocalyx has resinous leaves. We tested whether these beetles have differentiated onto the two host plants, and if not, whether the beetles prefer the better host plant and prefer mates who are from that host plant. In feeding tests, adult beetles strongly preferred eating E. trichocalyx regardless of which host they came from. In addition, females laid more eggs if they ate E. trichocalyx than E. crassifolium. So, E. trichocalyx is generally the better host. However, beetle mate preference was not in line with food choice. Males did not prefer to mate with females from E. trichocalyx. Females from E. crassifolium did prefer males from E. trichocalyx over males from E. crassifolium, but did not lay more eggs as a result of these matings. We conclude that the beetle populations we studied have not differentiated based on their host plants and may not have even adapted to the better host. Although to humans these host plant defenses differ dramatically, signs that they have caused evolution in the beetles are lacking. The case of T. eriodictyonis stands counter to many other studies that have seen the differentiation of ecotypes and/or adaptive coordination of an herbivore's life cycle based on host plant differences.     

Host race formation inTetranychus urticae: genetic differentiation, host plant preference, and mate choice in a tomato and a cucumber strain

The two-spotted spider mite,Tetranychus urticae Koch, occurs in two colour forms in greenhouses in the Netherlands: a red form on tomato and a green form on cucumber. The evolutionary status of these strains was analysed by studying genetic differentiation, host plant preference, and mate choice. Males of the tomato strain preferred the female (30 h-old) teleiochrysales from the same strain to those of the cucumber strain, independent of the host plants (tomato, cucumber, bean) on which the teleiochrysales were placed. In contrast, males of the cucumber strain were not selective. In a Y-tube olfactometer, females of the cucumber strain were not responsive to host plant volatiles alone. However, in two-choice disc experiments, where females were exposed to both volatile and contact cues, they settled on cucumber leaves in preference to tomato leaves. Females of the tomato strain preferred the odour of tomato leaves and settled on tomato leaves in preference to cucumber leaves. These experimental results provide the first evidence for (1) host-plant independent mate selection in male spider mites and (2) olfactory discrimination between host plants in female spider mites. Electrophoretic analysis showed much genetic differentiation at the phosphoglucose isomerase locus. The cucumber strain showed large variation with 5 alleles, whereas the tomato strain was fixed for the most common allele of the cucumber strain. The results suggest that the two strains represent host races. We hypothesize that the tomato strain has originated from the cucumber strain because (1) tomato represents a more hostile host plant to spider mites (due to toxic compounds and glandular hairs) and (2) the tomato strain is genetically impoverished suggesting that it passed through one or more bottlenecks.