Oviposition behaviour and host colony size discrimination in Episyrphus balteatus (Diptera: Syrphidae)

Three bioassays were used to examine the oviposition behaviour of the hoverfly, Episyrphus balteatus (Degeer) in which various stimuli purported to influence host assessment and choice were tested. Episyrphus balteatus failed to exhibit enhanced behavioural responses, in terms of approaches and landings, to artificial leaves with the highest numbers of aphids, suggesting that females are merely responding to the colour of the artificial leaves. A density-dependent oviposition response was reported in a second bioassay using whole bean plants, although there was an asymptotic relationship. Syrphid responses, which were measured by oviposition over a five-day period, were greatest towards those plants with the highest number of aphids. In a final series of trials, gravid female E. balteatus showed a behavioural preference for filter papers treated with the greatest honeydew concentrations (0.26 mg microl(-1) honeydew). More time was spent in the treated areas and syrphids demonstrated more proboscis and ovipositor extensions in these treatments. These bioassays provided an opportunity to isolate some of the individual components of decision-making by female E. balteatus during egg-laying behaviour. A greater understanding of natural enemy behaviour is essential before enhanced control of pest populations in the field can be established.     

Density-dependent and frequency-dependent selection by bumblebees Bombus terrestris (L.) (Hymenoptera: Apidae)

The behaviour of bumblebee workers foraging on arrays of artificial flowers of two colour morphs was observed. Experiments were conducted on arrays of varying morph frequencies and at three different total flower densities. Bumblebees consistently showed a preference for the commonest colour morph, and this behaviour was not significantly affected by changing density. In contrast, frequency-independent preferences changed significantly with density. At low densities, there was a strong bias towards the more conspicuous colour, whereas at higher densities there was no overall colour bias. Flight distances between flowers decreased significantly at high density. Bumblebees also visited flowers of similar colours sequentially, but this behaviour was not density-dependent. It is suggested that as densities increase, there is an increased probability that bumblebees detect yellow flowers, which were probably less conspicuous compared with blue flowers, and that this might be caused by changes in flight speed with flight distance. Where there is a positive relationship between pollinator visitation and the relative fitness of a floral morph, the observed behaviour would induce positive frequency-dependent selection on a plant population with two corolla colour morphs on which the bumblebees were foraging, which would result in stabilizing selection for a single corolla colour, irrespective of density. There was no indication that rare colour morphs would be preferred at high density. The probability of different corolla colour morphs going to fixation would, however, be affected by density.     

Pink flower preference in sunbirds does not translate into plant fitness differences in a polymorphic Erica species

Bird-pollinated plants typically have reddish flowers, but it is not clear whether this trait can be attributed to selection by birds. Here we experimentally test for the first time the foraging behaviour of sunbirds in relation to flower colour, using the Orange-breasted Sunbird Anthobaphes violacea (Nectariniidae) and the colour dimorphic Erica perspicua (Ericaceae). Pink and white flower morphotypes co-flower in intermixed populations and have similar nectar volumes and concentrations. Using floral arrays in a field aviary, we found that sunbirds preferred pink flowers; 95 % of their first choices were to pink inflorescences and they visited and probed more pink inflorescences and flowers, respectively. We also tested for flower constancy (the tendency to move between same colour rather than different colour morphotypes), but found no evidence for this in the sequence of their foraging choices, indicating that this mechanism did not maintain flower colour differences in sympatry. There was evidence for optimal foraging: 80 % of moves were to adjacent inflorescences. Unexpectedly, the preference for pink flowers observed in the aviary did not translate into a female fitness advantage for this morphotype in the field, since no difference is found in natural pollination rate, fruit or seed set. This may be because the minimization of flight distances between plants is the primary factor in sunbird foraging choices, overriding their colour preference. Antagonistic nectar robbers did not act as a selective force on the polymorphism, since nectar-robbing rates were equal between white and pink morphotypes in the field.     

Honeybee (Apis mellifera ligustica) Response to Differences in Handling Time, Rewards and Flower Colours

Free flying honeybees were tested outdoors on blue–white and blue–yellow dimorphic artificial flower patches to examine the influence of reward difference, flower handling‐time difference and flower colour choice on foraging decisions. We employed different flower‐well depths to vary handling times (costs), and differences in sucrose molarity to vary reward quality. Tests were performed with 2 and 6 μl rewards to vary quantity. We show that when handling time is correlated with flower‐colour morphs on a pedicellate artificial flower patch, a honeybee's foraging behaviour is dependent on the flower colours used in the choice tests. This supports a honeybee foraging model where constraints are a significant factor in decision making. Bees visiting blue–yellow flower patches exhibited flower constancy to colour, where they restricted most visits to a single flower colour, some bees to blue and others to yellow, irrespective of handing time differences. When offered a choice of equally rewarding blue or white flowers, bees were not constrained by flower colour and chose to visit flowers with a lower handling time. When reward molarity varied with well depth between blue and white flowers, foragers chose shallow‐well flowers (short‐handling time) with a smaller net harvest rate over deep‐well flowers (long‐handling time) with a greater net harvest rate. Results using the blue–white dimorphic flower patch suggest that when foraging options simultaneously involve reward and handling‐time choices, honeybee forager behaviour is inconsistent with an absolute method of evaluating profit.     

Odour and colour information in the foraging choice behaviour of the honeybee

1. Honeybees Apis mellifera ligustica were trained to work on a patch with artificial rewarding and non-rewarding flowers, coupled to an air extractor. The perceptual colour distance between the rewarding and the non-rewarding flowers was varied and the flower choice and the repellent scent-marking behaviour of the bees were recorded. 2. The discrimination between rewarding and non-rewarding flowers depended on their colour distance, improving with a greater colour difference. This task was guided thus visually and was not affected by activating the air extractor. 3. The scent-marking activity was only observable when the colour information of both groups of flowers was the same or very similar. This thus represents the first reported case of a modulation of an olfactory activity through the visual input provided by colour distances. When the air extractor was activated, rejections associated with the scent-marking behaviour disappeared, thus confirming the olfactory nature of this behaviour. 4. Honeybees are thus capable of using one or more sensory cues to enhance their foraging efficiency, according to the environmental situation. This great plasticity allows them to attain an enhanced efficiency while foraging. 5. We successfully applied the model of colour choice behaviour of the honeybee. Since the original theory was developed for Apis mellifera carnica, this work also constitutes the first attempt to describe the behaviour of the honeybee race, Apis mellifera ligustica, using the postulated model, and reaffirms thus its generality.     

Colour association influences honey bee choice between sucrose concentrations

Certain colours associated with floral food resources are more quickly learned by honey bees (Apis mellifera) than are other colours. But the impact of colour, and other floral cues, on bee choice behaviour has not yet been determined. In these experiments, colour association and sugar concentration of reward were varied to assess how they interact to affect bee choice behaviour. Thirty-five bees were individually given binary choices between blue and yellow artificial flowers that contained either the same rewards or rewards of different sucrose concentrations. Honey bee choice between sucrose concentrations was affected by colour association and this effect was greatest when absolute difference between rewards was the lowest. The honey bee's ability to maximize energetic profitability during foraging is constrained by floral cue effectiveness.     

Risk‐averse inflorescence departure in hummingbirds and bumble bees: could plants benefit from variable nectar volumes?

Most hermaphroditic, many-flowered plants should suffer reduced fitness from within-plant selfing (geitonogamy). Large inflorescences are most attractive to pollinators, but also promote many flower visits during a single plant visit, which may increase selfing and decrease pollen export. A plant might avoid the negative consequences of attractiveness through modification of the floral display to promote fewer flower visits, while retaining attractiveness. This report shows that increasing only the variance in nectar volume per flower results in fewer flower visits per inflorescence by wild hummingbirds ( Selasphorus rufus ) and captive bumble bees ( Bombus flavifrons ) foraging on artificial inflorescences. Inflorescences were either constant (all flowers contained the same nectar volume) or variable (half the flowers were empty, the other half contained twice as much nectar as in the constant flowers). Both types of inflorescence were simultaneously available to foragers. Risk-averse foraging behaviour was expressed as a patch departure preference: birds and bees visited fewer flowers on variable inflorescences, and this preference was expressed when resource variability could be determined only by concurrent sampling. When variance treatments were clearly labelled with colour and offered to hummingbirds, the departure effect was maintained; however, when preference was measured by inflorescence choice, birds did not consistently prefer to visit constant inflorescences. The reduced visitation lengths on variable inflorescences by both birds and bees documented in this study imply that variance in nectar production rates within inflorescences may represent an adaptive trait to avoid the costs of geitonogamy.     

Role of prey–host plant associations on Harmonia axyridis and Episyrphus balteatus reproduction and predatory efficiency

In order to predict possible locations of Harmonia axyridis Pallas (Coleoptera: Coccinellidae) and Episyrphus balteatus DeGeer (Diptera: Syrphidae) in the field, we studied their oviposition and prey preferences in relation to several host plant–prey associations under laboratory conditions. Oviposition preference of H. axyridis and E. balteatus females was determined for three aphid (Homoptera: Aphididae)–host plant associations: Microlophium carnosum Buckton on stinging nettle [ Urtica dioica L. (Urticaceae)], Acyrthosiphon pisum Harris on green pea [ Pisum sativum L. (Fabaceae)], and Sitobion avenae F. on wheat [ Triticum aestivum L. (Poaceae)]. Prey preference of H. axyridis and E. balteatus larvae was determined with the aphids M. carnosum , A. pisum , and S. avenae . Harmonia axyridis females showed a strong oviposition preference for the stinging nettle– M. carnosum association. The preferred association for ovipostion by E. balteatus was pea-hosting A. pisum , on which fecundity was also highest. Feeding behaviour was less restricted in H. axyridis , in which the preferred preys were M. carnosum and S. avenae . A lower specificity of predation was observed in E. balteatus larvae with respect to A. pisum . These laboratory experiments may help us to understand the spatial distribution of H. axyridis and E. balteatus in the field.     

Discrimination of parasitized aphids by a hoverfly predator: effects on larval performance, foraging, and oviposition behavior

The choice of oviposition site by female aphidophagous predators is crucial for offspring performance, especially in hoverflies whose newly hatched larvae are unable to move over large distance. Predator and parasitoid interactions within the aphidophagous guild are likely to be very important in influencing the choices made by predatory hoverfly females. In the present study, the foraging and oviposition behavior of the aphidophagous hoverfly Episyrphus balteatus DeGeer (Diptera: Syrphidae) was investigated with respect to the parasitized state of its aphid prey, Acyrthosiphon pisum Harris (Homoptera: Aphididae), that were parasitized by Aphidius ervi Haliday (Hymenoptera: Aphidiidae). We also recorded the number of eggs laid by hoverfly females when subjected to parasitized aphids. Furthermore, we studied the influence of being fed with parasitized aphids on hoverfly larval performance. Hoverfly females did not exhibit any preference for plants infested with unparasitized or aphids parasitized for 7 days. On the other hand, plants infested with mummies or exuvia were less attractive for E. balteatus . These results were correlated with (i) the number of eggs laid by E. balteatus females and (ii) larval performance. Thus, our results demonstrate that E. balteatus behavior is affected by parasitoid presence through their exploitation of aphid colonies. Indeed, hoverfly predators select their prey according to the developmental state of the parasitoid larvae.     

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.     

Innate and Learned Colour Preference in the Zebrafish, Danio rerio

Receiver biases towards specific sensory signals have been demonstrated in insects, birds and fish, both in the context of foraging and mate choice. In some cases, signals important in sexual selection appear to have evolved by exploiting a pre-existing bias in the sensory system. For instance, female preferences for male nuptial colouration may have arisen from selection on foraging practices. Using the zebrafish ( Danio rerio ), a species in which red is not a factor in mate choice, we tested for a foraging bias towards the colour red. We further investigated the plasticity of foraging biases by raising groups of fish on diets consisting solely of red, blue, green or white food. When we subsequently tested their colour preferences in a foraging context, each group responded most strongly to red, irrespective of the colour of food with which they had been conditioned. We also detected a significant effect of conditioning on colour preferences; fish responded more strongly to the colour that matched diet colour than to other colours. The observed receiver bias towards red may have evolved as an adaptive preference for carotenoid compounds in their diet. While the bias to red appears to be innate, our results indicate that learning is also important in shaping foraging biases.     

The role of pollinators in maintaining variation in flower colour in the Rocky Mountain columbine,Aquilegia coerulea

Background and Aims Flower colour varies within and among populations of the Rocky Mountain columbine, Aquilegia coerulea, in conjunction with the abundance of its two major pollinators, hawkmoths and bumble-bees. This study seeks to understand whether the choice of flower colour by these major pollinators can help explain the variation in flower colour observed in A. coerulea populations.Methods Dual choice assays and experimental arrays of blue and white flowers were used to determine the preference of hawkmoths and bumble-bees for flower colour. A test was made to determine whether a differential preference for flower colour, with bumble-bees preferring blue and hawkmoths white flowers, could explain the variation in flower colour. Whether a single pollinator could maintain a flower colour polymorphism was examined by testing to see if preference for a flower colour varied between day and dusk for hawkmoths and whether bumble-bees preferred novel or rare flower colour morphs.Key Results Hawkmoths preferred blue flowers under both day and dusk light conditions. Naïve bumble-bees preferred blue flowers but quickly learned to forage randomly on the two colour morphs when similar rewards were presented in the flowers. Bees quickly learned to associate a flower colour with a pollen reward. Prior experience affected the choice of flower colour by bees, but they did not preferentially visit novel flower colours or rare or common colour morphs.Conclusions Differences in flower colour preference between the two major pollinators could not explain the variation in flower colour observed in A. coerulea. The preference of hawkmoths for flower colour did not change between day and dusk, and bumble-bees did not prefer a novel or a rare flower colour morph. The data therefore suggest that factors other than pollinators may be more likely to affect the flower colour variation observed in A. coerulea.     

Flower constancy and learning in foraging preferences of the green-veined white butterfly Pleris napi

Abstract.

• 1 Evolutionary pressure should select for efficient foraging strategies, within the constraints of other selective forces. We assess the mechanisms underlying flower choice in the butterfly, Pieris napi (L.), which as an adult forages for nectar. Experiments were carried out on a laboratory colony, using artificial flowers of two colours, and replicated on two successive generations.
• 2 When nectar was freely available from all flowers, equal numbers of butterflies visited each colour, but individual butterflies exhibited flower constancy, showing a strong preference for one colour or the other.
• 3 Following 3 day conditioning periods in which nectar was available from flowers of one colour only, butterflies responded by developing a preference for this colour, which persisted when both flower colours were refilled. This preference could subsequently be switched to the other flower colour following a further 3 days of conditioning. These are interpreted as adaptive (learned) responses, which would have obvious selective benefits in the field, enabling butterflies to avoid flower species which experience has shown are poor sources of nectar, and to adapt to temporal and spatial changes in nectar availability.

     

Flower choice in honeybees: Effects of food value and flower density

To test the effects of food value on the flower choice, individual honeybees (Apis mellifera) were offered a choice of 25 % sucrose solution (SS) and 1 of 6 different SSs, ranging from 5 % to 50 % SS, at either a low or a high flower density. Artificial flowers were filled with each SS. The honeybees showed a stronger preference for a concentrated SS to a diluted SS at a high than at a low flower density, and the degree of preference was positively correlated to the difference in the sucrose concentration between paired SSs. These foraging patterns were consistent with qualitative predictions from optimal foraging theory. Furthermore, it was found that experience in feeding on a concentrated SS lowered the foraging motivation for a diluted SS at the high flower density, but not at the low flower density. I discuss the effects of food density, food profitability and experience on the foraging behaviour of honeybees.     

The spatial distribution of nonrewarding artificial flowers affects pollinator attraction

Many species of orchids that do not offer food rewards to pollinators bloom in clusters, early in the season, and are polymorphic for corolla colour. Previous studies suggest that the foraging behaviour of insect pollinators may select for early blooming and colour polymorphism. I tested whether pollinator behaviour can also favour aggregated flowering in these species, in a two-stage laboratory experiment on na?ve bumblebees, Bombus terrestris (L.). In the first stage, the bees were allowed to forage on three colours of artificial flowers that contained sucrose rewards. In the second stage, I added nonrewarding flowers of a fourth colour and recorded the bees' visits to them. The four types of artificial flowers were either arranged in spatially distinct clusters, or were randomly intermingled. I used two reward schedules for each spatial arrangement: constant refilling of reward-containing flowers and probabilistic refilling. Bees that foraged on clustered flowers flew more often to the nonrewarding patch, and made more visits to nonrewarding flowers, than bees that foraged on intermingled flowers. This tendency was obtained both in the constant reward and in the probabilistic reward schedules. The results support the hypothesis that pollinator attraction may select for clustered, synchronized blooming in flowers that do not contain nectar and pollen rewards. Copyright 2000 The Association for the Study of Animal Behaviour.     

Bumblebees (Bombus terrestris) and honeybees (Apis mellifera) prefer similar colours of higher spectral purity over trained colours

Differences in the concentration of pigments as well as their composition and spatial arrangement cause intraspecific variation in the spectral signature of flowers. Known colour preferences and requirements for flower-constant foraging bees predict different responses to colour variability. In experimental settings, we simulated small variations of unicoloured petals and variations in the spatial arrangement of colours within tricoloured petals using artificial flowers and studied their impact on the colour choices of bumblebees and honeybees. Workers were trained to artificial flowers of a given colour and then given the simultaneous choice between three test colours: either the training colour, one colour of lower and one of higher spectral purity, or the training colour, one colour of lower and one of higher dominant wavelength; in all cases the perceptual contrast between the training colour and the additional test colours was similarly small. Bees preferred artificial test flowers which resembled the training colour with the exception that they preferred test colours with higher spectral purity over trained colours. Testing the behaviour of bees at artificial flowers displaying a centripetal or centrifugal arrangement of three equally sized colours with small differences in spectral purity, bees did not prefer any type of artificial flowers, but preferentially choose the most spectrally pure area for the first antenna contact at both types of artificial flowers. Our results indicate that innate preferences for flower colours of high spectral purity in pollinators might exert selective pressure on the evolution of flower colours.     

Preferences for coloured bower decorations can be explained in a nonsexual context

The sensory bias model of sexual selection suggests that elaborate male secondary sexual traits evolved to exploit biases in the female's sensory system. Such biases may have evolved in a nonsexual context. Male bowerbirds build and decorate elaborate structures, bowers, which function as targets of female choice. The colour of decorations used on bowers appears to be important in determining a male's mating success. We tested two predictions made by the sensory bias model, using cache presentation experiments of artificially coloured grapes made to captive bowerbirds of five species. We first searched for evidence of ancestral biases for certain colours that could explain current colour preferences across species. We found no single parsimonious explanation for ancestral patterns of colour preference across the family. We next tested whether female preference patterns could be explained in a nonsexual, foraging, context. For both of the species where sufficient data could be collected, female foraging preferences for grapes were significantly related to male preferences for grapes used as bower decorations. Our results suggest that choice of bower decoration colour may have evolved to exploit a bias in the female's sensory system, originally shaped by selection on foraging practices. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Colour biases in learned foraging preferences in Trinidadian guppies

Learning allows animals to adaptively adjust their behaviour in response to variable but predictable environments. Stable aspects of the environment may result in evolved or developmental biases in the systems impacting learning, allowing for improved learning performance according to local ecological conditions. Guppies (Poecilia reticulata), like many animals, show striking colour preferences in foraging and mating contexts and guppy artificial selection experiments have found that the form and progress of evolved responses to coloured stimuli differ depending on stimulus colour. Blue colouration is thought to typically be a relatively unimportant food cue in guppies. This raises the possibility that learned foraging associations with blue objects are formed less readily than with other colours. Here, guppies were rewarded for foraging at green or blue objects in two experiments. Guppies readily foraged from these objects, but learning performance differed with rewarded object colour. With equal amounts of training, the preference for green objects became stronger than the preference for blue objects. These differences in performance were not attributable to differences in initial preferences or to foraging more on one colour during training. These findings suggest that associative pairings within a single sensory modality that do not have a historic relevancy can be more difficult for animals to learn even when there is no clear initial bias present.     

Conspicuousness, not colour as foraging cue in plant–animal signalling

The global prevalence of red and black fruits has still not been explained. Hypotheses based on innate consumer preferences have been tested and rejected. Though colour itself plays an important role in animal foraging, it is only one component of signals. Another major component are colour contrasts against background achieving the conspicuousness of signals. In order to evaluate which signal component determines consumers behaviour, we measured fruit colour and colour contrasts of 43 species against their natural background under ambient light conditions. Red and black fruits exhibit stronger contrasts and are therefore more conspicuous to consumers than fruits of other colours. Subsequently, trials were carried out to determine whether colour or conspicuousness influences avian food choice. Four bird species strongly preferred contrasting red–green or black–green over uni-coloured red, green, or black fruit displays, while no preference for particular hues was found. We therefore hypothesize that conspicuousness determines avian food selection and define the contrast hypothesis: Diurnal dispersers select fruit colours based on their conspicuousness and not their colour itself.
Because colour vision is an ancient trait, the entire heterogeneous group of frugivorous birds most likely perceives conspicuousness uniformly over evolutionary time spans. Conspicuousness has thus the potential to explain the global prevalence of red and black fruits.     

Colour choices of naive bumble bees and their implications for colour perception

The innate preferences of inexperienced bumble bees, Bombus terrestris, for floral colour stimuli were studied using artificial flowers. The artificial flowers provided a colour pattern and consisted of a star-shaped corolla and of central colour patches similar to the nectar guide of natural flowers. The innate choice behaviour was assessed in terms of the number of approach flights from some distance towards the artificial flowers and the percentage of approach flights terminating in antennal contact with the floral guide. The colours of the floral guide, the corolla and the background were varied. It was shown that the innate flower colour preference in bumble bees has two components. 1. The frequency of approaches from a distance is correlated with the colour difference between the corolla and the background against which it is presented. If the corolla colour was constant but its background colour varied, the relative attractiveness of the corolla increased with its colour difference to the background. The colour difference assessment underlying this behaviour on a perceptual basis can be attained by means of colour opponent coding, a system well-established in Hymenoptera. 2. The frequency of antennal contacts with the floral guides relative to that of approach flights cannot be accounted for by colour opponent coding alone. Whether the approach flights are interrupted, or whether they end in an antennal contact with the nectar guide is strongly dependent on the direction (sign) of the colour difference, not only its magnitude. The choice behaviour requires a unique perceptual dimension, possibly that of colour saturation or that of hue perception comparable to components of colour perception in humans.