Male mate searching behavior in the black chafer Holotrichia loochooana loochooana (Sawada) (Coleoptera: Scarabaeidae): response to pheromone lures of various colors and intensities

In order to understand the visual orientation mechanisms of the male black chafer Holotrichia loochooana loochooana to females, we examined male behavioral response to colored objects (lures) when female sex pheromone was present. In the field, males landed more frequently on darker lures than on lighter ones of the same color. Among lures of different colors but with the same brightness, males preferred landing on black, blue, or red ones to green or grey ones. The reflectance intensities of preferred lures were weaker than those of green and grey in the range of 540–580 nm. This range is considered to affect choice by males. When females perch on the leaf edges of the food plant to adopt a calling posture, the dominant background color is that of the leaves (i.e., green). This means that the landing target forms a silhouette (dark spot) which is prominent against a background of green leaves, providing maximum visual acuity for mating.     

Variation in Nectar and Pollen Availability,Sucrose Preference,and Daily Response in the Use of Flowers by <Emphasis Type="Italic">Heliconius erato phyllis</Emphasis>

The mechanisms mediating the use of flowers in the butterfly Heliconius erato phyllis (Nymphalidae) are poorly understood. Availability of nectar and pollen, nectar concentration, and abundance of Stachytarpheta cayennensis and Lantana camara (Verbenaceae), two flower species commonly used by H. erato phyllis in the Neotropics, as well as flower use by this butterfly species in the field were examined in southern Brazil. Under insectary conditions, the preference of H. erato phyllis for different sucrose concentrations (0, 10, 20, 40, and 80%) and the ability to associate sucrose concentrations with preferred and non-preferred flower colors were evaluated through choice tests. Lantana camara inflorescences were less abundant, but contained larger amounts of pollen and nectar than S. cayennensis, and H. erato phyllis utilized the flowers of the former species with higher frequency compared to the latter. In the choice tests, butterflies fed more intensely on 20 and 40% sucrose solutions, an interval in which the nectars of L. camara and S. cayennensis are situated, and were able to associate preferred sucrose concentrations with flower color efficiently within the color spectrum of L. camara flowers (i.e., preferred colors), but not within that of S. cayennensis (non-preferred colors). Thus, the greater use of L. camara flowers by H. erato phyllis is related to the plant’s superior floral rewards and not flower abundance, and to the cognitive abilities of these butterflies to adjust their feeding to the availability of pollen and nectar. To our knowledge, this is the first report showing sucrose preferences in a butterfly species.     

Why are fruits colorful? The relative importance of achromatic and chromatic contrasts for detection by birds

The colors of fruits and flowers are traditionally viewed as an adaptation to increase the detectability of plant organs to animal vectors. The detectability of visual signals increases with increasing contrasts between target and background. Contrasts consist of a chromatic aspect (color) and an achromatic aspect (light intensity), which are perceived separately by animals. To evaluate the relative importance of fruits’ chromatic and achromatic contrasts for the detection by avian fruit consumers we conducted an experiment with artificial fruits of four different colors in a tropical forest. We displayed the fruits against two different backgrounds, an artificial background and a natural one, because they differed in achromatic properties. We found no effect of the type of background on fruit detection rates. Detection rates differed for the four fruit colors. The probability of detection was explained by the chromatic contrast between fruits and their background, not by the achromatic contrasts. We suggest that birds attend primarily to chromatic contrast probably because these are more reliably detected under variable light conditions. Consistent with this hypothesis, we found habitat-specific differences in the conspicuousness of natural fruit colors in the study area. Fruits of understory species that are subjected to the variable light conditions within a forest displayed higher chromatic contrasts than species growing in the open restinga forest with constant bright illumination. There was no such difference for achromatic contrasts. In sum, we suggest that fruit colors differ between habitats because fruit colors that have strong chromatic contrasts against background can increase plants’ reproductive success, particularly under variable light conditions.     

UV bullseye contrast of Hemerocallis flowers attracts hawkmoths but not swallowtail butterflies

The color and patterns of animal‐pollinated flowers are known to have effects on pollinator attraction. In this study, the relative importance of flower color and color contrast patterns on pollinator attraction was examined in two pollinator groups, swallowtail butterflies and hawkmoths using two Hemerocallis species; butterfly‐pollinated H. fulva and hawkmoth‐pollinated H. citrina, having reddish and yellowish flowers in human vision, respectively. Flowers of both species have UV bullseye patterns, composed of UV‐absorbing centers and UV‐reflecting peripheries, known to function as a typical nectar guide, but UV reflectance was significantly more intense in the peripheries of H. citrina flowers than in those of H. fulva flowers. Comparison based on the visual systems of butterflies and hawkmoths showed that the color contrast of the bullseye pattern in H. citrina was more intense than that in H. fulva. To evaluate the relative importance of flower color and the color contrast of bullseye pattern on pollinator attraction, we performed a series of observations using experimental arrays consisting of Hemerocallis species and their hybrids. As a result, swallowtail butterflies and crepuscular/nocturnal hawkmoths showed contrasting preferences for flower color and patterns: butterflies preferred H. fulva‐like colored flower whereas the preference of hawkmoths was affected by the color contrast of the bullseye pattern rather than flower color. Both crepuscular and nocturnal hawkmoths consistently preferred flowers with stronger contrast of the UV bullseye pattern, whereas the preference of hawkmoths for flower color was incoherent. Our finding suggests that hawkmoths can use UV‐absorbing/reflecting bullseye patterns for foraging under light‐limited environments and that the intensified bullseye contrast of H. citrina evolved as an adaptation to hawkmoths. Our results also showed the difference of visual systems between pollinators, which may have promoted floral divergence.     

Priority of color over scent during flower visitation by adult <Emphasis Type="Italic">Vanessa indica</Emphasis> butterflies

Most flower visitors innately prefer a particular color and scent, and use them as cues for flower recognition and selection. However, in most cases, since color and scent serve as a combined signal, not only does the preference for an individual cue, but also the preference hierarchy among different cues, influence their flower visitation. In the present study, we attempted to reveal (1) the chromatic and (2) the olfactory cues that stimulate flower visiting, and (3) the preference hierarchy between these cues, using the naïve adult butterfly Vanessa indica. When we offered 12 different-colored (six chromatic and six achromatic) paper flower models, V. indica showed a color preference for yellow and blue. When we examined the proboscis extension reflex (PER) of V. indica towards 16 individual compounds identified in the floral scents from two nectar plants belonging to the family Compositae, Taraxacum officinale and Cirsium japonicum, six compounds were found to have relatively high PER-eliciting activities, including benzaldehyde, acetophenone, and (E+Z)-nerolidol. When we combined color and scent cues in two-choice bioassays, where butterflies were offered flower models that were purple (a relatively unattractive color), the models scented with these active compounds were significantly more attractive than the odorless controls. In addition, synthetic blends mimicking the floral scents of T. officinale and C. japonicum (at doses equivalent to that of ten flowers) enhanced the number of visits to the scented models. However, the effect of odorizing was not conspicuous in parallel bioassays when yellow flower models were used, and the butterflies also significantly preferred odorless yellow models to scented purple models. These results demonstrate that V. indica depends primarily on color and secondarily on scent during flower visitation.     

Effect of bright wing color of males on other males in Favonius taxila (Lepidoptera: Lycaenidae) with sexual dimorphism in wing color

To examine the effect of wing colors of Favonius taxila males on other males, an intact bright butterfly model or a dull butterfly model whose wing scales had been dislodged was placed in a potential territory, and the behavior of invading males was recorded. The invader showed decreased probability of landing there with a bright model in place compared with a dull model, and exhibited complex flights with frequent turning for a bright model, in contrast to simple flights with relatively linear paths for a dull model. These results support the idea of intrasexual selection in which bright colors of males' wings have evolved through male–male interactions.     

Simultaneous brightness contrast of foraging Papilio butterflies

This study focuses on the sense of brightness in the foraging Japanese yellow swallowtail butterfly, Papilio xuthus. We presented two red discs of different intensity on a grey background to butterflies, and trained them to select one of the discs. They were successfully trained to select either a high intensity or a low intensity disc. The trained butterflies were tested on their ability to perceive brightness in two different protocols: (i) two orange discs of different intensity presented on the same intensity grey background and (ii) two orange discs of the same intensity separately presented on a grey background that was either higher or lower in intensity than the training background. The butterflies trained to high intensity red selected the orange disc of high intensity in protocol 1, and the disc on the background of low intensity grey in protocol 2. We obtained similar results in another set of experiments with purple discs instead of orange discs. The choices of the butterflies trained to low intensity red were opposite to those just described. Taken together, we conclude that Papilio has the ability to learn brightness and darkness of targets independent of colour, and that they have the so-called simultaneous brightness contrast.     

Relationships between the age of northern Kantou plain (central Japan) coppice woods used for production of Japanese forest mushroom logs and butterfly assemblage structure

To discern mechanisms maintaining the diversity of grassland and forest butterflies in coppice woods managed for the production of Japanese forest mushroom logs, we investigated the butterfly fauna in cut-over land tracts shortly after felling and 5 year later, and in forest stands 10, 15, and 25 year after felling (here, we use the term “forests” when referring to the chronosequence of these treed stands). Butterfly species richness and diversity (H′) and the densities of individuals were highest in cut-over lands 5 year after clear-cutting, followed by 25-year-old forest stands. In forests, the richness and densities of forest butterfly species were higher than were those of grassland species. Among forest stands of different ages, forest butterfly species’ richness and the densities of individuals were highest in 25-year-old woods nearing felling time. Some forest butterfly species were observed only in forests. The species richness and densities of grassland butterflies were much higher in cut-over lands 0 and 5 year post felling than in forests; grassland species were rarely found in stands ≥10 year old. Thus, cut-over lands seem to function as temporary habitats for grassland species. Furthermore, the number of forest butterfly species was the same in cut-over lands 5 year after felling and in 25-year-old forest stands; the densities of forest butterfly species was higher in these cut-over lands than in the forest stands. Forest butterfly species living on cut-over land 5 year post felling sipped flower nectar, laid eggs on host plants, and practiced territorial behaviour involved in mate finding. Hence, these cut-over lands functioned as important habitats for various developmental stages of forest butterflies. In conclusion, traditional coppicing in woods for production of Japanese forest mushroom logs is very important for the maintenance of diversity in grassland and forest butterfly species.     

Genetic differences and phenotypic plasticity as causes of variation in oviposition preference in Battus philenor

Summary Bradshaw (1965) proposed that phenotypic plasticity would be more common than adaptive genetic variability in species for which environmental fluctuations occur over periods roughly equal to that species' generation time. In an effort to examine this notion, sources of seasonal variation in two components of oviposition behavior in an east Texas population of pipevine swallowtail butterflies (Battus philenor) were investigated under natural and seminatural conditions. Variability in a visually-based prealighting component involving orientation to leaf shape was primarily due to phenotypic plasticity in the form of adult learning; no seasonally-based genotypic differences in leaf-shape discrimination behavior were observed. By contrast, a chemotactile post-alighting component involving elicitation of oviposition after landing on the host plant was not phenotypically plastic, i.e., not susceptible to learning. In addition, only slight and nonsignificant seasonally-based differences in post-alighting responses to different host species were observed.     

Colour saturation triggers innate reactions to flower signals: Flower dummy experiments with bumblebees

Summary Innate behavioural reactions, i.e. reactions of untrained, flower-naive bumblebees (Bombus terrestris L., B. lucorum L.; Apidae) were observed in flower dummy experiments. It was proven that an innate releasing mechanism responds to optical flower signals: the spectral purity of corolla colour was found to be crucial for far attraction toward flower dummies. During the subsequent near orientation, that is when a bumblebee finally reaches a flower dummy, the bumblebee's antennae contact the part of highest spectral purity while the bee is still in flight. Guides such as stamen patches present in the center of flower dummies are used only for near orientation. Flower dummies receiving the greatest number of antennae reactions at the guide were always those with low spectral purity in the surrounding background colour, high spectral purity at the corolla colour and highest spectral purity at the guide colour. In contrast, dominant wavelength and intensity of flower dummy colours had no detectable influence on innate behavioural reactions, while colour contrast had some. These results are interpreted as follows: orientation toward guides is based upon a gradient of centripetally increasing, bee-subjective colour saturation which directs the bumblebee's approach toward the center of the flower dummy where additional factors may contribute to stimulating the landing reaction.     

Can flower constancy in nectaring butterflies be explained by Darwin’s interference hypothesis?

 When foraging for nectar many insects exhibit flower constancy (a preference for flower species which they have previously visited) and frequently ignore rewarding flowers of other species. Darwin proposed the favoured explanation for this behaviour, hypothesizing that learning of handling skills for one flower species interferes with the ability to recall handling skills for previously learned species. A crucial element of this hypothesis is that savings in handling time resulting from constancy must exceed increases in travelling time necessitated by ignoring other suitable species. A convincing quantification of this trade-off has not been achieved and tests to date on bumblebees indicate that savings in handling time are too small to offset an increase in travelling time. To assess further the validity of Darwin’s hypothesis, handling and flight times of the butterfly, Thymelicus flavus, were measured under natural conditions, and the abundance and reward provided by the available flower species quantified to enable estimation of foraging efficiency. Butterflies exhibited a mean increase in handling time of 0.85 s per flower associated with switching between flower species, although the magnitude of this difference varied greatly among flower species. Switching was not associated with a decrease in travelling time, contrary to expectation. Switching was more frequent following a lower than average reward from the last flower visited. In butterflies, flights serve functions other than movement between nectar sources, such as mate location (unlike worker bees). Hence constancy may be a viable strategy to reduce time spent in handling flowers and increase time available for other activities. Although savings in handling time may be small, Darwin’s interference hypothesis remains a valid explanation for flower constancy in foraging butterflies. Received: 27 January 1997 / Accepted: 5 June 1997     

Thermoregulation and flying habits of the Japanese sulfur butterfly Colias erate (Lepidoptera: Pieridae) in an open habitat

This study evaluated the impact of the thermal environment on the flying behavior of male Japanese sulfur butterflies Colias erate searching for females in an open habitat. Thoracic temperature was monitored before and after flight. Mean thoracic temperature of butterflies immediately after landing was consistently higher than, but independent of, ambient temperature. Although ground speed of flying butterflies was different between flight types, air speed against the butterfly was similar across flight types. The excess of thoracic over ambient temperature was lower in flying butterflies than in basking ones, as predicted by a model. This difference appeared to be due to air current, which enhanced heat loss. In a laboratory study, newly eclosed male butterflies were placed under an incandescent lamp to measure their thoracic temperature at different air current speeds. The excess of thoracic over ambient temperature decreased as the speed of air currents increased. When the air current was similar to the air speed against flying butterflies in the field, a substantial decrease occurred in the operative thoracic temperature.     

Butterflies and bumblebees in greenways and sown wildflower strips in southern Sweden

Greenways have recently been established in some intensively farmed areas of South Sweden in order to enhance recreation opportunities and biodiversity, but the effects of these green structures on biodiversity have not yet been determined. In this study, greenways and experimental sown wildflower strips were investigated for butterfly and bumblebee diversity. In total, 1,769 butterflies of 18 species and 1,216 foraging bumblebees of eight species were recorded. Sown wildflower strips proved to support much higher abundances and species numbers of butterflies and bumblebees than greenways, with 86% of all butterflies and 83% of all bumblebees being observed in the sown flower strips. However, in both types of green structure mostly common species were found. Counts of flower visits showed that Knautia, Centaurea and Cirsium were the most commonly visited plant species. The greenways studied did not seem to fulfil their function of enhancing biodiversity—at least not for butterflies and bumblebees. However, these greenways could easily be improved for common bumblebee and butterfly species by sowing wildflower strips along their margins.     

Effects of elevated CO2 on flowering phenology and nectar production of nectar plants important for butterflies of calcareous grasslands

Effects of elevated CO₂ on flowering phenology and nectar production were investigated in Trifolium pratense, Lotus corniculatus, Scabiosa columbaria, Centaurea jacea and Betonica officinalis, which are all important nectar plants for butterflies. In glasshouse experiments, juvenile plants were exposed to ambient (350 μl l⁻¹) and elevated (660 μl l⁻¹) CO₂ concentrations for 60–80 days. Elevated CO₂ significantly enhanced the development of flower buds in C. jacea. B. officinalis flowered earlier and L. corniculatus produced more flowers under elevated CO₂. In contrast, the number of flowers decreased in T. pratense. The amount of nectar per flower was not affected by elevated CO₂ in the tested legumes (T. pratense and L. corniculatus), but was significantly reduced (!) in the other forbs. Elevated CO₂ did not significantly affect nectar sugar concentration and composition. However, S. columbaria and C. jacea produced significantly less total sugar under elevated CO₂. The nectar amino acid concentration remained unaffected in all investigated plant species, whereas the total of amino acids produced per flower was reduced in all non-legumes. In addition, the amino acid composition changed significantly in all investigated species except for C. jacea. The observed effects are unexpected and are a potential threat to flower visitors such as most butterflies which have no alternative food resources to nectar. Changes in nectar production due to elevated CO₂ could also have generally detrimental effects on the interactions of flowers and their pollinators. Received: 12 September 1996 / Accepted: 9 September 1997     

Do foraging bumblebees scent-mark food sources and does it matter?

Summary The foraging of worker bees of Bombus terrestris visiting artificial feeders in a climatic test chamber was investigated. The behaviour of worker bees visiting rewarding and unrewarding feeders is completely different. Of all flower visits to rewarding feeders 94% are probing-visits, i.e. the bees land on the flower and probe for nectar. In contrast, only 0.3% of all visits to unrewarding feeders are probing-visits, whereas 47% are approach-visits, i.e., the bees approach the feeders without landing. Exchanging feeder discs proves that the signal used for discrimination must be associated with the plastic disc used as landing platform. Most probably it involves scent-marking of the rewarding feeders with components of high and low volatility. The mean foraging efficiency of bees in a scent-marked foraging arena is 5.7 mg sugar/min and drops to 2.8 mg sugar/min after the scent marked discs are replaced by clean ones. Three components generate this drop in foraging efficiency: (1) the between-flower flight time increases, i.e. the bees search for a longer time before landing on flowers, (2) the bees no longer discriminate between rewarding and unrewarding feeders, and (3) the bees probe empty feeders longer than necessary; obviously they expect to find nectar.     

Butterflies show flower colour preferences but not constancy in foraging at four plant species

1. The extent to which flower colour and other visual cues influence butterfly flower choice in the field is poorly understood, especially in comparison with choices by Hymenoptera. 2. Using a novel approach to studies of visitation behaviour by butterflies, flower colour of four Asteraceae species was phenotypically manipulated to decouple the influence of that trait from others (including morphology and nectar rewards) on visitation by Lycaena heteronea, Speyeria mormonia, Cercyonis oetus, and Phyciodes campestris. 3. Flower visits were recorded to experimental flower arrays in subalpine meadows to measure (i) spontaneous preference by butterflies for particular colours and other traits and (ii) flower constancy (longer than expected strings of visits made to flowers of the same species), a behaviour that can reduce interspecific gene flow in plants. 4. Over three field seasons, 3558 individual flower visits in 1386 foraging bouts were observed for free‐flying butterflies. All four butterfly species responded to the phenotypic manipulations of flower colour, although in different ways. Speyeria mormonia and L. heteronea also exhibited preferences based on other flower traits. Lycaena heteronea responded to combinations of traits such that the other traits it preferred depended upon the context of flower colour. 5. None of the butterfly species exhibited flower constancy in any of the arrays employed. 6. The observed preferences show that butterflies, like some other pollinators, are potentially capable of exerting selection on colour and other floral traits. Moreover, these flower preferences can depend on the context of other flower traits. The absence of constancy contrasts with reports of high constancy in many bees.     

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.

     

INTERPOPULATION DIFFERENCES IN HOST PREFERENCE AND THE EVOLUTION OF LEARNING IN THE BUTTERFLY,BATTUS PHILENOR

The host-discrimination behavior of the adult female pipevine swallowtail butterfly (Battus philenor) was investigated for two populations, an east Texas population that uses two host species with different leaf shapes and a Virginia montane population that uses one host species with a single leaf shape. While Texas and Virginia females exhibited similar chemotactile responses after landing on various host species, butterflies from each population landed more frequently on certain host species used by that particular population. Despite this difference in searching behavior, Texas and Virginia populations were equally capable of learning to search for the leaf shape of a particular host species in artificial enclosure arrays. In addition, learning of leaf-shape preference was confounded equally when each population was introduced into arrays containing all the host species found in both populations. The lack of differentiation in learning of leaf-shape preference between populations that differ in host breadth and host preference argues against the specific hypothesis that learning of leaf-shape preference represents a local adaptation that permits foraging butterflies in the east Texas population to respond to seasonal changes in the relative quality or abundance of host species that differ in leaf shape. Three hypotheses are proposed to explain the apparent absence of interpopulation genetic variation in learning traits. One hypothesis supposes that learning of leaf-shape discrimination is mediated by the same physiological mechanisms that permit females to learn to discriminate among suitable and unsuitable conspecific plants. Selection for restriction of learning of leaf-shape preference in the Virginia montane population may therefore be constrained by selection for learning of other types of discrimination behavior.     

High specialisation in the pollination system of Mandevilla tenuifolia (J.C. Mikan) Woodson (Apocynaceae) drives the effectiveness of butterflies as pollinators

Butterfly pollination in the tropics is considered somewhat effective or solely effective in a few plant species. In the present study, we tested the hypothesis that Mandevilla tenuifolia (Apocynaceae), which has floral attributes associated with psychophily, has strategies adapted to pollination by butterflies, restricting other floral visitors and making these insects act as efficient pollinators. We analysed the floral and reproductive biology of M. tenuifolia, as well as the frequency and efficiency of its flower visitors. M. tenuifolia is an herb whose flowers have strong herkogamy and secondary pollen presentation on the style head, which corresponds to 60.4% of pollen on the anthers. Flower longevity and the long period of receptivity of the stigmatic region associated with the large amount of pollen removed in the first visits suggest that flowers remain functionally female during part of anthesis. Butterflies, mainly of the families Nymphalidae and Pieridae, are the only pollinators of M. tenuifolia. Despite being self‐compatible, M. tenuifolia depends on biotic vectors for fruit production. A non‐significant difference in fruit set between controlled treatments and natural conditions suggests that the pollinators are efficient. The inclination resulting from the landing of butterflies on flowers, together with flower morphology, guiding the insect proboscis inside the floral tube, as well as the frequency and efficiency of butterfly visits, are evidence of the close relationship between butterflies and M. tenuifolia, and also of the efficiency of these insects as pollinators.     

The influence of ultraviolet reflectance differs between conspicuous aposematic signals in neotropical butterflies and poison frogs

Warning signals are often characterized by highly contrasting, distinctive, and memorable colors. Greater chromatic (hue) and achromatic (brightness) contrast have both been found to contribute to greater signal efficacy, making longwave colored signals (e.g., red and yellow), that are perceived by both chromatic and achromatic visual pathways, particularly common. Conversely, shortwave colors (e.g., blue and ultraviolet) do not contribute to luminance perception yet are also commonly found in warning signals. Our understanding of the role of UV in aposematic signals is currently incomplete as UV perception is not universal, and evidence for its utility is at best mixed. We used visual modeling to quantify how UV affects signal contrast in aposematic heliconiian butterflies and poison frogs both of which reflect UV wavelengths, occupy similar habitats, and share similar classes of predators. Previous work on butterflies has found that UV reflectance does not affect predation risk but is involved in mate choice. As the butterflies, but not the frogs, have UV‐sensitive vision, the function of UV reflectance in poison frogs is currently unknown. We found that despite showing up strongly in UV photographs, UV reflectance only appreciably affected visual contrast in the butterflies. As such, these results support the notion that although UV reflectance is associated with intraspecific communication in butterflies, it appears to be nonfunctional in frogs. Consequently, our data highlight that we should be careful when assigning a selection‐based benefit to the presence of UV reflectance.