Associative Learning of Color by Males of the Parasitoid Wasp Nasonia vitripennis (Hymenoptera: Pteromalidae)

Males of the parasitoid wasp Nasonia vitripennis showed no innate preference for blue versus yellow or for green versus brown. They learned to associate color with mates, but their ability to do so depended on the color used and the strength of the reward. Specifically, males learned to associate brown or green with a reward of many virgin females. With fewer females, fewer training periods, or mated females as the reward, males still learned a preference for green but not for brown. Males did not learn to associate color with rewards of honey or water. Previous studies of color preference and associative learning in parasitoid wasps have focused almost entirely on females. This is the first demonstration of associative learning in response to visual cues by male parasitoid wasps.     

Female Canaries (Serinus canaria) Associate more with Males that Contrast Strongly against the Background

Male birds with bright plumage colors which entail production costs and increased predation risks should be able to emphasize the handicap function of their ornaments when they present themselves against a contrasting background. Contrast with the background may accentuate the signal's costliness and make the male more conspicuous to predators. Both effects should contribute to increase male attractiveness. It is therefore conceivable that the extent to which the male plumage contrasts against the background modifies female choice behavior as it improves the discrimination of mates. We tested this hypothesis in domesticated canaries (Serinus canaria). In the first experiment, yellow females could choose between two yellow males presented in front of a yellow and a white background, respectively. In the second experiment we replaced the yellow males with white ones. In experiment 1 females associated significantly more with yellow males which contrasted strongly against the white background. In experiment 2 there was at least a trend for preferred associations with the white male in front of the yellow background. We found no support that male properties per se were chosen. We could further clarify that females associated with the contrasting male and not with a particular background color. Thus, our study demonstrates that not only inherent properties of the sender but also the interaction of bird color with the signaling environment may influence mate choice.     

Visual response of the tomato fruit fly, Neoceratitis cyanescens, to colored fruit models

To determine how mature females of the tomato fruit fly, Neoceratitis cyanescens (Bezzi) (Diptera: Tephritidae), detect host fruit after arriving in their host plant habitat, behavioral responses to colored models were observed in a laboratory flight chamber. Host‐seeking females oriented themselves preferentially towards bright orange spheres (3.7 cm in diameter), irrespective of their natal host fruit: tomato, bug weed, or black nightshade. Females oriented themselves preferentially towards the orange sphere when placed against a fluorescent yellow background as opposed to a black background, but the distribution of responses to the set of colored spheres did not vary significantly with background color. In a choice situation between bright orange spheres of various sizes (1.9, 3.7, and 7.5 cm in diameter), females landed preferentially on the bigger sphere. However, they preferred a yellow color when the latter was associated with two‐dimensional models, probably mimicking leaves. The attractiveness of orange spheres depended more on the proportion of reflected light in the spectral region around 610 nm than brightness of color in itself. Low light intensity significantly influenced the activity of the flies but not their visual preference. The strong response of females to bright orange spheres confirmed the importance of visual characteristics in short‐range mechanisms of host‐plant location in specialist insects. Responses to fruit visual stimuli are discussed relative to other Tephritidae, host‐finding strategy, and pest management.     

Yellow–green color polymorphism in males of a pea aphid clone and its genetic pattern

Although most aphid species living on leaves have a green body color, little is known regarding the biosynthetic pathways of green pigments. We found that a clone of the pea aphid, Acyrthosiphon pisum (Harris) produced both green- and yellow-colored males. The females of this clone were green in color, while 8.4% of the males produced were yellow. To date, yellow body color has been reported only in a single mutant clone in A. pisum. To explore the genetic pattern of yellow body color, green or yellow males were mated with green females of the same clone. The hatchability of the eggs sired by yellow males (26.2%) was less than half that of the eggs sired by green males (79.0%). The hatched foundresses of both groups were all green, with no yellow foundresses. Because aphids have an XX-XO sex determination system, color polymorphism in males suggests that male body color may be governed by an X-linked locus. If females possess heterozygosity at the putative locus, they can produce alternative phenotypes in males. The small proportion of yellow males and absence of yellow foundresses imply that the allele responsible for yellow body color has a deleterious effect. The present study suggests that this clone could be used to elucidate the biosynthetic pathways and underlying genetics of green pigments in aphids.     

The scent of food and defence: green leaf volatiles and toluquinone as sex attractant mediate mate finding in the European cockchafer Melolontha melolontha

Mate finding in the European cockchafer, Melolontha melolontha , has been investigated in the field. Our data suggest that mainly males show flight activity at dusk. In a landing cage bioassay, male cockchafers preferred cages baited with females over cages baited with males. Gas chromatographic analysis of beetle extracts with electronantennographic detection revealed the presence of electrophysiologically active compounds, among them toluquinone. In funnel trap bioassays none of these compounds was attractive towards males per se . A mixture of green leaf volatiles (GLV) mimicking the bouquet of mechanically damaged leaves was highly attractive to M. melolontha males. The attractiveness of the same GLV mixture was synergistically enhanced when toluquinone was added to the lure. GLV attract males to damaged leaves. Toluquinone as a sex pheromone indicates the presence of conspecific females and synergistically enhances the attractiveness of GLV.     

Delayed plumage maturation in green‐backed flycatchers (Ficedula elisae): An evidence of female mimicry

Delayed plumage maturation (DPM) is the delayed acquisition of an adult color and pattern of plumage until after the first potential breeding period. Among the hypotheses proposed to explain DPM, the female mimicry hypothesis (FMH) has received considerable attention. FMH predicts that after‐second‐calendar‐year (ASY) males should attack ASY males more than second‐calendar‐year (SY) males and females, while no difference between the two latter. Few studies have been thought as support for FMH, while in fact most of them give no conclusive evidence to the assumption of FMH that ASY males are unable to distinguish SY males from females. Thus, other support besides behavioral experiments, such as it is physiologically impossible to discriminate SY males and females from avian perspective, is important in testing FMH. We studied color differences in six plumage patches between ASY male, SY male and female by analyzing reflectance spectra in the avian visual system, and tested the prediction of FMH by conducting intrusion experiments on territorial males with three kinds of conspecifics (i.e., ASY males, SY males and females) as territory “intruders” in the green‐backed flycatcher (Ficedula elisae). It is physiologically impossible for ASY males to distinguish SY males and females through the plumage color of crown, mantle, rump, and throat patches, moreover difficult to distinguish through breast and belly patches even under reasonable viewing conditions. ASY males were more aggressive toward ASY males than SY males and females with no differences between the two latter. SY males showed a slightly but not significantly higher attack intensity on ASY males than SY males and females. Our results suggest that female mimicry is more likely to be the explanation for DPM in the green‐backed flycatcher. To our knowledge, this is the first study combining avian visual system and behavioral experiments in testing hypotheses of DPM.     

Responses to computer-generated visual stimuli by the male praying mantis, Sphodromantis lineola (Burmeister)

We tested male mantis' responses to square, computer-generated visual stimuli of various sizes to determine the stimulus parameters that affect their striking behaviour. Males in a high-hunger group displayed a sharp preference for solid black 12×12-degree squares moving linearly against a white background, especially when the squares moved downward (versus horizontally). Males in a low-hunger group visually tracked but, with rare exception, did not strike at any linearly moving squares. In contrast, when solid black squares moved erratically around visual field centre, males struck regularly at them irrespective of hunger level, even when the squares were as large as 47×47 degrees. Males also recognized black-and-white Julesz-patterned square stimuli moving against a similarly patterned background, indicating that they recognized the synchronous movement of a group of stimulus elements as a single moving object. Finally, we compared allometric and life history data between male and females S. lineola to elucidate the possible reasons for the differences between male response patterns and previously published data on females. These comparisons suggest that males employ a different behavioural strategy than do females when faced with large, erratically moving visual stimuli.     

Sexual Selection of Zorion guttigerum Westwood (Coleoptera: Cerambycidae: Cerambycinae) in Relation to Body Size and Color

Zorion guttigerum is a flower-visiting longhorned beetle endemic to New Zealand. Sexual selection of this species in relation to the body size and color form of different sexes was investigated in the field. The population sex ratio, based on censuses of feeding and mating sites (flowers), is male-biased. Females are significantly larger than males. Both sexes have antennae of similar length but the antennal length relative to the elytral length is greater in males than in females, and the antennal length of males increases more with an increase in body size than that of females. Both sexes have dark blue (DB) and yellowish-brown (YB) individuals. Both pair-bonded and solitary males are similar in elytral and antennal length. In pair-bonded males, DB individuals are significantly more numerous than YB ones, but in solitary males, the number of both color forms is similar. Males tend to have territory protection behavior, fighting with and chasing away rival males from feeding and mating sites. Larger males usually win the fight but the size-dependent fighting advantage does not translate into mating success. Male color plays an important role in mating success, with DB males having a significantly better chance to mate than YB males. Furthermore, male body size and color also have interactions in mating success: males of DB color morph obtain a greater mating advantage according to body size. Pair-bonded females are significantly larger and have longer antennae than solitary females, suggesting that males prefer larger females for mating. In addition, females of DB color morph with longer antennae are also preferred by males for mating. The significance of these findings is discussed.     

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.     

Sexual Selection in Black Morph Girardinus metallicus (Pisces: Poeciliidae): Females Can Spot a Winner (But We Cannot)

Male–male competition may interfere with the ability of females to choose mates by interrupting courtship or by favoring highly aggressive males who may damage females during mating attempts. Alternatively, females may benefit by mating with dominant males, and female choice and male–male competition may therefore act in unison. The same traits, including aggressiveness, may indicate male quality to females and to rivals. We investigated sexual selection in the black morph of the endemic Cuban poeciliid fish, Girardinus metallicus, to ascertain the links between morphological and behavioral traits and success in intra‐ and intersexual selection. Males conspicuously exhibit their black ventral surface and gonopodium to females during courtship. Dichotomous choice tests revealed female association preferences for certain males, and those same males were more successful in monopolizing access to females when the fish were allowed to directly interact. Dominant males followed, courted, and copulated with females more than subordinate males within a pair, and it appears that females could either assess dominance based on cues we did not measure, or could influence subsequent mating success by their behavior during the dichotomous choice trials. There was an interaction between black status (i.e., whether the male in each pair had more or less ventral black coloration than the other male in that pair) and dominance, such that low‐black dominant males courted early and then shifted to following females, whereas high‐black dominant males courted far more later in the observation period. These results hint at the importance for sexual selection of the interplay between a static morphological trait (black coloration) and a dynamic behavioral trait (aggressiveness), but the functional significance of the courtship display remains a mystery.     

Risky Courtship: Background Contrast,Ornamentation, and Display Behavior of Wolf Spiders Affect Visual Detection by Toad Predators

Males that search widely for females and perform conspicuous courtship displays run a high risk of being detected by their predators. Therefore, gains in reproductive success might be offset by increased mortality due to predation. Male brush‐legged wolf spiders (Schizocosa ocreata) with larger decorative traits (foreleg tufts) are preferred by females as mates, but are more readily detected by predators. However, predation risk may also be influenced by the interaction between components of signals and the environment in which signaling occurs. Courting male spiders were readily accepted as prey by a sympatric predator, the American toad (Anaxyrus americanus). We used video playback to tease apart the interactive effect between visual signals and the signaling environment on the ability of toads to detect courting spiders as a function of distance, background contrast, the presence or absence of male foreleg tufts, and behavioral activity. The response of toads to video sequences of male spiders was similar to their response to live male spiders. Toad response varied over distance toward spiders displayed against high contrast (sunny) vs. low contrast (shaded) backgrounds. Beyond 30 cm, more toads detected courting male spiders against light, ‘sunny’ backgrounds and detected them faster when compared to the same spider stimulus against darker, ‘shady’ backgrounds. In choice tests, toads oriented more often toward courting males with leg tufts than those without. Toad responses also varied with male spider behavior in that only videos of moving males were attacked. Latency to orient and detection by toads was significantly greater for walking males than courting males, and this effect was most evident at distances between 30 cm and 50 cm. Results supported that courting wolf spiders are at significant risk of predation by visually acute predators. Distance, background contrast, and the presence of foreleg decorations influence detection probability. Thus, the same complex visual signals that make males conspicuous and are preferred by females can make males more vulnerable as prey to toads.     

Relevance of visual and olfactory cues for host location in the mustard leaf beetle Phaedon cochleariae

The relevance of visual and olfactory cues for host‐plant location is investigated in males and females of the oligophagous mustard leaf beetle Phaedon cochleariae Fabricius (Coleoptera: Chrysomelidae). Different objects are offered in a walking arena and the behaviour of beetles is observed. Beetles orient toward vertically or horizontally striped black and white pattern independent of stripe orientation. The results suggest that contrast facilitates orientation in the field, whereas the pattern itself may be less important for host location in dense vegetation. The response to green and yellow objects is tested to investigate discrimination abilities between young (green) and mature (yellow) leaves. Beetles prefer green over yellow independent of material (cardboard or leaves of Nasturtium officinale R. Br., Brassicaceae). Preference behaviour tested in a dual‐choice contact assay coincides with visual preferences, where adults prefer young, more nutritious leaves for feeding and oviposition. Furthermore, females discriminate between visual cues of green leaves and green cardboard, whereas males do not, indicating that females are more sensitive in colour discrimination. Differences in colour wavelength influence the choice of beetle behaviour more strongly than differences in intensity. Both sexes of P. cochleariae prefer volatiles of the host plant N. officinale, whereas only females respond to the main volatile compound 2‐phenylethyl isothiocyanate. Given a choice between visual and olfactory cues, males orientate towards the colour cues, whereas females do not show any preferences. In males, visual cues may thus override olfactory cues, whereas, in females, both are equally important, which may reflect different ecological requirements and/or physiological abilities.     

Factors influencing the landing of maleEpiphyas postvittana (Walker) exhibiting pheromone-mediated flight (Lepidoptera: Tortricidae)

The effects of changes in various visual and olfactory properties of a white card surface on the landing position of male Epiphyas postvittanaexhibiting pheromone-mediated flight were studied in a wind tunnel. Males landed predominantly at the most downwind position of a surface in line with the pheromone source, regardless of the strength of the source. The position on the surface that males landed was strongly influenced by visual factors. The landing position of males appeared to be influenced by visual cues along all three axes of the surface. Decreases in either the dimension horizontally perpendicular to the wind direction or the vertical dimension resulted in greater numbers of males landing farther upwind on the surface than the downwind edge. Visual changes in the axis along the wind direction also affected the position at which males landed. For example, when presented with two white card surfaces with a 4- cm gap between them, males tended to land on the downwind edge of the upwind surface (on which the source was located). When the gap was bridged with clear Mylar, the landing pattern was significantly different, with the greater proportion of males landing on the downwind surface. However, when Mylar was placed on the plexiglass floor of the tunnel (in addition to bridging the gap), the landing pattern on the surface was not significantly different from that on the two surfaces without the Mylar bridge. It is suggested that during the prelanding and landing phases of pheromone-mediated flight, male moths orient to visual features of the surface containing the pheromone source rather than to visual features of the source (conspecific female moth) itself.     

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.     

Landing Surface Color Preferences of Spathius agrili (Hymenoptera: Braconidae), a Parasitoid of Emerald Ash Borer, Agrilus planipennis (Coleoptera: Buprestidae)

The color preferences for landing surfaces were examined for Spathius agrili (Hymenoptera: Braconidae), a parasitic wasp introduced for biocontrol of emerald ash borer, Agrilus planipennis (Coleoptera: Buprestidae). Lures with the 3-component pheromone blend of male S. agrili were used to activate upwind flight by virgin female S. agrili in a laminar flow wind tunnel. Paper discs with halves of two different colors (combination pairs of black, white, red, yellow, green, or purple), with the pheromone lure in the center, were tested to quantify preferences for landing on one color over another. Females landed preferentially on green, yellow, and white surfaces, and landed the least frequently on red, black, and purple surfaces. Changes in color preferences due to adjacent colors were observed and discussed.     

Indirect effects of phytochemicals on offspring performance of Queensland fruit fly,Bactrocera tryoni (Diptera: Tephritidae)

Phytochemical lures such as methyl eugenol (ME) and cue‐lure are used in the management of Bactrocera fruit flies for monitoring and control. These lures are not just attractants, but also trigger physiological changes in males that lead to enhanced mating success. Additionally, in the cue‐lure‐responsive Bactrocera tryoni, females mated with lure‐fed males exhibit changes in fecundity, remating receptivity and longevity. While the lures show current generation effects, no research has been carried out on possible multigenerational effects, although such effects have been hypothesized within a ‘sexy‐son’ sexual selection model. In this study, we test for indirect, cross‐generational effects of lure exposure in F1offspring of B. tryoni females mated with cue‐lure‐fed, zingerone‐fed and lure‐unfed (=control) males. The F1 attributes we recorded were immature development time, immature survival, adult survival and adult male lure foraging. No significant differences were found between treatments for any of the three life‐history measurements, except that the offspring sired by zingerone‐fed males had a longer egg development time than cue‐lure and control offspring. However, indirect exposure to lures significantly enhanced the lure‐foraging ability of F1 adult males. More offspring of cue‐lure‐fed males arrived at a lure source in both large flight cages and small laboratory cages over a 2‐h period than did control males. The offspring of zingerone‐fed males were generally intermediate between cue‐lure and control offspring. This study provides the first evidence of a next generation effect of fruit fly male lures. While the results of this study support a ‘sexy‐son’ sexual selection mechanism for the evolution of lure response in Bactrocera fruit flies, our discussion urges caution in interpreting our results in this way.     

Landing response of Aedes (Stegomyia) polynesiensis mosquitoes to coloured targets

Aedes polynesiensis Marks (Diptera: Culicidae) is the primary vector of lymphatic filariasis (LF) in the island countries and territories of the South Pacific. In the development of a novel control tool, the response of Ae. polynesiensis to six different colours (three solid fabrics, two patterned fabrics and a plastic tarp) was measured using a digital photographic system. Adult mosquitoes were placed into an environmental chamber and allowed to choose between a white target and one of six experimental targets. Mosquito landing frequency and landing duration were calculated. Adult female Ae. polynesiensis preferred all of the experimental targets to the white control target. Mosquito landing frequency was highest for the solid targets (black, navy blue and red) followed in turn by the two colour pattern targets and the polyethylene target. Mosquito landing duration was greater for experimental targets when compared with white control targets. Mosquito landing frequencies did not change over time during the course of the assay. The response of male Ae. polynesiensis was also measured when exposed to a 100% cotton black target. Male mosquitoes preferred the black target to the white control target, although at levels lower than that observed in female mosquitoes. The results suggest that future investigations evaluating the visual responses of Ae. polynesiensis mosquitoes are warranted, with a special emphasis on semi‐field and field‐based experiments.     

Variation in the Visual Habitat May Mediate the Maintenance of Color Polymorphism in a Poeciliid Fish

The conspicuousness of animal signals is influenced by their contrast against the background. As such, signal conspicuousness will tend to vary in nature because habitats are composed of a mosaic of backgrounds. Variation in attractiveness could result in variation in conspecific mate choice and risk of predation, which, in turn, may create opportunities for balancing selection to maintain distinct polymorphisms. We quantified male coloration, the absorbance spectrum of visual pigments and the photic environment of Poecilia parae, a fish species with five distinct male color morphs: a drab (i.e., grey), a striped, and three colorful (i.e., blue, red and yellow) morphs. Then, using physiological models, we assessed how male color patterns can be perceived in their natural visual habitats by conspecific females and a common cichlid predator, Aequidens tetramerus. Our estimates of chromatic and luminance contrasts suggest that the three most colorful morphs were consistently the most conspicuous across all habitats. However, variation in the visual background resulted in variation in which morph was the most conspicuous to females at each locality. Likewise, the most colorful morphs were the most conspicuous morphs to cichlid predators. If females are able to discriminate between conspicuous prospective mates and those preferred males are also more vulnerable to predation, variable visual habitats could influence the direction and strength of natural and sexual selection, thereby allowing for the persistence of color polymorphisms in natural environments.     

Traps baited with pear ester and acetic acid attract both sexes of Hedya nubiferana (Lepidoptera: Tortricidae)

This study was initiated with the objective of studying field responses of the green budworm moth, Hedya nubiferana (Haworth) (Lepidoptera: Tortricidae), to pear ester [PE; ethyl (E,Z)‐2,4‐decadienoate] and acetic acid (AA) with the aim of developing a lure attractive also for females. In the overwhelming majority of tests, traps baited with the PEAA lure (the combination of both PE and AA) caught more than traps baited with either of the constituents presented alone. PEAA lures were attractive to H. nubiferana no matter whether the two compounds were provided in separate dispensers or mixed together in a single one, and a large percentage (up to 71%) of trap catch consisted of females. Traps with PEAA lures caught (females plus males) on an average 30% of the catches in traps baited with the synthetic green budworm moth sex pheromone (only males). This suggested that the new PEAA lure had a trapping performance comparable with that of pheromone traps, which latter are used by farmers today. Consequently, the PEAA lure showed potential for future practical applications as a female‐targeted lure for H. nubiferana. To our knowledge, this is the first well‐documented report on the attraction of PEAA lure for a tortricid species other than codling moth.