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The eyes of insects display an incredible diversity of adaptations to enhance vision across the gamut of light levels that they experience. One commonly studied contrast is the difference in eye structure between nocturnal and diurnal species, with nocturnal species typically having features that enhance eye sensitivity such as larger eyes, larger eye facets, and larger ocelli. In this study, we compared eye structure between workers of closely related nocturnal and diurnal above ground foraging ant species (Hymenoptera: Formicidae) in four genera (Myrmecocystus, Aphaenogaster, Temnothorax, Veromessor). In all four genera, nocturnal species tend to have little cuticular pigment (pale), while diurnal species are heavily pigmented (dark), hence we could use cuticle coloration as a surrogate for activity pattern. Across three genera (Myrmecocystus, Aphaenogaster, Temnothorax), pale species, as expected for nocturnally active animals, had larger eyes, larger facet diameters, and larger visual spans compared to their dark, more day active congeners. This same pattern occurred for one pale species of Veromessor, but not the other. There were no consistent differences between nocturnal and diurnal species in interommatidial angles and eye parameters both within and among genera. Hence, the evolution of eye features that enhance sensitivity in low light levels do not appear to have consistent correlated effects on features related to visual acuity. A survey across several additional ant genera found numerous other pale species with enlarged eyes, suggesting these traits evolved multiple times within and across genera. We also compared the size of the anterior ocellus in workers of pale versus dark species of Myrmecocystus. In species with larger workers, the anterior ocellus was smaller in pale than in dark species, but this difference mostly disappeared for species with smaller workers. Presence of the anterior ocellus also was size-dependent in the two largest pale species. [ABSTRACT FROM AUTHOR]

Eclosion, the molt from pupae to adult stage, occurs at specific times of the day in many holometabolous insects. In this study, we explored the daily pattern of eclosion and the extent to which it varies both within and among six species of nymphalid butterflies. Our broad goal was not only to document the variation in eclosion times, but also to evaluate, through comparative analysis, predictions from several potential adaptive explanations for when during the day eclosions occur. For all species, we reared individuals to pupation, and, at the beginning of the pupal stage, placed the pupae in a greenhouse and recorded their eclosions with a video camera. From these recordings, we obtained for each pupa the time of day at which eclosion occurred. All six species exhibited eclosion distributions that were significantly clustered in the hours around and immediately after sunrise. Only females of the Monarch butterfly showed a non-clustered pattern of eclosion times. There were no sexual differences in eclosion times for any species. The data on interspecific differences in the daily pattern of eclosion do not fit predictions from the known interspecific differences in the daily timing of mating. The data do not fit the prediction that elevated avian predation in the morning will favor eclosion at other times during the day. [ABSTRACT FROM AUTHOR]

Light is a primary environmental factor used by aquatic invertebrates for depth selection behavior. Many branchiopod crustaceans live in ephemeral aquatic habitats. All branchiopod crustaceans studied to date express four or more visual opsins in their compound eyes. We asked whether two branchiopods, Triops longicaudatus, and Streptocephalus mackini, use multiple spectral channels to regulate their position in the water column. At the lowest intensities that elicited photonegative behavior, both species had broad spectral bandwidths, suggesting they use multiple spectral photoreceptor classes. Male Streptocephalus mackini were more likely to maintain a vertical position 8.0-12.0 cm below the surface than females, independently of whether females were present. Male photopositive behavior at low intensity was restricted to narrow bandwidth centered at 532nm, suggesting a single photoreceptor class is used to maintain position above females. We compared ephemeral pools from two regions in Arizona and found that diffuse light attenuation coefficients were two orders of magnitude greater than the most heavily attenuating coastal waters. At less than a meter of depth, pools were often dimmer than terrestrial habitats under starlight. Soil particle size distribution in each region affected spectral light environments, and behavioral responses of field-caught shrimp were adapted to the spectral properties of their region. The results suggest that branchiopods predominantly use luminance vision summed from multiple spectral photoreceptor classes for depth selection in dim, spectrally variable environments. The neuroanatomical basis for summation is described in a companion paper. [ABSTRACT FROM AUTHOR]

Animals that have true color vision possess several spectral classes of photoreceptors. Pancrustaceans (Hexapoda + Crustacea) that integrate spectral information about their reconstructed visual world do so from photoreceptor terminals supplying their second optic neuropils, with subsequent participation of the third (lobula) and deeper centers (optic foci). Here we describe experiments and correlative neural arrangements underlying convergent visual pathways in two species of branchiopod crustaceans that have to cope with a broad range of spectral ambience and illuminance in ephemeral pools, yet possess just two optic neuropils, the lamina and optic tectum. Electroretinographic recordings and multimodel inference based on modeled spectral absorptance were used to identify the most likely number of spectral photoreceptor classes in their compound eyes. Recordings from the retina provide support for four color channels. Neuroanatomical observations resolve arrangements in their laminas that suggest signal summation at low light intensities, incorporating chromatic channels. Neuroanatomical observations demonstrate that spatial summation in the lamina of the two species are mediated by quite different mechanisms, both of which allow signals from several ommatidia to be pooled at single lamina monopolar cells. We propose that such summation provides sufficient signal for vision at intensities equivalent to those experienced by insects in terrestrial habitats under dim starlight. Our findings suggest that despite the absence of optic lobe neuropils necessary for spectral discrimination utilized by true color vision, four spectral photoreceptor classes have been maintained in Branchiopoda for vision at very low light intensities at variable ambient wavelengths that typify conditions in ephemeral fresh water habitats. [ABSTRACT FROM AUTHOR]

In species wherein males display elaborate sexually selected traits or ornaments, conspecific females may also express these traits in full or partial form. This is called mutual ornamentation, and the co-occurrence of such traits in males and females remains poorly understood. In many sulphur butterflies (subfamily Coliadinae), males have a brilliant ultraviolet (UV) iridescence on their dorsal wing surfaces that functions as a sexual signal in courtship. In some of these sulphurs, such as the large grass yellow, Eurema hecabe , females also display dorsal iridescent patches, albeit smaller, restricted to the forewings and less bright than the male's, but the reasons for its occurrence in females are unknown. Here we present a study testing two functional hypotheses for the female UV-reflecting patch: an antiharassment hypothesis and a male mate choice hypothesis. The daily activity pattern of this species suggests that males are most likely to harass or choose among females from midday on. Observations made at this time of day on the characteristics of females related to male courtship duration suggest that males may preferentially court females with a large UV patch. Experiments with colour-manipulated models also suggest that males court with equal intensity females with and without a basal UV patch. Taken altogether the results provide no support for the antiharassment hypothesis. Because support for the male mate choice hypothesis was relatively weak and because of the limited potential for selection on female coloration due to male choice in these butterflies, the nonfunctional sexual correlation hypothesis remains a viable explanation for the female ornament. [ABSTRACT FROM AUTHOR]

Because of the important role sensory systems play in the behaviour of animals, information on sensory capabilities is of great value to behavioural ecologists in the development of hypotheses to explain behaviour. In compound eyes, interommatidial angles are a key determinant of visual acuity but methods for measuring these angles are often demanding and limited to live animals with a pseudopupil. Here we present a new technique for measuring interommatidial angles that is less demanding in terms of technology than other techniques but still accurate. It allows measurements in eyes without a pseudopupil such as dark eyes or even museum specimens. We call this technique the radius of curvature estimation ( RCE) method. We describe RCE and validate the method by comparing results from RCE with those from pseudopupil analysis for the butterfly Asterocampa leilia. As an application of RCE we measure the eyes of the butterfly Battus philenor, a species whose visually guided behaviour is well known but whose eye structure and visual acuity are unknown. We discuss the results of the eye morphology in B. philenor in relation to their behaviour and ecology. We contend that RCE fills a gap in the repertoire of techniques available to study peripheral determinants of spatial resolution in compound eyes, because it can be applied on species with dark eyes. RCE then opens up for sampling a larger number of specimens, which, in combination with being able to use museum specimens, makes it possible to quantitatively test ecologically and evolutionarily driven hypotheses about vision in animals in a new way. [ABSTRACT FROM AUTHOR]

Warning colours can deter predators from attacking unpalatable or toxic prey. These colours are avoided by predators, either innately or by learned response through trial and error. Predators that have learned these signals can also generalize their responses to similar colours. Much of the previous research on warning colours has examined long-wavelength colours (e.g. orange and red) and little is known about short-wavelength coloration (e.g. blue) in warning signals, despite its presence in unpalatable animals in many different taxa. We determined whether prey with only blue coloration, only orange coloration, and with both colours elicited innate avoidance from a naïve predator (Gambel's quail chicks, Callipepla gambelii), as well as how effectively the signals were learned and generalized. Prey were composed of pieces of mealworm (palatable or unpalatable) beneath a coloured paper tent. Blue did not elicit any innate avoidance, whereas orange and a combination of both colours (orange-and-blue) were effective at deterring naïve predators. Predators were able to learn the association between colour and unpalatability most effectively for orange prey, but also learned this association for blue and for orange-and-blue prey. Predators showed asymmetrical generalization, in which birds were more likely to generalize from the colour they learned as unpalatable (conditioned stimulus) to orange prey and orange-and-blue prey than to blue prey. We conclude that orange is likely a more effective warning colour to these birds, but blue can still be learned alone and when displayed adjacent to orange. [ABSTRACT FROM AUTHOR]

Color ornaments are often viewed as products of countervailing sexual and natural selection, because more colorful, more attractive individuals may also be more conspicuous to predators. However, while evidence for such countervailing selection exists for vertebrate color ornaments (e.g., Trinidadian guppies), similar studies have yet to be reported in invertebrates. Indeed, evidence for female mate choice based on extant variation in male coloration is limited in invertebrates, and researchers have not explicitly asked whether more attractive males are also more conspicuous to predators. Here we provide evidence that more chromatic male cabbage white butterflies (Pieris rapae) are more attractive to females but should also be more conspicuous to predators. Female P. rapae preferentially mate with more chromatic males when choosing from populations of males with naturally occurring or commensurate, experimentally induced color variation. Mathematical models of female color vision confirm that females should be able to discriminate color differences between prospective mates. Further, chromatic and luminance contrast scores from female visual system models better predicted male mating success than did measures of male color derived more directly from color spectra. Last, models of avian color vision suggest that preferred males should be more conspicuous to known avian predators. [ABSTRACT FROM AUTHOR]

1. Iridescent colour signals are directional but, like diffusely reflected colours, vary within and among species in ways that may be adaptations to specific types of receivers in specific light environments. 2. The hindwings of pipevine swallowtail butterflies exhibit brilliant blue and iridescent colour patches on the ventral surface in both sexes and on the dorsal wing surface in males. Evidence suggests that the ventral iridescent blue is a component of the warning coloration of this distasteful species, while the dorsal blue iridescent wing area is a sexual signal. Given differences in the function and ecological context of signal production, we analysed reflectance spectra from the iridescent blue areas of both field-caught and laboratory-reared animals to test several predictions about the iridescent colour patches on these wing surfaces. 3. The ventral blue patches in the warning coloration of males and females should be most visible early and late in the day, due to wing orientation relative to sun angle. We therefore predicted that these iridescent colour patches would be brighter and of longer wavelengths than the male dorsal blue patch displayed during midday courtships. The prediction about reflectance intensity was supported but the prediction about hue was not. 4. We predicted that the sexually selected dorsal hindwing iridescence of males would be more variable among individuals and condition dependent than the naturally selected ventral iridescent colour patches. To assess variation and condition dependence, laboratory-reared and field-captured individuals were compared. The prediction about variation was not supported, but only the dorsal wing surfaces showed evidence of being condition dependent. 5. We investigated whether development of dorsal and ventral blue iridescence was coupled by determining correlations in colour properties between the wing surfaces. Our finding of positive correlations indicated a potential developmental constraint in the evolution of colour differences between the two wing surfaces. 6. Results of this study suggest that some properties of iridescent coloration on the hindwing of the pipevine swallowtail (especially intensity) may have been fine-tuned by evolution in response to prevailing ambient light conditions and viewing perspectives that differ among types of signal receivers. [ABSTRACT FROM AUTHOR]

Understanding the evolution of herbivore–plant interactions requires detailed information on proximate responses to relevant dietary variability and genetic variance, if any, associated with these responses. We measured the behavioral and developmental responses of Pieris rapae larvae to variation in the nitrogen (N) and carbohydrate (CARB) content of chemically-defined diets, using differences in the average responses of sibling groups to estimate underlying genetic variance. Larval P. rapae responded to dietary reductions in both N and CARB with increased feeding, but these responses were inadequate to compensate for dietary N deficiencies within the range of N found in host plants. As a result, larvae on reduced N diets exhibited lower relative growth rates and longer development times, whereas larvae on reduced CARB diets maintained normal developmental trajectories. We also report evidence for genetic variation underlying (a) compensatory feeding responses to CARB and N availability, (b) N-driven variation in growth rate and (c) CARB-driven variation in larval duration. Our results highlight N availability as a key factor in the growth and development of this herbivorous butterfly, with CARB availability being less constraining and sufficiently addressed by changes to larval consumption rate. Furthermore, our study reveals standing genetic variance associated with larval responses to macronutrient availability, suggesting continued potential for herbivore–plant co-evolution in this system despite a putative history of strong directional selection. [ABSTRACT FROM AUTHOR]

Abstract: In some species of sulphur butterflies (Pieridae: Coliadinae) females as well as males display bright structural reflectance on their dorsal wing surfaces, although comparatively little attention has been paid to this coloration in females. We examined the spectral properties of female dorsal coloration and scale structure in three species of sulphurs for which published images show bright UV reflectance in females: the Neotropical Anteos clorinde and two species of Indo-Australian Eurema, E. hecabe and E. candida. In A. clorinde and E. hecabe, female UV reflectance is iridescent and produced by thin film interference in a system of ridges and lamellae, as it is in conspecific males. Female A. clorinde exhibit the same spatial distribution and chromaticity of UV reflectance as seen in males, but the UV reflectance in female E. hecabe is much smaller in area compared to that of conspecific males and is both less bright and less chromatic than observed in males. In contrast, UV reflectance in E. candida females is diffuse, and arises from a lack of pterin pigments in the wings, which permits a broad-band scattered reflection to be seen. This is the mechanism that is known to produce bright UV reflectance in females of the confamilial whites. Our results highlight the diversity of UV reflectances and underlying mechanisms in sulphurs and suggest multiple evolutionary pathways leading to this diversity in female sulphur butterflies. [Copyright &y& Elsevier]

Many of nature’s most striking animal colours are iridescent, exhibiting a high degree of spectral purity and strong angular dependence of intensity and hue. Although a growing number of studies have detailed the intricate mechanisms responsible for producing iridescent colours, few attempts have been made to describe their dynamic appearance in ecologically and behaviourally realistic contexts. We suggest that the optical properties unique to iridescent structural colours are important for understanding how they function as signals during behavioural interactions. Using males of the orange sulphur butterfly, Colias eurytheme, which exhibit an iridescent ultraviolet (UV) reflectance on their dorsal wing surfaces, we develop a holistic framework for inferring the appearance of this signal to conspecifics under field conditions that incorporate data on their spectral sensitivity. We show that, during flight, the UV signal is brightest within a wing beat cycle when viewed from directly above the male. Spectral properties of the signal under natural lighting indicate that male wing colour should be readily perceived and distinguished from that of females and from the dark green visual background of UV-absorbing vegetation. Finally, our analyses permit predictions regarding how signal senders and receivers should orientate themselves for maximal transmission and reception of this ultraviolet iridescent signal. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90, 349–364. [ABSTRACT FROM AUTHOR]

Structural colors result from an interaction between light and the fine-scale physical structure of a surface, and are often extremely bright, chromatic, and iridescent. Given that these visual features depend upon the aggregate abundance and architectural precision of photonic structures, structurally colored sexual ornaments seem well placed to indicate a range of mate quality characteristics. We tested this hypothesis by investigating the signaling potential of structural coloration in the sexually dimorphic butterfly Colias eurytheme. Males of this species display iridescent ultraviolet (UV) markings (arising from multilayer thin films) that overlay a broad area of yellowish-orange pigmentation on their dorsal wing surface. Only the structural UV has demonstrated function as a sexual signal; hence we predicted that it should contain more reliable phenotypic and/or genetic quality information, which would be indicated by phenotypic and/or genetically mediated condition dependence. In two split-family breeding experiments we manipulated condition by exposing full siblings to different stressors at two different juvenile life-history stages: (1) reduced larval host-plant quality and (2) transient heat/cold shocks during metamorphosis. Both stressors had profound effects on key developmental and life-history traits. Each stressor also significantly affected male dorsal coloration; thus, the expression of both structural and pigmentary coloration is phenotypically condition dependent. As predicted, the strongest condition dependence was evident in the brightness and angular visibility (i.e., iridescence) of the UV. Characteristics of both the iridescent UV and pigmentary orange also exhibited moderate-high and significant heritability (H² ∼ h² ∼ 0.4-0.9). However, genetic and residual variances did not increase under stress; thus, the observed condition dependence was not genetically mediated as predicted if wing color trait signals "good" genes for the ability to either withstand or circumvent developmental stress. The heightened stress sensitivity of the iridescent UV suggests that it offers an informative lifetime indicator of juvenile environments and, henceforth, adult male phenotypic condition, which may be salient to females seeking a highly fertile and/or nutritious male ejaculate. [ABSTRACT FROM AUTHOR]

Adaptations that enhance fitness in one sex may be harmful to members of the opposite sex and lead to antagonistic coevolution between the sexes. In fruit flies, for example, selection for fertilization success has rendered the male ejaculate slightly toxic to females. Here we investigated whether mating imposes a cost upon female fitness in a polyandrous pierid butterfly ( Colias eurytheme) by comparing life history traits between once-mated females and virgins. Mated females laid relatively more eggs early in their adult life, but suffered a reduction in longevity relative to virgins held under identical experimental conditions. The effect of mating on female survivorship was statistically independent of lifetime and early life fecundity. Moreover, lifetime fecundity co-varied positively with longevity across all females, and across females within each treatment group, hence there was no phenotypic trade-off between survival and reproduction. These results suggest that the observed longevity difference between virgin and mated females represents a true cost of mating, possibly arising from a toxic side effect of the male ejaculate. However, irrespective of this cost, virgin and mated females laid an equivalent lifetime number of eggs. Female C. eurytheme are also known to use nutrients from the male ejaculate to supplement their reproductive output, hence it is presently unclear how the observed longevity cost may have influenced the evolution of lifetime mating schedules in this polyandrous species. [ABSTRACT FROM AUTHOR]

When females mate multiple times it presents an intriguing problem for evolutionary biologists due to the high costs and not very apparent benefits. Yet, this behaviour must have higher benefits than costs to be maintained by natural selection. We studied possible benefits for multiple mating by female leaf beetles, Leptinotarsa decemlineata, both with multiple males (polyandry) and multiple times with the same male (repeated matings). For polyandry, we tested the material benefits hypothesis, as well as fertility insurance, by mating females to varying numbers of males and looking at sperm received and fecundity. Although there was a relationship between amount of stored sperm and number of matings, we did not find the predicted positive relationship between fecundity and number of matings. Thus, the material benefits hypothesis was not supported. For repeated matings, we tested the sperm and nonsperm material benefit hypotheses by mating females to a single male varying numbers of times on one mount. For this set of tests neither sperm number nor fecundity was positively correlated with number of matings, therefore neither hypothesis was supported. Interestingly, in both experiments, there was a significant decrease in hatch rate with an increase in matings, demonstrating a cost of polyandry. These findings, along with previous research, suggest a cost of multiple mating with a possible role of seminal fluids in the reduction in female fecundity. [Copyright &y& Elsevier]

Male Empress Leilia butterflies (Asterocampa leilia) use a sit-and-wait tactic to locate mates. To see how vision might influence male behavior, we studied the morphology, optics, and receptor physiology of their eyes and found the following. (1) Each eye's visual field is approximately hemispherical with at most a 10° overlap in the fields of the eyes. There are no large sexual differences in visual field dimensions. (2) In both sexes, rhabdoms in the frontal and dorsal ommatidia are longer than those in other eye regions. (3) Interommatidial angles are smallest frontally and around the equator of the eye. Minimum interommatidial angles are 0.9–1° in males and 1.3–1.4° in females. (4) Acceptance angles of ommatidia closely match interommatidial angles in the frontal region of the eye. We conclude that vision in these butterflies is mostly monocular and that males have more acute vision than females, especially in the frontal region (large facets, small interommatidial angles, small acceptance angles, long rhabdoms, and a close match between interommatidial angles and acceptance angles). This study also suggests that perched males direct their most acute vision where females are likely to appear but show no eye modifications that appear clearly related to a mate-locating tactic. [ABSTRACT FROM AUTHOR]

Male Hypolimnas bolina (L.) (Lepidoptera: Nymphalidae) defend perch sites as a method of visually locating receptive females. In this study we charted the spatio-temporal activity of perching males in tropical Australia to investigate (a) the extent to which male activity is thermally constrained, and (b) the physical attributes of selected perching sites. Butterflies were surveyed along two 5-15m wide open corridors through dense vegetation, and this allowed the prediction that males should favour narrower corridor sections in order to maximize their visual search capability. This prediction was supported. Beyond corridor width, the distribution of favoured perches was not explained any further by patterns of shading, larval food plant distribution, or the presence of nectar resources. Males were active from 0800 to 1700 h, but the number of perching individuals varied throughout the day, and this pattern varied between the two transects. Most individuals perched along one transect in the morning (0900-1100 h), whereas activity along the other peaked around midday (1100-1300 h). This between-transect difference in male activity followed changes in shading between the transects, however this variable did not predict male distribution at the territory scale. Although ambient and black body temperatures were significantly related to population-level activity, these variables only predicted 15-55% of the variance in male counts in individual transects. This result, viewed in conjunction with the limited available information on female receptivity, suggests that the timing of mate location in H. bolina may be primarily influenced by the daily pattern of female availability. [ABSTRACT FROM AUTHOR]

We experimentally investigated proximate factors influencing the visual detection of flying conspecifics by male butterflies (Asterocampa leilia) engaged in a sit-and-wait mate-searching tactic. Model butterflies were presented to perched males in the field using an apparatus that permitted us to control the path and speed of a model while varying minimum distance of the model from the male, height of the model above the ground, and model size. The dependent variable in all cases was whether or not the male left his perch and pursued the model. Males responded to normal-size models up to but not beyond distances of 3 m, and, because doubling the model surface area increased the distance at which males responded, we conclude that males do not detect conspecifics if they are more than 3 m away. At distances of 2 m or less males perched on the ground were more likely to detect conspecifics than males perched off the ground. This is likely to be due to differences either in the background against which the perched male typically views conspecifics or how large an angle conspecifics subtend from a perched male's perspective. These results suggest that thermally-driven changes during the activity period in perch preferences have consequences for success in mate detection that may be evolutionarily significant. [ABSTRACT FROM AUTHOR]

Males of the nymphalid butterfly Asterocampa leilia perch and wait during the morning at places where females are likely to appear. Males leave their perches to court passing females and chase away intruding males. As air and ground temperatures rise during the morning, males switch from perching on the ground to perching off the ground (average height = 0.87 m) for thermoregulatory reasons. To evaluate how this switch in perch location might affect mate detection, I have investigated how the three-dimensional posture of the male's body and head varies with perch location and how conspecifics fly through male perching areas. The body posture of males varies with perch location, as measured by pitch and roll relative to gravity, and yaw relative to the sun. Moreover, the pitch and roll of the head relative to the body is adjusted in a way that compensates for variation in body pitch and roll. These results, along with information on conspecific flight altitudes, suggest that when a male is perched on the ground his visual system is positioned in such a way that he is less likely to detect conspecifics flying nearby than when he is perched off the ground. Hence, it appears that early in the morning visual detection of mates and intruding males may be compromised by thermoregulatory concerns. [ABSTRACT FROM AUTHOR]

Evaluates the impact of thermal environment on the behavior of male insects waiting for females at encounter sites. Courting and mating between male and female butterflies; Effects of body temperature on mating behavior; Examination of four characteristics of male perch preference, body orientation and posture changes.

Eclosions in populations of some species of holometabolous insects only occur during a restricted time each day, often within as little as an hour or two. We have described the daily pattern of eclosions in a desert population of the Pipevine Swallowtail ( Battus philenor) so that we might better understand the daily pattern of this species' mating behavior. Our anecdotal impression that these animals eclose only around sunrise was confirmed both in a greenhouse with a natural light cycle and in an incubator with a semi natural 16:8 light:dark cycle. Our experiments indicate that on the day of eclosion, levels of short wavelength light at sunrise reach a point that triggers eclosion and that an internal clock contributes to the pattern of when eclosions occur during the day. Finally, we show that the internal clock is set by the light cycle to which the pupa is exposed during the last few days of the 11 to 12 day pupal stage. [ABSTRACT FROM AUTHOR]

Warning coloration deters predators from attacking distasteful or toxic prey. Signal features that influence warning color effectiveness are not well understood, and in particular, we know very little about how effective short-wavelength and iridescent colors are as warning color elements in nature and how warning signal effectiveness changes throughout the day. We tested the effect of these factors on predation risk in nature using specimens of the distasteful pipevine swallowtail butterfly, Battus philenor. B. philenor adults display both iridescent blue and diffusely reflecting orange components in their warning signal. We painted B. philenor wings to create five different model types: all-black, only-iridescent-blue, only-orange, iridescent-blue-and-orange (intact signal), and matte-blue-and-orange. We placed 25 models in each of 14 replicate field sites for 72 h and checked for attacks three times each day. Model type affected the likelihood of attack; only-orange models were, the only model attacked significantly less than the all-black model. Iridescence did not enhance or decrease warning signal effectiveness in our experiment because matte-blue-and-orange models were attacked at the same rate as iridescent-blue-and-orange models. Time of day did not differentially affect model type. Video recordings of attacks revealed that insectivorous birds were responsible. The results of this experiment, when taken with previous work, indicate that the response to blue warning coloration is likely dependent on predator experience and context, but that iridescence per se does not affect warning signals in a natural context. [ABSTRACT FROM AUTHOR]

Warning colouration functions to deter predators from attacking unprofitable (e.g., unpalatable or toxic) prey items. While warning colours are often characterized by long-wavelength components (e.g., orange, red or yellow), many distasteful animals across the animal kingdom display orange or red colour patches adjacent to or within a field of short-wavelength colouration such as blue (e.g., strawberry poison dart frogs, pipevine swallowtail and pyjama nudibranch), which yields a multicomponent visual warning signal. Here we show that, in such signals, blue and orange patches can function as redundant signal components; avian predators trained not to attack the intact blue and orange colouration of the pipevine swallowtail (Battus philenor) recognised the butterflies as distasteful even when the blue and orange were presented individually. Our results demonstrate that blue colouration and potential multiple, unimodal, signal components should be considered in research on visual warning signals, including in well-studied animals, such as dendrobatid frogs and swallowtail butterflies. [ABSTRACT FROM AUTHOR]

Predation on distasteful animals should favor warning coloration that is relatively conspicuous and phenotypically invariable. However, even among similarly colored individuals there can be variation in their warning signals. In butterflies, individual differences in larval feeding history could cause this variation. The warning signal of the pipevine swallowtail butterfly, Battus philenor L. (Lepidoptera: Papilionidae) consists of both a blue iridescent patch and pigmentbased orange spots on the ventral hindwing. B. philenor males also display a dorsal surface iridescent patch that functions as a sexual indicator signal. A previous study of iridescence in B. philenor found that the iridescent blue on both the dorsal and ventral hind wings is variable and significantly different between lab-reared and field-caught individuals. These differences could be the result of larval food deprivation in the field. Through experimental manipulation of larval diet, larval food deprivation was evaluated as a potential cause of the differences observed between lab and field individuals, and if food deprivation is a source of inter-individual variation in warning signals. B. philenor larvae were food restricted starting at two points in the last larval instar, and one group was fed through pupation. Adult coloration was then compared. Food deprivation led to poorer adult condition, as indicated by lower adult body mass, forewing length, and fat content of stressed individuals. As the level of food deprivation increased, the hue of the iridescent patches on both the dorsal and ventral hind wing shifted to shorter wavelengths, and the chroma of the orange spots decreased. The shifts in iridescent color did not match the differences previously found between lab and field individuals. However, the treatment differences indicate that food deprivation may be a significant source of warning color variation. The differences between the treatment groups are likely detectable by predators, but the effect of the variation on signal effectiveness and function remains to be empirically explored. [ABSTRACT FROM AUTHOR]

In butterflies, wing colour may simultaneously be under sexual selection in the context of mating selection and natural selection in the context of thermoregulation. In the present study, we collected mated females of the green-veined white butterfly ( Pieris napi) from locations spanning 960 km of latitude across Fennoscandia, and investigated sex-specific latitudinal wing colour variation in their offspring raised under identical conditions. We measured wing colour characteristics, including reflectance at wavelengths 300-700 nm and the degree of wing melanization. At all latitudes, females reflected more light in the short wavelengths (< 400 nm) and less in the long wavelengths (> 450 nm), and they were more melanized than males. However, female wing colour varied more with latitude than that of males. Among females, long wavelength reflectance decreased, whereas short wavelength reflectance and melanization increased, towards the north. By contrast, among males, latitudinal variation was found only in the ventral hindwing melanization. These results are consistent with the idea that the balance between natural and sexual selection acting on wing colour changes with latitude differently in males than females. The dark wing colour of females in the north may be a thermoregulatory adaptation, although males may be constrained from evolving the dark dorsal wing colour favoured by natural selection because of constant sexual selection across latitudes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ••, ••-••. [ABSTRACT FROM AUTHOR]

Studies of compound eyes have revealed that variation in eye structure can substantially affect visual performance. Here, we investigate the degree to which a stressful rearing environment, which decreases body size, affects the eye phenotype. Full siblings of the Orange Sulphur butterfly, Colias eurytheme, were collected from known parents and split within families among two diet treatments that varied in quality. In both sexes, individuals reared on the high-quality diet had larger eye height and anterior facet diameter, and therefore, by inference, superior vision. However, relative to their reduced body size, individuals reared on low-quality diet had proportionally larger eyes and facets than individuals reared on high-quality diet. We interpret this finding as evidence that butterflies encountering nutritional stress increased proportional investment in eye development to reduce loss of visual performance. We also found significant broad-sense genetic variation underlying eye structure in both males and females, and report novel heritability estimates for eye height and facet diameter. Surprisingly, there was greater genetic variation in eye height among males than among females, despite apparently stronger directional selection on male vision. We discuss the implications of these data for our understanding of eye development and evolution. [ABSTRACT FROM AUTHOR]

In sexual selection, multimodal signals elicit mate choice when more than one sensory modality is activated. However, determining the relative use of each signal is difficult because it requires a comprehensive understanding of the mating system and how this system works under natural conditions. We examined the role of structural ultraviolet (UV) reflectance and pheromones in the butterfly Colias eurytheme. Both traits are important in mediating interspecific interactions and pheromones have been implicated in intraspecific mate choice. UV reflectance, which arises from the presence of a multilayer thin-film interference array, has potential as an honest indicator of male condition, viability and/or age. We investigated the relevance of these signal traits to courtship success by releasing virgin females in the path of free-flying males until each female had rejected and accepted at least one male. This design facilitated a within-subjects (females) analysis of mate choice, thus controlling for potentially confounding variation in intrinsic female receptivity. Principal component analysis indicated that variation across males in UV brightness and pheromones was essentially orthogonal. Females preferred younger males (as subjectively adjudged by wing wear), and while age covaried with UV brightness and almost all pheromone descriptors, UV brightness emerged as the best and most general predictor of male mating success. Our results suggest that this trait serves as an important intraspecific sexual signal in C. eurytheme, and they provide the clearest evidence to date regarding the functional relevance of structural coloration to female mate choice in butterflies. We discuss the preferential use of one secondary sexual characteristic (UV reflectance) over another (pheromones) with regard to evolutionary strategies. [Copyright &y& Elsevier]

• We tested predator behavior that could shape the diversity of warning signals. • We measured innate avoidance and learning in quail and two breeds of chicken. • The innate response to blue but not orange varied among the three predator types. • Blue was learned less effectively than orange in some predator types. • Chicken breeds should be carefully considered in warning color experiments. Warning coloration deters predators from attacking unpalatable prey, and is often characterized by long-wavelength colors, such as orange and red. However, warning colors in nature are more diverse and include short-wavelength colors, like blue. Blue has evolved as a primary defense in some animals but is not common. One hypothesis for the maintenance of this diversity is interspecific variation in predator responses to signals. We tested this hypothesis with galliform birds: Gambel's quail (Callipepla gambelii) and two domestic chicken breeds (Gallus gallus domesticus; Plymouth Rocks, Cochin Bantams). We measured innate avoidance and learning responses to only blue prey, only orange prey, and orange-and-blue prey, where the blue was iridescent to represent the natural coloration of the pipevine swallowtail butterfly (Battus philenor). We predicted birds would have similar responses to orange, but vary in response to blue. Upon first encounter, Cochin Bantams did not attack blue and Gambel's quail readily attacked, indicating innate avoidance by Cochin Bantams. Plymouth Rocks had no innate aversion to any color, lower attack latencies and attacked most prey items. Cochin Bantams and Gambel's quail both learned orange and orange-and-blue quicker than blue. Our results support the hypothesis that interspecific variation in predator response could maintain warning color diversity. [ABSTRACT FROM AUTHOR]

Many organisms have evolved adaptive coloration that reduces their risk of predation. Cryptic coloration reduces the likelihood of detection/recognition by potential predators, while warning or aposematic coloration advertises unprofitability and thereby reduces the likelihood of attack. Although some studies show that aposematic coloration functions better at decreasing attack rate than crypsis, recent work has suggested and demonstrated that crypsis and aposematism are both successful strategies for avoiding predation. Furthermore, the visual environment (e.g. ambient lighting, background) affects the ability for predators to detect prey. We investigated these 2 related hypotheses using 2 well-known visually aposematic species of Heliconius butterflies, which occupy different habitats (open-canopy vs. closed-canopy), and one palatable, cryptic, generalist species Junonia coenia. We tested if the differently colored butterflies differ in attack rates by placing plasticine models of each of the 3 species in 2 different tropical habitats where the butterflies naturally occur: disturbed, open-canopy habitat and forested, closed-canopy habitat. The cryptic model had fewer attacks than one of the aposematic models. Predation rates differed between the 2 habitats, with the open habitat having much higher predation. However, we did not find an interaction between species and habitat type, which is perplexing due to the different aposematic phenotypes naturally occurring in different habitats. Our findings suggest that during the Panamanian dry season avian predation on perched butterflies is not a leading cause in habitat segregation between the 2 aposematic species and demonstrate that cryptically colored animals at rest may be better than aposematic prey at avoiding avian attacks in certain environments. [ABSTRACT FROM AUTHOR]

A key determinant of the intensity of sexual selection acting on a trait is how variation in that trait is related to variance in reproductive success of individuals. This connection compels efforts to assess lifetime mating number and how it varies among individuals in a population. In the Lepidoptera, female mating success can be assessed relatively easily by counting by the number of spermatophores in the female’s copulatory sac but male mating success in the field can often only be documented by observing copulations. Here we report a method for identifying whether or not males have recently mated that relies on the effect of mating on the appearance of the male’s reproductive tract in the pipevine swallowtail, Battus philenor. In this species laboratory experiments reveal that during mating, components of a male’s reproductive tract become shorter, decrease in mass, and change in appearance, irrespective of male age. These changes persist for at least two days after mating. After documenting these indicators of recent mating, we examined the reproductive tract of 68 field-caught males and found that twelve (17.6%) showed strong evidence of having mated recently. We found that older males were more likely to have recently mated. In addition, the color of the dorsal hindwing, a feature that females use in mate choice, was significantly greener in males, that according to our criteria, had recently-mated than in males that had not. [ABSTRACT FROM AUTHOR]

Many animals search for potential mates or prey using a perch-and-sally strategy. The success of such a strategy will depend on factors that affect the observer's ability to detect a passing resource item. Intrinsic factors (e.g., eye structure and physiology) have received much recent attention, but less is known about effects on object detection in nature and extrinsic factors such as size, coloration, and speed of a passing object and the background against which the object is viewed. Here, we examine how background affects the detection of butterfly models by perched males of the butterfly Asterocampa leilia in the field. We test the hypothesis that male choice of perch site in nature will influence the contrast between the object and background against which it is viewed and that this will influence success in detecting the object. We also test the effect of contrast by manipulating the brightness of the object and presenting butterfly models of different reflectance (ranging from black to white). We found an effect of model luminance, with dark models being most likely to elicit a response regardless of background. Further, there was an effect of background type with models viewed against blue sky eliciting the highest response. Perceived luminance contrast correlates to behavior; highly contrasting objects are more frequently detected. This study expands our understanding of visual system performance and has implications for our understanding of the behavior and evolutionary ecology of perching species. [ABSTRACT FROM AUTHOR]

Abstract: Sensory information plays a critical role in determining an animal''s behavior on both proximate and evolutionary timescales. Butterflies, like many other insects, use vision extensively over their lifetimes, and yet relatively little work has been published to date on their visual capabilities. We describe the visual system of a pierid butterfly, Colias eurytheme, with the ultimate goal of better understanding its role in shaping the behavior of this animal. We made several measurements: visual field dimensions, eye surface area, interommatidial angle (Δϕ), facet diameter (D), and eye parameter (p). C. eurytheme had a large visual field and considerable regional variation in visual acuity, as inferred by Δϕ and D. When compared to females, males had larger eye surface areas, smaller Δϕ, and larger D in all regions except ventrally. Both sexes had proportionally large eye surface areas compared to other butterflies. Minimum p in males was small, indicating that some regions of their eyes may operate close to the diffraction limit. Finally, we found that both eye surface area and D scaled positively, but with negative allometry to body size. We discuss the relevance of these visual characteristics to the biology and behavior of C. eurytheme. [Copyright &y& Elsevier]