Wing Patches Research Articles

SEX CHROMOSOME LINKED GENETIC VARIANCE AND THE EVOLUTION OF SEXUAL DIMORPHISM OF QUANTITATIVE TRAITS

Theory predicts that sex chromsome linkage should reduce intersexual genetic correlations thereby allowing the evolution of sexual dimorphism. Empirical evidence for sex linkage has come largely from crosses and few studies have examined how sexual dimorphism and sex linkage are related within outbred populations. Here, we use data on an array of different traits measured on over 10,000 individuals from two pedigreed populations of birds (collared flycatcher and zebra finch) to estimate the amount of sex-linked genetic variance (h(2)z ). Of 17 traits examined, eight showed a nonzero h(2)Z estimate but only four were significantly different from zero (wing patch size and tarsus length in collared flycatchers, wing length and beak color in zebra finches). We further tested how sexual dimorphism and the mode of selection operating on the trait relate to the proportion of sex-linked genetic variance. Sexually selected traits did not show higher h(2)Z than morphological traits and there was only a weak positive relationship between h(2)Z and sexual dimorphism. However, given the relative scarcity of empirical studies, it is premature to make conclusions about the role of sex chromosome linkage in the evolution of sexual dimorphism.

Long-term effects of yolk androgens on phenotype and parental feeding behavior in a wild passerine

Early growth conditions, such as exposure to maternally derived androgens in bird eggs, have been shown to shape offspring in ways that may have important long-term consequences for phenotype and behavior. Using an experimental approach, we studied the long-term effects of yolk androgens on several phenotypic traits and parental behavior in adult and female collared flycatchers (Ficedula albicollis). We elevated yolk androgen levels and monitored the experimental recruits the following breeding seasons. Androgen treatment had a sex-dependent effect on adult body condition, yolk androgen-treated males being heavier than control males when controlling for size, a result which may be caused potentially by selective mortality, physiological differences, or different life-history strategies. Androgen treatment did not however affect the expression of sexually selected plumage ornaments (forehead and wing patch size), UV coloration, or parental feeding rate in either sex. Our results suggest that yolk androgens are unlikely to affect sexual selection via plumage characteristics or contribute to breeding success via altered parental care. Yolk androgens do not seem to act as a means for female collared flycatchers to enhance the attractiveness of their sons. The lower return rate previously observed for androgen-treated male offspring compared to controls may therefore not be due to lower mating or breeding success, but may rather reflect lower survival or higher dispersal propensity of yolk androgen-treated males.

A photonic heterostructure produces diverse iridescent colours in duck wing patches

The colours of birds are diverse but limited relative to the colours they can perceive. This mismatch may be partially caused by the properties of their colour-production mechanisms. Aside from pigments, several classes of highly ordered nanostructures (thin films, amorphous three-dimensional arrays) can produce a range of colours. However, the variability of any single nanostructural class has rarely been explored. Dabbling ducks are a speciose clade with substantial interspecific variation in the iridescent coloration of their wing patches (specula). Here, we use electron microscopy, spectrophotometry, polarization and refractive index-matching experiments, and optical modelling to examine these colours. We show that, in all species examined, speculum colour is produced by a photonic heterostructure consisting of both a single thin-film of keratin and a two-dimensional hexagonal lattice of melanosomes in feather barbules. Although the range of possible variations of this heterostructure is theoretically broad, only relatively close-packed, energetically stable variants producing more saturated colours were observed, suggesting that ducks are either physically constrained to these configurations or are under selection for the colours that they produce. These data thus reveal a previously undescribed biophotonic structure and suggest that both physical variability and constraints within single nanostructural classes may help explain the broader patterns of colour across Aves.

Winter body condition in relation to age, sex and plumage ornamentation in a migratory songbird

Winter body condition may play important roles in the life history of migratory birds, but it is difficult to estimate. We used the growth rate of winter‐grown tail feathers of Collared Flycatchers Ficedula albicollis as an indicator of winter body condition, comparing this trait between age classes and sexes and relating it to plumage ornamentation (forehead and wing patch sizes). Adults and males were in better nutritional condition during winter, as indicated by their faster tail feather growth rates, than were yearlings and females, respectively, which could indicate differences in individual quality and foraging ability with age, or age‐ and sex‐related winter habitat segregation. However, feather growth rate was related neither to the size of the winter‐grown forehead patch nor to the size of the summer‐grown wing patch, suggesting weak condition‐dependence for the winter‐grown ornament and complex life‐history consequences for the summer‐grown ornament.

Experimental manipulation shows that the white wing patch in collared flycatchers is a male sexual ornament

Descriptive analysis suggests that a conspicuous white wing patch in dichromatic (black and white) pied and collared flycatchers is under sexual selection. Here, we use an experimental approach to test whether this trait is indeed the target of selection. We caught 100 collared flycatcher Ficedula albicollis males soon after their arrival on the breeding site. We reduced (blackened) part of the white wing patch in half of these males and recorded their mating success and within and extra-pair offspring production. Reduction of the size of the white wing patch lowered a male's probability to attract a secondary social female, but not a primary female. However, primary females paired to males with a reduced wing patch were smaller (in tarsus), suggesting that male choice of partner or female–female competition over mates occurs in this species. The probability of pairing with a primary female (but not other components of male reproductive success) declined with arrival time (proxied by the date of capture). Males with a reduced wing patch size tended to sire less extra-pair offspring, although this relationship was reversed in one of the three study plots, suggesting that mating dynamics are context dependent. While our findings show that wing patch size is the target of sexual selection, the pathways and the strength of selection on this ornament differed markedly from a previous descriptive study. Nonexperimental studies of sexual selection in the wild may overestimate its importance because male fitness and ornamentation both depend positively on environmental conditions.

A ROLE FOR LEARNING IN POPULATION DIVERGENCE OF MATE PREFERENCES

Learning and other forms of phenotypic plasticity have been suggested to enhance population divergence. Mate preferences can develop by learning, and species recognition might not be entirely genetic. We present data on female mate preferences of the banded demoiselle (Calopteryx splendens) that suggest a role for learning in population divergence and species recognition. Populations of this species are either allopatric or sympatric with a phenotypically similar congener (C. virgo). These two species differ mainly in the amount of wing melanization in males, and wing patches thus mediate sexual isolation. In sympatry, sexually experienced females discriminate against large melanin wing patches in heterospecific males. In contrast, in allopatric populations within the same geographic region, females show positive ("open-ended") preferences for such large wing patches. Virgin C. splendens females do not discriminate against heterospecific males. Moreover, physical exposure experiments of such virgin females to con- or hetero-specific males significantly influences their subsequent mate preferences. Species recognition is thus not entirely genetic and it is partly influenced by interactions with mates. Learning causes pronounced population divergence in mate preferences between these weakly genetically differentiated populations, and results in a highly divergent pattern of species recognition at a small geographic scale.

Melanin coloration has temperature‐dependent effects on breeding performance that may maintain phenotypic variation in a passerine bird

Fluctuating selection pressure may maintain phenotypic variation because of different types of individuals being adapted to different environmental conditions. We show that the extensive variation in the coloration of male pied flycatchers (Ficedula hypoleuca) can be maintained through differences in the reproductive success of male phenotypes under different conditions. The effects of weather conditions on the relative success of different male phenotypes varied between different phases of breeding. The reproductive output of black males was the highest when it was cold during egg-laying but warm during the nestling period, whereas the fledgling production of brown males was highest when it was continuously warm. In addition, male forehead and wing patch sizes had context-dependent effects on timing of breeding and nestling mortality, respectively. These results indicate that environmental heterogeneity plays a role in maintaining phenotypic variation. As melanin-based coloration is heritable, climate change may alter phenotype frequencies depending on the patterns of warming.

Butterfly wing colors: glass scales of Graphium sarpedon cause polarized iridescence and enhance blue/green pigment coloration of the wing membrane

The wings of the swordtail butterfly Graphium sarpedon nipponum contain the bile pigment sarpedobilin, which causes blue/green colored wing patches. Locally the bile pigment is combined with the strongly blue-absorbing carotenoid lutein, resulting in green wing patches and thus improving camouflage. In the dorsal forewings, the colored patches lack the usual wing scales, but instead have bristles. We have found that on the ventral side most of these patches have very transparent scales that enhance, by reflection, the wing coloration when illuminated from the dorsal side. These glass scales furthermore create a strongly polarized iridescence when illuminated by obliquely incident light from the ventral side, presumably for intraspecific signaling. A few ventral forewing patches have diffusely scattering, white scales that also enhance the blue/green wing coloration when observed from the dorsal side.

Nutritional correlates and mate acquisition role of multiple sexual traits in male collared flycatchers

The information content of a sexual signal may predict its importance in a multiple signal system. Many studies have correlated sexual signal expression with the absolute levels of nutrient reserves. In contrast, the changes of nutrient reserves associated with signal expression are largely unknown in the wild due to technical limitations although they are important determinants of signal information content. We compared two visual and eight acoustic sexual traits in male collared flycatchers to see whether the nutritional correlates of expression predict the role of the signal in sexual selection. We used single point assays of plasma lipid metabolites to estimate short-term changes in nutritional state in relation to sexual trait expression during courtship. As a measure of sexual selection, we estimated the relationship with pairing latency after arrival in a 4-year dataset. Males which found a mate rapidly were characterized by large wing and forehead patches, but small song strophe complexity and small figure repertoire size. Traits more strongly related to pairing latency were also more closely related to changes in nutrient reserves. This indicates a link between signal role and information content. Small wing patches and, surprisingly, complex songs seemed to indicate poor phenotypic quality and were apparently disfavoured at mate acquisition in our population. Future studies of the information content of sexual traits, especially dynamic traits such as song, may benefit from the use of plasma metabolite profiles as non-invasive indicators of short-term changes in body condition.

Distinguishing between male and territory quality: females choose multiple traits in the pied flycatcher

To understand how sexual selection works, it is important to distinguish between the mechanisms underlying mating success. One of the basic questions is to differentiate between the importance of territory-related resources (gained through male–male competition) and male quality in female choice. Few studies, however, have tried to separate these two often intercorrelated variables. We randomized territory quality among male pied flycatchers, Ficedula hypoleuca, and studied which male traits were associated with their success in female mate choice. Females appeared to choose multiple male traits in the wild when the males were not allowed to choose territories before female choice for males. Males with a long tarsus, large white wing patch and versatile song were favoured by females. In addition, the contrast of ultraviolet (UV) reflectance in the white wing patch relative to its background, that is, the dorsal brown-black coloration of the males, appeared to be an important mate choice cue. The UV reflectance had a stronger effect on pairing success in dark males than in brown ones. An experimental manipulation of the UV reflectance revealed that the UV coloration of the males appeared to be used in female choice early but not late in the pairing season, which suggests time-dependent plasticity in female choice. Our study thus exemplifies the complexity of mate choice; it is both temporally constrained and dependent on multiple ornaments.

Microelectrical Mechanical Systems Switch for Designing Multi-Band Antenna

Problem statement: In this study Microelectrical Mechanical System (MEMS) switches were proposed to design a reconfigurable/multi-band antenna to replaced PIN diode switches or semiconductor switches due to lower insertion losses, good isolation, much lower intermodulation distortion, and lower power consumption. The antenna is able to operate at very high frequencies. Approach: A reconfigurable antenna that is capable to operate at several frequencies was proposed by introducing two adjacent patches along with main radiating patch and two MEMS switches. Parametric analysis of the size of the wing patches was done for finding optimum size. A comparative study was done for Alumina, SiN, GaAs and Teflon as MEMS bridge materials for finding better results in terms of return loss and number of bands. The design was performed by using 3D electromagnetic simulator HFSS considering ideal MEMS switches. Results: It was found that SiN as MEMS bridge material makes the antenna to operate at 16.76, 23.56 and 27.7 GHz in the &#34OFF&#34 states and operate at 20.9 and 21.91 GHz in the &#34ON&#34 states of MEMS switches. Conclusion/Recommendations: MEMS cantilever beam material played an important role for providing antenna to operate at multi-band frequencies. The proposed multiband/reconfigurable antenna can be implemented with easy fabrication process steps by the Sandwich method of fabrication.

Structural colours create a flashing cue for sexual recognition and male quality in a Neotropical giant damselfly

Summary1. Structural coloration is common among animals that produce sexual displays involving motion or ultraviolet reflection. Different sources of colour may provide multicomponent signals that indicate the location, sex, and fitness of a potential mate or rival. We investigated the proximate basis and ultimate function of the wing coloration of the territorial damselflyMegaloprepus caerulatus,which produces a dynamic, high contrast display during flight. The wings of both sexes have blue and white bands, but the location of the white patches are sex specific.2. Wax filaments produce diffuse, white areas through broadband scattering of wavelengths between 300 and 700 nm. Blue bands reflect wavelengths between 300 and 500 nm (λmax = 398 nm) and shift in hue with viewing angle, the result of thin layer interference produced by layers of cuticle and pigment within the wing membrane. Both wing bands strongly reflect UV wavelengths.3. Both the iridescent UV‐blue and white wing patches provide high contrast against the vegetation in forest light gaps where mating occurs. Moreover, the iridescent signal oscillates during flight. Angle‐dependent UV‐blue iridescence is periodically extinguished during each wing beat cycle, in contrast to the white areas, which remain bright.4. Males distinguish potential mates from rivals by the presence of a female’s white wing tip. Blackening the white wing bands of males and adding white wing tips to resemble a female elicits a sexual rather than aggressive response from males. Conversely, blackening the white wing tips of females reduces sexual responses.5. The proportional area of the white wing bands of males is indicative of wing symmetry, correlated with body size, and in turn, territory residency suggesting that it may serve as a signal of male condition during intra‐ and intersexual interactions.6. We propose that the flashing iridescent UV‐blue wing bands provide a beacon to potential mates across forest light gaps, whereas the white patches serve in mate recognition and may indicate male quality or territorial status. Our study identifies a unique combination of interference and broadband reflectors that provide a dynamic multicomponent signal.

Patterns of Phenotypic Divergence in Wing Covariance Structure of Calopterygid Damselflies

Comparing species differences in covariance patterns of traits subject to divergent selection pressures can increase our understanding to the mechanisms of phenotypic divergence. Different species of calopterygid damselflies have diverged in the melanized wing patch of males. This trait serves multiple ecological functions and has behavioral consequences in terms of sexual selection, interspecific interactions, reproductive isolation. We compared the phenotypic variance-covariance matrices (P) of wing traits among nine populations of four European species of calopterygid damselflies. We found modest divergence in covariance structure among populations of the same species, but strong divergence between species. Interestingly, the orientation of the first eigenvector of P (P (max) ) differed more between closely related species than between distantly related species, although this pattern was absent when overall covariance structures were compared. We also found that distantly related species but geographically closer had converged towards a similar covariance structure. Finally, divergence in covariance structure was correlated with divergence in wing patch length, but not with other wing traits. This last finding suggests that divergent selection on wing patch length might have affected the stability of P. These results indicate that P might not only reflect ancestral developmental pathways but might also be influenced by current ecology.

Different traits affect gain of extrapair paternity and loss of paternity in the pied flycatcher, Ficedula hypoleuca

Extrapair paternity may create marked variation in the reproductive success of males and thus contribute to the evolution of sexually selected traits in socially monogamous mating systems. We examined the distribution of extrapair paternity in a passerine bird, the pied flycatcher, in relation to plumage coloration and male age. The frequency of extrapair paternity was found to be low in our study population in Southwest Finland (38 of 857 (4.4%) offspring in 25 of 191 (13%) broods). We were able to identify the sire for 53% of the extrapair young and in each case the extrapair sire was at least 2 years old. The probability of siring extrapair young was not associated with pigment-based brown-black dorsal plumage coloration, structural ultraviolet (UV) coloration or forehead patch size. The probability of being cuckolded, however, increased with decreasing UV chroma of the white wing patch. Our results thus indicate that there are multiple factors influencing when and with whom females engage in extrapair copulations. They also indicate that the male traits influencing the decision to cuckold the social mate are different from those used for choosing an extrapair mate.

Individual quality and reproductive effort mirrored in white wing plumage in both sexes of south polar skuas

It is well established that female choice may lead to sexual selection on quality-revealing ornaments in males. However, in many species, both sexes display conspicuous characters, which may reflect individual quality and condition. We examined the correlations between measures of individual condition, reproductive performance and variation in size and whiteness of white wing patches in both sexes of the south polar skua (Catharacta maccormicki). Females with a whiter patch had a lower mean clutch size and higher survival, and males with whiter patches had higher immune responses against the injected immunogen tetanus. Birds with a larger white patch, on the other hand, had a higher mass loss, and females with large white patches laid larger eggs and had reduced survival probability. Thus, variation in 2 measured aspects of the wing patches in the south polar skua seems to mirror a gradient of reproductive investment strategies; from 1) reduced reproductive investment, better immune defense, and higher survival in birds with more intensely white wing patches to 2) high reproductive investment but survival costs in birds with larger patches.

Female ornamentation and territorial conflicts in collared flycatchers (Ficedula albicollis)

Female ornaments in species with conventional sex roles often indicate individual quality, but the evolutionary forces maintaining them are less clear. Sexual competition for breeding opportunities may represent an important role for female signals, especially in polygynous species, but there is little experimental evidence for this. The wing patch size (WPS) of female collared flycatchers indicates age and body condition and predicts social mating patterns. We challenged nest-building females with decoy females of varying WPS and found that the aggressive response of residents increased with decoy WPS, suggesting a role for this female ornament in territorial competition. Our results explain why female WPS predicts territorial distances when mated to a polygynous male and indicate that the role of WPS in female competitive interactions is similar to that in males of the same population.

Intrasexual selection on multiple plumage ornaments in the lark bunting

Competition over resources can lead to fights and injury, so many species have evolved badge-of-status signals to settle conflicts without resorting to overt aggression. Most studies of status signals have focused on a single trait, under the assumption that aggression is univariate and therefore multiple signals would be redundant. We examined the relationship between male–male social dominance and several plumage traits (wing patch size and colour, body colour and coverage of black feathers on the body or the rump) in lark buntings, Calamospiza melanocorys, using a combination of observation and experimental territorial intrusions. Large body size was correlated with elevated aggression in several assays. Our observational data suggest that some traits function as close-range badges of status, because males with a larger proportion of black feathers on the body and rump win escalated contests, whereas other traits function at longer distances, because males with larger wing patches experience fewer territorial intrusions. Experimental tests of social dominance further suggest that different male plumage traits convey different information: some traits were associated with physical aggression, whereas other traits were associated with the intensity of approach. Together, these results show that selection can favour multiple badges of status to accommodate different levels of escalation or context-dependent dominance interactions.

The roles of ecological factors and sexual selection in the evolution of white wing patches in ducks

Most studies of the evolution of sexual ornamentation have dealt with plumage attributes. White plumage patches are widespread in birds, the disproportionate role of wearing costs makes their evolutionary trajectories unique, and their visual assessment is less biased than that of other color categories. Still, comparative studies of white patches are very rare. We examined the evolution of white wing patches in ducks (Anatinae), assessing both sex-specific trait expression and dichromatism. Habitat openness, nest site (cavity or open), or the length of the incubation period did not predict white patch expression. Patch size on the wing coverts of females increased with parental care contribution by males. Covert patch size relative to the wing surface was positively related to body size in males, suggesting a role in sexual competition. White wing patch expression was unrelated to the frequency of social mate change or testis size, a measure of general sperm competition intensity. However, covert patch size in both sexes showed strong negative correlation with the phallus length of males, an indicator of the prevalence of forced copulations. Further studies are needed to clarify the role of flight feather patches and the factors that limit the exaggeration of white wing patches in this and other groups.

Phenotypic plasticity in a conspicuous female plumage trait: information content and mating patterns

Explaining sexual ornamentation in the limiting sex, usually females, requires information on the proximate background of ornaments and their consequences for sexual selection. Phenotypic variation within individuals has received little attention in either of these research directions. We used 6 years of data to examine the information content and potential role of white wing patch size in female collared flycatchers, Ficedula albicollis. Female wing patch size differed among years. Yearling females had smaller wing patches than older females. The negative effect of original patch size on intraindividual patch size change was stronger in yearling than in older birds, which may reflect an age-dependent trade-off. Change in wing patch size was strongly positively related to the summer North Atlantic Oscillation (NAO) index. Clutch size laid in the previous breeding season had a negative effect on patch size change, but only in high-NAO years. Thus, the immediate effect of poor climate during moult apparently overrode the more indirect influence of reproductive effort. Two sexually selected ornaments of mates were unrelated to female wing patch size at the population level, but intraindividual changes of female wing patch size significantly predicted differences in wing patch size between mates obtained in the 2 years. Our data suggest that significant mating advantages to more ornamented females may not be detected from population-level mating patterns. Research on potential female ornaments should also pay more attention to age-dependent phenotypic plasticity, the trade-off between current and future ornament size and the costs of reproduction to apparent future attractiveness.

A role for female ornamentation in the facultatively polygynous mating system of collared flycatchers

In a polygynous mating system, females settling with already mated males often experience low mating success due to the reduced parental contribution of the male. However, there are numerous factors that may still make it advantageous for some females to choose this mating status. Facultative polygyny is believed to be dominated by male advertisement and female choice. Although quality differences and competition among females are increasingly recognized as important determinants of polygynous settlement patterns, the importance of signals of female quality in this mating system is largely unknown. Here we examined the relationship of the white wing patch size (WPS) of female collared flycatchers, a phenotypically plastic and age-dependent ornament, with social mating status, while controlling for settlement date and age. At the population level, monogamous, primary, and secondary females did not differ in WPS. However, the primary female of individual males was more ornamented than the secondary female, and this difference declined with increasing distance between primary and secondary nests. Secondary female ornamentation increased, whereas that of the primary female did not change with nest distance. These results suggest a subtle role for female ornamentation at polygynous mating. Future studies should therefore take into account mating status when assessing the costs and benefits of female signals. Moreover, patterns in quality-indicating female traits may contribute to the explanation of differences in reproductive success among females of different mating status. Key words: female sexual ornament, individual quality, polygyny, settlement date, territorial distance. [Behav Ecol]