Wing Patch Size Research Articles

Looks or personality: what drives damselfly male mating success in the wild?

Understanding the connection between personality and fitness is an important topic in both behavioural and evolutionary ecology. Most of our current knowledge stems from lab-studies despite that studies in natural populations for most parts generate more ecologically relevant information on behavioural variation and mating success. Here, we quantified Calopteryx splendens male mating success across two behavioural axes: courtship and boldness-aggression, testing personality and plasticity of behavioural traits in a natural population situated in the central-core of the species distribution. We also measured wing patch area and body size as these traits are important in damselfly sexual selection. The only behavioural trait that positively affected mating success was consistency in courtship effort. Among morphological traits, smaller wing patch size was associated with greater male mating success, while thorax size was a better predictor of male mating success than either personality or plasticity in boldness and activity. Our results demonstrate the importance of behavioural consistency over plasticity in a wild insect population and support prior studies showing the importance of body size for mating success.

Variability of a consistent trait: The size of the white wing patch in European Stonechats (Saxicola rubicola rubicola)

Numerous bird species exhibit striking white patches on their plumages that have been reported as signals of individual quality in the context of sexual selection. Whereas differences in white plumage traits between individuals have been well studied, phenotypic variation and the factors affecting their consistency within the individual have received less attention. Here, we studied the consistency in the size of the white wing patch and its components of variation in a Spanish population of European Stonechats Saxicola rubicola rubicola monitored over 6 years (2007–2012). The wing patch was larger in males than in females for all age‐classes, and first‐year individuals exhibited smaller wing patches than older birds, particularly males. This trait also varied within the plumage‐year at both population and individual level, with slight changes from the moulting period to the breeding season and a sharp decrease afterwards. In addition, patch size varied both between and within individuals across years. The daily mean temperature experienced in the month immediately prior to the moult (i.e. May) had a positive effect on male wing patch size. In females, however, no variable was found to explain wing patch variation satisfactorily. Despite being a dynamic trait throughout the moults, baseline and adjusted repeatabilities indicated a moderately high consistency for white wing patch size in both sexes. Therefore, the white wing patch in Stonechats is a dynamic trait that changes throughout individuals' lives, but is also a consistent trait at the individual level. These results indicate that this depigmented patch could convey information about both the long‐term quality and the current state of individuals, harmonizing the existence of intra‐individual changes in the size of sexual traits with its potential role as a genetic quality indicator.

The extent of white plumage patches in female Pied Flycatchers Ficedula hypoleuca is negatively associated with corticosterone concentration in partly unpigmented feathers

Sexually and socially selected signals are predicated to express the present and past condition of individuals and thereby their capacity to cope with environmental challenges. The concentration of corticosterone in feathers (CORTf) has been validated as a marker of activation of the hypothalamic-pituitary axis in birds during feather development. Measurements of CORTf can thus express the physiological stress of migratory birds during non-breeding periods of the annual cycle when feathers moulted in the wintering range are analysed. Thus, negative trends of ornament expression with CORTf may be predicted. Unpigmented plumage patches may constitute signals of phenotypic quality and are used in many species in social interactions by both sexes. Pied Flycatchers of both sexes Ficedula hypoleuca show white forehead and wing patches during the breeding season, of which the forehead and part of the wing patch (tertials) are moulted on the wintering grounds before spring migration. These patches are used as signals in social and sexual interactions and their expression has been previously related to several indicators of physiological status. We collected tertials of females during nestling provisioning in a Spanish population and determined their CORTf in the laboratory. CORTf was thus measured in some of the prenuptially moulted feathers used as signals of social dominance by breeding females. According to AICc model selection, CORTf turned up in all the best models explaining forehead patch size and in most best models for wing patch size when controlling for age and size. Both forehead and wing patch sizes were negatively related to CORTf, although the trend for wing patch size was not significant. This indicates that breeding females expressing large unpigmented patches experienced low levels of stress during the prenuptial moult in their winter quarters, or that strongly signalling individuals present a lower level of activation of the HPA axis.

Preference for supernormal stimuli tends to override initially learned associations for conspicuous prey traits: implications from a laboratory study

How predators select on conspicuous prey traits is not well understood. We used a laboratory setup to investigate the role of learning in predator choice of conspicuous visual traits. We used a generalist predator, the great tit, and coloured wings of males of two species of damselflies as prey. Wing pigmentation differs between the species in colour (green vs. blue) and size (large vs. small). Wing pigmentation is a sexually selected trait that experiences negative selection by avian predators. Inexperienced great tits showed no preference for the colour or the size of wing pigmentation. Great tits were then repeatedly exposed to rewarded wings with either large or small wing patch size. When these experienced birds were exposed to both wing patch sizes for the first time, they tended to prefer the wings with the large patch, irrespective of their previous experience. Our results suggest that the choice of the predator was based on an initial association of the trait to a reward followed by a preference for a supernormal stimulus, probably due to a larger sensory stimulation. We discuss the implications of our laboratory results in the light of previous estimates of damselfly predation risk under field conditions.

Ornaments and condition: plumage patch sizes, nutritional reserve state, reserve accumulation, and reserve depletion

Condition-dependence is considered as a dominant mechanism ensuring the fitness benefits of continued mate choice for heritable sexual signal traits, but crucial questions remain concerning the underlying physiological pathways. For example, it is unclear whether condition-dependence is mediated by the different amount of resource obtained, some uncheatable functional link with nutritional status, or the adverse effect of nutritional stress experienced by some individuals. Furthermore, the pattern of change in nutritional reserves in relation to ornamental traits has recently been proposed as a critical pillar of the condition-dependence concept, but this pattern is virtually unknown in natural populations. We quantified separate measures for actual body condition, lipid reserve accumulation rate, and lipid reserve depletion, and applied these measures to two white plumage ornaments of male collared flycatchers during courtship, during nestling rearing, and before the summer moult. Neither actual condition nor reserve accumulation rate before moult predicted the subsequent change of ornament sizes, but reserve depletion was accompanied by the reduction of forehead patch size to the following year. Wing patch size, a trait important in territoriality, was negatively related to both reserve accumulation and reserve depletion in the courtship period, but not related to current condition. Finally, irrespective of breeding phase, measures of current condition, and recent nutritional reserve depletion were negatively correlated, but both were unrelated to resource accumulation rate. These results indicate that measuring nutritional reserve trajectories in addition to actual condition may reveal functionally important processes underlying signal-condition correlations.Significance statementOrnamental signal characters are known to convey honest information to signal receivers through their dependence on nutritional condition. Here, we show that such signals can also indicate the temporal trends of condition. We examine plumage patch sizes and separate measures of actual condition, nutritional reserve accumulation, and nutritional reserve depletion in three contexts: during courtship, during nestling rearing, and before the summer moult. The results suggest nutritional stress effects on signal expression, and predictable reserve dynamics in relation to signal expression, thereby highlighting the usefulness of dynamic nutritional measures in clarifying the fundamental concept of condition-dependent signalling.

Mutual plumage ornamentation and biparental care: consequences for success in different environments

Lay SummaryParental care and reproductive success of collared flycatchers were related to own and partner’s sexual ornaments in partial interaction with environmental conditions. In a brood size manipulation experiment, male wing patch size and female plumage brightness correlated with the partner’s feeding rate. Nestling quality correlated with male forehead patch size depending on breeding environment. These findings reveal a complex picture of relationships between the sexual ornament expression of parents and their parental care and success.

Interspecific interactions and learning variability jointly drive geographic differences in mate preferences

Co-occurrence of closely related species can cause behavioral interference in mating and increase hybridization risk. Theoretically, this could lead to the evolution of more species-specific mate preferences and sexual signaling traits. Alternatively, females can learn to reject heterospecific males, to avoid male sexual interference from closely related species. Such learned mate discrimination could also affect conspecific mate preferences if females generalize from between species differences to prefer more species-specific mating signals. Female damselflies of the banded demoiselle (Calopteryx splendens) learn to reject heterospecific males of the beautiful demoiselle (C. virgo) through direct premating interactions. These two species co-occur in a geographic mosaic of sympatric and microallopatric populations. Whereas C. virgo males have fully melanized wings, male C. splendens wings are partly melanized. We show that C. splendens females in sympatry with C. virgo prefer smaller male wing patches in conspecific males after learning to reject heterospecific males. In contrast, allopatric C. splendens females with experimentally induced experience with C. virgo males did not discriminate against larger male wing patches. Wing patch size might indicate conspecific male quality in allopatry. Co-occurrence with C. virgo therefore causes females to prefer conspecific male traits that are more species specific, contributing to population divergence and geographic variation in female mate preferences.

Carry-over effects of conditions at the wintering grounds on breeding plumage signals in a migratory bird: roles of phenotypic plasticity and selection.

To understand the consequences of ever-changing environment on the dynamics of phenotypic traits, distinguishing between selection processes and individual plasticity is crucial. We examined individual consistency/plasticity in several male secondary sexual traits expressed during the breeding season (white wing and forehead patch size, UV reflectance of white wing patch and dorsal melanin coloration) in a migratory pied flycatcher (Ficedula hypoleuca) population over an 11-year period. Furthermore, we studied carry-over effects of three environmental variables (NAO, a climatic index; NDVI, a vegetation index; and rainfall) at the wintering grounds (during prebreeding moult) on the expression of these breeding plumage traits of pied flycatcher males at individual and population levels. Whereas NAO correlates negatively with moisture in West Africa, NDVI correlates positively with primary production. Forehead patch size and melanin coloration were highly consistent within individuals among years, whereas the consistency of the other two traits was moderate. Wing patch size decreased with higher NAO and increased with higher rainfall and NDVI at the individual level. Interestingly, small-patched males suffered lower survival during high NAO winters than large-patched males, and vice versa during low NAO winters. These counteracting processes meant that the individual-level change was masked at the population level where no relationship was found. Our results provide a good example of how variation in the phenotypic composition of a natural population can be a result of both environment-dependent individual plasticity and short-term microevolution. Moreover, when plasticity and viability selection operate simultaneously, their impacts on population composition may not be evident.

Bird predation selects for wing shape and coloration in a damselfly.

Wing shape is related to flight performance, which is expected to be under selection for improving flight behaviours such as predator avoidance. Moreover, wing conspicuousness, usually involved in sexual selection processes, is also relevant in terms of predation risk. In this study, we examined how predation by a passerine bird, the white wagtail Motacilla alba, selects wing shape and wing colour patch size in males of the banded demoiselle Calopteryx splendens. The wing colour patch is intra- and intersexually selected in the study species. In a field study, we compared wings of live damselflies to wings of predated damselflies which are always discarded after predation. Based on aerodynamic theory and a previous study on wing shape of territorial tactics in damselflies, we predicted an overall short and broad wing, with a concave front margin shape to be selected by predation. This shape would be expected to improve escaping ability. Moreover, we predicted that wing patch size should be negatively selected by predation. We found that selection operated differently on fore- and hindwings. In contrast to our predictions, predation favoured a slender general forewing shape. However, the predicted wing shape was favoured in hindwings. We also found selection favouring a narrower wing colour patch. Our results suggest different roles of fore- and hindwings in flight, as previously suggested for Calopteryx damselflies and shown for butterflies and moths. Forewings would be more involved in sustained flight and hindwings in flight manoeuvrability. Our results differ somehow from a recently published work in the same study system, but using another population, suggesting that selection can fluctuate across space, despite the simplicity of this predator-prey system.

The relationship between maternal ornamentation and feeding rate is explained by intrinsic nestling quality

In altricial birds, parental feeding is essential, and its amount may depend on the quality of both parents. A relationship between parental quality and feeding rate is generally attributed to an active adjustment by parents in order to retain good quality mates or ensure high fitness through raising high-quality offspring. However, the behaviour and need of young may also change with parental quality, and this may affect parental behaviour. A further problem is that most studies have investigated post-hatching parental investment in relation to the secondary sexual signals of males, but not females. In a cross-fostering experiment, we examined the feeding rates of rearing parents in relation to the size and ornamentation of both original and rearing parents in collared flycatchers (Ficedula albicollis). Using this setup, we could examine whether the observed feeding patterns were the results of the decision of the parents based on their own and their partner’s traits or the constraints imposed by the behaviour or need of offspring. When correcting for clutch size and year, we found that feeding rate of both foster parents correlated with the wing patch size of the original female. This implies that original maternal quality had an offspring-mediated indirect effect on investment of foster parents, that is intrinsic nestling quality may constrain parental feeding decisions. This explanation should not be overlooked in future studies on preferential parental investment, and our results also point out that maternal ornaments deserve more attention in such studies.

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.

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.

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.

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]

Selective Predation on Wing Morphology in Sympatric Damselflies

Although predation is thought to affect species divergence, the effects of predator-mediated natural selection on species divergence and in nonadaptive radiations have seldom been studied. Wing melanization in Calopteryx damselflies has important functions in sexual selection and interspecific interactions and in species recognition. The genus Calopteryx and other damselfly genera have also been put forward as examples of radiations driven by sexual selection. We show that avian predation strongly affects natural selection on wing morphology and male wing melanization in two congeneric and sympatric species of this genus (Calopteryx splendens and Calopteryx virgo). Predation risk was almost three times higher for C. virgo, which has an exaggerated degree of wing melanization, than it was for the less exaggerated, sympatric congener C. splendens. Selective predation on the exaggerated species C. virgo favored a reduction and redistribution of the wing melanin patch. There was evidence for nonlinear selection involving wing patch size, wing patch darkness, and wing length and width in C. splendens but weaker nonlinear selection on the same trait combinations in C. virgo. Selective predation could interfere with species divergence by sexual selection and may thus indirectly affect male interspecific interactions, reproductive isolation, and species coexistence in this genus.

Selective Predation on Wing Morphology in Sympatric Damselflies

Although predation is thought to affect species divergence, the effects of predator‐mediated natural selection on species divergence and in nonadaptive radiations have seldom been studied. Wing melanization in Calopteryx damselflies has important functions in sexual selection and interspecific interactions and in species recognition. The genus Calopteryx and other damselfly genera have also been put forward as examples of radiations driven by sexual selection. We show that avian predation strongly affects natural selection on wing morphology and male wing melanization in two congeneric and sympatric species of this genus (Calopteryx splendens and Calopteryx virgo). Predation risk was almost three times higher for C. virgo, which has an exaggerated degree of wing melanization, than it was for the less exaggerated, sympatric congener C. splendens. Selective predation on the exaggerated species C. virgo favored a reduction and redistribution of the wing melanin patch. There was evidence for nonlinear selection involving wing patch size, wing patch darkness, and wing length and width in C. splendens but weaker nonlinear selection on the same trait combinations in C. virgo. Selective predation could interfere with species divergence by sexual selection and may thus indirectly affect male interspecific interactions, reproductive isolation, and species coexistence in this genus.

Determinants of male territorial behavior in a Hungarian collared flycatcher population: plumage traits of residents and challengers

In intrasexual conflicts, contestants can rely on relative or absolute size of status badges to mediate aggressive behavior. Most studies focus on the response of focal animals to variation in status badges of their competitors; few have simultaneously considered the traits of both participants under experimental conditions. By simulating territorial intrusions, we tested the importance of two sexual traits [forehead patch size (FPS) and wing patch size (WPS)] in territorial behavior of males in a Hungarian population of the collared flycatcher, Ficedula albicollis. We presented a stimulus male to an unpaired resident male to examine whether the characteristics of the territory owner or those of the challenger were associated with the latency of the first attack of the owner, which is a good predictor of the territorial behavior in general. WPS of the stimulus male was a significant determinant of the latency of the first attack, as males with a larger WPS elicited quicker attacks from the residents than males with a small WPS. From the residents’ perspective, age appeared to influence their territorial behavior, as yearlings had shorter attack latencies than older males. Additionally, latency could be considered an individual-specific attribute because it varied consistently among males, even when the WPS of the stimulus male was controlled, and it was associated with pairing success. Contrary to findings in a Swedish population, FPS seemed to be unimportant in male–male competition in our population, which suggests population differences in the role of the two plumage traits. Our results indicate that in a territorial conflict, the characteristics of both participants are important.