Soil and preen waxes influence the expression of carotenoid-based plumage coloration

The signaling function of carotenoid-based plumage is mainly determined by the concentration of pigments in feathers. For this reason, most studies of the proximate control of coloration focus on processes during and preceding moult. In great tits Parus major, past research demonstrates that carotenoid-based plumage coloration honestly indicates male quality and, thus, may be a sexually selected signal. In this study, we investigate how dirt and preen oil influence the coloration of carotenoid-based feathers in the great tit. We collected six feathers from each individual bird; three feathers served as controls while the remaining three feathers were washed with a chloroform/methanol mixture to remove soil and preen waxes. We assessed plumage coloration using digital photography. This cleaning procedure slightly enhanced ornamentation; the experimentally cleaned feathers expressed hues shifted towards shorter wavelengths and expressed brighter overall coloration than control feathers. This is the first experimental study conducted on wild birds demonstrating that, in addition to pigment concentration, the presence of preen waxes and soils on feathers may contribute to variation in coloration.     

Permanent exposure versus facultative concealment of sexual traits: an experimental study in the house sparrow

In many bird species the cryptic winter plumage is due to the presence of light feather tips that conceal conspicuous colorations. The gradual abrasion of these tips that makes sexual traits visible has been interpreted as a strategy to improve mating success (here referred as the permanent exhibition hypothesis). However, under some circumstances, the maintenance of a full plumage that facultatively enables the bird to exhibit or cover aggression-inducing traits has proved to be advantageous (the coverable badge hypothesis). In a population of house sparrows where black throat patches (here called badges) are used in intrasexual competition, the degree of abrasion of dull feather tips that conceal bright colour early in the breeding season correlated neither with badge size nor with traits indicating morphology and body condition. These results are only in accordance with predictions of the coverable badge hypothesis. In 1992 experimental clipping of badge feather tips, which forced permanent exposure of badges, negatively affected birds with the largest badges in terms of nest acquisition, but an opposite trend was indicated for birds with the smallest badges. In 1993, when more novel birds in the study area were competing for nesting sites than in 1992, only badge size, but not the experimental manipulation, affected nesting success. These results suggest that the loss of the ability to conceal badges was disadvantageous, and more so if most competing individuals had already been resident in the colony in previous years and the larger their badges were. There is a striking contrast between the results reported for this study population, where badges mainly signal fighting ability, and those reported for another house sparrow population, where badges are mainly used in mate choice. In this last population, tip abrasion is advantageous for dominant individuals showing the largest badges. This suggests that the trade-off between conserving versus wearing off the feathers that conceal ornaments could have different optimal resolutions depending on the relative importance of inter and intrasexual selection on indicator traits. Received: 29 September 1995/Accepted after revision: 14 July 1996     

Delayed maturation in plumage colour: Evidence for the female-mimicry hypothesis in the kestrel

Summary In many sexually dichromatic species, young males have female-like plumage during their first potential breeding year. The female-mimicry hypothesis (FMH) supposes that by possessing female-like plumage young males deceive older conspicuous males into believing that they are females, thus reducing competition from adult males. The status-signalling hypothesis (SSH) supposes that adult males can distinguish sex, but postulates that young males reduce competition from adult males by reliably signaling low status with their dull plumage. We tested these hypotheses in the European kestrel (Falco tinnunculus). Female-like young males settled to breed closer to adult males than did other adult males (Figs. 1a, b). By settling near adult males, young males seemed to increase their chance of mating with adult females. Adult female-young male pairs had better reproductive success than yearling-yearling pairs. These results suggest that there is an adaptive value in possessing a female-like plumage colour in the breeding season. To test the FMH, we measured sexual preference of adult males when adult females and young males were simultaneously shown in an aviary. Adult males were unable to recognize sex, because in half the cases they preferred young males (Fig. 3). However, when adult males and females were shown simultaneously, males preferred females (Fig. 2). Our results support the FMH rather than the SSH, because young males successfully deceived older males by their plumage.     

Sexual selection and cuckoldry in a monogamous songbird: implications for sexual selection theory

Sexual selection is generally assumed to be weaker in monogamous than in polygynous animals. Recently, though, extra-pair fertilizations have been hailed as an important force in generating variance in reproductive success among males in socially monogamous species, thereby increasing the intensity of sexual selection. To see if extra-pair copulations contribute to variance in male reproductive success in the house finch (Carpodacus mexicanus), we used DNA fingerprinting to determine the paternity of chicks from 35 nests. This species is a socially monogamous passerine in which plumage brightness serves as a sexually selected indicator of male quality. Out of 119, nestlings 10 (8.3%) were fathered by a male other than the attending male, but cuckoldry occurred randomly with respect to the plumage colouration, size, or age of the attending male. Thus extra-pair fertilizations do not generate variance in male reproductive success with respect to plumage colour. On the other hand, a strongly male-biased sex ratio and asynchronous breeding by females may generate substantial variance in male reproductive success and could explain the evolution of ornamental colouration.     

Diet quality affects an attractive white plumage pattern in dark-eyed juncos (Junco hyemalis)

Sexually selected traits that act as signals of quality often display some degree of condition dependence. In birds, condition dependence of ornamental plumage is often mediated by production costs related to acquisition or allocation of dietary resources. White plumage ornaments, however, have often been assumed to be inexpensive because their production requires neither pigment nor specialized feather structure. In male dark-eyed juncos (Junco hyemalis), the size of a white patch on the tail contributes to attractiveness and mating success. Using captive males, we examined the effects of diet quality on the size and brightness of the tail-white patch. After removing four tail feathers to induce replacement, we maintained subjects on a subsistence (low-protein) or enriched (high-protein) diet while induced feathers grew. Birds that received an enriched diet grew their feathers more quickly and grew larger, brighter white patches. Feather growth rate was positively correlated with the increase in the size of the tail-white patch, a relationship that was stronger in the subsistence diet group. However, within diet treatments, faster-grown feathers were slightly duller. Taken together, these results suggest that variation in diet quality may lead to condition-dependent expression of tail white and that condition dependence may be stronger in more stressful environments. We suggest a mechanism by which increased feather growth rate may lead to an increase in the size of the tail-white patch and discuss potential trade-offs between signal size and brightness.     

Brood sex ratio and male UV ornamentation in blue tits (<Emphasis Type="Italic">Cyanistes caeruleus</Emphasis>): correlational evidence and an experimental test

Sex-allocation theory predicts that females paired to attractive males should bias the brood sex ratio towards male offspring, as these would inherit the attractiveness of their father. We studied sex allocation based on male ornamentation in blue tits. Brood sex ratios varied with male UV coloration in an age-dependent manner. For juvenile males, the proportion of sons increased with increasing UV ornamentation, which is in agreement with previous findings from a Swedish population. However, the relationship between UV ornamentation and brood sex ratio was reversed for adult males, with females paired to less UV-ornamented adult males producing more sons. This pattern fits with the observation that, in our population, less UV-ornamented adult males sire the majority of extra-pair young. To test the causality of the association between brood sex ratio and male coloration, we experimentally manipulated crown colour largely within the natural range. We created two groups of males: one with higher and one with lower UV reflectance, UV(+) and UV(−), respectively. Contrary to our expectations, there was no significant treatment effect. However, in UV(−), but not UV(+) males, the proportion of sons was negatively correlated with male coloration before manipulation. This suggests that the UV(−) treatment caused males that were more UV ornamented to decline more in attractiveness, as shown in a similar experiment in Sweden. However, given that correlational patterns differ between these populations, similarities in experimental results should not be taken as evidence for consistent patterns of adaptive sex allocation in this species.     

Hummingbird with modern feathering: an exceptionally well-preserved Oligocene fossil from southern France

Hummingbirds (Trochilidae) today have an exclusively New World distribution, but their pre-Pleistocene fossil record comes from Europe only. In this study, we describe an exceptionally preserved fossil hummingbird from the early Oligocene of southeastern France. The specimen is articulated, with a completely preserved beak and feathering. Osteological characters allow to identify it as Eurotrochilus sp. This genus is a stem group representative of Trochilidae and was recently described from the early Oligocene of southern Germany. The new fossil reveals that these European Trochilidae were remarkably modern in size, skeletal proportions and the shape of the wing, tail and beak and hyoid bones. These features confirm the early acquisition of the abilities of hovering and nectarivory in hummingbirds, probably before the Oligocene. In several morphological characteristics, they resemble members of the 'true hummingbirds' (subfamily Trochilinae) and differ from hermits (Phaethornithinae). These features, which include a short and square tail and a moderately long, almost straight beak, appear to be primitive within the family Trochilidae.     

Testis size variation in the greenfinch Carduelis chloris : relevance for some recent models of sexual selection

Interspecific evidence that testis size responds to selection caused by sperm competition has been obtained from many taxa. However, little is known about the sources of intraspecific variation in testis size, although such variation may have functional significance. Variation in testis size and asymmetry was studied within and between eight geographically separated (and genetically differentiated) populations of greenfinches Carduelis chloris. The relationships between testis size and plumage brightness (degree of yellowness) and the prevalence of haematozoan infections were also investigated in three of these populations, as they related to the predictions of the immunocompetence handicap hypothesis, and Møller's hypothesis relating directional testis asymmetry to phenotypic quality. There were large differences between populations in testis size, with males from northern populations having larger testes than those from southern populations. Within populations, large testes were associated with larger body size and greater age. When the influence of these factors was removed statistically, males with large testes were more likely to be infected with haematozoan parasites, and had brighter yellow plumage. No evidence was found that directional asymmetry in testis size was related to either of these measures of phenotypic quality. These results are consistent with the hypothesis that males with large testes, while signalling higher phenotypic quality as revealed by increased plumage brightness, also pay a cost in terms of reduced immunocompetence, revealed by the increased probability of infection in these males. That these patterns were similar in three different populations adds further strength to these conclusions. Our results suggest that studying the sources of variation in testis size among individuals can reveal interesting processes in sexual selection.     

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.     

Plumage characteristics, reproductive investment and assortative mating in tree swallows Tachycineta bicolor

Elaborate ornamental plumage has been associated with various measures of individual quality in many species of birds. Male plumage characteristics, which have been relatively well studied, have been shown to reflect past reproductive investment, as well as the potential for reproductive investment in the current breeding attempt. In contrast, the signalling functions of female traits remain largely unexplored. In this study, we investigated the relationship between plumage attributes of breeding adult tree swallows and past reproductive investment, current reproductive investment and social mate pairing strategy. Both males and older females possess metallic green to metallic blue iridescent plumage on their dorsal surface, making this a suitable species for this type of investigation. We did not find any effects of past reproductive investment and success on the plumage attributes of returning breeders. In contrast, female plumage hue covaried with fledging success, and female plumage brightness was positively associated with mean clutch egg mass. In addition, we found that social pairs mated assortatively with respect to plumage brightness. We argue that since plumage characteristics vary with age in both male and female tree swallows, plumage attributes in this species are indicative of breeding experience and may be honest signals of quality. Positive assortative pairing could be the result of mutual mate choice or intra-sexual competition for nest sites by both males and females.