Covariation of sexual dichromatism and plumage colours in lekking and non-lekking birds: A comparative analysis

The extreme polygyny expressed by male lekking birds leads to the expectation that sexual dimorphism should be greater in lekkers than related non-lekkers. However, evidence for this association is weak, and many lekkers are actually monomorphic in size and plumage. To better understand the kinds of plumages associated with lekking, I characterized plumage variation for combinations of sexual dichromatism and colourfulness-and-conspicuousness (COCO) among lekking and related non-lekking birds. Compared in this way, the plumages of lekkers and non-lekkers differ dramatically for both sexes. Correlations between sexual dichromatism and COCO for phylogenetically independent contrasts are significant for male lekkers (positive) and female non-lekkers (negative), but not for female lekkers or male non-lekkers. Moreover, the total number of character–state combinations, and multivariate measures of variability, are greater in non-lekkers than lekkers.The characteristic plumages of lekkers (duller monochromatic, brighter dichromatic and intermediate between these extremes) comprise just a subset of those observed among non-lekkers, and exclude extremely dull dichromatic and extremely bright monochromatic plumages. I suggest that predation, and foraging behaviours compatible with lekking, may restrict plumage variation among lekkers. Thus ecological rather than overt sexual characteristics may explain monomorphism in birds under intense mate competition, as well as the paradox of strong female mate preferences on leks, where males appear to contribute only sperm to female reproductive efforts.     

Ornament and body size variation and their measurement in natural populations

Measurement of intrapopulation variation in secondary sexual traits is a priority in the testing of sexual selection models. However, it is important to take care in the choice of materials and delimitation of populations. The use of museum skins to study variation in male tail ornaments may substantially underrepresent the real degree of intrapopulation variation. Data from live animals in specific areas provide more realistic estimates, and should be used whenever possible. I use as an example field data on male ornament length and body size in Vidua macroura (Aves: Ploceidae), a promiscuous, parasitic African finch with elongated tail plumes. Individual males differ in the timing and rate of ornament growth, and females are therefore faced with a large degree of phenotypic variation in male ornament size, even though genetic variation may not be great. By correcting for seasonal variation in the ornament lengths of males caught at different times, I show that mid-season coefficients of variation in ornament length of breeding males in two populations are as high as 18% and 55%. By contrast, tarsus, wing and unornamented tail lengths of the same males vary from 2 to 4%.     

Carotenoids, colour and conservation in an endangered passerine, the hihi or stitchbird (Notiomystis cincta)

Carotenoids are essential dietary components utilized not only in pigmentation but also as immuno-stimulants and antioxidants. Reduced availability can have consequences on individual health and survival, thus making carotenoids a good indicator of environmental stress. We compared carotenoid profiles and plumage colour characteristics of an endangered passerine species in New Zealand, between its remnant island source population and two reintroduced island populations. Circulating carotenoids were predominantly lutein (mean of 82.2%) and zeaxanthin (mean of 14.8%), and these were the major carotenoids present as yellow pigments in the males' plumage. There were clear differences in total carotenoid concentrations and plumage colour among the three populations. Circulating carotenoid concentration was significantly higher in one of the reintroduced populations, and the yellow plumage of males was significantly higher in both reintroduced populations in comparison with the remnant population (reflected as a significant increase in hue). Understanding how these differences arise may be of importance to this species given the health benefits carotenoids impart and our ability to select plant species containing these compounds or artificially supplement them.     

High brood patch temperature of less colourful,less pheomelanic female Barn Swallows Hirundo rustica

In birds, even a minor difference in egg temperature (1–1.5 °C) has been shown to affect the fitness of offspring by changing hatching success, incubation period and nestling quality. Female, but not male, passerines develop brood patches. Thus if there are traits, such as plumage ornamentation, that indicate optimal egg temperature, males should pair with females that exhibit those traits. However, no study has yet investigated the relationship between female brood patch temperature, which would directly affect egg temperature, and female plumage ornamentation. In this study, we examined the surface temperature of female brood patches during nocturnal incubation and examined its relationship with female plumage ornaments in Asian Barn Swallows Hirundo rustica gutturalis. After controlling for ambient air temperature, brood patch temperature was negatively associated with colour saturation of the female throat patch. No other female ornaments, such as tail‐length, white tail spots or throat patch size, predicted brood patch temperature. When oral (mouth) temperature was statistically controlled, females with less colourful throats and longer tails showed higher brood patch temperature, indicating that these females had hotter brood patches in relation to the temperature of other body parts. Furthermore, we found a negative relationship between pheomelanin pigmentation and brood patch temperature after controlling for ambient air temperature or oral temperature. To our knowledge, this is the first study to show that female ornaments can predict the absolute/relative thermal investment in brood patches. This relationship, together with other aspects of female quality, may affect male mate preference and female ornamentation.     

Mutual ornamentation and the parental behaviour of male and female Collared Flycatchers Ficedula albicollis during incubation

One of the benefits of mate choice based on sexually selected traits is the greater investment of more ornamented individuals in parental care. The choosy individual can also adjust its parental investment to the sexual signals of its partner. Incubation is an important stage of avian reproduction, but the relationship between behaviour during incubation and mutual ornamentation is unclear. Studying a population of Collared Flycatchers Ficedula albicollis, we monitored the behaviour of both sexes during incubation in relation to their own and their partner's plumage traits, including plumage‐level reflectance attributes and white patch sizes. There was a marginally positive relationship between male feeding rate during incubation and female incubation rate. Female but not male behavioural traits were associated with the laying date of the first egg and clutch size. The behaviour of the two sexes jointly determined the relative hatching speed of clutches and the hatching success of eggs. Females with larger white wing patches spent less time incubating eggs and left the nestbox more frequently. Males with larger white wing patches fed females less frequently, whereas males with brighter white plumage areas visited the nestbox more regularly without feeding. Females tended to leave the nest less often when mated to males with larger wing patches, and females spent less time incubating when males had more UV chromatic plumage. The behaviour of both partners during incubation therefore predicted hatching patterns and was correlated with their own and sometimes with their partner's plumage ornamentation. These results call for further studies of mutual ornamentation and reproductive effort during incubation.     

Females mate with males with diminished pheomelanin‐based coloration in the Eurasian nuthatch Sitta europaea

Sexual selection can drive the evolution of phenotypic traits because of female preferences for exaggerated trait expression in males. Sexual selection can also lead to the evolutionary loss of traits, a process to which female preferences for diminished male trait expression are hypothesized to contribute. However, empirical evidence of female preferences for diminished male traits is virtually lacking. Eurasian nuthatches Sitta europaea provide an opportunity to test this possibility, as a chestnut flank patch produced by the pigment pheomelanin is present since the first plumage of these birds and its color is more intense in nestlings in poor condition in our study population. It has been proposed that developing birds in poor condition may increase their production of pheomelanin as a detoxifying strategy. Female nuthatches may thus prefer mating with males showing flank feathers of diminished color, as this could indicate that males experienced good conditions early in development, which can positively affect the fitness of future generations. Here we show results according with this prediction in a wild population of Eurasian nuthatches, as adult males with lighter chestnut feathers paired earlier in the season, while chestnut coloration had no effect on female mating success. Chestnut color expression was not affected by the body condition of birds, suggesting that females obtain information on the body condition in early life of their potential mates and not on their current body condition. This constitutes one of the few examples of females mating with males showing diminished traits and provides the only explanation so far by which this process can occur.     

Variation in melanin pigmentation of a sexually selected plumage trait and its adaptive value in the Common Snipe Gallinago gallinago

There is increasing evidence that melanin‐based plumage coloration correlates with different components of fitness and that it may act as a social or sexual signal of individual quality. We analysed variation in melanin pigmentation in the outermost tail feathers of the Common Snipe Gallinago gallinago. During courtship flights, male Snipe use their outermost tail feathers to generate a drumming sound, which plays a role in territory establishment and mate choice. As the outermost tail feathers are displayed to females during these flights, we predicted that conspicuous variation in their rusty‐brown (pheomelanin‐based) coloration may act as an honest signal of individual quality. To test this prediction, we spectrophotometrically measured brightness (an indicator of total melanin content) and red chroma (an indicator of pheomelanin content) of the outermost tail feathers in 180 juvenile and adult Common Snipe. An age‐related decline in feather brightness was found exclusively in females, suggesting that melanization could have evolved by natural selection to camouflage incubating birds. In both sexes, brightness of the tail feathers was inversely correlated with their structural quality (as measured with mass–length residuals), suggesting that melanization could increase mechanical properties of feathers and, in males, enhance the quality of courtship sonation. Red chroma positively correlated with total plasma protein concentration, supporting our prediction that pheomelanin pigmentation of tail feathers may act as an honest signal of condition. Our study indicated that variation in the melanin‐based coloration of the outermost tail feathers in the Common Snipe could have evolved as a result of several different selection pressures and it emphasizes the complexity of the processes that underlie the evolution of melanin‐based plumage coloration in birds.     

Do trace metals select for darker birds in urban areas? An experimental exposure to lead and zinc

Trace metals from anthropogenic activities are involved in numerous health impairments and may therefore select for detoxification mechanisms or a higher tolerance. Melanin, responsible for the black and red colourations of teguments, plays a role in metal ion chelation and its synthesis is positively linked to immunity, antioxidant capacity and stress resistance due to pleiotropic effects. Therefore, we expected darker birds to (1) store higher amounts of metals in their feathers, (2) maintain lower metal concentrations in blood and (3) suffer less from metal exposure. We exposed feral pigeons (Columba livia) exhibiting various plumage darkness levels to low, but chronic, concentrations of zinc and/or lead, two of the most abundant metals in urban areas. First, we found negative and positive effects of lead and zinc, respectively, on birds' condition and reproductive parameters. Then, we observed positive relationships between plumage darkness and both zinc and lead concentrations in feathers. Interestingly, though darker adults did not maintain lower metal concentrations in blood and did not have higher fitness parameters, darker juveniles exhibited a higher survival rate than paler ones when exposed to lead. Our results show that melanin‐based plumage colouration does modulate lead effects on birds' fitness parameters but that the relationship between metals, melanin, and fitness is more complex than expected and thus stress the need for more studies.     

Blood parasites, leucocytes and plumage brightness in the Cirl Bunting, Emberiza cirlus

1. Although a female preference for pairing with brightly plumaged males has been reported in many species, the reasons for this choice are not fully understood.
2. Parasites have been proposed as playing an important role in shaping these preferences because, by pairing with brightly coloured individuals, females can obtain parasite-free and/or more healthy mates.
3. In this paper one of the predictions of this hypothesis is tested, namely a higher health level in brightly coloured individuals, by analysing the relationships among blood parasites, leucocyte levels and plumage brightness in the Cirl Bunting, Emberiza cirlus .
4. Two species of blood parasites were detected. Whereas a lower body condition was detected in individuals infected by Leucocytozoon cambournaci, no such differences were associated with Plasmodium relictum infections. Infected individuals showed higher total leucocyte counts than non-infected individuals.
5. Colour intensity of carotenoid derived colorations was negatively correlated to the relative proportion of lymphocytes and positively correlated to the relative presence of heterophils. Furthermore, the size of yellow feathered areas was positively correlated with the absolute number of leucocytes and the relative presence of heterophils. Only some of these relationships were found for non-carotenoid derived traits.
6. These results suggest that male plumage yellow coloration in the Cirl Bunting is a reliable indicator of health status and supports the hypothesis that females obtain more parasite-free mates if they pair with brightly coloured individuals.     

Red coloration of male northern cardinals correlates with mate quality and territory quality

I investigated how mate quality and territory quality influencean extravagant ornament in a socially monogamous species thatdefends multipurpose territories. Northern cardinals (Cardinaliscardinalis) are a highly dichromatic, socially monogamous species,and males are a brilliant red. I conducted a 3-year field studyof northern cardinals and found that redder males produced moreoffspring in a breeding season. Two selective factors mediatedthis fitness gain. Redder males were paired with earlier breedingfemales, an established measure of mate quality in birds. Second,redder males obtained territories of higher quality, as measuredby vegetation density. Interactions among these factors werealso important in explaining variance in male reproductive success.Multivariate analysis indicated that earlier breeding increasedreproductive success independent of territory quality. In turn,territory quality contributed to male reproductive success throughits effect on nest survival and possibly through its role in attractingan earlier breeding female.