Correlates of timing of spring migration in birds: a comparative study of trans-Saharan migrants

The evolution of migratory strategies in birds is likely to have been influenced by ecological as well as socio-sexual factors in both wintering and breeding areas. In this comparative study, we analysed timing of spring passage of 38 long-distance migratory bird species that winter south of the Sahara desert and breed in Europe, in relation to wintering and breeding latitudes, moult strategy, nesting site (open vs. cavity), and sexual dimorphism in size and coloration, which may reflect intensity of sexual selection. We employed a large data set consisting of more than 190 000 individuals ringed during spring migration in the Mediterranean Sea. We found that the species that migrated earlier were those wintering farther north, nesting in cavities and showing the largest degree of sexual size dimorphism (SSD). However, sexual dichromatism was not related to migration date. Among passerine species, moulting wing-feathers in Africa delayed migration. We found no support for the energetic constraint hypothesis, which proposes that early arrival selects for large male size, since early arriving species were not larger than late arriving ones. Thus, the observed associations suggest that variation in migration schedules at the interspecific level may have evolved in relation to ecological factors and SSD, possibly reflecting the intensity of mating competition.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 199–210.     

Parasite-mediated sexual selection and species divergence in Lake Victoria cichlid fish

We investigate the role of parasite-mediated sexual selection in the divergence of two species of Lake Victoria cichlids. Pundamilia pundamilia and Pundamilia nyererei represent a common pattern of male nuptial colour divergence between haplochromine sister species: metallic grey–blue in P. pundamilia and bright yellow and red in P. nyererei . Female mating preferences for different male colours maintain the genetic and phenotypic differentiation of the two species in clear water. Previous work indicated that the red coloration of P. nyererei males, which is subject to directional sexual selection, may be a carotenoid-dependent signal of parasite infestation rate. In the present study, we find a parallel result for P. pundamilia : bright blue males are infected with fewer species of parasites. We also find that parasite infestation rates differ quantitatively between the two species in a way that is consistent with species differences in diet and microhabitat. We conclude that parasite-mediated sexual selection may have contributed to the divergence of female mating preferences between P. pundamilia and P. nyererei , and may currently strengthen reproductive isolation between these species.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 94 , 53–60.     

Sexual, seasonal, and environmental variation in plasma carotenoids in great tits, Parus major

In many birds, carotenoids have dual functions as irreversible plumage pigments and as physiologically essential vitamins and antioxidants. They must be obtained through the diet and may therefore be a limiting resource, a constraint that is likely to vary with factors such as sex, habitat, and time of year. In the present study, we investigated signs of carotenoid limitation in great tits, Parus major , in relation to sex, season, year, and within an urban versus a rural habitat. The two main carotenoids, lutein and zeaxanthin, were analysed by high-performance liquid chromatography in the plasma and in the yellow carotenoid-based breast feathers. We found that plasma carotenoid concentrations were significantly influenced by sex, season, and year, but not by urban versus rural habitat. At moult, plasma concentration was positively correlated with feather pigmentation, independent of body condition and sex. During the breeding season, however, circulating carotenoid concentrations were negatively related to the feather pigmentation (i.e. from previous autumn moult). We suggest that great tits are carotenoid deprived before leaf emergence, and that carotenoid utilization and limitations are sex-specific, but that there are neither any obvious honesty-maintaining costs of pigmentation, nor any fitness consequences of the colour variation.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 521–527.     

Carotenoids bolster immunity during moult in a wild songbird with sexually selected plumage coloration

Carotenoid‐based colours in animals are valuable models for testing theories of sexual selection and life‐history trade‐offs because the pigments used in coloration are chemically tractable in the diet and in the body, where they serve multiple purposes (e.g. health enhancement, photoprotection). An important assumption underlying the hypothesized signalling value of carotenoid coloration is that there is a trade‐off in carotenoid pigment allocation, such that not all individuals can meet the physiological/morphological demands for carotenoids (i.e. carotenoids are limited) and that only those who have abundant supplies or fewer demands become the most colourful. Studies of carotenoid trade‐offs in colourful animals have been limited largely to domesticated species, which may have undergone artificial selection that changed the historical/natural immunomodulatory roles of carotenoids, to young animals lacking carotenoid‐based signals or to species displaying carotenoid‐based skin and bare parts. We studied the health benefits of carotenoids during moult in house finches (Carpodacus mexicanus), which display sexually selected, carotenoid‐based plumage coloration. We manipulated dietary carotenoid availability during both winter (nonmoult) and autumn (moult) in captive males and females and found that carotenoid‐supplemented birds mounted stronger immune responses (to phytohemagglutinin injection and to a bacterial inoculation in blood) than control birds only during moult. This study provides experimental, seasonal support for a fundamental tenet of Lozano's ‘carotenoid trade‐off’ hypothesis and adds to a growing list of animal species that benefit immunologically from ingesting higher dietary carotenoid levels. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 560–572.     

Effects of genetics and light environment on colour expression in threespine sticklebacks

The genetic basis of traits that are under sexual selection and that are involved in recognizing conspecific mates is poorly known, even in systems in which the phenotypic basis of these traits has been well studied. In the present study, we investigate genetic and environmental influences on nuptial colour, which plays important roles in sexual selection and sexual isolation in species pairs of limnetic and benthic threespine sticklebacks ( Gasterosteus aculeatus species complex). Previous work demonstrated that colour differences among species correlate to differences in the ambient light prevalent in their mating habitat. Red fish are found in clear water and black fish in red-shifted habitats. We used a paternal half-sib split-clutch design to investigate the genetic and environmental basis of nuptial colour. We found genetic differences between a red and a black stickleback population in the expression of both red and black nuptial colour. In addition, the light environment influenced colour expression, and genotype by environment interactions were also present. We found evidence for both phenotypic and genetic correlations between our colour traits; some of these correlations are in opposite directions for our red and black populations. These results suggest that both genetic change and phenotypic plasticity underlie the correlation of male colour with light environment.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 663–673.     

Rapid divergence of social signal coloration across the White Sands ecotone for three lizard species under strong natural selection

Animal social signals are important for population recognition, communication, and mate choice. Although natural selection often favours cryptic coloration, sexual selection can underlie patterns of coloration that function in inter- or intrasexual communication. We compared social signal coloration of three lizard species across a substrate colour ecotone in New Mexico. These species exhibit cryptic blanched dorsal coloration on the gypsum dunes of White Sands and dark coloration on the surrounding desert soils. We detected corresponding population divergence in colour used for intra- ( Aspidoscelis inornata , Sceloporus undulatus ) or inter- ( Holbrookia maculata ) sexual signalling. Although the magnitude and direction of change in coloration varied among taxa, differences in hue and chroma accounted for more variation in social coloration than for dorsal coloration. The relative conspicuousness of social signals also varied across the ecotone. We discuss the possibilities that divergent signalling colours in this system are the result of: (1) stochastic processes, (2) direct selection, and/or (3) a correlated response to natural selection on dorsal coloration.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 243–255.     

Testing the role of coadapted genes versus bet-hedging for mating strategies in colour polymorphic pygmy grasshoppers

Recent investigations of mate choice indicate that the genetic effect of sires on offspring fitness may depend on the interaction between maternal and paternal genotypes and the environmental conditions experienced by the offspring. Alternative colour morphs of the pygmy grasshopper, Tetrix subulata , represent ecological strategies that differ in body size, life history, thermoregulatory behaviour, and habitat selection. The hypothesis that selection promotes behaviours maintaining coadapted gene complexes predicts individuals to mate assortatively with respect to colour morph. On the other hand, the bet-hedging hypothesis predicts that the temporal variability of the environment inhabited by these animals may select for disassortative mating behaviour resulting in heterogeneous offspring. To distinguish between these competing hypotheses, we investigated mating behaviours using dual-choice experiments. Our results were not in agreement with the prediction of assortative mating but suggest instead that matings were random with regard to colour morph. Polyandry was common, and females mated with the second male regardless of whether the first mating was assortative or disassortative. Polyandry also was equally frequent among females in triads in which the two males belonged to different colour morphs as in triads where both males belonged to the same colour morph. A field experiment confirmed that polyandry occurred also among free-ranging individuals, and uncovered variation in mating success among male colour morphs, probably due to indirect effects of coloration on activity or habitat use. The consequences of this random and polyandrous mating strategy for the evolutionary dynamics of the colour polymorphism remain to be explored.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 491–499.     

Condition and brightness of structural blue‐green: motmot tail‐racket brightness is related to speed of feather growth in males,but not in females

Coloration plays an important role in sexual and social communication, and in many avian species both males and females maintain elaborate colours. Recent research has provided strong support for the hypothesis that elaborate female traits can be maintained by sexual or social selection; however, most research on female ornamentation has focused on pigment‐based colours, and less is known about how structural colours are maintained. Both sexes of the turquoise‐browed motmot (Eumomota superciliosa) have a blue‐green racket‐tipped tail, and it remains unknown if tail coloration serves as a sexual or social signal in one or both sexes. Here, we describe sexual dichromatism in the blue‐green portion of the tail racket, and we test for a relationship between coloration and condition, as indicated by growth bars. Tail colour of both sexes has a similar spectral shape, and there is significant, although moderate, sexual dichromatism: males are brighter than females, and males have marginally greater blue‐green saturation than females. The length of feather grown per day is positively related to overall feather brightness, but this relationship is only present in males. The relationship between male coloration and condition suggests that tail colour has the potential to convey information about individual quality during mate choice or contest competition. The lack of a similar relationship in females suggests that female tail colour does not convey the same condition‐dependent information that we suggest may be reflected by male colour. Female tail colour may therefore reflect other aspects of condition, be involved in other (non‐condition‐dependent) forms of communication, or be expressed as a non‐functional byproduct of genetic correlation between the sexes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 673–681.     

Colour composition of nest lining feathers affects hatching success of barn swallows,Hirundo rustica (Passeriformes: Hirundinidae)

Many bird species use feathers as lining material, and its functionality has traditionally been linked to nest insulation. However, nest lining feathers may also influence nest detection by predators, differentially affect reproductive investment of mates in a post‐mating sexual selection process, and affect the bacterial community of the nest environment. Most of these functions of nest lining feathers could affect hatching success, but the effect might vary depending on feather coloration (i.e. pigmented versus white feathers). This would be the case if coloration is related to: (1) thermoregulatory properties; (2) attractiveness of feathers in the nest for mates; (3) eggshell bacterial density. All of these hypothetical scenarios predict that feathers of different colours would differentially affect the hatching success of birds, and that birds should preferentially choose the most beneficial feather colour for lining their nests. Results from two different experiments performed with a population of Danish barn swallow, Hirundo rustica, were in accordance with these predictions. First, H. rustica preferentially selected white experimentally offered feathers for lining their nests. Second, the experimental manipulation of the feather colour composition of nests of H. rustica had a significant effect on hatching success. Experimental nests with more white feathers added at the beginning of incubation had a lower probability of hatching failures, suggesting differential beneficial effects of lining nests with feathers of this colour. We discuss the relative importance of hypothetical functional scenarios that predicted the detected associations, including those related to sexual selection or to the community of microorganisms associated with feathers of different colours. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 67–74.     

Large‐scale geographic variation in iridescent structural ornaments of a long‐distance migratory bird

Iridescent colours produced during moult likely play an important role in pair formation in birds. We sought to quantify geographic variation in such colouration in a duck species, Eurasian teal Anas crecca, in winter (when mating occurs) to evaluate whether this variation reflects birds’ breeding origins or differential individual migration strategies in both males and females. We combined information on feather production region and individual attributes (body size, sex and age) of Eurasian teal from 82 wintering sites in France. Feather production region (moult site or natal origin) was inferred using feather deuterium values (δD_f). We performed spectral measurements to evaluate speculum colour and brightness contrasts for 1052 teal collected over four years. Colouration differed strongly among wintering regions, with birds wintering in eastern France exhibiting higher colour contrast than those wintering in the west. Body size and colouration were positively related. There were no differences in cohort‐specific δD_f values between separate wintering regions in France, indicating that within a winter quarter teal originated from areas across the entire breeding range. Overall, patterns of spatial variation in feather colouration were related most closely to body size which was consistent with predictions of a differential migration hypothesis, with larger and more colour‐contrasting birds wintering closer to their breeding grounds. Because moult speed is also known to affect colour production, early breeders or individuals that skipped reproduction may have invested more or earlier in their feather quality to gain potential advantages in monopolizing future mates.     

Migratory behaviour affects the trade-off between feather growth rate and feather quality in a passerine bird

Migratory birds have less time for moulting than sedentary birds, which may force them to produce their feathers faster at the expense of reducing feather quality. However, the effects of migration on the trade-off between moult speed and plumage quality remain to be studied in natural populations. We analysed the relationship between growth rate and quality of individual feathers, taking advantage of natural variation between migratory and sedentary populations of blackcaps Sylvia atricapilla . As predicted by life-history theory, individual blackcaps showed variable individual quality, which was revealed by positive correlations between feather growth rate and feather mass within populations. However, migrants grew up their feathers faster, producing lighter feathers than sedentary blackcaps. These results support the idea that feather growth rate and feather quality are traded against each other in blackcaps. Such a trade-off is apparently caused by different selection associated to migratory and sedentary life styles, which opens new insights into the diversification of moult patterns in birds.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 98–105.     

Size-dependent mating strategies and the risk of cannibalism

The role of sexual selection in determining the nature and direction of sexual size dimorphism may depend upon the timing of sexual selection, and this may also influence the variation in male size. For example, selection through sperm competition favours smaller males in the highly sexually size dimorphic orb-weaving spider Nephila edulis , whereas larger males are better able to exclude their smaller rivals from the central hub of the web where mating takes place. We investigate experimentally the role of body size and hub tenure in determining male fertilization success when males of different sizes compete for a single female over a 24-h period that includes a period of darkness. Our results confirm that small and large males obtain similar paternity share but that, in contrast with previous studies, hub tenure does not translate into greater paternity share. Unexpectedly, smaller males are at greater risk of postmating sexual cannibalism than larger males, suggesting that natural selection through sexual cannibalism may place a lower limit on male size.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 355–363.     

Population differences in the structure and coloration of great tit contour feathers

Contour feathers cover most of the avian body and play critical roles in insulation, social communication, aerodynamics, and water repellency. Feather production is costly and the development of the optimum characteristics for each function may be constrained by limited resources or time, and possibly also lead to trade‐offs among the different characteristics. Populations exposed to different environmental conditions may face different selective pressures, resulting in differences in feather structure and coloration, particularly in species with large geographical distributions. Three resident populations of great tit Parus major L. from different latitudes differed in feather structure and coloration. Individuals from the central population exhibited less dense and longer contour feathers, with a higher proportion of plumulaceous barbs than either northern or southern birds, which did not differ in their feather structure. Ultraviolet reflectance and brightness of the yellow of the contour feathers of the breast was higher for the southern than for the northern population. Birds with greener plumage (higher hue) had less dense but longer feathers, independently of the population of origin. Differences in feather structure across populations appear to be unrelated to the contour feather colour characteristics except for hue. Nutritional and time constraints during molt might explain the pattern of feather structure, whereas varying sexual selection pressure might underlie the coloration patterns observed. Our results suggest that different selective pressures or constraints shape contour feather traits in populations exposed to varying environmental conditions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 82–91.     

Population differences in density and resource allocation of ornamental tail feathers in the barn swallow

Many organisms show well‐defined latitudinal clines in morphology, which appear to be caused by spatially varying natural selection, resulting in different optimal phenotypes in each location. Such spatial variability raises an interesting question, with different prospects for the action of sexual selection on characters that have a dual purpose, such as locomotion and sexual attraction. The outermost tail feathers of barn swallows (Hirundo rustica) represent one such character, and their evolution has been a classic model subject to intense debate. In the present study, we examined individuals from four European populations to analyze geographical variation in the length and mass of tail feathers in relation to body size and wing size. Tail feather length differed between sexes and populations, and such variation was a result of the effects of natural selection, acting through differences in body size and wing size, as well as the effects of sexual selection that favours longer tails. The extra enlargement of the tail promoted by sexual selection (i.e. beyond the natural selection optimum) could be achieved by increasing investment in ornaments, and by modifying feather structure to produce longer feathers of lower density. These two separate processes accounting for the production of longer and more costly tail feathers and less dense feathers, respectively, are consistent with the hypothesis that both Zahavian and Fisherian mechanisms may be involved in the evolution of the long tails of male barn swallows. We hypothesize that the strength of sexual selection increases with latitude because of the need for rapid mating as a result of the short duration of the breeding season at high latitudes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 925–936.     

Winter Plumage Coloration in Male American Goldfinches: Do Reduced Ornaments Serve Signaling Functions in the Non‐Breeding Season?

The signaling role of sexual ornaments that are displayed during the mating season is well known for many species, but dimorphisms that occur in the non‐breeding season have received much less attention, particularly when individuals only partially express their breeding condition during reproductively inactive periods. I assessed variation in the expression of colorful breeding and non‐breeding plumages in male American goldfinches (Carduelis tristis), a species in which males molt out of their colorful breeding ornaments in the fall but still display reduced carotenoid‐ and melanin‐based sexual dichromatism during the winter. I found that variability in the saturation of carotenoid‐based plumage pigmentation did not differ significantly between the breeding and non‐breeding seasons. Moreover, the area of melanin coloration in the cap was more variable in winter than when it is fully displayed during breeding. I also detected a significant positive correlation between the extent of melanin coloration during the winter and the saturation of non‐breeding carotenoid‐based plumage. Because of such variation in and correlated expression of these two color ornaments in winter, it is conceivable that male goldfinches display these hints of non‐breeding coloration for use as conspecific social or sexual signals. Natural selection pressures like predation and energetic demands are traditionally thought of as factors that restrict sex ornaments to breeding times alone, but this should not preclude animals from simply reducing their exaggerated features during winter and finding an expression optimum that balances signal costs and value. Such ‘remnant’ winter ornaments might be expected to evolve not only in animals that live in large non‐breeding groups (e.g. status‐signaling systems) but also in those where mates begin associating before breeding onset.     

A sexual conflict in collared flycatchers,Ficedula albicollis: early male moult reduces female fitness

A sexual conflict over levels of parental care occurs in most animals with biparental care, and studies of sexual differences in levels of parental care have usually focused on its intra-annual fitness consequences. We investigated inter-annual fitness consequences of a sexual difference in timing of feather replacement (moult) in collared flycatchers (Ficedula albicollis). In this study, males overlapped reproduction and moult more often than females, they also initiated their moult at an earlier stage of breeding than females. Females mated to males with a moult-breeding overlap had significantly lowered survival chances than females mated with males initiating moult after breeding. Furthermore, females mated with moulting males risked a lowered future fecundity in terms of a delayed start to breeding in the following season. However, early moulting males achieved a similar reproductive success as males initiating moult after breeding. Likewise, male survival probability to the following breeding season did not differ between early and late moulting individuals, nor was there any evidence that males gained or lost in future mating advantages by moulting early. These results show not only that a sexual conflict over timing of moult may operate, but also that it can impose severe fitness consequences, in terms of reduced future fecundity and survival probability, upon the ''losing'' sex.     

Visual conditions and habitat shape the coloration of the Eurasian perch (Perca fluviatilis L.): a trade‐off between camouflage and communication?

In theory, selection for effective camouflage (i.e. dull coloration) in fish should be strongest when the conditions for visual predation are most favourable, such as in structurally simple pelagic habitats. By contrast, in more sheltered (e.g. littoral) habitats, selection may favour effective intra‐specific communication (i.e. bright coloration) (at the expense of crypsis). Poor transparency, as in highly humic waters, should constrain colour adaptations. We investigated phenotypic variation in body coloration of Eurasian perch (Perca fluviatilis L.) in littoral and pelagic habitats of four humic boreal lakes. Perch from the most transparent lake had the lightest and less coloured belly and perch were more colourful in the littoral habitats than in the pelagic areas, with the pattern being clearest in the most transparent lake. In addition, perch in the most transparent lake exhibited sexual dichromatism, with males having a more colourful belly than the females, whereas no indications of sexual dichromatism were found in more humic lakes. Moreover, in the most transparent lake, the condition of fish correlated with bright belly coloration in the littoral, but with dull belly coloration in the pelagic habitat. The results obtained in the present study suggest that selection on perch coloration may differ between lakes as a result of visual properties of the water, and within lakes as a result of divergent selection for camouflage and communication in pelagic and littoral habitats. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 47–59.     

Consistent annual schedules in a migratory shorebird

Many migratory birds start prebreeding moult and premigratory fuelling some months before the breeding season and face severe time constraints, while travelling up to 15,000 km between non-breeding and breeding grounds. Shorebirds typically leave Southern Hemisphere non-breeding areas over a 3-4 week period, but whether they benefit from interannually consistent timing of departure is unknown. Here, I show that individual bar-tailed godwits (Limosa limosa baueri) from New Zealand are highly consistent in their migratory scheduling. Most birds left within the same week each year (between-year repeatability, r, of 0.83) and adult males, which moult into a bright breeding plumage, were also highly repeatable in the extent of their prebreeding moult (r=0.86). This is consistent with the hypothesis that birds have individually optimized migration schedules. Within adult males, but not females, smaller birds tended to migrate earlier than large birds. Whether this reflects differences in size-related migration speed, optimal breeding time at different sites or size-related natural or sexual selection pressures, remains unknown.     

Preen waxes do not protect carotenoid plumage from bleaching by sunlight

The plumage coloration of wild birds often changes during the breeding season. One of the possible reasons for this is that sunlight, and particularly ultraviolet (UV) wavelengths, degrades the pigments responsible for plumage coloration. It has been suggested that birds may apply preen wax to feathers to protect feathers from bleaching. This hypothesis is tested by exposing carotenoid-based breast feathers of Great Tits to ambient light, light filtered to exclude UV and darkness. Preen waxes were experimentally removed from feather samples and the effect of light on coloration of treatment and control feathers compared. Ambient light had an effect on feather colour but preen wax did not. Feathers exposed to sun gradually became less saturated and hues shifted towards shorter wavelengths. This was not apparent in control feathers kept in darkness. Feathers exposed to full-spectra sunlight faded more than those that were kept in light with UV wavelengths removed. There was a decrease in brightness of feathers in both experimental and control groups, which was assumed to be an effect of dirt accumulation. This experiment confirmed earlier suspicions regarding the detrimental effects of UV irradiation on carotenoid-based coloration of avian feathers but failed to show any protective function of preen waxes. The possible consequences of these mechanisms of colour change for birds with regard to mating strategies are discussed.     

Environmental-induced acquisition of nuptial plumage expression: a role of denaturation of feather carotenoproteins?

Several avian species show a bright carotenoid-based coloration during spring and following a period of duller coloration during the previous winter, despite carotenoids presumably being fully deposited in feathers during the autumn moult. Carotenoid-based breast feathers of male linnets (Carduelis cannabina) increased in hue (redness), saturation and brightness after exposing them to outdoor conditions from winter to spring. This represents the first experimental evidence showing that carotenoid-based plumage coloration may increase towards a colourful expression due to biotic or abiotic environmental factors acting directly on full-grown feathers when carotenoids may be fully functional. Sunlight ultraviolet (UV) irradiation was hypothesized to denature keratin and other proteins that might protect pigments from degradation by this and other environmental factors, suggesting that sunlight UV irradiation is a major factor in the colour increase from winter to spring. Feather proteins and other binding molecules, if existing in the follicles, may be linked to carotenoids since their deposition into feathers to protect colourful features of associated carotenoids during the non-breeding season when its main signalling function may be relaxed. Progress towards uncovering the significance of concealment and subsequent display of colour expression should consider the potential binding and protecting nature of feather proteins associated with carotenoids.