Calls,colours, shape,and genes: a multi‐trait approach to the study of geographic variation in the Amazonian frog Allobates femoralis

Evolutionary divergence in behavioural traits related to mating may represent the initial stage of speciation. Direct selective forces are usually invoked to explain divergence in mate‐recognition traits, often neglecting a role for neutral processes or concomitant differentiation in ecological traits. We adopted a multi‐trait approach to obtain a deeper understanding of the mechanisms behind allopatric divergence in the Amazonian frog, Allobates femoralis. We tested the null hypothesis that geographic distance between populations correlates with genetic and phenotypic divergence, and compared divergence between mate‐recognition (acoustic) and ecological (coloration, body‐shape) traits. We quantified geographic variation in 39 phenotypic traits and a mitochondrial DNA marker among 125 individuals representing eight populations. Geographic variation in acoustic traits was pronounced and tracked the spatial genetic variation, which appeared to be neutral. Thus, the evolution of acoustic traits tracked the shared history of the populations, which is unexpected for pan‐Amazonian taxa or for mate‐recognition traits. Divergence in coloration appeared uncorrelated with genetic distance, and might be partly attributed to local selective pressures, and perhaps to Batesian mimicry. Divergence in body‐shape traits was low. The results obtained depict a complex evolutionary scenario and emphasize the importance of considering multiple traits when disentangling the forces behind allopatric divergence. ©2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 826–838.     

Phenotypic variation in nestlings of a bird of prey under contrasting breeding and diet conditions

Environmental conditions often vary in space and time, and this may explain variation in the expression of phenotypic traits related to individual quality, such as ornamental coloration. Furthermore, the direction and strength of the relationship between coloured trait expression and individual quality might vary under contrasting conditions. These issues have been explored in adult birds but much less so in nestlings, which are more likely to experience different selective pressures and different physiological trade‐offs than adults. Here, we empirically investigated the effects of contrasting breeding and diet conditions on the expression of carotenoid‐based colour traits displayed by marsh harrier (Circus aeruginosus) nestlings. We studied the variation in coloration, body condition, and immune responsiveness of nestlings in four populations over a 5‐year period. We characterized spatiotemporal differences in rearing conditions experienced by C. aeruginosus nestlings in terms of breeding (laying date, clutch size, and number of nestlings hatched and fledged) and diet (percentage of mammal in diet and prey diversity) conditions. We found that breeding conditions influenced the co‐variation between coloration and immune responsiveness in female nestlings, and that diet conditions influenced the condition‐dependence of nestling coloration in later‐hatched nestlings. In addition, breeding conditions influenced nestling body condition and immune responsiveness, whereas diet conditions influenced nestling coloration and body condition. Our study highlights that nestling phenotype (levels of signalling, circulating carotenoids, and immunity) varies both spatially and temporally, and that some of this variation is related to differences in breeding and diet conditions. Moreover, under contrasting conditions, the direction of the relationships between nestling carotenoid‐based coloration and nestling quality may also vary. In order to fully understand the evolution and maintenance of colour traits in nestling birds, studies and experiments should ideally be replicated under contrasting rearing conditions. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Fecundity selection does not vary along a large geographical cline of trait means in a passerine bird

Local environmental and ecological conditions are commonly expected to result in local adaptation, although there are few examples of variation in phenotypic selection across continent‐wide spatial scales. We collected standardized data on selection with respect to the highly variable plumage coloration of pied flycatcher (Ficedula hypoleuca Pall.) males from 17 populations across the species' breeding range. The observed selection on multiple male coloration traits via the annual number of fledged young was generally relatively weak. The main aim of the present study, however, was to examine whether the current directional selection estimates are associated with distance to the sympatric area with the collared flycatcher (Ficedula albicollis Temminck), a sister species with which the pied flycatcher is showing character displacement. This pattern was expected because plumage traits in male pied flycatchers are changing with the distance to these areas of sympatry. However, we did not find such a pattern in current selection on coloration. There were no associations between current directional selection on ornamentation and latitude or longitude either. Interestingly, current selection on coloration traits was not associated with the observed mean plumage traits of the populations. Thus, there do not appear to be geographical gradients in current directional fecundity selection on male plumage ornamentation. The results of the present study do not support the idea that constant patterns in directional fecundity selection would play a major role in the maintenance of coloration among populations in this species. By contrast, the tendency for relatively weak mosaic‐like variation in selection among populations could reflect just a snapshot of temporally variable, potentially environment‐dependent, selection, as suggested by other studies in this system. Such fine‐grained variable selection coupled with gene flow could maintain extensive phenotypic variation across populations. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 808–827.     

Geographical variation in male advertisement calls and female preference of the wide‐ranging canyon treefrog,Hyla arenicolor

We surveyed the geographical variation in male advertisement calls of the wide‐ranging canyon treefrog, Hyla arenicolor, and found large call differences among geographically distant lineages that had been characterized by a recent phylogeographical study. To test whether these call differences were biologically relevant and could allow reproductive isolation of different lineages should they come into secondary contact, we assessed female preference in a lineage occurring in southern Utah and north‐western Arizona, USA. These females exhibited a strong preference for their own lineage's call type over the calls of two Mexican lineages, but not over the calls from the geographically nearest lineage. We also identified traits that female frogs probably use to discriminate between lineage‐specific advertisement calls. Our behavioural results, together with recent molecular estimates of phylogenetic relationships among lineages, will guide future work addressing the evolutionary forces that have led to this biologically significant variation in male sexual signals. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Cross‐fostering reveals that among‐brood differences in ornamental mouth coloration mostly reflect rearing conditions in nestling house sparrows

Dependent offspring use specialized traits to attract parental care. In birds, this includes morphological ornaments (e.g. colourful plumage or mouthparts) that are associated with nestling condition and shape the allocation of parental care. Ornament expression often differs among broods, even after differences in individual condition are accounted for statistically. Understanding how this variation arises is important for understanding the information content of these signals, their functional importance, and their evolution. The present study used a cross‐fostering experiment to assess the relative contributions of parental effects to among‐brood differences in the mouth coloration of nestling house sparrows, specifically the carotenoid‐richness, overall brightness, and ultraviolet (UV) coloration of rictal flanges. The expression of carotenoid‐based coloration was explained by synchronous breeding, nest‐of‐rearing and nest‐of‐origin. Brightness and relative UV intensity, however, were explained only by synchronous breeding, and there was substantial unexplained variation in all three colour parameters. Among‐brood variation in mouth coloration, then, may primarily contain information about the environment in which offspring are reared. At the individual level, ontogenetic changes in the carotenoid‐richness and brightness of flanges positively reflected mass gain (a proxy for food intake). Larger and yellower chicks gained more mass, consistent with parental preferences for these traits. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 169–179.     

Female mating preferences and male coloration covary with water transparency in a Lake Victoria cichlid fish

Rapid speciation in Lake Victoria cichlid fish of the genus Pundamilia may be facilitated by sexual selection: female mate choice exerts sexual selection on male nuptial coloration within species and maintains reproductive isolation between species. However, declining water transparency coincides with increasingly dull coloration and increasing hybridization. In the present study, we investigated the mechanism underlying this pattern in Pundamilia nyererei, a species that interbreeds with a sister species in turbid but not in clear water. We compared measures of intraspecific sexual selection between two populations from locations that differ in water transparency. First, in laboratory mate‐choice experiments, conducted in clear water and under broad‐spectrum illumination, we found that females originating from turbid water have significantly weaker preferences for male coloration than females originating from clear water. Second, both the hue and body coverage of male coloration differ between populations, which is consistent with adaptation to different photic habitats. These findings suggest that the observed relationship between male coloration and water transparency is not mediated by environmental variation alone. Rather, female mating preferences are indicated to have changed in response to this variation, constituting the first evidence for intraspecific preference‐trait co‐evolution in cichlid fish. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 398–406.     

The evolution of release and advertisement calls in green toads (Bufo viridis complex)

In the present paper we tested the hypothesis that differences in the acoustic communication system of diploid and tetraploid green toads ( Bufo viridis complex) might be due to selection for reproductive character displacement. We recorded two acoustic signals of the toad repertoire − the advertisement call (a long range mate-attracting signal) and the release call (a short-range signal mediating male–male interactions) − from six sympatric Central Asian populations (three diploid and three tetraploid populations) as well as from three allopatric diploid populations from Italy, and compared their patterns of variation with the pattern of among-population genetic distances. Although release and advertisement calls share the same morpho-physiological constraints, they show significantly different patterns of variation. Release calls vary congruently with the pattern of genetic distances, suggesting that mutation and genetic drift have been the major forces responsible for their change both in time and space. By contrast, the pattern of advertisement-call variation is not consistent with the phylogeny of the group, because the advertisement calls of Asian diploid and tetraploid populations differ from each other more than their genetic distances would predict. These results strongly support the hypothesis that selection acted on the advertisement calls of either or both Asian taxa , possibly, to favour reproductive isolation.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77, 379–391.     

Individual quality and carotenoid‐based plumage ornaments in male red bishops (Euplectes orix): plumage is not all that counts

Males in many bird species develop elaborate carotenoid‐based plumage ornaments that play an important role as signals of individual quality in intra‐ or intersexual selection. In the present study, we investigated which of several factors related to male condition and health affect the brightness and coloration of the carotenoid‐based orange–red breeding plumage in males of the red bishop (Euplectes orix), a polygynous and sexually dimorphic weaverbird species. The study revealed a very complex pattern, with the relationships between plumage traits and both heterophil‐to‐lymphocyte ratio and blood parasite load varying considerably among seasons, suggesting a strong influence of environmental conditions. Furthermore, overall condition of males strongly affected the association pattern between plumage traits and other factors, with males in bad condition being forced to allocate resources away from plumage elaboration to body maintenance or the enhancement of immune functions, whereas males in good condition can afford to invest in plumage ornamentation without obvious detrimental effects on health. Thus, females cannot rely on plumage characteristics alone to gather information on male quality, but have to assess additional traits that advertise general male health status. Perhaps surprisingly, testosterone levels were not related to male plumage characteristics. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 384–397.     

Melodic males and flashy females: Geographic variation in male and female reproductive behavior in red‐eyed treefrogs (Agalychnis callidryas)

Geographic variation in courtship behavior can affect reproductive success of divergent phenotypes via mate choice. Over time, this can lead to reproductive isolation and ultimately to speciation. The Neotropical red‐eyed treefrog (Agalychnis callidryas) exhibits high levels of phenotypic variation among populations in Costa Rica and Panama, including differences in color pattern, body size, and skin peptides. To test the extent of behavioral premating isolation among differentiated populations, we quantified male advertisement calls from six sites and female responses to male stimuli (acoustic and visual signals) from four sites. Our results show that both male advertisement calls and female behavior vary among populations: Discriminant function analyses can predict the population of origin for 99.3% ± 0.7 of males based on male call (dominant frequency and bandwidth) and 76.1% ± 6.6 of females based on female response behavior (frequency and duration of visual displays). Further, female mate choice trials (n = 69) showed that population divergence in male signals is coupled with female preference for local male stimuli. Combined, these results suggest that evolved differences among populations in male call properties and female response signals could have consequences for reproductive isolation. Finally, population variation in male and female behavior was not well explained by geographic or genetic distance, indicating a role for localized selection and/or drift. The interplay between male courtship and female responses may facilitate the evolution of local variants in courtship style, thus accelerating premating isolation via assortative mating.     

Wing pigmentation in Calopteryx damselflies: a role in thermoregulation?

Body melanization may show adaptive variation related to thermoregulation ability, and it is to be expected that the degree of melanization will change among populations or closely related species across environmental gradients of solar radiation and/or environmental temperature. Some melanized secondary sexual traits may also play a role in sexual selection, leading to interpopulation variation, which would not be predicted by thermoregulation pressures alone. We studied the relationships between the interpopulation variation in wing pigmentation level (i.e. melanized secondary sexual trait) of two closely related species of Calopteryx damselfly, and both solar radiation and maximum environmental temperature estimates. Wing pigmentation differs between these species, is gender specific and is used in species' discrimination. Only Calopteryx virgo meridionalis males showed a significant negative partial correlation between wing pigmentation degree and temperature. However, C. virgo meridionalis females showed a positive significant partial correlation between wing pigmentation degree and solar radiation. Wing pigmentation in Calopteryx xanthostoma males was not related to solar radiation or temperature. Thus, thermoregulation pressures poorly explained the observed variations in wing pigmentation between populations, although they might have an adaptive significance at the species' level. As wing pigmentation showed important latitudinal variation, several other selection pressures which might act on melanized traits are briefly discussed. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 36–44.     

No species barrier by call in an avian hybrid zone between California and Gambel's quail (Callipepla californica and C. gambelii)

Acoustic signals sometimes act as premating isolating barriers between animal species, but we know little about the circumstances that dictate the presence and strength of these barriers. Among insects, barriers to backcrossing are strengthened by acoustic signals that are under genetic control. Hybrid signals tend to be intermediate to parental signals, and signals are recognized only by like‐types, which results in reinforced species boundaries. This is not typically the case in avian taxa. Instead, acoustic signal transmission is controlled by some combination of genes and learning, and perhaps as a consequence of this variation, vocalizations play a diversity of roles in avian hybrid zones. I used California and Gambel's quail (Callipepla californica and C. gambelii), hybridizing birds that do not learn to vocalize, to explore whether genetically determined vocalizations function as a species barrier. Using spectral analysis, I measured temporal features of calls of uniquely colour‐banded quail that were recorded across one area of the California and Gambel's quail hybrid zone. Species discrimination is known to occur under captive conditions, though its basis is unexplored. Here I show that differences in the calls of parental species are likely great enough to permit species discrimination. Hybrid call components were intermediate to those of the parental species and covaried with genetic traits, as assessed with seven highly polymorphic microsatellite loci. Contrary to expectation, males as frequently called in response to unlike‐ as like‐type females who had initiated antiphonal calling, which is a courtship call between a female and a male. Furthermore, paired males and females did not share like‐type assembly calls, nor was there a correlation between the female's genetic or plumage traits and her mate's advertisement call. Based on these results, I conclude that California and Gambel's quail recognize each other and hybrids as potential mates and backcrossing occurs frequently. Thus, compatible mating signals could contribute to increased mixing of gene pools and slow the rate of speciation. I suggest that selection to respond to wide signal variation within species and imprinting on calls of mixed‐species coveys may cause mating signal compatibility between classes within the area of hybridization. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 253–264.     

The interplay between multiple predators and prey colour divergence

Evolutionary divergence in the coloration of toxic prey is expected when geographic variation in predator composition and behavior favours shifts in prey conspicuousness. A fundamental prediction of predator‐driven colour divergence is that the local coloration should experience lower predation risk than novel prey phenotypes. The dorsal coloration of the granular poison frog varies gradually from populations of conspicuous bright red frogs to populations of dull green and relatively cryptic frogs. We conducted experiments with clay models in four populations to examine the geographic patterns of taxon‐specific predation. Birds avoided the local phenotype while lizards consistently selected for decreased conspicuousness and crab predation did not depend on frog coloration. Importantly, birds and lizards favoured low conspicuousness in populations where relatively cryptic green morphs have evolved. This study provides evidence for the interplay among distinct selective pressures, from multiple‐predator taxa, acting on the divergence in protective coloration of prey species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 580–589.     

Rapid genetic differentiation between ex situ and their in situ source populations: an example of the endangered Silene otites (Caryophyllaceae)

Ex situ cultivation in botanic gardens could be one possibility to preserve plant species diversity and genetic variation. However, old ex situ populations are often sparsely documented. We were able to retrieve three different ex situ populations and their source in situ populations of the endangered plant species Silene otites after 20–36 years of isolation. Furthermore, three additional wild populations were included in the analysis. Population genetic diversity and differentiation were analysed using AFLP markers. Genetic variation in the ex situ populations was lower than the variation found in the in situ populations. Strong differentiation (F_ST = 0.21–0.36) between corresponding in situ and ex situ populations was observed. Bayesian clustering approach also showed a distinct genetic separation between in situ and ex situ populations. The high genetic differentiation and loss of genetic diversity during spatial and temporal isolation in the ex situ populations can be attributable to small population sizes and unconscious selection during cultivation. Therefore, adequate sampling prior to ex situ cultivation and large effective population sizes are important to preserve genetic diversity. Near‐natural cultivation allowing for generation overlap and interspecific competition without artificial selection is recommended as being best for the maintenance of the genetic constitution. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, ??, ??–??.     

Morphological adaptation to coastal marshes in spite of limited genetic structure in the Neotropical passerine Spartonoica maluroides (Aves: Furnariidae)

Tidal marshes present profound adaptive challenges to terrestrial vertebrates. For example, North American sparrows have relatively longer and thinner bills and darker dorsal plumage in coastal saltmarshes than in interior marshes. Bay‐capped wren‐spinetail (Furnariidae; Spartonoica maluroides) show a strong association with South American saltmarshes. We hypothesized that bay‐capped wren‐spinetail have similar morphological adaptations to North American sparrows to the saltmarsh environment, which would be indicative of the generality of selection on these traits in the coastal saltmarsh ecosystem. We captured individuals of S. maluroides from coastal saltmarshes and interior marshes. Populations were compared based on morphology and molecular markers. We found significant phenotypic differences in bill shape and plumage coloration (melanism) between S. maluroides populations from coastal and inland marshes. The low levels of genetic variation, weak geographical structure and shallow divergences, based on mitochondrial DNA and microsatellite data, suggest that coastal populations had a recent demographic expansion. Our results are consistent with the pattern of morphological divergence found between North American Emberizids. The possibility of convergent evolutionary adaptations between saltmarsh North American Emberizids and South American Furnariids suggests that there are strong selective pressures associated with saltmarsh environments on the beak, leading to adaptations for food acquisition, and on plumage coloration for better camouflage for predator avoidance (melanism). © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 78–91.     

A role for ecology in male mate discrimination of immigrant females in Calopteryx damselflies?

Sexual selection against immigrants is a mechanism that can regulate premating isolation between populations but, so far, few field studies have examined whether males can discriminate between immigrant and resident females. Males of the damselfly Calopteryx splendens show mate preferences and are able to force pre‐copulatory tandems. We related male mate responses to the ecological characteristics of female origin, geographic distances between populations, and morphological traits of females to identify factors influencing male mate discrimination. Significant heterogeneity between populations in male mate responses towards females was found. In some populations, males discriminated strongly against immigrant females, whereas the pattern was reversed or nonsignificant in other populations. Immigrant females were particularly attractive to males when they came from populations with similar predation pressures and densities of conspecifics. By contrast, immigrant females from populations with strongly dissimilar predation pressures and conspecific densities were not attractive to males. Differences in the abiotic environment appeared to affect mating success to a lesser degree. This suggests that male mate discrimination is context‐dependent and influenced by ecological differences between populations, a key prediction of ecological speciation theory. The results obtained in the present study suggest that gene‐flow is facilitated between ecologically similar populations. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 506–518.     

Genotype × environment interaction in the allometry of body,genitalia and signal traits in Enchenopa treehoppers (Hemiptera: Membracidae)

Developmental plasticity may promote divergence by exposing genetic variation to selection in novel ways in new environments. We tested for this effect in the static allometry (i.e. scaling on body size) of traits in advertisement signals, body and genitalia. We used a member of the Enchenopa binotata species complex of treehoppers – a clade of plant‐feeding insects in which speciation is associated with colonization of novel environments involving marked divergence in signals, subtle divergence in body size and shape, and no apparent divergence in genitalia. We found no change in mean allometric slopes across environments, but substantial genetic variation and genotype × environment interaction (G × E) in allometry. The allometry of signal traits showed the most genetic variation and G × E, and that of genitalia showed the weakest G × E. Our findings suggest that colonizing novel environments may have stronger diversifying consequences for signal allometry than for genitalia allometry. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 187–196.     

Genetic and morphologic diversity of European fan palm (Chamaerops humilis L.) populations from different environments from Sicily

Chamaerops humilis is decreasing in abundance in Mediterranean Europe, which has induced the European community to call for its protection in Special Areas of Conservation. However, information about its genetic and morphological variability, which is crucial to the development of any conservation strategies, is insufficient. The present study aimed to investigate the genetic and morphological variability of C. humilis in Sicily, which was selected as a model because of the high number of dense populations. The relationships between morphological traits and climatic variables were studied to highlight patterns of adaptation to the environment, along with the genetic similarity among the populations. Ten natural populations were sampled, analyzed using 28 specifically designed SSR primers, and evaluated based on 29 morphological traits. The populations were clustered similarly based on genetic and morphological traits. Heterozygosity was high and inbreeding coefficients were low. These results, along with higher intra‐ than inter‐population differentiation, suggest that C. humilis populations in Sicily differentiated from a common ancestor and that inter‐population variation arose from secondary evolution processes induced by ecological adaptation. The correlations between climatic variables and morphological traits suggest that the morphological adaptation to arid environments depends more on summer temperatures than on evapotranspiration or rainfall and that autumn and winter temperatures are determinants of the species establishment at new sites. Considering the response of C. humilis to seasonal temperatures, the present results indicate this species as a candidate for tracking climatic changes in Europe. Further studies are needed to highlight the adaptation of C. humilis to cold environments. Palaeo‐climatological and ‐ecological studies could help clarify its strategies for the colonization of new sites. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 66–81.     

Spawning coloration and sperm quality in a large lake population of Arctic charr (Salmonidae: Salvelinus alpinus L.)

The modern theories of sexual selection predict that male sexual ornaments may have evolved as reliable signals of male fertilization efficiency. However, among the studies of fishes with external fertilization, the results have yielded ambiguous evidence. In the present study, we present data on the phenotypic relationships between red spawning coloration and ejaculate quality (spermatocrit, sperm motility) from Arctic charr, Salvelinus alpinus. We studied two generations (F₁ and F₂) of males from a large lake population, reared in a standardized hatchery environment, to determine whether differential hatchery history, or duration of hatchery selection, affected the variation in ejaculate characteristics or abdominal coloration. After controlling for body length, there was no difference between the hatchery generations in these traits. However, the degree of redness increased with fish size. We found a positive correlation between sperm velocity and sperm longevity, indicating a functional integration between these sperm features across generations. Sperm velocity was also positively correlated with male redness. Therefore, the finding obtained in the present study suggests that the carotenoid‐based ornamentation in Arctic charr may provide information about differences between males in their fertilization potential. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 794–802.     

Melanin‐based coloration of sneaker male Atlantic salmon is linked to viability and emergence timing of their offspring

The ‘good genes’ hypothesis of sexual selection predicts that male ornaments are favoured by female mate choice because male ornament reveals genetic quality. In species with different male reproductive tactics, variation in genetic quality among ‘sneaking’ males has rarely been investigated, as usually ‘sneakers’ are thought not to be chosen by females. Here we focused on the alternative reproductive tactic in Atlantic salmon (Salmo salar Linnaeus, 1758) to test whether the skin colour of sneakers may reveal the performance traits of their offspring. A fully factorial breeding design was realized between 20 sneakers and two females using in vitro fertilization. We quantified the red and dark colorations of males and measured the survival of their progeny under semi‐natural conditions. In addition, the size of offspring and their emergence timing from the gravel nest were monitored in the laboratory. We found that darker males sired more viable offspring, whereas red coloration was negatively correlated with offspring survival. Nevertheless, darker and redder male pigmentations were linked to a delay in offspring emergence. These results demonstrate that colours can reveal individual genetic quality in an alternative male reproductive tactic, with male melanin‐based coloration being linked to both beneficial and detrimental effects for the offspring. Our results imply that sneaker ornaments may potentially play a role in both intra‐ and intersexual selection. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 126–135.     

Carotenoid limitation and allocation priorities in asynchronous raptor nestlings

The evolution and maintenance of conspicuous animal traits and communication signals have long fascinated biologists. Many yellow–red conspicuous traits are coloured by carotenoid pigments, and in some species they are displayed at a very young age. In nestling birds, the functions and proximate mechanisms of carotenoid‐pigmented traits are probably different and not as well known as those of adults. Here we investigated how Montagu's harrier (Circus pygargus) nestlings within structured families used a limited resource, carotenoid pigments, and whether they used these for increasing coloration (deposition in integuments) or for mounting a response to a phytohaemagglutinin (PHA) challenge, which measures pro‐inflammatory potential and aspects of cellular immune responsiveness. We manipulated carotenoid availability, using dietary carotenoid supplementations, and show that when supplemented, nestlings primarily allocated supplemental carotenoids to increase their coloration, irrespective of their sex, but depending of their position within the brood. Responses to PHA challenge were condition‐dependent, but depending on carotenoid availability. Moreover, how nestlings allocated carotenoids depended on their rank within the brood, which in turn influenced their level of carotenoid limitation (first‐hatched nestlings being less constrained than later‐hatched nestlings). We discuss why nestlings would use supplemental carotenoids for increasing bare parts coloration rather than for responding to a PHA challenge, and the potential benefits for doing so in a parent–offspring communication context. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 13–24.