Carotenoids,immune response and the expression of sexual ornaments in male greenfinches (<Emphasis Type="Italic">Carduelis chloris</Emphasis>)

Allocation trade-offs of carotenoids between their use in the immune system and production of sexual ornaments have been suggested as a proximate mechanism maintaining honesty of sexual signals. To test this idea, we experimentally examined whether carotenoid availability in the diet was related to variation in antibody response to novel antigens in male greenfinches (Carduelis chloris aurantiiventris), a species with extensive carotenoid-dependent plumage colouration. We also measured the cost of mounting a humoral response in terms of circulating carotenoids. Finally, we examined the relationship between plumage colour, immune response and circulating carotenoids. We found that males with carotenoid-supplemented diets showed stronger antibody response than non-supplemented birds. We also found that activation of the immune system significantly reduced circulating carotenoids (24.9% lower in immune-challenged birds than in control birds). Finally, intensity (chroma) of ventral plumage colouration of males, a character directly related to concentration of total carotenoids in feathers, was negatively correlated with the immune response and circulating carotenoids in winter. These results support the idea that carotenoids are a limiting resource and that males trade ornamental colouration against immune response.     

Colour change in a structural ornament is related to individual quality,parasites and mating patterns in the blue tit

Carry-over effects refer to processes that occur in one season and influence fitness in the following. In birds, two costly activities, namely reproduction and moult, are restricted to a small time window, and sometimes overlap. Thus, colour in newly moulted feathers is likely to be affected by the costs of reproduction. Using models of bird vision we investigated male colour change in a free-living population of blue tits (Cyanistes caeruleus) in three sampling occasions: spring 1, winter and spring 2. We related crown, tail, breast and cheek feather colouration after the moult (winter) to the intensity of infections by blood parasites during reproduction (spring 1). In the following spring (spring 2), we explored mating patterns with respect to changes in feather colour (springs 1 vs. 2). Males that were less intensely infected by the malaria parasite Plasmodium while breeding showed purer white cheek feathers in winter, which may indicate higher feather quality. Increased brightness in the white cheek was associated with better body condition during reproduction. In the following season, males with brighter cheeks paired with females that had noticeably brighter cheek patches compared to the male’s previous mate. These results suggest that the conditions experienced during reproduction are likely to affect moult and thus feather colouration, at least in the white patch. High quality individuals may allocate resources efficiently during reproduction increasing future reproductive success through variation in mating patterns. Carry-over effects from reproduction might extend not only to the non-breeding phase, but also to the following breeding season.     

The effect of growth rate and ageing on colour variation of European pond turtles

Many chelonians have colourful dots, patches and stripes throughout their body that are made up, at least in part, of carotenoids. Therefore, turtles are very suitable models to study the evolution and functionality of carotenoid-based colouration. Recent studies suggested a close link between colouration and immune system in these taxa. However, more research is needed to understand the role of these colourful stripes and patches in turtle visual signalling. The purpose of this study was to explore the relationship between growth rate and colouration in European pond turtles. In particular, we wanted to answer the question of whether there is a trade-off between growth and colour expression. We also aimed to explore the effect of body size and age on colour variation. Turtles from a reintroduction-breeding program were recaptured, weighed and measured over an 8-year period to estimate their growth rates and age. We also measured with a spectrometer the reflectance of colour patches in two different body parts: shell and forelimb. We found that turtles with a faster growth rate had brighter limb stripes independently of their age. On the other hand, shell colouration was related to body size with larger turtles having brighter shell stripes and higher values of carotenoid chroma. Our results suggest that fast-growers may afford to express intense colourful limb stripes likely due to their higher intake of carotenoids that would modulate both growth and colour expression. However, shell colouration was related to body size probably due to ontogenetic differences in the diet, as juveniles are strictly carnivorous while adults are omnivorous. Alternatively, shell colouration might be involved in crypsis as the shell is visually exposed to predators.     

Ornamental comb colour predicts T-cell-mediated immunity in male red grouse <Emphasis Type="Italic">Lagopus lagopus scoticus</Emphasis>

Sexual ornaments might reliably indicate the ability to cope with parasites and diseases, and a better ability to mount a primary inflammatory response to a novel challenge. Carotenoid-based ornaments are amongst the commonest sexual signals of birds and often influence mate choice. Because carotenoids are immuno-stimulants, signallers may trade-off allocating these to ornamental colouration or using them for immune responses, so carotenoid-based ornaments might be particularly useful as honest indicators of immuno-compentence. Tetraonid birds, such as the red grouse Lagopus lagopus scoticus, exhibit supra-orbital yellow-red combs, a conspicuous ornament which functions in intra- and inter-sexual selection. The colour of combs is due to epidermal pigmentation by carotenoids, while their size is testosterone-dependent. In this study, I investigated whether comb characteristics, and in particular, comb colour, indicated immuno-competence in free-living male red grouse. I assessed T-cell-mediated immunity using a standardised challenge with phytohaemagglutinin. Red grouse combs reflect in the red and in the ultraviolet spectrum of light, which is not visible to humans but that grouse most likely see, so I measured comb colour across the whole bird visible spectrum (300-700 nm) using a reflectance spectrometer. I found that males with bigger and redder combs, but with less ultraviolet reflectance, had greater T-cell-mediated immune response. Comb colour predicted T-cell-mediated immune response better than comb size, indicating that the carotenoid-based colouration of this ornament might reliably signal this aspect of male quality.     

Assortative mating by carotenoid-based plumage colour: a quality indicator in American goldfinches,<Emphasis Type="Italic"> Carduelis tristis</Emphasis>

We investigated the mating patterns in 22 breeding pairs of wild American goldfinches (Carduelis tristis) with respect to their body size, condition, and carotenoid-based, yellow plumage colour. Using reflectance spectrometry, we objectively quantified plumage colours across the bird-visible wavelengths, revealing a unexpected UV peak in the reflectance spectrum from yellow feathers. We summarized our colour measurements using a principal components analysis to create a single variable, carotenoid PC1, that represents the intensity of this carotenoid-based yellow colour, a measure of phenotypic quality in this species. We found no evidence of assortative mating with respect to measures of body size or condition but there was positive assortative mating by carotenoid PC1, such that the yellow plumage colours of males and females were significantly correlated within pairs. We argue that the yellow carotenoid coloration of goldfinches may be important in mutual mate choice and, thus, that sexual selection in this species may act upon female ornamentation, as well as the more obvious plumage signals of males. Because assortative mating results in an increase in genetic variance, we suggest that this might be a mechanism that maintains variance in ornamental traits in spite of the variance-eroding effects of sexual selection.     

Negative correlation between nuptial throat colour and blood parasite load in male European green lizards supports the Hamilton–Zuk hypothesis

During female mate choice, conspicuous male sexual signals are used to infer male quality and choose the best sire for the offspring. The theory of parasite-mediated sexual selection (Hamilton–Zuk hypothesis) presumes that parasite infection can influence the elaboration of sexual signals: resistant individuals can invest more energy into signal expression and thus advertise their individual quality through signal intensity. By preferring these males, females can provide resistance genes for their offspring. Previous research showed that nuptial throat colour of male European green lizard, Lacerta viridis, plays a role in both inter- and intrasexual selections as a condition-dependent multiple signalling system. The aim of this study was to test the predictions of the Hamilton–Zuk hypothesis on male European green lizards. By blood sampling 30 adult males during the reproductive season, we found members of the Haemogregarinidae family in all but one individual (prevalence?=?96 %). The infection intensity showed strong negative correlation with the throat and belly colour brightness in line with the predictions of the Hamilton–Zuk hypothesis. In addition, we found other correlations between infection intensity and other fitness-related traits, suggesting that parasite load has a remarkable effect on individual fitness. This study shows that throat patch colour of the European green lizards not only is a multiple signalling system but also possibly acts as an honest sexual signal of health state in accordance with the Hamilton–Zuk hypothesis.     

Do feather-degrading bacteria actually degrade feather colour? No significant effects of plumage microbiome modifications on feather colouration in wild great tits

Parasites are known to exert selective pressures on host life history traits since the energy and nutrients needed to mount an immune response are no longer available to invest in other functions. Bird feathers harbour numerous microorganisms, some of which are able to degrade feather keratin (keratinolytic microorganisms) and affect feather integrity and colouration in vitro. Although named “feather-degrading” microorganisms, experimental evidence for their effects on feathers of free-living birds is still lacking. Here, we tested whether (i) keratinolytic microorganisms can degrade feathers in vivo and thus modify the colour of feathers during the nesting period and (ii) whether feather microorganisms have a long-term effect on the investment in colouration of newly moulted feathers. We designed treatments to either favour or inhibit bacterial growth, thus experimentally modifying plumage bacterial communities, in a wild breeding population of great tits (Parus major). Our analyses revealed no significant effects of the treatments on feather colours. Moreover, we found that differences in bacterial exposure during nesting did not significantly affect the colouration of newly moulted feathers. Our results suggest that significant feather degradation obtained during in vitro studies could have led to an overestimation of the potential of keratinolytic microorganisms to shape feather colouration in free-living birds.     

Laying date and polygyny as determinants of annual reproductive success in male collared flycatchers (Ficedula albicollis): a long-term study

Annual reproductive success (ARS) is one of the main components of lifetime reproductive success, a reliable measure of individual fitness. Previous studies often dealt with ARS and variables potentially affecting it. Among them, long-term studies that consider multiple factors at the same time are particularly important in understanding the adaptive value of different phenotypes. Here, we used an 18-year dataset to quantify the ARS of male collared flycatchers (Ficedula albicollis) on the basis of recruited offspring. We simultaneously assessed the effect of start of breeding, age, polygyny, body size and the expression of forehead patch (a sexually selected trait). The success of early breeding individuals was appreciably higher than late birds; however, breeding too early was also disadvantaged, and males that bred around the yearly median breeding date had the highest ARS. Polygynous males were more successful in years with good food supply, while in years with low food availability, they did not produce more recruits than monogamous males. The age of males, their forehead patch size and body size did not affect the number of recruits. Our findings support the importance of breeding date and suggest stabilizing selection on it in the long term. We also show that polygyny is not always advantageous for males, and its fitness pay-off may depend on environmental quality.     

Immune challenges and visual signalling in tree frogs

In animals, mate-choice is often based on sexual signals that carry information and help the receiver make the best choice to improve the receiver’s fitness. Orange visual sexual signals have been hypothesised to carry immune information because they are often due to carotenoid pigments which are also involved in immunity response. Although many studies have focused on the direct relationships between coloration and immunocompetence, few studies have simultaneously studied immunocompetent response and coloration variation after an immune challenge. We tested this hypothesis on starved and ad libitum-fed males of the European tree frog Hyla arborea. Our results show that male coloration is not a reliable indicator of its immune response capacity in this species. However, after an immune challenge induced by a PHA (Phaseolus vulgaris phytohaemagglutinin) injection, starved males presented a significant coloration loss and this alteration was related to the immune response intensity. Taken together, these results suggest that the brighter (lighter) coloration may be used as a cue by female to exclude males with a recent immune challenge, due to diseases or parasites for example.     

Growth conditions affect carotenoid-based plumage coloration of great tit nestlings

 Carotenoid-based integument colour in animals has been hypothesised to signal individual phenotypic quality because it reliably reflects either foraging efficiency or health status. We investigated whether carotenoid-derived yellow plumage coloration of fledgling great tits (Parus major) reflects their nestling history. Great tit fledglings reared in a poor year (1998) or in the urban habitat were less yellow than these reared in a good year (1999) or in the forest. The origin of nestlings also affected their coloration since nestlings from a city population did not improve their coloration when transferred to the forest. Brood size manipulation affected fledgling colour, but only in the rural population, where nestlings from reduced broods developed more yellow coloration than nestlings from increased and control broods. Effect of brood size manipulation on fledgling plumage colour was independent of the body mass, indicating that growth environment affects fledgling body mass and plumage colour by different pathways. Received: 20 March 2000 / Accepted in revised form: 4 September 2000     

Melanin- and carotenoid-dependent signals of great tits (<Emphasis Type="Italic">Parus major</Emphasis>) relate differently to metal pollution

Due to their high phenotypic plasticity, the expression of secondary sexual characteristics is particularly sensitive to stress. Here, we investigated the expression of two conspicuous visual signals in great tits (Parus major) in a metal pollution gradient. In three study sites with marked differences in metal contamination (mainly lead, cadmium, copper and zinc), we compared melanin and carotenoid colouration of great tits. While carotenoid colouration (yellow breast) was negatively related to metal pollution, the size of a melanin trait (breast stripe) was larger in the most polluted sites. Environmental pollutants not only affect the expression of conspicuous signals but may even enhance, directly or indirectly, a signal of male quality such as breast stripe. Our results also support the multiple messages hypothesis predicting that different signals highlight different aspects of geno- and phenotypic condition of the bearer.     

Integrated plumage colour variation in relation to body condition, reproductive investment and laying date in the collared flycatcher

The possible integration of different sexual ornaments into a composite system, and especially the information content of such ornament complexes, is poorly investigated. Many bird species display complex plumage coloration, but whether this represents one integrated or several independent sexual traits can be unclear. Collared flycatchers (Ficedula albicollis) display melanised and depigmented plumage areas, and the spectral features (brightness and UV chroma) of these are correlated with each other across the plumage. In a 5-year dataset of male and female plumage reflectance, we examined some of the potential information content of integrated, plumage-level colour attributes by estimating their relationships to previous and current year body condition, laying date and clutch size. Females were in better condition the year before they became darker pigmented, and males in better current year condition were also darker pigmented. Female pigment-based brightness was positively, while male structurally based brightness was negatively related to current laying date. Finally, the overall UV chroma of white plumage areas in males was positively associated with current clutch size. Our results show that higher degree of pigmentation is related to better condition, while the structural colour component is associated with some aspects of reproductive investment. These results highlight the possibility that correlated aspects of a multiple plumage ornamentation system may reflect together some aspects of individual quality, thereby functioning as a composite signal.     

Colour mimicry and sexual deception by Tongue orchids (Cryptostylis)

Typically, floral colour attracts pollinators by advertising rewards such as nectar, but how does colour function when pollinators are deceived, unrewarded, and may even suffer fitness costs? Sexually deceptive orchids are pollinated only by male insects fooled into mating with orchid flowers and inadvertently transferring orchid pollinia. Over long distances, sexually deceptive orchids lure pollinators with counterfeit insect sex pheromones, but close-range deception with colour mimicry is a tantalising possibility. Here, for the first time, we analyse the colours of four sexually deceptive Cryptostylis orchid species and the female wasp they mimic (Lissopimpla excelsa, Ichneumonidae), from the perspective of the orchids’ single, shared pollinator, male Lissopimpla excelsa. Despite appearing different to humans, the colours of the orchids and female wasps were effectively identical when mapped into a hymenopteran hexagonal colour space. The orchids and wasps reflected predominantly red-orange wavelengths, but UV was also reflected by raised bumps on two orchid species and by female wasp wings. The orchids’ bright yellow pollinia contrasted significantly with their overall red colour. Orchid deception may therefore involve accurate and species-specific mimicry of wavelengths reflected by female wasps, and potentially, exploitation of insects’ innate attraction to UV and yellow wavelengths. In general, mimicry may be facilitated by exploiting visual vulnerabilities and evolve more readily at the peripheries of sensory perception. Many sexually deceptive orchids are predominantly red, green or white: colours that are all potentially difficult for hymenoptera to detect or distinguish from the background.     

Dietary antioxidants, lipid peroxidation and plumage colouration in nestling blue tits Cyanistes caeruleus

Carotenoid pigments are responsible for many of the red, yellow and orange plumage and integument traits seen in birds. One idea suggests that since carotenoids can act as antioxidants, carotenoid-mediated colouration may reveal an individual's ability to resist oxidative damage. In fact, there is currently very little information on the effects of most dietary-acquired antioxidants on oxidative stress in wild birds. Here, we assessed the impacts on oxidative damage, plasma antioxidants, growth and plumage colouration after supplementing nestling blue tits Cyanistes caeruleus with one of three diets; control, carotenoid treatment or α-tocopherol treatment. Oxidative damage was assessed by HPLC analysis of plasma levels of malondialdehyde (MDA), a by-product of lipid peroxidation. Contrary to predictions, we found no differences in oxidative damage, plumage colouration or growth rate between treatment groups. Although plasma lutein concentrations were significantly raised in carotenoid-fed chicks, α-tocopherol treatment had no effect on concentrations of plasma α-tocopherol compared with controls. Interestingly, we found that faster growing chicks had higher levels of oxidative damage than slower growing birds, independent of treatment, body mass and condition at fledging. Moreover, the chromatic signal of the chest plumage of birds was positively correlated with levels of MDA but not plasma antioxidant concentrations: more colourful nestlings had higher oxidative damage than less colourful individuals. Thus, increased carotenoid-mediated plumage does not reveal resistance to oxidative damage for nestling blue tits, but may indicate costs paid, in terms of oxidative damage. Our results indicate that the trade-offs between competing physiological systems for dietary antioxidants are likely to be complex in rapidly developing birds. Moreover, interpreting the biological relevance of different biomarkers of antioxidant status represents a challenge for evolutionary ecology.     

A comparative analysis of colour preferences in temperate and tropical social bees

The spontaneous occurrence of colour preferences without learning has been demonstrated in several insect species; however, the underlying mechanisms are still not understood. Here, we use a comparative approach to investigate spontaneous and learned colour preferences in foraging bees of two tropical and one temperate species. We hypothesised that tropical bees utilise different sets of plants and therefore might differ in their spontaneous colour preferences. We tested colour-naive bees and foragers from colonies that had been enclosed in large flight cages for a long time. Bees were shortly trained with triplets of neutral, UV-grey stimuli placed randomly at eight locations on a black training disk to induce foraging motivation. During unrewarded tests, the bees’ responses to eight colours were video-recorded. Bees explored all colours and displayed an overall preference for colours dominated by long or short wavelengths, rather than a single colour stimulus. Naive Apis cerana and Bombus terrestris showed similar choices. Both inspected long-wavelength stimuli more than short-wavelength stimuli, whilst responses of the tropical stingless bee Tetragonula iridipennis differed, suggesting that resource partitioning could be a determinant of spontaneous colour preferences. Reward on an unsaturated yellow colour shifted the bees’ preference curves as predicted, which is in line with previous findings that brief colour experience overrides the expression of spontaneous preferences. We conclude that rather than determining foraging behaviour in inflexible ways, spontaneous colour preferences vary depending on experimental settings and reflect potential biases in mechanisms of learning and decision-making in pollinating insects.     

Sexually dichromatic coloration reflects size and immunocompetence in female Spanish terrapins, Mauremys leprosa

Many studies have shown the importance of colorful ornamentation in mate choosiness or intrasexual conflict. However, research on color ornaments has focused mainly on birds, lizards or fish, but remains practically unknown in other animal groups such as turtles. In addition, female ornaments and their relation with sexual selection also remain almost unknown. Here, we measured the coloration of the shell and the limb stripes of male and female Spanish terrapins Mauremys leprosa and explored the existence of sexual dichromatism and the relation of color characteristics with body size and health state estimated from the immune response to the injection of an antigen (phytohaemagglutinin test). Our results showed that shell coloration, which could be constrained by natural selection to be cryptic, changed with body size, but did not differ between sexes. In contrast, females had brighter and less ultraviolet-saturated and more orange-saturated limb stripes than males. In females, interindividual variation in limb stripe coloration was related with body size and immune response suggesting that this coloration may inform honestly about multiple traits that could be important in sexual selection. In contrast, coloration of limb stripes of males was duller than in females, and was not related with any trait suggesting that coloration is not important in sexual selection for males.     

Temporal variation in size-assortative mating and male mate choice in a spider with amphisexual care

Males should be more selective when they have a high investment in reproduction, especially in species with biparental or paternal care. In this context, male mate choice can promote size-assortative mating (SAM) when (1) large males win intrasexual disputes, (2) large females are more fecund, and (3) males prefer larger females to smaller ones. In the spider Manogea porracea, males exhibit high reproductive investment by building their webs above those of females and exhibiting extended care of offspring in the absence of females. Under these circumstances, we expect the occurrence of SAM and male preference for large females. Herein, we performed observations and experiments in the field to evaluate the hypotheses that (1) M. porracea mates assortatively by size and (2) SAM is influenced by male mate choice. Furthermore, we measured variables that could affect mating patterns, the sex ratios, and densities of both sexes. Pairing in M. porracea was positively size-assortative in 2012, but not in 2013. Large males won most disputes for mates and preferred larger females, which produced more eggs. The inconsistency in detection of SAM was due to population dynamics, namely variations in sex ratio and population density across the breeding season. Furthermore, we found that the significance of male mate choice on sexual selection of body size in M. porracea strongly depends on the competition intensity for mating opportunities. The traditional sexual selection hypothesis of SAM needs to be reviewed and must include measures of competition intensity.     

Colour preferences influences odour learning in the hawkmoth, Macroglossum stellatarum

The hummingbird hawkmoth, Macroglossum stellatarum, learns colour fast and reliably. It has earlier been shown to spontaneously feed from odourless artificial flowers. Now, we have studied odour learning. The moths were trained to discriminate feeders of the same colour but marked with different odours. They did not learn to discriminate two natural flower odours when they were presented with the innately preferred colour blue, but they did learn this discrimination combined with yellow or green colours that are less attractive to the moth. The yellow colour could be trained to become as attractive as the innately preferred blue colour and the blue colour could be trained to become less attractive. This is the first proof of odour learning in a diurnal moth. The results show that M. stellatarum can use more than one modality in their foraging behaviour and that the system is plastic. By manipulating the preferences for the different colours, their influence on odour learning could be changed.     

Blue-green eggshell coloration is not a sexually selected signal of female quality in an open-nesting polygynous passerine

It has been proposed that blue-green egg colours have evolved as a post-mating signal of female quality, selected by males allocating their parental effort in response to the strength of this signal. We tested two main assumptions of the sexually selected egg coloration hypothesis: (1) whether the intensity of eggshell blue-green chroma (BGC) reflects female quality; and (2) whether males make their decisions on the level of parental care that they provide according to the intensity of eggshell BGC. As a model species, we chose the facultatively polygynous great reed warbler (Acrocephalus arundinaceus). In this species, females simultaneously paired with the same male, compete for his nest attendance and could benefit from signalling their quality through egg coloration. However, we found no association between the variation in eggshell BGC and the measures of female quality (physical condition, mean egg volume and age). Moreover, great reed warbler males did not adjust their investment (as measured in terms of nest defence against a brood parasite) in relation to the eggshell BGC. We conclude that blue-green egg coloration in this open-nesting passerine is unlikely to have a signalling function. Rather, the large colour variation among clutches of individual females may depend on yearly fluctuations in environmental conditions.