A link between eumelanism and calcium physiology in the barn owl

In many animals, melanin-based coloration is strongly heritable and is largely insensitive to the environment and body condition. According to the handicap principle, such a trait may not reveal individual quality because the production of different melanin-based colorations often entails similar costs. However, a recent study showed that the production of eumelanin pigments requires relatively large amounts of calcium, potentially implying that melanin-based coloration is associated with physiological processes requiring calcium. If this is the case, eumelanism may be traded-off against other metabolic processes that require the same elements. We used a correlative approach to examine, for the first time, this proposition in the barn owl, a species in which individuals vary in the amount, size, and blackness of eumelanic spots. For this purpose, we measured calcium concentration in the left humerus of 85 dead owls. Results showed that the humeri of heavily spotted individuals had a higher concentration of calcium. This suggests either that plumage spottiness signals the ability to absorb calcium from the diet for both eumelanin production and storage in bones, or that lightly spotted individuals use more calcium for metabolic processes at the expense of calcium storage in bones. Our study supports the idea that eumelanin-based coloration is associated with a number of physiological processes requiring calcium.     

Covariation between eumelanic pigmentation and body mass only under specific conditions

Identifying the factors that mediate covariation between an ornament and other phenotypic attributes is important to determine the signaling function of ornaments. Sign and magnitude of a covariation may vary across environments if the expression of the ornament or of its linked genes regulating correlated phenotypes is condition-dependent. I investigated in the barn owl Tyto alba whether sign and magnitude of covariation between body mass and two heritable melanin-based plumage ornaments change with food supply, along the reproductive cycle and from the morning to the evening. Using a dataset of 1,848 measurements of body mass in 336 breeding females, I found that females displaying large black spots were heavier than conspecifics with smaller spots in the afternoon (i.e., a long time after the last feeding) but not in the morning (i.e., a short time after the last feeding). This is consistent with the recently proposed hypothesis that eumelanin-based ornaments are associated with the ability to maintain energy balance between food intake and energy expenditure. Thus, covariation between melanin-based coloration and body mass can be detected only under specific conditions potentially explaining why it has been reported in only ten out of 28 vertebrate species. The proposition that ornamented individuals achieve a higher fitness than drab conspecifics only in specific environments should be tested for other ornaments.     

Condition-dependent expression of melanin-based coloration in the Eurasian kestrel

Melanin is the most common pigment in animal integuments and is responsible for some of the most striking ornaments. A central tenet of sexual selection theory states that melanin-based traits can signal absolute individual quality in any environment only if their expression is condition-dependent. Significant costs imposed by an ornament would ensure that only the highest quality individuals display the most exaggerated forms of the signal. Firm evidence that melanin-based traits can be condition-dependent is still rare in birds. In an experimental test of this central assumption, we report condition-dependent expression of a melanin-based trait in the Eurasian kestrel (Falco tinnunculus). We manipulated nestling body condition by reducing or increasing the number of nestlings soon after hatching. A few days before fledging, we measured the width of sub-terminal black bands on the tail feathers. Compared to nestlings from enlarged broods, individuals raised in reduced broods were in better condition and thereby developed larger sub-terminal bands. Furthermore, in 2?years, first-born nestlings also developed larger sub-terminal bands than their younger siblings that are in poorer condition. This demonstrates that expression of melanin-based traits can be condition-dependent.     

Plumage coloration and nutritional condition in the great tit<Emphasis Type="Italic"> Parus major</Emphasis>: the roles of carotenoids and melanins differ

The size and coloration of some body characters seem to influence mate choice in many species. Most animal colours are either structural or based on melanin or carotenoid pigments. It has recently been suggested that carotenoid-based or structural coloration may be a condition-dependent trait, whereas melanin-based coloration is not; a difference that may be highly relevant when studying the evolution of multiple mating preferences. We tested this hypothesis in the great tit (Parus major). The size of the melanin breast band was not correlated to nutritional condition as estimated by the rate of tail growth (ptilochronology), controlling for locality, age, sex, year and season effects. However, the correlation was significant for the hue of yellow breast (carotenoid-based coloration), and the slopes of the regressions of the two pigments to growth bars differed significantly. These results suggest that the expression of the two traits may be regulated by different mechanisms.     

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.     

Habitat structure is associated with the expression of carotenoid-based coloration in nestling blue tits Parus caeruleus

We investigated how the expression of carotenoid-based plumage coloration (lightness and chroma) in nestling blue tits Parus caeruleus is associated with forest structure in oak forests of central Spain. We found evidence of a reduced expression of carotenoid-based coloration in nestlings growing up in successionally young and structurally simple forest territories. Our results suggest that breast feather coloration can be used as an indicator of nestling quality because nestlings with more intense yellow plumage coloration had larger body size and stronger immune responses to the injection of phytohemagglutinin (PHA). Given the association of forest structural complexity with carotenoid-based plumage coloration, our findings suggest that variation in habitat structure may have a significant impact on forest birds in their first stages of life which has implications for forest management practices.     

David and Goliath: potent venom of an ant-eating spider (Araneae) enables capture of a giant prey

It is rare to find a true predator that repeatedly and routinely kills prey larger than itself. A solitary specialised ant-eating spider of the genus Zodarion can capture a relatively giant prey. We studied the trophic niche of this spider species and investigated its adaptations (behavioural and venomic) that are used to capture ants. We found that the spider captures mainly polymorphic Messor arenarius ants. Adult female spiders captured large morphs while tiny juveniles captured smaller morphs, yet in both cases ants were giant in comparison with spider size. All specimens used an effective prey capture strategy that protected them from ant retaliation. Juvenile and adult spiders were able to paralyse their prey using a single bite. The venom glands of adults were more than 50 times larger than those of juvenile spiders, but the paralysis latency of juveniles was 1.5 times longer. This suggests that this spider species possesses very potent venom already at the juvenile stage. Comparison of the venom composition between juvenile and adult spiders did not reveal significant differences. We discovered here that specialised capture combined with very effective venom enables the capture of giant prey.     

Does contrast between eggshell ground and spot coloration affect egg rejection?

Obligate avian brood parasitic species impose the costs of incubating foreign eggs and raising young upon their unrelated hosts. The most common host defence is the rejection of parasitic eggs from the nest. Both egg colours and spot patterns influence egg rejection decisions in many host species, yet no studies have explicitly examined the role of variation in spot coloration. We studied the American robin Turdus migratorius, a blue-green unspotted egg-laying host of the brown-headed cowbird Molothrus ater, a brood parasite that lays non-mimetic spotted eggs. We examined host responses to model eggs with variable spot coloration against a constant robin-mimetic ground colour to identify patterns of rejection associated with perceived contrast between spot and ground colours. By using avian visual modelling, we found that robins were more likely to reject eggs whose spots had greater chromatic (hue) but not achromatic (brightness) contrast. Therefore, egg rejection decision rules in the American robin may depend on the colour contrast between parasite eggshell spot and host ground coloration. Our study also suggests that egg recognition in relation to spot coloration, like ground colour recognition, is tuned to the natural variation of avian eggshell spot colours but not to unnatural spot colours.     

Natal departure timing from spatially varying environments is dependent of individual ontogenetic status

Natal departure timing represents one of the first crucial decisions for juveniles born in spatially varying environments that ultimately disappear, but our knowledge on its determinants is limited. The present study aimed at understanding the determinants of juvenile natal departure by releasing individually tagged juvenile pike (Esox lucius L.) with variable body size and trophic position in a temporary flooded grassland. Specifically, we investigated whether natal departure depends on individual competitive status (‘competition hypothesis’), physiological tolerance to environmental conditions (‘physiological hypothesis’) or individual trophic position and the spatial heterogeneity of trophic resources (‘trophic hypothesis’). The results indicated that departure timing was negatively correlated with body size at release, showing that the dominance status among competing individuals was not the main trigger of juvenile departure. A positive correlation between departure timing and individual body size at departure was observed, suggesting that inter-individual variability in physiological tolerance did not explain departure patterns. While individual growth performances were similar irrespective of the timing of natal departure, stable isotope analyses revealed that juveniles with higher trophic position departed significantly earlier than individuals with lower trophic position. Therefore, the trade-off driving the use of spatially varying environments was most likely dependent upon the benefits associated with energetic returns than the costs associated with inter-individual competition or physiological stress. This result highlighted how ontogeny, and particularly ontogenetic niche shift, can play a central role in juvenile’s decision to depart from natal habitats in a predatory species.     

Intraspecific competition and high food availability are associated with insular gigantism in a lizard

Resource availability, competition, and predation commonly drive body size evolution. We assess the impact of high food availability and the consequent increased intraspecific competition, as expressed by tail injuries and cannibalism, on body size in Skyros wall lizards (Podarcis gaigeae). Lizard populations on islets surrounding Skyros (Aegean Sea) all have fewer predators and competitors than on Skyros but differ in the numbers of nesting seabirds. We predicted the following: (1) the presence of breeding seabirds (providing nutrients) will increase lizard population densities; (2) dense lizard populations will experience stronger intraspecific competition; and (3) such aggression, will be associated with larger average body size. We found a positive correlation between seabird and lizard densities. Cannibalism and tail injuries were considerably higher in dense populations. Increases in cannibalism and tail loss were associated with large body sizes. Adult cannibalism on juveniles may select for rapid growth, fuelled by high food abundance, setting thus the stage for the evolution of gigantism.     

The fractal dimension of a conspicuous ornament varies with mating status and shows assortative mating in wild red-legged partridges (<Emphasis Type="Italic">Alectoris rufa</Emphasis>)

Complex body designs, such as plumage ornaments in birds, can be described by fractal geometry. These complex patterns could have a role as visual signals during courtship and social interactions, but an empirical validation in the wild is currently lacking. Here, we investigated whether the fractal dimension (FD) of a complex plumage pattern displayed by red-legged partridges Alectoris rufa could function as a potential sexual signal. We captured wild birds early in the breeding season and tested if mated and unmated birds differed in the FD of their conspicuous melanin-based black bib. We also tested if the FD of the black bib was correlated within the pair, looking for evidence of assortative mating based on the expression of this trait. We simultaneously assessed similar effects in other ornamental traits (black bib size, white throat patch and black flank band surface, redness of the eye rings and bill). Mated birds showed higher black bib FD values than unmated ones. Mated males, but not females, also displayed a larger black bib. Moreover, the black bib FD (but not the trait size) and the white throat patch surface showed assortative mating. Finally, females with higher black bib FD showed smaller black flank band surface, suggesting a trade-off in the expression of the two melanin-pigmented plumage traits. This provides unique and novel indication for the shape complexity of a pigmented trait, here described by its fractal dimension, to be potentially under sexual selection in a wild animal.     

Agri-environment schemes and foraging of barn owls Tyto alba

The barn owl Tyto alba has been in decline throughout much of its range largely as a consequence of loss of foraging habitat resulting from agricultural intensification. The recent introduction of agri-environment schemes in the UK has resulted in the reinstatement of substantial areas of rough grassland which may be of benefit to small mammals and their avian predators. In this study mature grasslands provided by agri-environmental schemes (farm woodlands, permanent set-aside, 2 and 6 m field margins) were compared to better understand which hold the greatest prey numbers and how new areas of grassland targeted at barn owl conservation should be created and managed. In all habitats apart from farm woodlands, small mammal numbers were positively associated with taller swards. This study suggests that areas cut every 2 or 3 years supported more individuals and species than areas cut annually, although this trend was only significant for common shrews. With the introduction of the new Environmental Stewardship scheme in 2005, our results suggest that the area of suitable foraging habitat for barn owls is likely to increase.     

Inverse Rensch's rule in a frog with female-biased sexual size dimorphism

Rensch's rule claims that sexual size dimorphism (SSD) increases with body size when males are larger but decreases with body size when males are smaller. Chinese wood frog Rana chensinensis is a medium-sized species with female-biased size dimorphism. Using data on body size and age in 27 populations covering the full known size range of the species, we tested the consistency of allometric relationships between the sexes with Rensch's rule and evaluated the hypothesis that SSD is largely a function of age differences between the sexes. The results showed that level of female-biased SSD increased with increasing mean size, supporting the inverse of Rensch's rule. Moreover, most of the variation in SSD can be explained in terms of differences in age between the sexes in populations.     

Now you see me,now you don’t: iridescence increases the efficacy of lizard chromatic signals

The selective forces imposed by primary receivers and unintended eavesdroppers of animal signals often act in opposite directions, constraining the development of conspicuous coloration. Because iridescent colours change their chromatic properties with viewer angle, iridescence offers a potential mechanism to relax this trade-off when the relevant observers involved in the evolution of signal design adopt different viewer geometries. We used reflectance spectrophotometry and visual modelling to test if the striking blue head coloration of males of the lizard Lacerta schreibeiri (1) is iridescent and (2) is more conspicuous when viewed from the perspective of conspecifics than from that of the main predators of adult L. schreibeiri (raptors). We demonstrate that the blue heads of L. schreiberi show angle-dependent changes in their chromatic properties. This variation allows the blue heads to be relatively conspicuous to conspecific viewers located in the same horizontal plane as the sender, while simultaneously being relatively cryptic to birds that see it from above. This study is the first to suggest the use of angle-dependent chromatic signals in lizards, and provides the first evidence of the adaptive function of iridescent coloration based on its detectability to different observers.     

Mass predicts web asymmetry in Nephila spiders

The architecture of vertical aerial orb webs may be affected by spider size and gravity or by the available web space, in addition to phylogenetic and/or developmental factors. Vertical orb web asymmetry measured by hub displacement has been shown to increase in bigger and heavier spiders; however, previous studies have mostly focused on adult and subadult spiders or on several size classes with measured size parameters but no mass. Both estimations are suboptimal because (1) adult orb web spiders may not invest heavily in optimal web construction, whereas juveniles do; (2) size class/developmental stage is difficult to estimate in the field and is thus subjective, and (3) mass scales differently to size and is therefore more important in predicting aerial foraging success due to gravity. We studied vertical web asymmetry in a giant orb web spider, Nephila pilipes, across a wide range of size classes/developmental stages and tested the hypothesis that vertical web asymmetry (measured as hub displacement) is affected by gravity. On a sample of 100 webs, we found that hubs were more displaced in heavier and larger juveniles and that spider mass explained vertical web asymmetry better than other measures of spider size (carapace and leg lengths, developmental stage). Quantifying web shape via the ladder index suggested that, unlike in other nephilid taxa, growing Nephila orbs do not become vertically elongated. We conclude that the ontogenetic pattern of progressive vertical web asymmetry in Nephila can be explained by optimal foraging due to gravity, to which the opposing selective force may be high web-building costs in the lower orb. Recent literature finds little support for alternative explanations of ontogenetic orb web allometry such as the size limitation hypothesis and the biogenetic law.     

Colorful displays signal male quality in a tropical anole lizard

Parasites influence colorful ornaments and their behavioral display in many animal hosts. Because coloration and display behavior are often critical components of communication, variation in these traits may have important implications for individual fitness, yet it remains unclear whether such traits are signals of quality in many taxa. We investigated the association between ectoparasitic mite load and the color and behavioral use of the throat fan (dewlap) by male Anolis brevirostris lizards. We found that heavily parasitized lizards exhibited lower body condition, duller dewlaps, and less frequent dewlap displays than less parasitized individuals. Our results thus suggest that highly parasitized individuals invest less in both ornamental color and behavioral display of that color. Because the two components of the signal simultaneously provide information on male quality, this study provides novel support for the long-standing hypothesis that colorful traits may function as social or sexual signals in reptiles.     

Color polymorphism in a land snail Cepaea nemoralis (Pulmonata: Helicidae) as viewed by potential avian predators

Avian predation is one of the most probable factors maintaining polymorphism of shell coloration in Cepaea nemoralis. This assumption is justified by the fact that birds frequently forage on snails and their prey choice varies with morph coloration. However, in all preceding studies, the conspicuousness of morphs was determined only by using human vision which is significantly different from birds’ visual perception. In this study, we assessed how birds perceive colors of four Cepaea nemoralis morphs using physiological models of avian color vision. We calculated combined chromatic and achromatic contrast between shells and three habitat background types as a measure of shell conspicuousness. The degree of background color matching in Cepaea nemoralis depended on both shell morph and habitat type. On average, banded morphs were more conspicuous than unbanded morphs. Morphs were the most cryptic against dry vegetation and the most conspicuous on bare ground. We also found a significant interaction between habitat type and color morph. The relative conspicuousness of shell morphs depended on habitat and was the most variable against green vegetation. Our study provides the first insight into how potential avian predators view Cepaea nemoralis morphs. The results are discussed in light of multiple hypotheses explaining selective predation on Cepaea nemoralis morphs.     

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.     

Nestling rearing is antioxidant demanding in female barn swallows (Hirundo rustica)

Reproduction is a demanding activity, since organisms must produce and, in some cases, protect and provision their progeny. Hence, a central tenet of life-history theory predicts that parents have to trade parental care against body maintenance. One physiological cost thought to be particularly important as a modulator of such trade-offs is oxidative stress. However, evidence in favour of the hypothesis of an oxidative cost of reproduction is contradictory. In this study, we manipulated the brood size of wild barn swallows Hirundo rustica soon after hatching of their nestlings to test whether an increase in nestling rearing effort translates into an increased oxidative damage and a decreased antioxidant protection at the end of the nestling rearing period. We found that, while plasma oxidative damage was unaffected by brood size enlargement, females rearing enlarged broods showed a decrease in plasma non-enzymatic antioxidants during the nestling rearing period. This was not the case among females rearing reduced broods and among males assigned to either treatment. Moreover, individuals with higher plasma oxidative damage soon after the brood size manipulation had lower plasma non-enzymatic antioxidants at the end of the nestling rearing period, suggesting that non-enzymatic antioxidants were depleted to buffer the negative effects of high oxidative damage. Our findings point to antioxidant depletion as a potential mechanism mediating the cost of reproduction among female birds.     

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.