Colour-specific diet specialization is associated with differences in owlet weight in a polymorphic owl: influence of the trophic quality variation.

The niche divergence hypothesis proposes that the evolution and maintenance of colour polymorphism is based on a mechanism of disruptive selection. In a trophic context, the hypothesis predicts that individuals differing in colour vary in their trophic niche, either because they differ in foraging efficiency or feed in different habitats. A major evolutionary conundrum is how these expectations are affected by variation in trophic quality. Using an owl species with colour plumage polymorphism, the Eurasian scops owl Otus scops, we examined diet and habitat segregation during reproduction in relation to plumage colouration and trophic quality. Intensive sampling revealed that trophic quality for scops owls (i.e. abundance of grasshoppers and locusts) varied more among territories than between years, but scops owls did not segregate among territories of different quality by their colouration. However, we found that sex, plumage colouration and territory differences in trophic quality explained differences in the degree of dietary specialization. Brownish males delivered a higher diversity of prey to the nest than greyish ones in high trophic quality territories. We also found that the more diverse the diet provided by males, the heavier the owlets at fledging. Our study provides evidence for a different sensitivity to trophic quality of the colour morphs with potential fitness consequences in scops owls. We highlight the importance of studying the mechanisms leading to the persistence of colour polymorphism in patchy environments, since segregation may pass otherwise unnoticed if only habitats or years with similar conditions are considered.

Geographical variation in pace-of-life in a long-distance migratory bird: implications for population management.

Life-history theory predicts that animals should develop adaptive trade-offs between survival and reproduction to maximize their fitness. This results in a continuum of life-history strategies among species, ranging from slow to fast paces-of-life. The optimal pace-of-life has been shown to vary within environmental gradients, with a commonly observed pattern of a slow-to-fast continuum from the tropics to the poles. Within species, pace-of-life variability has however received much less attention. In this study, we investigated whether or not the pace-of-life of populations within a species follows the expected slow-fast continuum associated with latitude. We analysed the variability of life-history strategies among populations of the European roller Coracias garrulus, a long-distance migratory species, comparing breeding parameters and adult survival between populations across a latitudinal gradient. The findings showed a negative correlation between survival and clutch size in roller populations, with a slower pace-of-life in the northern populations and a faster pace-of-life in the southern populations: a reverse gradient to what might be expected from inter-specific studies. These results suggest that northern populations would benefit from measures enhancing adult survival probability, such as reduction in harvesting rates, while southern populations would respond better to actions favouring reproductive success, such as nesting site provisioning. This study highlights that life-history traits can vary substantially between populations of a single species with a large latitudinal breeding range, and pinpoint how knowledge about this variability may be key in anticipating different populations' responses to threats as well as to conservation strategies.

Avian and rodent responses to the olfactory landscape in a Mediterranean cavity community.

Animals rely on cues informing about future predation risk when selecting habitats to breed in. Olfactory information may play a fundamental role in the assessment of predation threats, because predators produce characteristic body odours, but the role of odours in habitat selection has seldom been considered. Here, we test whether fear of predation induced by odour cues may affect the settlement pattern of a Mediterranean cavity-dependent community of rodents and non-excavator hole-nesting birds. To test this hypothesis, we experimentally manipulated the perception of predation risk on a scale of patch by applying either odours of a carnivore predator (risky odour treatment), lemon essence (non-risky odour treatment) and a control non-odorous treatment and studied bird and rodent settlement patterns. Nest-box occupation probability differed across treatments so that species in the community settled in more numbers in control than in non-risky and than in risky odour-treated nest boxes. Concerning settlement patterns, control nest boxes were occupied more rapidly than nest boxes with odour information. Birds and rodents settled earlier in control than in risky odour-treated nest boxes, but their settlement pattern did not significantly vary between risky odour and non-risky odour-treated nest boxes. Our findings demonstrate that olfactory cues may be used to assess habitat quality by settling species in this community, but we cannot pinpoint the exact mechanism that has given rise to the pattern of preference by nest boxes.

Colour polymorphism in owls is linked to light variability.

Owls show an astonishing variation in their degree of colour polymorphism, although the exact mechanisms driving such variation remain controversial. Here we address this fundamental question by considering information on all extant owls and recent advances in comparative methods in the frame of three mutually non-exclusive evolutionary scenarios. In addition, we study for the first time whether the evolution of influential ecological characters facilitated the evolution of colour polymorphism (or vice versa). In agreement with the niche divergence hypothesis, we found that species living under more variable luminal conditions, i.e., species with diurnal and crepuscular habits and those inhabiting in a mixture of open and closed habitats, were more likely to show colour polymorphism. Correlated evolution analyses revealed that a change in the luminal niche might be a fundamental requisite for the evolution of colour polymorphism. Moreover, polymorphism was more frequent among owl species occupying lower trophic levels, which could be explained by a particularly high selection for crypsis on small predator owls. Our results, thus, provide support for the idea that colour polymorphism in owls is an adaptive character likely maintained by the selective advantage of morphs under different environmental conditions via disruptive selection mechanisms.

Receivers matter: the meaning of alarm calls and competition for nest sites in a bird community.

Animal communities may constitute information networks where individuals gain information on predation risk by eavesdropping on alarm calls of other species. However, communities include species in different trophic levels, and it is not yet known how the trophic level of the receiver influences the informative value of a call. Furthermore, no empirical study has yet tested how increased competition may influence the value of alarm calls for distinct receivers. Here, we identify the importance of alarm calls emitted by a small owl, the little owl (Athene noctua), on the structure of a cavity-nesting bird community including mesopredators and primary prey under variable levels of competition for nest holes. Competitors sharing top predators with the callers and prey of the callers interpreted alarm and non-alarm calls differently. Competitors chose preferentially alarm and non-alarm patches over control patches to breed, while prey selected alarm patches. In contrast, competition for nest sites affected habitat selection of prey species more than that of competitors of the callers. This study provides support for a changing value of alarm calls and competition for nest sites for distinct receivers related to niche overlapping among callers and eavesdroppers, therefore, calling attention to possible cascading effects by the use of information in natural communities.

Alarm calls modulate the spatial structure of a breeding owl community.

Animals should continuously assess the threat of predation. Alarm calls inform on predation risk and are often used as cues to shape behavioural responses in birds and mammals. Hitherto, however, the ecological consequences of alarm calls in terms of organization of animal communities have been neglected. Here, we show experimentally that calls of a resident nocturnal raptor, the little owl Athene noctua, triggered a response in terms of breeding habitat selection and investment in current reproduction in conspecifics and heterospecifics. Little owls preferred to settle in territories where calls of conspecifics, irrespective of their type (i.e. alarm versus contact calls), were broadcasted, indicating that either conspecific attraction exists or calls are interpreted as foreign calls, eliciting settlement as a mode of defence against competitors. Also, we found that little owls seemed to invest more in current reproduction in safe territories as revealed by conspecific calls. Innovatively, we reported that a second owl species, the migratory scops owl Otus scops, preferred to breed in safe territories as indicated by little owls' calls. These results evidence that the emission of alarm calls may have, apart from well-known behavioural effects, ecological consequences in natural communities by inducing species-specific biases in breeding habitat selection. This study demonstrates a previously unsuspected informative role of avian alarm calls which may modulate the spatial structure of species within communities.

Sex recognition by odour and variation in the uropygial gland secretion in starlings.

1. Although a growing body of evidence supports that olfaction based on chemical compounds emitted by birds may play a role in individual recognition, the possible role of chemical cues in sexual selection of birds has been only preliminarily studied. 2. We investigated for the first time whether a passerine bird, the spotless starling Sturnus unicolor, was able to discriminate the sex of conspecifics by using olfactory cues and whether the size and secretion composition of the uropygial gland convey information on sex, age and reproductive status in this species. 3. We performed a blind choice experiment during mating, and we found that starlings were able to discriminate the sex of conspecifics by using chemical cues alone. Both male and female starlings preferred male scents. Furthermore, the analysis of the chemical composition of the uropygial gland secretion by using gas chromatography-mass spectrometry (GC-MS) revealed differences between sexes, ages and reproductive status. 4. In conclusion, our study reveals for first time that a passerine species can discriminate the sex of conspecifics by relying on chemical cues and suggests that the uropygial gland secretion may potentially function as a chemical signal used in mate choice and/or intrasexual competition in this species.

Predation risk determines breeding territory choice in a Mediterranean cavity-nesting bird community.

Non-direct effects of predation can be an important component of the total effect of predation, modulating animal population and community dynamics. The isolated effects of predation risk on the spatial organisation of the breeding bird community, however, remains poorly studied. We investigated whether an experimentally increased predation risk prior to reproduction affected breeding territory selection and subsequent reproductive strategies in three Mediterranean cavity-nesting birds, i.e., the little owl Athene noctua, European roller Coracias garrulus and scops owl Otus scops. We found that territories used the previous year were more likely to be re-occupied when they belonged to the safe treatment rather than to the risky treatment. The first choice of breeders of all three species was for safe territories over risky ones. When all breeding attempts in the season (i.e., final occupation) were considered, breeders also preferred safe to risky sites. In addition, little owls laid larger eggs in risky territories than in safe territories. Our study provides experimental evidence of a rapid preventive response of the three most abundant species in a cavity-nesting bird community to a short-term manipulation of predation risk. This response highlights the key role of the non-direct effects of predation in modulating avian community organisation.

Visual cues and parental favouritism in a nocturnal bird.

Visual signals are crucial for parent-offspring communication, although their functioning has been neglected for nocturnal birds. Here, we investigated parental preference for nestling coloration in nocturnal conditions--a question hitherto unexplored--in a nocturnal raptor, the scops owl (Otus scops). We assessed how parents allocated food during the night in relation to a manipulation of ultraviolet (UV) reflectance of the cere (skin above the beak) of their offspring. Reflectance of the cere shows a marked peak in the UV part of the spectrum, and location of the UV peak is related to nestling body mass (i.e. heavier nestlings have a UV peak at lower wavelengths). We found evidence of parental bias in favour of lighter offspring: UV-reduced nestlings gained more weight during the night than their control siblings. This study provides the first experimental evidence of the use of visual cues for parent-offspring communication in a nocturnal bird.

Do spotless starlings place feathers at their nests by ultraviolet color?

A considerable number of bird species carry feathers to their nests. Feathers' presence in the nests has traditionally been explained by their insulating properties. Recently, however, it has been suggested that feathers carried to the nests by females of the spotted starling (Sturnus unicolor L.) could have an ornamental function based on their ultraviolet (300-400 nm) and human-visible longer wavelength (400-700 nm) coloration. In our population, 95.7% of feathers found inside next-boxes occupied by nesting starlings were rock dove fly feathers. Of these feathers, 82.7% were naturally positioned with their reverse side oriented toward the entrance hole and 42.4% of all found feathers were situated within the nest-cup. Here we experimentally assess the signaling function of ultraviolet coloration of feathers in nests of spotless starlings by providing nests with a number of pigeon flight feathers that were respectively treated on their obverse, reverse, both, or neither side with a UV blocker. Starlings placed 42.5% of the experimental feathers in the nest-cup irrespective of the UV block treatment. Orientation of feathers toward the entrance hole was not related with their ultraviolet radiation. However, feathers placed within the nest-cup were more likely found with their reverse side oriented toward the entrance hole confirming our correlative findings. These results suggest a minor role of ultraviolet coloration on feather location by spotless starlings.

Melanism influences the use of social information in a polymorphic owl.

Social information use has well-known fitness benefits. However, causes underlying the apparent inter-individual variability in the propensity to use social information are poorly studied. Melanins are pigments responsible for most of intra-specific color variation in vertebrates and their variation is often associated with changes in behaviour. Here, we explored whether melanism is related to individual propensity to use social information in the color polymorphic scops owl Otus scops. We manipulated social information on predation risk at nests by broadcasting calls of the sympatric little owl Athene noctua and found that owlets of brownish females exposed to alarm calls had lower levels of natural antibodies than those of greyish females. In parallel, we found changes in parental behaviour contingent on coloration because when exposed to the risky treatment brownish females returned earlier to nests than greyish females and owlets raised by brownish females were fed with smaller prey than those raised by greyish ones. These results provide support for a previous ignored role of melanins on the propensity to use social information, which may help to explain the maintenance of melanin-based color polymorphisms wherever social environments are variable.

Phaeomelanin matters: Redness associates with inter-individual differences in behaviour and feather corticosterone in male scops owls (Otus scops).

Individuals within populations often show consistent variation in behavioural and physiological traits which are frequently inter-correlated, potentially leading to phenotypic integration. Understanding the mechanisms behind such integration is a key task in evolutionary ecology, and melanin based colouration has been suggested to play a pivotal role. In birds, most of plumage colour variation is determined by two types of melanin, eumelanin and phaeomelanin, but the role of phaeomelanin in avian phenotype integration has been barely investigated. Here, we test for covariation between phaeomelanin-based colouration, behavioural traits (i.e. nest territoriality, aggressiveness, breath rate and parental behaviour) and corticosterone in feathers in the polymorphic scops owl Otus scops, a bird species in which more phaeomelanic individuals display reddish colourations. In males, we observed that reddish males took longer to return to their nests and showed higher levels of feather CORT than more greyish ones. Behaviour and feather CORT were not associated to plumage colour in females. The found associations between redness, behaviour and feather CORT in males, but not in females, might suggest the existence of a sex-specific integrated phaeomelanic phenotype in scops owls.

Methionine supplementation influences melanin-based plumage colouration in Eurasian kestrel, Falco tinnunculus, nestlings.

The extent to which the expression of melanin-based plumage colouration in birds is genetically or environmentally determined is controversial. Here, we performed a between-nest design supplementation with either the sulphur amino acid dl-methionine or with water to investigate the importance of the non-genetic component of melanin-based plumage colouration in the Eurasian kestrel, Falco tinnunculus. Methionine affects growth and immunity, thus we aimed to modify nestling growth and immunity before feather development. Then, we measured the effect of the experiment on colouration of two melanin-based plumage patches of nestling kestrels. We found that methionine slowed down nestling growth through treatment administration and that nestlings compensated by speeding up their growth later. We did not find any effects of methionine on nestling immunity (i.e. lymphocyte counts, natural antibody levels or complement-mediated immunity). Effects on growth seemed to be mirrored by changes in nestling colouration in the two sexes: methionine-nestlings showed less intense brown plumage on their backs compared with control nestlings. These results provide support for a non-genetic determination of a melanin-based plumage patch in the two sexes of nestling kestrels.

Immunity and fitness in a wild population of Eurasian kestrels Falco tinnunculus.

The immune system of vertebrates consists of several components that partly interact and complement each other. Therefore, the assessment of the overall effectiveness of immune defence requires the simultaneous measurement of different immune components. In this study, we investigated intraspecific variability of innate [i.e. natural antibodies (NAb) and complement] and acquired (i.e. leucocyte profiles) immunity and its relationship with fitness correlates (i.e. blood parasite load and reproductive success in adults and body mass and survival until fledging in nestlings) in the Eurasian kestrel Falco tinnunculus. Immunity differed between nestlings and adults and also between adult males and females. Adult kestrels with higher levels of complement were less parasitised by Haemoproteus, and males with higher values of NAbs showed a higher reproductive success. In nestlings, the H/L ratio was negatively related to body mass. Survival until fledging was predicted by all measured immunological variables of nestlings as well as by their fathers' level of complement. This is the first time that innate immunity is linked to survival in a wild bird. Thus, intraspecific variation in different components of immunity predicts variation in fitness prospects in kestrels, which highlights the importance of measuring innate immune components together with components of the acquired immunity in studies assessing the effectiveness of the immune system in wild animals.

Adaptive responses of animals to climate change are most likely insufficient.

Biological responses to climate change have been widely documented across taxa and regions, but it remains unclear whether species are maintaining a good match between phenotype and environment, i.e. whether observed trait changes are adaptive. Here we reviewed 10,090 abstracts and extracted data from 71 studies reported in 58 relevant publications, to assess quantitatively whether phenotypic trait changes associated with climate change are adaptive in animals. A meta-analysis focussing on birds, the taxon best represented in our dataset, suggests that global warming has not systematically affected morphological traits, but has advanced phenological traits. We demonstrate that these advances are adaptive for some species, but imperfect as evidenced by the observed consistent selection for earlier timing. Application of a theoretical model indicates that the evolutionary load imposed by incomplete adaptive responses to ongoing climate change may already be threatening the persistence of species.

Blue Tits use fledgling quantity and quality as public information in breeding site choice.

Public information (PI), which is the information that can be derived from the behavior and performance of conspecifics, has been demonstrated to be used in many fitness-enhancing decisions. In the context of breeding habitat choice, PI use has been called "habitat copying." We experimentally tested the existence of habitat copying in the Blue Tit (Cyanistes caeruleus), a nonmigratory, short-lived hole-nesting bird. We manipulated the mean number of fledglings raised locally (quantity) and their condition (quality) as components of PI by transferring nestlings from Decreased (D) patches to Increased (I) patches. Our manipulations caused a negative relationship between fledgling quantity and quality that does not exist naturally: I patches had a higher number of fledglings that were in poorer condition, whereas D patches had a lower number in better condition. Control (C) patches, whether manipulated or not, had intermediate levels in terms of fledgling quantity and quality. Adult emigration the following year was higher from D than from C or I patches. Similarly, adult dispersal distance decreased for individuals coming from D to C to I patches. This suggests that resident breeders rely mainly on fledgling quantity to make emigration decisions. Emigration patterns of juveniles did not vary in relation to our patch manipulation. Immigration rates were higher and similar in I and D patches than in C patches. Hence, immigrant Blue Tits seem to rely on one of the manipulated components of PI and are insensitive to the discrepancy between fledgling quantity and quality. This shows that even nonmigratory species, such as Blue Tits, may use PI in their dispersal decisions but weigh its components differently for emigration and immigration. Differences among species in the importance of PI in breeding habitat choices may be explained by differences in life histories.

An experimental test of host's life history traits modulation in response to cuckoo parasitism risk.

Hosts can counteract parasites through defences based on resistance and/or tolerance. The mechanistic basis of tolerance, which involve defensive mechanisms minimizing parasite damage after a successful parasitic attack, remains poorly explored in the study of cuckoo-host interactions. Here, we experimentally explore the possibility that the risk of great spotted cuckoo Clamator glandarius parasitism may induce tolerance defences in magpie Pica pica hosts through plasticity in life-history traits. We predict that magpies exposed to auditory cues indicating high parasitism risk will more likely exhibit resistance and/or modify their life-history traits to minimize parasitism costs (i.e. tolerance) compared to magpies under low parasitism risk. We found that manipulating the perceived parasitism risk did not affect host resistance (i.e. rejection of parasitic eggs) nor host life-history traits. Unexpectedly, host's egg volume increased over the season in nests exposed to auditory cues of control non-harmful hoopoes Upupa epops. Our results do not provide support for inducible defences (either based on resistance or tolerance) in response to risk of parasitism in magpie hosts. Even so, we encourage studying plastic expression of breeding strategies in response to risk of cuckoo parasitism to achieve a better understanding of the mechanistic basis of tolerance defences.

Sex Difference in Condition Dependence of Carotenoid Gapes in the Eurasian Roller (Coracias garrulus).

In altricial birds, sex differentiation can start early in the ontogeny in the form of color, physiology, and/or growth and may potentially result in sex-specific condition dependence of traits mediating parent-offspring communication. Carotenoids have long been hypothesized to modulate the expression of gape coloration, but their sex-specific role enforcing honesty of gape coloration remains poorly studied. In a within-nest design, we provided carotenoid supplementation to nestlings of the Eurasian roller (Coracias garrulus) and measured the response in circulating carotenoids, coloration of the gape, cutaneous immune responsiveness to phytohemagglutinin, and growth while accounting for the sex of nestlings. Male nestlings supplemented with carotenoids displayed enhanced pigmentation of their gapes and grew faster than control nestlings, but there was no within-individual correlation between gape color and growth in either carotenoid-supplemented or control males. Female nestlings, however, diverted most supplemented carotenoids into growing fast at the expense of reducing their level of circulating carotenoids and displaying less-pigmented gapes. Nestling cutaneous immune response was not affected by carotenoid supplementation in either sex. Our results provide only weak support for the hypothesis that carotenoids enforce the honesty of gape color signals in nestling rollers and demonstrate sex specificity in how nestlings divert a surplus of carotenoids into different physiological functions.

Exploratory behaviour modulates the relationship between colony familiarity and helping in a cooperative bird.

Individuals within animal groups may differ in personality and degree of familiarity raising the question of how this influences their social interactions. In Iberian magpies Cyanopica cooki, a portion of first-year males engage in cooperative behaviours and dispersal, allowing addressing this question. In this study, we first investigate the relationship between colony familiarity (native versus foreign) and reproductive status (breeding versus helping) of males during 21 years. Secondly, we measure the exploratory behaviour and monitor reproductive status of a sample of individuals with different colony familiarity during 2 years. Long-term monitoring revealed that foreign individuals were more likely breeders. The analysis on the subset of individuals in which exploratory behaviour was measured revealed a mediatory effect of exploratory behaviour in the association between colony familiarity and helping behaviour. Specifically, among foreign individuals, higher explorative males were more frequently involved in helping behaviour than lower explorative ones. Conversely, among native males, breeders were more explorative than helpers. Our results suggest that aspects of personality may mediate the value of familiarity in reproductive tasks in social species.

Are female starlings able to recognize the scent of their offspring?

BACKGROUND: Although there is growing evidence that birds may have individual chemical profiles that can function in several social contexts, offspring recognition based on olfactory cues has never been explored. This ability should be more likely evolved in colonial birds and/or species suffering brood parasitism, in which the risk of being engaged in costly misdirected parental care is high. METHODOLOGY/PRINCIPAL FINDINGS: We performed a choice experiment to examine whether females of the spotless starling, Sturnus unicolor, a species that is colonial, and where a fraction of the population is exposed to intraspecific brood parasitism, can discriminate between the scent of their offspring and that of unrelated nestlings. We also explored whether the development of the uropygial gland secretion may play a role in such olfactory discrimination by performing the choice experiments to females rearing nestlings of two different ages, that is, without and with developed uropygial glands. Results showed that female starlings did not preferentially choose the scent of their offspring, independently of whether the gland of nestlings was developed or not. CONCLUSIONS/SIGNIFICANCE: Our results suggest that female starlings do not have or do not show the ability to distinguish their offspring based on olfaction, at least up to 12-14 days of nestling age. Further research is needed to examine whether odour-based discrimination may function when fledgling starlings leave the nest and the risk of costly misidentification is likely to increase.