Social rules govern vocal competition in the barn owl

To resolve the share of limited resources, animals often compete through exchange of signals about their relative motivation to compete. When two competitors are similarly motivated, the resolution of conflicts may be achieved in the course of an interactive process. In barn owls, Tyto alba, in which siblings vocally compete during the prolonged absence of parents over access to the next delivered food item, we investigated what governs the decision to leave or enter a contest, and at which level. Siblings alternated periods during which one of the two individuals vocalized more than the other. Individuals followed turn-taking rules to interrupt each other and momentarily dominate the vocal competition. These social rules were weakly sensitive to hunger level and age hierarchy. Hence, the investment in a conflict is determined not only by need and resource-holding potential, but also by social interactions. The use of turn-taking rules governing individual vocal investment has rarely been shown in a competitive context. We hypothesized that these rules would allow individuals to remain alert to one another's motivation while maintaining the cost of vocalizing at the lowest level.

Habitat, breeding performance, diet and individual age in Swiss barn owls (Tyto alba)

Intensification of farming over the past 50 years has homogenised the landscape structure and contributed to the decline of bird populations in Europe. To better target the conservation of the Barn Owl Tyto alba, we assessed the influence of the landscape structure on breeding performance in western Switzerland. The analyses considered a 23-year data set of breeding parameters collected in an area dominated by intensive agriculture. Using a Geographic Information System approach, landscape characteristics were described around 194 nest sites. Our analyses showed that nest-box occupancy, laying date, clutch and brood size, egg volume and probability of producing a second annual clutch were not significantly associated with any of the eight principal landscape variables (agricultural land, woodland, urban area, hedgerows, cereals, sugar beet, maize and meadow). Nevertheless, the probability that a breeding pair occupied a nest-box decreased the more roads there were surrounding the nest-box. The absence of strong associations between habitat features and breeding parameters suggests that prey availability may be relatively similar between the different breeding sites. In our study area Barn Owls can always find suitable foraging habitats around most nest-boxes.

Female- and male-specific signals of quality in the barn owl

Most bird studies of female signalling have been confined to species in which females display a male-ornament in a vestigial form. However, a great deal of benefit may be gained from considering phenotypic traits that are specific to females. This is because (1) sex-specific traits may signal sex-specific qualities and (2) females may develop a male-ornament not because they are selected to do so, but because fathers transmit to daughters the underlying genes for its expression (genetic correlation between the sexes). We investigated these two propositions in the barn owl Tyto alba, a species in which male plumage is lighter in colour and less marked with black spots than that of females. Firstly, we present published evidence that female plumage spottiness reflects parasite resistance ability. We also show that male plumage coloration is correlated with reproductive success, male feeding rate and heart mass. Secondly, cross-fostering experiments demonstrate that plumage coloration and spottiness are genetically correlated between the sexes. This implies that if a given trait value is selected in one sex, the other sex will indirectly evolve towards a similar value. This prediction is supported by the observation that female plumage coloration and spottiness resembled that of males, in comparisons at the level of Tyto alba alba populations, Tyto alba subspecies and Tyto species. Our results therefore support the hypothesis that sex-specific traits signal sex-specific qualities and that a gene for a sex-specific trait can be expressed in the other sex as the consequence of a genetic correlation between the sexes.

Effects of corticosterone on innate and humoral immune functions and oxidative stress in barn owl nestlings.

The costs of coping with stressful situations are traded-off against other functions such as immune responses. This trade-off may explain why corticosterone secretion reduces immune reactions. Corticosterone differentially affects various immunity components. However, which component is suppressed varies between studies. It remains unclear whether the trade-off in energy, nutrition, autoimmunity or oxidative stress accounts for differential immunosuppression. In this study, we investigated whether corticosterone differentially affects the constitutive innate and humoral acquired immunity. We used barn owl nestlings, implanting 50% with a corticosterone-releasing pellet and the other 50% with a placebo pellet. To measure the effect on humoral immunity we vaccinated 50% of the corticosterone-nestlings and 50% of the placebo-nestlings with the antigens 'Tetravac' and the other 50% were injected with PBS. To assess the costs of elevated corticosterone, we measured body mass and resistance to oxidative stress. Administration of corticosterone increased corticosterone levels whereas vaccination induced the production of antibodies. Corticosterone reduced the production of antibodies, but it did not significantly affect the constitutive innate immunity. Corticosterone reduced body growth and resistance to oxidative stress. Under stressful conditions barn owl nestlings seem to keep the constitutive innate immunity, whereas elevated corticosterone levels negatively affected inducible immune responses. We found evidence that mounting a humoral immune reaction is not costly in terms of growth, but reduces the resistance to oxidative stress independently of corticosterone administration. We suggest that humoral immunity is suppressed because the risk of immunopathologies may be disproportionately high when mounting an antibody response under stressful situations.

Divorce in the barn owl: securing a compatible or better mate entails the cost of re-pairing with a less ornamented female mate.

Two nonmutually exclusive hypotheses can explain why divorce is an adaptive strategy to improve reproductive success. Under the 'better option hypothesis', only one of the two partners initiates divorce to secure a higher-quality partner and increases reproductive success after divorce. Under the 'incompatibility hypothesis', partners are incompatible and hence they may both increase reproductive success after divorce. In a long-term study of the barn owl (Tyto alba), we address the question of whether one or the two partners derive fitness benefits by divorcing. Our results support the hypothesis that divorce is adaptive: after a poor reproductive season, at least one of the two divorcees increase breeding success up to the level of faithful pairs. By breeding more often together, faithful pairs improve coordination and thereby gain in their efficiency to produce successful fledglings. Males would divorce to obtain a compatible mate rather than a mate of higher quality: a heritable melanin-based signal of female quality did not predict divorce (indicating that female absolute quality may not be the cause of divorce), but the new mate of divorced males was less melanic than their previous mate. This suggests that, at least for males, a cost of divorce may be to secure a lower-quality but compatible mate. The better option hypothesis could not be formally rejected, as only one of the two divorcing partners commonly succeeded in obtaining a higher reproductive success after divorce. In conclusion, incompatible partners divorce to restore reproductive success, and by breeding more often together, faithful partners improve coordination.

Personality and melanin-based colouration traits in barn owls, tawny owl and kestrels

Individuals need to adapt to their local environment in order to survive. When selection pressures differ in local populations, polymorphism can evolve. Colour polymorphism is one of the most obvious polymorphisms since it is readily observable. Different sources of colouration exist, but melanin-based colouration is one of the most common in birds. The melanocortin system produces this colouration and because the melanocortin system has pleiotropic effects on behavioural and physiological traits, it is a good candidate to be an underlying mechanism to explain the maintenance of colour polymorphism. In this thesis I studied three different raptors which all display melanin-based colouration; barn owls (Tyto alba), tawny owls (Strix aluco) and Eurasian kestrels (Falco tinnunculus). The main question was if there was a relationship between melanin-based colouration and individual behavioural differences. The underlying hypothesis is that colour could be a signal of certain adaptive traits. Our goal was to find evolutionary explanations for the persistence of colour polymorphism. I found that nestling kestrels and barn owls differ in anti-predatory behaviour, with respect to their melanic colouration (chapters 1 and 2). Darker individuals show less reaction to human handling, but in kestrels aggression and colouration are related in opposite ways than in barn owls. More reddish barn owls travel greater distances in natal dispersal and this behaviour is repeatable between parents and same sex offspring (chapter 3). Dark reddish tawny owls defend their nests more intensely against intruders and appear to suffer less from nest predation (chapter 4). Finally I show that polymorphism in the Melanocortin 1 receptor gene (MC1R), which is strongly correlated with reddish colouration in the barn owl, is related to natal dispersal distance, providing a first indication for a genetic basis of the relation between this behaviour and colouration (chapter 5). My results demonstrate a clear link between melanin-based colouration and animal personality traits. I demonstrated this relation in three different species, which shows there is most likely a general underlying mechanism responsible. Different predation pressures might have shaped the reactions to predation, but also differences in sex-related colouration. Male-like and female-like colouration might signal more or less aggressive behaviour. Fluctuating environmental conditions might cause different individual strategies to produce equal reproductive success. The melanocortin system with its pleiotropic effects might be an underlying mechanism, as suggested by the results from the genetic polymorphism, the similar results found in these three species and by the similar relations reported in other species. This thesis demonstrates that colouration and individual differences are correlated and it provides the first glimpse of an underlying system. We can now conduct a more directed search for underlying mechanisms and evolutionary explanations with the use of quantitative genetic methods.

Sexually dimorphic melanin-based colour polymorphism, feather melanin content, and wing feather structure in the barn owl (Tyto alba)

Feathers confer protection against biophysical agents and determine flying ability. The geometry and arrangement of the barbs, together with the keratin and pigments deposited in the feathers, determine the mechanical stability of the vane, and its stiffness and resistance to abrasive agents. In colour-polymorphic species, individuals display alternative colour morphs, which can be associated with different foraging strategies. Each morph may therefore require specific flying abilities, and their feathers may be exposed to different abrasive agents. Feathers of differently coloured individuals may thus have a specific structure, and colour pigments may help resist abrasive agents and improve stiffness. We examined these predictions in the barn owl (Tyto alba), a species for which the ventral body side varies from white to dark reddish pheomelanic, and in the number and size of black spots located at the tip of the feathers. White and reddish birds show different foraging strategies, and the size of black feather spots is associated with several phenotypic attributes. We found that birds displaying a darker reddish coloration on the ventral body side deposit more melanin pigments in their remiges, which also have fewer barbs. This suggests that wear resistance increases with darkness, whereas feathers of lighter coloured birds may bend less easily. Accordingly, individuals displaying a lighter reddish coloration on the ventral body side, and those displaying larger black spots, displayed more black transverse bars on their remiges: as larger-spotted individuals are heavier and longer-winged birds also have more transverse bars, these bars may reduce feather bending when flying. We conclude that differently coloured individuals produce wing feathers of different strengths to adopt alternative behavioural and life history strategies

Selection on a eumelanic ornament is stronger in the tropics than in temperate zones in the worldwide-distributed barn owl.

Spatial variation in the pattern of natural selection can promote local adaptation and genetic differentiation between populations. Because heritable melanin-based ornaments can signal resistance to environmentally mediated elevation in glucocorticoids, to oxidative stress and parasites, populations may vary in the mean degree of melanic coloration if selection on these phenotypic aspects varies geographically. Within a population of Swiss barn owls (Tyto alba), the size of eumelanic spots is positively associated with survival, immunity and resistance to stress, but it is yet unknown whether Tyto species that face stressful environments evolved towards a darker eumelanic plumage. Because selection regimes vary along environmental gradients, we examined whether melanin-based traits vary clinally and are expressed to a larger extent in the tropics where parasites are more abundant than in temperate zones. To this end, we considered 39 barn owl species distributed worldwide. Barn owl species living in the tropics displayed larger eumelanic spots than those found in temperate zones. This was, however, verified in the northern hemisphere only. Parasites being particularly abundant in the tropics, they may promote the evolution of darker eumelanic ornaments.

El Cancionero del Barón de Claret: fama de Lope de Vega y Calderón de la Barca en la Cataluña de la segunda mitad del siglo XVII

Comentari del llibre Cancionero del Barón de Claret, compilat a Catalunya. El comentari es centra en la influència de Lope de Vega i de Calderón de la Barca sobre aquesta obra. Resulta més influent Lope de Vega, malgrat que en el moment de la compilació de la obra ja no estava de moda, que Calderón de la Barca

Sib-sib communication and the risk of prey theft in the barn owl Tyto alba

Conflicts among siblings are widespread and their resolution involves complex physical and communication tools. Observations in the barn owl Tyto alba showed that siblings vocally communicate in the absence of parents to negotiate priority of access to the impending food resources that parents will bring. In the present paper, we hypothesize and provide correlative evidence that after a parent brought a food item to their progeny, sibling competition involves vocal sib-sib communication. A food item takes a long time to be entirely consumed, and hence siblings continue to compete over prey monopolization even after parents gave a food item to a single offspring. When physical competition is pronounced and thereby the risk of prey theft is high, the individual that received a prey item consumes it in a concealed place. Concomitantly, nestlings vocalize intensely probably to indicate their motivation to siblings to not share their food item, since this vocal behaviour was particularly frequent in younger individuals for which the risk of being robbed is higher than in their older siblings. Furthermore, nestlings consumed more rapidly a food item when their siblings vocalized intensely presumably because the intensity of siblings' vocalizations is associated with a risk of prey theft. Our correlative study suggests that sibling competition favoured the evolution of sib-sib communication under a wide range of situations.

Dominant nestlings displaying female-like melanin coloration behave altruistically in the barn owl

When competing over parental resources, young animals may be typically selfish to the point of siblicide. This suggests that limited parental resources promote the evolution of sibling competition rather than altruistic or cooperative behaviours. In striking contrast, we show here that in 71% of experimental three-chick broods, nestling barn owls, Tyto alba, gave food to their siblings on average twice per night. This behaviour prevailed in the first-born dominant nestlings rather than the last-born subordinate nestlings. It was also more prevalent in individuals displaying a heritable dark phaeomelanin-based coloration, a typical female-specific plumage trait (owls vary from dark reddish to white, females being on average darker reddish than males). Stealing food items from siblings, which occurred in 81% of the nests, was more frequent in light than dark phaeomelanic dominant nestlings. We suggest that food sharing has evolved in the barn owl because parents store prey items in their nest that can be used by the offspring to feed their nestmates to derive indirect (kin selection) or direct benefits (pseudoreciprocity or by-product mutualism). The cost of feeding siblings may be relatively low for dominant individuals while the indirect genetic benefits could be high given that extrapair paternity is infrequent in this species. Thus, in situations in which young animals have access to more food resources than they currently need, they can altruistically share them with their siblings.

Similar patterns of local barn owl adaptation in the Middle East and Europe with respect to melanic coloration

The maintenance of phenotypic variation is a central question in evolutionary biology. A commonly suggested mechanism is that of local adaptation, whereby different phenotypes are adapted to alternative environmental conditions. A recent study in the European barn owl (Tyto alba) has shown that natural selection maintains a strong clinal variation in reddish pheomelanin-based coloration. Studies in the region where phenotypic variation in this owl is the highest in Europe have further demonstrated that dark-reddish and pale-reddish owls exploit open and wooded habitats, predate voles and wood mice, and are long-tailed and short-tailed, respectively. However, it remains unclear as to whether these traits evolved as a consequence of allopatric evolution of dark colour in northern Europe and white colour in southern Europe, during which owls could have also evolved different morphologies and foraging behaviour. This scenario implies that covariation between coloration and foraging behaviour could be a specificity of the European continent, which is not found in other worldwide-distributed populations. To investigate this issue we studied a barn owl population in the Middle East. Our results show that, as in Central Europe, dark-reddish female owls breed more often in the open landscape than their pale-reddish female conspecifics, their offspring are fed with more voles than Muridae, and they are longer-winged and longer-tailed. These findings indicate that in the barn owl the association in females between pheomelanin-based coloration and foraging behaviour and morphology is not restricted to the European continent but may well evolve in sympatry in many barn owl populations worldwide.

Selection on a eumelanic ornament is stronger in the tropics than in temperate zones in the worldwide-distributed barn owl.

Spatial variation in the pattern of natural selection can promote local adaptation and genetic differentiation between populations. Because heritable melanin-based ornaments can signal resistance to environmentally mediated elevation in glucocorticoids, to oxidative stress and parasites, populations may vary in the mean degree of melanic coloration if selection on these phenotypic aspects varies geographically. Within a population of Swiss barn owls (Tyto alba), the size of eumelanic spots is positively associated with survival, immunity and resistance to stress, but it is yet unknown whether Tyto species that face stressful environments evolved towards a darker eumelanic plumage. Because selection regimes vary along environmental gradients, we examined whether melanin-based traits vary clinally and are expressed to a larger extent in the tropics where parasites are more abundant than in temperate zones. To this end, we considered 39 barn owl species distributed worldwide. Barn owl species living in the tropics displayed larger eumelanic spots than those found in temperate zones. This was, however, verified in the northern hemisphere only. Parasites being particularly abundant in the tropics, they may promote the evolution of darker eumelanic ornaments.

A record of communal nesting in the barn owl (Tyto alba)

We report a unique case of two female Barn Owls laying eggs and incubating together in a single nest cup in a communal nest. A trio of two females and one male bred in an abandoned water tower in 2013 in Israel. Both females incubated/brooded together in the communal nest, and all three individuals brought food to the communal family. The two females laid 20 eggs, of which 19 hatched and 16 fledged.

Wildflower areas within revitalized agricultural matrices boost small mammal populations but not breeding Barn Owls

Agro-ecosystems have recently experienced dramatic losses of biodiversity due to more intensive production methods. In order to increase species diversity, agri-environment schemes provide subsidies to farmers who devote a fraction of their land to ecological compensation areas (ECA). Several studies have shown that invertebrate biodiversity is actually higher in ECA than in nearby intensively cultivated farmland. It remains poorly understood, however, to what extent ECA also favour vertebrates, such as small mammals and their predators, which would contribute to restore functioning food chains within revitalized agricultural matrices. We studied small mammal populations among eight habitat types - including wildflower areas, a specific ECA in Switzerland - and habitat selection (radiotracking) by the barn owl Tyto alba, one of their principal predators. Our prediction was that habitats with higher abundances of small mammals would be more visited by foraging Barn owls during the period of chicks' provisioning. Small mammal abundance tended to be higher in wildflower areas than in any other habitat type. Barn owls, however, preferred to forage in cereal fields and grassland. They avoided all types of crops other than cereals, as well as wildflower areas, which suggests that they do not select their hunting habitat primarily with respect to prey density. Instead of prey abundance, prey accessibility may play a more crucial role: wildflower areas have a dense vegetation cover, which may impede access to prey for foraging owls. The exploitation of wildflower areas by the owls might be enhanced by creating open foraging corridors within or around wildflower areas. Wildflower areas managed in that way might contribute to restore functioning food chains within agro-ecosystems.