Evolutionary predictors of the specific colors of birds.

Animal coloration is one of the most conspicuous aspects of human-perceived organismal diversity, yet also one of the least understood. In particular, explaining why species have specific colors (e.g., blue vs. red) has proven elusive. Here, we quantify for nearly all bird species, the proportion of the body covered by each of 12 human-visible color categories, and test whether existing theory can predict the direction of color evolution. The most common colors are black, white, gray and brown, while the rarest are green, blue, purple, and red. Males have more blue, purple, red, or black, whereas females have more yellow, brown, or gray. Sexual dichromatism is partly due to sexual selection favoring ornamental colors in males but not in females. However, sexual selection also correlated positively with brown in both sexes. Strong social selection favors red and black, colors used in agonistic signaling, with the strongest effects in females. Reduced predation risk selects against cryptic colors (e.g., brown) and favors specific ornamental colors (e.g., black). Nocturnality is mainly associated with brown. The effects of habitat use support the sensory drive theory for camouflage and signaling. Darker colors are more common in species living in wet and cold climates, matching ecogeographical rules. Our study unambiguously supports existing theories of color evolution across an entire class of vertebrates, but much variation remains unexplained.

Extra-pair paternity and sexual dimorphism in birds.

Differences in the strength of sexual selection between males and females can lead to sexual dimorphism. Extra-pair paternity (EPP) can increase the variance in male reproductive success and hence the opportunity for sexual selection. Previous research on birds suggests that EPP drives the evolution of dimorphism in plumage colour and in body size. Because EPP increases the intensity of sexual selection in males, it should lead to increased dimorphism in species with larger or more colourful males, but decreased dimorphism in species with larger or more colourful females. We explored the covariation between EPP and sexual dimorphism in wing length and plumage colouration in 401 bird species, while controlling for other, potentially confounding variables. Wing length dimorphism was associated positively with the frequency of EPP, but also with social polygamy, sex bias in parental behaviour and body size and negatively with migration distance. The frequency of EPP was the only predictor of plumage colour dimorphism. In support of our prediction, high EPP levels were associated with sexual dichromatism, positively in species in which males are more colourful and negatively in those in which females are more colourful. Contrary to our prediction, high EPP rates were associated with increased wing length dimorphism in species with both male- and female-biased dimorphism. The results support a role for EPP in the evolution of both size and plumage colour dimorphism. The two forms of dimorphism were weakly correlated and predicted by different reproductive, social and life-history traits, suggesting an independent evolution.

The evolution of carotenoid-based plumage colours in passerine birds.

Many birds use carotenoids to colour their plumage yellow to red. Because birds cannot synthesise carotenoids, they need to obtain these pigments from food, although some species metabolise dietary carotenoids (which are often yellow) into derived carotenoids (often red). Here, we study the occurrence of yellow and red carotenoid-based plumage colours in the passerines, the largest bird radiation and quantify the effects of potential ecological and life-history drivers on their evolution. We scored the presence/absence of yellow and red carotenoid-based plumage in nearly 6,000 species and use Bayesian phylogenetic mixed models to assess the effects of carotenoid-availability in diet, primary productivity, body size, habitat and sexual selection. We also test the widespread assumption that red carotenoid-based colours are more likely to be the result of metabolization. Finally, we analyse the pattern of evolutionary transitions between yellow and red carotenoid-based plumage colours to determine whether, as predicted, the evolution of yellow carotenoid-based colours precedes red. We show that, as expected, both colours are more likely to evolve in smaller species and in species with carotenoid-rich diets. Yellow carotenoid-based plumage colours, but not red, are more prevalent in species that inhabit environments with higher primary productivity and closed vegetation. In general, females were more likely to have yellow and males more likely to have red carotenoid-based plumage colours, closely matching the effects of sexual selection. Our analyses also confirm that red carotenoid-based colours are more likely to be metabolised than yellow carotenoid-based colours. Evolutionary gains and losses of yellow and red carotenoid-based plumage colours indicate that red colours evolved more readily in species that already deposited yellow carotenoids, while the reverse was rarely the case. Our study provides evidence for a general, directional evolutionary trend from yellow to red carotenoid-based colours, which are more likely to be the result of metabolization. This may render them potentially better indicators of quality, and thus favoured by sexual selection.

Sexual selection for extreme physical performance in a polygynous bird is associated with exceptional sex differences in oxygen carrying capacity.

In many animals, males compete for access to fertile females. The resulting sexual selection leads to sex differences in morphology and behaviour, but may also have consequences for physiology. Pectoral sandpipers are an arctic-breeding polygynous shorebird in which males perform elaborate displays around-the-clock and move over long distances to sample potential breeding sites, implying the need for physiological adaptations to cope with extreme endurance. We examined the oxygen carrying capacity of pectoral sandpipers, measured as the volume percentage of red blood cells in blood (haematocrit, Hct). We found a remarkable sex difference in Hct levels, with males having much higher values (58.9 ± 3.8 s.d.) than females (49.8 ± 5.3 s.d.). While Hct values of male pectoral sandpipers are notable for being among the highest recorded in birds, the sex difference we report is unprecedented and more than double that of any previously described. We also show that Hct values declined after arrival to the breeding grounds in females, but not in males, suggesting that males maintain an aerobic capacity during the mating period equivalent to that sustained during trans-hemispheric migration. We conclude that sexual selection for extreme physical performance in male pectoral sandpipers has led to exceptional sex differences in oxygen carrying capacity.

Breeding site sampling across the Arctic by individual males of a polygynous shorebird.

Males of many polygynous species compete for access to fertile females without providing them with resources other than sperm and without investing in care for the offspring (male dominance polygyny). In such systems, local competition for access to females is intense and typically only a few males obtain matings, leading to strong sexual selection. Sampling multiple breeding areas could then provide a mechanism for males to increase their chances to reproduce. However, little is known about such sampling behaviour and about the spatial scale at which males compete. Here we show that most males of a migratory, polygynous shorebird, the pectoral sandpiper (Calidris melanotos), that arrived at a known breeding location in northern Alaska subsequently moved through a considerable part of the entire species' breeding range (up to 13,045 km in a four-week period), sampling as many as 23 additional potential breeding sites. Our data suggest that males do not have a final breeding destination after migration from their wintering quarters, but make nomadic movements that are probably not a consequence of breeding failure. Tenure, the duration of stay at a site, correlated strongly with the number of breeding females at the site, suggesting that decisions to leave are dependent on local mating opportunities. Nomadic movements may allow males to display and sire offspring at multiple sites within a single breeding season. Sexual selection may then favour high-performance males that are able to reduce sleep to compete locally and to fly long distances between breeding sites, leading to a population with unrestricted interbreeding and without local adaptation and speciation.

Scrutinizing assortative mating in birds.

It is often claimed that pair bonds preferentially form between individuals that resemble one another. Such assortative mating appears to be widespread throughout the animal kingdom. Yet it is unclear whether the apparent ubiquity of assortative mating arises primarily from mate choice ("like attracts like"), which can be constrained by same-sex competition for mates; from spatial or temporal separation; or from observer, reporting, publication, or search bias. Here, based on a conventional literature search, we find compelling meta-analytical evidence for size-assortative mating in birds (r = 0.178, 95% CI 0.142-0.215, 83 species, 35,591 pairs). However, our analyses reveal that this effect vanishes gradually with increased control of confounding factors. Specifically, the effect size decreased by 42% when we used previously unpublished data from nine long-term field studies, i.e., data free of reporting and publication bias (r = 0.103, 95% CI 0.074-0.132, eight species, 16,611 pairs). Moreover, in those data, assortative mating effectively disappeared when both partners were measured by independent observers or separately in space and time (mean r = 0.018, 95% CI -0.016-0.057). Likewise, we also found no evidence for assortative mating in a direct experimental test for mutual mate choice in captive populations of Zebra finches (r = -0.020, 95% CI -0.148-0.107, 1,414 pairs). These results highlight the importance of unpublished data in generating unbiased meta-analytical conclusions and suggest that the apparent ubiquity of assortative mating reported in the literature is overestimated and may not be driven by mate choice or mating competition for preferred mates.

Unexpected diversity in socially synchronized rhythms of shorebirds.

The behavioural rhythms of organisms are thought to be under strong selection, influenced by the rhythmicity of the environment. Such behavioural rhythms are well studied in isolated individuals under laboratory conditions, but free-living individuals have to temporally synchronize their activities with those of others, including potential mates, competitors, prey and predators. Individuals can temporally segregate their daily activities (for example, prey avoiding predators, subordinates avoiding dominants) or synchronize their activities (for example, group foraging, communal defence, pairs reproducing or caring for offspring). The behavioural rhythms that emerge from such social synchronization and the underlying evolutionary and ecological drivers that shape them remain poorly understood. Here we investigate these rhythms in the context of biparental care, a particularly sensitive phase of social synchronization where pair members potentially compromise their individual rhythms. Using data from 729 nests of 91 populations of 32 biparentally incubating shorebird species, where parents synchronize to achieve continuous coverage of developing eggs, we report remarkable within- and between-species diversity in incubation rhythms. Between species, the median length of one parent's incubation bout varied from 1-19 h, whereas period length-the time in which a parent's probability to incubate cycles once between its highest and lowest value-varied from 6-43 h. The length of incubation bouts was unrelated to variables reflecting energetic demands, but species relying on crypsis (the ability to avoid detection by other animals) had longer incubation bouts than those that are readily visible or who actively protect their nest against predators. Rhythms entrainable to the 24-h light-dark cycle were less prevalent at high latitudes and absent in 18 species. Our results indicate that even under similar environmental conditions and despite 24-h environmental cues, social synchronization can generate far more diverse behavioural rhythms than expected from studies of individuals in captivity. The risk of predation, not the risk of starvation, may be a key factor underlying the diversity in these rhythms.

A global analysis of song frequency in passerines provides no support for the acoustic adaptation hypothesis but suggests a role for sexual selection.

Animals use acoustic signals for communication, implying that the properties of these signals can be under strong selection. The acoustic adaptation hypothesis predicts that species in dense habitats emit lower-frequency sounds than those in open areas because low-frequency sounds propagate further in dense vegetation than high-frequency sounds. Signal frequency may also be under sexual selection because it correlates with body size and lower-frequency sounds are perceived as more intimidating. Here, we evaluate these hypotheses by analysing variation in peak song frequency across 5,085 passerine species (Passeriformes). A phylogenetically informed analysis revealed that song frequency decreases with increasing body mass and with male-biased sexual size dimorphism. However, we found no support for the predicted relationship between frequency and habitat. Our results suggest that the global variation in passerine song frequency is mostly driven by natural and sexual selection causing evolutionary shifts in body size rather than by habitat-related selection on sound propagation.

The macroecology of extra-pair paternity in birds.

Extra-pair paternity (EPP) is a key aspect of the mating behaviour of birds and its frequency varies widely among populations and species. Several hypotheses predict patterns of geographical variation in the occurrence and frequency of EPP, but a global-scale study on variation in this trait is still lacking. We collected data on EPP from 663 populations of 401 avian species and explored the geographical variation in the frequency of EPP among populations, species and species assemblages. We modelled the variation in the frequency of EPP within the species' breeding range accounting for the specific ecological context of each population, and used the model predictions to compute frequencies of EPP at the level of species assemblages. A global map of assemblage-level EPP rates shows clear differences between zoogeographical realms, with the highest EPP values in the Nearctic realm. Our results show that the frequency of EPP (1) decreases with latitude and increases with the distance from the breeding range boundary within the species' breeding range, (2) is negatively associated with generation length and pair-bond duration among species, and (3) decreases with latitude at assemblage level. The latitudinal decline of EPP is consistent across zoogeographical realms.

The effects of life history and sexual selection on male and female plumage colouration.

Classical sexual selection theory provides a well-supported conceptual framework for understanding the evolution and signalling function of male ornaments. It predicts that males obtain greater fitness benefits than females through multiple mating because sperm are cheaper to produce than eggs. Sexual selection should therefore lead to the evolution of male-biased secondary sexual characters. However, females of many species are also highly ornamented. The view that this is due to a correlated genetic response to selection on males was widely accepted as an explanation for female ornamentation for over 100 years and current theoretical and empirical evidence suggests that genetic constraints can limit sex-specific trait evolution. Alternatively, female ornamentation can be the outcome of direct selection for signalling needs. Since few studies have explored interspecific patterns of both male and female elaboration, our understanding of the evolution of animal ornamentation remains incomplete, especially over broad taxonomic scales. Here we use a new method to quantify plumage colour of all ~6,000 species of passerine birds to determine the main evolutionary drivers of ornamental colouration in both sexes. We found that conspecific male and female colour elaboration are strongly correlated, suggesting that evolutionary changes in one sex are constrained by changes in the other sex. Both sexes are more ornamented in larger species and in species living in tropical environments. Ornamentation in females (but not males) is increased in cooperative breeders--species in which female-female competition for reproductive opportunities and other resources related to breeding may be high. Finally, strong sexual selection on males has antagonistic effects, causing an increase in male colouration but a considerably more pronounced reduction in female ornamentation. Our results indicate that although there may be genetic constraints to sexually independent colour evolution, both female and male ornamentation are strongly and often differentially related to morphological, social and life-history variables.

Wind conditions influence breeding season movements in a nomadic polygynous shorebird.

Nomadism is a behaviour where individuals respond to environmental variability with movements that seem unpredictable in timing and direction. In contrast to migration, the mechanisms underlying nomadic movements remain largely unknown. Here, we focus on a form of apparent nomadism in a polygynous shorebird, the pectoral sandpiper (Calidris melanotos). Local mating opportunities are unpredictable and most males sampled multiple sites across a considerable part of their breeding range. We test the hypothesis that individuals decided which part of the breeding range to sample in a given season based on the prevailing wind conditions. Using movement data from 80 males in combination with wind data from a global reanalysis model, we show that male pectoral sandpipers flew with wind support more often than expected by chance. Stronger wind support led to increased ground speed and was associated with a longer flight range. Long detours (loop-like flights) can be explained by individuals flying initially with the wind. Individuals did not fly westwards into the Russian Arctic without wind support, but occasionally flew eastwards into the North American Arctic against strong headwinds. Wind support might be less important for individuals flying eastwards, because their autumn migration journey will be shorter. Our study suggests that individuals of a species with low site fidelity choose their breeding site opportunistically based on the prevailing wind conditions.

Body size and climate as predictors of plumage colouration and sexual dichromatism in parrots.

Psittaciformes (parrots, cockatoos and lorikeets) comprise one of the most colourful clades of birds. Their unique pigments and safe cavity nesting habits are two potential explanations for their colourful character. However, plumage colour varies substantially between parrot species and sometimes also between males and females of the same species. Here, we use comparative analyses to evaluate what factors correlate with colour elaboration, colour diversity and sexual dichromatism. Specifically, we test the association between different aspects of parrot colouration and (a) the intensity of sexual selection and social interactions, (b) variation along the slow-fast life-history continuum and (c) climatic variation. We show that larger species and species that live in warm environments display more elaborated colours, yet smaller species have higher levels of sexual dichromatism. Larger parrots tend to have darker and more blue and red colours. Parrots that live in wetter environments are darker and redder, whereas species inhabiting warm regions have more blue plumage colours. In general, each of the variables we considered explain small to moderate amounts of variation in parrot colouration (up to 15%). Our data suggest that sexual selection may be acting more strongly on males in small, short-lived parrots leading to sexual dichromatism. More elaborate colouration in both males and females of the larger, long-lived species with slow tropical life histories suggests that mutual mate choice, social selection and reduced selection for crypsis may be important in these species, as has been shown for passerines.

Morph-dependent fitness and directional change of morph frequencies over time in a Dutch population of Common buzzards Buteo buteo.

How genetic polymorphisms are maintained in a population is a key question in evolutionary ecology. Previous work on a plumage colour polymorphism in the common buzzard Buteo buteo suggested heterozygote advantage as the mechanism maintaining the co-existence of three morphs (light, intermediate and dark). We took advantage of 20 years of life-history data collected in a Dutch population to replicate earlier studies on the relationship between colour morph and fitness in this species. We examined differences between morphs in adult apparent survival, breeding success, annual number of fledglings produced and cumulative reproductive success. We found that cumulative reproductive success differed among morphs, with the intermediate morph having highest fitness. We also found assortative mating for colour morph, whereby assortative pairs were more likely to produce offspring and had longer-lasting pair bonds than disassortative pairs. Over the 20-year study period, the proportion of individuals with an intermediate morph increased. This apparent evolutionary change did not just arise from selection on individual phenotypes, but also from fitness benefits of assortative mating. The increased frequency of intermediates might also be due to immigration or drift. We hypothesize that genetic variation is maintained through spatial variation in selection pressures. Further studies should investigate morph-dependent dispersal behaviour and habitat choice.

Timing of arrival in the breeding area is repeatable and affects reproductive success in a non-migratory population of blue tits.

Events in one part of the annual cycle often affect the performance (and subsequently fitness) of individuals later in the season (carry-over effects). An important aspect of this relates to the timing of activities. For example, many studies on migratory birds have shown that relatively late-spring arrival in the breeding area reduces both the likelihood of getting a mate or territory and reproductive success. In contrast, relatively little is known about the movements of individuals in non-migratory populations during the non-breeding season. Few studies have investigated the timing of arrival at the breeding area in such species, possibly due to the assumption that most individuals remain in the area during the non-breeding season. In this study, we used 4 years of data from a transponder-based automated recording system set up in a non-migratory population of blue tits Cyanistes caeruleus to describe individual variation in arrival at the breeding site. We investigated whether this variation can be explained by individual characteristics (sex, body size or status), and we assessed its effect on aspects of reproductive success in the subsequent breeding season. We found substantial variation in arrival date and demonstrate that this trait is individual-specific (repeatable). Females arrived later than males, but the arrival dates of social pair members were more similar than expected by chance, which suggests that individuals may mate assortatively depending on their arrival in the breeding area. Arrival predicted both whether an individual would end up breeding that season and several aspects of its breeding success. Our study suggests that individuals of non-migratory species leave the breeding area during the non-breeding season. Hence, it may be useful to consider variation in the scale of movements between breeding and non-breeding sites, rather than using a simple dichotomy between 'resident' and 'migratory' species. We conclude that the timing of pre-breeding events, in particular arrival date, may be an overlooked, but important, fitness-relevant trait in non-migratory species.

Exposure to predator models during the fertile period leads to higher levels of extra-pair paternity in blue tits.

The perceived risk of predation can affect breeding behaviour and reduce reproductive success in prey species. Individuals exposed to predators may also adopt different mating tactics with potential consequences for the distribution of paternity in socially monogamous species that engage in extra-pair copulations. We experimentally increased perceived predation risk during the fertile period in blue tits Cyanistes caeruleus. Every morning between nest completion and the onset of egg laying, we presented a model of either a predator or a non-predator (control) near active nestboxes. Broods from pairs exposed to predators had higher levels of extra-pair paternity than control broods. This mainly resulted from a higher proportion of extra-pair offspring in broods with at least one extra-pair young. Females exposed to predators first emerged from the nestbox later in the morning, stayed away from the nestbox for longer and were less likely to be visited at the nest by their social mate, but we detected no behavioural differences once the model was removed. Our results suggest that the higher rates of extra-pair paternity resulted from the disruption of morning routines, which may have inhibited within-pair copulations or increased opportunities for females to engage in extra-pair copulations. We conclude that the perceived risk of predation can have substantial effects on levels of extra-pair paternity.

Partial or complete? The evolution of post-juvenile moult strategies in passerine birds.

Moulting strategies in birds have evolved to avoid overlap with, or prepare for, other demanding parts of the annual cycle, such as reproduction or migration. When moulting for the first time after leaving the nest, young birds replace their typically poor-quality plumage during the post-juvenile moult. The extent of this moult varies between species from partial to complete. Earlier studies, restricted to Western Palearctic birds, suggest that in most species a complete post-juvenile moult may not be possible simply because young birds are constrained by not having the same access to resources as adults, unless environmental conditions are favourable. These studies also show that complete post-juvenile moult is more common in species with poor-quality nest-grown plumage. We expanded the spatial and taxonomic scope of previous studies to 1,315 species of passerines from across the world and considered both the role of constraints, plumage quality and other selective pressures favouring a complete post-juvenile moult. Thus, we test whether complete moult is more prevalent in species where nest-grown feathers are presumably of poor quality (shorter nestling period), that live in environments that foster quick plumage degradation (open habitats, high insolation and humidity), and where males are under strong sexual selection. Our data reveal that 24% of species carry out a complete post-juvenile moult, and that this trait has a strong phylogenetic signal. Complete moult is more common in species that inhabit warmer regions and open habitats, show no delayed plumage maturation and have higher levels of sexual dichromatism (indicative of strong sexual selection). Neither the presumed quality of the nest-grown plumage nor living in regions with high insolation correlates with complete moult. In conclusion, the evolution of complete post-juvenile moult not only depends on whether birds can perform a complete moult (i.e. suitable environmental conditions) but also on the strength of selection associated with the need of a complete moult. In particular, the necessity to keep the plumage in good condition in challenging environments and the benefits associated with producing adult-like plumage colours to attract mates or deter rivals seem to play an important role.

Reconciling ecogeographical rules: rainfall and temperature predict global colour variation in the largest bird radiation.

Ecogeographical rules that associate climate with organismal form and function can reveal patterns of climatic adaptation. Two rules link animal coloration with climate: Gloger's rule (darker coloration where wet and warm), and Bogert's rule (darker coloration where cold). Whereas Gloger's rule was proposed for endotherms, and Bogert's rule for ectotherms, both rules may apply more broadly, despite their seemingly opposing effects. Here, we test this contradiction on a global scale across passerine birds. Consistent with Gloger's rule, birds were darker in wetter areas and, following Bogert's rule, lighter where warm, although birds became lighter again at very low temperatures. Rainfall and temperature had antagonistic or additive effects depending on their pattern of covariation, and this predicted whether birds followed the rules. We integrate both rules into a general framework to explain heterogeneity in climatic effects on coloration, which has implications to understand patterns of diversification, climatic adaptation and climate change impacts.

Evolutionary drivers of seasonal plumage colours: colour change by moult correlates with sexual selection, predation risk and seasonality across passerines.

Some birds undergo seasonal colour change by moulting twice each year, typically alternating between a cryptic, non-breeding plumage and a conspicuous, breeding plumage ('seasonal plumage colours'). We test for potential drivers of the evolution of seasonal plumage colours in all passerines (N = 5901 species, c. 60% of all birds). Seasonal plumage colours are uncommon, having appeared on multiple occasions but more frequently lost during evolution. The trait is more common in small, ground-foraging species with polygynous mating systems, no paternal care and strong sexual dichromatism, suggesting it evolved under strong sexual selection and high predation risk. Seasonal plumage colours are also more common in species predicted to have seasonal breeding schedules, such as migratory birds and those living in seasonal climates. We propose that seasonal plumage colours have evolved to resolve a trade-off between the effects of natural and sexual selection on colouration, especially in seasonal environments.

Sperm morphology and evidence for sperm competition among parrots.

Sperm competition is an important component of post-copulatory sexual selection that has shaped the evolution of sperm morphology. Previous studies have reported that sperm competition has a concurrently directional and stabilizing effect on sperm size. For example, bird species that show higher levels of extrapair paternity and larger testes (proxies for the intensity of sperm competition) have longer sperm and lower coefficients of variation in sperm length, both within and between males. For this reason, these sperm traits have been proposed as indexes to estimate the level of sperm competition in species for which other measures are not available. The relationship between sperm competition and sperm morphology has been explored mostly for bird species that breed in temperate zones, with the main focus on passerine birds. We measured sperm morphology in 62 parrot species that breed mainly in the tropics and related variation in sperm length to life-history traits potentially indicative of the level of sperm competition. We showed that sperm length negatively correlated with the within-male coefficient of variation in sperm length and positively with testes mass. We also showed that sperm is longer in sexually dichromatic and in gregarious species. Our results support the general validity of the hypothesis that sperm competition drives variation in sperm morphology. Our analyses suggest that post-copulatory sexual selection is also important in tropical species, with more intense sperm competition among sexually dichromatic species and among species that breed at higher densities.

Interference competition pressure predicts the number of avian predators that shifted their timing of activity.

Being active at different times facilitates the coexistence of functionally similar species. Hence, time partitioning might be induced by competition. However, the relative importance of direct interference and indirect exploitation competition on time partitioning remains unclear. The aim of this study was to investigate the relative importance of these two forms of competition on the occurrence of time-shifting among avian predator species. As a measure of interference competition pressure, we used the species richness of day-active avian predator species or of night-active avian predator species (i.e. species of Accipitriformes, Falconiformes and Strigiformes) in a particular geographical area (assemblage). As an estimate of exploitation competition pressure, we used the total species richness of avian predators in each assemblage. Estimates of the intensity of interference competition robustly predicted the number of Accipitriformes species that became crepuscular and the number of Strigiformes species that became day-active or strictly crepuscular. Interference competition pressure may depend on body size and on the total duration of the typical active period (day or night length). Our results support-to some extent-that smaller species are more likely to become time-shifters. Day length did not have an effect on the number of time-shifter species in the Accipitriformes. Among the large Strigiformes, more time-shifter species occur in areas where nights are shorter (i.e. where less of the typical time resource is available). However, in the small Strigiformes, we found the opposite, counterintuitive effect: more time-shifters where nights are longer. Exploitation competition may have had an additional positive effect on the number of time-shifters, but only in Accipitriformes, and the effect was not as robust. Our results thus support the interference competition hypothesis, suggesting that animals may have shifted their time of activity, despite phylogenetic constraints on the ability to do so, to reduce the costs of direct interactions. Our findings also highlight the influence of body size as a surrogate of competitive ability during encounters on time partitioning, at least among avian predators.