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Within species, bird parents defend their nests with different levels of intensity. Previous studies have examined several hypotheses for this variation by looking at factors such as morphological characteristics of the parents and the characteristics of the brood, but few have controlled for multiple factors simultaneously. Here we examined a broad range of factors that may influence the nest defense behaviour of Mountain Bluebirds ( Sialia currucoides (Bechstein, 1798)), including the adult's morphology (sex, age, plumage colour, body size, and body condition) and characteristics of their brood (size, hatch date, and stage of nestling development). Parental aggression was triggered using a taxidermy mount of an American red squirrel ( Tamiasciurus hudsonicus (Erxleben, 1777)). We found no relationship between nest defense and any morphological characteristic of adults, but the intensity of nest defense was positively correlated with the stage of offspring development, clutch size, and hatch date consistent with parental investment theory. Males also defended nests more strongly than females, consistent with the premise of the renesting hypothesis that males have fewer future opportunities to breed and, therefore, are more invested in the current brood. Our results provide strong support for parental investment theory and renesting as the primary drivers of nest defense intensity in breeding birds.

Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young., This meta-analysis was financed by the grant of the Polish National Science Centre (Narodowe Centrum Nauki) (no. 2017/27/B/NZ8/00465) awarded to Lucyna Hałupka.

Highly mobile predators can show strong numerical responses to pulsed resources, sometimes resulting in irruptions where large numbers of young invade landscapes at a continental scale. High production of young in irruption years may have a strong influence on the population dynamics unless immature survival is reduced compared to non-irruption years. This could occur if subordinate individuals (mainly immatures) are forced into suboptimal habitats due to density-dependent effects in irruption years. To test whether irruptive individuals had lower survival than non-irruptive ones, we combined necropsy results (N = 365) with telemetry (N = 185) from more than 20 years to record timing and causes of mortality in snowy owls (Bubo scandiacus), which irrupt into eastern North America during winter following high breeding output caused by lemming peaks in the Arctic. Mortality was more than four times higher in irruption years than non-irruption years, but only for immatures, and occurred disproportionately in early winter for immatures, but not adults. Mortality was also higher in eastern North America, where owl abundance fluctuates considerably between years, compared to core winter regions of the Arctic and Prairies where populations are more stable. Most mortality was not due to starvation, but rather associated with human activity, especially vehicle collisions. We conclude that immature snowy owls that irrupt into eastern North America are limited by density-dependent factors, such as increased competition forcing individuals to occupy risky human-altered habitats. For highly mobile, irruptive animals, resource pulses may have a limited impact on population dynamics due to low subsequent survival of breeding output during the nonbreeding season.

Genetic analysis can provide valuable information for conservation programs by unraveling the demographic trajectory of populations, by estimating effective population size, or by inferring genetic differentiation between populations. Here, we investigated the genetic differentiation within the Snowy Owl (Bubo scandiacus), a species identified as vulnerable by the IUCN, to (i) quantify connectivity among wintering areas, (ii) to evaluate current genetic diversity and effective population size and (iii) to infer changes in the historical effective population size changes from the last millennia to the recent past. The Snowy Owl, a highly mobile top predator, breeds across the Arctic tundra which is a region especially sensitive to current climate change. Using SNP-based analyses on Snowy Owls sampled across the North American nonbreeding range, we found an absence of genetic differentiation among individuals located up to 4,650 km apart. Our results suggest high genetic intermixing and effective dispersal at the continental scale despite documented philopatry to nonbreeding sites in winter. Reconstructing the population demographic indicated that North American Snowy Owls have been steadily declining since the Last Glacial Maximum ca 20,000 years ago and concurrently with global increases in temperature. Conservation programs should now consider North American Snowy Owls as a single, genetically homogenous continental-wide population which is most likely sensitive to the long-term global warming occurring since the Last Glacial Maximum.

Divorce is widespread among species of birds and may either be an adaptive strategy to secure a better mate or territory or be a nonadaptive result of a failure to maintain the pairbond. I examined the causes and consequences for divorce in the Northern Flicker (Colaptes auratus), a migratory woodpecker with a high annual mortality rate. In a long-term population study of 1,793 breeding pairs over 17 years, the within-season divorce rate was 4.6% and the between-season divorce rate was 15.5%. Retained pairs within a season initiated their renest 5 days faster than divorced birds that had no greater fledgling production, suggesting that within-season divorce was making the best of a bad job with severe time constraints. Poor performance in the year prior to divorce was not strongly associated with divorce, and analysis of multiple breeding stages revealed that divorcing individuals in the subsequent year had later laying dates, smaller clutches, and fewer fledglings than retained pairs but no better performance than widowed individuals. Analyzing the data separately by sex showed that neither males nor females benefitted from divorce. Thus, there is a reproductive cost linked to finding a new partner per se, but no reproductive advantage associated with divorce. New mates after divorce were usually not older (not higher quality) than previous mates, so intrasexual competition was probably not driving partnership splits. The most plausible explanation seems to be a “bet-hedging” hypothesis in which birds re-pair rapidly in spring if their previous mate does not quickly arrive during spring migration. Divorce in Northern Flickers does not appear to be adaptive and future studies on arrival and interactions of individuals in spring will elucidate proximate constraints on relocating the previous partner.

The allocation of prey within broods of altricial nestlings may be the result of parents targeting certain offspring preferentially and/or the result of intrabrood competition related to features of nestlings such as body size. We investigated two mechanisms that could influence allocation of food within broods of Mountain Bluebirds (Sialia currucoides) by filming the size and type of prey that parents fed nestlings. According to the gape size constraint hypothesis, the inability of small nestlings to physically swallow large prey items might place junior brood members at a higher risk of starvation than older siblings if parents do not bring enough small items. Consistent with this idea, the frequency of ‘testing’, placing an item in the gape of a nestling but then withdrawing it, was positively correlated with prey size. The junior nestling of a brood was fed less often when parents brought larger prey items, and nestlings were tested more in the early nestling stage (0–4 days old) when their gapes were smaller than at older stages. Parents of many cavity-nesting species may feed the brood from the nest entrance and the resulting scramble competition among brood members to access the entrance hole also potentially increases the mortality of the least competitive (smallest) nestlings. Although the frequency of feeding from the entrance hole was prevalent during the late nestling stage (≥ 12 days old), feeding from the entrance was not associated with mortality rate in bluebird broods and the mortality of junior nestlings was not linked to the testing of prey. The results confirm that a mechanism of gape size constraint does direct large prey away from junior nestlings, but apparently small-sized prey items were abundant enough during our study so that this did not result in increased mortality of nestlings.

Migrating animals occur along a continuum from species that spend the nonbreeding season at a fixed location to species that are nomadic during the nonbreeding season, essentially continuously moving. Such variation is likely driven by the economics of territoriality or heterogeneity in the environment. The Snowy Owl (Bubo scandiacus) is known for its complex seasonal movements, and thus an excellent model to test these ideas, as many individuals travel unpredictably along irregular routes during both the breeding and nonbreeding seasons. Two possible explanations for this large variation in the propensity to move are (1) dominance hierarchies in which dominant individuals (adult females in this case) monopolize some key, consistent resources, and move less than subdominants; and (2) habitat heterogeneity in which individuals foraging in rich and less heterogenic environments are less mobile. We analyzed fine-scale telemetry data (global positioning system [GPS]/global system for mobile communication [GSM]) from 50 Snowy Owls tagged in eastern and central North America from 2013 to 2019, comparing space use during the winter period according to sex and age, and to land cover attributes. We used variograms to classify individuals as nomadic (58%) or range-resident (42%), and found that nomadic owls had ten times larger wintering areas than range-resident owls. The frequency of nomadism was similar in socially-dominant adult females, immatures, and males. However, nomadism increased from west to east, and north to south, and was positively associated with the use of water and negatively associated with croplands. We conclude that many individual Snowy Owls in Eastern North America are nomadic during the nonbreeding season and that movement patterns during this time are driven primarily by extrinsic factors, specifically heterogeneity in habitat and prey availability, as opposed to intrinsic factors associated with spacing behavior, such as age and sex.

Woodpeckers and other excavators create most of the holes used by secondary cavity nesters (SCNs) in North American temperate mixedwood forests, but the degree to which excavators release SCNs from nest-site limitation is debated. Our goal was to quantify how excavators maintain the diversity and abundance of secondary cavity nesters in a temperate forest through the creation of tree cavities. We examined the short- and long-term (legacy) effects of excavators (principally woodpeckers, but also red-breasted nuthatches and black-capped chickadees) on forest biodiversity using longitudinal monitoring data (1,732 nest cavities, 25 sites, 16 years) in British Columbia, Canada. Sites with higher densities of excavator nests had more cavities available, higher species richness of SCNs and higher nest density of SCNs, indicating the importance of a standing stock of cavities. Years with higher nesting densities of excavators were followed by years with higher SCN diversity, indicating that the creation of nesting opportunities through fresh excavation releases SCNs from community-wide nest-site limitation. We also show that excavators leave a 'legacy' of biodiversity (species richness and abundance) at a site by accumulating cavities at rates faster than they become unusable by decay or destruction. By quantifying site-level effects of cavity excavation on the SCN community, our study highlights the key role of excavators as ecosystem engineers that maintain forest wildlife biodiversity.Si, dans la forêt mixte tempérée d’Amérique du Nord, les pics et autres excavateurs créent la plupart des cavités utilisées par les cavernicoles secondaires (SCN), la question de savoir s'ils augmentent la disponibilité des sites de nidification des SCN reste ouverte. Notre objectif était d’évaluer et quantifier l'impact qu'ont les excavateurs lorsqu'ils creusent des cavités dans les arbres, sur la diversité et l'abondance de cavernicoles secondaires dans une forêt mixte tempérée. Nous avons examiné les effets des excavateurs (principalement les pics, mais aussi la sittelle à poitrine rousse et la mésange à tête noire) à court et à long terme (survivance), sur la biodiversité de la forêt en Colombie-Britannique, au Canada, en utilisant des données longitudinales de surveillance (1,732 cavités ressource, 25 sites, 16 ans). Les sites à plus fortes densités de nids d'excavateurs avaient une plus grande abondance en cavités disponibles, une richesse en espèce SCN plus élevée, et une plus forte densité de nids de SCN, ce qui montre l'importance des arbres à cavités utilisables, vivants ou morts sur pied. Les années à plus fortes densités de nids d'excavateurs étaient suivies par des années à plus fortes diversités d'espèces SCN, ce qui signifie que la disponibilité de sites de nidification à partir de nouveaux creusements augmente le potentiel de nidification des espèces SCN pour l'ensemble de la communauté qui en dépend. Nous démontrons aussi que, sur un site donné, les excavateurs laissent une ‘survivance’ de biodiversité (richesse en espèce et abondance) en formant des cavités à un taux plus élevé que le taux avec lequel ces dernières deviennent inutilisables suite à leur décomposition ou destruction. En quantifiant les effets du creusement de cavités au niveau de chaque site par rapport à la communauté des SCN, notre étude met en évidence le rôle clé des excavateurs comme ingénieurs de l’écosystème favorisant la diversité de la faune forestière.

Nest sites of animals are often concealed to keep vulnerable offspring from being detected by predators. Parents may use landmarks near the nest to relocate it quickly. We allowed blue tits (Cyanistes caeruleus) to choose between two nest boxes fixed on the same tree with either none, the same, or different white painted markings. Surprisingly, the female brought material to both boxes and sometimes laid eggs in both. In a second experiment, we let pied flycatchers (Ficedula hypoleuca) and great tits (Parus major) become familiar with a marking on the initial nest box and then let them choose between two new nest boxes erected on different, nearby trees. Neither species preferred the box with the matching mark. In nature, the birds may locate the correct entrance of a cavity using other landmarks near the nest opening, like branches and the height of the cavity opening above the ground.

Among species that use similar resources, an individual may benefit by observing and copying the behavioural decision of a heterospecific. We tested the hypothesis of heterospecific social learning in passerine birds, namely that a migrant species, the Pied Flycatcher Ficedula hypoleuca, uses external markings on the nest cavities of a resident species, the Great Tit Parus major, as cues when choosing a nest site. Others have suggested that prospecting flycatchers assess the clutch size of tit “demonstrators” by entering their nest boxes and, assuming that a large clutch indicates a high-quality individual, will copy the nest appearance of tits with large, but not small clutches. During a 4-year period in Norway, we designed a similar study but did not find that flycatchers based their nest choice on the clutch size of tits. Neither were there any relationships between clutch size of the tit and its laying date, incubation behaviour, or the number of eggs visible through nest material during egg-laying so Pied Flycatchers did not use these indirect cues to assess quality of the tutor. Filming of tit nests showed that prospecting flycatchers did not enter tit nest boxes to assess the content. Indeed, incubating female tits only left their nest boxes for short bouts of unpredictable duration so there was little opportunity for flycatchers to inspect the nest contents unnoticed. Our study calls into question the mechanism of using the content of tit nests as public information for choosing traits of nest sites based on external characteristics. We suggest that similar studies of nest site choice in relation to possible social information transfer be replicated more widely.

Many animals require tree cavities for breeding and these sites may be reused by a diversity of secondary cavity nesters over a timespan of decades. It is unknown whether the reuse of holes changes their desirability as nest sites. We hypothesized that some species, “cavity destroyers,” degrade the quality of holes by filling them with coarse nest material or waste whereas excavating species, “cavity cleaners,” might prolong the use of a hole by removing debris or enlarging the hole. Using data gathered during 22 years from a field study in central British Colombia, we analyzed long-term patterns of cavity occupancy in relation to their sequential use by bird and mammal species, grouped by traits. Patterns of cavity occupancy were variable with 49% of 875 large-sized holes (excavated by northern flickers Colaptes auratus and pileated woodpeckers Dryocopus pileatus) and 19% of 652 smaller-sized holes incorporating runs of sequential use that lasted to 18 years. About 11% of large and 25% of small cavities also had gaps of 3–13 years between occupancies. Mammals, raptors and European starlings, consistent with the hypothesis, were cavity destroyers, occupying cavities as terminal users and before gaps more often than expected by chance. The pattern of occupancy by northern flickers was random in relation to gaps or prior use by other species. Although flickers did not target old holes to clean, neither did they avoid them. Small cavities that were renovated by flickers into larger cavities were reused at twice the rate after renovation. Runs of cavity occupancy that involved only cavity-destroying species were shorter than runs that involved periodic use by flickers, suggesting the woodpecker, through its cleaning and renovation, prolonged the use of such holes. Our study contributes insights on additional ecological factors, namely previous users, that can influence the use and availability of cavities over time. Fil: Wiebe, Karen L.. University of Saskatchewan; Canadá Fil: Cockle, Kristina Louise. University of British Columbia; Canadá. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Instituto de Biología Subtropical. Instituto de Biología Subtropical - Nodo Puerto Iguazú | Universidad Nacional de Misiones. Instituto de Biología Subtropical. Instituto de Biología Subtropical - Nodo Puerto Iguazú; Argentina Fil: Trzcinski, M. Kurtis. University of British Columbia; Canadá Fil: Edworthy, Amanda B.. University of British Columbia; Canadá Fil: Martin, Kathy. University of British Columbia; Canadá

Natal dispersal and local recruitment are affected by factors both intrinsic and extrinsic to juveniles and may affect fitness. Understanding the relationship between dispersal and population density in birds has been hindered by a lack of long-term studies and a focus on resident species has neglected the role of weather operating at large spatial scales. I studied local recruitment and the reproductive consequences of natal dispersal distance within a population of Northern Flickers (Colaptes auratus), a migratory woodpecker. During a field study spanning 16 yr in British Columbia, 8,272 fledglings were banded and 138 males and 105 females recruited locally. The average annual local recruitment rate for males (3.36%) was greater than that for females (2.55%) and the propensity to recruit locally was positively correlated with an early hatch date and high body condition. Annual local recruitment was not associated with population density in the year of hatch but was positively correlated with population density in the year of settlement. Local recruitment was also positively correlated with warmer springs during migration, consistent with the phenology hypothesis that the location of settlement is affected by weather along the route. Among local recruits, natal dispersal distance was independent of the presence of parents and so dispersal was not a behavior to prevent inbreeding. However, settling closer to the natal site led to reproductive benefits in terms of earlier laying dates and better nest success. Therefore, juveniles may gain useful information about the location of nesting substrates, predation risk, and patchy food resources by exploring the landscape around their natal site during the post-fledging period and then settling in the familiar area after returning from migration.

Interspecific competition is expected when two species share resource needs. For example, secondary cavity‐nesting birds may compete vigorously for suitable nest sites both within and among species. However, little is known of whether monogamous species defend more than one nest site on their territory after breeding has begun, and in particular whether they are aggressive against other species. Defending extra nest sites may be adaptive because they may be used for renesting after a failure or to produce a second brood. We studied interactions between a monogamous, resident bird, the great tit Parus major and a migrant, the pied flycatcher Ficedula hypoleuca, providing nest boxes in woodlands in Norway. Agonistic behaviours may be subtle and easily overlooked so we experimentally placed caged male pied flycatchers near nests of great tits and at a dyad of empty nest boxes erected in the territory 25 m away. We filmed interactions between species at 21 sites in the egg‐laying period of the tit, and at 26 nest sites during incubation. Male great tits showed aggression towards the caged flycatcher both at their own nest box and at the nest boxes erected at a distance. We manipulated the external appearance of the nest boxes with painted designs around the entrances but the intensity of aggression at the empty nest boxes did not depend on whether those boxes matched the nest box with the tit nest and was not correlated with tit clutch size. Neither was the intensity of display activity at each nest box by the flycatchers that settled associated with these variables. The results are discussed in relation to hypotheses for nest site choice involving interspecific social learning and aggression.

Age-related improvement in reproductive performance is widespread in vertebrates and constraints at young ages are a common cause. The sex that invests energetically more in reproduction, typically the female, is predicted to show stronger age-related performance but the effect of the male's age on reproduction has often been ignored. I studied age-related reproduction of both sexes in northern flickers, in which males invest more parental care than females, predicting that the effect of age would be stronger in males than in females. Longitudinal data on individuals collected during an 18-year field study confirmed this prediction. Laying dates for females improved only between the first 2 years of her life and no other reproductive parameter changed over her lifetime when the male's age was statistically controlled. In contrast, males improved up to age five for laying date, clutch size, hatching success and fledging success. Partner familiarity (fidelity) was further associated with earlier laying, larger clutches, improved fledging success and more fledglings. There was significant assortative pairing by age but there is apparently little benefit for males to choose older females, but a benefit to females with older males. Females appear to strategically lay larger clutches when paired to old males which invest more in paternal care than younger males. This is the first example of clutch size being influenced by only male age and not female age in any bird and suggests that sex roles in parental care are important determinants of aging patterns in vertebrates with diverse life histories.

Meeting energy requirements during the nonbreeding season is important for many animals and some defend winter territories to secure a food supply. In birds of prey, females, the larger and competitively dominant sex, may monopolize areas with higher prey abundance than males. We thus predicted that female Snowy Owls (Bubo scandiacus (Linnaeus, 1758)), which might acquire the high-quality habitats and individuals in better body condition, would be able to persist on smaller home ranges, travel shorter distances, and spend proportionally more time on a home range than males, during the winter. On the prairies in central Saskatchewan, we deployed satellite transmitters on 11 male and 12 female Snowy Owls over two winters. There were no significant differences between the sexes in home-range size or the amount of travelling during the winter months. Mean first home range (95% minimum convex polygon) size was 54.4 km2 for males, 31.9 km2 (estimated marginal means) for females, and 53.8 km2 for the sexes combined. However, within sexes, home-range size was negatively correlated with body condition as predicted. A lack of defense of home-range perimeters against conspecifics could increase variation in home-range size and movement patterns and reduce differences between the sexes.

Natal dispersal is common in animals but the fitness cost of moving from the natal area is not well understood. One reason for a fitness cost is that foraging skills and prey preference learned early in life may be less efficient if the individual settles in a new, unfamiliar habitat. In a 4-year study, we found that immigrant parent blue tits, Cyanistes caeruleus, and great tits, Parus major, were inferior food providers compared to local recruits. In blue tits, immigrants provided smaller prey items than local recruits, whereas in great tits, the immigrants provided fewer green larvae, but relatively more brown larvae, to the offspring than local recruits. We also found that immigrant females laid later or smaller clutches than females locally recruited. The results are consistent with the hypothesis that natal dispersal carries costs related to the learning of foraging skills. However, alternative explanations are that the differences were caused by genetic and/or quality differences between the two groups of birds. We discuss various ecological and behavioural traits that may influence, and be influenced by, the mismatch of foraging between natal and breeding habitats. In altricial birds, yearlings will not have previous foraging experience during breeding and, in addition, immigrants will not have spent a long postfledging period in the new local habitat with their parents. If there are foraging-habitat mismatches as a result of dispersal, researchers should include natal origin in models of optimal foraging, time budgets, reproductive success and survival because performance may be directly related to the early learning environment rather than genetic differences.

Snowy Owls Bubo scandiacus often face the sun when sitting on the ground or when perched. Such sun-orienting has been suggested to represent a visual display to conspecifics but other explanations have not been thoroughly examined. We observed the orientation of wintering Snowy Owls to both the sun and the wind, and their perching behaviour during two winters in central Saskatchewan, Canada. We proposed three new explanations for sun orientation in addition to the display hypothesis: thermoregulation, hunting and defense against predators. On sunny days, 44% of 710 owls faced the sun; this was non-random because few did so on overcast days. Sun as opposed to wind orienting was strongly associated with weather conditions. Logistic regressions indicated that at temperatures below -13 °C and at wind speeds greater than about 18 km/ hr, owls tended to orient to the wind rather than the sun. The likelihood of wind-orienting increased if the owl perched above the ground whereas the likelihood of sun-orientating increased slightly when the owl was sitting on the ground. There was no difference between the sexes in orienting behaviour. Snowy Owls seemed to prioritize wind-orienting for thermoregulation but the results are also consistent with the idea that sun-orientation can reduce heat loss. Facing into the sun did not support the hunting explanation because the birds would have been blinded and not able to see prey but was consistent with the protection explanation if it helps to increase vigilance against enemies. Although we cannot completely rule out the display explanation, the spatial context of sunning owls and a lack of a sex effect makes it unlikely that this is the main function. Instead, owls seem to trade off wind versus sun orienting according to the prevailing weather conditions and do so mainly to thermoregulate and perhaps to maintain vigilance. This article is protected by copyright. All rights reserved.

Migratory species display a range of migration patterns between irruptive (facultative) to regular (obligate), as a response to different predictability of resources. In the Arctic, snow directly influences resource availability. The causes and consequences of different migration patterns of migratory species as a response to the snow conditions remains however unexplored. Birds migrating to the Arctic are expected to follow the spring snowmelt to optimise their arrival time and select for snow-free areas to maximise prey encounter en-route. Based on large-scale movement data, we compared the migration patterns of three top predator species of the tundra in relation to the spatio-temporal dynamics of snow cover. The snowy owl, an irruptive migrant, the rough-legged buzzard, with an intermediary migration pattern, and the peregrine falcon as a regular migrant, all followed, as expected, the spring snowmelt during their migrations. However, the owl stayed ahead, the buzzard stayed on, and the falcon stayed behind the spatio-temporal peak in snowmelt. Although none of the species avoided snow-covered areas, they presumably used snow presence as a cue to time their arrival at their breeding grounds. We show the importance of environmental cues for species with different migration patterns.

Woodpeckers are considered ecosystem engineers because they excavate tree cavities which are used subsequently by many species of secondary cavity nesters for breeding. Woodpeckers have the choice of excavating a new hole or reusing an existing one, and this propensity to excavate (e) may affect community dynamics but has rarely been investigated. Using 18 years of data on a population of northern flickers Colaptes auratus, I tested six hypotheses to explain the propensity to excavate (e) in a landscape which experienced two types of disturbance: pine beetles and wildfires. Woodpecker age, breeding experience and mate retention had little influence on e which varied between 13-39% annually and averaged 23% for 1843 first nests over the 18 years. Body size and body condition of males and females were not associated with e but rates of excavation declined seasonally, suggesting time rather than energy costs limited excavation effort. Reduced cavity availability mediated through high conspecific density coupled with wildfires triggered relatively high excavation rates, up to 39% but e decreased to baseline levels three years after the landscape disturbances. Nearly 2/3 of males did not excavate in their lifetime but apparently, e is great enough to balance the average rate of cavity tree loss in this forest which is 11% annually. Excavation propensity in flickers is flexible, but the birds reduce their work levels if there is a surplus of holes available. This article is protected by copyright. All rights reserved.

Tree cavities are a critical multi-annual resource that can limit populations and structure communities of cavity-nesting vertebrates. We examined the regional and local factors influencing lifetime productivity (number and richness of occupants) of individual tree cavities across two divergent forest ecosystems: temperate mixed forest in Canada and subtropical Atlantic Forest, Argentina. We predicted that (1) species would accumulate more rapidly within cavities in the species-rich system (Argentina: 76 species) than the poorer system (Canada: 31 species), (2) cavity characteristics associated with nest-site selection in short-term studies would predict lifetime cavity productivity, and (3) species would accumulate more rapidly across highly used cavities than across cavities used only once, and in Argentina than in Canada. We monitored and measured nesting cavities used by birds and mammals over 22 breeding seasons (1995–2016) in Canada and 12 breeding seasons (2006–2017) in Argentina. Cavities were used an average of 3.1 times by 1.7 species in Canada and 2.2 times by 1.4 species in Argentina. Species richness within cavities increased with number of nesting events at similar rates in Canada and Argentina, in both cases much slower than expected if within-cavity species assemblages were random, suggesting that lifetime richness of individual cavities is more strongly influenced by local ecological factors (nest site fidelity, nest niche) than by the regional species pool. The major determinant of lifetime cavity productivity was the cavity’s life span. We found only weak or inconsistent relationships with cavity characteristics selected by individuals in short-term nest-site selection studies. Turnover among (vs. within) cavities was the primary driver of diversity at the landscape scale. In Canada, as predicted, species accumulation was fastest when sampling across high-use cavities. In Argentina, the rates of species accumulation were similar across high- and low-use cavities, and fastest when both high- and low-use cavities were pooled. These findings imply that biodiversity of cavity nesters is maintained by a mix of long-lived (highly productive, legacy trees) and many high-turnover (singleuse, fast decaying) tree cavities. Conservation of both long-lasting and single-use cavities should be incorporated into decisions about stand-level forest management, regional land use policies, and reserve networks. Fil: Cockle, Kristina Louise. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Instituto de Biología Subtropical. Instituto de Biología Subtropical - Nodo Puerto Iguazú | Universidad Nacional de Misiones. Instituto de Biología Subtropical. Instituto de Biología Subtropical - Nodo Puerto Iguazú; Argentina. University of British Columbia; Canadá Fil: Trzcinski, Mark Kurtis. University of British Columbia; Canadá Fil: Wiebe, Karen L.. University of Saskatchewan; Canadá Fil: Edworthy, Amanda B.. University of British Columbia; Canadá Fil: Martin, Kathy. Environment and Climate Change Canadá. Science & Technology Branch; Canadá. University of British Columbia; Canadá

Hybridization can bring in single individuals alleles that were never designed to work together, which can result in unexpected or transgressive phenotypes. The Yellow-shafted (auratus group) and Red-shafted (cafer group) subspecies groups of the Northern Flicker (Colaptes auratus) differ conspicuously in the coloration of their flight feathers, but hybridize freely where their ranges overlap in western North America. The difference in color is largely the result of the Red-shafted form harboring ketolated products at C4(4′) of the carotenoids found in the Yellow-shafted form. Characterizing the carotenoid pigments in a series of birds of intermediate color (presumed hybrids) revealed that most accumulated a product of β-cryptoxanthin with a keto group on its hydroxylated ring (3-hydroxy-echinenone), while a few accumulated the product with a keto group on the unhydroxylated ring (3′-hydroxy-echinenone). Surprisingly, the latter group also had feather barbs that were noticeably yellower than the associated rachis, corresponding to a lower level of ketolation at C4(4′). We assessed possible biochemical explanations for the differences by probing the relative carotenoid concentration data in individuals of varying color. The difference between the hybrids could not be explained by the general level of ketolation of carotenoids or a particular selectivity of the 4-ketolase involved. We present a testable genetic explanation that invokes incompatibilities between divergent alleles of the two parental forms at interacting loci. Because the idiosyncrasies affect oxidation, they may be the product of mitonuclear incompatibilities.

Birds that winter in cold northern climates experience harsh conditions, including reduced food availability and increased energy demands. In raptors, the ability to forage and maintain body condition may be related to age (hunting experience) or the ability to defend good-quality territories (dominance). We examined the effects of age and sex on body condition and various sources of mortality in wintering Snowy Owls (Bubo scandiacus) on the Canadian prairies. Because of reversed sexual size dimorphism, we predicted that females, the dominant sex, would be in better condition than males and that adults would be in better condition than juveniles. Consistent with these predictions, data from 537 live Snowy Owls trapped over 18 winter field seasons showed that adults were heavier than juveniles for a given body size and carried more fat reserves. We found that 56% of males lacked furcular and wing-pit fat, whereas only 31% of females lacked such fat; and females, but not males, tended to put on fat...

Interspecific competition over nest sites is common among cavity-nesting birds, but little is known about what determines the outcome of such contests, particularly whether or not prior ownership plays a role. Using a box removal and replacement experiment, I tested whether Mountain Bluebird (Sialia currucoides) or Tree Swallow (Tachycineta bicolor) had the higher resource holding potential (RHP). A different box-blocking experiment with paired boxes on territories measured the extent to which prior ownership influenced contest outcomes for both species. Behavioral observations in the pre-laying period showed more physical aggression by Mountain Bluebird against Tree Swallow. Nevertheless, Tree Swallow won 70% of boxes when neither species had prior ownership, suggesting Tree Swallow had a higher RHP. When their boxes were blocked, 24% of Tree Swallow usurped boxes from Mountain Bluebird, which did not differ statistically from the 33% of boxes usurped by bluebirds from swallows; thus, prior owne...

Carotenoid-based colouration in feathers is widely accepted to be a reliable signal of the health of an individual, but the condition-dependence of melanin-based plumage ornaments has been highly debated. Using broods that were manipulated in size, we tested whether nutritional stress during rearing affected the carotenoid pigmentation in secondary feathers and the size, shape, and symmetry of melanin spots on breast plumage of northern flicker Colaptes auratus nestlings. Two measures of carotenoid colour (chroma and brightness) of secondary flight feathers did not vary according to brood size treatment, but in a larger dataset from the population, carotenoid chroma was positively associated with nestling mass. Nestlings from experimentally enlarged broods had smaller melanin spots than those from reduced broods, which is some of the first experimental evidence that melanin ornament size in growing nestlings is condition-dependent. However, the shape and symmetry of the melanin breast spots was not associated with nestling mass. Sexual dimorphism was apparent in both types of pigmentation and future studies should investigate whether there are any trade-offs for nestlings between investing in carotenoid colouration and melanisation and whether trade-offs differ between the sexes.

Optimal foraging theory suggests that avian parents should prefer the most energetically efficient (largest) prey items when delivering food to offspring at a central place. However, during periods of high demand, selectivity of prey may decline, leading to the delivery of smaller and/or less nutritious items. We compared foraging trade-offs between great tits (Parus major) which had a wider feeding niche than blue tits (Cyanistes caeruleus). We also compared the foraging efficiency of cross-fostered young, which had learned the spatial foraging niche and prey size of the foreign species, to that of control conspecifics. Mean delivery rates did not differ between control and cross-fostered parents of either species but as delivery rates increased, prey size declined for both species and both treatment groups. However, across the range of increasing delivery rates, parents were able to increase the total biomass of prey delivered. Cross-fostering did not alter the proportion of different prey taxa in the diet, but cross-fostered birds shifted the size of the prey taken to that of their foster species. Consistent with their broader feeding niche, great tits, but not blue tits, incorporated more unpalatable items (flies) as delivery rates increased. Although great tits foraged less efficiently in the blue tit niche, paradoxically, blue tits seem to deliver more prey biomass when foraging in the great tit niche.

Nest sanitation is an understudied form of parental care that may improve offspring fitness by reducing microbes and ectoparasites in the nest. Many species clean nests, but it is unknown whether parents respond flexibly to various costs and benefits when deciding how much effort to invest. We experimentally manipulated brood size in the short term and in the long term in a cavity nester, the northern flicker, Colaptes auratus, to test whether parents alter their sanitation effort in response to brood demands. Males generally removed more faecal sacs than females at all treatments and in most cases, faecal removal rates were proportional to feeding rates in control and experimental broods. The rate of sac removal was negatively correlated with feather corticosterone in females but not in males. Males maintained higher faecal removal rates than females to enlarged broods and were more flexible than females in altering their sanitation effort relative to their feeding rate. Single males, but not females, removed fewer faecal sacs than paired males, suggesting that males reduce nest sanitation effort in times of high nestling demands if it helps maximize fledging success. Across taxa, the sex that invests the most in other forms of parental care also seems to perform more nest sanitation, and future studies should test how parents value it differently depending on brood demands and intrinsic factors.

Elucidating the processes that create species differences is a central goal of evolutionary biology. The Northern Flicker (Colaptes auratus) exists as two well-differentiated subspecies groups in North America, the Yellow-shafted (auratus group) and Red-shafted Flickers (cafer group), which differ strikingly in the color of the underside and rachises of flight feathers, and of malar and nuchal patches. We investigated the physiological basis of these conspicuous phenotypic differences by identifying and quantifying the pigments involved. The yellow feathers of auratus contained carotenoids commonly found in nature (lutein, β-cryptoxanthin, zeaxanthin and β-carotene). The orange to red shafts/vanes of cafer and hybrids contained these carotenoids as well as mono- and diketo-carotenoids (notably adonirubin, α-doradexanthin, canthaxanthin, astaxanthin), representing oxygenated products at carbon C4(4') of the carotenoids present in auratus. Oxygenation of feather carotenoids at C4(4') correlated closely with shaft/vane redness. Carotenoid hydroxylation at C3(3') and the proportion of carotenoids with ε end-rings also varied with color and belie differences in the activity of several carotenoid-modifying enzymes between the two subspecies groups. Curiously, occasional yellow feathers in red-shafted individuals had the carotenoids of auratus, hence the differences are not constitutive in cafer, underscoring regulatory differences. The red malar stripe of cafer, the black malar stripe and red nuchal patch of auratus all contained similar types and amounts of carotenoids, mostly 3-hydroxy-4-keto-carotenoids. The biochemical differences between two strongly differentiated forms we uncovered shed light on how plumage coloration can change over evolutionary time and point to further avenues of research.

Melanin is a common pigment in the plumage of birds, but the extent to which its deposition in feathers is condition dependent and can be used as a reliable signal of quality and reproductive performance is still much debated. In addition, the existence and function of melanin ornaments in female birds or in birds of different age classes has rarely been addressed. We studied the size and color of 4 melanin ornaments in the Northern Flicker (Colaptes auratus), a woodpecker, and found that the plumage patches became larger or darker with age class in both sexes. Consistent with the partly reversed sex roles of Northern Flickers and sexual selection in both sexes, several melanin ornaments were the same size in males and females and were correlated with body size or body condition. There was assortative pairing according to melanin ornaments when controlling for age, and 1 or more ornaments predicted laying date or clutch size of the pair independent of age. The results suggest that melanin ornamen...

This research has been accepted and published in the Journal of Avian Biology. © 2017 Wiley

Foraging with others can reduce predation risk but may increase competition for resources. Species, sex, or age classes may use different niches to reduce competition for food, which may be enforced by social dominance or may lead to ecological specialization. An alternate hypothesis is that individuals using resource patches that renew over time intentionally avoid recently depleted patches by avoiding each other. We tested these hypotheses by simultaneously radio-tracking northern flicker (Colaptes auratus) mates. Members of a pair rarely foraged in the same patch, spending over 90 % of the time >50 m apart. Counter to the ecological specialization hypothesis, the types of ground cover used for foraging and the distance of foraging patches from cover and edge did not differ between males and females. Mates were more spatially segregated on the home range than random and males foraged closer to the nest than females, but we never saw agonistic behaviors counter to the dominance hypothesis. Both sexes avoided returning to the same foraging location repeatedly, perhaps because of prey depletion. We suggest that members of a pair avoided foraging close to each other to reduce encounters with depleted prey patches. We emphasize the importance of simultaneous tracking of mates to identify foraging strategies and potential niche segregation at the home range scale.

Brood enlargement experiments have been conducted in several species of birds to investigate how parents of both sexes adjust their investment in the current breeding attempt. We studied parental feeding effort in the Northern Flicker (Colaptes auratus), a species with partially reversed sex roles where males invest more in parental care than females and in which there is facultative polyandry and no extra-pair young. By experimentally manipulating brood sizes to be either larger or smaller by about 40%, we tested the flexibility of provisioning responses by each sex and whether the upper limit to provisioning corresponded to the maximum clutch size in the population. Both male and female parents increased feeding rates to enlarged broods, but per-nestling provisioning declined so that they fledged lighter nestlings with shorter wings. Mortality in reduced broods was lower than in control broods, but there was no difference in the incidence of mortality between control and enlarged broods. Parent...

The importance of cavities as roost sites in migratory species is often unknown because it is challenging to monitor cavity use during the non-breeding period. We documented cavity use throughout the annual cycle of a woodpecker using light-level geolocators. Northern Flickers Colaptes auratus spent 63–90% of nights roosting in a cavity throughout the year, including during migration. The high frequency of year-round cavity use by Flickers suggests that cavities provide benefits beyond nest-sites. Our work highlights the potential use of geolocators to examine cavity use.

Census-based studies document changes in population size of animal species in response to wildfires, but mechanisms involving behavior of individuals and the effects on reproductive success are usually unknown. I developed a conceptual model explaining the persistence of cavity-nesting birds on the landscape after fires depending on carrying capacity (food supply) of the habitat and the philopatry and territoriality of the species. The breeding density, nest site characteristics, and reproductive success of Northern Flickers Colaptes auratus was studied before and after low- to moderate-severity fires on replicated plots. The density and spatial distribution of nests did not change in a consistent way after fires, but the rates of cavity excavation increased and characteristics of nest sites changed as such decay class of the tree. Laying dates were delayed and clutches were smaller in freshly excavated vs. reused cavities on the burned sites. Breeding philopatry caused a shift to older age classes in the population. Depredation of nests by small mammals increased during the first three years after fires, reducing the number of young produced per nest attempt on burned plots. The study shows that, even when fire does not reduce the density of breeding pairs, there may be detrimental effects detected only by monitoring the behavior and reproduction of individuals.

Manipulations of brood size measure the willingness or ability of parents to invest in offspring and different reproductive roles may lead to differences in feeding effort between the sexes. Parental investment in birds is usually assessed by quantifying feeding rates, but this provides an incomplete picture of parental effort because it fails to account for how parents collect food on the landscape. We studied northern flickers (Colaptes auratus), a woodpecker in which males provide the majority of parental care and used a repeated measures design and short-term (24 h) brood enlargements (N = 35) and reductions (N = 27) to assess effects of treatment on feeding rates to nestlings and parental foraging behaviour. Parents of enlarged broods did not significantly increase feeding rate, resulting in a decline in nestling mass. Parents of reduced broods decreased their feeding rates by 84%, but increased per capita feeding rates, resulting in nestling mass gain. The variation in feeding rates to enlarged broods was not influenced by feather corticosterone, body condition, feather re-growth rate or mass change between the incubation and nestling periods. Foraging pattern on the landscape remained the same during the enlarged treatment for both sexes. We conclude that flickers respond to proximate cues in brood demands, but do not increase feeding rates to enlarged broods, at least in the short term. A literature review suggested that this lack of response is atypical for short-lived species. We hypothesize that parents in species with large home ranges and long nestling periods face energetic limitations that constrain their ability to respond to enlarged broods. We encourage future studies to assess foraging behaviour on the landscape to document important trade-offs for parents such as predation risk and energy expenditure while feeding offspring.

Sexual conflict is magnified during the post-fledging period of birds when the sexes face different trade-offs between continuing parental care or investing in self maintenance or other mating opportunities. Species with reversed sex roles provide a unique opportunity to study the relationship between mating systems and investment in parental care. Here, we provide the first detailed study of the length of care by males versus females (n = 24 pairs) during the post-fledging period, assessing factors that may promote care within and between the sexes. In the northern flicker Colaptes auratus, a species with partly reversed sex roles, males cared longer than females (average 16 versus 12 days, respectively). Overall, 36 % of females but no males deserted the brood prior to fledgling independence. Parents that provisioned nestlings at a high rate also spent more days feeding fledglings. Among males, age and nestling feeding rates were positively associated with the length of care. Among females, a low level of feather corticosterone (CORTf) was associated with a longer length of care. About 45 % of fledglings died within the first week, but fledglings with intermediate body mass had the highest survival suggesting stabilizing selection on mass. Fledgling survival was also higher in individuals with larger broods and lower levels of CORTf. We demonstrate that because females can be polyandrous they often desert the brood before males, and that the sexes respond to different cues relating to their energy balance when deciding the length of care given to their offspring.