Density-dependent winter survival of immatures in an irruptive raptor with pulsed breeding.

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.

Woodpeckers and other excavators maintain the diversity of cavity-nesting vertebrates.

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.

No evidence that nest site choice in Pied Flycatchers is mediated by assessing the clutch size of a heterospecific, the Great Tit.

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.

Egg covering in cavity nesting birds may prevent nest usurpation by other species.

ABSTRACT: Some birds cover their eggs with nest material when they leave to forage. It has been suggested that such egg-covering aids thermoregulation or prevents predation but here we present a new hypothesis, that secondary cavity-nesting species cover their eggs to prevent nest usurpation by other birds. When the bottom of the cavity is dark, as when eggs are covered by nest material, it may be difficult for a prospecting competitor to see whether a defending nest owner or a predator is hiding inside the cavity. Competitors may therefore hesitate to enter dark cavities. We filmed 21 great tit (Parus major) nests during the egg-laying period and found that the female spent bouts of highly variable length outside the nest box (range 0.3-250 min, n = 51), so prospecting small passerines would have difficulty predicting whether an aggressive tit owner was in the box or would soon return. We presented prospecting male pied flycatchers (Ficedula hypoleuca) with a dyad of boxes (n = 93), each containing a great tit nest but only one with visible eggs. Flycatchers hesitated more to enter a nest box with no visible tit eggs than a box with exposed eggs. This was most evident for nest boxes with dark versus light interior paint, supporting the idea that better interior illumination makes prospecting birds more confident about entering an unfamiliar cavity. The usurpation and predation hypotheses are not mutually exclusive because both competitors and small predators may hesitate to enter dark, enclosed spaces if visibility is low. SIGNIFICANCE STATEMENT: Some birds deposit a layer of material on top of the eggs when they leave the nest. Several hypotheses have been proposed for such egg covering, for example that it may insulate the eggs and reduce the risk of nest predation. We propose a new hypothesis, namely that secondary hole-nesting birds cover their eggs when they leave the nest to prevent usurpation of the cavity by other birds. Great tits that we filmed at the nest during the egg-laying period could be absent for long periods. To test the hypothesis, we presented male pied flycatchers, potential nest competitors, with a dyad of nest boxes, each containing a great tit nest but only one with visible tit eggs. In support of the prediction, prospecting flycatchers hesitated to enter dark cavities with dark floors relative to boxes with exposed, reflective eggs.

Nest decoration: birds exploit a fear of feathers to guard their nest from usurpation.

Many species of birds incorporate feathers into their nest as structural support and to insulate the eggs or offspring. Here, we investigated the novel idea that birds reduce the risk of nest usurpation by decorating it with feathers to trigger a fear response in their rivals. We let prospecting birds choose between a dyad of nest-boxes in the wild, both containing some nest materials, but where one had a few white feathers and the other had none. All three species of cavity-nesting birds studied, the pied flycatcher Ficedula hypoleuca, the blue tit Cyanistes caeruleus, and the tree swallow Tachycineta bicolor, hesitated to enter boxes with white feathers. A similar avoidance of white feathers was found when the alternative nest-box of a dyad held black feathers. However, the birds readily collected white feathers that we placed in front of their nest-box, showing the fear of such feathers was context-dependent. We suggest that naive prospecting birds may perceive feathers in nests as the result of a predation event, and that owners decorate nests with bright feathers that can be seen from the opening to deter others from entering.

Disruptions of feather carotenoid pigmentation in a subset of hybrid northern flickers (Colaptes auratus) may be linked to genetic incompatibilities.

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 ß-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.

Author Correction: Arctic avian predators synchronise their spring migration with the northern progression of snowmelt.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Arctic avian predators synchronise their spring migration with the northern progression of snowmelt.

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.

Lifetime productivity of tree cavities used by cavity-nesting animals in temperate and subtropical forests.

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 (single-use, 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.

Age-related improvements in fecundity are driven by the male in a bird with partially reversed sex roles in parental care.

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.

Plumage pigment differences underlying the yellow-red differentiation in the Northern Flicker (Colaptes auratus).

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, ß-cryptoxanthin, zeaxanthin and ß-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.

Determinants of parental care and offspring survival during the post-fledging period: males care more in a species with partially reversed sex roles.

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.

Males migrate farther than females in a differential migrant: an examination of the fasting endurance hypothesis.

Patterns of migration including connectivity between breeding and non-breeding populations and intraspecific variation in the distance travelled are important to study because they can affect individual fitness and population dynamics. Using data from 182 band recoveries across North America and 17 light-level geolocators, we examined the migration patterns of the northern flicker (Colaptes auratus), a migratory woodpecker. This species is unusual among birds because males invest more in parental care than females. Breeding latitude was positively correlated to migration distance because populations in the north appeared to travel farther distances than southern populations to find wintering locations with little snow cover. Connectivity was strong for populations west and east of the Continental Divide. Contrary to the three main hypotheses for intraspecific variation in migration distance, females wintered, on average, farther north than males, although there was overlap throughout their non-breeding range. This pattern contradicts those of other species found to date and is most consistent with the fasting endurance hypothesis if investment in parental care depletes the energy reserves of male flickers more than females. We thus propose a new factor, parental effort, which may influence optimal wintering areas and migration strategies within birds, and encourage future experimental studies to test the relationship between parental care roles and migration strategies of the sexes.

Survival analysis of a critical resource for cavity-nesting communities: patterns of tree cavity longevity.

Tree cavities are a vital multi-annual resource used by cavity-nesting birds and mammals for nesting and shelter. The abundance of this resource will be influenced by the rates at which cavities are created and destroyed. We applied the demographic concepts of survival and longevity to populations of tree holes to investigate rates of loss for cavities in three tree species, as well as how characteristics of nest trees, habitat type, and species of excavator affected the persistence of tree cavities in trembling aspen, Populus tremuloides (95% of cavities were in aspen trees), in interior British Columbia, Canada. By modeling survival of 1635 nesting cavities in aspen over a time span of 16 years, we found that the decay stage of the nest tree was the most important factor determining cavity longevity. Cavities in trees with advanced decay had a relatively short median longevity of 7 years (95% CI 6-9 years), whereas those in living trees had a median longevity of more than 15 years. We found that cavity longevity was greater in continuous forest than in aspen grove habitat. Interestingly, cavities formed by weak excavators survived as long as those created by Northern Flickers (Colaptes auratus), despite occurring in more decayed tree stems. Thus, weak excavators may be selecting for characteristics that make a tree persistent, such as a broken top. Our results indicate that retention of cavities in large, live aspen trees is necessary to conserve persistent cavities, and that cavity longevity will have a large effect on the structure and function of cavity-using vertebrate communities.

Climate change, breeding date and nestling diet: how temperature differentially affects seasonal changes in pied flycatcher diet depending on habitat variation.

1. Climate warming has led to shifts in the seasonal timing of species. These shifts can differ across trophic levels, and as a result, predator phenology can get out of synchrony with prey phenology. This can have major consequences for predators such as population declines owing to low reproductive success. However, such trophic interactions are likely to differ between habitats, resulting in differential susceptibility of populations to increases in spring temperatures. A mismatch between breeding phenology and food abundance might be mitigated by dietary changes, but few studies have investigated this phenomenon. Here, we present data on nestling diets of nine different populations of pied flycatchers Ficedula hypoleuca, across their breeding range. This species has been shown to adjust its breeding phenology to local climate change, but sometimes insufficiently relative to the phenology of their presumed major prey: Lepidoptera larvae. In spring, such larvae have a pronounced peak in oak habitats, but to a much lesser extent in coniferous and other deciduous habitats. 2. We found strong seasonal declines in the proportions of caterpillars in the diet only for oak habitats, and not for the other forest types. The seasonal decline in oak habitats was most strongly observed in warmer years, indicating that potential mismatches were stronger in warmer years. However, in coniferous and other habitats, no such effect of spring temperature was found. 3. Chicks reached somewhat higher weights in broods provided with higher proportions of caterpillars, supporting the notion that caterpillars are an important food source and that the temporal match with the caterpillar peak may represent an important component of reproductive success. 4. We suggest that pied flycatchers breeding in oak habitats have greater need to adjust timing of breeding to rising spring temperatures, because of the strong seasonality in their food. Such between-habitat differences can have important consequences for population dynamics and should be taken into account in studies on phenotypic plasticity and adaptation to climate change.

Social learning in birds and its role in shaping a foraging niche.

We briefly review the literature on social learning in birds, concluding that strong evidence exists mainly for predator recognition, song, mate choice and foraging. The mechanism of local enhancement may be more important than imitation for birds learning to forage, but the former mechanism may be sufficient for faithful transmission depending on the ecological circumstances. To date, most insights have been gained from birds in captivity. We present a study of social learning of foraging in two passerine birds in the wild, where we cross-fostered eggs between nests of blue tits, Cyanistes caeruleus and great tits, Parus major. Early learning causes a shift in the foraging sites used by the tits in the direction of the foster species. The shift in foraging niches was consistent across seasons, as showed by an analysis of prey items, and the effect lasted for life. The fact that young birds learn from their foster parents, and use this experience later when subsequently feeding their own offspring, suggests that foraging behaviour can be culturally transmitted over generations in the wild. It may therefore have both ecological and evolutionary consequences, some of which are discussed.

Negotiation of parental care when the stakes are high: experimental handicapping of one partner during incubation leads to short-term generosity.

1. Most game theoretical models of biparental care predict that a reduction in care by one partner should not be fully compensated by increased work of its mate but this may not be true for incubating birds because a reduction in care could cause the entire brood to fail. 2. I performed the first handicapping experiment of both males and females during incubation, by placing small lead weights on the tails of male and female northern flickers Colaptes auratus, a woodpecker in which males do most of the incubation. 3. Females responded to the acute stressor (handling and handicapping) by tending to abandon more readily than males and staying away from the nest longer in the first incubation bout. Among pairs that persisted, both males and females compensated fully for a handicapped partner, keeping the eggs covered nearly 100% of the time. 4. Partners did not retaliate by forcing their handicapped mate to sit on the eggs with a long incubation bout length subsequent to having a long bout length themselves. Instead, during the 24 h immediately after handicapping, males behaved generously by relieving handicapped females early. 5. Such generosity was probably not energetically sustainable as these male partners took on less incubation in the 72 h following handicapping compared to female partners of handicapped males. Males and females are probably generous in the short-term because of the high cost of nest failure during incubation but maintaining increased work loads in the longer term is probably limited by body condition and abandonment thresholds consistent with game theory models.

Isolation and characterization of polymorphic microsatellite loci for the northern flicker (Colaptes auratus; Aves).

To investigate the mating system of northern flickers (Colaptes auratus), we developed primers for 14 microsatellite loci and screened them in 68 unrelated adults and their offspring. All markers were highly polymorphic with 9 to +36 alleles per locus. One marker was Z-chromosome linked; one marker exceeded the size standard range and could not be analysed further. We checked the other 12 markers for Mendelian inheritance in 36 broods for which the social parents were known. Seven markers showed evidence for the presence of null alleles, and three of those showed significant deviation from Hardy-Weinberg equilibrium. Markers were generally unlinked.

Learning the ecological niche.

A cornerstone of ecological theory is the ecological niche. Yet little is known about how individuals come to adopt it: whether it is innate or learned. Here, we report a cross-fostering experiment in the wild where we transferred eggs of blue tits, Cyanistes caeruleus, to nests of great tits, Parus major, and vice versa, to quantify the consequences of being reared in a different social context, but in an environment otherwise natural to the birds. We show that early learning causes a shift in the feeding niche in the direction of the foster species and that this shift lasts for life (foraging conservatism). Both species changed their feeding niches, but the change was greater in the great tit with its less specialized feeding behaviour. The study shows that cultural transmission through early learning is fundamental to the realization of ecological niches, and suggests a mechanism to explain learned habitat preference and sympatric speciation in animals.

Evolution of clutch size in cavity-excavating birds: the nest site limitation hypothesis revisited.

There are two major competing hypotheses for variation in clutch size among cavity-nesting species. The nest site limitation hypothesis postulates that nesting opportunities are more limited for weak excavators, which consequently invest more in each breeding attempt by laying larger clutches. Alternatively, clutch size may be determined by diet; the clutch sizes of strong excavators may be smaller because they are able to specialize on a more seasonally stable prey. We built a conceptual model that integrated hypotheses for interspecific variation in clutch size and tested it with comparative data on life-history traits of woodpeckers (Picidae) and nuthatches (Sittidae). In most analyses, diet explained more variation in clutch size among species than did propensity to excavate. Migratory status was positively associated with clutch size but was difficult to distinguish from diet since resident species consumed more bark beetles (a prey available in winter) and had smaller clutches than migratory species. The literature suggests that cavities are not limited in natural, old-growth forests. Although our data do not rule out nest site limitation, we conclude that annual stability of food resources has a larger impact on the evolution of clutch sizes in excavators than does limitation of nest sites.