Poor nutritional condition as a consequence of high dominance status in the Coal Tit Parus ater

The costs and benefits of dominance status have been investigated in the past and it has generally been reported that subdominant birds are at a nutritional disadvantage owing to their low dominance status. The nutritional condition of birds during winter can be important in determining their likelihood of survival. This is particularly so in small passerines that are sensitive to severe weather conditions. Ptilochronology is an accurate method used to produce a long-term estimate of body condition spanning the number of days that it takes to grow a new feather. Ptilochronology was used during this study to estimate the nutritional condition of Coal Tits Parus ater during one winter and how condition was affected by dominance status. Dominant Coal Tits produced poorer quality feathers, which they grew at a slower rate, than did subdominant conspecifics. This study highlights a nutritional cost to high dominance status that could have long-term consequences because the induced tail feathers will not be replaced for at least 5 months.     

Territory acquisition during winter by juvenile Willow Tits Parus montanus

Willow Tits Parus montanus live within exclusive territories in non-kin winter flocks with a dominance hierarchy in which males dominate females and adults dominate juveniles. Because they live in habitats that are saturated with dominant territory owners, the options of subordinate juveniles surviving the winter are: (1) to become a dominant owner of a vacated territory, (2) to stay in an occupied territory and hope for ownership or (3) to leave the area and hope for territory acquisition elsewhere. Removal experiments during December to February revealed that particularly the most dominant of the juvenile pairs left their own flock territory and replaced birds experimentally removed from a neighbouring flock, if by doing so they increased their own dominance position. It is suggested that juvenile Willow Tits updated their information about the composition and dominance structure of adjacent winter flocks by regularly visiting these territories and attempted to become dominant owners during winter, thereby increasing their chances of breeding.     

Advantages of social foraging of Willow Tits Parus montanus

The mean number of Willow Tits Parus montanus in single-species flocks was significantly larger than in mixed-species flocks of Willow and Coal Tits P. ater. Both flock size and the tendency of Willow Tits to join mixed-species flocks were negatively correlated with ambient temperature, probably because each bird, when the metabolic rate of the birds increased, could allocate more time to foraging due to improved predator detection by many eyes. The vigilance time of Willow Tits decreased with flock size and was determined by the total number of individuals in a flock rather than by the number of Willow Tits in mixed-species flocks of Willow and Coal Tits.     

Seeing without being seen: a removal experiment with mixed flocks of Willow and Crested Tits Parus montanus and cristatus

This paper tests the hypothesis that foraging site selection reflects a trade-off between the various needs for concealment from predators, to find food, and for the individual to maintain some view of its surroundings. After removal of Crested Tits Parus cristatus (the dominant species in mixed flocks), Willow Tits P. montanus did not decrease their foraging heights as expected but remained in the most exposed parts of young pines. In contrast, after removal of Willow Tits, Crested Tits increased their foraging height from the sheltered lower canopy to sites previously occupied by Willow Tits. When flock size was reduced, the birds maintained the same high levels of vigilance without concealing themselves in dense vegetation. I suggest that flock members may benefit from foraging in sites that afford good anti-predator vigilance.     

DIFFERENTIATION OF FORAGING NICHE AMONG TITS, PARUS SPP., IN NORWAY DURING WINTER

The winter foraging behaviour of Willow Tit Parus montanuss, Coal Tit P. ater and Crested Tit P. cristatus was studied in several spruce-dominated, coniferous forest areas in which either Willow Tit alone, Willow Tit and Coal Tit, or all three species together were present. In the areas where Crested Tits were absent, Willow Tits foraged significantly more often in the outermost parts of spruce branches and less frequently on the parts of branches closer to the trunk, than in the area in which Crested Tits were present. In the area in which all three species were present, the same change in foraging pattern of Willow Tits was found in mixed flocks in which Crested Tits were lacking. The foraging pattern of Coal Tits was unaffected by the presence of members of either or both the other two tit species. It is suggested that the expansion of the feeding niche of Willow Tit in the absence of Crested Tit is a result of ecological release, due to a reduction in the degree of interspecific competition.     

Subordination in mixed-age bird flocks–a removal study

During the winters of 1979 to 1986, flocks (7–17 per season, 78 total) of Willow Tits Parus montanus (generally consisting of an adult pair and four juveniles) were studied in a subalpine mixed forest in Central Norway. The mean survival rate between December and early April for juveniles (0.32) was significantly lower than that found for adults (0.74). As it has previously been found that adult Willow Tits, being dominant, prevent the juveniles in a flock from using preferred foraging sites, a removal experiment was made to see if the survival of juveniles improved when living in flocks with no adults present. After removal of the adult pair from 13 flocks in early January, the juveniles from these flocks survived significantly less well, up to early April, than either the juveniles in 11 flocks in which the adults were present (controlled for flock size) or those in 14 flocks consisting of an adult pair and four juveniles. Despite a skewed access to the resources in favour of dominant adults, the cost of being a subordinate juvenile is probably outweighed by the improved predator evasion and improved food finding efficiency due to the greater experience of the adult pair possessing the territory.     

Ptilochronology proves unreliable in studies of nestling Pied Flycatchers

Ptilochronology does not appear to be a reliable measure of the daily growth rate of contour feathers or the nutritional state of nestling Pied Flycatchers Ficedula hypoleuca . Growth bars on primary remiges, which according to ptilochronology represent a day's increment of feather growth, are only about half as wide as the actual daily increase in the length of these feathers while they are growing. The average width of the growth bars on primaries was also uncorrelated with other commonly used measures of growth or nutritional status (increase in body mass or in the size of the wing or tarsus), although these were highly correlated with each other. In adult flycatchers, the average width of the growth bars on tertials was unrelated to the average bar width on greater coverts, although both feathers are replaced during the winter (prenuptial) moult. This suggests that the growth bars either do not reflect the nutritional status of adults during normal periods of moult or that contour feathers in different tracts vary in their sensitivity to the nutritional status of the moulting bird. To our knowledge, this is the first time that anyone has attempted to apply ptilochronology to nestlings. It is noteworthy that a method of measuring growth and nutritional state that has shown promise when applied to induced feathers of adult birds seems to be unreliable when applied to the developing plumage of nestlings, and perhaps the normal (not-induced) replacement plumage of adults.     

Juvenile coloration of Florida Scrub-Jays ( Aphelocoma coerulescens ) is sexually dichromatic and correlated with condition

The Florida Scrub-Jay is a monogamous cooperative breeder in which both males and females display extensive structurally based blue plumage. Juveniles of this species exhibit blue tail and wing feathers that they begin growing as nestlings, and some of these feathers are retained throughout their first year. Although the birds appear to be sexually monochromatic, we assessed whether cryptic dichromatism exists in both the magnitude and pattern of coloration in tail feathers of juvenile Florida Scrub-Jays. We then determined whether variation in plumage coloration is associated with nutritional condition during molt. Tails of juvenile male Florida Scrub-Jays exhibit a greater proportion of UV reflectance than those of females. Mass at age 11 days and ptilochronology of the juvenile tail feathers were used as measures of individual nutritional condition during feather growth, and the latter was found to be positively associated with UV chroma. These data demonstrate that Florida Scrub-Jays are sexually dichromatic and suggest that variation in plumage color may be condition dependent, although we cannot rule out alternative explanations. Juvenile plumage coloration, therefore, has the potential to function as a signal of individual quality in both males and females.     

STRUCTURE AND FORAGING PATTERNS OF FLOCKS OF TITS AND ASSOCIATED SPECIES IN AN ENGLISH WOODLAND DURING THE WINTER

Blue Tits were the commonest and most frequent members of mixed-species insectivorous flocks during the winter at Wytham Wood, Oxford. Six common flocking species (Blue Tit, Longtailed Tit, Great Tit, Coal Tit, Marsh Tit and Golderest) made up nearly two-thirds of the birds in the study area. A tentative interspecific social dominance hierarehy was constructed: Great Tit > Blue Tit > Marsh Tit > Coal Tit > Long-tailed Tit and Goldcrest. Blue Tits permitted conspecifics to approach them more closely than did other flock members. Blue Tits most frequently led the flocks, but no more than would be predicted by their abundance. Great Tits, and Marsh Tits and Coal Tits to a lesser extent, defended territories during the winter. Two of the six common species frequented the inner parts of branches (Great Tit, Coal Tit), two the outer parts of branches (Blue Tit, Marsh Tit), and two the twigs (Long-tailed Tit, Goldcrest). Members of each of these species-pairs showed marked differences in height of foraging and/or species of tree frequented. Because of their abundance, the impact of Blue Tits outside of their most highly frequented foraging zones may exceed that of species concentrating in these other zones (e. g., twigs high in trees). The Coal Tit foraged most diversely, the Great Tit least diversely. Species that foraged diversely in one of the three foraging categories (species of tree, substrate, height) usually foraged less diversely than most other species in the other dimensions. Species usually overlapped least in the zones exploited (inner parts of branches, etc.). Great Tits overlapped with other species less than did any other common flock member, and Coal Tits were the next lowest in this regard.     

Increase of feather quality during moult: a possible implication of feather deformities in the evolution of partial moult in the great tit Parus major

Here we investigate the change in feather quality during partial post‐juvenile and complete post‐breeding moult in great tit Parus major by measuring the change in the number of fault bars and feather holes on wing and tail feathers. Feathers grown during ontogeny usually are of lower quality than feathers grown following subsequent moults at independence. This is reflected by higher number of fault bars and feather holes on juveniles compared to adults. Fault bars are significantly more common on tail and proximal wing feathers than on the distal remiges, indicating a mechanism of adaptive allocation of stress induced abnormalities during ontogeny into the aerodynamically less important flight feathers. On the contrary, feather holes produced probably by chewing lice have a more uniform distribution on wing and tail feathers, which may reflect the inability of birds to control their distribution, or the weak natural selection imposed by them. The adaptive value of the differential allocation of fault bar between groups of feathers seems to be supported by the significantly higher recapture probability of those juvenile great tits which have fewer fault bars at fledging on the aerodynamically most important primaries, but not on other groups of flight feathers. The selection imposed by feather holes seems to be smaller, since except for the positive association between hatching date, brood size and the number of feather holes at fledging, great tits' survival was not affected by the number of feather holes. During post‐juvenile moult, the intensity of fault bars drops significantly through the replacement of tail feathers and tertials, resulting in disproportional reduction of the total number of fault bars on flight feathers related to the number of feathers replaced. The reduction in the number of fault bars during post‐juvenile moult associated with their adaptive allocation to proximal wing feathers and rectrices may explain the evolution of partial post‐juvenile moult in the great tit, since the quality of flight feathers can be increased significantly at a relatively small cost. Our results may explain the widespread phenomenon of partial post‐juvenile moult of flight feathers among Palearctic passerines. During the next complete post‐breeding moult, the total number of fault bars on flight feathers has remained unchanged, indicating the effectiveness of partial post‐juvenile moult in reducing the number of adaptively allocated fault bars. The number of feather holes has also decreased on groups of feathers replaced during partial post‐juvenile moult, but the reduction is proportional with the number of feathers moulted. In line with this observation, the number of feather holes is further reduced during post‐breeding moult on primaries and secondaries, resulting in an increase in feather quality of adult great tits.     

Social rank in winter flocks of Willow Tits Parus montanus

The social hierarchies in winter of ten flocks of Willow Tits Parus montanus were studied when the birds were foraging naturally and when visiting feeders. All the flocks consisted of one adult mated pair together with two juvenile males and two juvenile females (probably pairs). All flocks studied had a stable composition and the hierarchies remained constant throughout the study period. The hierarchies were linear and unilateral. The adults of each sex dominated the respective juveniles and within each age group the male dominated the female. The dominance relationships between the age and sex groups were not consistent. Although the males dominated all the females in six flocks, in one flock the adult female dominated both the juvenile males, but only one of them in three other flocks. The degree of aggression between flock-members was 0.8 encounters per hour, and males initiated 94% of all attacks. Body-weight explained 77% of the variation in dominance rank. It is suggested that the dominance rank of a male is also a function of his seniority, while the rank of a juvenile female is correlated with the rank of her mate.     

Winter territoriality and flock cohesion in the black-capped chickadee Parus atricapillus

In northwest Massachusetts, black-capped chickadees shifted foraging groups repeatedly each day during the winter. As a result, there was no clear distinction between flock members and floaters, but instead a continuous gradation from birds that formed relatively stable associations to those that shifted groups more frequently. Group territories were absent and individual territories were at most poorly defended, if present at all. This picture differs from other recently studied populations, where flock membership is stable and flock territories are clear. Comparison with other populations indicates that winter social structure in the black-capped chickadee may be sensitive to both winter food levels and the proportion of year-round residents. This indicates that cost-benefit analyses of winter social organization should consider factors acting throughout the year as well as the residence status of the wintering birds. The data suggest that flock size, population density and home range size are sensitive to winter food on the study area, whereas territoriality and flock cohesion may respond more clearly to the local availability of breeding resources or the proportion of year-round residents.     

Plasma levels of luteinizing hormone and steroid hormones in free-living winter groups of willow tits (Parus montanus)

Plasma levels of LH, DHT, testosterone, and corticosterone were measured for all members in free-living winter flocks of willow tits, Parus montanus. Hormonal data were related to (1) flock size and (2) age/sex differences. The winter flock defends a large winter territory and shows a well-established social hierarchy in which adults consistently dominate first-year birds. One winter group normally consists of four individuals, two adults and two juveniles. In flocks containing four or five members juvenile willow tits had significantly higher corticosterone values than adults. In small-sized groups, containing three members, all individuals had high plasma levels of corticosterone. No other effects of flock size was found. When data were treated on an age/sex basis, i.e., flock size was not considered, juvenile females were found to have significantly higher plasma levels of testosterone than adult birds, and also significantly higher levels of DHT than juvenile males and adult females. Also, juvenile willow tits had significantly higher plasma levels of corticosterone than adult birds.     

Sex-dependent frequency and consequences of natural handicaps in American Kestrels

Natural selection theory suggests that there should be fitness consequences for animals that possess morphological abnormalities such as missing or deformed appendages and other structures. As these characters are expected to be detrimental, morphological abnormalities could be considered “handicaps”. However, little information exists about the nature, prevalence or consequences of naturally occurring handicaps in birds, even though experimental handicapping has been used in some behavioural studies. Here, we document and describe the frequency of natural handicaps in American Kestrels Falco sparverius, and investigate their influence on condition and survival. We considered three different categories of handicap that should negatively impact falcons: missing remiges and rectrices, broken remiges and rectrices, and foot handicaps. Broken feathers, followed by missing feathers, were the most common handicaps detected in kestrels and their prevalence increased throughout the breeding season. Foot handicaps were relatively rare and showed no seasonal trends. There was no effect of any handicap type on body condition or return rates of male kestrels. In contrast, females with either foot handicaps or broken feathers were in significantly poorer condition than control females, and females with broken feathers were less likely than control females to return to the study area in years following initial capture. Given that female kestrels with broken feathers have more fault bars than those without broken feathers, and fault bar formation is related to stresses in a bird's past, we suggest that female kestrels that had broken feathers were of poorer quality than those without broken feathers or those with missing feathers. Females with broken feathers may therefore have been in poorer condition initially than other birds, or if broken feathers are indicative of poor quality, then females with broken feathers may have been less able to cope with the negative impacts of feather loss on flight performance compared to birds with missing feathers.     

Winter survival and breeding success of dominant and subordinate Willow Tits Parus montanus

We studied the effect of winter rank on survival rate and reproductive success in Willow Tits Parus montanus, a resident passerine living in dominance-structured flocks during the nonbreeding season, in 6 years. Winter survival was dependent on both the birds' age and rank. Adults survived better than first-year birds, and within first-year males, dominants survived better than subordinates. In other sex and age classes, rank did not contribute to survival. Although first-year males were in excess among nonbreeders, no connection existed between breeding status and rank. Female rank did not explain the variation in the start of laying, clutch size, number of fledglings or recruit production. We conclude that social status in Willow Tits affects individual fitness mainly through rank-dependent survival. Acquiring a high rank position seems to be most important for first-year birds, especially first-year males.     

Winter body condition in relation to age,sex and plumage ornamentation in a migratory songbird

Winter body condition may play important roles in the life history of migratory birds, but it is difficult to estimate. We used the growth rate of winter‐grown tail feathers of Collared Flycatchers Ficedula albicollis as an indicator of winter body condition, comparing this trait between age classes and sexes and relating it to plumage ornamentation (forehead and wing patch sizes). Adults and males were in better nutritional condition during winter, as indicated by their faster tail feather growth rates, than were yearlings and females, respectively, which could indicate differences in individual quality and foraging ability with age, or age‐ and sex‐related winter habitat segregation. However, feather growth rate was related neither to the size of the winter‐grown forehead patch nor to the size of the summer‐grown wing patch, suggesting weak condition‐dependence for the winter‐grown ornament and complex life‐history consequences for the summer‐grown ornament.     

Ontogeny of food storing in titmice Parus spp.

The ontogeny of food storage in Crested Tits Parus cristatus and Willow Tits P. montanus starts during the period of parental care after the young have left the nest and develops gradually. In southern Norway, the storing behaviour becomes fully developed during August-September, i.e. at a time when most juveniles have completed their dispersal period and settled in permanent winter flocks. The food storing behaviour is largely innate, but is improved by practice and experience.     

Winter feeding influences the cost of living in boreal passerines

The plastic regulation of internal energy reserves is acknowledged as the main adaptive response to winter conditions of resident small birds in northern latitudes, a strategy that may be altered whenever human‐supplemented food is available. We investigated the effects of supplementary feeding on the energy management strategy of two wild passerine species, the Willow Tit Poecile montanus and Blue Tit Cyanistes caeruleus, wintering in boreal conditions by measuring body mass and the energy cost of living, i.e. basal metabolic rate. Individuals of both species were heavier, larger and exhibited a higher energy cost of living when captured at the feeders than were individuals captured away from feeders. Fed Willow Tits expended more energy in maintenance, although this difference disappeared once mass was accounted for. Conversely, Blue Tits at feeders had higher mass‐adjusted energy cost of living, but only at low ambient temperatures. The results indicate that winter feeding has species‐specific effects on overall energy management strategy and modifies the response to environmental conditions of wintering passerines.     

Factors influencing patch occupancy and within‐patch habitat use in an apparently stable population of Willow Tits Poecile montanus kleinschmidti in Britain

Capsule Birds selected younger woodland patches with fewer conifers and higher soil moisture content; within these patches, they selected areas with higher vegetation cover at 2–4 m, and less bracken.

Aims For an apparently stable population, to investigate habitat differences between occupied and unoccupied woodland patches and habitat selection within occupied woodlands.

Methods We surveyed woodland patches for breeding Willow Tits in April 2006 in an area with an apparently stable population. Habitat was measured at points spread throughout each woodland patch and at points where Willow Tits were located. We compared habitat between occupied and unoccupied sites. In addition, within occupied sites, we examined habitat differences between Willow Tit locations and the points spread throughout the wood.

Results We surveyed 65 woodland patches, of which 29 were occupied. Willow Tits were more likely to be found in deciduous woodland that was younger and had higher levels of soil moisture. Probability of occupancy fell from 60% for woods aged 20–25 years to only around 15–30% for woods aged over 80 years. Within occupied woods, Willow Tits were more likely to be found in areas with more vegetation cover at the 2–4 m level and in areas with less bracken.

Conclusions Habitat management for Willow Tits should involve provision of young woodland patches with moist soils. Changes to the age structure and/or the soil moisture content of woods may be implicated in the dramatic decline of Willow Tits in Britain.     

Fault bars timing and duration: the power of studying feather fault bars and growth bands together

Growth bands and fault bars, widespread features of feathers that form during regeneration, have largely been studied independently. Growth bands result from normal regeneration: each pair of dark/light bands forms every 24 h. Fault bars are a response to stress during regeneration, creating a translucent line that can break the feather. We studied the relative position and width of these two structures in feathers of nestling and adult white storks Ciconia ciconia. We first confirmed that one growth band represents 24 h of feather regeneration. Fault bars did not occur at random within growth bands: 65.7% (in nestlings) and 45.6% (in adults) of them occurred in one out of six defined segments within a growth band, namely that segment corresponding to the first one‐third of night time hours. The width of fault bars relative to growth bands suggested that fault bars were produced during a median (range) of 7.0 h (2.7–27.0) in nestlings and 3.7 h (1.8–7.9) in adults. Fault bars were concentrated at feather tips in nestlings, but at central locations in adult feathers. Our results suggest that, in general, fault bars are a discrete event of a finite duration occurring mainly during the night (particularly in nestlings). This, along with current knowledge, suggests that acute stressors, rather than chronic ones, are responsible for fault bar formation. Thus, such acute punctual stressors (a matter of minutes) can have long‐lasting (months?years) physiological effects due to the wing load increase from feather breakage caused by fault bars.