Dawn chorus and female behaviour in the Willow Tit Parus montanus

We studied the effect of female behaviour on the dawn chorus of Willow Tits Parus montanus. Ten females were temporarily kept in their nest holes in order to delay their dawn emergence. Simultaneously, their mates' singing behaviour was measured. We compared the control dawn behaviour (during which the females emerged when they wanted) with that during the experimental delay. The dawn song by the males lasted longer when the females' emergences were delayed. Our results show that the duration of the dawn chorus is directly related to the female's behaviour. We suggest that at least one function of the dawn song is intrapair signalling, which is often linked to sperm competition.     

Cost of reproduction: parental survival and production of recruits in the Willow Tit Parus montanus

Summary Brood sizes of the Willow Tit were altered experimentally by subtracting or adding two nestlings in 1986 and 1987 in the vicinity of Oulu, northern Finland. The manipulated broods were within the normal range observed in natural conditions. Unaltered broods were used as controls. Data from natural broods from 1978–1985 were available for comparison. When the nestlings were 13 days old they were ringed and weighed and their tarsus, wing, and tail lengths were measured. On the same day the parents were caught, weighed, and measured. In 1986 there were no differences in nestling mortality between the reduced, control, or enlarged broods; i.e. parents were able to fledge the two extra young. In 1987 starvation was most pronounced in the enlarged broods. This resulted in the number of fledglings being practically the same in each manipulation category. Especially the body weight, but also the other indices of body size, decreased as a function of the brood size category, suggesting that there may be quality differences between the young reared in different experimental groups. In 1986 there was a non-significant trend towards lower body weight of the parents attending reduced, control, or enlarged broods, in that order. In 1987 the differences were much smaller. These results were not due to size differences between the groups, so possibly the increased reproductive effort of raising extra young was responsible for the trend observed in 1986. There were no significant differences in parental survival associated with the manipulation category, although the trend in the females was consistent with the hypothesis of reproductive cost. It is possible that environmental conditions in 1986 were so favourable that the tits were not unduly stressed even when attending two extra young. Correlative data from 1978–1985 did not support the cost hypothesis either. A non-significant trend towards reduced post-fledging survival and recruitment of the young was observed with increased brood size. The average fitness value of parents, incorporating parental survival and number of recruits, showed that the success of the adults raising enlarged broods may be lower than that of others. It seems that the reproductive cost, if it exists, decreases individual fitness value by reducing the chances of recruiting descendants into the next generation. The reproductive stress may be insufficient to reduce the subsequent survival of parents. More data are however needed to confirm these results.     

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.     

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.     

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.     

Brood size manipulations within the natural range did not reveal intragenerational cost of reproduction in the Willow Tit Parus montanus

To test for the existence of a reproductive cost, we manipulated brood sizes (-2 and +2 nestlings) over 6 years in a northern population of Willow Tits Parus montanus breeding in natural holes. Possible effects were sought in subsequent survival and fecundity of the parents. Parents given extra chicks made more feeding visits than did parents with reduced and control broods. However, this was not reflected in differences in parental body-weight between groups at the end of the nestling period. Brood size manipulation did not significantly affect female or male survival. In 4 out of 6 years, there was a weak and nonsignificant effect on male survival, consistent with a cost of reproduction. Female and male fecundity in the year following the experiment was not affected by the manipulations. Thus, the data do not give evidence of an intragenerational cost of reproduction in the Willow Tit. Parents appeared unwilling to increase their breeding effort to a level which jeopardized their own survival or future breeding success. It is possible that, because of the time constraints in northern latitudes, females work under their capacity and lay smaller clutches than would otherwise be most profitable. Thus, no costs to the parents would be expected as a consequence of manipulations. These results suggest that the current reproductive rate is not maintained by reproductive cost in the Willow Tit. However, the results do not rule out the possibility that selection has operated outside the current range of reproductive rates during evolutionary history of the species.     

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.     

Phylogeography of the Eurasian Willow Tit (Parus montanus) based on DNA sequences of the mitochondrial cytochrome b gene

The phylogeographic relationships of the trans-Palearctic Willow Tit assemblage were studied by obtaining sequence data from the mitochondrial cytochrome b gene from 34 specimens representing nine subspecies from across the species range. Four distinct genetic groups were identified: Parus montanus weigoldicus, P. m. affinis, P. m. songarus, and a clade containing six Eurasian subspecies (ssp. baicalensis, borealis, montanus, restrictus, rhenanus, and sachalinensis). P. m. weigoldicus, P. m. affinis, and P. m. songarus were reciprocally monophyletic and separated from each other and other subspecies by uncorrected genetic distances between 1.9 and 5.8%. The remaining six subspecies were closely related and shared mitochondrial haplotypes, despite marked morphological and acoustical differences, suggesting a rapid evolution of distinct vocalization patterns. The current classification splitting the species into the songarus and montanus subspecies groups is not concordant with our phylogeny. Also, the four regiolect groups, Lowland, Alpine, Siberian, and Sino-Japanese, are not fully mirrored in the phylogeny. Our data suggest that the mono-frequency song type may be ancestral and was retained over a long evolutionary time in certain populations, but was altered or camouflaged by learning processes in others.     

Initial winter flock formation in the Willow Tit Parus montanus. Do immigrating juveniles assess the quality of territorial adults?

In the Willow Tit Parus montanus , forming of winter flocks starts shortly after the juveniles have become independent of their parents. In the early phase of flock formation the juveniles roam about prospecting for suitable winter territories. They visit several occupied territories in succession and seek contact with the territory holders. I describe the behaviour of juveniles during such encounters with adult conspecifics. I suggest that, by approaching adults closely and behaving specifically, juveniles gain important information on the adults' innate qualities; this may affect the settling juveniles' future survival.     

Long‐Distance Calling by the Willow Tit,Poecile montanus,Facilitates Formation of Mixed‐Species Foraging Flocks

The occurrence of mixed‐species foraging flocks is a worldwide phenomenon in terrestrial bird communities. Previous studies suggest that individuals participating in flocks might derive benefits in terms of improved feeding efficiency and/or reduced risk of predation. However, very little is known about how individuals establish mixed‐species flocks. Here, I provide the first experimental evidence that long‐distance calling by the willow tit, Poecile montanus, facilitates the establishment of mixed‐species flocks at a foraging patch. Observations at experimental foraging patches showed that willow tits that find a food source produce long‐distance calls, particularly when they are isolated from conspecific flockmates. The probability of long‐distance calling was negatively correlated with the number of heterospecific foraging individuals near the food source. A playback experiment confirmed that calls attract both conspecific and heterospecific members of foraging flocks. This study demonstrates that willow tits use long‐distance calls to attract conspecific flockmates to foraging patches, and these calls can also facilitate the formation of mixed‐species flocks on patches.     

Social rank and antipredator behaviour of Willow Tits Parus montanus in winter flocks

Field experiments on three free-ranging Willow Tit winter flocks, each consisting of one adult pair and two male and two female juveniles (first-year birds), were performed to examine whether preferences for feeding site and antipredator behaviour are related to social rank. The dominance structure was the same in all flocks; adult male > juvenile male 1 > juvenile male 2 > adult female > juvenile female 1 > juvenile female 2.
The proportion of time spent scanning for predators was positively correlated with distance from cover, and adults scanned relatively more than juveniles at the same controlled distance from cover, especially in the afternoon. Given a choice between feeders placed 1 m, 3 m, 5 m, 10 m and 20 m from the forest edge, the tits preferred feeders close to cover. Low-ranking individuals used feeders farther from cover indicating that higher ranked tits prevented them from using the feeders close to cover by means of social dominance. When only the 10m and 20 m feeders were baited, only low-ranked juveniles visited the feeders, subordinate females slightly more than males. The subordinate juveniles increased their use of exposed feeders at low ambient temperature, suggesting that they are prepared to take greater risks during cold periods.
The sequence of return to a feeder, after a life-like stuffed predator model mounted 1 m from a feeder opening was removed, was positively correlated with dominance status, revealing that subordinates take the greatest predation risks.     

Social dominance in free-living Willow Tits Parus montanus: determinants and some implications of hierarchy

Several features of social dominance among Willow Tit Parus montanus winter flocks were examined during a four-winter study. Birds of both sexes were evenly distributed over the 33 flocks studied. In nearly half of the flocks there was an adult pair accompanied by yearlings, but one-third of the flocks consisted of more than two adults with yearlings. Males were found to be dominant over and larger than females. Within a sex, yearlings were usually subordinate to adults. The effect of size on dominance, after controlled for sex and age, remained obscure in our field data. The hierarchical status of an individual was found to rise or at least stay the same in different years, which supports the "hopeful dominants" hypothesis, i.e. birds stay in flocks hoping to achieve a higher status in the future. The ranks of mates correlated highly significantly, implying that high-ranking birds were paired with other high-ranking birds and low-ranking birds with other low-ranking birds. Birds of different age and sex did not show any differences in the proportion of initiated aggressive encounters directed at other individuals. However, males were more aggressive to other males than to females and also tended to behave less aggressively towards their own mates than towards other individuals in the flock. This could be a male strategy to strengthen the pair-bond and to enhance mate protection described earlier in     

Factors influencing Willow Tit Poecile montanus site occupancy: a comparison of abandoned and occupied woods

The British Willow Tit Poecile montanus kleinschmidti underwent a decline of 85% between 1970 and 2003. The cause of this decline is unknown. However, several hypotheses have been put forward to account for it: competition from other tit species, predation by Great Spotted Woodpeckers Dendrocopos major and habitat change. In order to test these, woods that are currently occupied by Willow Tits were paired with woods (within 50 km) that had been abandoned by Willow Tits five or more years previously. Point counts for other tit species (potential competitors) and woodpecker species (potential predators) were carried out at ten evenly spaced points throughout each wood. Habitat variables were collected within a 50-m radius of where a Willow Tit was located (in the occupied woods) or where maps showed a Willow Tit had been located (for abandoned woods). No evidence was found for differences in numbers of potential competitor or potential predator species in abandoned and occupied woods. Soil water content was found to be higher at occupied sites. No other habitat features differed between the two categories of site. The drying up of British woods could therefore be implicated in the Willow Tit decline and this warrants further investigation.     

Age Differences in the Response of Willow Tits (Parus montanus) to Conspecific Alarm Calls

Predation is an important mortality factor in wintering birds. To counter this, birds produce alarm calls in the presence of predators which serve to warn conspecifics. In social hierarchical bird flocks, adults survive the winter better than juveniles and therefore survival strategies probably vary with social status. This study examined the differential responses to alarm calls by free-living willow tits, Parus montanus, in dominance-structured winter flocks in Finland. To explore the age-dependent differences in response to conspecific alarm calls, a series with three alarm calls was played to focal adults and juveniles while they sat in the middle section of a spruce branch. Immediately after the playback, juvenile willow tits moved more often, flew longer distances and changed branches more often than did adults. Previous mammal studies have shown that juveniles are more likely to flee than adults after hearing conspecific alarm calls. The current study demonstrates that similar age-dependent responses to conspecific alarm calls occur in birds also. These findings reflect an increased vulnerability to predators or lack of experience of young birds.     

Effect of Willow Tit Poecile montanus alarm calls on attack rates by Pygmy Owls Glaucidium passerinum

One suggested anti‐predator function of alarm calls is to deliver a message to a predator that it has been detected. Moreover, giving the alarm call could provide a signal to the predator that capturing the individual giving the alarm is more difficult than capturing its silent group members, as the caller is probably the most aware of the predator's location. In an aviary experiment using stuffed dummy Willow Tits Poecile montanus, we assessed whether an authentic alarm call given by Willow Tit affected Pygmy Owl Glaucidium passerinum prey preference. In the experiment, the Owls attacked only the ‘silent’ dummy individuals, suggesting that alarm calling could offer direct fitness benefits to the caller by decreasing the attack risk of the caller relative to its group members.     

Willow tit Parus montanus extrapair offspring are more heterozygous than their maternal half‐siblings

Through extrapair matings, males can sire additional offspring with low cost and females may look for direct benefits in form of food or additional paternal care or gain genetic benefits that increase offspring fitness. We studied the patterns of female mate choice and frequency of extrapair paternity in the socially monogamous willow tit Parus montanus using microsatellites. We also examined the effect of heterozygosity on the growth rate and survival of the chicks. We found 25 mixed‐paternity broods out of 117 broods of which both parents were sampled. Altogether, 6.7% of sampled chicks were classified as extrapair young. The pairwise relatedness of social pairs did not correlate with the percentage of extrapair young in the brood and there was no difference in heterozygosity between promiscuous and monogamous parents. However, the extrapair young were more heterozygous than the within‐pair young in the mixed‐paternity broods. The maternal half‐siblings in mixed paternity broods were similar in body size. Thus, there was no indication for different growth rate between the siblings, but there were indications that heterozygosity affects survival.     

Winter spacing and non-breeding social system of the Coal Tit Parus ater in a subalpine forest

In the genus Parus, non-breeding social territoriality has often been attributed to the presence of food hoarding which, by increasing the value of an area, is thought to favour the existence of exclusive territories. To test this, it is necessary to analyse the social structure of hoarding populations inhabiting different environments. I studied Coal Tit spacing patterns and winter social systems in a colour-ringed hoarding population inhabiting a mountain coniferous forest in the Pyrenees. Using data on home range, range overlap and social coherence, the social system was found to be based on adult birds that had previously bred together and which occupied relatively small home ranges. Large home ranges of resident juveniles overlapped extensively with each other and with those of adults. However, resident juveniles were resighted much closer to their ringing place than was expected at random, suggesting restricted floater ranges. Associations, composed mainly of juveniles, seemed to be rather large and loose, with little coherence. No territorial defence by any group of birds was detected during the study. Therefore, the social organization described for the present Coal Tit population is similar to the loose, non-territorial basic flock system of non-hoarding species. Resident social patterns seemed to be linked primarily to the availability of breeding resources, probably leading to the high site fidelity of adults and the floater life of resident juveniles. On the other hand, short-term survival chances may attract transients to a rich food supply and thus affect flocking strategies of birds. High availability of resources allowing a high population density seems to be the main proximate factor influencing the spacing patterns described.     

Evidence for cuckoldry in the Great Tit Parus major

We estimated the frequency of cuckoldry (the proportion of offspring resulting from extra-pair copulations) in Great Tits Parus major using heritability estimates based on the resemblance of offspring tarsus-length to that of their parents. Our results suggest that cuckoldry occurs and may be relatively common in certain years.