Feather mite abundance increases with uropygial gland size and plumage yellowness in Great Tits Parus major

Plumicolous feather mites are ectosymbiotic organisms that live on bird feathers. Despite their abundance and prevalence among birds, the ecology of the interaction between these organisms and their hosts is poorly known. As feather mites feed on oil that birds spread from their uropygial gland, it has been hypothesized, but never tested, that the number of feather mites increases with the size of the uropygial gland of their hosts. In this study the number of feather mites is considered with respect to uropygial gland size in a breeding population of Great Tits Parus major in order to test this hypothesis. As predicted, the number of feather mites correlated positively with the uropygial gland size of their hosts, showing for the first time that uropygial gland size can explain the variance in feather mite load among conspecifics. Previous studies relating feather mite load to plumage colour have suggested that feather mites may be parasitic or neutral. To confirm this, the yellowness of breast feathers was also assessed. However, the results ran in the opposite direction to that expected, showing a positive correlation between mite load and plumage yellowness, which suggests that further work is needed to give clear evidence for a specific nature of feather mites. However, Great Tits with higher mite loads had lower hatching and breeding success, which may support the idea that feather mites are parasites, although this effect must be taken with caution because it was only found in males. Age or sex effects were not found on the number of feather mites, and it is proposed that hormonal levels may not be sufficient to explain the variation in feather mite loads. Interestingly, a positive correlation was detected between uropygial gland size and plumage brightness, which could be a novel factor to take into account in studies of plumage colour.     

Ultraviolet reflectance of plumage for parent-offspring communication in the great tit (Parus major)

Ultraviolet (UV) reflectance has been implicated in mate selection.Yet, in some bird species the plumage of young varies in UVreflectance already in the nest and long before mate choiceand sexual selection come into play. Most birds molt the juvenilebody plumage before reaching sexual maturity, and thus, someconspicuous traits of the juvenile body plumage may rather haveevolved by natural selection, possibly via predation or parentalpreference. This second hypothesis is largely untested and predictsa differential allocation of food between fledging and totalindependence, which is a time period of 2–3 weeks whereoffspring mortality is also highest. Here, we test the predictionthat parents use the individual variation in UV reflectanceamong fledglings for differential food allocation. We manipulatedUV reflectance of the plumage of fledgling great tits Parusmajor by treating chest and cheek feathers with a lotion thateither did or did not contain UV blockers and then recordedfood allocation by parents in an outdoor design simulating postfledgingconditions. The visible spectrum was minimally affected by thistreatment. Females were found to feed UV-reflecting offspringpreferentially, whereas males had no preference. It is the firstevidence showing that the UV reflectance of the feathers ofyoung birds has a signaling function in parent–offspringcommunication and suggests that the UV traits evolved via parentalpreference.     

Pair bond and breeding success in Blue Tits Parus caeruleus and Great Tits Parus major

Data from 939 nests of the Blue Tit Parus caeruleus and 1008 nests of the Great Tit P. major from nestboxes provided in superabundance in mixed forest study sites between 1976 and 2001 were analysed to examine the effects of mate retention on breeding success and the relationship between mate fidelity and site fidelity. Most birds retained their former partner (76% in Great Tits and 65% in Blue Tits). The probability of a pair divorcing was affected by male age in Great Tits, divorce being more likely in pairs with first‐year males. Great Tit pairs breeding together for a second season bred earlier, but had no higher breeding success than pairs breeding together for the first time. In Blue Tits laying date and start of incubation tended to be earlier in pairs breeding together for a second season, but hatching and fledging dates were not earlier than in other pairs. Great Tit pairs breeding together for two consecutive seasons bred earlier in the second season than in the first, but breeding success did not differ significantly between years. In both species, breeding performance did not differ between pairs that divorced after a season and pairs that stayed together. Thus breeding success did not determine whether a pair divorced or bred together again. Neither Blue Tits nor Great Tits improved their breeding performance through divorce. Blue Tit females even had fewer fledglings in the year after divorce than in the year before. Mate retention affected breeding site fidelity. Blue Tit females had greater breeding dispersal distances between consecutive years when re‐mating than when breeding again with the same mate. In Great Tits both males and females dispersed more when re‐mating than when retaining the former partner, suggesting that mate retention increased the chance of retaining the breeding site. In both species, breeding dispersal distances did not differ between pairs that divorced and pairs in which one mate disappeared. Because no major advantage of mate retention was evident, we suggest that mate retention evolved under different conditions than those found in study sites with high breeding densities and a superabundance of artificial nesting sites.     

Interactive vocal communication at the nest by parent Great Tits Parus major

Although most bird species show monogamous pair bonds and bi‐parental care, little is known of how mated birds coordinate their activities. Whether or not partners communicate with each other to adjust their behaviour remains an open question. During incubation and the first days after hatching, one parent – generally the female – stays in the nest for extended periods, and might depend on acoustic communication to exchange information with its mate outside. The Great Tit Parus major is an interesting study system to investigate intra‐pair communication at the nest because males address songs to their mate while she is in the nest cavity, and females answer the male from the cavity with calls. However, the function of this communication remains unknown. In this study, we recorded the vocalizations and observed the resulting behaviour of Great Tit pairs around the nest at different breeding stages (laying, incubation and chick‐rearing). We observed vocal exchanges (vocalization bouts, alternated on the same tempo, between the female inside the nest and her male outside) in three contexts with different outcomes: (1) the female left the nest, (2) the male entered the box with food, and the female then used specific call types, (3) mates stopped calling but did not leave or enter the nest. The structure of vocal exchanges was globally stable between contexts, but females used calls with an up‐shifted spectrum during exchanges, at the end of which they left the nest or the male entered the nest. Birds vocalized more and at higher tempo during exchanges that ended up in feeding inside the nest. Birds also vocalized more during exchanges taking place during laying – a period of active mate guarding – than during incubation. We conclude that vocal exchanges could signal the females’ need for food and the males’ mate guarding behaviour, and discuss other possible functions of this communication.     

Body temperature responses of Great Tits Parus major to handling in the cold

Animals typically respond to stressful stimuli such as handling by increasing core body temperature. However, small birds in cold environments have been found to decrease body temperature (T_b) when handled over longer periods, although there are no data extending beyond the actual handling event in such birds. We therefore measured both the initial T_b decrease during ringing and standardized T_b sampling, and subsequent recovery of T_b after this handling protocol in wild Great Tits Parus major roosting in nestboxes in winter. Birds reduced their T_b by 2.3 °C during c. 4 min of handling. When birds were returned to their nestboxes after handling, T_b decreased by a further 1.9 °C over c. 2 min, reaching a T_b of 34.6 °C before taking 20 min to rewarm to 2.5 °C above their initial T_b. The T_b reduction during handling could be a consequence of increased heat loss rate from disrupted plumage insulation, whereas T_b reduction after handling might reflect reduced heat production. These are important factors to consider when handling small birds in the cold.     

The significance of clutch overlap in Great Tits Parus major

We studied the effects of manipulation of the size of first broods in the Great Tit Parus major on the size and breeding success of second clutches and its relation to the degree of clutch overlap. The rearing of first brood fledglings always overlapped with the laying of the second clutch and in most cases also with the incubation period of the latter. The degree of clutch overlap depended on the size of the first brood, being less when the first brood was large. Clutch overlap also increased with season. Mechanisms affecting the timing of laying of second clutches are discussed. A large first brood imposed reproductive costs. It affected the size of the second clutch by causing it to be delayed; second clutch size decreases with season. It affected the post-fledging survival of second brood young as, in this population, this decreases with fledging date. The breeding success of second clutches was, however, not affected by the size of the first brood, but instead by the weight of the female when rearing the first brood.     

Early morning fledging improves recruitment in Great Tits Parus major

A potential key event linking the nestling phase to first‐year survival is fledging (nest leaving) because this process is characterized by a major change of environments and therefore a sudden shift in selective forces. Here we assessed whether different facets of fledging predicted subsequent survival (measured as local recruitment) in Great Tits Parus major. Nestlings had a higher recruitment probability when they fledged early in the morning and when they were heavy. The existence of selection for fledging early in the day has been suggested before, but here we provide the first empirical evidence in support of that prediction.     

Temperature and foraging success of Great Tits Parus major hunting for spiders

The number of spiders caught by caged Great Tits Parus major in a 10 minute test increased in a sigmoid fashion with ambient temperature between 2 and 13°C. In control tests with immobile prey Calliphora pupae there was no significant effect of temperature. These results indicate that increasing activity of arthropod prey with temperature (Kacelnik 1979b) renders them more vulnerable to predators. We suggest that one reason why small temperate-zone birds such as Great Tits do not forage but sing when they get up in the morning is because low ambient temperatures reduce prey activity and hence the profitability of foraging.     

Variation in plumage colour of the Great tit Parus major in relation to habitat, season and food

The plumage colour of nestling and adult Great tits Parus major major L. was studied at Trondheim (63 °N, 10 °E), Norway. Nestlings reared in deciduous woodland were yellower than those reared in coniferous woodland, those reared in 1983 were yellower than those in 1982, and the later in the breeding season they were reared the more yellow they were. Furthermore, adults were yellower than nestlings, and for both groups the males were yellower than the females. Experimental interchange of eggs between nests showed that the variation in plumage colour observed between broods was related primarily to the food and not to genetic differences. Sampling, by the neck-collar method, of the food brought by the parent birds showed that the yellower nestlings received a greater proportion of lepidopteran larvae, in particular greenish-coloured ones, than did the nestlings with a paler plumage. A speculation is whether or not the variation in plumage colour has some adaptive value (viz. background matching).
In avian systematics, plumage coloration has been used to distinguish and describe subspecies. The present study shows that such a classification should only be made with certain reservations, until possible variation in the food quality of the subspecies is known to have no influence on plumage coloration.     

The timing of breeding and offspring size in Great Tits Parus major

The relationship between the timing of breeding, offspring weight and offspring recruitment was investigated in two populations of Great Tits Parus major. In two of six seasons a significant negative correlation was found between laying date and fledgling weight, in three seasons the opposite was true and in one season no significant correlation existed between these variables. Offspring recruitment was quantified in five seasons, and late-nesting birds were found to produce significantly fewer recruiting offspring in two of these, the same two seasons when late fledglings had significantly lower weights. No significant correlation existed between the number of recruits and laying date in the three seasons when late fledglings were as heavy as or heavier than early fledglings. The potential detriment to reproductive success of lateness thus was offset when late-nesting parents managed to produce heavy young. Between-year variations in the seasonal pattern of reproductive success has previously been suggested to contribute to the maintenance of variability in the timing of breeding in Great Tits. In this paper we suggest that producing heavy offspring late in the season might be a proximate factor influencing the seasonal patterns of reproductive success. Producing heavy offspring late in the season is probably not a "strategy" but rather an effect of chance differences between years in the seasonal availability of food or the seasonality of other factors important for the nesting birds.     

Plumage coloration can be perceived as a multiple condition‐dependent signal by Great Tits Parus major

The evolution of multiple signals can be explained because they enhance the perception of a general message by recipients. Plumage coloration frequently acts as a condition‐dependent signal, so that species displaying different colour patches have the potential to transmit information on condition through a multiple signalling system. The Great Tit Parus major exhibits plumage colour patches generated by the main types of colour production, some of which, particularly those based on melanins and carotenoids, are known to be related to body condition. However, the colour expressions of all of the different colour types have never been investigated simultaneously to determine whether they function as multiple signals of condition. In addition, visual perception models have never been applied to a multiple signalling system in a wild population of birds. Here I present information that links body condition with the colour expression of almost all of the different plumage patches of male Great Tits captured during the winter. Birds in better condition had greater reflectance values at short wavelengths in all plumage patches, and this was especially so in the white (i.e. structural) colour of the cheeks. Plumage colour characteristics were calculated by means of avian visual models, suggesting that Great Tits have the capacity to perceive information contained in the plumage coloration of conspecifics. These results show that short‐wavelength reflectance has great potential to transmit biologically significant information on the body condition of birds, even in achromatic plumage patches.     

Extra-pair paternity among Great Tits Parus major following manipulation of male signals

Female Great Tits are known to eavesdrop on the singing behaviour of males. It is unknown, however, whether manipulation of these signals is sufficient to influence extra-pair copulations, or whether such potentially costly reproductive decisions are unaffected by altering short-term signals of condition. Using interactive playbacks, we systematically engaged males in territorial contests in which we could control whether the focal male won or lost the interaction, regardless of the males' true potential. We then determined the levels and patterns of extra-pair paternity among experimental and neighbouring pairs using DNA microsatellite analysis. Extra-pair young were found in equal frequency among the nests of males allowed to win interactions as among those who lost interactions. However, cuckolded males were significantly less variable in allele sizes across the five microsatellite alleles tested than males who were not cuckolded. As measures of genetic variability are increasingly being found to correlate with individual fitness, this may suggest that females are attentive to underlying measures of condition when making extra-pair decisions. Short-term modification of the males' perceived quality may be insufficient to cause females to alter extra-pair decisions.     

Car traffic along hedgerows affects breeding success of Great Tits Parus major

Capsule?In hedgerows near roads with fast and frequent traffic, the mortality of Great Tit Parus major broods was higher than in hedgerows with less traffic and hedgerows with no disturbance.     

Dominance and anti-predator behaviour of Great Tits Parus major: a field study

During the winter of 1981–82 the hierarchical system of free-living Great Tits Parus major that visited a feeder supplied with sunflower seeds was studied in relation to the appearance of a predator. Without a predator, dominant birds were more successful in feeding and had to wait less before feeding. A significant negative correlation was found between dominance and the sequence of initial arrival.
After a Sparrowhawk Accipiter nisus had flown over the feeder, a correlation was found between the birds' dominance and the sequence of their return to the feeder. This field study supports the results of an analogous investigation on anti-predator behaviour and dominance in captive Blue Tits P. caeruleus.     

Environmental drivers of roosting body mass variation in boreal Great Tits Parus major

Small wintering passerines adaptively modulate daily body mass acquisition as part of their energy management policy. However, whether birds optimize overnight mass loss or body mass at dawn remains poorly understood. We studied environmental correlates of individual variation in body mass at dusk, overnight mass loss and body mass at dawn in a wild population of Great Tits Parus major wintering in northern Fennoscandia. Body mass at dusk, overnight mass loss and body mass at dawn were independent of prevailing conditions despite extremely low night ambient temperatures. Body mass at dusk was higher in males than in females, and decreased throughout winter and when snowfall was higher in the previous month. Overnight mass loss increased with precipitation during the previous week and tended to be higher in mid‐winter, when nights were longest. However, birds reduced overnight mass loss with higher temperatures in the previous week and higher precipitation in the previous 2 weeks. Dawn body mass was strongly correlated with dusk body mass and overnight mass loss, and showed only mild associations with weather variables once dusk mass was accounted for. Body mass in roosting boreal Great Tits seems to be constrained by recent snowfall as the winter progresses, but otherwise appears to be mostly unaffected by previous and current temperatures, suggesting a regular use of facultative hypothermia.     

Activity patterns of parent Great Tits Parus major feeding their young during rainfall

The influence of rainfall on the foraging patterns of Great Tit Parus major parents while feeding chicks at the nest was investigated using automated nest monitoring with electronic balances and photography. Great Tit females significantly reduced their visit rate to the nest during all rain intensities, while male feeding frequency did not significantly change. The female response was probably due to increased brooding requirements of young since the reduction in visit rate was most apparent at early nestling stages. At this time the chicks are incapable of thermoregulation and females significantly increased their nestbox occupancy time during rain. There was no indication that parents were compensating for periods of female inactivity during rainfall: there was no significant increase in visit rate following rainfall and no significant increase in prey size delivered to the nest during periods of rain. An analysis of data from six consecutive years revealed that the proportion of wet hours within the first week of the nestling period significantly influenced fledging weight in this species.     

The effect of old age on the reproduction of Great Tits Parus major and Blue Tits P. caeruleus

The results from two data sets show that in Great Tits Parus major and Blue Tits P. caeruleus reproduction is influenced by age. In both species, laying dates become earlier while clutch- and brood-size increase between the ages of 1 and 2, but there is no change in nesting success or post-fledging juvenile survival. Great Tits aged 5 or older are 'old' in that laying starts later and nesting success, brood-size and post-fledging juvenile survival decrease. Blue Tits become 'old' 1 year earlier than Great Tits: females aged 4 or older lay later, have a lower nesting success and smaller brood-size, and their young show a decreased post-fledging survival. The effect on laying date, in the Blue Tit, becomes apparent only for females aged 6 and older. It is concluded that ageing in small passerines is to be expected more generally, but that no extrapolations can be made as to the timing of the effect from one species to another.