Decision-Making in the Wild: Urgency and Complexity Drive Feeding Decision Speed and the Likelihood of Revising a Choice in a Sex-Dependent Manner in Great Tit (Parus major) Parents.

The fat reserves of small birds are built up daily as insurance against starvation. They are believed to reflect a trade-off between the risks of starvation and predation such that in situations of high predation risk birds are expected either to reduce their fat reserves in response to mass-dependent predation risk or to increase them in response to foraging interruptions. We assessed the effect on fat reserves of experimentally altering the perceived (but not the actual) risk of predation of wild great tits at a winter feeding site. The perceived predation risk was alternated between 'safe' and 'risky'. Increasing the perceived risk of predation involved 'swooping' a model sparrowhawk over the feeder at four unpredictable times each day using a remote mechanism We produce evidence that the experiment was suceessfull in altering the perceived risk of predation. As predicted from the hypothesis of mass-dependent predation risk, great tits (Parus major) carried significantly reduced fat reserves during the 'risky' treatment. Furthermore, dominant individuals were able to reduce their reserves more than subordinates. As birds returned to feeders within seconds after a predator 'attack', the reduction in fat reserves cannot be attributed to an interruption in feeding.

Dispersal is a major determinant of the dynamics and genetic structure of populations, and its consequences depend not only on average dispersal rates and distances, but also on the characteristics of dispersing and philopatric individuals. We investigated whether natal dispersal correlated with a predisposed behavioural trait: exploratory behaviour in novel environments. Wild great tits were caught in their natural habitat, tested the following morning in the laboratory using an open field test and released at the capture site. Natal dispersal correlated positively with parental and individual exploratory behaviour, using three independent datasets. First, fast-exploring parents had offspring that dispersed furthest. Second, immigrants were faster explorers than locally born birds. Third, post-fledging movements, comprising a major proportion of the variation in natal dispersal distances, were greater for fast females than for slow females. These findings suggest that parental behaviour influenced offspring natal dispersal either via parental behaviour per se (e.g. via post-fledging care) or by affecting the phenotype of their offspring (e.g. via their genes). Because this personality trait has a genetic basis, our results imply that genotypes differ in their dispersal distances. Therefore, the described patterns have profound consequences for the genetic composition of populations.

Theoretical models of host–parasite coevolution assume a partially genetic basis to the variability in susceptibility to parasites among hosts, for instance as a result of genetic variation in immune function. However, few empirical data exist for free–living vertebrate hosts to support this presumption. In a cross–fostering experiment with nestling great tits, by comparing nestlings of the same origin we investigated (i) the variance in host resistance against an ectoparasite due to a common genetic origin, (ii) the effect of ectoparasite infestation on cell–mediated immunity and (iii) the variance in cell–mediated immunity due to a common genetic origin. Ectoparasitic hen fleas can impair the growth of nestling great tits and nestling growth was therefore taken as a measure of host susceptibility. A common origin did not account for a significant part of the variation in host susceptibility to fleas. There was no significant overall effect of fleas on nestling growth or cell–mediated immunity, as assessed by a cutaneous hypersensitivity response. A common rearing environment explained a significant part of the variation in cell–mediated immunity among nestlings, mainly through its effect on nestling body mass. The variation in cell–mediated immunity was also related to a common origin. However, the origin–related variation in body mass did not account for the origin–related differences in cell–mediated immunity. The results of the present study thus suggest heritable variation in cell–mediated immunity among nestling great tits.

IntroductionAvian poxvirus infections are widespread in the domestic poultry population but are also reported in wild birds. In poultry, these infections cause significant economic losses, while wild birds may be a reservoir for poxvirus which affects breeding poultry. However, wild birds may also exhibit characteristic anatomopathological changes. This study concerns the infection of wild-living great tits (Parus major) with the avian poxvirus in Poland.Material and MethodsSamples of internal organs and skin collected from great tits were homogenised and total cellular DNA was isolated. In PCR, the primers complementary to gene encoding the core protein 4b of the HP44 strain of fowl poxvirus (FPV) were used.ResultsAfter electrophoresis in 2% agarose gel, the PCR product of 578 bp characteristic for FPV was obtained in DNA samples isolated from skin lesions and the heart. The analysis of the nucleotide sequence of the virus strain showed 99% similarity to many poxviruses previously isolated from great tits and other free birds at various sites in the world.ConclusionsThis paper is the first clinically documented evidence obtained in laboratory conditions of avian poxvirus cases in great tits in Poland.

To date, song research has focused primarily on the interactions of conspecifics. However, frequent interactions of songbirds with heterospecifics may necessitate adequate communication outside the species boundary. In this study, we focus on heterospecific communication behaviour of two small sympatric congeneric passerines, great and blue tits (Parus major and Parus caeruleus), which breed in overlapping territories and compete for food and nesting cavities. By means of a first playback experiment, we show that (1) heterospecific matching (imitating songs of the other species) is a strategy frequently used by great tits but not by blue tits, (2) both blue tit trilled and untrilled song can be accurately matched by great tits and that (3) almost half of the great tits in our study population match at least one blue tit song across all studied breeding stages, indicating that this heterospecific matching behaviour is a common feature in this population. A second playback experiment showed that these great tit imitations of blue tit songs do not function in intraspecific communication between male great tits. Hence, these heterospecific imitations appear to be designed for interspecific communication with blue tits. These findings suggest a strong heterospecific influence on the vocal learning process, repertoire composition and repertoire use in great tits and provide a possible mechanism that can drive song convergence in songbirds.

Great and blue tit feathers as biomonitors for heavy metal pollution

Great and Blue Tits as Indicators of Heavy Metal Contamination in Terrestrial Ecosystems

We tested the hypothesis that the weight lost by female Great and Blue Tits Parus major and P. caeruleus while raising their first brood influences their ability to start a second brood. The evening weight of female parents was recorded when the nestlings were 5 and 13 days old, in different years and habitats. Several predictions were tested: (1) both species lose weight while raising nestlings and Great Tit females which start a second brood lose less weight than females which do not; (2) differences in the average weight lost between years and areas correlate with differences in the proportion of second broods; (3) the relative weight loss in Blue Tits, which only rarely undertake second broods, is higher than in Great Tits in which second broods are more common. Other factors also are related to the probability of undertaking a second brood: more second broods are undertaken by more successful females, adult females and females that lay earlier.The comparison of Great and Blue Tits suggests that the two species use different reproductive strategies.

Maternal effects, in which differences in parental state cause differences in offspring fitness, are important in trade-offs influencing an individual's optimal reproductive strategy. In zebra finches (Taeniopygia guttata) we manipulated the nutritional state for four weeks before the start of breeding through protein supplementation. Zebra finches were kept on identical diets during the rest of the experiment. We then tested the effects of maternal state on offspring size, survival and fecundity. In order to separate the effects of maternal state occurring through egg production, incubation and chick-rearing, we used a cross-fostering experiment. We show that a protein-rich diet prior to laying improved maternal body weight prior to breeding compared with birds on a protein-poor diet. Poorer maternal state prior to breeding gave rise to offspring with lower fecundity than offspring from birds in a better nutritional state. Maternal state is thought to affect the conditions developing offspring experience through the bird's ability to produce and incubate eggs. Male and female embryos differed in their responses to conditions at different developmental stages. This shows that embryonic developmental conditions and sex differences in vulnerability to these conditions need to be incorporated into future models of selection, life-history evolution and sex-ratio theory.

BackgroundIn egg-laying animals, mothers can influence the developmental environment and thus the phenotype of their offspring by secreting various substances into the egg yolk. In birds, recent studies have demonstrated that different yolk substances can interactively affect offspring phenotype, but the implications of such effects for offspring fitness and phenotype in natural populations have remained unclear. We measured natural variation in the content of 31 yolk components known to shape offspring phenotypes including steroid hormones, antioxidants and fatty acids in eggs of free-living great tits (Parus major) during two breeding seasons. We tested for relationships between yolk component groupings and offspring fitness and phenotypes.ResultsVariation in hatchling and fledgling numbers was primarily explained by yolk fatty acids (including saturated, mono- and polyunsaturated fatty acids) - but not by androgen hormones and carotenoids, components previously considered to be major determinants of offspring phenotype. Fatty acids were also better predictors of variation in nestling oxidative status and size than androgens and carotenoids.ConclusionsOur results suggest that fatty acids are important yolk substances that contribute to shaping offspring fitness and phenotype in free-living populations. Since polyunsaturated fatty acids cannot be produced de novo by the mother, but have to be obtained from the diet, these findings highlight potential mechanisms (e.g., weather, habitat quality, foraging ability) through which environmental variation may shape maternal effects and consequences for offspring. Our study represents an important first step towards unraveling interactive effects of multiple yolk substances on offspring fitness and phenotypes in free-living populations. It provides the basis for future experiments that will establish the pathways by which yolk components, singly and/or interactively, mediate maternal effects in natural populations.

Temperature-induced variation in yolk androgen and thyroid hormone levels in avian eggs

Urbanization and climate change are two environmental factors that have most prominently affected breeding phenology of birds during recent decades. We examined such relationships in rural, suburban and urban nest box populations of Great Tits Parus major and Blue Tits P. caeruleus in the capital region of Finland in the 1980s and 1990s. We expected that mild winters and high spring temperatures may advance the breeding season of tits, but less so in urban habitats, where breeding should in any case start earlier than elsewhere. On average, Blue Tits began egg laying a few days earlier in spring than Great Tits. Contrary to expectations, tits bred later in the urban parks of the city than in other habitats, whereas breeding was earliest in suburban areas. It seems that these intermediate habitats, in some way, offer the advantages of both rural and urban habitats. During the study period, the timing of breeding in tits showed advancing temporal trends in rural habitats, and in the Blue Tit also in urban habitats. The effect of increasing winter temperatures on laying dates was mainly minor, but a significant delay emerged in urban Great Tits. The main effect of increasing April temperatures on laying dates was a significant advancement. In urban habitats, however, the advancing effect was in Great Tits significantly stronger and in Blue Tits significantly weaker than in other habitats. The results suggest that breeding of tits may advance also with warming climate but some urban populations might be more resistant to climate change than rural ones.

Understanding the selective pressures shaping the number of offspring per breeding event is a key area in the study of life-history strategies. However, in species with parental care, costs incurred in offspring production, rather than rearing, have been largely ignored in both theoretical and empirical studies until relatively recently. Furthermore, the few experimental studies that have manipulated production costs have not yet teased apart effects that operate via the parental phenotype from effects on the quality of the resulting young. To examine whether increased egg production influences parental brood rearing capacity independently of effects operating via egg quality, we experimentally increased egg production in gulls and then examined their capacity to rear a control clutch. We found that the capacity of parents to rear the control brood was substantially reduced solely as a consequence of having themselves produced one extra egg. The paradox that, in many species, parents apparently aim for fewer young per breeding event than the experimentally and theoretically demonstrated optimum, has partly arisen from the failure to take into account the constraints imposed by production costs.

Why do people feed wild birds? It is intuitively clear that humans feel delighted by the presence of birds that they can manage to attract by offering ordinary seeds at a feeder. According to surveys on human motivations for feeding wild birds in Australia, some people may gain experiential knowledge from observing the birds in their gardens, or feed the birds in return for the massive habitat destruction caused by humans ( Howard & Jones, 2004 ; Ishigame & Baxter, 2007 ). However, outside Australia, the largely unstudied motivations of people feeding birds are just one example for our gaps of knowledge with respect to bird feeding ( Jones & Reynolds, 2008 ). It is in sharp contrast to our limited knowledge on the habit that wild bird feeding is probably the most widespread and popular form of human–wildlife interaction throughout the world ( Jones, 2011 ), and, at least in northern temperate regions, the largest wildlife management activity ( Martinson & Flaspohler, 2003 ). In the UK, suffi cient commercial wild bird foods are sold to support a hypothetical number of over 30 million great tits ( Parus major ; Robb et al., 2008a ), which is many more than the 2 million pairs of great tits that are actually present (www.bto.org). Surveys have found that 64% of households provide supplementary food for birds in the UK ( Davies et al., 2012 ), and 43% in the USA ( Martinson & Flaspohler, 2003 ). In suburban and rural environments of Australia, estimated household feeding rates range from 36% to 48% ( Ishigame & Baxter, 2007 ). Although BirdLife Australia does not encourage supplementary feeding of wild birds ( Bird Observation and Conservation Australia, 2010 ), northern organizations such as the British Trust for Ornithology (BTO), the Royal Society for the Protection of Birds (RSPB) and the Cornell Laboratory of Ornithology now recommend feeding birds for promoting nature conservation ( Jones, 2011 ). In Germany, a renowned ornithologist recently wrote a popular book on bird feeding that sold 50,000 copies in the fi rst 1.5 years, in which he advocates feeding the birds year-round and on a massive scale ( Berthold & Mohr, 2008 ). Indeed, wild bird feeding is not only here to stay ( Jones, 2011 ), it also seems to be increasing, at least in the UK ( Chamberlain et al., 2005 ). Today, in many countries, the huge effort in providing supplementary food for birds may be one of the largest human infl uences on bird populations, in addition to habitat loss and change, humaninduced climate change and hunting. As I will show in this chapter, we now have a sound basis of knowledge about the impacts that supplementary feeding can have on birds. However, as I will also show, our knowledge mostly comes from smallscale experiments that researchers did in natural and rural habitats. Surprisingly little is known on the impact of feeding birds in our urban gardens and backyards. However, humans are infl uencing urban food supply for birds not only directly by providing feeders, but also via waste treatment and by creating, changing, or destroying urban or natural habitats and food sources in our cities ( Chace & Walsh, 2006 ). Because such additional human infl uences are usually weaker in rural landscapes, the effects of feeding wild birds are likely

Mycoplasma spp. are important pathogens in poultry and cause high economic losses for poultry industry worldwide. In other bird species (e.g. white storks, birds of prey, and several waterfowl species), Mycoplasma spp. are regularly found in healthy individuals, hence, considered apathogenic or part of the microbiota of the upper respiratory tract. However, as Mycoplasma spp. are absent in healthy individuals of some wild bird species, they might play a role as respiratory pathogen in these bird species, e.g. Mycoplasma gallisepticum in house finches. The knowledge on the occurrence of Mycoplasma spp. in wild birds is limited. To evaluate the relevance of Mycoplasma spp. in free-ranging nightingales and tits, 172 wild caught birds were screened for the presence of mycoplasmas. The birds were sampled via choanal swabs and examined via molecular methods (n = 172) and, when possible, via culture (n = 142). The Mycoplasma sp. was determined by sequencing the 16S rRNA gene and 16S-23S Intergenic Transcribed Spacer Region. All birds were tested negative for mycoplasmas via PCR and/or mycoplasmal culture. Hence, free-ranging nightingales and tits do not show any mycoplasma in their microbial flora of the respiratory tract. Therefore, these songbird species may suffer from clinical mycoplasmosis when being infected. We hypothesize that birds relying on their vocal ability for reproduction have excluded mycoplasmas from their respiratory flora compared to other bird species.