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Major histocompatibility complex (Mhc) genes are believed to play a key role in the genetic basis of disease control. Although numerous studies have sought links between Mhc and disease prevalence, many have ignored the ecological and epidemiological aspects of the host–parasite interaction. Consequently, interpreting associations between prevalence and Mhc has been difficult, whereas discriminating alleles for qualitative resistance, quantitative resistance and susceptibility remains challenging. Moreover, most studies to date have quantified associations between genotypes and disease status, overlooking the complex relationship between genotype and the properties of the Mhc molecule that interacts with parasites. Here, we address these problems and demonstrate avian malaria (Plasmodium) parasite species-specific associations with functional properties of Mhc molecules (Mhc supertypes) in a wild great tit (Parus major) population. We further show that correctly interpreting these associations depends crucially on understanding the spatial variation in risk of infection and the fitness effects of infection. We report that a single Mhc supertype confers qualitative resistance to Plasmodium relictum, whereas a different Mhc supertype confers quantitative resistance to Plasmodium circumflexum infections. Furthermore, we demonstrate common functional properties of Plasmodium-resistance alleles in passerine birds, suggesting this is a model system for parasite–Mhc associations in the wild.

One consequence of climate change is an increasing mismatch between timing of food requirements and food availability. Such a mismatch is primarily expected in avian long-distance migrants because of their complex annual cycle, and in habitats with a seasonal food peak. Here we show that insectivorous long-distance migrant species in The Netherlands declined strongly (1984–2004) in forests, a habitat characterized by a short spring food peak, but that they did not decline in less seasonal marshes. Also, within generalist long-distance migrant species, populations declined more strongly in forests than in marshes. Forest-inhabiting migrant species arriving latest in spring declined most sharply, probably because their mismatch with the peak in food supply is greatest. Residents and short-distance migrants had non-declining populations in both habitats, suggesting that habitat quality did not deteriorate. Habitat-related differences in trends were most probably caused by climate change because at a European scale, long-distance migrants in forests declined more severely in western Europe, where springs have become considerably warmer, when compared with northern Europe, where temperatures during spring arrival and breeding have increased less. Our results suggest that trophic mismatches may have become a major cause for population declines in long-distance migrants in highly seasonal habitats.

The hatching success of a bird's egg is one of the key determinants of avian reproductive success, which may be compromised by microbial infections causing embryonic death. During incubation, outer eggshell bacterial communities pose a constant threat of pathogen translocation and embryo infection. One of the parental strategies to mitigate this threat is the incorporation of maternal immune factors into the egg albumen and yolk. It has been suggested that habitat changes like forest fragmentation can affect environmental factors and life-history traits that are linked to egg contamination. This study aims at investigating relationships between microbial pressure, immune investment and hatching success in two abundant forest bird species and analyzing to what extent these are driven by extrinsic (environmental) factors. We here compared (1) the bacterial load and composition on eggshells, (2) the level of immune defenses in eggs, and (3) the reproductive success between great (Parus major) and blue (Cyanistes caeruleus) tits in Belgium and examined if forest fragmentation affects these parameters. Analysis of 70 great tit and 34 blue tit eggshells revealed a similar microbiota composition (Enterobacteriaceae, Lactobacillus spp., Firmicutes and Bacteroidetes), but higher bacterial loads in great tits. Forest fragmentation was not identified as an important explanatory variable. Although a significant negative correlation between hatching success and bacterial load on the eggshells in great tits corroborates microbial pressure to be a driver of embryonic mortality, the overall hatching success was only marginally lower than in blue tits. This may be explained by the significantly higher levels of lysozyme and IgY in the eggs of great tits, protecting the embryo from increased infection pressure. Our results show that immune investment in eggs is suggested to be a species-specific adaptive trait that serves to protect hatchlings from pathogen pressure, which is not directly linked to habitat fragmentation.

The hatching success of a bird's egg is one of the key determinants of avian reproductive success, which may be compromised by microbial infections causing embryonic death. During incubation, outer eggshell bacterial communities pose a constant threat of pathogen translocation and embryo infection. One of the parental strategies to mitigate this threat is the incorporation of maternal immune factors into the egg albumen and yolk. It has been suggested that habitat changes like forest fragmentation can affect environmental factors and life-history traits that are linked to egg contamination. This study aims at investigating relationships between microbial pressure, immune investment and hatching success in two abundant forest bird species and analyzing to what extent these are driven by extrinsic (environmental) factors. We here compared (1) the bacterial load and composition on eggshells, (2) the level of immune defenses in eggs, and (3) the reproductive success between great (Parus major) and blue (Cyanistes caeruleus) tits in Belgium and examined if forest fragmentation affects these parameters. Analysis of 70 great tit and 34 blue tit eggshells revealed a similar microbiota composition (Enterobacteriaceae, Lactobacillus spp., Firmicutes and Bacteroidetes), but higher bacterial loads in great tits. Forest fragmentation was not identified as an important explanatory variable. Although a significant negative correlation between hatching success and bacterial load on the eggshells in great tits corroborates microbial pressure to be a driver of embryonic mortality, the overall hatching success was only marginally lower than in blue tits. This may be explained by the significantly higher levels of lysozyme and IgY in the eggs of great tits, protecting the embryo from increased infection pressure. Our results show that immune investment in eggs is suggested to be a species-specific adaptive trait that serves to protect hatchlings from pathogen pressure, which is not directly linked to habitat fragmentation.

Climate change has led to phenological shifts in many species, but with large variation in magnitude among species and trophic levels. The poster child example of the resulting phenological mismatches between the phenology of predators and their prey is the great tit (Parus major), where this mismatch led to directional selection for earlier seasonal breeding. Natural climate variability can obscure the impacts of climate change over certain periods, weakening phenological mismatching and selection. Here, we show that selection on seasonal timing indeed weakened significantly over the past two decades as increases in late spring temperatures have slowed down. Consequently, there has been no further advancement in the date of peak caterpillar food abundance, while great tit phenology has continued to advance, thereby weakening the phenological mismatch. We thus show that the relationships between temperature, phenologies of prey and predator, and selection on predator phenology are robust, also in times of a slowdown of warming. Using projected temperatures from a large ensemble of climate simulations that take natural climate variability into account, we show that prey phenology is again projected to advance faster than great tit phenology in the coming decades, and therefore that long-term global warming will intensify phenological mismatches.

Social network analysis has become a popular tool for characterising the social structure of populations. Animal social networks can be built either by observing individuals and defining links based on the occurrence of specific types of social interactions, or by linking individuals based on observations of physical proximity or group membership, given a certain behavioural activity. The latter approaches of discovering network structure require splitting the temporal observation stream into discrete events given an appropriate time resolution parameter. This process poses several non-trivial problems which have not received adequate attention so far. Here, using data from a study of passive integrated transponder (PIT)-tagged great tits Parus major, we discuss these problems, demonstrate how the choice of the extraction method and the temporal resolution parameter influence the appearance and properties of the retrieved network and suggest a modus operandi that minimises observer bias due to arbitrary parameter choice. Our results have important implications for all studies of social networks where associations are based on spatio-temporal proximity, and more generally for all studies where we seek to uncover the relationships amongst a population of individuals that are observed through a temporal data stream of appearance records.

Colours are commonly used as visual cues when measuring animals’ cognitive abilities. However, animals can have innate biases towards certain colours that depend on ecological and evolutionary contexts, therefore potentially influencing their performance in experiments. For example, when foraging, the colour red can advertise profitable fruits or act as a warning signal about chemically defended prey, and an individual’s propensity to take food of that colour may depend on experience, age or physical condition. Here, we investigate how these contexts influence blue tits’ (Cyanistes caeruleus) and great tits’ (Parus major) responses to red-coloured almond flakes. We found that juvenile birds preferred red both when it was presented simultaneously with green, and when it was presented with three alternative colours (orange, purple, green). Adult birds, however, only preferred red after a positive experience with the colour, or when it was presented with the three alternative colours. We then tested whether colour influenced avoidance learning about food unpalatability. Despite the prediction that red is a more salient warning signal than green, we found only weak evidence that birds discriminated red unpalatable almonds from a green palatable alternative more quickly than when the colours were reversed. Our results suggest that biases towards red food may depend on birds’ age and previous experience, and this might influence their performance in experiments that use red stimuli. Considering the ecological relevance of colours is, therefore, important when designing experiments that involve colour cues.

Changes in the timing of life‐history events (phenology) are a widespread consequence of climate change. Predicting population resilience requires knowledge of how phenology is likely to change over time, which can be gained by identifying the specific environmental cues that drive phenological events. Cue identification is often achieved with statistical testing of candidate cues. As the number of methods used to generate predictions increases, assessing the predictive accuracy of different approaches has become necessary. This study aims to (a) provide an empirical illustration of the predictive ability of five commonly applied statistical methods for cue identification (absolute and relative sliding time window analyses, penalized signal regression, climate sensitivity profiles and a growing degree‐day model) and (b) discuss approaches for implementing cue identification methods in different systems. Using a dataset of mean clutch initiation timing in wild great tits (Parus major), we explored how the days of the year identified as most important, and the aggregate statistic identified as a cue, differed between statistical methods and with respect to the time span of data used. Each method's predictive capacity was tested using cross‐validation and assessed for robustness to varying sample size. We show that the identified critical time window of cue sensitivity was consistent across four of the five methods. The accuracy and precision of predictions differed by method with penalized signal regression resulting in the most accurate and most precise predictions in our case. Accuracy was maximal for near‐future predictions and showed a relationship with time. The difference between predictions and observations systematically shifted across the study from preceding observations to lagging. This temporal trend in prediction error suggests that the current statistical tools either fail to capture a key component of the cue–phenology relationship, or the relationship itself is changing through time in our system. These two influences need to be teased apart if we are to generate realistic predictions of phenological change. We recommend future phenological studies to challenge the idea of a static cue–phenology relationship and should cross‐validate results across multiple time periods. The authors give a case study of the predictive performance of five different phenological cue identification methods. They identify a temporal bias in prediction error, suggesting that the current statistical tools either fail to capture a key component of the cue–phenology relationship, or the relationship itself is changing through time. Photo credit: Emily G. Simmonds.

Birds that are foraging in tree canopies can cause a substantial decrease in arthropod numbers. Trees may benefit from avian insectivores attacking insect herbivores. In a field study, we tested whether the intensity of bird predation on caterpillars is linked quantitatively to leaf damage caused by insect herbivores, a hypothesized relationship that previously was poorly investigated. Artificial caterpillars were placed in the lower part of oak trees (Quercus robur) in urban and suburban sites across the city of Gothenburg, Sweden. Two days later, we recorded the survival: the pooled predation rate was 11.5% (5.7% day-1). Mean predation rate per tree was 10.4%. Mean leaf damage, i. e. leaf area eaten by insect herbivores, per tree was 5.7% but there was large variation between trees. We found a significant negative relationship between survival probability of caterpillars and leaf damage in an analysis using a mixed model logistic regression. This suggests that caterpillars are at high risk of bird attacks in trees with a high degree of leaf damage and avian insectivores may increase the foraging effort in the foliage of such oak trees. Our findings concerning the quantitative relationship between the predator–prey interactions and plant damage suggested tentatively that the survival probability of caterpillars decreases rapidly at 15–20 % leaf damage in lower part of oak canopies. Furthermore, our findings add credence to the idea of using artificial caterpillars as a means to obtain standardized comparisons of predation rates in various habitats.

Habitat quality has direct effects on the evolutionary fitness of breeding organisms, which is why it is believed that animals tend to have an evolved preference for the best possible habitats. However, some animals may mistakenly choose to reproduce in habitats that decrease their fitness, resulting in ‘ecological traps’. In this study, we tested whether great tits (Parus major) attracted to areas affected by outbreaks of the great web-spinning sawfly (Acantholyda posticalis) had fitness detriments characteristic of ecological traps. Sawfly larvae consume pine needles, which decreases resource availability for birds co-habiting the forest. Using artificial nesting sites, we found that great tits inhabiting areas of sawfly outbreaks had similar clutch sizes as tits breeding in healthy forest patches; however, the fledgling number was significantly lower, and fledgling condition was worse in the damaged forests. While moth larvae are the most important food for bird nestlings, the forest patches damaged by sawflies had lower larval biomass. Although most ecological traps occur in environments altered by humans, this study shows that pest insects can lower habitat quality, forming ecological traps. Our results indicate that attracting cavity-nesting birds should be done with caution because it may negatively impact birds’nutritional status and reproductive fitness.