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In cooperatively breeding species, group members may derive multiple benefits from helping to raise other individuals' offspring, yet not all individuals do so. In this study, we tested predictions from the "kin selection", "pay-to-stay", "group augmentation" and "skills" hypotheses, to explain why group members feed nestlings of breeding placid greenbuls (Phyllastrephus placidus). In our study population, about 70% of the breeding pairs were accompanied by subordinates, and in 60% of these cases at least one subordinate helped in provisioning nestlings. In total, 80% of the subordinates were related to one or both breeders. In accordance with the "kin selection" hypothesis, and contrary to the "pay-to-stay" hypothesis, all the helpers were first-order kin of the breeding female (although relatedness to the breeding male did not explain variation in helping) and the presence of helpers was associated with increased survival of the breeding pair. However, the propensity to help varied among group members, as 46% of group members related to the breeding female did not feed nestlings. Younger helpers fed offspring more often than older ones, supporting the "skills" and "group augmentation" hypotheses. However, support for the "group augmentation" hypothesis was mixed since subordinate sex and group size did not explain additional variation in helping propensity and effort. We argue that in addition to indirect and direct benefits, also the costs of helping as well as other types of helping aside from provisioning must be considered to better understand variation in helping behavior. Relatedness is thought to be an important factor explaining helping at the nest by non-breeding group members. We show that placid greenbul helpers are always related to the breeding females, supporting this view. However, many related groups members never helped suggesting that other factors shape helping decision in this species.

Many studies have investigated how spring temperature affects laying dates and how this in turn affects the synchrony between nestling food demands and the insect food peak that follows tree budburst. While there is strong evidence that temperature itself acts as a cue for this plasticity in annual timing, the exact nature of the cue and response remains to be elucidated. Here we use long-term data on Great and Blue Tits and an unprecedented dataset on the location and phenology of 1396 trees to investigate whether small-scale variation in laying date can be explained by local tree phenology, and/or by tree species composition around nestboxes. Individual trees maintained their relative timing of budburst between years, while differences among tree species were more variable between springs. Contrary to expectation, we found no relation between first-egg dates and average budburst date at different distances around the nestbox. This can at least partly be explained by the very low degree of spatial autocorrelation in tree budburst. We did find an effect of local tree composition whereby Blue Tits, but not Great Tits, laid earlier in nestboxes surrounded by more oaks and fewer beeches. Although Blue Tit nest failure rate was higher in territories with more beech trees, independently of laying date, we found no evidence for preferential occupation of oak-dominated territories. Thus although we found some evidence for fine-scale effects of tree species on timing of breeding, the underlying mechanism remains unclear.

Urbanization has been shown to strongly affect community composition of various taxa with potentially strong shifts in ecological interactions, including those between hosts and parasites. We investigated the effect of urbanization on the composition of arthropods in nests of great tits in Flanders, Belgium. These nests contain taxonomically and functionally diverse arthropod communities including parasites, predators, detritivores and accidental commensals. Using a standardized hierarchical sampling design with subplots (200 m x 200 m) nested in plots (3 km x 3 km) of varying urbanization levels, we collected arthropods from nests of resident great tits after the young had fledged. Arthropods were extracted, identified to Primary Taxonomical Groups (PTG) and counted. Using generalized linear mixed models (GLMMs) we found diverging effects of urbanization on PTG occurrences and abundances at various levels, but we did not find an overall signal in arthropod diversity or richness. Also, visual inspection of non-metric multidimensional scaling (NMDS) plots did not reveal any community differences between urbanization levels at plot or subplot scales. Land use and environmental variables at different distances around nestboxes did not contribute much to the variation between communities. Our results indicate that arthropod nestbox communities are generally not adversely affected by urbanization, and even city gardens and parks harbor comparable communities to forests and suburban areas. We thus found no evidence for a parasite release effect due to urbanization, nor an increased risk of parasitism in human-dominated environments.

The distribution of ticks in the Ixodes ricinus species complex is partly driven by climate, with temperature and relative humidity affecting survival. These variables are driven by macroclimate, but vary locally due to microclimate buffering. This buffering has been suggested to be one of the driving forces behind variation in tick survival and density in time and space. In order to understand the role of the herb layer with respect to this variation, we deployed I. ricinus within an existing experimental setup studying the response of forest under -storey to micrometeorological changes. This allowed for the analysis of both direct effects of warming on tick survival in controlled field conditions, as well as indirect effects through changes in herb layer biomass. Herb layer biomass estimates were observed to be higher in plots that had been experimentally warmed, with a trend towards higher survival in these warmed plots. This marginal increase in survival rate may be due to increased microclimate buffering. Comparing our results to literature implies that canopy and shrub layer vegetation have a larger effect on climate buffering, and therefore also on tick survival. Since the herb layer biomass is expected to increase due to global warming and increased frequency of disturbance-induced canopy gaps, survival in forested habitats may increase in the future. This would increase the difference in survival compared to that in open habitats.

Wild birds are frequently exposed to the zoonotic tick-borne bacteria Borrelia burgdorferi sensu lato (s.l.), and some bird species act as reservoirs for some Borrelia genospecies. Studying the tropism of Borrelia in the host, how it is sequestered in different organs, and whether it is maintained in circulation and/or in the host's skin is important to understand pathogenicity, infectivity to vector ticks and reservoir competency.We evaluated tissue dissemination of Borrelia in blackbirds (Turdus merula) and great tits (Parus major), naturally and experimentally infected with Borrelia genospecies from enzootic foci. We collected both minimally invasive biological samples (feathers, skin biopsies and blood) and skin, joint, brain and visceral tissues from necropsied birds. Infectiousness of the host was evaluated through xenodiagnoses and infection rates in fed and moulted ticks. Skin biopsies were the most reliable method for assessing avian hosts' Borrelia infectiousness, which was supported by the agreement of infection status results obtained from the analysis of chin and lore skin samples from necropsied birds and of their xenodiagnostic ticks, including a significant correlation between the estimated concentration of Borrelia genome copies in the skin and the Borrelia infection rate in the xenodiagnostic ticks. This confirms a dermatropism of Borrelia garinii, B. valaisiana and B. turdi in its avian hosts. However, time elapsed from exposure to Borrelia and interaction between host species and Borrelia genospecies may affect the reliability of skin biopsies. The blood was not useful to assess infectiousness of birds, even during the period of expected maximum spirochetaemia. From the tissues sampled (foot joint, liver, spleen, heart, kidney, gut and brain), Borrelia was detected only in the gut, which could be related with infection mode, genospecies competition, genospecies-specific seasonality and/or excretion processes. This study received financial support from Fundação para a Ciência e a Tecnologia by the strategic program of MARE (MARE - UID/MAR/04292/2019), the fellowship to Ana Cláudia Norte (SFRH/BPD/108197/2015) and from the Portuguese National Institute of Health Doutor Ricardo Jorge. Dieter Heylen is funded by the Marie Sklodowska-Curie Actions (EU-Horizon 2020, Individual Global Fellowship, project no. 799609), the Fund for Scientific Research – Flanders (FWO) and the Bill & Melinda Gates Foundation. info:eu-repo/semantics/publishedVersion

Individual variation in behaviour can contribute to the heterogeneous distribution of parasites among hosts for example by affecting the probability of encountering infective stages (larvae). Here, we investigated the relationship between host boldness/exploration tendency and gastro-intestinal helminth infection in invasive Eastern grey squirrels (Sciurus carolinensis). We used direct helminth counts, data rarely available in host-parasite studies that often used less reliable indirect estimates of infection. We predicted that bolder animals with a strong exploration tendency will have higher parasite load than shy, less explorative hosts. We examined two parameters of parasite infection: infection status and intensity of infection. Individual personality of 207 grey squirrels was assessed by capture-mark-recapture (CMR), calculating the trappability and trap diversity indices as estimates of boldness and exploration, respectively. Since both indices were strongly correlated, we used PCA to derive a single score (first component) which had a high value for bold, exploring animals. At the end of the study, 77 individuals were euthanized and gastro-intestinal helminths were identified and counted. Overall 73% of grey squirrels were infected by Strongyloides robustus with the intensity of infection varying from 1 to 86 worms (mean ± SE = 10.7 ± 2.1 helminths per host). We found that bolder, more explorative animals were more likely to be infected by S. robustus. However, host personality was not related to parasite intensity, which was instead positively associated with host body mass. Our results confirm that differences in personality-related host behaviour can influence the distribution of infections within host populations and stimulate further questions on whether such personality-parasite relationships may affect the invasion process.

Simple Summary Nocturnal light pollution is a worldwide growing problem, threatening nocturnal biodiversity. We studied the impact of streetlights on mating success of the female common glow-worm, a bioluminescent nocturnal beetle (Lampyris noctiluca L.) that uses light signals for sexual communication. We monitored individual females daily and assumed that when they stopped glowing, they had effectively mated. We found that females in dark surroundings typically stopped glowing after one night, while females in illuminated areas glowed for significantly more nights, in some cases up to 15 nights. Our study confirms previous hypotheses that females exposed to artificial light suffer from a reduced mate attraction success, which can lead to population declines. Our findings represent valuable information that can be used by policy makers and managers to conserve the iconic glow-worms. Abstract Nocturnal light pollution from anthropogenic origin is increasing worldwide and is recognised as a major threat for nocturnal biodiversity. We studied the impact of artificial light on the mate attraction success of female common glow-worms (Lampyris noctiluca L.) by daily monitoring their glowing status in the field, acting as a proxy for mating status throughout the mating season. We found that females in dark surroundings typically stopped glowing after one night, indicating that they had mated, while females in illuminated areas glowed for significantly more nights, in some cases up to 15 nights. Our study confirms previous findings and hypotheses that females exposed to artificial light suffer from a reduced mate attraction success with a negative impact on populations.

Background: Variation in parasite burdens among hosts is typically related to differences in adaptive immunity. Comprehension of underlying mechanisms is hence necessary to gain better insights into endemic transmission cycles. Here we investigate whether wild songbirds that have never been exposed to ticks develop adaptive humoral immunity against endemic Ixodes ricinus ticks. Methods: Blue tits were exposed three times in succession to wild Ixodes ricinus ticks. For each infestation, serum samples were obtained. An enzyme-linked immunosorbent assay was developed, using tick salivary antigens, in order to quantify the bird’s IgY response against ticks. In addition, at every sampling occasion the birds’ body weight (corrected for body size) and haematocrit level was determined. Results: Individual IgY levels against the ticks’ salivary proteins increased over three consecutive tick infestations, and large among-individual variation was observed. The responses were specifically directed against I. ricinus; cross-reactivity against the congeneric tree-hole tick Ixodes arboricola was negligibly low. IgY responses did not impinge on tick feeding success (engorgement weight and attachment success). Yet, those birds with the highest immune responses were more capable to reduce the acute harm (blood depletions) by compensating erythrocyte loss. Furthermore, at the end of the experiment, these birds had gained more body weight than birds with lower IgY levels. Conclusions: Latter observations can be considered as an effect of host quality and/or tolerance mechanisms. Birds anticipate the (future) costs of the activation of the immune system by ticks and/or ongoing tick-borne pathogen infections. Furthermore, although unsuccessful against tick feeding, the IgY responses may indirectly protect birds against tick-borne disease by acting against salivary protein secretions on which pathogens rely for transmission. Grant Information: This project has received funding from the European Union’s Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement No [799609], (project ECODIS). DH is also funded by the Fund for Scientific Research – Flanders (FWO). BB is funded by the European Community Action Scheme for the Mobility of University Students Programme. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Open Access

The pace-of-life syndrome (POLS) theory provides an evolutionary explanation for the existence of consistent among-individual variation in behaviour, or animal personality. Herein, individuals with a fast lifestyle are considered to be bolder and should take more risks resulting in a lower life expectancy compared to shyer individuals with a slower lifestyle. However, this assumption depends on the levels of intra-specific competition that the individuals experience which has rarely been tested in species that experience large changes in competition on a very short time scale. We used the multimammate mice (Mastomys natalensis) as a model system to study the POLS assumption by investigating the effects of two personality traits (exploration and stress-sensitivity) on survival, maturation (a proxy for reproductive investment) and recapture probability during one population cycle (N

The phenology of many species shows strong sensitivity to climate change; however, with few large scale intra-specific studies it is unclear how such sensitivity varies over a species’ range. We document large intra-specific variation in phenological sensitivity to temperature using laying date information from 67 populations of two European songbirds covering a large part of their breeding range. Populations inhabiting deciduous habitats showed stronger phenological sensitivity compared with those in evergreen and mixed habitats. Strikingly, however, the lowest sensitivity was seen in populations that had experienced the greatest change in climate. Therefore, we predict that the strongest phenological advancement will not occur in those populations with the highest sensitivity. Our results show that to effectively assess the impact of climate change on phenology across a species range it will be necessary to account for intra-specific variation in phenological sensitivity, climate change exposure, and the ecological characteristics of a population.

Consistent among-individual variation in behavior, or animal personality, is present in a wide variety of species. This behavioral variation is maintained by both genetic and environmental factors. Parental effects are a special case of environmental variation and are expected to evolve in populations experiencing large fluctuations in their environment. They represent a non-genetic pathway by which parents can transmit information to their offspring, by modulating their personality. While it is expected that parental effects contribute to the observed personality variation, this has rarely been studied in wild populations. We used the multimammate mouse Mastomys natalensis as a model system to investigate the potential effects of maternal personality on offspring behavior. We did this by repeatedly recording the behavior of individually housed juveniles which were born and raised in the lab from wild caught females. A linear correlation, between mother and offspring in behavior, would be expected when the personality is only affected by additive genetic variation, while a more complex relationship would suggests the presence of maternal effects. We found that the personality of the mother predicted the behavior of their offspring in a non-linear pattern. Exploration behavior of mother and offspring was positively correlated, but only for slow and average exploring mothers, while this correlation became negative for fast exploring mothers. This may suggests that early maternal effects could affect personality in juvenile M. natalensis, potentially due to density-dependent and negative frequency-dependent mechanisms, and therefore contribute to the maintenance of personality variation.

Human activities impact upon natural habitats used by birds for breeding and foraging, and lead to changes in the composition and spatial distribution of predator communities, mainly through loss, fragmentation and disturbance of formerly pristine habitat. Yet possible fitness consequences of such changes through impacts on bird nest‐site selection remain poorly known. Here we study nest‐site selection and reproductive success of Placid Greenbuls Phyllastrephus placidus in the Taita Hills, southeast Kenya. We show that habitat features associated with nest‐site selection by this insectivorous, open‐cup‐nesting bird species vary among forest fragments that are exposed to different levels of habitat disturbance. Such differences in sites selected for breeding result from a plastic response to fragment‐specific conditions or may be driven by fragment‐specific variation in the distribution and availability of certain habitat features. Given the overall high nest predation rates in our study area, we expected variation in nest‐site selection to correlate with reproductive success and nestling condition, but detected no such relationship. Because predator density and nest predation rates may vary strongly in space and time, a better understanding of spatio‐temporal variation in predator communities is needed to assess the possible adaptive value of nest‐site selection strategies for reducing the high predation rates that are typical for this and many other open‐cup‐nesting tropical passerines.

Humans fundamentally affect dispersal, directly by transporting individuals and indirectly by altering landscapes and natural vectors. This human-mediated dispersal (HMD) modifies long-distance dispersal, changes dispersal paths, and overall benefits certain species or genotypes while disadvantaging others. HMD is leading to radical changes in the structure and functioning of spatial networks, which are likely to intensify as human activities increase in scope and extent. Here, we provide an overview to guide research into HMD and the resulting rewiring of spatial networks, making predictions about the ecological and evolutionary consequences and how these vary according to spatial scale and the traits of species. Future research should consider HMD holistically, assessing the range of direct and indirect processes to understand the complex impacts on eco-evolutionary dynamics.

An increasing number of studies have investigated the consequences of biodiversity loss for the occurrence of vector-borne diseases such as Lyme borreliosis, the most common tick-borne disease in the northern hemisphere. As host species differ in their ability to transmit the Lyme borreliosis bacteria Borrelia burgdorferi s.l. to ticks, increased host diversity can decrease disease prevalence by increasing the proportion of dilution hosts, host species that transmit pathogens less efficiently. Previous research shows that Lyme borreliosis risk differs between forest types and suggests that a higher diversity of host species might dilute the contribution of small rodents to infect ticks with B. afzelii, a common Borrelia genospecies. However, empirical evidence for a dilution effect in Europe is largely lacking. We tested the dilution effect hypothesis in 19 Belgian forest stands of different forest types along a diversity gradient. We used empirical data and a Bayesian belief network to investigate the impact of the proportion of dilution hosts on the density of ticks infected with B. afzelii, and identified the key drivers determining the density of infected ticks, which is a measure of human infection risk. Densities of ticks and B. afzelii infection prevalence differed between forest types, but the model indicated that the density of infected ticks is hardly affected by dilution. The most important variables explaining variability in disease risk were related to the density of ticks. Combining empirical data with a model-based approach supported decision making to reduce tick-borne disease risk. We found a low probability of a dilution effect for Lyme borreliosis in a north-western European context. We emphasize that under these circumstances, Lyme borreliosis prevention should rather aim at reducing tick-human contact rate instead of attempting to increase the proportion of dilution hosts.

Urbanisation represents one of the most radical forms of terrestrial land use change and has been shown to lead to alterations in ecosystem functioning and community dynamics and changes in individual phenotypic traits. While the recent surge in microbiome studies has brought about a paradigm shift by which individuals cannot truly be considered independently of the bacterial communities they host, the role of gut microbiota in organismal response to human-induced environmental change is still scarcely studied. Here, we applied a metabarcoding approach to examine the impact of urbanisation on the gut microbiota of Passer domesticus. We found urbanisation to be associated to lower microbiota species diversity, modifications in taxonomic composition and community structure, and changes in functional composition. The strength of these relationships, however, depended on the spatial scale and season at which they were considered. Such spatio-temporal effect suggests that urbanisation may dampen the natural seasonal variation of the gut microbiota observed in more pristine habitats, potentially influencing the fitness of urban organisms. Our results hence shed light on a hitherto little considered perspective, i.e. that the negative effects of urbanisation on city-dwelling organisms may extend to their microbiomes, causing potential dysbioses.

Dispersal is a process of central importance for the ecological and evolutionary dynamics of populations and communities, because of its diverse consequences for gene flow and demography. It is subject to evolutionary change, which begs the question, what is the genetic basis of this potentially complex trait? To address this question, we (i) review the empirical literature on the genetic basis of dispersal, (ii) explore how theoretical investigations of the evolution of dispersal have represented the genetics of dispersal, and (iii) discuss how the genetic basis of dispersal influences theoretical predictions of the evolution of dispersal and potential consequences. Dispersal has a detectable genetic basis in many organisms, from bacteria to plants and animals. Generally, there is evidence for significant genetic variation for dispersal or dispersal-related phenotypes or evidence for the micro-evolution of dispersal in natural populations. Dispersal is typically the outcome of several interacting traits, and this complexity is reflected in its genetic architecture: while some genes of moderate to large effect can influence certain aspects of dispersal, dispersal traits are typically polygenic. Correlations among dispersal traits as well as between dispersal traits and other traits under selection are common, and the genetic basis of dispersal can be highly environment-dependent. By contrast, models have historically considered a highly simplified genetic architecture of dispersal. It is only recently that models have started to consider multiple loci influencing dispersal, as well as non-additive effects such as dominance and epistasis, showing that the genetic basis of dispersal can influence evolutionary rates and outcomes, especially under non-equilibrium conditions. For example, the number of loci controlling dispersal can influence projected rates of dispersal evolution during range shifts and corresponding demographic impacts. Incorporating more realism in the genetic architecture of dispersal is thus necessary to enable models to move beyond the purely theoretical towards making more useful predictions of evolutionary and ecological dynamics under current and future environmental conditions. To inform these advances, empirical studies need to answer outstanding questions concerning whether specific genes underlie dispersal variation, the genetic architecture of context-dependent dispersal phenotypes and behaviours, and correlations among dispersal and other traits.

While numerous studies have reported negative effects of urbanisation on birds, few have examined the role of urban scale in influencing breeding success. Furthermore, many studies have relied on qualitative rather than quantitative assessments of urbanisation. This study sought to address these issues by testing the effects of urbanisation, measured at two spatial scales, on the breeding success of great tits Parus major. A nested study design, incorporating over 400 nestboxes, was used in study sites across northern Belgium with a priori quantified degrees of urbanisation at both local and regional scales. All measured breeding parameters were found to vary at one or both spatial scales of urbanisation; in more urbanised areas great tits displayed advanced laying dates but lower breeding success compared to rural areas, with smaller clutch sizes, lower nestling masses and fewer fledglings per egg. Importantly, urbanisation effects were not limited to big cities as birds breeding in gardens or parks in small towns also had comparatively low success. We found that both regional- and local-scale urbanisation had consistent significant effects on laying date, clutch size and nestling mass, while the number of fledglings per egg was negatively influenced by local-scale urbanisation only. Results of this study therefore highlight the importance of utilising multiple spatial scales in analysing urbanisation effects, as well as the potential negative impact of local urbanisation on breeding success. This calls for further investigation into mechanisms driving urbanisation effects and how these may vary at different scales.

A major aim of evolutionary biology is to understand why patterns of genomic variation vary among populations and species. Large-scale genomic studies of widespread species are useful for studying how the environment and demographic history shape patterns of genomic divergence, and with the continually decreasing cost of sequencing, such studies are now becoming feasible. Here, we carry out one of the most comprehensive surveys of genomic variation in a wild vertebrate to date; the great tit ( Parus major ) HapMap project. We screened ca 500,000 SNP markers across 647 individuals from 29 populations, spanning almost the entire geographic range of the European great tit subspecies. We found that genome-wide variation was consistent with a recent colonisation across Europe from a single refugium in the Balkans and/or Turkey, with bottlenecks and reduced genetic diversity in island populations. Differentiation across the genome was highly heterogeneous, with clear “islands of differentiation” even among populations which are ostensibly panmictic. These islands of differentiation were consistently found in regions of low recombination, suggesting that background selection can rapidly promote population differentiation among even the most recently colonised populations. We also detected genomic outlier regions that were unique to peripheral great tit populations, most likely as a result of recent directional selection at the range edges of this species. These “unique” outlier regions contained candidate genes for morphology, thermal adaptation and colouration, supporting previous research in this species, and providing avenues for future investigation. Our study suggests that comprehensive screens of genomic variation in wild organisms can provide unique insights into evolution. Author summary Studying patterns of genetic variation is a useful way of determining why populations and species differ in nature. Genetic variation is shaped by natural selection, but also by the present and past size of populations, the amount of migration, and by features of the genome, such as variation in recombination rate, of the organism being studied. Teasing apart the effects of these different processes on genomic diversity is difficult, but one way that this can be achieved is by studying genomic variation across the entire range of a species. We performed a continental-scale analysis of genetic variation in the great tit - a widespread songbird that has been the focus of extensive ecological research. We first used genomic data to reconstruct the historical colonisation of great tits across Europe, and showed that during the last ice age, this species was likely restricted to a single region in Eastern Europe, from which they spread across the continent. We then studied how patterns of variation differ along the genome, and show that recombination rate is a key driver of variation among all populations. Importantly, by comparing many populations we were able to identify genes that have been subject to natural selection in specific geographical regions. We found that natural selection appeared to be strongest in populations on the edges of the great tit’s range acting on traits such as morphology, stress response and colouration. Large-scale genetic analyses such as ours are therefore useful approaches for understanding how evolution operates in the wild.

The risk of tick-borne disease in humans depends on the exposure to pathogen-infected ticks, which in turn is driven by local tick population densities, pathogen prevalence and human activity. Variation in tick densities and pathogen prevalence between green spaces differing in habitat characteristics, location and geography has been well documented. In contrast, variation within green spaces, although vital for management and prevention of disease risk, remains poorly understood. Studying this variation may lead to a better understanding of the drivers of small-scale tick distribution and reveal priority locations for tick management. We sampled ticks within green spaces at three location types representing different green space infrastructures and levels of human activity: (1) a structural element (e.g. bench); (2) the ecotone 40 m further along the adjacent trail and; (3) the interior of the associated forest stand, 20 m perpendicular to the trail, between (1) and (2). Drag sampling took place in 2018 and 2019 at 36 locations in 10 green spaces located in the Campine region of Flanders, Belgium. The density of questing nymphs (DON) was lowest at structural elements and slightly higher adjacent to trails. The highest tick densities were recorded in the forest interior. DON was higher in deciduous than in coniferous forests as well as in stands with a more developed shrub layer. This was true for all location types, as we observed a strong correlation of DON between location types within forest stands. This enables the prediction of DON within forest stands, thus enabling the prediction of DON near infrastructure based on the associated forest stand characteristics. Prevention and management efforts should be focused on infrastructure in or adjacent to deciduous, structure-rich forest stands, although large variation in DON at all location types indicates overall, factual risk while using green space infrastructure. This work was supported by the Research Foundation - Flanders [grant number 60897] Van Gestel, M (corresponding author), Campus Drie Eiken,Room D1-46,Univ Pl 1, B-2610 Antwerp, Belgium. Mats.vangestel@uantwerpen.be

SUMMARYLyme disease is caused by bacteria of theBorrelia burgdorferigenospecies complex and transmitted by Ixodid ticks. In North America only one pathogenic genospecies occurs, in Europe there are several. According to the dilution effect hypothesis (DEH), formulated in North America, nymphal infection prevalence (NIP) decreases with increasing host diversity since host species differ in transmission potential. We analysedBorreliainfection in nymphs from 94 forest stands in Belgium, which are part of a diversification gradient with a supposedly related increasing host diversity: from pine stands without to oak stands with a shrub layer. We expected changing tree species and forest structure to increase host diversity and decrease NIP. In contrast with the DEH, NIP did not differ between different forest types. Genospecies diversity however, and presumably also host diversity, was higher in oak than in pine stands. Infected nymphs tended to harbourBorrelia afzeliiinfection more often in pine stands whileBorrelia gariniiandBorrelia burgdorferiss. infection appeared to be more prevalent in oak stands. This has important health consequences, since the latter two cause more severe disease manifestations. We show that the DEH must be nuanced for Europe and should consider the response of multiple pathogenic genospecies.

The red squirrel (Sciurus vulgaris) population in the United Kingdom has declined over the last century and is now on the UK endangered species list. This is the result of competition from the eastern grey squirrel (S. carolinensis) which was introduced in the 19th century. However, recent evidence suggests that the rate of population decline is enhanced by squirrelpox disease, caused by a viral infection carried asymptomatically by grey squirrels but to which red squirrels are highly susceptible. Population genetic diversity provides some resilience to rapidly evolving or exotic pathogens. There is currently no data on genetic diversity of extant UK squirrel populations with respect to genes involved in disease resistance. Diversity is highest at loci involved in the immune response including genes clustered within the major histocompatibility complex (MHC). Using the class II DRB locus as a marker for diversity across the MHC region we genotyped 110 red squirrels from locations in the UK and continental Europe. Twenty-four Scvu-DRB alleles at two functional loci; Scvu-DRB1 and Scvu-DRB2, were identified. High levels of diversity were identified at both loci in the continental populations. In contrast, no diversity was observed at the Scvu-DRB2 locus in the mainland UK population while a high level of homozygosity was observed at the Scvu-DRB1 locus. The red squirrel population in the UK appears to lack the extensive MHC diversity associated with continental populations, a feature which may have contributed to their rapid decline.

Dispersal and migration are superficially similar large-scale movements, but which appear to differ in terms of inter-individual behavioural synchronization. Seasonal migration is a striking example of coordinated behaviour, enabling animal populations to track spatio-temporal variation in ecological conditions. By contrast, for dispersal, while social context may influence an individual's emigration and settlement decisions, transience is believed to be mostly a solitary behaviour. Here, we review differences in drivers that may explain why migration appears to be more synchronized than dispersal. We derive the prediction that the contrast in the importance of behavioural synchronization between dispersal and migration is linked to differences in the selection pressures that drive their respective evolution. Although documented examples of collective dispersal are rare, this behaviour may be more common than currently believed, with important consequences for eco-evolutionary dynamics. Crucially, to date, there is little available theory for predicting when we should expect collective dispersal to evolve, and we also lack empirical data to test predictions across species. By reviewing the state of the art in research on migration and collective movements, we identify how we can harness these advances, both in terms of theory and data collection, to broaden our understanding of synchronized dispersal and its importance in the context of global change.

Urban sprawl increasingly affects the ecology of natural populations, including host-microbiota interactions, with observed differences in the gut microbiota between urban and rural hosts. While different mechanisms could explain this pattern, dietary uptake constitutes a likely candidate. To assess the contribution of diet in explaining urban-rural variation in gut microbiota, we performed an aviary experiment in which urban and rural house sparrows were fed with mimics of urban or rural diets. Before the experiment, rural sparrows hosted more diverse gut communities, with a higher relative abundance of Enterococcaceae and Staphylococcaceae and lower abundance of genes involved in xenobiotic degradation and lipid metabolism than their urban counterparts. The experimental diets significantly altered gut microbiota alpha- and beta-diversity and taxonomic composition, with the strongest shifts occurring in individuals exposed to contrasting diets. Overall, diet-induced shifts resembled initial differences between free-ranging urban and rural hosts. Furthermore, rural diet had a positive impact on urban host body mass but only in hosts with the highest initial gut diversity. Overall, our results indicate that diet constitutes an important factor contributing to differences in gut microbiota along the urbanization gradient and provide new insights on possible fitness consequences of a reduced gut diversity in urban settings.

Aim To mitigate the threat invasive species pose to ecosystem functioning, reliable risk assessment is paramount. Spatially explicit predictions of invasion risk obtained through bioclimatic envelope models calibrated with native species distribution data can play a critical role in invasive species management. Forecasts of invasion risk to novel environments, however, remain controversial. Here, we assess how species’ association with human-modified habitats in the native range and within-taxon niche structure shape the distribution of invasive populations at biogeographical scales and influence the reliability of predictions of invasion risk. Location Africa, Asia and Europe. Methods We use ~1200 native and invasive ring-necked parakeet (Psittacula krameri) occurrences and associated data on establishment success in combination with mtDNA-based phylogeographic structure to assess niche dynamics during biological invasion and to generate predictions of invasion risk. Niche dynamics were quantified in a gridded environmental space while bioclimatic models were created using the biomod2 ensemble modelling framework. Results Ring-necked parakeets show considerable niche expansion into climates colder than their native range. Only when incorporating a measure of human modification of habitats within the native range do bioclimatic envelope models yield credible predictions of invasion risk for parakeets across Europe. Invasion risk derived from models that account for differing niche requirements of phylogeographic lineages and those that do not achieve similar statistical accuracy, but there are pronounced differences in areas predicted to be susceptible for invasion. Main conclusions Information on within-taxon niche structure and especially association with humans in the native range can substantially improve predictive models of invasion risk. To provide policymakers with robust predictions of invasion risk, including these factors into bioclimatic envelope models is recommended.

Males of the speckled wood butterfly Pararge aegeria L. (Satyrinae), actively search for females (“patrolling”) or wait for them at particular places (“perching”). Darker males are more likely to patrol than pale ones, which are mainly territorial perchers. We studied whether this morphological variation relates to thermoregulatory differences. The relationship between thoracic temperature and ambient temperature differed between the colour types under natural conditions: darker males had on average lower body temperatures than paler males. Different activities (e.g. resting, flying) and behavioural strategies (perching or patrolling) were associated with differences in thoracic temperature: patrolling males which mainly engaged in long flights and periods of basking afterwards, had lower thoracic temperatures than perching males which engaged in very short flights, fights and basking. When resting for a while thoracic temperatures did not differ between males practising different strategies. Under laboratory conditions, darker males heated up faster than pale males but there was no difference in the thoracic temperature at which they started to fly. These results indicate that thermal requirements (or general conditions) differ between the behavioural strategies, and that behavioural differences between phenotypes (colour types) relate to differences in thermal ecology. This supports the idea that darker males are better adapted to patrolling. There is no evidence that one mate-locating strategy is always superior to the other, which coincides with the observation that both strategies co-exist. More generally, this study shows that relatively small differences in colour can have a considerable effect on thermoregulation and hence on the behavioural strategies a heliothermic insect will adopt.

We studied the costs of lamellae autotomy with respect to growth and survival of Lestes sponsa damselfly larvae in field experiments. We manipulated predation risk by Aeshna cyanea dragonfly larvae and lamellae status of L. sponsa larvae in field enclosures and compared differences in numbers, size and mass of survivors among treatments. In the absence of a free-ranging A. cyanea larva, about 29% of the L. sponsa larvae died. This was probably due to cannibalism. The presence of a free-ranging A. cyanea reduced larval survival by 68% compared to treatments in which it was absent or not permitted to forage on L. sponsa damselflies. Across all predator treatments, lamellae autotomy reduced survival by about 20%. The mean head width and mass of survivors was lower in the enclosures with a free-ranging A. cyanea compared to the other two predator treatments. This suggested that larvae grew less in the presence of a free-ranging predator, indicating that increased antipredator behaviours were more important in shaping growth responses than reduced population density. Mass, but not head width, of survivors was also reduced after autotomy. The fitness consequences of these effects for the adults may be pronounced. In general, these field data strongly suggest that lamellae autotomy affects population regulation of damselflies.

A growing number of studies have recently reported links between personality and dispersal, yet the adaptive significance of personality-dependent dispersal strategies still remain poorly understood. Here, we tested whether birds differing in exploratory behaviour (EB, quantified by a novel environment test), gain differential fitness benefits when remaining in or leaving their natal area in a natural population of great tits (Parus major). Using data on lifetime fecundity, we found no evidence that particular combinations of EB and dispersal lead to a higher fitness after settlement. In males, we found positive selection acting on EB through first-year survival, resulting in a positive relationship between EB and number of lifetime fledglings. However, contrary to expectation, these effects were only present in local recruits and not in immigrants. Male immigrants produced fewer fledglings during their lifetime than did local recruits, suggesting selection against dispersal at a scale exceeding the size of our study area. In females, we found selection on EB through recruitment with respect to year (i.e., negative, positive and stabilizing selection), but neither the strength nor the direction differed among birds with different dispersal strategies. We found no evidence in either sex for lifetime selection acting on EB using recruits as a measure of fitness. In conclusions, our results do not support the hypothesis that the co-expression of personality and dispersal is shaped by selection after settlement.

The survival of large and mobile species in the face of habitat loss and fragmentation depends on several factors, including the landscape configuration of subpopulations and the dispersal capabilities of the species. We performed a landscape analysis of the Bolivian Tropical Andes to determine whether remaining habitat patches were suitable in terms of ecological characteristics and potential connectivity for the long-term survival of the Andean bear (Tremarctos ornatus). First we built a ruled-based model to identify key patches or areas large enough to sustain a viable population by using knowledge of Andean bear habitat requirements and movement patterns. Second, we estimated potential functional connectivity among these areas applying a cost-distance analysis based on estimates of the resistance to movement through the landscape. Finally, we quantified the proportion of these key patches and corridor habitats within the Bolivian protected area system. The rule-based model identified 13 key patches covering 21,113 km(2) corresponding to a maximum estimated population of 3,165 adult bears. Using cost-distance analysis, all 13 key patches were potentially connected to >= 1 other key patch. Twelve of the patches were at least partially protected by national parks, and 40% of areas considered suitable as corridors were included within a protected area. Although the current protected area system includes suitable bear habitat, large portions of key patches and corridors are unprotected, which could eventually lead to fragmentation and habitat loss if these areas are not protected.

Animal personalities can play an important role in how individuals deal with specific social situations, which may have important fitness consequences. A link between personality and social skills seems obvious from a human perspective and it is therefore surprising that relatively few studies have looked at the effect of animal personality on communicative interactions. In this study we tested the evidence for personality-related variation in birds during vocal interactions in the field. We scored individual variation in exploratory behaviour of great tits, Parus major, in repeated trials of a novel environment test in the laboratory and related this score to approach and vocal behaviour in response to playback of conspecific songs in the field. Contrary to our expectation, we found males with low exploration scores approached the speaker more closely than birds with high exploration scores. Birds with high exploration scores seemed to respond less aggressively while dealing with the simulated intruder vocally by switching song types more often. Our results suggest that slow-exploring birds in the laboratory respond more strongly during natural interactions in the field. An independent and very similar study also revealed a correlation between personality score and playback response, but with a seemingly contrasting outcome. A comparison of the two studies provides insights into the complexity of interpreting playback results and territorial response patterns in general.

Our study tested whether two European bird-specialized ticks, Ixodes arboricola and I. frontalis, can act as vectors in the transmission cycles of Borrelia burgdorferi s.l. The ticks have contrasting ecologies but share songbird hosts (such as the great tit, Parus major) with the generalist I. ricinus which may therefore act as a bridging vector. In the first phase of the experiment, we obtained Borrelia-infected ornithophilic nymphs by exposing larvae to great tits that had previously been exposed to I. ricinus nymphs carrying a community of genospecies (Borrelia garinii, valaisiana, afzelii, burgdorferi s.s., spielmanii). Skin samples showed that birds selectively amplified B. garinii and B. valaisiana. The spirochetes were transmitted to the ornithophilic ticks and survived moulting, leading to infection rates of 16% and 27% in nymphs of I. arboricola and I. frontalis respectively. In the second phase, pathogen-free great tits were exposed to the Borrelia-infected ornithophilic nymphs. None of these ticks were able to infect the birds, as indicated by the tissue samples. Analysis of xenodiagnostic I. ricinus larvae found no evidence for co-feeding or systemic transmission of B. burgdorferi s.l. These outcomes do not support the occurrence of enzootic cycles of Borrelia burgdorferi s.l. involving songbirds and their specialized ornithophilic ticks.

We investigated to what extent a European songbird (Parus major) selectively transmits and amplifies Borrelia burgdorferi s.l. bacteria. Borrelia-naive birds were recurrently exposed to Ixodes ricinus nymphs carrying a community of more than 34 5S-23S genotypes belonging to five genospecies (Borrelia garinii, Borrelia valaisiana, Borrelia afzelii, B. burgdorferi s.s. and Borrelia spielmanii). Fed ticks were screened for Borrelia after moulting. We found evidence for co-feeding transmission of avian and possibly also mammalian genotypes. Throughout the course of infestations, the infection rate of B. garinii and B. valaisiana increased, indicating successful amplification and transmission, while the infection rate for B. afzelii, B. burgdorferi s.s and B. spielmanii tended to decrease. Within the B. garinii and B. valaisiana genotype communities, certain genotypes were transmitted more than others. Moreover, birds were able to host mixed infections of B. garinii and B. valaisiana, as well as mixed infections of genotypes of the same genospecies. We experimentally show that resident songbirds transmit a broad range of Borrelia genotypes, but selectively amplify certain genotypes, and that one bird can transmit simultaneously several genotypes. Our results highlight the need to explicitly consider the association between genotypes and hosts, which may offer opportunities to point out which hosts are most responsible for the Borrelia presence in questing ticks.

Aim Niche conservatism, or the extent to which niches are conserved across space and time, is of special concern for the study of non-native species as it underlies predictions of invasion risk. Based on the occurrence of 28 non-native birds in Europe, we assess to what extent Grinnellian realized niches are conserved during invasion, formulate hypotheses to explain the variation in observed niche changes and test how well species distribution models can predict non-native bird occurrence in Europe.

Summary We examined the Borrelia burgdorferi sensu lato circulation in a tick community consisting of three species (Ixodes ricinus, I. frontalis, I. arboricola) with contrasting ecologies, but sharing two European songbird hosts (Parus major and Cyanistes caeruleus). Parus major had the highest infestation rates, primarily due to larger numbers of I. ricinus, and probably because of their greater low-level foraging. The prevalence of Borrelia in feeding ticks did not significantly differ between the two bird species; however, P. major in particular hosted large numbers of Borrelia-infected I. frontalis and I. ricinus larvae, suggesting that the species facilitates Borrelia transmission. The low but significant numbers of Borrelia in questing I. arboricola ticks also provides the first field data to suggest that it is competent in maintaining Borrelia. Aside from Borrelia garinii, a high number of less dominant genospecies was observed, including several mammalian genospecies and the first record of Borrelia turdi for North-Western Europe. Borrelia burgdorferi sensu lato IGS genotypes were shared between I. arboricola and I. ricinus and between I. frontalis and I. ricinus, but not between I. arboricola and I. frontalis. This suggests that the Borrelia spp. transmission cycles can be maintained by bird-specific ticks, and bridged by I. ricinus to other hosts outside bird-tick cycles.

Cavity shortage limits secondary cavity-nesting birds (SCN) assemblages in intensively-managed forests with few cavities, but whether cavity availability limits SCN in cavity-rich forests remains unclear. To address this issue, we used a replicated before-after-control-impact design in which the availability of cavities was experimentally reduced and increased in cavity-rich Quercus pyrenaica forests. Cavity-blocking and nest-box addition led to reduction by 66% and increase by 185%, respectively, of nest densities of blue (Cyanistes caeruleus) and great (Parus major) tits. Analysis of cavity characteristics suggests that tits occupied less suitable cavities in blocking plots compared to control and nest-box addition plots after treatments, further indicating that the availability of suitable nest-sites may limit tit populations. No effect of cavity manipulation was found for short-toed treecreepers (Certhia brachydactyla) and Eurasian nuthatches (Sitta europaea), which defend larger territories than tits and whose numbers may be more influenced by territoriality than by nest-sites. Alternatively, high plasticity in nest-site use of treecreepers may enable them to cope with reductions in cavity availability and increased interspecific competition. Spotless starlings (Sturnus unicolor) were little affected by cavity manipulation. Often foraging on pastures, starlings may be more limited through reduced foraging opportunities in this forested landscape than by nest-sites. In addition, starlings may exclude other species from their preferred cavities to cope with reductions in numbers of suitable cavities. Community responses to the variation in cavity abundance in cavity-rich Pyrenean oak forests may therefore vary considerably among species, depending on their plasticity in nest-site use, behavioral dominance, foraging conditions or social structure. Thus, management actions aimed to increase cavity abundance and suitability for SCN communities in cavity-rich forests should consider inter-specific interactions and species-specific characteristics of the targeted species.

Persistence of forest-dependent species in fragmented landscapes strongly relies on sufficient dispersal between patches, making it important to understand how animal movements are affected by the intervening matrix. Movements can be influenced through selection or avoidance of land cover based on their perceived suitability for foraging or providing cover. The composition and configuration of the matrix will, therefore, most likely be an important factor to consider when estimating connectivity between patches. To address this, we performed translocation experiments to understand how forest birds used different land cover types in a fine-grained matrix of a fragmented Afromontane biodiversity hotspot (Taita Hills, Kenya). Our results revealed that use of land cover types for both the forest specialist Cabanis’s greenbul Phyllastrephus cabanisi and for the forest generalist white-starred robin Pogonocichla stellata was disproportional to their availability. However, this effect was influenced by matrix configuration; in patchy matrices, land cover selection was more pronounced compared with more uniform matrices, especially for the forest specialist. At the scale of movement steps, risk avoidance seemed to be a strong factor in the route decisions for both species. Observed steps contained on average lower proportions of open land cover and did less frequently intersect built-up areas than expected. P. stellata did not differentiate between the alternative land cover types, whereas P. cabanisi preferred steps that contained more indigenous forest. The observed negative relationship between degree of forest dependency and matrix permeability implies that for members of the Taita bird community, which are even more dependent on intact forest habitat (i.e. the critically endangered Taita thrush Turdus helleri), current permeability of the matrix may be even lower. Matrix restoration to improve connectivity may, therefore, be a crucial instrument for the long-term survival of forest-dependent species in these fragmented Afromontane landscapes.

Dispersal costs can be classified into energetic, time, risk and opportunity costs and may be levied directly or deferred during departure, transfer and settlement. They may equally be incurred during life stages before the actual dispersal event through investments in special morphologies. Because costs will eventually determine the performance of dispersing individuals and the evolution of dispersal, we here provide an extensive review on the different cost types that occur during dispersal in a wide array of organisms, ranging from micro-organisms to plants, invertebrates and vertebrates. In general, costs of transfer have been more widely documented in actively dispersing organisms, in contrast to a greater focus on costs during departure and settlement in plants and animals with a passive transfer phase. Costs related to the development of specific dispersal attributes appear to be much more prominent than previously accepted. Because costs induce trade-offs, they give rise to covariation between dispersal and other life-history traits at different scales of organismal organisation. The consequences of (i) the presence and magnitude of different costs during different phases of the dispersal process, and (ii) their internal organisation through covariation with other life-history traits, are synthesised with respect to potential consequences for species conservation and the need for development of a new generation of spatial simulation models.

1. Interest in the evolutionary origin and maintenance of individual behavioural variation and behavioural plasticity has increased in recent years. 2. Consistent individual behavioural differences imply limited behavioural plasticity, but the proximate causes and wider consequences of this potential constraint remain poorly understood. To date, few attempts have been made to explore whether individual variation in behavioural plasticity exists, either within or between populations. 3. We assayed 'exploration behaviour' among wild-caught individual great tits Parus major when exposed to a novel environment room in four populations across Europe. We quantified levels of individual variation within and between populations in average behaviour, and in behavioural plasticity with respect to (i) repeated exposure to the room (test sequence), (ii) the time of year in which the assays were conducted and (iii) the interval between successive tests, all of which indicate habituation to novelty and are therefore of functional significance. 4. Consistent individual differences ('I') in behaviour were present in all populations; repeatability (range: 0.34-0.42) did not vary between populations. Exploration behaviour was also plastic, increasing with test sequence - but less so when the interval between subsequent tests was relatively large - and time of year; populations differed in the magnitude of plasticity with respect to time of year and test interval. Finally, the between-individual variance in exploration behaviour increased significantly from first to repeat tests in all populations. Individuals with high initial scores showed greater increases in exploration score than individuals with low initial scores; individual by environment interaction ('I × E') with respect to test sequence did not vary between populations. 5. Our findings imply that individual variation in both average level of behaviour and behavioural plasticity may generally characterize wild great tit populations and may largely be shaped by mechanisms acting within populations. Experimental approaches are now needed to confirm that individual differences in behavioural plasticity (habituation) - not other hidden biological factors - caused the observed patterns of I × E. Establishing the evolutionary causes and consequences of this variation in habituation to novelty constitutes an exciting future challenge.

Secondary cavity-nesting birds (SCN), which cannot create their own breeding cavities, are expected to be influenced by habitat alteration caused by forest management practices, but the mechanisms underlying the distribution pattern of SCN subjected to different management systems are poorly known. To improve our knowledge on these mechanisms, we examine cavity abundance, cavity occupation and reproductive performance of SCN in Pyrenean oak ( Quercus pyrenaica ) forests subjected to two management systems: (i) dense “young forests”, maintained at such stage by clear-cuttings and burns, and (ii) “old forest”, subjected to extensive traditional grazing and scarce firewood extraction by selective cutting. Young forests had considerably lower density of cavities (1.29 ± 0.71 vs 15.09 ± 2.00 cavities ha −1 ), SCN species (0.18 ± 0.11 vs 0.61 ± 0.07 species ha −1 ) and nests (0.40 ± 0.27 vs 2.67 ± 0.25 nests of all SCN ha −1 ) than old forests, indicating that a low availability of cavities may limit SCN assemblages in young oak forests. However, reproductive parameters of great ( Parus major ) and blue ( Cyanistes caeruleus ) tits associated with the availability of food (laying date, clutch size, nestling number and weight, adult weight) did not differ between both forest types, suggesting that food supply was not reduced in young forests, at least for tits during the breeding season. Large diameter (up to 170 cm dbh) decayed trees were the most likely to hold cavities, but birds preferred smaller living cavity-trees for nesting (90% of nests in 21–65 cm dbh trees). The preservation of cavity-trees within traditionally managed old oak forests is crucial in providing nesting opportunities to SCN. Besides, the protection of these traditionally managed forests would also benefit to other forest organisms that depend on old and open oak forests.

Habitat fragmentation can restrict geneflow, reduce neighbourhood effective population size, and increase genetic drift and inbreeding in small, isolated habitat remnants. The extent to which habitat fragmentation leads to population fragmentation, however, differs among landscapes and taxa. Commonly, researchers use information on the current status of a species to predict population effects of habitat fragmentation. Such methods, however, do not convey information on species-specific responses to fragmentation. Here, we compare levels of past population differentiation, estimated from microsatellite genotypes, with contemporary dispersal rates, estimated from multi-strata capture-recapture models, to infer changes in mobility over time in seven sympatric, forest-dependent bird species of a Kenyan cloud forest archipelago. Overall, populations of sedentary species were more strongly differentiated and clustered compared to those of vagile ones, while geographical patterning suggested an important role of landscape structure in shaping genetic variation. However, five of seven species with broadly similar levels of genetic differentiation nevertheless differed substantially in their current dispersal rates. We conclude that post-fragmentation levels of vagility, without reference to past population connectivity, may not be the best predictor of how forest fragmentation affects the life history of forest-dependent species. As effective conservation strategies often hinge on accurate prediction of shifts in ecological and genetic relationships among populations, conservation practices based solely upon current population abundances or movements may, in the long term, prove to be inadequate.

Parental care has been widely studied in birds and mammals, but variation in space use in family groups has received less attention, despite its potential importance for both survival and subsequent dispersal of offspring. In this study, we evaluate factors affecting postfledging family space use in a small territorial songbird, the great tit (Parus major). Family space use was monitored using radio tracking. Our main objectives were 1) to quantify in detail the temporal and spatial scale of family movements, 2) to test behavioral hypotheses explaining when and how frequently families leave their breeding territory, and 3) to test to what extent movements were based on familiarity with the environment. We found that variation in space use was to a large extent due to some families, but not others, regularly undertaking foraging excursions of up to more than a kilometer away. Daily excursion probability was higher for families occupying low-quality territories, and consequently, these families covered larger areas during foraging. Excursion behavior and range use also strongly depended on maternal breeding experience and personality. We further present some striking examples of inexperienced mothers moving toward previously visited areas, suggesting that familiarity with the environment plays an important role in patterns of space use. Overall, our results suggest that variation in family movements reflects different foraging strategies in relation to parental characteristics. Copyright 2011, Oxford University Press.

Many bird species start laying their eggs earlier in response to increasing spring temperatures, but the causes of variation between and within species have not been fully explained. Moreover, synchronization of the nestling period with the food supply not only depends on first-egg dates but also on additional reproductive parameters including laying interruptions, incubation time and nestling growth rate. We studied the breeding cycle of two sympatric and closely related species, the blue tit Cyanistes caeruleus and the great tit Parus major in a rich oak-beech forest, and found that both advanced their mean first-egg dates by 11-12 days over the last three decades. In addition, the time from first egg to fledging has shortened by 2-3 days, through a decrease in laying interruptions, incubation time (not statistically significant) and nestling development time. This decrease is correlated with a gradual increase of temperatures during laying, suggesting a major effect of the reduction in laying interruptions. In both species, the occurrence of second clutches has strongly decreased over time. As a consequence, the average time of fledging (all broods combined) has advanced by 15.4 and 18.6 days for blue and great tits, respectively, and variance in fledging dates has decreased by 70-75%. Indirect estimates of the food peak suggest that both species have maintained synchronization with the food supply. We found consistent selection for large clutch size, early laying and short nest time (laying to fledging), but no consistent changes in selection over time. Analyses of within-individual variation show that most of the change can be explained by individual plasticity in laying date, fledging date and nest time. This study highlights the importance of studying all components of the reproductive cycle, including second clutches, in order to assess how natural populations respond to climate change.

Little is known about the foraging ecology of invasive bird species in Europe. We used radio-telemetry to assess home-ranges of breeding male Ring-necked Parakeets in Brussels. Results indicate that parakeets primarily forage in parks and gardens while avoiding forests. This can probably be explained by the higher food availability in anthropogenic habitats and fits a general pattern that invasive species generally select heavily altered environments.

Birdsong can play a critical role in establishing a territory and finding a mate among individuals from local and foreign populations. Variation in birdsong among populations can be influenced by habitat fragmentation and might affect successful dispersal among habitat fragments. We studied variation in great tit song in a long-term study population distributed over nine forest fragments. All individual males recorded had a known dispersal history within the fragmented forest habitat. We found spatial structure of declining song-type sharing with distance, with a marked drop from an individual’s own forest fragment to another across a habitat gap. We also found decreasing song similarity among increasingly distant fragments in terms of temporal and spectral characteristics of shared song types. The change in acoustic structure was more gradual and seemed less affected by habitat discontinuity but also showed a tight correlation with dispersal index among forest fragments. Immigrant birds shared fewer song types with neighbouring birds that were born within the same forest fragment, but not less compared to birds born in another forest fragment within the study area. Our data provide detailed insight into the relationship between song differentiation and male dispersal and contribute to our understanding of the potential role of song in reproductive exchange and avian speciation. The fact that birds in small forest fragments shared more songs than birds in larger forest fragments confirms that song analysis has potential as a tool for conservation in rare species.

Summary 1. Biological invasions are a major threat to biodiversity. Given the large number of invasive species, we need to be able to identify invaders with large effects in order to prioritize management efforts. Here, we present a framework using species distribution modelling to predict how abundance of native species will change as a result of competition with an invasive species using ring-necked parakeets Psittacula krameri and nuthatches Sitta europaea as a case study. 2. Ring-necked parakeets are widely introduced throughout Europe and compete with native cavity-nesters. Using the relationship between parakeet abundance and environmental variables in Brussels (Belgium), we predict parakeet abundance across Flanders. A competition coefficient, quantifying the parakeets’ impact on nuthatches, was obtained by a regression on a data set of parakeet and nuthatch abundance. An estimate of the number of nuthatches that will be lost when parakeets have occupied all suitable sites was calculated by superimposing the abundance maps of the two species and applying the competition coefficient. 3. Our results predict a potential population of about 22 000 parakeet pairs, indicating that they could become one of the most numerous cavity-nesters in the region. Parakeet abundance is the highest in older, more fragmented forest in urban areas whereas nuthatches prefer larger, old and oak-dominated forests. 4. Our models indicate that throughout much of their range, and in a variety of habitats, parakeets and nuthatches will compete for nesting cavities, but as the competition strength is only moderate, the total impact of parakeets on nuthatch populations will be limited, with at most one-third of the population at risk. 5. Synthesis and applications. Species distribution models combined with empirical estimates of competition strength can be used as a general tool to make an assessment of the potential impact of established invasive species. Such information is required to make effective decisions on how to prioritize management effort and resources across the multitude of invasive species that currently threaten native ecosystems. For the ring-necked parakeet, our results indicate that there is no compelling evidence indicating that parakeets pose a threat large enough to justify an eradication campaign where they are currently present.

Conditions experienced during early development may affect both adult phenotype and performance later during life. Phenotypic traits may hence be used to indicate past growing conditions and predict future survival probabilities. Relationships between phenotypic markers and future survival are, however, highly heterogeneous, possibly because poor- and high-quality individuals cannot be morphologically discriminated when developing under good environmental conditions. Sub-optimal breeding conditions, in contrast, may unmask poor-quality individuals in a measurable way at the morphological level. We thus predict stronger associations between phenotype and performance under stress. In this field study, we test this hypothesis, experimentally challenging the homeostasis of great tit (Parus major) nestlings by short-term deprivation of parental care, which had no immediate effect on nestling fitness. The experiment was replicated during two subsequent breeding seasons with contrasting ambient weather conditions. Experimental (short-term) stress affected tarsus growth but not residual mass at fledging, whereas ambient (continuous) stress affected residual mass but not tarsus growth. Short-term stress effects on tarsus length and tarsus fluctuating asymmetry were only apparent when ambient conditions were unfavourable. Residual mass and hatching date, but none of the other phenotypic traits, predicted local survival, whereby the strength of the relationship did not vary between both years. Because effects of stress on develop- mental homeostasis are likely to be trait-specific and condition-dependent, studies on the use of phenotypic markers for individual fitness should integrate multiple traits comprising different levels of developmental complexity. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 103-110.

Animal dispersal and subsequent settlement is a key process in the life history of many organisms, when individuals use demographic and environmental cues to target post-dispersal habitats where fitness will be highest. To investigate the hypothesis that environmental disturbance (habitat fragmentation) may alter these cues, we compared dispersal patterns of 60 red squirrels (Sciurus vulgaris) in three study sites that differ in habitat composition and fragmentation. We determined dispersal distances, pre- and post-dispersal habitat types and survival using a combination of capturemarkrecapture, radio-tracking and genetic parentage assignment. Most (75%) squirrels emigrated from the natal home range with mean dispersal distance of 1,014 ± 925 m (range 514,118 m). There were no sex-related differences in dispersal patterns and no differences in average dispersal distance, and the proportion of dispersers did not differ between sites. In one of the sites, dispersers settled in patches where density was lower than in the natal patch. In the least fragmented site, 90% of animals settled in the natal habitat type (habitat cuing) against 4454% in the more strongly fragmented sites. Overall, more squirrels settled in the natal habitat type than expected based on habitat availability, but this was mainly due to individuals remaining within the natal wood. In the highly fragmented landscape, habitat cuing among emigrants did not occur more frequently than expected. We concluded that increased habitat fragmentation seemed to reduce reliable cues for habitat choice, but that dispersing squirrels settled in patches with lower densities of same-sex animals than at the natal home range or patch, independent of degree of fragmentation.

Polymorphisms in the dopamine receptor D4 gene (DRD4) have been related to individual variation in novelty-seeking or exploratory behaviour in a variety of animals, including humans. Recently, the human DRD4 orthologue was sequenced in a wild bird, the great tit (Parus major) and a single nucleotide polymorphism in exon 3 of this gene (SNP830) was shown to be associated with variation in exploratory behaviour of lab-raised individuals originating from a single wild population. Here we test the generality of this finding in a large sample of free-living individuals from four European great tit populations, including the originally sampled population. We demonstrate that the association between SNP830 genotype and exploratory behaviour also exists in free-living birds from the original population. However, in the other three populations we found only limited evidence for an association: in two populations the association appeared absent; while in one there was a nonsignificant tendency. We could not confirm a previously demonstrated interaction with another DRD4 polymorphism, a 15 bp indel in the promoter region (ID15). As yet unknown differences in genetic or environmental background could explain why the same genetic polymorphism (SNP830) has a substantial effect on exploratory behaviour in one population, explaining 4.5-5.8% of the total variance-a large effect for a single gene influencing a complex behavioural trait-but not in three others. The confirmation of an association between SNP830 genotype and personality-related behaviour in a wild bird population warrants further research into potential fitness effects of the polymorphism, while also the population differences in the strength of the association deserve further investigation. Another important future challenge is the identification of additional loci influencing avian personality traits in the wild.

Aim Some invasive species succeed particularly well and manage to establish populations across a wide variety of regions and climatic conditions. Understanding how biotic and environmental factors facilitate their invasion success remains a challenge. Here, we assess the role of two major hypotheses explaining invasion success: (1) enemy-release, which argues that invasive species are freed from their native predators and parasites in the new areas; and (2) climate-matching, which argues that the climatic similarity between the exotic and native range determines the success of invasive populations. Location India, Israel and the UK. Methods We studied the reproductive success of one of the most successful avian invaders, the rose-ringed parakeet (Psittacula krameri), in its native range (India) and in two introduced regions, varying in their climate conditions (Israel and the UK). We combined literature and field data to evaluate the role of predation pressure and climatic conditions in explaining the differences in reproductive success between the three regions. Results We found significant differences in reproductive success between regions. In accordance with the enemy-release hypothesis, we discovered that while predation was the main factor responsible for the reduction of fecundity in India, it did not significantly affect the fecundities of parakeet populations in the two introduced regions. In accordance with the climate-matching hypothesis, we found that in the colder temperate UK, egg infertility was high, resulting in lower fecundities. Populations in both the warmer Mediterranean climate of Israel and in the native Indian range had significantly lower egg infertility and higher fecundities than the UK populations. Main conclusions Our findings support both the enemy-release and the climate-matching hypotheses. While release from predators facilitates the reproductive success and therefore the invasiveness of parakeets in both the UK and in Israel, colder climate impedes reproduction and therefore the spread of parakeets in the UK.

When habitats become fragmented, variation in patch size and quality are expected to impose changes on the spacing pattern and social organization of animal populations. General theory predicts different possible responses including shrinking home ranges (fission response), increasing range overlap (fusion) and incorporation of multiple patches in the home range (expansion response) as fragmentation increases. We studied space use and social organization in a metapopulation of red squirrels ( Sciurus vulgaris ) in 15 woodland fragments differing in size and tree species composition. Home ranges and core areas of males were larger than females, and fragmentation had different and complex effects on the spacing pattern of both sexes. In food-supplemented patches, high densities led to increased intra-sexual overlap. In linear-shaped patches, squirrels used smaller home ranges and core areas and had lower male–male and male–female overlap levels, independent of patch quality or size. Home range and core area size of males increased with patch size, and male core areas overlapped extensively those of other males and females. Hence males seemed to show a fission response only in some patches. In contrast, home range and core area size of females was not related with patch size, but decreased with habitat quality, supporting predictions of a fusion response and intra-sexual defense of food-based core areas. Hence, where patch size and shape strongly affected space use of male red squirrels, social organization of females was only affected in small, food-supplemented patches, suggesting that the basic spatio-social organization of adult females is very resistant to fragmentation.

Summary 1 Little information is available on the ecological interactions between ticks and their hosts under natural conditions, and particularly so for avian hosts. To understand this host–parasite interaction it is necessary to assess the physiological harm ticks can do to their host. 2 We combined observational and experimental (field and laboratory) data to examine the effects of a common tick species with major economic importance, the sheep tick (Ixodes ricinus), on the health status of a common passerine bird, the great tit (Parus major). 3 In the laboratory experiment a parallel group design was carried out in which the birds of the experimental group were infested with 3–10 nymphs, whereas the birds of the control group were kept free of ticks and received a sham treatment. Both groups were stratified according to age and sex. Health parameters were measured the day before and 3 days after infestation or sham treatment: haematocrit level, erythrocyte sedimentation rate, leucocyte concentration and general body condition (body mass corrected for body size). 4 No effects of age were observed on any of the health parameters. The decrease in haematocrit level in the experimental group was significantly greater than in the control group. Moreover, infested males suffered more blood depletion than infested females. The increase in sedimentation rate was greater in the experimental group than in the control group. Surprisingly, no treatment effects were found on leucocyte concentrations, which may indicate immunoregulation by the ticks on components of the birds’ cellular immune response. Also no difference in general body condition between the treatment groups was found. None of the infested birds died during infestation. 5 Lower haematocrit levels in infested birds, but unaffected leucocyte concentrations and general body condition are confirmed by field data (experimental and observational) of adult birds during breeding season. 6 Neither haematocrit level nor general body condition was associated with parasite intensity among infested birds, suggesting that immature Ixodes ricinus are not resource limited at high natural densities. Still, the measurable direct harm caused by sheep tick infestations calls for further study on its importance for the evolutionary ecology of passerine hosts.