Search

Background: Mate choice is a fundamental element of sexual selection and has the potential to shape the evolution of traits. Mate choice based on body size has been shown to be a common trait in several arthropod species. In hard ticks, a taxon of medical and veterinary importance, engorgement weight is positively correlated with reproductive output but it is unknown whether adult males show mate choice. Here, we experimentally investigated whether males (i) use chemical cues to choose their mating partner, (ii) consistently choose for the same female individual and (iii) prefer females with highest weight after feeding., Methods: We used two experimental setups which allowed chemical communication between ticks: (i) a horizontal tube preventing physical contact with the female and (ii) an arena where tactile cues were allowed. In total, we tested 62 different triads in 124 tests (66 tests in the horizontal tube and 58 in the arena) composed of one male that could choose between two engorged females. Specifically, we tested 42 triads in the tube and 46 in the arena; 24 triads were repeatedly tested in the tube while 38 triads were tested in both setups., Results: We found no preference for individual or heavier females in either setup. However, in the horizontal tube setup, males significantly preferred females that were not visited by them in the previous test., Conclusions: Our results suggest a lack of male mate choice despite heavier females having higher fecundity. However, future studies should take into account that males may recognize the potential mating partners they previously met., (© 2022. The Author(s).)

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 co-familial European songbirds, the great tit (Parus major) and blue tit (Cyanistes caeruleus), covering a large part of their breeding range. Populations inhabiting deciduous habitats showed stronger phenological sensitivity than those in evergreen and mixed habitats. However, populations with higher sensitivity tended to have experienced less rapid change in climate over the past decades, such that populations with high phenological sensitivity will not necessarily exhibit the strongest phenological advancement. 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., (© 2022. The Author(s).)

Little is known about the intraspecific variation of parasite life-history traits and how this variation may affect parasite fitness and evolution. We investigated how life-history traits predict success of individual tree-hole ticks Ixodes arboricola and estimated their evolutionary potential, as well as genetic correlations within stages and phenotypic correlations within and across stages. Ticks were followed individually over two generations while allowed to feed on great tits Parus major. After accounting for host and tick maternal effects, we found that short feeding times and high engorgement weights strongly increased molting success. Molting time was also positively correlated with feeding success in adults. In larvae and nymphs, we found negative phenotypic correlations between engorgement weight and both feeding and molting time, the latter supported by a negative genetic correlation. We found sex-related differences in feeding time (longer in male nymphs) and molting time (longer in male larvae but shorter in male nymphs). Also, time since the last feeding event (set experimentally) reduced larval and nymphal fitness, whereas it increased adult female fitness. Furthermore, we found significant heritability and evolvability, that is, the potential to respond to selection, for engorgement weight and molting time across all stages but no significant heritability for feeding time. Our findings suggest that variation in tick fitness is shaped by consistent individual differences in tick quality, for which engorgement weight is a good proxy, rather than by life-history trade-offs., (© 2022 The Authors. Evolution published by Wiley Periodicals LLC on behalf of The Society for the Study of Evolution.)

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., Competing Interests: No competing interests were disclosed., (Copyright: © 2021 Heylen D et al.)

The integration and synthesis of the data in different areas of science is drastically slowed and hindered by a lack of standards and networking programmes. Long-term studies of individually marked animals are not an exception. These studies are especially important as instrumental for understanding evolutionary and ecological processes in the wild. Furthermore, their number and global distribution provides a unique opportunity to assess the generality of patterns and to address broad-scale global issues (e.g. climate change). To solve data integration issues and enable a new scale of ecological and evolutionary research based on long-term studies of birds, we have created the SPI-Birds Network and Database (www.spibirds.org)-a large-scale initiative that connects data from, and researchers working on, studies of wild populations of individually recognizable (usually ringed) birds. Within year and a half since the establishment, SPI-Birds has recruited over 120 members, and currently hosts data on almost 1.5 million individual birds collected in 80 populations over 2,000 cumulative years, and counting. SPI-Birds acts as a data hub and a catalogue of studied populations. It prevents data loss, secures easy data finding, use and integration and thus facilitates collaboration and synthesis. We provide community-derived data and meta-data standards and improve data integrity guided by the principles of Findable, Accessible, Interoperable and Reusable (FAIR), and aligned with the existing metadata languages (e.g. ecological meta-data language). The encouraging community involvement stems from SPI-Bird's decentralized approach: research groups retain full control over data use and their way of data management, while SPI-Birds creates tailored pipelines to convert each unique data format into a standard format. We outline the lessons learned, so that other communities (e.g. those working on other taxa) can adapt our successful model. Creating community-specific hubs (such as ours, COMADRE for animal demography, etc.) will aid much-needed large-scale ecological data integration., (© 2020 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

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.

Infection probability, load, and community structure of helminths varies strongly between and within animal populations. This can be ascribed to environmental stochasticity or due to individual characteristics of the host such as their age or sex. Other, but understudied, factors are the hosts' behavior and co-infection patterns. In this study, we used the multimammate mouse ( Mastomys natalensis ) as a model system to investigate how the hosts' sex, age, exploration behavior, and viral infection history affects their infection risk, parasitic load, and community structure of gastrointestinal helminths. We hypothesized that the hosts' exploration behavior would play a key role in the risk for infection by different gastrointestinal helminths, whereby highly explorative individuals would have a higher infection risk leading to a wider diversity of helminths and a larger load compared to less explorative individuals. Fieldwork was performed in Morogoro, Tanzania, where we trapped a total of 214 individual mice. Their exploratory behavior was characterized using a hole-board test after which we collected the helminths inside their gastrointestinal tract. During our study, we found helminths belonging to eight different genera: Hymenolepis sp., Protospirura muricola, Syphacia sp., Trichuris mastomysi, Gongylonema sp., Pterygodermatites sp., Raillietina sp., and Inermicapsifer sp. and one family: Trichostrongylidae . Hierarchical modeling of species communities (HMSC) was used to investigate the effect of the different host-related factors on the infection probability, parasite load, and community structure of these helminths. Our results show that species richness was higher in adults and in females compared to juveniles and males, respectively. Contrary to our expectations, we found that less explorative individuals had higher infection probability with different helminths resulting in a higher diversity, which could be due to a higher exposure rate to these helminths and/or behavioral modification due to the infection., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Vanden Broecke, Bernaerts, Ribas, Sluydts, Mnyone, Matthysen and Leirs.)

Heterogeneous selection is often proposed as a key mechanism maintaining repeatable behavioral variation ("animal personality") in wild populations. Previous studies largely focused on temporal variation in selection within single populations. The relative importance of spatial versus temporal variation remains unexplored, despite these processes having distinct effects on local adaptation. Using data from >3,500 great tits ( Parus major ) and 35 nest box plots situated within five West-European populations monitored over 4 to 18 y, we show that selection on exploration behavior varies primarily spatially, across populations, and study plots within populations. Exploration was, simultaneously, selectively neutral in the average population and year. These findings imply that spatial variation in selection may represent a primary mechanism maintaining animal personalities, likely promoting the evolution of local adaptation, phenotype-dependent dispersal, and nonrandom settlement. Selection also varied within populations among years, which may counteract local adaptation. Our study underlines the importance of combining multiple spatiotemporal scales in the study of behavioral adaptation., Competing Interests: The authors declare no competing interest.

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 individuals  = 201). Such a cycle consists of two phases in which the levels of intra-specific competition vary drastically. We found that only one personality trait, namely stress-sensitivity, had a negative effect on both survival and recapture probability but none of them affected maturation. This suggests that less stress-sensitive individuals take more risks in the wild and have a higher survival probability compared to high stress-sensitive individuals. However, the effect of personality on survival was only present during the population decrease phase, when the levels of intra-specific competition are high due to a scarcity of food. This suggests that seasonal changes in competition might be important in the evolution and maintenance of animal personalities in species whose population dynamics have a clear seasonal component.

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., (© The Author(s) (2020). Published by Oxford University Press on behalf of Editorial Office, Current Zoology.)

It remains poorly understood how effects of anthropogenic activity, such as large-scale habitat fragmentation, impact sociality in animals. In cooperatively breeding species, groups are mostly formed through delayed offspring dispersal, and habitat fragmentation can affect this process in two opposite directions. Increased habitat isolation may increase dispersal costs, promoting delayed dispersal. Alternatively, reduced patch size and quality may decrease benefits of philopatry, promoting dispersal. Here, we test both predictions in a cooperatively breeding bird (placid greenbul, Phyllastrephus placidus ) from an Afrotropical cloud forest archipelago. Males born in fragmented forest dispersed about 1 year earlier than those born in continuous forest. Contrary to females, males also started to reproduce earlier and mostly settled within their natal patch. Females only rarely delayed their dispersal for more than 1 year, both in fragmented and continuous forests. Our results suggest that early male dispersal and reproduction is jointly driven by a decrease in the value of the natal territory and an increase in local breeding opportunities in fragmented forest. While plasticity in dispersal strategies of cooperative breeders in response to anthropogenic change is believed to optimize reproduction-survival trade-offs, to what extent it shapes the ability of species to respond to rapid environmental change remains to be studied.

Candidatus Rickettsia vini was originally detected in Ixodes arboricola ticks from Spain, and subsequently reported from several other Western Palearctic countries including Belgium. Recently, the bacterium was isolated in mammalian (Vero) cell culture from macerated male I. arboricola from Czech Republic, but there have been no reports of propagation in tick cells. Here we report isolation in a tick cell line of three strains of Ca. R. vini from I. arboricola collected from nests of great tits (Parus major) in Belgium. Internal organs of one male and two engorged female ticks were dissected aseptically, added to cultures of the Rhipicephalus microplus cell line BME/CTVM23 and incubated at 28 °C. Rickettsia-like bacteria were first seen in Giemsa-stained cytocentrifuge smears between 2 and 15 weeks later. Two of the isolates grew rapidly, destroying the tick cells within 2-4 weeks of onward passage in BME/CTVM23 cells, while the third isolate grew much more slowly, only requiring subculture at 4-5-month intervals. PCR amplification of bacterial 16S rRNA and Rickettsia gltA, sca4, ompB, ompA and 17-kDa genes revealed that all three isolates were Ca. R. vini, with 100 % identity to each other and to published Ca. R. vini sequences from other geographical locations. Transmission electron microscopy revealed typical single Rickettsia bacteria in the cytoplasm of BME/CTVM23 cells. The Ca. R. vini strain isolated from the male I. arboricola tick, designated Boshoek1, was tested for ability to grow in a panel of Ixodes ricinus, Ixodes scapularis and R. microplus cell lines and in Vero cells. The Boshoek1 strain grew rapidly, causing severe cytopathic effect, in the R. microplus line BME26, the I. ricinus line IRE11 and Vero cells, more slowly in the I. ricinus line IRE/CTVM19, possibly established a low-level infection in the I. ricinus line IRE/CTVM20, and failed to infect cells of any of four I. scapularis lines over a 12-week observation period. This study confirmed the applicability of the simple tick organ-cell line co-cultivation technique for isolation of tick-borne Rickettsia spp. using BME/CTVM23 cells., (Copyright © 2020 The Authors. Published by Elsevier GmbH.. All rights reserved.)

We explored the inter-individual variability in bud-burst and its potential drivers, in homogeneous mature stands of temperate deciduous trees. Phenological observations of leaves and wood formation were performed weekly from summer 2017 to summer 2018 for pedunculate oak, European beech and silver birch in Belgium. The variability of bud-burst was correlated to previous' year autumn phenology (i.e. the onset of leaf senescence and the cessation of wood formation) and tree size but with important differences among species. In fact, variability of bud-burst was primarily related to onset of leaf senescence, cessation of wood formation and tree height for oak, beech and birch, respectively. The inter-individual variability of onset of leaf senescence was not related to the tree characteristics considered and was much larger than the inter-individual variability in bud-burst. Multi-species multivariate models could explain up to 66% of the bud-burst variability. These findings represent an important advance in our fundamental understanding and modelling of phenology and tree functioning of deciduous tree species., Competing Interests: The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (© 2020 The Author(s).)

Sustainable land-use management must account for the potential trade-offs between biodiversity conservation, productive land uses and ecosystem services. In this study, we used Marxan with Zones to generate land use plans that optimize conservation, farming and forestry land uses to reach biodiversity targets while minimizing the opportunity cost for local communities in an inhabited but data-poor National Park in the Andes of Bolivia. Based on six alternative land-use plans, we identified the synergies and trade-offs between the biodiversity benefits achieved in the different plans and the delivery of four locally important water-related ecosystem services modeled with the web-based tool AguAAndes. Although we find synergies between the conservation of high altitude Polylepis woodlands and their associated avifauna and three of the ecosystem services investigated, soil erosion levels were actually higher in scenarios with higher achieved biodiversity benefits. Our study shows how systematic conservation planning and ecosystem service delivery modelling can be used to solve land-use conflicts and identify trade-offs between biodiversity conservation and ecosystem services in a data-poor tropical area.

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.

The increasing urbanization process is hypothesized to drastically alter (semi-)natural environments with a concomitant major decline in species abundance and diversity. Yet, studies on this effect of urbanization, and the spatial scale at which it acts, are at present inconclusive due to the large heterogeneity in taxonomic groups and spatial scales at which this relationship has been investigated among studies. Comprehensive studies analysing this relationship across multiple animal groups and at multiple spatial scales are rare, hampering the assessment of how biodiversity generally responds to urbanization. We studied aquatic (cladocerans), limno-terrestrial (bdelloid rotifers) and terrestrial (butterflies, ground beetles, ground- and web spiders, macro-moths, orthopterans and snails) invertebrate groups using a hierarchical spatial design, wherein three local-scale (200 m × 200 m) urbanization levels were repeatedly sampled across three landscape-scale (3 km × 3 km) urbanization levels. We tested for local and landscape urbanization effects on abundance and species richness of each group, whereby total richness was partitioned into the average richness of local communities and the richness due to variation among local communities. Abundances of the terrestrial active dispersers declined in response to local urbanization, with reductions up to 85% for butterflies, while passive dispersers did not show any clear trend. Species richness also declined with increasing levels of urbanization, but responses were highly heterogeneous among the different groups with respect to the richness component and the spatial scale at which urbanization impacts richness. Depending on the group, species richness declined due to biotic homogenization and/or local species loss. This resulted in an overall decrease in total richness across groups in urban areas. These results provide strong support to the general negative impact of urbanization on abundance and species richness within habitat patches and highlight the importance of considering multiple spatial scales and taxa to assess the impacts of urbanization on biodiversity., (© 2019 John Wiley & Sons Ltd.)

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 α- and β-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.

Mechanisms of on-host habitat selection of parasites are important to the understanding of host-parasite interactions and evolution. To this end, it is important to separate the factors driving parasite micro-habitat selection from those resulting from host anti-parasite behaviour. We experimentally investigated whether tick infestation patterns on songbirds are the result of an active choice by the ticks themselves, or the outcome of songbird grooming behaviour. Attachment patterns of three ixodid tick species with different ecologies and host specificities were studied on avian hosts. Ixodes arboricola, Ixodes ricinus and Ixodes frontalis were put on the head, belly and back of adult great tits (Parus major) and adult domestic canaries (Serinus canaria domestica) which were either restricted or not in their grooming capabilities. Without exception, ticks were eventually found on a bird's head. When we gave ticks full opportunities to attach on other body parts - in the absence of host grooming - they showed lower attachment success. Moreover, ticks moved from these other body parts to the host's head when given the opportunity. This study provides evidence that the commonly observed pattern of ticks feeding on songbirds' heads is the result of an adaptive behavioural strategy. Experimental data on a novel host species, the domestic canary, and a consistent number of published field observations, strongly support this hypothesis. We address some proximate and ultimate causes that may explain parasite preference for this body part in songbirds. The link found between parasite micro-habitat preference and host anti-parasite behaviour provides further insight into the mechanisms driving ectoparasite aggregation, which is important for the population dynamics of hosts, ectoparasites and the micro-pathogens for which they are vectors., (Copyright © 2019 Australian Society for Parasitology. All rights reserved.)

Conspecific density and animal personality (consistent among-individual differences in behavior) may both play an important role in disease ecology. Nevertheless, both factors have rarely been studied together but may provide insightful information in understanding pathogen transmission dynamics. In this study, we investigated how both personality and density affect viral infections both direct and indirectly, using the multimammate mice ( Mastomys natalensis ) and Morogoro arenavirus (MORV) as a model system. Using a replicated semi-natural experiment, we found a positive correlation between MORV antibody presence and density, suggesting that MORV infection is density-dependent. Surprisingly, slower explorers were more likely to have antibodies against MORV compared to highly explorative individuals. However, exploration was positively correlated with density which may suggest a negative, indirect effect of density on MORV infection. We have shown here that in order to better understand disease ecology, both personality and density should be taken into account., (© 2019 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Biological responses to climate change have been widely documented across taxa and regions, but it remains unclear whether species are maintaining a good match between phenotype and environment, i.e. whether observed trait changes are adaptive. Here we reviewed 10,090 abstracts and extracted data from 71 studies reported in 58 relevant publications, to assess quantitatively whether phenotypic trait changes associated with climate change are adaptive in animals. A meta-analysis focussing on birds, the taxon best represented in our dataset, suggests that global warming has not systematically affected morphological traits, but has advanced phenological traits. We demonstrate that these advances are adaptive for some species, but imperfect as evidenced by the observed consistent selection for earlier timing. Application of a theoretical model indicates that the evolutionary load imposed by incomplete adaptive responses to ongoing climate change may already be threatening the persistence of species.

Many vector-borne diseases are transmitted through complex pathogen-vector-host networks, which makes it challenging to identify the role of specific host groups in disease emergence. Lyme borreliosis in humans is now the most common vector-borne zoonosis in the Northern Hemisphere. The disease is caused by multiple genospecies of Borrelia burgdorferi sensu lato bacteria transmitted by ixodid (hard) ticks, and the major host groups transmit Borrelia genospecies with different pathogenicity, causing variable clinical symptoms in humans. The health impact of a given host group is a function of the number of ticks it infects as well as the pathogenicity of the genospecies it carries. Borrelia afzelii, with mainly small mammals as reservoirs, is the most common pathogen causing Lyme borreliosis, and it is often responsible for the largest proportion of infected host-seeking tick nymphs in Europe. The bird-borne Borrelia garinii, though less prevalent in nymphal ticks, is more likely to cause Lyme neuroborreliosis, but whether B. garinii causes disseminated disease more frequently has not been documented. Based on extensive data of annual disease incidence across Norway from 1995 to 2017, we show here that 69% of disseminated Lyme borreliosis cases were neuroborreliosis, which is three times higher than predicted from the infection prevalence of B. garinii in host-seeking ticks (21%). The population estimate of migratory birds, mainly of thrushes, explained part of the annual variation in cases of neuroborreliosis, with a one-year time lag. We highlight the important role of the genospecies' pathogenicity and the host associations for understanding the epidemiology of disseminated Lyme borreliosis.

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., (Copyright © 2018 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Despite the increasing knowledge on the processes involved in the acquisition and development of the gut microbiota in model organisms, the factors influencing early microbiota successions in natural populations remain poorly understood. In particular, little is known on the role of the rearing environment in the establishment of the gut microbiota in wild birds. Here, we examined the influence of the nesting environment on the gut microbiota of Great tits ( Parus major ) by performing a partial cross-fostering experiment during the intermediate stage of nestling development. We found that the cloacal microbiota of great tit nestlings underwent substantial changes between 8 and 15 days of age, with a strong decrease in diversity, an increase in the relative abundance of Firmicutes and a shift in the functional features of the community. Second, the nesting environment significantly influenced community composition, with a divergence among separated true siblings and a convergence among foster siblings. Third, larger shifts in both microbiota diversity and composition correlated with lower nestling body condition. Our results shed new light on the dynamics of microbial diversity during the ontogeny of avian hosts, indicating that the nest environment continues to shape the gut microbiota during the later stages of nestling development and that the increase in gut diversity between hatching and adulthood may not be as linear as previously suspected. Lastly, the microbiota changes incurred during this period may have implications for nestling body condition which can lead to long-term consequences for host fitness.

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., (Copyright © 2018 Elsevier GmbH. All rights reserved.)

Body size is intrinsically linked to metabolic rate and life-history traits, and is a crucial determinant of food webs and community dynamics 1,2 . The increased temperatures associated with the urban-heat-island effect result in increased metabolic costs and are expected to drive shifts to smaller body sizes 3 . Urban environments are, however, also characterized by substantial habitat fragmentation 4 , which favours mobile species. Here, using a replicated, spatially nested sampling design across ten animal taxonomic groups, we show that urban communities generally consist of smaller species. In addition, although we show urban warming for three habitat types and associated reduced community-weighted mean body sizes for four taxa, three taxa display a shift to larger species along the urbanization gradients. Our results show that the general trend towards smaller-sized species is overruled by filtering for larger species when there is positive covariation between size and dispersal, a process that can mitigate the low connectivity of ecological resources in urban settings 5 . We thus demonstrate that the urban-heat-island effect and urban habitat fragmentation are associated with contrasting community-level shifts in body size that critically depend on the association between body size and dispersal. Because body size determines the structure and dynamics of ecological networks 1 , such shifts may affect urban ecosystem function.

The human pathogens Borrelia afzelii, which causes Lyme borreliosis and B. miyamotoi, which causes relapsing fever, both circulate between Ixodes ricinus ticks and rodents. The spatiotemporal dynamics in the prevalence of these pathogens have not yet been fully elucidated, but probably depend on the spatiotemporal population dynamics of small rodents. We aimed to evaluate the effect of different forest types on the density of infected nymphs in different years and to obtain more knowledge about the spatial and temporal patterns of ticks and tick-borne pathogens. We analysed unfed nymphal ticks from 22 stands of four different forest types in Belgium in 2009, 2010, 2013 and 2014 and found that the density of nymphs in general and the density of nymphs infected with B. afzelii and B. miyamotoi varied yearly, but without temporal variation in the infection prevalence. The yearly variation in density of infected nymphs in our study thus seems to be caused most by the variation in the density of nymphs, which makes it a good predictor of disease risk. The risk for rodent-associated tick-borne diseases also varied between forest types. We stress the need to elucidate the contribution of the host community composition to tick-borne disease risk., (Copyright © 2017 Elsevier GmbH. All rights reserved.)

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., (© 2017 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

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., (Copyright © 2017 Elsevier B.V. All rights reserved.)

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., (© 2016 Cambridge Philosophical Society.)

For avian group living to be evolutionary stable, multiple fitness benefits are expected. Yet, the difficulty of tracking fledglings, and thus estimating their survival rates, limits our knowledge on how such benefits may manifest postfledging. We radio-tagged breeding females of the Afrotropical cooperatively breeding Placid greenbul ( Phyllastrephus placidus) during nesting. Tracking these females after fledging permitted us to locate juvenile birds, their parents, and any helpers present and to build individual fledgling resighting datasets without incurring mortality costs or causing premature fledging due to handling or transmitter effects. A Bayesian framework was used to infer age-specific mortality rates in relation to group size, fledging date, maternal condition, and nestling condition. Postfledging survival was positively related to group size, with fledglings raised in groups with four helpers showing nearly 30% higher survival until independence compared with pair-only offspring, independent of fledging date, maternal condition or nestling condition. Our results demonstrate the importance of studying the early dependency period just after fledging when assessing presumed benefits of cooperative breeding. While studying small, mobile organisms after they leave the nest remains highly challenging, we argue that the telemetric approach proposed here may be a broadly applicable method to obtain unbiased estimates of postfledging survival.

Lyme disease is caused by bacteria of the Borrelia burgdorferi genospecies 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 analysed Borrelia infection 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 harbour Borrelia afzelii infection more often in pine stands while Borrelia garinii and Borrelia burgdorferi ss. 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.

As biodiversity hotspots are often characterized by high human population densities, implementation of conservation management practices that focus only on the protection and enlargement of pristine habitats is potentially unrealistic. An alternative approach to curb species extinction risk involves improving connectivity among existing habitat patches. However, evaluation of spatially explicit management strategies is challenging, as predictive models must account for the process of dispersal, which is difficult in terms of both empirical data collection and modelling.Here, we use a novel, individual-based modelling platform that couples demographic and mechanistic dispersal models to evaluate the effectiveness of realistic management scenarios tailored to conserve forest birds in a highly fragmented biodiversity hotspot. Scenario performance is evaluated based on the spatial population dynamics of a well-studied forest bird species.The largest population increase was predicted to occur under scenarios increasing habitat area. However, the effectiveness was sensitive to spatial planning. Compared to adding one large patch to the habitat network, adding several small patches yielded mixed benefits: although overall population sizes increased, specific newly created patches acted as dispersal sinks, which compromised population persistence in some existing patches. Increasing matrix connectivity by the creation of stepping stones is likely to result in enhanced dispersal success and occupancy of smaller patches. Synthesis and applications . We show that the effectiveness of spatial management is strongly driven by patterns of individual dispersal across landscapes. For species conservation planning, we advocate the use of models that incorporate adequate realism in demography and, particularly, in dispersal behaviours.

Multiple mating is common among ticks, a large group of haematophagous ectoparasites, but multiple paternity has rarely been investigated. Multiple paternity may be common because the resultant increased genetic diversity allows ticks to rapidly evolve in relation to host responses and increases colonisation potential in novel habitats. Knowledge concerning mating systems is important because ticks may have profound effects on their hosts and are the principal transmitters of many pathogenic agents. In the current study, we investigated the mating system of the nidicolous tick Ixodes arboricola. These ticks attach to their bird hosts in the nest, which restricts gene flow but facilitates finding a partner off-host. Having genetically variable offspring may be beneficial for ticks which may encounter very different conditions when dispersed to the nest of another host type. We conducted an experiment in which female ticks fed on great tit nestlings and mated with two males in three treatments of the females: mating with both males before feeding, mating with one male before and the other male after feeding, or mating with both males after feeding. We investigated paternity with microsatellites. In a complementary experiment we investigated male preference for unfed or engorged females, and measured mating duration. We predicted (i) there would be multiple mating by I. arboricola males and females, leading to multiple paternity, and (ii) males would prefer to mate with engorged females and those matings would last longer because engorged females present a higher probability of successful reproduction. We found multiple paternity within clutches but no indications of sperm precedence. Males preferred to mate with engorged females and those matings lasted significantly longer, even including attachment beyond egg deposition. We suggest such mate guarding and male preference for mating after feeding is adaptive because there is no first male precedence. Male preference for mating after feeding may also be adaptive because dispersal is low and females are available after the blood meal., (Copyright © 2016 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

Similar to many other parasites, the distribution of ticks among hosts is strongly skewed, with few hosts harbouring the majority of parasites. Because parasite-induced impairment of host health, parasite population growth and pathogen transmission are density-dependent, understanding why tick distributions are skewed is important for the population and evolutionary dynamics of both parasite and host. However, there is currently no knowledge concerning parasites that strongly depend on individual hosts. Here, we investigated the effects of tick density on feeding performance in the nidicolous tree-hole tick, Ixodes arboricola, which feeds on cavity-nesting birds and is the carrier of several tick-borne pathogens. Nidicolous ticks reside in or close to their hosts' nests and therefore depend strongly on individual hosts and their offspring. Increased feeding success at higher densities (facilitation) may therefore be detrimental to the ticks themselves. We investigated the effects of tick density on feeding performance of I. arboricola by infesting great tit nestlings with one to five adult ticks, which is within the natural range. There was no effect of tick density on initial attachment success, attachment after 48 h or engorgement weight, but tick recovery rates increased significantly with tick density. We also found a modest increase in nestling body mass with tick density, suggesting that birds over-compensate resource drainage by the ticks and, by doing this, anticipate the costs of a tick-rich environment. Our results indicate that nidicolous ticks perform better when feeding in aggregation. This may have important consequences for population dynamics and consequently pathogen transmission., (Copyright © 2015 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.)

Lyme disease cases caused by Borrelia burgdorferi s.l. bacteria is increasing steadily in Europe, in part due to the expansion of the vector, Ixodes ricinus. Wild reservoir hosts are typically recurrently infested. Understanding the impact of these cumulative parasite exposures on the host's health is, therefore, central to predict the distribution of tick populations and their pathogens. Here, we have experimentally investigated the symptoms of disease caused by recurrent infestations in a common songbird (Parus major). Birds were exposed three times in succession to ticks collected in a Borrelia endemic area. Health and immune measures were analyzed in order to investigate changes in response to tick infestation and Borrelia infection rate. Nitric oxide levels increased with the Borrelia infection rate, but this effect was increasingly counteracted by mounting tick infestation rates. Tick infestations equally reduced haematocrit during each cycle. But birds overcompensated in their response to tick feeding, having higher haematocrit values during tick-free periods depending on the number of ticks they had been previously exposed to. Body condition showed a similar overshooting response in function of the severity of the Borrelia infection. The observed overcompensation increases the bird's energetic needs, which may result in an increase in transmission events.

Conservation actions often focus on restoration or creation of natural areas designed to facilitate the movements of organisms among populations. To be efficient, these actions need to be based on reliable estimates or predictions of landscape connectivity. While circuit theory and least-cost paths (LCPs) are increasingly being used to estimate connectivity, these methods also have proven limitations. We compared their performance in predicting genetic connectivity with that of an alternative approach based on a simple, individual-based "stochastic movement simulator" (SMS). SMS predicts dispersal of organisms using the same landscape representation as LCPs and circuit theory-based estimates (i.e., a cost surface), while relaxing key LCP assumptions, namely individual omniscience of the landscape (by incorporating perceptual range) and the optimality of individual movements (by including stochasticity in simulated movements). The performance of the three estimators was assessed by the degree to which they correlated with genetic estimates of connectivity in two species with contrasting movement abilities (Cabanis's Greenbul, an Afrotropical forest bird species, and natterjack toad, an amphibian restricted to European sandy and heathland areas). For both species, the correlation between dispersal model and genetic data was substantially higher when SMS was used. Importantly, the results also demonstrate that the improvement gained by using SMS is robust both to variation in spatial resolution of the landscape and to uncertainty in the perceptual range model parameter. Integration of this individual-based approach with other developing methods in the field of connectivity research, such as graph theory, can yield rapid progress towards more robust connectivity indices and more effective recommendations for land management.

Invasive species present a major threat to global biodiversity. Understanding genetic patterns and evolutionary processes that reinforce successful establishment is paramount for elucidating mechanisms underlying biological invasions. Among birds, the ring-necked parakeet (Psittacula krameri) is one of the most successful invasive species, established in over 35 countries. However, little is known about the evolutionary genetic origins of this species and what population genetic signatures tell us about patterns of invasion. We reveal the ancestral origins of populations across the invasive range and explore the potential influence of climate and propagule pressure from the pet trade on observed genetic patterns. Ring-necked parakeet samples representing the ancestral native range (n = 96) were collected from museum specimens, and modern samples from the invasive range (n = 855) were gathered from across Europe, Mauritius and Seychelles, and sequenced for two mitochondrial DNA markers comprising 868 bp of cytochrome b and control region, and genotyped at 10 microsatellite loci. Invasive populations comprise birds that originate predominantly from Pakistan and northern areas of India. Haplotypes associated with more northerly distribution limits in the ancestral native range were more prevalent in invasive populations in Europe, and the predominance of Asian haplotypes in Europe is consistent with the higher number of Asian birds transported by the pet trade outside the native range. Successful establishment of invasive species is likely to be underpinned by a combination of environmental and anthropogenic influences., (© 2015 John Wiley & Sons Ltd.)

Climate change is expected to have profound ecological effects, yet shifts in competitive abilities among species are rarely studied in this context. Blue tits (Cyanistes caeruleus) and great tits (Parus major) compete for food and roosting sites, yet coexist across much of their range. Climate change might thus change the competitive relationships and coexistence between these two species. Analysing four of the highest-quality, long-term datasets available on these species across Europe, we extend the textbook example of coexistence between competing species to include the dynamic effects of long-term climate variation. Using threshold time-series statistical modelling, we demonstrate that long-term climate variation affects species demography through different influences on density-dependent and density-independent processes. The competitive interaction between blue tits and great tits has shifted in one of the studied sites, creating conditions that alter the relative equilibrium densities between the two species, potentially disrupting long-term coexistence. Our analyses show that long-term climate change can, but does not always, generate local differences in the equilibrium conditions of spatially structured species assemblages. We demonstrate how long-term data can be used to better understand whether (and how), for instance, climate change might change the relationships between coexisting species. However, the studied populations are rather robust against competitive exclusion., (© 2015 The Author(s) Published by the Royal Society. All rights reserved.)

Host specialisation in parasites can be due to either limited exposure or limited adaptation to different host types. When the first barrier is lifted experimentally, the degree of adaptive specialisation can be studied. The tree-hole tick Ixodes arboricola is an endophilic parasite with a narrow host range, found in nest boxes used mainly by great and blue tits (Parus major, Cyanistes caeruleus) and to a lesser extent by pied flycatchers (Ficedula hypoleuca) and nuthatches (Sitta europaea). In the current study, we exposed two nestlings per nestbox of pied flycatchers (N = 14), blue tits (N = 18), great tits (N = 14), and nuthatches (N = 16) to ten I. arboricola nymphs each. We found no differences in attachment success 2 days after infestation (56 ± 4% across species) nor were there any differences in tick engorgement weight (1.95 ± 0.03 mg across species), and moulting success was >90% for ticks from all bird species. Hence, our data suggest that all bird species investigated here are suitable host species. This may enhance the ticks' chances for persistence in cavities and dispersal among cavities inhabited by multiple host species, and supports the hypothesis that host use by ticks is limited by host ecology rather than by host specialisation.

Investigation of endo-macroparasite infections in living animals relies mostly on indirect methods aimed to detect parasite eggs in hosts' faeces. However, faecal flotation does not provide quantitative information on parasite loads, whereas faecal egg count (FEC) techniques may not give reliable estimates of parasite intensity, since egg production may be affected by density-dependent effects on helminth fecundity. We addressed this issue using Eastern grey squirrels (Sciurus carolinensis) and their gastrointestinal nematode Strongyloides robustus to assess the performance of coprological techniques and to investigate factors affecting parasite fecundity. We compared results of gut examination, flotation and McMaster FECs in 65 culled grey squirrels. Sensitivity and specificity of flotation were 81.2% (Confidence Interval, CI 54.3-95.9%) and 85.7% (CI 72.7-94.1%), respectively, resulting in low positive predictive values when infection prevalence is low. Individual parasite fecundity (no. of eggs/adult female worm) was negatively affected by S. robustus intensity, leading to a non-linear relationship between parasite load and eggs/gram of faeces (EPG). As a consequence, whereas flotation may be a valid method to perform the first screening of infection status, FECs are not a reliable method to estimate S. robustus intensity, since diverse values of EPG may correspond to the same number of parasites. Neither the amount of analysed faeces nor the season had any effect on EPG, indicating that the observed reduction in helminth fecundity is likely caused exclusively by density-dependent processes such as competition among worms or host immune response.

We investigated to what extent a European songbird (Parus major) selectively transmits and amplifies Borrelia burgdorferi s.l. bacteria. Borrelia-naïve 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., (© 2013 Society for Applied Microbiology and John Wiley & Sons Ltd.)

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., (© 2013 Society for Applied Microbiology and John Wiley & Sons Ltd.)

While the importance of personality in explaining individual dispersal strategies is increasingly recognized, limited information is still available on how patterns of personality-dependent dispersal may develop, hampering our understanding of the ecological significance of behavioural dispersal syndromes. Here, we examine the relative importance of personality at different stages of dispersal in the great tit, by analysing the sex-specific relationship between exploratory behaviour (EB; quantified in a novel environment) and dispersal distances in different seasons over the course of the first year of life (summer, autumn, winter, and until the first breeding attempt). In females, we found that EB was an important predictor of dispersal distances in summer and autumn, but only a weak to moderate correlation remained for females captured in winter or for natal dispersal distances based on first breeding records. We obtained a contrasting pattern at the population level, whereby male (but not female) immigrants captured in summer and autumn had higher EB scores than locally born birds, while this was not the case in birds captured in winter and those recruited as breeders into the population. In addition to providing further evidence for the existence of a behaviour dispersal syndrome in birds, our results show that correlations between EB and dispersal appear strongest at the early stages of the dispersal process, rather than being developed gradually. These findings show the importance of analysing the effect of phenotypic attributes on dispersal across different stages of the dispersal phenomenon and in each sex separately.

Background: Climate change potentially has important effects on distribution, abundance, transmission and virulence of parasites in wild populations of animals., Methodology/principal Finding: Here we analyzed paired information on 89 parasite populations for 24 species of bird hosts some years ago and again in 2010 with an average interval of 10 years. The parasite taxa included protozoa, feather parasites, diptera, ticks, mites and fleas. We investigated whether change in abundance and prevalence of parasites was related to change in body condition, reproduction and population size of hosts. We conducted analyses based on the entire dataset, but also on a restricted dataset with intervals between study years being 5-15 years. Parasite abundance increased over time when restricting the analyses to datasets with an interval of 5-15 years, with no significant effect of changes in temperature at the time of breeding among study sites. Changes in host body condition and clutch size were related to change in temperature between first and second study year. In addition, changes in clutch size, brood size and body condition of hosts were correlated with change in abundance of parasites. Finally, changes in population size of hosts were not significantly related to changes in abundance of parasites or their prevalence., Conclusions/significance: Climate change is associated with a general increase in parasite abundance. Variation in laying date depended on locality and was associated with latitude while body condition of hosts was associated with a change in temperature. Because clutch size, brood size and body condition were associated with change in parasitism, these results suggest that parasites, perhaps mediated through the indirect effects of temperature, may affect fecundity and condition of their hosts. The conclusions were particularly in accordance with predictions when the restricted dataset with intervals of 5-15 years was used, suggesting that short intervals may bias findings.

Although bird-tick systems affect the human risk of tick-borne diseases, very little is known about the ecological factors that shape the spatio-temporal variation of tick infestations in terrestrial songbirds. We present a risk model that explains the levels of infestation of Ixodes ricinus, the main vector of Borrelia burgdorferi s.l., during the breeding season of the great tit (Parus major), one of Europe's most abundant avian reservoir hosts of B. burgdorferi s.l. Tit tick burden were modelled as a function of variables summarising vegetation, climate, proxies for mammal abundance and characteristics of individual birds and their nests. Tick loads were positively associated with the relative humidity prior to capture of the bird and the cover of bracken inside its territory. The number of cold winter days prior to the bird's breeding season showed a negative association with tick loads. None of the proxies for mammal abundance correlated with tick loads. Tick loads decreased with age in female tits, whereas they increased with age in male tits. Tick burdens in the parental tits were positively associated with their brood size and negatively correlated with the average nestling body weight. Possible mechanisms include: how tit foraging influences tick encounter rates, host tick resistance mechanisms and the environmental conditions that simultaneously affect tick exposure risk and brood characteristics. We believe this study provides the first detailed insights into the ecological factors that shape tick burden in a terrestrial songbird., (Copyright © 2013 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.)

The assessment of genetic architecture and selection history in genes for behavioural traits is fundamental to our understanding of how these traits evolve. The dopamine receptor D4 (DRD4) gene is a prime candidate for explaining genetic variation in novelty seeking behaviour, a commonly assayed personality trait in animals. Previously, we showed that a single nucleotide polymorphism in exon 3 of this gene is associated with exploratory behaviour in at least one of four Western European great tit (Parus major) populations. These heterogeneous association results were explained by potential variable linkage disequilibrium (LD) patterns between this marker and the causal variant or by other genetic or environmental differences among the populations. Different adaptive histories are further hypothesized to have contributed to these population differences. Here, we genotyped 98 polymorphisms of the complete DRD4 gene including the flanking regions for 595 individuals of the four populations. We show that the LD structure, specifically around the original exon 3 SNP is conserved across the four populations and does not explain the heterogeneous association results. Study-wide significant associations with exploratory behaviour were detected in more than one haplotype block around exon 2, 3 and 4 in two of the four tested populations with different allele effect models. This indicates genetic heterogeneity in the association between multiple DRD4 polymorphisms and exploratory behaviour across populations. The association signals were in or close to regions with signatures of positive selection. We therefore hypothesize that variation in exploratory and other dopamine-related behaviour evolves locally by occasional adaptive shifts in the frequency of underlying genetic variants., (© 2013 Blackwell Publishing Ltd.)

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., (© 2012 Society for Applied Microbiology and Blackwell Publishing Ltd.)

One aspect of animal personality that has been well described in captivity, but received only little attention in studies in the wild, is that personality types may vary in their behavioural flexibility towards environmental changes. A fundamental factor underlying such differences is believed to be the degree to which individual behavior is guided by environmental stimuli. We tested this hypothesis in the wild using free-ranging great tits. Personality variation was quantified using exploratory behaviour in a novel environment, which has previously been shown to be repeatable and correlated with other behaviours in this and other populations of the same species. By temporarily removing food at feeding stations we examined whether birds with different personality differed in returning to visit empty feeders as this may provide information on how birds continue to sample their environment after a sudden change in conditions. In two summer experiments, we found that fast-exploring juveniles visited empty feeders less often compared to slow-exploring juveniles. In winter, sampling behaviour was sex dependent but not related to personality. In both seasons, we found that birds who sampled empty feeders more often were more likely to rediscover food after we again re-baited the feeding stations, but there was no effect of personality. Our results show that personality types may indeed differ in ways of collecting environmental information, which is consistent with the view of personalities as different styles of coping with environmental changes. The adaptive value of these alternative behavioural tactics, however, needs to be further explored.

Dispersal is a major determinant of the ecological and evolutionary dynamics of natural populations. Individuals differ greatly in the likelihood and distance of dispersal, but it is generally unclear to what extent intrinsic, possibly genetic, differences contribute. Here we present the first explicit empirical evidence for genetic coupling of local dispersal and exploratory behaviour, a key 'animal personality' trait. Using relatedness data from a multi-generation pedigree of free-living great tits (Parus major), we find quantitative genetic variation for both the distance of local dispersal within our study area and the rate at which individuals explore a novel environment. Moreover, we find a strongly positive genetic correlation between local dispersal distance and exploration rate, despite a weak and non-significant phenotypic correlation. These findings demonstrate a potentially important behavioural mechanism underlying heritable differences in local dispersal and highlight the potential for concerted evolution of dispersal and animal personality in response to selection.