European Bird Species Research Articles

Life‐history characteristics of European birds

AbstractMotivationBirds are an extremely diverse group in terms of adaptations to environmental conditions, which is reflected in their life histories and ecological traits. Recently, functional aspects of avian diversity have been used frequently in comparative analyses as well as in community ecology studies; thus, open access to complete datasets of traits will be valuable. We focused on European bird species and compiled information about crucial ecological traits. This dataset is thus useful for research investigating large‐scale patterns in European avifauna and can be used in various analyses in evolutionary ecology, macroecology or conservation biology.Main types of variables containedWe chose several types of avian traits, such as morphological (e.g., weight, wing, bill or tarsus length), reproductive (e.g., clutch size, egg mass, incubation period or type of young) and behavioural traits (type of nest, mating system or type of parental care), dietary (e.g., folivore, granivore, insectivore or carnivore) and habitat preferences (e.g., deciduous/coniferous forest, reed or grassland).Spatial location and grainEurope; all breeding bird species (n = 499).Major taxa and level of measurementIn total, we created a dataset for 499 bird species breeding in Europe and 34 key life‐history traits represented in 85 variables. As a primary source of information we used the comprehensive handbook The birds of the Western Palearctic. The traits provide information about species‐specific mean values. We did not record values for different geographical subspecies (i.e., the trait value always represents the average for the whole breeding area of a particular species).Software formatThe data file is in ASCII text, tab delimited, not compressed.

Wing morphology, flight type and migration distance predict accumulated fuel load in birds.

Birds often accumulate large fat and protein reserves to fuel long-distance flights. While it is well known that species that fly the longest accumulate the largest amounts of fuel, considerable cross-species variation in fuel load is seen after controlling for overall migration distance. It remains unclear whether this variation can be explained by aerodynamic attributes of different species, despite obvious ecological and conservation implications. Here, we collected data on wing morphology, flight type, migration distance and fuel load from 213 European bird species and explored three questions: (1) does maximum fuel load relate to migration distance across species?; (2) does wing morphology, as described by wing aspect ratio and wing loading, influence maximum fuel load?; and (3) does flight type influence maximum fuel load? Our results indicate that maximum fuel load increases with migration across species, but residual variance is high. The latter variance is explained by aspect ratio and flight type, while wing loading and body mass explain little variance. Birds with slender wings accumulate less fuel than species with low wing aspect ratio when covering a similar migration distance. Continuously flapping species accumulate the largest amounts of fuel, followed by flapping and soaring species and flapping and gliding species, while the smallest fuel loads were observed in birds with passerine-type flight. These results highlight complex eco-evolutionary adaptations to migratory behaviour, pointing toward the importance of energy minimisation.

Continent-wide test of the efficiency of the European union’s conservation legislation in delivering population benefits for bird species

Birds are among the most important organisms for indicating the state of environmental health and their population changes may be thus informative for assessments of country-level conservation tools. One such tool applied in the European Union (EU) is the Birds Directive which (together with general protection of all bird species) lists a number of species under its Annex I and these species enjoy specific protection conditions. Although some previous studies found indications of the efficiency of the Annex I in delivering benefits for the listed species, the assessments where either confined to the so called old member states (i.e. countries entered EU before 2004) or did not include countries outside EU as a suitable control group. Therefore, it remains unclear whether this tool is efficient also in the new EU-member states (i.e. countries entered EU from 2004 onwards). For this purpose, we used publicly available information source and assembled a dataset providing country-level population trends of 252 European breeding bird species estimated for the time period 2001–2012 in 33 European countries containing old member states, new member states and non-member states. We predicted that if efficient, then listing the species under Annex I would result in significantly positive population trends of the listed species in EU countries irrespective to the time of their enterance, while no such pattern should be observed in non-EU countries. We tested this prediction using linear mixed effect models controlling for the effects of 11 species’ traits reflecting the influence of other factors (e.g. climate change, land cover change, proximity to range edges) on trends and including the species and country indentifiers as random effects. We found that the listing under the Annex I had significantly positive effect on bird trends in both old and new member states, whereas no such effect was observed in the non-member states. Although the positive influence of listing was larger in the old and than in the new member states, the difference was not statistically significant. Our results imply that the Annex I of the Birds Directive is an important tool for bird conservation in Europe and that its positive influence on bird populations is detectable even in the new EU members entering EU relatively recently. As birds are often used as indicators also for other groups of organisms, these results suggest that not only birds may benefit from EU’s conservation legislation but comprehensive assessments are needed.

Geographic variations of the bird-borne structural risk of West Nile virus circulation in Europe.

The structural risk of West Nile Disease results from the usual functioning of the socio-ecological system, which may favour the introduction of the pathogen, its circulation and the occurrence of disease cases. Its geographic variations result from the local interactions between three components: (i) reservoir hosts, (ii) vectors, both characterized by their diversity, abundance and competence, (iii) and the socio-economic context that impacts the exposure of human to infectious bites. We developed a model of bird-borne structural risk of West Nile Virus (WNV) circulation in Europe, and analysed the association between the geographic variations of this risk and the occurrence of WND human cases between 2002 and 2014. A meta-analysis of WNV serosurveys conducted in wild bird populations was performed to elaborate a model of WNV seropositivity in European bird species, considered a proxy for bird exposure to WNV. Several eco-ethological traits of bird species were linked to seropositivity and the statistical model adequately fitted species-specific seropositivity data (area under the ROC curve: 0.85). Combined with species distribution maps, this model allowed deriving geographic variations of the bird-borne structural risk of WNV circulation. The association between this risk, and the occurrence of WND human cases across the European Union was assessed. Geographic risk variations of bird-borne structural risk allowed predicting WND case occurrence in administrative districts of the EU with a sensitivity of 86% (95% CI: 0.79–0.92), and a specificity of 68% (95% CI: 0.66–0.71). Disentangling structural and conjectural health risks is important for public health managers as risk mitigation procedures differ according to risk type. The results obtained show promise for the prevention of WND in Europe. Combined with analyses of vector-borne structural risk, they should allow designing efficient and targeted prevention measures.

How weather instead of urbanity measures affects song trait variability in three European passerine bird species.

Previous studies detected an influence of urban characteristics on song traits in passerine birds, that is, song adjustments to ambient noise in urban areas. Several studies already described the effect of weather conditions on the behavior of birds, but not the effect on song traits. We investigate, if song trait variability changes along a continuous urbanity gradient in Frankfurt am Main, Germany. We examined, for the first time on a larger scale, the influence of weather on song parameters. We made song recordings of three common passerine species: the blue and great tit (Cyanistes caeruleus (Linnaeus, 1758) and Parus major Linnaeus, 1758) and the European blackbird (Turdus merula Linnaeus, 1758). We measured different song traits and performed statistical analyses and modeling on a variety of variables—among them urbanity and weather parameters. Remarkably, we found only few cases of a significant influence of urbanity parameters on song traits. The influence of weather parameters (air pressure, atmospheric humidity, air and soil temperatures) on song traits was highly significant. Birds in Frankfurt face high noise pollution and might show different adaptations to high noise levels. The song trait variability of the investigated species is affected more by weather conditions than by urban characteristics in Frankfurt. However, the three species react differently to specific weather parameters. Smaller species seem to be more affected by weather than larger species.

Morphological constraints on changing avian migration phenology.

Many organisms at northern latitudes have responded to climate warming by advancing their spring phenology. Birds are known to show earlier timing of spring migration and reproduction in response to warmer springs. However, species show heterogeneous phenological responses to climate warming, with those that have not advanced or have delayed migration phenology experiencing population declines. Although some traits (such as migration distance) partly explain heterogeneity in phenological responses, the factors affecting interspecies differences in the responsiveness to climate warming have yet to be fully explored. In this comparative study, we investigate whether variation in wing aspect ratio (reflecting relative wing narrowness), an ecomorphological trait that is strongly associated with flight efficiency and migratory behaviour, affects the ability to advance timing of spring migration during 1960-2006 in a set of 80 European migratory bird species. Species with larger aspect ratio (longer and narrower wings) showed smaller advancement of timing of spring migration compared to species with smaller aspect ratio (shorter and wider wings) while controlling for phylogeny, migration distance and other life-history traits. In turn, migration distance positively predicted aspect ratio across species. Hence, species that are better adapted to migration appear to be more constrained in responding phenologically to rapid climate warming by advancing timing of spring migration. Our findings corroborate the idea that aspect ratio is a major evolutionary correlate of migration, and suggest that selection for energetically efficient flights, as reflected by high aspect ratio, may hinder phenotypically plastic/microevolutionary adjustments of migration phenology to ongoing climatic changes.

Reproductive timing as a constraint on invasion success in the Ring-necked parakeet (Psittacula krameri)

Climate similarity favors biological invasion, but a match between seasonality in the novel range and the timing of life cycle events of the invader also influences the outcome of species introduction. Yet, phenology effects on invasion success have generally been neglected. Here we study whether a phenological mismatch limits the non-native range of a globally successful invader, the Ring-necked parakeet, in Europe. Given the latitudes at which parakeets have established across Europe, they breed earlier than expected based on breeding dates from the native Asian range. Moreover, comparing the breeding dates of European populations to those of parakeets in the native Asian range, to five native breeding bird species in Europe and to the start of the growing season of four native European trees shows that the discrepancy between expected and actual breeding phenology is greater in northern Europe. In northern European populations, this temporal mismatch appears to have negative effects on hatching success, and on population growth rates in years that are colder than average in the first six months. Phenological mismatch also can explain why parakeets from African populations (that are more likely to breed in autumn) have been poor invaders compared to parakeets from Asia. These lines of evidence support the hypothesis that the reproductive phenology of the Ring-necked parakeet can be a limiting factor for establishment and range expansion in colder climates. Our results provide growing support for the hypothesis that the match between climate seasonality and timing of reproduction (or other important life cycle events) can affect the establishment success, invasive potential and distribution range of introduced non-native species, beyond the mere effect of climate similarity.

Vocal behaviour of mates at the nest in the White-throated Dipper Cinclus cinclus: contexts and structure of vocal interactions, pair-specific acoustic signature

Birdsong has been extensively studied in the context of sexual selection, and intersexual vocal communication prior to pair formation has been the focus of most interest. With the exception of conspicuous vocal duets of tropical bird species, descriptions of intra-pair vocal communication are missing for most other bird species. This communication can take different forms, fulfil different functions and be more widespread than previously described. Females and males of the White-throated Dipper Cinclus cinclus, a European bird species, have been reported to sing during pair formation, but vocal communication between mates after this phase has not been investigated to date. We report here our study of the vocal behaviour between mates at the nest and describe the specific contexts and structure of the various vocal interactions both during incubation and while brooding hatchlings (N = 23 pairs). Vocal interactions were mainly composed of two vocalization types: ‘trills’ and ‘notes’, with trills being more common at the beginning of the vocal sequence. Both the acoustic features of the vocalizations and the temporal organization of the vocal sequences changed between the two breeding stages. In particular, compared to the trills and notes produced during the incubation period, those produced while brooding were lower in pitch, and female–male vocal sequences were composed of more notes and songs, with a lower overlap rate. This difference may reflect changes in parental activities. Trills and notes carried a pair-specific acoustic signature. This first description of acoustic communication between mates during breeding in the White-throated Dipper lays the basis for future investigation of the functions of the different vocalizations.

Colonial, more widely distributed and less abundant bird species undergo wider population fluctuations independent of their population trend.

Understanding temporal variability in population size is important for conservation biology because wide population fluctuations increase the risk of extinction. Previous studies suggested that certain ecological, demographic, life-history and genetic characteristics of species might be related to the degree of their population fluctuations. We checked whether that was the case in a large sample of 231 European breeding bird species while taking a number of potentially confounding factors such as population trends or similarities among species due to common descent into account. When species-specific characteristics were analysed one by one, the magnitude of population fluctuations was positively related to coloniality, habitat, total breeding range, heterogeneity of breeding distribution and natal dispersal, and negatively related to urbanisation, abundance, relative number of subspecies, parasitism and proportion of polymorphic loci. However, when abundance (population size) was included in the analyses of the other parameters, only coloniality, habitat, total breeding range and abundance remained significantly related to population fluctuations. The analysis including all these predictors simultaneously showed that population size fluctuated more in colonial, less abundant species with larger breeding ranges. Other parameters seemed to be related to population fluctuations only because of their association with abundance or coloniality. The unexpected positive relationship between population fluctuations and total breeding range did not seem to be mediated by abundance. The link between population fluctuations and coloniality suggests a previously unrecognized cost of coloniality. The negative relationship between population size and population fluctuations might be explained by at least three types of non-mutually exclusive stochastic processes: demographic, environmental and genetic stochasticity. Measurement error in population indices, which was unknown, may have contributed to the negative relationship between population size and fluctuations, but apparently only to a minor extent. The association between population size and fluctuations suggests that populations might be stabilized by increasing population size.

A phylogenetic comparative analysis reveals correlations between body feather structure and habitat

SummaryBody feathers ensure both waterproofing and insulation in waterbirds, but how natural variation in the morphological properties of these appendages relates to environmental constraints remains largely unexplored. Here, we test how habitat and thermal condition affect the morphology of body feathers, using a phylogenetic comparative analysis of five structural traits [i.e., total feather length, the lengths of the pennaceous (distal) and plumulaceous (proximal) sections, barb density, and pennaceous barbule density] from a sample of 194 European bird species.Body feather total length is shorter in aquatic than in terrestrial birds, and this difference between groups is due to the shorter plumulaceous feather section in aquatic birds. Indeed, a reduced plumulaceous section in feather length probably reflects the need to limit air trapped in the plumage to adjust the buoyancy of aquatic birds. In contrast, the high pennaceous barbule density of aquatic birds compared to their terrestrial counterparts reflects water resistance of the plumage in contact with water.Our results show that birds living in environments with low ambient temperature have long plumulaceous feather lengths, low barb density, and low pennaceous barbule density. Data also suggest that plumage probably has limited function in reducing the heat absorption of species living in hot environments.Our results have broad implications for understanding the suite of selection pressures driving the evolution of body feather functional morphology. It remains to be tested, however, how other feather traits, such as the density of plumage (feathers per unit area) and the relative number of different feather types, for example downy feathers, are distributed amongst birds with different water resistance and thermoinsulative needs.Alay summaryis available for this article.

Inefficient co-feeding transmission of Borrelia afzelii in two common European songbirds

The spirochete bacterium Borrelia afzelii is the most common cause of Lyme borreliosis in Europe. This tick-borne pathogen can establish systemic infections in rodents but not in birds. However, several field studies have recovered larval Ixodes ricinus ticks infected with B. afzelii from songbirds suggesting successful transmission of B. afzelii. We reviewed the literature to determine which songbird species were the most frequent carriers of B. afzelii-infected I. ricinus larvae and nymphs. We tested experimentally whether B. afzelii is capable of co-feeding transmission on two common European bird species, the blackbird (Turdus merula) and the great tit (Parus major). For each bird species, four naïve individuals were infested with B. afzelii-infected I. ricinus nymphal ticks and pathogen-free larval ticks. None of the co-feeding larvae tested positive for B. afzelii in blackbirds, but a low percentage of infected larvae (3.33%) was observed in great tits. Transstadial transmission of B. afzelii DNA from the engorged nymphs to the adult ticks was observed in both bird species. However, BSK culture found that these spirochetes were not viable. Our study suggests that co-feeding transmission of B. afzelii is not efficient in these two songbird species.

Large-brained birds suffer less oxidative damage.

Large brains (relative to body size) might confer fitness benefits to animals. Although the putative costs of well-developed brains can constrain the majority of species to modest brain sizes, these costs are still poorly understood. Given that the neural tissue is energetically expensive and demands antioxidants, one potential cost of developing and maintaining large brains is increased oxidative stress ('oxidation exposure' hypothesis). Alternatively, because large-brained species exhibit slow-paced life histories, they are expected to invest more into self-maintenance such as an efficacious antioxidative defence machinery ('oxidation avoidance' hypothesis). We predict decreased antioxidant levels and/or increased oxidative damage in large-brained species in case of oxidation exposure, and the contrary in case of oxidation avoidance. We address these contrasting hypotheses for the first time by means of a phylogenetic comparative approach based on an unprecedented data set of four redox state markers from 85 European bird species. Large-brained birds suffered less oxidative damage to lipids (measured as malondialdehyde levels) and exhibited higher total nonenzymatic antioxidant capacity than small-brained birds, whereas uric acid and glutathione levels were independent of brain size. These results were not altered by potentially confounding variables and did not depend on how relative brain size was quantified. Our findings partially support the 'oxidation avoidance' hypothesis and provide a physiological explanation for the linkage of large brains with slow-paced life histories: reduced oxidative stress of large-brained birds can secure brain functionality and healthy life span, which are integral to their lifetime fitness and slow-paced life history.

Differences in shifts of wintering and breeding ranges lead to changing migration distances in European birds

Studies on the impact of climate change on the distributions of bird species in Europe have largely focused on one season at a time, especially concerning summer breeding ranges. We investigated whether migratory bird species show consistent range shifts over the past 55 yr in both breeding and wintering areas or if these shifts are independent. We then analyzed whether patterns in changing migration distances of Finnish breeding birds could be explained by habitat use, phylogeny or body size. We used long‐term datasets from the Finnish ringing centre to analyze the mean wintering latitudes of 29 species of Finnish breeding birds, then used breeding distribution data to make predictions as to whether certain species were migrating shorter or longer distances based on the comparative shifts in the wintering and breeding grounds. Our data reveal species‐specific differences in changing migration distances. We show that for many species, long‐term shifts in wintering ranges have not followed the same patterns as those in the breeding range, leading to differences in migration distances over time. We conclude that species are not adjusting predictably to climate change in their wintering grounds, leading to changing migration distances in some, but not all, species breeding in Finland. This research fills an important gap in the current climate change biology literature, focusing on individuals’ entire life histories and revealing new complexities in range shift patterns.

Survival and dispersal of the Cyprus wheatear Oenanthe cypriaca, an endemic migrant

Many populations of European migrant bird species are declining and this may be driven by survival rates; however, there are few studies that can estimate true survival rates. Cyprus wheatears Oenanthe cypriaca are an endemic migrant that winter in East Africa: populations are probably not declining but are annually variable. We recorded territory occupation and reoccupation in a colour-ringed population of 45–69 pairs over a 4-year period (2010–2013) from April to August to measure apparent survival and determine how it varied with sex, age, breeding productivity and year. We then estimated true survival by correcting apparent survival for dispersal by recording territory shifts and how this also varied by sex, age, breeding productivity and year. Apparent annual survival rate varied significantly by sex, age and year (males 2011, 2012, 2013: 0.70, 0.50, 0.62; females: 0.56, 0.34, 0.47; chicks: 0.35, 0.19, 0.28) but was not affected by the productivity of a territory. An average of 1.1 % of males and 8.2 % of females were lost during breeding, where 5/7 lost females were found depredated during incubation. Adults did not usually change territories between years (87 % were resident and 99 % moved less than four territories between years) regardless of sex, productivity or year; chicks, independent of their sex, moved on average three territories away from their natal territory. After correcting apparent survival for the probability of dispersal, males had the highest true minimum annual survival compared to females which were very similar to chicks (males 2011, 2012, 2013: 0.77, 0.50, 0.65; females: 0.65, 0.35, 0.50; chicks: 0.64, 0.34, 0.49). The results indicate a very high survival rate for a small passerine migrant, although they are probably sufficiently annually variable to profoundly affect annual population dynamics. If females have a lower survival rate, the sex ratio at birth may be female-biased to compensate; alternatively, females may have longer range dispersal than we could measure, particularly if they respond to their mates not returning by moving territories, leading to underestimation of their true survival. A high survival rate may be due to the rarity of sparrowhawks on Cyprus and the wheatears relatively short distance migration.

Morphological Adaptations to Migration in Birds

Migratory flight performance has direct or carry-over effects on fitness. Therefore, selection is expected to act on minimizing the costs of migratory flight, which increases with the distance covered. Aerodynamic theory predicts how morphological adaptations improve flight performance. These predictions have rarely been tested in comparative analyses that account for scaling and phylogenetic effects. We amassed a unique dataset of 149 European bird species and 10 morphological traits. Mass-adjusted aspect ratio increased, while mass-adjusted heart weight and wing loading decreased with increasing migration distance. These results were robust to whether the analyses were based on the entire species pool or limited to passerines or migrants. Our findings indicate that selection due to migration acts on wing traits that reduce the energetic cost of transportation to increase the flight range. Consequently, the demands for high ‘exercise organ’ performance might be low, and hence such energetically expensive tissues are not associated (pectoral muscle) or are inversely associated (heart) with migration distance.

The drivers of avian abundance: patterns in the relative importance of climate and land use

AbstractAimClimate and land use can have important effects on the local abundances of species, but few studies have investigated the relative impacts of these factors. Here, we quantify the relative importance of climate, land use and surrounding population size in determining the abundances of birds across a continent.LocationEurope.MethodsWe used species abundance models to identify the relative importance of different environmental predictors for estimating the local abundances of 342 species of European breeding birds. Models controlling for phylogeny were used to relate species life history and ecological traits to the climate:land use importance ratio. The mean of this ratio, across all species occurring in a given area, was mapped to explore spatial variation in the major drivers of abundance.ResultsAt the scale examined, climate is generally more important than land use in determining species abundances. However, the abundance of species in neighbouring areas is also a major correlate. Among climate variables, temperature is of greater importance than moisture availability in determining abundances. The relative importance of these variables varies with latitude, with temperature being most important in the north, and moisture availability in the south. Differences in the importance of specific drivers are related to species ecological traits: climate is more important for determining the abundance of species that have larger global ranges or a smaller body mass.Main conclusionsAbundances of species occurring in northern Europe, an area predicted to experience climatic changes of high magnitude, are most sensitive to climate, particularly temperature. Given the greater confidence in future projections of temperature than precipitation, this increases confidence in projections of the impacts of climate change on species in the north, whilst attempts to predict future populations in central and southern Europe may be dependent on less predictable changes in land use and precipitation.

Is research effort associated with the conservation status of European bird species?

Research effort is critical to the success of interventions which aim to reduce de clines in biodiversity and so can be considered as a conservation resource. European bird species are among the best studied and most popular groups of animals, yet 1 in 10 are currently globally threatened or near threatened, and many have been prioritised by various multilateral environmental agreements. To investigate how research effort is directed towards European bird species, and in particular whether conservation listing prompts additional research, the number of outputs from a Web of Science search related to conservation was used to develop an index of research effort for each species. This index was then analysed against a set of population, morphological, ecological and socioeconomic parameters collated from the literature, plus measures of threat level. A series of generalised linear models revealed that the most important factors in explaining the distribution of research effort amongst European bird species included ‘European population size’, ‘potential research investment’ and ‘habitat type’, which were linked to ease of study. Also important were the species’ generation length, 1990−2000 European population trend and migration status. Research effort was not well targeted with respect to either European or global threat status, and there was little support for the suggestion that inclusion of species in legislative instruments such as Annex I of the EU Birds Directive might stimulate research. Research effort must become better prioritised to achieve the greatest net conservation benefit.

Interspecific variation in the structural properties of flight feathers in birds indicates adaptation to flight requirements and habitat

Summary The functional significance of intra‐ and interspecific structural variations in the flight feathers of birds is poorly understood. Here, a phylogenetic comparative analysis of four structural features (rachis width, barb and barbule density and porosity) of proximal and distal primary feathers of 137 European bird species was conducted. Flight type (flapping and soaring, flapping and gliding, continuous flapping or passerine type), habitat (terrestrial, riparian or aquatic), wing characteristics (wing area, S and aspect ratio, AR) and moult strategy were all found to affect feather structure to some extent. Species characterized by low wing‐beat frequency flight (soaring and gliding) have broader feather rachises (shafts) and feather vanes with lower barb density than birds associated with more active flapping modes of flight. However, the effect of flying mode on rachis width disappeared after controlling for S and AR, suggesting that rachis width is primarily determined by wing morphology. Rachis width and feather vane density are likely related to differences in force distribution across the wingspan during different flight modes. An increase in shaft diameter, barb density and porosity from the proximal to distal wing feathers was found and was highest in species with flapping flight indicating that aerodynamic forces are more biased towards the distal feathers in flapping flyers than in soarers and gliders. Habitat affected barb and barbule density, which was greatest in aquatic species, and within this group, barb density was greater in divers than non‐divers, suggesting that the need for water repellency and resistance to water penetration may influence feather structure. However, we found little support for the importance of porosity in water repellency and water penetration, because porosity was similar in aquatic, riparian and terrestrial species and among the aquatic birds (divers and non‐divers). We also found that barb density was affected by moult pattern. Our results have broad implications for the understanding of the selection pressures driving flight feather functional morphology. Specifically, the large sample size relative to any previous studies has emphasized that the morphology of flight feathers is the result of a suite of selection pressures. As well as routine flight needs, constraints during moulting, habitat (particularly aquatic) and migratory requirements also affect flight feather morphology. Identifying the exact nature of these trade‐offs will perhaps inform the reconstruction of the flying modes of extinct birds.

DNA barcoding and minibarcoding as a powerful tool for feather mite studies.

Feather mites (Astigmata: Analgoidea and Pterolichoidea) are among the most abundant and commonly occurring bird ectosymbionts. Basic questions on the ecology and evolution of feather mites remain unanswered because feather mite species identification is often only possible for adult males, and it is laborious even for specialized taxonomists, thus precluding large-scale identifications. Here, we tested DNA barcoding as a useful molecular tool to identify feather mites from passerine birds. Three hundred and sixty-one specimens of 72 species of feather mites from 68 species of European passerine birds from Russia and Spain were barcoded. The accuracy of barcoding and minibarcoding was tested. Moreover, threshold choice (a controversial issue in barcoding studies) was also explored in a new way, by calculating through simulations the effect of sampling effort (in species number and species composition) on threshold calculations. We found one 200-bp minibarcode region that showed the same accuracy as the full-length barcode (602bp) and was surrounded by conserved regions potentially useful for group-specific degenerate primers. Species identification accuracy was perfect (100%) but decreased when singletons or species of the Proctophyllodes pinnatus group were included. In fact, barcoding confirmed previous taxonomic issues within the P.pinnatus group. Following an integrative taxonomy approach, we compared our barcode study with previous taxonomic knowledge on feather mites, discovering three new putative cryptic species and validating three previous morphologically different (but still undescribed) new species.

Characterisation of Toll-like receptors 4, 5 and 7 and their genetic variation in the grey partridge.

Toll-like receptors (TLRs) are a cornerstone of vertebrate innate immunity. In this study, we identified orthologues of TLR4, TLR5 and TLR7 (representing both bacterial- and viral-sensing TLRs) in the grey partridge (Perdix perdix), a European Galliform game bird species. The phylogeny of all three TLR genes follows the known phylogeny of Galloanserae birds, placing grey partridge TLRs (PePeTLRs) in close proximity to their turkey and pheasant orthologues. The predicted proteins encoded by the PePeTLR genes were 843, 862-863 and 1,047 amino acids long, respectively, and clearly showed all TLR structural features. To verify functionality in these genes we mapped their tissue-expression profiles, revealing generally high PePeTLR4 and PePeTLR5 expression in the thymus and absence of PePeTLR4 and PePeTLR7 expression in the brain. Using 454 next-generation sequencing, we then assessed genetic variation within these genes for a wild grey partridge population in the Czech Republic, EU. We identified 11 nucleotide substitutions in PePeTLR4, eight in PePeTLR5 and six in PePeTLR7, resulting in four, four and three amino acid replacements, respectively. Given their locations and chemical features, most of these non-synonymous substitutions probably have a minor functional impact. As the intraspecific genetic variation of the three TLR genes was low, we assume that either negative selection or a bottleneck may have reduced TLR population variability in this species.