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Cryptochromes, blue-light absorbing proteins involved in the circadian clock, have been proposed to be the receptor molecules of the avian magnetic compass. In birds, several cryptochromes occur: Cryptochrome 2, Cryptochrome 4 and two splice products of Cryptochrome 1, Cry1a and Cry1b. With an antibody not distinguishing between the two splice products, Cryptochrome 1 had been detected in the retinal ganglion cells of garden warblers during migration. A recent study located Cry1a in the outer segments of UV/V-cones in the retina of domestic chickens and European robins, another migratory species. Here we report the presence of cryptochrome 1b (eCry1b) in retinal ganglion cells and displaced ganglion cells of European Robins, Erithacus rubecula. Immuno-histochemistry at the light microscopic and electron microscopic level showed eCry1b in the cell plasma, free in the cytosol as well as bound to membranes. This is supported by immuno-blotting. However, this applies only to robins in the migratory state. After the end of the migratory phase, the amount of eCry1b was markedly reduced and hardly detectable. In robins, the amount of eCry1b in the retinal ganglion cells varies with season: it appears to be strongly expressed only during the migratory period when the birds show nocturnal migratory restlessness. Since the avian magnetic compass does not seem to be restricted to the migratory phase, this seasonal variation makes a role of eCry1b in magnetoreception rather unlikely. Rather, it could be involved in physiological processes controlling migratory restlessness and thus enabling birds to perform their nocturnal flights.

Seed dispersal allows plants to colonise new sites and escape from pathogens and intraspecific competition, maintaining plant genetic diversity and regulating plant distribution. Conversely, most plant species form mutualistic associations with arbuscular mycorrhizal (AM) fungi in a symbiosis established immediately after seed germination. Because AM fungi are obligate symbionts, using the same dispersal vector as their host should be highly advantageous for their survival, but the co-dispersal of seeds and AM fungal spores has never been confirmed. We aim to clarify the potential role of European birds, essential dispersers for many plant species, as co-dispersers of seeds and AM fungal spores. In total, 63 bird droppings with intact seeds were placed in sterilised soil and maintained for 4 months in a protected environment to avoid contamination. Additionally, 173 bird droppings and 729 gauze swabs used to clean birds’ feet were inspected for AM fungal spores. Although no spores were detected by direct observation of these samples, seven Rubus ulmifolius seedlings obtained from four independent droppings of Erithacus rubecula and Sylvia melanocephala were colonised by AM fungi. Our results show that birds can effectively co-disperse viable seeds and AM fungal spores, potentially over long distances, providing a pivotal mechanism to understand the cosmopolitan distribution of AM fungi.

Background Birds can act as reservoirs of tick-borne pathogens and can also disperse pathogen-containing ticks to both nearby and remote localities. The aims of this study were to estimate tick infestation patterns on migratory birds and the prevalence of different Borrelia species and tick-borne encephalitis virus (TBEV) in ticks removed from birds in south-eastern Sweden. Methods Ticks were collected from resident and migratory birds captured at the Ottenby Bird Observatory, Öland, Sweden, from March to November 2009. Ticks were molecularly identified to species, and morphologically to developmental stage, and the presence of Borrelia bacteria and TBEV was determined by quantitative real-time PCR. Results A total of 1339 ticks in the genera Haemaphysalis, Hyalomma , and Ixodes was recorded of which I. ricinus was the most abundant species. Important tick hosts were the European robin ( Erithacus rubecula ), Blackbird ( Turdus merula ), Tree pipit ( Anthus trivialis ), Eurasian wren ( Troglodytes troglodytes ) , Common redstart ( Phoenicurus phoenicurus ) , Willow warbler ( Phylloscopus trochilus ), and Common whitethroat ( Sylvia communis ). Borrelia bacteria were detected in 25% (285/1,124) of the detached ticks available for analysis. Seven Borrelia species ( B. afzelii , B. burgdorferi ( s.s. ), B. garinii , B. lusitaniae , B. turdi, B. valaisiana , and B. miyamotoi ) were identified. B. turdi was recorded for the first time in ticks in Sweden. The number of Borrelia cells per tick ranged from 2.0 × 10 0 to 7.0 × 10 5 . B. miyamotoi -containing ticks contained a significantly higher median number of Borrelia cells than B. burgdorferi ( s.l. )-containing ticks. B. garinii and B. miyamotoi were the most prevalent Borrelia species in tick larvae. Larvae of I. ricinus with B. garinii were removed from seven bird species, particularly S. communis and A. trivialis , which may suggest that the larvae had contracted the Borrelia bacteria from or via these birds. Also, a high percentage of tick larvae containing B. miyamotoi was removed from E. rubecula. All ticks were negative for TBEV. Conclusions The results corroborate the view that the contributions of birds to human disease are substantial, particularly as blood hosts for ticks and for their short-, medium-, and long-distance dispersal. Moreover, several ground-foraging bird species appear to be important for the maintenance and dispersal of Borrelia species. The absence of TBEV in the ticks conforms to other similar studies. Graphical Abstract

Magnetic compass orientation in night-migratory songbirds is embedded in the visual system and seems to be based on a light-dependent radical pair mechanism. Recent findings suggest that both broadband electromagnetic fields ranging from ~2 kHz to ~9 MHz and narrow-band fields at the so-called Larmor frequency for a free electron in the Earth’s magnetic field can disrupt this mechanism. However, due to local magnetic fields generated by nuclear spins, effects specific to the Larmor frequency are difficult to understand considering that the primary sensory molecule should be organic and probably a protein. We therefore constructed a purpose-built laboratory and tested the orientation capabilities of European robins in an electromagnetically silent environment, under the specific influence of four different oscillating narrow-band electromagnetic fields, at the Larmor frequency, double the Larmor frequency, 1.315 MHz or 50 Hz, and in the presence of broadband electromagnetic noise covering the range from ~2 kHz to ~9 MHz. Our results indicated that the magnetic compass orientation of European robins could not be disrupted by any of the relatively strong narrow-band electromagnetic fields employed here, but that the weak broadband field very efficiently disrupted their orientation.

The increasing urbanisation has important impacts on natural soundscapes, through habitat loss and noise pollution (anthrophony), affecting acoustic communication in wildlife. Birds living in cities often adjust their songs to avoid masking by low‐frequency traffic noise, but little is still known about how multiple song traits vary during the day. In this study, we investigated how anthrophony affects song traits in European robins (Erithacus rubecula) over a 10‐h timeframe by comparing urban and rural populations. We recorded 59 robins in the city of Edinburgh, Scotland, and 54 robins in the Midlothian countryside and compared their songs with linear mixed models. Minimum frequency in rural robins decreased over the day, whereas it remained elevated and constant in urban robins. Urban songs were also longer, less complex and had narrower frequency ranges. These patterns suggest plastic regulation of minimum frequency in response to background noise, anthrophony in urban areas and biophony in rural areas, while suggesting potential constraints on other song traits. Although the fitness consequences of reduced complexity and frequency range remain unclear, behavioural plasticity may help preserve signal reliability in urban habitats. Our study investigated how anthrophony affected birdsong by comparing European robin songs from urban and rural populations in Midlothian, Scotland. The minimum frequency showed a decreasing pattern in rural robins, but remained higher and constant in urban robins, which also had longer, less complex songs with narrower frequency ranges. Overall, the results suggest a plastic regulation of minimum frequency in response to background noise, although fitness consequences remain unclear.

Haemosporidian blood parasites are widely used in evolutionary ecological research when exploring the effects of parasites on different life-history traits of their bird hosts. However, their roles in bird migration are less studied. If these parasites deteriorate the body condition of the birds strongly, they might negatively affect the whole migration phenology and the survival of the birds as well. In our study, we tested the relationships between infection for parasite genera (Haemoproteus or Plasmodium), the three most frequent parasite lineages and body condition (body mass, fat deposit), and the timing of autumn migration in the European Robin (Erithacus rubecula). We found that mean body mass and fat scores did not differ between parasitized and non-parasitized individuals, but infected juveniles arrived later than their non-infected counterparts. The difference in the arrival time of parasitized and non-parasitized birds was greater in the case of Haemoproteus infections. However, when we analysed the effects of the distinct parasite lineages separately, we found that prevalence of parasite lineages correlated with the body mass, fat storage, and timing of autumn migration of the birds in a different direction. Our results therefore emphasize the importance of testing the impacts of the different parasites individually, because possible lineage-specific effects on bird condition during migration might exist.

Cryptochromes are photolyase-related blue-light receptors acting as core components of the mammalian circadian clock in the cell nuclei. One or more members of the cryptochrome protein family are also assumed to play a role in avian magnetoreception, but the primary sensory molecule in the retina of migratory birds that mediates light-dependent magnetic compass orientation has still not been identified. The mRNA of cryptochrome 2 (Cry2) has been reported to be located in the cell nuclei of the retina, but Cry2 localisation has not yet been demonstrated at the protein level. Here, we provide evidence that Cry2 protein is located in the photoreceptor inner segments, the outer nuclear layer, the inner nuclear layer and the ganglion cell layer in the retina of night-migratory European robins, homing pigeons and domestic chickens. At the subcellular level, we find Cry2 both in the cytoplasm and the nucleus of cells residing in these layers. This broad nucleic expression rather points to a role for avian Cry2 in the circadian clock and is consistent with a function as a transcription factor, analogous to mammalian Cry2, and speaks against an involvement in magnetoreception.

Many partially migratory species show phenotypically divergent populations in terms of migratory behaviour, with climate hypothesized to be a major driver of such variability through its differential effects on sedentary and migratory individuals. Based on long‐term (1947–2011) bird ringing data, we analysed phenotypic differentiation of migratory behaviour among populations of the European robin Erithacus rubecula across Europe. We showed that clusters of populations sharing breeding and wintering ranges varied from partial (British Isles and Western Europe, NW cluster) to completely migratory (Scandinavia and north‐eastern Europe, NE cluster). Distance migrated by birds of the NE (but not of the NW) cluster decreased through time because of a north‐eastwards shift in the wintering grounds. Moreover, when winter temperatures in the breeding areas were cold, individuals from the NE cluster also migrated longer distances, while those of the NW cluster moved over shorter distances. Climatic conditions may therefore affect migratory behaviour of robins, although large geographical variation in response to climate seems to exist.

Cryptochromes, blue-light absorbing proteins involved in the circadian clock, have been proposed to be the receptor molecules of the avian magnetic compass. In birds, several cryptochromes occur: Cryptochrome 2, Cryptochrome 4 and two splice products of Cryptochrome 1, Cry1a and Cry1b. With an antibody not distinguishing between the two splice products, Cryptochrome 1 had been detected in the retinal ganglion cells of garden warblers during migration. A recent study located Cry1a in the outer segments of UV/V-cones in the retina of domestic chickens and European robins, another migratory species. Here we report the presence of cryptochrome 1b (eCry1b) in retinal ganglion cells and displaced ganglion cells of European Robins, Erithacus rubecula. Immuno-histochemistry at the light microscopic and electron microscopic level showed eCry1b in the cell plasma, free in the cytosol as well as bound to membranes. This is supported by immuno-blotting. However, this applies only to robins in the migratory state. After the end of the migratory phase, the amount of eCry1b was markedly reduced and hardly detectable. In robins, the amount of eCry1b in the retinal ganglion cells varies with season: it appears to be strongly expressed only during the migratory period when the birds show nocturnal migratory restlessness. Since the avian magnetic compass does not seem to be restricted to the migratory phase, this seasonal variation makes a role of eCry1b in magnetoreception rather unlikely. Rather, it could be involved in physiological processes controlling migratory restlessness and thus enabling birds to perform their nocturnal flights.

Background Songbirds following distinct migration strategies (e.g. long- vs. short- to medium-distance migrants) often differ in their speed of migration during autumn and, thus, are assumed to face different time constraints. During migration, most songbird species alternate migratory flights with stopover periods. Many of them restrict these migratory flights to the night, i.e., they are nocturnal migrants. At stopover, nocturnal migrants need to select a specific night (night-to-night decision) and time of night (within-night decision) to resume migration. These departure decisions, which largely determine the speed of migration, are jointly affected by a set of intrinsic and extrinsic factors, i.e., departure cues. Here we aim to assess whether the set of intrinsic and extrinsic factors and the magnitude of their respective effects on stopover departure decisions differs between nocturnally migrating songbird species, depending on their migration strategy and associated time constraints. Methods We radio-tracked migrating Northern Wheatears ( Oenanthe oenanthe ; long-distance migrant), European robins ( Erithacus rubecula ) and Common Blackbirds ( Turdus merula ; both medium-distance migrants) during autumn stopover and analysed their night-to-night and within-night departure timing in relation to intrinsic and extrinsic factors. Results Species generally differed in their departure timing on both temporal scales, with shortest stopovers and earliest nocturnal departures in the long-distance migrant. Some factors, such as day of year, fuel load, cloud cover and crosswind, had consistent effects on stopover departure decisions in all three species. However, species differed in the effects of tailwind assistance, change in atmospheric pressure and air temperature on their stopover departure decisions. Whereas night-to-night decisions were affected by these extrinsic factors in either both or one of the medium-distance migrants, such effects were not found in the long-distance migrant. Conclusions Our results suggest that the general timing of departures in songbirds is affected by the species-specific migration strategy and associated time constraints. Further, they imply that the assessment and usage of specific extrinsic factors, i.e., weather conditions, as departure cues is adjusted based on this migration strategy, with the long-distance migrants being least selective at departure. Other intrinsic and extrinsic factors, however, seem to be used as departure cues independent of migration strategy.