A refined magnetic pulse treatment method for magnetic navigation experiments with adequate sham control: a case study on free-flying songbirds.

Migratory songbirds may navigate by extracting positional information from the geomagnetic field, potentially with a magnetic-particle-based receptor. Previous studies assessed this hypothesis experimentally by exposing birds to a strong but brief magnetic pulse aimed at remagnetizing the particles and evoking an altered behaviour. Critically, such studies were not ideally designed because they lacked an adequate sham treatment controlling for the induced electric field that is fundamentally associated with a magnetic pulse. Consequently, we designed a sham-controlled magnetic-pulse experiment, with sham and treatment pulse producing a similar induced electric field, while limiting the sham magnetic field to a value that is deemed insufficient to remagnetize particles. We tested this novel approach by pulsing more than 250 wild, migrating European robins (Erithacus rubecula) during two autumn seasons. After pulsing them, five traits of free-flight migratory behaviour were observed, but no effect of the pulse could be found. Notably, one of the traits, the migratory motivation of adults, was significantly affected in only one of the two study years. Considering the problem of reproducing experiments with wild animals, we recommend a multi-year approach encompassing large sample size, blinded design and built-in sham control to obtain future insights into the role of magnetic-particle-based magnetoreception in bird navigation.

The urge to breed early: Similar responses to environmental conditions in short- and long-distance migrants during spring migration.

Birds migrating different distances experience different temporal, energetic, physiological, and physical constraints throughout migration, which is reflected in their migration strategy. Consequently, we predict different behavioral decisions to similar environmental cues between short- and long-distance migrants, which has been documented for autumn migration. Here, we focus on the question whether trade-off decisions regarding departure, routing, and landing when alternating between migratory endurance flights and stopovers also differ during spring migration. As early arrivals at the breeding grounds should be ultimately favored regardless of migration distance, selection may favor more similar behavioral decisions in spring than in autumn. We radio-tagged short- and long-distance migratory songbirds at stopover sites along the German North Sea coast during spring and automatically tracked their migratory behavior using a large-scale network of receiver stations. Once departed, birds could either cross the sea or detour along the coast. We corrected for spatially biased detection data, using a hierarchical multistate model to assess how birds respond to variation in environmental conditions in their day-to-day departure decisions and route selection. The day-to-day departure probability was higher in long-distance migrants independently of the routing decision. Irrespective of migration distance, all species more likely departed under light winds and rainless conditions, while the influence of air pressure change and relative humidity was species-specific. By accounting for detection probabilities, we estimated that about half of all individuals of each species crossed the sea but did not find differences between short- and long-distance migrants. Offshore flights were more likely when winds blew offshore and began earlier within the night compared with onshore flights. Our results suggest that selection more similarly affects birds of different migration distances in spring than in autumn. These findings put the focus toward how ultimate mechanisms may shape departure and routing decisions differently between migration seasons.

Departure, routing and landing decisions of long-distance migratory songbirds in relation to weather.

Migrating birds flexibly adjust their individual migratory decisions, i.e. departing, routing and landing, based on intrinsic (e.g. energy stores) and extrinsic (e.g. landscape features and weather) factors modulating the endogenous stimuli. So far, these decisions have mostly been studied separately. Notably, we lack information on which factors landing decisions during active flight are based on. Therefore, we simultaneously recorded all three decisions in free-flying long-distance migratory songbirds in a coastal stopover area via regional-scale radio-telemetry and related them to the prevailing weather. Birds departed under favourable weather conditions resulting in specific nights with increased departure probability. Once departed, birds could either fly offshore or take a route along the coast, which was predicted by wind support. Radio-tracking revealed that departed individuals more likely interrupted their migratory endurance flight under overcast or headwind conditions. Studying departure, routing and landing decisions in concert, we highlight the importance of weather as a common driver across all migratory decisions. By radio-tracking individuals between stopovers, we provide evidence that avoidance of adverse weather conditions is an important function of stopover. Understanding how birds adjust migratory decisions and how they affect the timing of migration and survival is key to link migration performance to individual fitness.

A magnetic pulse does not affect free-flight navigation behaviour of a medium-distance songbird migrant in spring.

Current evidence suggests that migratory animals extract map information from the geomagnetic field for true navigation. The sensory basis underlying this feat is elusive, but presumably involves magnetic particles. A common experimental manipulation procedure consists of pre-treating animals with a magnetic pulse, with the aim of re-magnetising particles to alter the internal representation of the external field prior to a navigation task. Although pulsing provoked deflected bearings in caged songbirds, analogous studies with free-flying songbirds yielded inconsistent results. Here, we pulsed European robins (Erithacus rubecula) at an offshore stopover site during spring migration and monitored their free-flight behaviour with a regional-scale network of radio-receiving stations. We found no pulse effect on departure probability, nocturnal departure timing departure direction or consistency of flight direction. This suggests either no use of the geomagnetic map by our birds, or that magnetic pulses do not affect the sensory system underlying geomagnetic map detection.

No apparent effect of a magnetic pulse on free-flight behaviour in northern wheatears (Oenanthe oenanthe) at a stopover site.

Naïve migrants reach their wintering grounds following a clock-and-compass strategy. During these inaugural migrations, birds internalise, among others, cues from the Earth's magnetic field to create a geomagnetic map, with which they navigate to destinations familiar to them on subsequent migrations. Geomagnetic map cues are thought to be sensed by a magnetic-particle-based receptor, which can be specifically affected by a magnetic pulse. Indeed, the orientation of experienced but not naïve birds was compromised after magnetic pulsing, indicating geomagnetic map use. Little is known about the importance of this putative magnetoreceptor for navigation and decision-making in free-flying migrants. Therefore, we studied in unprecedented detail how a magnetic pulse would affect departure probability, nocturnal departure timing, departure direction and consistency in flight direction over 50-100 km in experienced and naïve long-distant migrant songbirds using a large-scale radio-tracking system. Contrary to our expectations and despite a high sample size (ntotal = 137) for a free-flight study, we found no significant after-effect of the magnetic pulse on the migratory traits, suggesting the geomagnetic map is not essential for the intermediate autumn migration phase. These findings warrant re-thinking about perception and use of geomagnetic maps for migratory decisions within a sensory and ecological context.

Days to visit an offshore island: effect of weather conditions on arrival fuel load and potential flight range for common blackbirds Turdus merula migrating over the North Sea.

BACKGROUND: Crossing open water instead of following the coast(line) is one way for landbirds to continue migration. However, depending on prevailing weather and the birds' physiological conditions, it is also a risky choice. To date, the question remains as to which interplay between environmental and physiological conditions force landbirds to stop on remote islands. We hypothesise that unfavourable winds affect lean birds with low energy resources, while poor visibility affects all birds regardless of their fuel loads. METHODS: To test this hypothesis, we caught 1312 common blackbirds Turdus merula stopping over on Helgoland during autumn and spring migration. Arrival fuel load was measured using quantitative magnetic resonance technology. Weather parameters (wind and relative humidity as a proxy for visibility) were interpolated for the night before arrival. Further, we calculated whether caught individuals would have successfully crossed the North Sea instead of landing on Helgoland, depending on wind conditions. RESULTS: Both wind and relative humidity the night before arrival were correlated with arrival fuel load. After nights with strong headwinds, birds caught the following day were mostly lean, most of which would not have managed to cross the sea if they had not stopped on Helgoland. In contrast, fat birds that could have successfully travelled on were caught mainly after nights with high relative humidity (≥ 80%). Furthermore, the rate of presumably successful flights was lower due to wind: although only 9% of all blackbirds captured on Helgoland had insufficient fuel loads to allow safe onward migration in still air, real wind conditions would have prevented 30% of birds from successfully crossing the sea during autumn and 21% during spring migration. CONCLUSIONS: We were able to decipher how physiological condition, wind and relative humidity partially force blackbirds to stop on a remote island. Adverse winds tend to affect lean birds with low energy resources, while poor visibility can affect blackbirds, regardless of whether the arrival fuel load was sufficient for onward flight. Our findings will help to understand different migratory strategies and explain further questions like migration timing.

Are movements of daytime and nighttime passerine migrants as different as day and night?

Even after decades of research, the migration of songbirds still holds numerous secrets. Distinct stopover and routing behavior of diurnally and nocturnally migrating songbirds has been stated in the 1960s, but empirical confirmation is yet lacking widely. We studied the behavior of individual diurnally migrating dunnocks and nocturnally migrating blackcaps by means of large-scale automated radio-telemetry. Birds were radio-tagged during their stopover at the German North Sea coast. Our data indicate longer initial stopover duration in the diurnally migrating dunnocks, opposing the hypothesis of nocturnal migrants needing more time to recover due to their longer migratory flights. Nonetheless, dunnocks stopped over more often along their tracks as when compared to the nocturnally migrating blackcaps. Behavior en route did not differ as clearly between species challenging the general view of contrasting routings of diurnal and nocturnal migrants with regard to landscape and open water. Our results imply additional factors of relevance other than differences in species or daily migration timing per se. We discuss and highlight the need of detailed and individual based data to better understand stopover and routing behavior of songbirds in the environmental context.

Weather at the winter and stopover areas determines spring migration onset, progress, and advancements in Afro-Palearctic migrant birds.

Climate change causes changes in the timing of life cycle events across all trophic groups. Spring phenology has mostly advanced, but large, unexplained, variations are present between and within species. Each spring, migratory birds travel tens to tens of thousands of kilometers from their wintering to their breeding grounds. For most populations, large uncertainties remain on their exact locations outside the breeding area, and the time spent there or during migration. Assessing climate (change) effects on avian migration phenology has consequently been difficult due to spatial and temporal uncertainties in the weather potentially affecting migration timing. Here, we show for six trans-Saharan long-distance migrants that weather at the wintering and stopover grounds almost entirely (∼80%) explains interannual variation in spring migration phenology. Importantly, our spatiotemporal approach also allows for the systematic exclusion of influences at other locations and times. While increased spring temperatures did contribute strongly to the observed spring migration advancements over the 55-y study period, improvements in wind conditions, especially in the Maghreb and Mediterranean, have allowed even stronger advancements. Flexibility in spring migration timing of long-distance migrants to exogenous factors has been consistently underestimated due to mismatches in space, scale, time, and weather variable type.

Occurrence of Gastrointestinal Parasites in Small Mammals from Germany.

An increase in zoonotic infections in humans in recent years has led to a high level of public interest. However, the extent of infestation of free-living small mammals with pathogens and especially parasites is not well understood. This pilot study was carried out within the framework of the "Rodent-borne pathogens" network to identify zoonotic parasites in small mammals in Germany. From 2008 to 2009, 111 small mammals of 8 rodent and 5 insectivore species were collected. Feces and intestine samples from every mammal were examined microscopically for the presence of intestinal parasites by using Telemann concentration for worm eggs, Kinyoun staining for coccidia, and Heidenhain staining for other protozoa. Adult helminths were additionally stained with carmine acid for species determination. Eleven different helminth species, five coccidians, and three other protozoa species were detected. Simultaneous infection of one host by different helminths was common. Hymenolepis spp. (20.7%) were the most common zoonotic helminths in the investigated hosts. Coccidia, including Eimeria spp. (30.6%), Cryptosporidium spp. (17.1%), and Sarcocystis spp. (17.1%), were present in 40.5% of the feces samples of small mammals. Protozoa, such as Giardia spp. and amoebae, were rarely detected, most likely because of the repeated freeze-thawing of the samples during preparation. The zoonotic pathogens detected in this pilot study may be potentially transmitted to humans by drinking water, smear infection, and airborne transmission.

To cross or not to cross - thrushes at the German North Sea coast adapt flight and routing to wind conditions in autumn.

BACKGROUND: Although many aspects of passerine migration are genetically determined, routing appears to be flexibly adjusted to the conditions experienced on each individual journey. This holds especially true for routing decisions taken when confronted with large bodies of water. Once taken, these decisions can be hardly altered or revised. In this paper, we analysed stopover and routing decisions taken by three species of thrushes, blackbirds, redwings and song thrushes, at the German North Sea coast. METHODS: Birds were equipped with radio-telemetry tags at stopover sites along the coast during autumn migration and subsequently tracked by an automated receiver network covering the coastline and islands of the German Bight. RESULTS: The thrushes resumed migration in nights with a favourable northward wind component and clear skies. About 40% of the tagged individuals have taken an offshore instead of an alongshore oriented flight route. Routing decisions were influenced by the strength of the eastward wind component with offshore oriented flights taking place primarily under weak winds or winds blowing towards the west. Thrushes that took an offshore oriented route stopped over at the coast longer than those flying alongshore. Interestingly, offshore as well as alongshore oriented flights co-occurred within single nights and under comparable weather conditions. CONCLUSIONS: Migratory flight and routing decisions of thrushes at the German North Sea coast are highly dependent on weather, in particular wind. Still, we found evidence that weather may not be the sole reason for individual routes taken. Physical condition, morphology or animal personality lend themselves as possible additional factors of influence. Enabling a more detailed understanding of thrush migration over and along the German North Sea, our data help to better judge risks that migratory birds are facing when en route conditions are altered, for example by artificial obstacles such as offshore wind turbines.

Autumn bird migration phenology: A potpourri of wind, precipitation and temperature effects.

Climate change has caused a clear and univocal trend towards advancement in spring phenology. Changes in autumn phenology are much more diverse, with advancement, delays, and 'no change' all occurring frequently. For migratory birds, patterns in autumn migration phenology trends have been identified based on ecological and life-history traits. Explaining interspecific variation has nevertheless been challenging, and the underlying mechanisms have remained elusive. Radar studies on non-species-specific autumn migration intensity have repeatedly suggested that there are strong links with weather. In long-term species-specific studies, the variance in autumn migration phenology explained by weather has, nevertheless, been rather low, or a relationship was even lacking entirely. We performed a spatially explicit time window analysis of weather effects on mean autumn passage of four trans-Saharan and six intra-European passerines to gain insights into this apparent contradiction. We analysed data from standardized daily captures at the Heligoland island constant-effort site (Germany), in combination with gridded daily temperature, precipitation and wind data over a 55-year period (1960-2014), across northern Europe. Weather variables at the breeding and stopover grounds explained up to 80% of the species-specific interannual variability in autumn passage. Overall, wind conditions were most important. For intra-European migrants, wind was even twice as important as either temperature or precipitation, and the pattern also held in terms of relative contributions of each climate variable to the temporal trends in autumn phenology. For the trans-Saharan migrants, however, the pattern of relative trend contributions was completely reversed. Temperature and precipitation had strong trend contributions, while wind conditions had only a minor impact because they did not show any strong temporal trends. As such, understanding species-specific effects of climate on autumn phenology not only provides unique insights into each species' ecology but also how these effects shape the observed interspecific heterogeneity in autumn phenological trends.

The influence of weather on avian spring migration phenology: What, where and when?

Over the past decades, spring arrival and passage of most short- and medium-distance migrating birds in the Northern Hemisphere have advanced. Changes in spring temperature at the passage or arrival area have been most frequently shown to be related to these changes in spring migration phenology. In most studies, preliminary assumptions are made on both the spatial location and the specific time frame of the weather influencing spring migration phenology. We performed a spatially explicit time-window analysis of the effect of weather on mean spring passage dates of nine short- and medium-distance passerines. We analysed data from standardized daily captures at the Helgoland (Germany) constant-effort site, in combination with gridded daily temperature, precipitation and wind data from the NCEP data set over a 55-year period (1960-2014), across the whole of West Europe and North Africa. Although we allowed for a time window of any length at any location, nevertheless incorporating various measures to avoid spurious correlations, time windows at the likely wintering or spring stopover grounds were almost exclusively selected as the best predicting variables (96%-100% of identified variables). The weather variables at the wintering and stopover grounds explain up to 77% of the interannual variability in spring passage. Yet, the response of spring migration phenology to weather at the winter or stopover areas does not fully explain the observed trends. Spring migration phenology is, hence, strongly driven by weather at the wintering and stopover grounds, but additional mechanisms are needed to fully explain the advancement of spring migration. Our results also clearly show that previously illustrated correlations, or the lack thereof, between spring migration phenology and weather at the passage or arrival location are due to spatio-temporal correlations in the weather data. This spatial mismatch might have led to false conclusions, especially the further away the wintering or stopover sites are.

Challenging a 15-year-old claim: The North Atlantic Oscillation index as a predictor of spring migration phenology of birds.

Many migrant bird species that breed in the Northern Hemisphere show advancement in spring arrival dates. The North Atlantic Oscillation (NAO) index is one of the climatic variables that have been most often investigated and shown to be correlated with these changes in spring arrival. Although the NAO is often claimed to be a good predictor or even to have a marked effect on interannual changes in spring migration phenology of Northern Hemisphere breeding birds, the results on relations between spring migration phenology and NAO show a large variety, ranging from no, over weak, to a strong association. Several factors, such as geographic location, migration phase, and the NAO index time window, have been suggested to partly explain these observed differences in association. A combination of a literature meta-analysis, and a meta-analysis and sliding time window analysis of a dataset of 23 short- and long-distance migrants from the constant-effort trapping garden at Helgoland, Germany, however, paints a completely different picture. We found a statistically significant overall effect size of the NAO on spring migration phenology (coefficient = -0.14, SE = 0.054), but this on average only explains 0%-6% of the variance in spring migration phenology across all species. As such, the value and biological meaning of the NAO as a general predictor or explanatory variable for climate change effects on migration phenology of birds, seems highly questionable. We found little to no definite support for previously suggested factors, such as geographic location, migration phenology phase, or the NAO time window, to explain the heterogeneity in correlation differences. We, however, did find compelling evidence that the lack of accounting for trends in both time series has led to strongly inflated (spurious) correlations in many studies (coefficient = -0.13, SE = 0.019).

True navigation in migrating gulls requires intact olfactory nerves.

During migratory journeys, birds may become displaced from their normal migratory route. Experimental evidence has shown that adult birds can correct for such displacements and return to their goal. However, the nature of the cues used by migratory birds to perform long distance navigation is still debated. In this experiment we subjected adult lesser black-backed gulls migrating from their Finnish/Russian breeding grounds (from >60°N) to Africa (to < 5°N) to sensory manipulation, to determine the sensory systems required for navigation. We translocated birds westward (1080 km) or eastward (885 km) to simulate natural navigational challenges. When translocated westwards and outside their migratory corridor birds with olfactory nerve section kept a clear directional preference (southerly) but were unable to compensate for the displacement, while intact birds and gulls with the ophthalmic branch of the trigeminal nerve sectioned oriented towards their population-specific migratory corridor. Thus, air-borne olfactory information seems to be important for migrating gulls to navigate successfully in some circumstances.

Climate warming, ecological mismatch at arrival and population decline in migratory birds.

Climate is changing at a fast pace, causing widespread, profound consequences for living organisms. Failure to adjust the timing of life-cycle events to climate may jeopardize populations by causing ecological mismatches to the life cycle of other species and abiotic factors. Population declines of some migratory birds breeding in Europe have been suggested to depend on their inability to adjust migration phenology so as to keep track of advancement of spring events at their breeding grounds. In fact, several migrants have advanced their spring arrival date, but whether such advancement has been sufficient to compensate for temporal shift in spring phenophases or, conversely, birds have become ecologically mismatched, is still an unanswered question, with very few exceptions. We used a novel approach based on accumulated winter and spring temperatures (degree-days) as a proxy for timing of spring biological events to test if the progress of spring at arrival to the breeding areas by 117 European migratory bird species has changed over the past five decades. Migrants, and particularly those wintering in sub-Saharan Africa, now arrive at higher degree-days and may have therefore accumulated a 'thermal delay', thus possibly becoming increasingly mismatched to spring phenology. Species with greater 'thermal delay' have shown larger population decline, and this evidence was not confounded by concomitant ecological factors or by phylogenetic effects. These findings provide general support to the largely untested hypotheses that migratory birds are becoming ecologically mismatched and that failure to respond to climate change can have severe negative impacts on their populations. The novel approach we adopted can be extended to the analysis of ecological consequences of phenological response to climate change by other taxa.

North Atlantic Oscillation and timing of spring migration in birds.

Migrant birds have been trapped on the island of Helgoland (southeastern North Sea) since 1909, with methods and sampling effort remaining unchanged throughout the last four decades. In 12 short/medium-distance migrants and 12 long-distance migrants (23 passerines plus the European woodcock) sample sizes were sufficient to calculate mean spring passage (msp) times and to relate these to climate change. All but one species, passing Helgoland en route to their breeding areas (mainly in Scandinavia), show a trend towards earlier msp-time, which is significant in 7 short/medium-distance migrants and 10 long-distance migrants. The msp-times advanced by 0.05-0.28 days per year, short/medium-distance migrants not differing from long-distance migrants. In 23 out of the 24 species, earlier msp-times coincide with local warmer msp-temperatures (significantly in 11 and 7 species of the two groups, respectively). Even more striking is the relation to a large-scale phenomenon, the North Atlantic Oscillation (NAO), during the last four decades. Again, in 23 out of the 24 species, an earlier msp-time coincides with higher NAO indices (significantly in 9 and 12 species, respectively). The NAO index can also explain differences and similarities in spring migration strategies, as well as migration routes within Europe.