Compass orientation and possible migration routes of passerine birds at high arctic latitudes

The use of celestial or geomagnetic orientation cues can lead migratory birds along different migration routes during the migratory journeys, e.g. great circle routes (approximate), geographic or magnetic loxodromes. Orientation cage experiments have indicated that migrating birds are capable of detecting magnetic compass information at high northern latitudes even at very steep angles of inclination. However, starting a migratory journey at high latitudes and following a constant magnetic course often leads towards the North Magnetic Pole, which means that the usefulness of magnetic compass orientation at high latitudes may be questioned. Here, we compare possible long‐distance migration routes of three species of passerine migrants breeding at high northern latitudes. The initial directions were based on orientation cage experiments performed under clear skies and simulated overcast and from release experiments under natural overcast skies. For each species we simulated possible migration routes (geographic loxodrome, magnetic loxodrome and sun compass route) by extrapolating from the initial directions and assessing a fixed orientation according to different compass mechanisms in order to investigate what orientation cues the birds most likely use when migrating southward in autumn. Our calculations show that none of the compass mechanisms (assuming fixed orientation) can explain the migration routes followed by night‐migrating birds from their high Nearctic breeding areas to the wintering sites further south. This demonstrates that orientation along the migratory routes of arctic birds (and possibly other birds as well) must be a complex process, involving different orientation mechanisms as well as changing compass courses. We propose that birds use a combination of several compass mechanisms during a migratory journey with each of them being of a greater or smaller importance in different parts of the journey, depending on environmental conditions. We discuss reasons why birds developed the capability to use magnetic compass information at high northern latitudes even though following these magnetic courses for any longer distance will lead them along totally wrong routes. Frequent changes and recalibrations of the magnetic compass direction during the migratory journey are suggested as a possible solution.     

SUNSET AND THE ORIENTATION BEHAVIOUR OF MIGRATING BIRDS

1. Migratory birds integrate information from a wide array of environmental sources. As our knowledge of migratory orientation depends heavily upon the results of cage-experiments with nocturnal migrants, it is essential that the results of these cage studies be interpreted in the light of field observations of migratory behaviour and experiments with free-flying migrants. When this is done, the impression emerges that night-migrating birds integrate directional information prior to departure, probably during the transition between daylight and darkness. At this time, information gained from the sun, in conjunction with other references, becomes especially valuable. 2. Despite intensive work with a few species, how migrants integrate information in the selection and maintenance of a direction is not well understood. The relationship between magnetic stimuli and solar cues at sunset in the selection process, for example, remains to be resolved, as does the contribution of skylight polarization patterns at sunset. Once a migratory heading is selected, birds probably use the stars or winds aloft to maintain that direction. How migrants integrate information is largely a matter of unravelling the complex causal relations among the different environmental stimuli that serve as orientation cues. Imagine a hypothetical migrant that departs on a migratory flight around the time of sunset. Given the uncertain relationship among variables (orientation cues) that might influence her migratory orientation, a path diagram is a useful device for displaying graphically the pattern of causal relations among the set of variables (see Fig. 1). This technique is adopted from path analysis, which is a statistical method developed by Sewall Wright for studying the direct and indirect causal relations among variables (see Kerlinger & Pedhazur, 1973). The pattern depicted in the figure is less a specific model of causal relations than it is a summary of possible relationships among the several cues based on current understanding. Causal flow in this ‘model’ is unidirectional, i.e. at any given point in time a variable cannot be both a cause and an effect of another variable. For example, variable 3 is dependent on variables 1 and/or 2, and is one of the independent variables in relation to variable 5 (orientation of migratory activity). Although the value of path analysis to the study of migratory orientation may be largely heuristic at this point, ‘one virtue of the method is that in order to apply it the researcher is required to make explicit the theoretical framework within which he operates’ (Kerlinger & Pedhazur, 1973). For instance, path diagrams (and path analysis, to the degree that correlations between variables can be specified) would help researchers study (i) the apparent redundancy built into the orientation process (see Fig. 1), (ii) alternative or competing causal models of orientation and navigation, or (iii) the ontogenetic changes that affect the relationship among orientation variables. Imagine, for example, how path coefficients might change in value with migratory experience. 3. Migrants probably redetermine preferred directions soon after landing or shortly before their next departure rather than while aloft. Cage-orientation results as well as observations of free-flying migrants suggest that solar-related information is involved in the morning orientation of ongoing migratory flight and possibly the re-determination of direction following night-time displacement. 4. Evidence is not clear on whether migrants respond to sunset by constant-angle orientation (menotaxis) or constant-azimuth orientation. 5. How migrants correctly identify sunset as a reference stimulus is an unresolved question. Identification might be based upon the characteristic spectral distribution of sunset, its pattern of illumination, or some other feature, such as the characteristic pattern of skylight polarization at sunset. 6. Several lines of evidence suggest that migrants learn to use the setting sun and associated skylight features as orientation cues. 7. The setting sun functions not only as a source of directional information but also as an environmental stimulus that influences the likelihood of migratory activity.     

Orientation Cage and Release Experiments with Migratory Wheatears (Oenanthe oenanthe) in Scandinavia and Greenland: The Importance of Visual Cues

Migratory orientation of Scandinavian and Greenland wheatears was recorded during the autumn migration periods of 1988 and 1989. Orientation cage tests were conducted under clear sunset skies, to investigate the importance of different visible sky sections on orientation performance. In addition, wheatears were released under clear starry skies and under total overcast to examine the orientation of free-flying birds. The following results were obtained:

• 1 Wheatears tested with a restricted visible sky section (90° centered around zenith) in orientation cages, showed a mean orientation towards geographic W/geomagnetic NW (Greenland) and towards geographic and magnetic WNW-NW (Sweden). These mean directions are clearly inconsistent with the expected autumn migration directions, SW-SSW in Scandinavia and SE in Greenland, as revealed by ringing recoveries for the two populations.
• 2 When the birds were allowed a much more extensive view of the sky, almost down to the horizon (above 10° elevation), Scandinavian wheatears chose headings in agreement with ringing data. Greenland birds were not significantly oriented.
• 3 Release experiments under clear starry skies resulted in mean vanishing directions in good agreement with ringing data from both sites. Greenland wheatears released under total overcast showed a similar orientation as under clear skies, indicating that a view of the stars may not be of crucial importance for selecting a seasonally accurate migratory direction.

The results suggest that an unobstructed view of the sky, including visual cues low over the horizon, is important, possibly in combination with geomagnetic cues, for the orientation of migratory naive wheatears. Furthermore, the birds showed remarkably similar orientation responses in Greenland and Scandinavia, respectively, indicating that they use basically the same orientation system, despite considerable differences in visual and geomagnetic orientation premises at the two different geographic and magnetic latitudes.     

Field studies of avian nocturnal migratory orientation I. interaction of sun,wind and stars as directional cues

Tracking radar and visual observation techniques were used to observe the orientation of free-flying passerine nocturnal migrants in situations in which potentially usable directional cues were absent or gave conflicting information. When migrants had seen the sun near the time of sunset and/or the stars, they oriented in appropriate migratory directions even when winds were opposed. Under solid overcast skies that prevented a view of both sun and stars, the birds headed downwind in opposing winds and thus moved in seasonally inappropriate directions. The data point to the primacy of visual cues over wind direction, with either sun or stars being sufficient to allow the birds to determine the appropriate migration direction.     

A long-distance avian migrant compensates for longitudinal displacement during spring migration

In order to perform true bicoordinate navigation, migratory birds need to be able to determine geographic latitude and longitude. The determination of latitude is relatively easy from either stellar or magnetic cues [1-3], but the determination of longitude seems challenging [4, 5]. It has therefore been suggested that migrating birds are unable to perform bicoordinate navigation and that they probably only determine latitude during their return migration [5]. However, proper testing of this hypothesis requires displacement experiments with night-migratory songbirds in spring that have not been performed. We therefore displaced migrating Eurasian reed warblers (Acrocephalus scirpaceus) during spring migration about 1000 km toward the east and found that they were correcting for displacements by shifting their orientation from the northeast at the capture site to the northwest after the displacement. This new direction would lead them to their expected breeding areas. Our results suggest that Eurasian reed warblers are able to determine longitude and perform bicoordinate navigation. This finding is surprising and presents a new intellectual challenge to bird migration researchers, namely, which cues enable birds to determine their east-west position.     

Virus isolation and molecular epidemiology of highly pathogenic avian influenza A(H5N8) from an outbreak in free-ranging wild birds,India 2016

We report the molecular epidemiology of highly pathogenic avian influenza (HPAI) virus involved in an outbreak causing death in free-ranging wild birds at Mysore, Karnataka state of India. The virus was typed as HPAI A(H5N8) by conventional and TaqMan probe based real-time PCR assays. Six isolates of HPAI virus were recovered in 9-day-old embryonated chicken eggs. Haemagglutinin gene-based phylogeny of virus isolates showed >?99.9% nucleotide sequence identity with HPAI A(H5N8) isolates from migratory birds and domestic poultry from China and Korea indicating either these wild birds have routed their migration through Korea and/or eastern China or these dead birds must have directly or indirectly contacted with wild birds migrating from Eastern China and/or Korean regions. The study emphasises the role of migratory wild birds in spread of HPAI across the globe.     

Can vector summation describe the orientation system of juvenile ospreys and honey buzzards? – An analysis of ring recoveries and satellite tracking

Juvenile bird migrants are generally believed to use a clock‐and‐compass migratory orientation strategy. According to such a strategy migrants accomplish their migration by flying a number of successive flight steps with direction and number of steps controlled by an endogenous programme. One powerful way of testing this is by comparing predictions from a model of such a strategy with observed patterns. We used data from ringing and satellite‐based radio telemetry to investigate the orientation system of juvenile ospreys (Pandion haliaetus) and honey buzzards (Pernis apivorus) migrating from Sweden to tropical west Africa. The ring recoveries showed a much larger scatter in the orientation of ospreys than of honey buzzards, but there was only a slight such difference in the satellite tracks. These tracks of individuals of both species were rather straight with a high directional concentration per step. The honey buzzard data showed a close fit to a simple vector summation model, which is expected if birds follow a clock‐and‐compass strategy. However, the osprey data did not fit such a simple model, as ring recoveries showed a significantly greater deviation at short distances than predicted on the basis of long distance data. Satellite tracking also indicated less concentrated orientation on short distances. The pattern observed for the osprey can generally be explained by an extended vector summation model, including an important element of pre‐migration dispersal. The existence of extensive dispersal in the osprey stands in contrast to the apparent absence of such dispersal in the honey buzzard. The explanation for this difference between the species is unclear. The model of orientation by vector summation is very sensitive to the existence of differences in mean direction between individuals. Assuming such differences, as tentatively indicated by the satellite tracking data, makes simple compass orientation by vector summation inconsistent with the distribution of ring recoveries at long distances, with a high proportion of misoriented birds falling outside the normal winter range.     

Nocturnal orientation of robins, Erithacus rubecula: birds caught during migratory flight are disoriented

For conventional experiments on the orientation behavior of migrant birds in funnels, either hand-raised birds or birds caught during resting periods at stopover sites are generally used. Topographic circumstances at the Alpine pass Col de Bretolet at the Switzerland/France border allow the capture of birds during active migratory flight during the whole night. These birds are in full migratory disposition. We expected them to orient in the seasonally appropriate direction more accurately than birds that had not experienced migration just before the funnel experiment. Experiments with robins, however, revealed a strong influence of the moon on the orientation behavior. The birds did not orient in the seasonally expected migratory direction but showed positive phototaxis, usually toward the lightest part of the funnel in the direction opposite to the moon. When the moon was absent the robins were disoriented. Sunset experiments with robins caught during the night before the experiment revealed a strong phototactic reaction toward the setting sun. As reasons for this poor orientation in the absence of light stimuli, the influence of the topography of the mountainous region and magnetic anomalies can be excluded. It is concluded that freshly caught birds are too stressed when tested immediately after capture or that the migration direction cannot be maintained. Testing of night migrants in complete darkness is also of disadvantage. Received: 23 December 1998 / Received in revised form: 16 July 1999 / Accepted: 30 July 1999     

Comparative migration strategies of wild and captive‐bred Asian Houbara Chlamydotis macqueenii

For migratory species, the success of population reintroduction or reinforcement through captive‐bred released individuals depends on survivors undertaking appropriate migrations. We assess whether captive‐bred Asian Houbara Chlamydotis macqueenii from a breeding programme established with locally sourced individuals and released into suitable habitat during spring or summer undertake similar migrations to those of wild birds. Using satellite telemetry, we compare the migrations of 29 captive‐bred juveniles, 10 wild juveniles and 39 wild adults (including three birds first tracked as juveniles), examining migratory propensity (proportion migrating), timing, direction, stopover duration and frequency, efficiency (route deviation), and wintering and breeding season locations. Captive‐bred birds initiated autumn migration an average of 20.6 (±4.6 se) days later and wintered 470.8 km (±76.4) closer to the breeding grounds, mainly in Turkmenistan, northern Iran and Afghanistan, than wild birds, which migrated 1217.8 km (±76.4), predominantly wintering in southern Iran and Pakistan (juveniles and adults were similar). Wintering locations of four surviving captive‐bred birds were similar in subsequent years (median distance to first wintering site = 70.8 km, range 6.56–221.6 km), suggesting that individual captive‐bred birds (but not necessarily their progeny) remain faithful to their first wintering latitude. The migratory performance of captive‐bred birds was otherwise similar to that of wild juveniles. Although the long‐term fitness consequences for captive‐bred birds establishing wintering sites at the northern edge of those occupied by wild birds remain to be quantified, it is clear that the pattern of wild migrations established by long‐term selection is not replicated. If the shorter migration distance of young captive‐bred birds has a physiological rather than a genetic basis, then their progeny may still exhibit wild‐type migration. However, as there is a considerable genetic component to migration, captive breeding management must respect migratory population structure as well as natal and release‐site fidelity.     

Juvenile songbirds compensate for displacement to oceanic islands during autumn migration

To what degree juvenile migrant birds are able to correct for orientation errors or wind drift is still largely unknown. We studied the orientation of passerines on the Faroe Islands far off the normal migration routes of European migrants. The ability to compensate for displacement was tested in naturally occurring vagrants presumably displaced by wind and in birds experimentally displaced 1100 km from Denmark to the Faroes. The orientation was studied in orientation cages as well as in the free-flying birds after release by tracking departures using small radio transmitters. Both the naturally displaced and the experimentally displaced birds oriented in more easterly directions on the Faroes than was observed in Denmark prior to displacement. This pattern was even more pronounced in departure directions, perhaps because of wind influence. The clear directional compensation found even in experimentally displaced birds indicates that first-year birds can also possess the ability to correct for displacement in some circumstances, possibly involving either some primitive form of true navigation, or 'sign posts', but the cues used for this are highly speculative. We also found some indications of differences between species in the reaction to displacement. Such differences might be involved in the diversity of results reported in displacement studies so far.     

Individuality in bird migration: routes and timing

The exploration of animal migration has entered a new era with individual-based tracking during multiple years. Here, we investigated repeated migratory journeys of a long-distance migrating bird, the marsh harrier Circus aeruginosus, in order to analyse the variation within and between individuals with respect to routes and timing. We found that there was a stronger individual repeatability in time than in space. Thus, the annual timing of migration varied much less between repeated journeys of the same individual than between different individuals, while there was considerable variation in the routes of the same individual on repeated journeys. The overall contrast in repeatability between time and space was unexpected and may be owing to strong endogenous control of timing, while short-term variation in environmental conditions (weather and habitat) might promote route flexibility. The individual variation in migration routes indicates that the birds navigate mainly by other means than detailed route recapitulation based on landmark recognition.     

Spatio-temporal distribution of migrating raptors: a comparison of ringing and satellite tracking

We describe the migration performance of three long-distance migrating raptors, osprey Pandion haliaetus , honey buzzard Pernis apivorus and marsh harrier Circus aeruginosus , and one short-distance migrating raptor, common buzzard Buteo buteo based on Swedish ringing recoveries and satellite telemetry, respectively. Tracking by satellite can provide detailed information about the exact timing of migration, migration speed, migration directions, stopover sites, and detours, thereby overcoming many of the potential biases found in ring recoveries. Comparison of the results from these two methods revealed agreement in the geographical distribution of the studied Swedish raptor populations during autumn migration and the winter period. Satellite tracking, nevertheless, provided much more detailed information in Africa and revealed significantly faster migration progress than indicated by ring recoveries. The implications of our findings for interpretation of migratory connectivity and the understanding of migration are discussed.     

The Orientation Behaviour of Chaffinches, Fringilla coelebs, Caught during Active Migratory Flight, in Relation to the Sun

The few orientation studies that have been carried out with day-migrating birds show that they are able to use solar and magnetic orientation cues for orientation. Previous orientation experiments in Emlen funnels have been carried out either with hand-raised birds or with birds caught during resting periods at stop-over sites. The aim of our study was to test whether birds caught during active flight show a higher concentration of migratory activity in the seasonally appropriate migratory direction in the funnels than birds that had not experienced migration just before the funnel experiments. The topography at the alpine pass Col de Bretolet at the border of Switzerland and France allowed us to capture birds during active migratory flight. These birds were in full migration disposition. Orientation experiments with chaffinches suggested an influence of the sun because chaffinches did not orient in the seasonally expected direction, but probably showed positive phototaxis towards the light of the sun at the opposite side of the funnel. Chaffinches tested under overcast conditions oriented to the north-west which probably was a 'nonsense' orientation and not a reverse migration or compensatory behaviour. We conclude that freshly caught birds are too stressed to show appropriate orientation when tested immediately after catching.     

Wind selection and drift compensation optimize migratory pathways in a high-flying moth

Numerous insect species undertake regular seasonal migrations in order to exploit temporary breeding habitats [1]. These migrations are often achieved by high-altitude windborne movement at night [2-6], facilitating rapid long-distance transport, but seemingly at the cost of frequent displacement in highly disadvantageous directions (the so-called "pied piper" phenomenon [7]). This has lead to uncertainty about the mechanisms migrant insects use to control their migratory directions [8, 9]. Here we show that, far from being at the mercy of the wind, nocturnal moths have unexpectedly complex behavioral mechanisms that guide their migratory flight paths in seasonally-favorable directions. Using entomological radar, we demonstrate that free-flying individuals of the migratory noctuid moth Autographa gamma actively select fast, high-altitude airstreams moving in a direction that is highly beneficial for their autumn migration. They also exhibit common orientation close to the downwind direction, thus maximizing the rectilinear distance traveled. Most unexpectedly, we find that when winds are not closely aligned with the moth's preferred heading (toward the SSW), they compensate for cross-wind drift, thus increasing the probability of reaching their overwintering range. We conclude that nocturnally migrating moths use a compass and an inherited preferred direction to optimize their migratory track.     

Migrating Birds as Dispersal Vehicles for West Nile Virus

Whereas migrating birds have been implicated in the spread of West Nile virus (WNV), there is no direct evidence of birds actively migrating while infectious. The role of birds in WNV dispersal is difficult to assess in the field. However, this role can be evaluated experimentally because birds in migratory disposition display increased locomotor activity or restlessness under captive conditions. We tested the following hypotheses: (1) migrating passerine birds continue to exhibit migratory activity while infectious with WNV and (2) the migratory state of the individual affects the magnitude of viremia. We examined the migratory activity of two neoarctic-neotropical passerine migrants, Swainson’s thrush (Catharus ustulatus) and gray catbird (Dumetella carolinensis), during acute WNV infection. All gray catbirds and six of nine Swainson’s thrushes exhibited migratory activity while infectious. Moreover, migratory status did not appear to influence viremia titers, as might be expected if individuals were immunosuppressed during migration. Therefore, we demonstrate that migrating passerine birds are potential dispersal vehicles for WNV.     

Geomagnetic information modulates nocturnal migratory restlessness but not fueling in a long distance migratory songbird

Geomagnetic cues have been shown to influence migratory orientation and migratory fuelling in night‐migratory songbird species. Here, we used captive‐bred northern wheatears Oenanthe oenanthe from the southern Norwegian population to show that other aspects of the birds’ migratory program can be influenced by magnetic cues as well. We observed that the amount of migratory restlessness increased strongly with progression of the migratory season when the birds were kept constantly in the magnetic field of northern Germany, but the amount of migratory restlessness decreased when the magnetic field changed along the birds’ natural flyway are simulated. Thus, the Earth's magnetic field can also act as a ‘signpost’ cue for fine‐tuning the spatio‐temporal course of migration.     

Migratory Orientation of Blackcaps (Sylvia atricapilla): Population-specific Shifts of Direction during the Autumn

The population-specific orientation of two groups of blackcaps (Sylvia atricapilla), one from southwest Germany, the other from easternmost Austria, was studied outdoors in Emlen funnels. We investigated whether a seasonal shift in the migratory direction — as expected for the Austrian birds from ringing recoveries — occurs under experimental conditions and in a seasonally constant magnetic field. The West German birds, for which no shift was expected, oriented southwest during the entire season. The Austrian birds oriented southeast in October and southsouthwest in November. The clockwise shift by about 60° occurred within a 10-day period. The results indicate that in this species seasonal changes of migratory direction are probably based on an endogenous program, occur without the birds actually migrating and are independent of changes in the magnetic field. Our results provide further evidence that directional shifts in Sylvia warblers may be controlled by a different mechanism than in pied flycatchers (Ficedula hypoleuca).     

Empirical evidence for differential organ reductions during trans-oceanic bird flight

Since the early 1960s it has been held that migrating birds deposit and use only fat as fuel during migratory flight, with the non-fat portion of the body remaining homeostatic. Recent evidence from field studies has shown large changes in organ sizes in fuelling birds, and theory on fuel use suggests protein may be a necessary fuel during flight. However, an absence of information on the body condition of migrants before and after a long flight has hampered understanding of the dynamics of organs during sustained flight. We studied body condition in a medium-sized shorebird, the great knot (Calidris tenuirostris), before and after a flight of 5400 km from Australia to China during northward migration. Not only did these birds show the expected large reduction in fat content after migration, there was also a decrease in lean tissue mass, with significant decreases in seven organs. The reduction in functional components is reflected in a lowering of the basal metabolic rate by 42% [corrected]. Recent flight models have tried to separate the 'flexible' part of the body from the constant portion. Our results suggest that apart from brains and lungs no organs are homeostatic during long-distance flight. Such organ reductions may be a crucial adaptation for long-distance flight in birds.     

Antennal regulation of migratory flight in the neotropical moth Urania fulgens

Migrating insects use their sensory systems to acquire local and global cues about their surroundings. Previous research on tethered insects suggests that, in addition to vision and cephalic bristles, insects use antennal mechanosensory feedback to maintain their airspeeds. Owing to the large displacements of migratory insects and difficulties inherent in tracking single individuals, the roles of these sensory inputs have never been tested in freely migrating insects. We tracked individual uraniid moths (Urania fulgens) as they migrated diurnally over the Panama Canal, and measured airspeeds and orientation for individuals with either intact or amputated flagella. Consistent with prior observations that antennal input is necessary for flight control, 59 per cent of the experimental moths could not fly after flagella amputation. The remaining fraction (41%) was flight-capable and maintained its prior airspeeds despite severe reduction in antennal input. Thus, maintenance of airspeeds may not involve antennal input alone, and is probably mediated by other modalities. Moths with amputated flagella could not recover their proper migratory orientations, suggesting that antennal integrity is necessary for long-distance navigation.     

Homing pigeons as a model for avian navigation?

The findings on the navigational mechanisms of homing pigeons and the available data on those of wild birds, in particular migrants, are compared. There are important parallels in the use of the magnetic field and the sun for directional orientation. Also the findings on the navigational ‘map’, its preferred use by experienced birds and the strategy of using route information to acquire the necessary knowledge to establish the ‘map’, obtained in pigeons studies, can probably be generalized to wild birds and migrants in their home region. It seems that birds share a common navigational system. Special development of migratory birds, however, is the innate migration program that enables young first‐time migrants to reach their still unknown wintering area.