Influence of fuel load and weather on timing of nocturnal spring migratory departures in European robins, <Emphasis Type="Italic">Erithacus rubecula</Emphasis>

Time of departure and landing of nocturnal migrants are of great importance for understanding migratory strategy used by birds. It allows us to estimate flying time and hence the distance that migrants cover during a single night. In this paper, I studied the temporal schedule of nocturnal departures of European robins during spring migration. The study was done on the Courish Spit on the Baltic Sea in 1998–2003 by retrapping 51 ringed birds in high mist nets during nocturnal migratory departure. Take-offs of individual birds occurred between the first and tenth hour after sunset (median 176 min after sunset). Departure time was not related to fuel stores at arrival and departure, stopover duration and progress of the season. The results suggest that one reason for temporal variation in take-off time was differential response of European robins with high and low motivation to depart to such triggers as air pressure and its trend. If these parameters reach a certain minimum threshold shortly before sunset, robins with a high migratory motivation take off in the beginning of the night. When air pressure or its trend reaches a maximum, it may trigger to take off later during the night birds with lower initial motivation for departure, including those that have low refuelling efficiency. In regulation of timing of take-offs of robins, an important role is also played by their individual endogenous circadian rhythm of activity which is related to the environment in a complex way.     

Wind selectivity of migratory flight departures in birds

Optimal migration theory predicts that birds minimizing the overall time of migration should adjust stopover duration with respect to the rate of fuel accumulation. Recent theoretical developments also take into account the wind situation and predict that there is a time window (a set of days) during which birds should depart when assisted by winds but will not do so if there are head winds. There is also a final day when birds will depart irrespective of wind conditions. Hence, the wind model of optimal migration theory predicts that birds should be sensitive to winds and that there should be a correlation between departures and winds blowing towards the intended migration direction. We tested this assumption by tracking the departures of radio-tagged passerines during autumn migration in southern Sweden. Our birds were moderately to very fat when released and therefore energetically ready for departure. There was a significant correlation between direction of departure and wind direction. We also found that during days when birds departed there was a significantly larger tail wind component than during days when birds were present but did not depart. Our results show that passerines do take the current wind situation into account when departing on migratory flights. We also briefly discuss possible clues that birds use when estimating wind direction and strength. The inclusion of wind is an important amendment to optimal migration theory of birds and should be explored further. Received: 1 March 1999 / Received in revised form: 4 October 1999 / Accepted: 16 October 1999     

The strategy of fly-and-forage migration,illustrated for the osprey (<Emphasis Type="Italic">Pandion haliaetus</Emphasis>)

Migrating birds often alternate between flight steps, when distance is covered and energy consumed, and stopover periods, when energy reserves are restored. An alternative strategy is fly-and-forage migration, useful mainly for birds that hunt or locate their prey in flight, and thus, enables birds to combine foraging with covering migration distance. The favourability of this strategy in comparison with the traditional stopover strategy depends on costs of reduced effective travel speed and benefits of offsetting energy consumption during migration flights. Evaluating these cost-benefit effects, we predict that fly-and-forage migration is favourable under many conditions (increasing total migration speed), both as a pure strategy and in combination with stopover behaviour. We used the osprey (Pandion haliaetus) as test case for investigating the importance of this strategy during spring and autumn migration at a lake in southern Sweden. The majority, 78%, of passing ospreys behaved according to the fly-and-forage migration strategy by deviating from their migratory track to visit or forage at the lake, while 12% migrated past the lake without response, and 10% made stopovers at the lake. Foraging success of passing ospreys was almost as good as for birds on stopover. Timing of foraging demonstrated that the birds adopted a genuine fly-and-forage strategy rather than intensified foraging before and after the daily travelling period. We predict that fly-and-forage migration is widely used and important among many species besides the osprey, and the exploration of its occurrence and consequences will be a challenging task in the field of optimal migration.     

Factors influencing the movement biology of migrant songbirds confronted with an ecological barrier

Whether or not a migratory songbird embarks on a long-distance flight across an ecological barrier is likely a response to a number of endogenous and exogenous factors. During autumn 2008 and 2009, we used automated radio tracking to investigate how energetic condition, age, and weather influenced the departure timing and direction of Swainson’s thrushes (Catharus ustulatus) during migratory stopover along the northern coast of the Gulf of Mexico. Most birds left within 1 h after sunset on the evening following capture. Those birds that departed later on the first night or remained longer than 1 day were lean. Birds that carried fat loads sufficient to cross the Gulf of Mexico generally departed in a seasonally appropriate southerly direction, whereas lean birds nearly always flew inland in a northerly direction. We did not detect an effect of age or weather on departures. The decision by lean birds to reorient movement inland may reflect the suitability of the coastal stopover site for deposition of fuel stores and the motivation to seek food among more extensive forested habitat away from the barrier.     

Stopover behaviour and departure decision of northern wheatears, Oenanthe oenanthe, facing different onward non-stop flight distances

On the small North Sea island Helgoland (54°11' N, 07°55' E) we studied the stopover ecology of two subspecies of northern wheatear, Oenanthe oenanthe, during spring migration. Birds heading for Scandinavia (O. o. oenanthe) face only short flights across an ecological barrier (50-500 km) whereas those originating from Greenland and Iceland (O. o. leucorhoa) have to cover between 1,000 and 2,500 km in the impending flight. Colour-ringed individuals showed that 90% of Scandinavian birds left on the day of ringing while 40% (males) and 30% (females) of Greenland/Icelandic birds stayed at least 1 night. The birds who remained were thus mostly O. o. leucorhoa. They often established desirable feeding territories on the beach and had a high rate of body mass increase (1.7 g/day). However, subspecies did not differ in habitat choice and in foraging effort, but O. o. leucorhoa had a higher success rate in pecking. Departure decisions were analysed by comparing (a) conditions on the day of ringing between departing and staying birds and (b) for birds staying between the day of departure and the preceding day. The factors that were probably important in the decision to depart differed between subspecies. In O. o. leucorhoa, few birds departed with bad or deteriorating weather conditions (tailwind component, cloud cover), whereas departures of O. o. oenanthe seemed to be little affected by those factors. A few O. o. oenanthe stayed early in the spring migration season and/or had low fat reserves. Interference during foraging seemed to play a role because both subspecies tended to leave when the densities of northern wheatears were high. Other factors related to refuelling conditions (food supply, foraging effort, predation risk) failed to show differences between staying and departing individuals. In summary, almost all Scandinavian birds departed quickly and irrespective of refuelling and weather conditions, whereas many (but not all) Greenland/Icelandic birds seemed to prepare for a long-distance flight and carefully adjusted departure to weather conditions. The observed differences in stopover behaviour and departure decisions in the two subspecies of northern wheatear indicate that the distance to the next stopover site or to the goal area has to be considered when applying optimal migration models.     

Cover Caption

Cover: Radar imagery can be used to map the locations from which migratory birds depart and to highlight important stopover sites. Shown are movements of long-distance migrants, such as the Blackburnian Warbler ( Dendroica fusca ), detected during spring migration near Buffalo, New York. Warbler photo by Charles Eiseman. See pages 440–448.     

Differential timing of spring migration in northern wheatears Oenanthe oenanthe: hurried males or weak females?

In a field experiment on the island of Helgoland (southeast North Sea), we investigated whether migration strategy or competition between the sexes cause the differential timing of spring migration of male and female northern wheatears (Oenanthe oenanthe) (males migrating earlier). The study included two subspecies, heading towards Greenland/Iceland and Scandinavia, respectively, and is based on colour-ringing and remote weighing of individuals. Despite food offered ad libitum, most Scandinavian birds left the island on the day of arrival or stayed only 1–3 days, whereas more than half of Greenlandic/Icelandic birds stayed for up to 12 days and refuelled rapidly. In the latter subspecies, males showed a positive correlation of departure fuel load and fuel deposition rate, resembling time-minimizers in optimal migration theory. In contrast, females departed irrespective of fuel deposition rate, with an approximately constant level of fuel stores. This level was lower than in males, but sufficient to enable by-passing of stopover sites en route, allowing us to regard females as time-minimizers also. Since females are not able to reach Greenland without additional refuelling elsewhere and males appeared to have a larger potential for by-passing stopover sites, time-selection seems to be more pronounced in males and may be the reason for earlier migration of males. Intraspecific aggressive interactions between colour-ringed birds were predominantly won by the initiator, by males and by larger birds, whereas fuel load and subspecies did not affect the outcome. Although compared to females, males were more often dominant at the feeding stations or held territories, refuelling patterns could not be explained by dominance. Subordinate or non-territorial birds did not refuel at a lower rate or depart with lower fuel loads than dominant or territorial birds. In non-territorial birds, the restricted access to feeding stations was made up with larger doses of food taken per visit, leading to the same energy intake as that of dominant and territorial birds. Therefore, competition during stopover could be eliminated as the reason for differential timing of migration of male and female wheatears, but this result may be species-specific.Communicated by W. Wiltschko     

Corticosterone levels in relation to migratory readiness in red-eyed vireos (<Emphasis Type="Italic">Vireo olivaceus</Emphasis>)

We examined the relationship between plasma levels of corticosterone and the migratory activity and directional preference of red-eyed vireos during fall migration at the northern coast of the Gulf of Mexico. Corticosterone is thought to play a role in physiological and behavioural processes before, during, and after long-distance migratory flights. An increase in corticosterone at the onset of migratory flights can be expected in birds that are energetically prepared to migrate in a seasonally appropriate southerly direction. Red-eyed vireos ( Vireo olivaceus) were tested in orientation cages under clear twilight skies. Just prior to the orientation experiments, blood was sampled to assay baseline corticosterone levels. Average corticosterone level for all birds was 22.8 ng/ml. Red-eyed vireos with higher than average baseline levels of corticosterone were significantly more active in orientation cages compared to birds with lower levels of corticosterone. Moreover, birds with higher than average levels oriented in a southwesterly direction, which is consistent with a trans-Gulf flight, whereas individuals with levels below average showed a NNW mean direction. Although there was no significant difference in baseline levels of corticosterone between fat and lean birds, individual mass loss between capture and test was negatively correlated with corticosterone levels. Results from this study clearly demonstrate that corticosterone influences departure decisions and the choice of direction during migration.     

Reduced take-off ability in robins (Erithacus rubecula) due to migratory fuel load

Recent studies have shown that large fuel loads in small birds impair flying ability. This is the first study to show how migratory fuel load affects flying ability, such as velocity and height gained at take-off in a predator escape situation, in a medium-distance migrant, and whether they adjust their take-off according to predator attack angle. First-year robins (Erithacus rubecula) were subjected to simulated attacks from a model merlin (Falco columbarius), and take-off velocity and angle were analysed. Robins with a wing load of 0.19 g cm⁻² took off at a 39% lower angle than robins with a wing load of 0.13 g cm⁻², while velocity remained unaffected. The robins did not adjust their angle of ascent in accordance with the predator's angle of attack. Since many predators rely on surprise attacks, a difference in flight ability due to varying fuel loads found in migrating robins can be important for birds' chances of survival when actually attacked. Received: 28 October 1998 / Received in revised form: 12 January 1999 / Accepted: 30 January 1999     

Fuel loss and flexible fuel deposition rates in a long-distance migrant

Most migrating birds alternate flight bouts with stopovers, during which they rest and replenish the fuel used during flight (refueling). The rate of refueling (fuel deposition rate, FDR) affects stopover duration, and hence is an important determinant of the overall time required for migration. Although environmental and endogenous factors affect FDR, the urge to refuel depends on the anticipated distance to be travelled and possibly also on the amount of fuel used during the flight preceding stopover. Combining a field study with a fasting–refueling experiment on long-term captive songbirds, we tested whether the extent of fuel loss prior to refueling indeed affects FDR. In the field study, we took a comparative approach and determined FDR in two subspecies of northern wheatear (Oenanthe oenanthe) that differ greatly in the distance flown, and thus the extent of fuel used to reach our study (stopover) site. As both winter in western sub-Sahelian Africa, they face the same remaining migration distance. We found that FDR was higher in the subspecies that uses more fuel to get to our study site. Solidifying this result, in the experiment on captive northern wheatears, we found that the extent of fuel loss as a consequence of fasting explained most of the variation in subsequent FDR. The observation that experimental birds losing little fuel did not maximize their FDR suggests there are costs to rapid refueling. Our study shows that FDR is shaped not just by current environmental and endogenous conditions but also by fuel loss prior to refueling.     

Timing and flight mode of departure in migrating European bee-eaters in relation to multi-scale meteorological processes

Understanding departure decisions of migratory birds and the environmental factors affecting them is important for predicting their distribution, abundance, and arrival times to breeding and wintering areas. In the past, methodological difficulties to obtain fine-scale bird departure and meteorological data have limited testing the multi-scale effects of meteorology on bird departure during migration. We investigated departure timing of European bee-eaters (Merops apiaster) staging in southern Israel, identified their departure flight mode (flapping or soaring) using radio telemetry, and measured local meteorological conditions to study if bird departure was affected by these. Departure timing was examined using a timescale analysis design. The conditions before, during, and after the time of departure were compared using timescales of 24 h, 6 h, 1 h, and 10 min and in relation to bird flight mode. At the between-days timescale, barometric pressure at departure time was significantly lower compared with 2–1 day earlier, whereas temperature at departure was significantly higher compared with 3–2 days earlier. Temperature at departure was also higher compared with 6 h and 3–2 h earlier. Tailwind assistance had no significant effect at any timescale. Soaring birds departed at significantly higher temperature compared with flapping birds. We suggest that bee-eater departure is tuned to the infrequent passage of warm atmospheric depressions at the between-days timescale and with an increasing temperature trend within these days enabling the birds to use energetically cheap soaring flight. We thus suggest that energetic considerations dictate the departure decisions of migrating European bee-eaters.     

Is there a different response to winds during migration between soaring and flapping raptors? An example with the Montagu’s harrier and the lesser kestrel

During migrations, birds have to cope with varying meteorological conditions, which shape their migratory routes and affect their performance. Amongst these, wind is the main meteorological agent influencing behaviour of birds in their migration journeys. Here we analyze the effect of winds during migrations of adult individuals of two raptor species tracked with satellite telemetry, the Montagu’s harrier (Circus pygargus) and the lesser kestrel (Falco naumanni). While harriers use mostly soaring flight, kestrels principally use flapping flight and thus, wind can differently affect these birds. We found that both forward and perpendicular winds significantly affected the movements of the Montagu’s harrier, which were drifted from their intended direction but also took advantage of tailwinds. On the contrary, lesser kestrels moved more regardless of forward winds, despite they were highly drifted by crosswinds. Our results also support that the drifting effect of winds at the onset of the spring migration may explain the loop migration observed for both species, with birds compensating later the effect of crosswinds to arrive to their breeding areas. Results presented here illustrate how winds can differently affect migrating birds according to their flight modes.     

Sunset orientation of robins,Erithacus rubecula,with different fields of sky vision

Summary Migratory orientation of robins (Erithacus rubecula) at sunset was recorded using orientation cages, under clear autumn skies. The aim of the experiments was to examine the importance of different visible sky sections for the orientation of robins. I obtained the following results: (1) Robins tested with the visible sky section limited to 90° around zenith (45° above the horizon) showed a mean orientation that coincided with the average sunset azimuth, with little scatter around the mean angle (Fig. 2). (2) When the birds were allowed a more extensive field of sky vision (maximum 160°), they chose headings on an approximate north-south axis, significantly different from tests with a restricted view of the sky (Fig. 3). (3) Experiments were also performed in which the response of robins to a mirror deflection (about 120° counterclockwise) of visual cues in the lower parts of the sunset sky was examined. The outcome indicated that visual information in the lower part of the sky may be critical for the orientation of robins (Fig. 5). These results, together with recent findings that robins captured and tested at two nearby sites show distinctly different orientation behavior in relation to experimental manipulations of the magnetic field, suggest that priorities among orientation cues may differ depending on the migratory situation encountered.     

Obligatory barrier crossing and adaptive fuel management in migratory birds: the case of the Atlantic crossing in Northern Wheatears (<Emphasis Type="Italic">Oenanthe oenanthe</Emphasis>)

Behaviour on migration was often suggested to be selected for time-minimising strategies. Current optimality models predict that optimal fuel loads at departure from stopover sites should increase with increasing fuel deposition rates. We modified such models for the special case of the east Atlantic crossing of the Northern Wheatear (Oenanthe oenanthe). From optimality theory, we predict that optimal time-minimising behaviour in front of such a barrier should result in a positive correlation between fuel deposition rates and departure fuel loads only above a certain threshold, which is the minimum fuel load (f _min) required for the barrier crossing. Using a robust range equation, we calculated the minimum fuel loads for different barrier crossings and predict that time-minimising wheatears should deposit a minimum of 24% fuel in relation to lean body mass (m ₀ ) for the sea crossing between Iceland and Scotland. Fuel loads of departing birds in autumn in Iceland reached this value only marginally but showed positive correlation between fuel deposition rate (FDR) and departure fuel load (DFL). Birds at Fair Isle (Scotland) in spring, which were heading towards Iceland or Greenland, were significantly heavier and even showed signs of overloading with fuel loads up to 50% of lean body mass. Departure decisions of Icelandic birds correlated significantly with favourable wind situations when assuming a migration direction towards Spain; however, the low departure fuel loads contradict a direct non-stop flight.     

Autumn migratory fuelling: a response to simulated magnetic displacements in juvenile wheatears, Oenanthe oenanthe

Recent experiments exposing migratory birds to altered magnetic fields simulating geographical displacements have shown that the geomagnetic field acts as an external cue affecting migratory fuelling behaviour. This is the first study investigating fuel deposition in relation to geomagnetic cues in long-distance migrants using the western passage of the Mediterranean region. Juvenile wheatears (Oenanthe oenanthe) were exposed to a magnetically simulated autumn migration from southern Sweden to West Africa. Birds displaced parallel to the west of their natural migration route, simulating an unnatural flight over the Atlantic Ocean, increased their fuel deposition compared to birds experiencing a simulated migration along the natural route. These birds, on the other hand, showed relatively low fuel loads in agreement with earlier data on wheatears trapped during stopover. The experimental displacement to the west, corresponding to novel sites in the Atlantic Ocean, led to a simulated longer distance to the wintering area, probably explaining the observed larger fuel loads. Our data verify previous results suggesting that migratory birds use geomagnetic cues for fuelling decisions and, for the first time, show that birds, on their first migration, can use geomagnetic cues to compensate for a displacement outside their normal migratory route, by adjusting fuel deposition.     

Characteristics of Important Stopover Locations for Migrating Birds: Remote Sensing with Radar in the Great Lakes Basin

Abstract:  A preliminary stage in developing comprehensive conservation plans involves identifying areas used by the organisms of interest. The areas used by migratory land birds during temporal breaks in migration (stopover periods) have received relatively little research and conservation attention. Methodologies for identifying stopover sites across large geographic areas have been, until recently, unavailable. Advances in weather-radar technology now allow for evaluation of bird migration patterns at large spatial scales. We analyzed radar data (WSR-88D) recorded during spring migration in 2000 and 2001 at 6 sites in the Great Lakes basin (U.S.A.). Our goal was to link areas of high migrant activity with the land-cover types and landscape contexts corresponding to those areas. To characterize the landscapes surrounding stopover locations, we integrated radar and land-cover data within a geographic information system. We compared landscape metrics within 5 km of areas that consistently hosted large numbers of migrants with landscapes surrounding randomly selected areas that were used by relatively few birds during migration. Concentration areas were characterized by 1.2 times more forest cover and 9.3 times more water cover than areas with little migrant activity. We detected a strong negative relationship between activity of migratory birds and agricultural land uses. Examination of individual migration events confirmed the importance of fragments of forested habitat in highly altered landscapes and highlighted large concentrations of birds departing from near-shore terrestrial areas in the Great Lakes basin. We conclude that conservation efforts can be more effectively targeted through intensive analysis of radar imagery.     

Migratory fuelling in blackcaps (Sylvia atricapilla) under perceived risk of predation

It has been argued that the body mass levels achieved by birds are determined by the trade-off between risks of starvation and predation. Birds have also been found to reduce body mass in response to an increased predation risk. During migration, the need of extra fuel for flights is obvious and crucial. In this study, migratory blackcaps (Sylvia atricapilla) were subject to an experimental stopover situation where the predation risk was manipulated by exposure to a stuffed predator. Blackcaps that perceived an imminent risk of predation increased their food intake and fuel deposition rate during the first period of stopover compared with a control group. The pattern of night activity indicates that birds that were exposed to the predator also chose to leave earlier than birds in the control group. Since there was no cover present at the stopover site, birds might have perceived the risk of predation as higher regardless of whether they were foraging or not. Under such circumstances it has been predicted that birds should increase their foraging activity. The findings in this study clearly indicate that birds are able to adjust their stopover behaviour to perceived predation risk. Received: 8 January 1997 / Accepted after revision: 11 April 1997     

The good and the bad stopover: behaviours of migrant reed warblers at two contrasting sites

Successful migration for passerine birds depends largely on the quality of stopover habitats, but we still lack complete knowledge of how migrants search for habitats en route and how they behave when landing at poor quality stopover sites. We compared the distance of exploratory movements and stopover durations of the reed warbler Acrocephalus scirpaceus, a reedbed habitat specialist, released at suitable (reed bed) and unsuitable (sand dune) stopover sites. Birds tape-lured during nocturnal migration to a sand dune were captured, radio-tagged, released and tracked at two sites of contrasting habitat quality. Lean birds were found to move further in the dunes (max. 300 m) than in reeds (max. 200 m), whereas ‘fat’ individuals at both sites remained stationary. Birds spent just 1 day in the dunes and up to 13 days in the reeds. Our results suggest that some nocturnal migrants with restricted diurnal exploratory movements depend on stopover site selection when ceasing nocturnal flight.     

Optimal Management of a Multispecies Shorebird Flyway under Sea‐Level Rise

Every year, millions of migratory shorebirds fly through the East Asian–Australasian Flyway between their arctic breeding grounds and Australasia. This flyway includes numerous coastal wetlands in Asia and the Pacific that are used as stopover sites where birds rest and feed. Loss of a few important stopover sites through sea‐level rise (SLR) could cause sudden population declines. We formulated and solved mathematically the problem of how to identify the most important stopover sites to minimize losses of bird populations across flyways by conserving land that facilitates upshore shifts of tidal flats in response to SLR. To guide conservation investment that minimizes losses of migratory bird populations during migration, we developed a spatially explicit flyway model coupled with a maximum flow algorithm. Migratory routes of 10 shorebird taxa were modeled in a graph theoretic framework by representing clusters of important wetlands as nodes and the number of birds flying between 2 nodes as edges. We also evaluated several resource allocation algorithms that required only partial information on flyway connectivity (node strategy, based on the impacts of SLR at nodes; habitat strategy, based on habitat change at sites; population strategy, based on population change at sites; and random investment). The resource allocation algorithms based on flyway information performed on average 15% better than simpler allocations based on patterns of habitat loss or local bird counts. The Yellow Sea region stood out as the most important priority for effective conservation of migratory shorebirds, but investment in this area alone will not ensure the persistence of species across the flyway. The spatial distribution of conservation investments differed enormously according to the severity of SLR and whether information about flyway connectivity was used to guide the prioritizations. With the rapid ongoing loss of coastal wetlands globally, our method provides insight into efficient conservation planning for migratory species. Gestión Óptima de una Ruta Migratoria de Múltiples Especies de Aves Costeras Sometida a Incremento del Nivel del Mar