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.     

Flexibility in daily travel routines causes regional variation in bird migration speed

We used novel Global Positioning System-based satellite telemetry to reconstruct daily time budgets on travelling days of a long-distance migrant, the Osprey Pandion haliaetus, to reveal how landscape affects migratory performance. We compared daily travel routines between the Ospreys’ passage of Europe and the Sahara. In Europe, where feeding habitat is abundant, Ospreys fed both before–after flights and during interruptions, thus, combining migration with foraging. This resulted in a 2.7-h shorter daily flight period in Europe than in the Sahara. A calculated energy budget indicated that a ‘fly-and-forage migration strategy’ is favourable in Europe because associated benefits (energy intake) more than outweigh costs (reduced flight time). The much shorter flight time in Europe was the main explanation why Ospreys covered on average 78 km less distance on a travelling day in Europe than in the Sahara. In addition, there were regional differences in hourly flight speeds that are most probably the result of variation in thermal soaring conditions. We conclude that landscape properties have a profound effect on migration through regional variation in daily routines.     

The nature of the migration route shapes physiological traits and aerodynamic properties in a migratory songbird

Migration distance is supposed to represent an important selection pressure shaping physiological and morphological properties. Previous work has focussed on this effect, while the importance of ecological barriers in this context has been rarely considered. We studied two subspecies of a migratory songbird, the northern wheatear (Oenanthe oenanthe oenanthe and O. o. leucorhoa L.), on an island in the North Sea. The former subspecies reaches their Scandinavian breeding areas after a short sea crossing, whereas leucorhoa northern wheatears cross the North Atlantic towards Iceland, Greenland or Canada. Physiological traits (fuel deposition rate) and wings’ aerodynamic properties (wing pointedness independent of body size), both affecting migration speed, were hypothesized to be more pronounced in leucorhoa than in oenanthe northern wheatears. Within subspecies, the physiological and aerodynamic properties were hypothesized to explain arrival date at the stopover site with “fast migrants” arriving early. Physiological and aerodynamic properties in leucorhoa northern wheatears lead to a faster and less costly migration, favouring a sea crossing, but in trade-off lower flight manoeuvrability than in oenanthe birds. Wings’ aerodynamic properties affected the seasonal occurrence of leucorhoa females, whereas the physiological traits significantly influenced arrival date in oenanthe individuals. The less risky migration route in oenanthe birds with few short sea crossings may have favoured higher flight manoeuvrability for foraging (less pointed wings), in trade-off an energetically more costly flight. Hence, not the migration distance itself, but the presence/absence of a sea barrier presents an important selection pressure in migratory land birds favouring low flight costs.     

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.     

The fly-and-social foraging hypothesis for diurnal migration: why American crows migrate during the day

Most migratory species migrate at night, and the benefits associated with nocturnal migration have been well reviewed; however, less attention has been paid to the benefits associated with diurnal migration. There are two theories for diurnal migration: (1) diurnal migration allows for the use of thermals and therefore reduces energy loss, and (2) the fly-and-forage hypothesis, which suggests that diurnal migrants can search for prey and forage as they migrate. We investigated whether American crows (Corvus brachyrhynchos) engage in the fly-and-forage strategy of diurnal migration as they migrated north in the spring. We tracked eight radio-tagged crows as they migrated hundreds of kilometers in the spring to determine if they were diurnal migrants, whether they migrated with conspecifics, whether they stopped to forage, and if they did stop, whether they forage with conspecifics. All crows migrated during the day, and while on several occasions crows were seen migrating in close association with conspecifics, all crows were also observed migrating alone. On average, crows migrated approximately 300 km the day they left their wintering grounds, and over the course of this day, they stopped twice and foraged at these locations for 35 min. On all but one occasion, the stops made during migration were to forage with groups of conspecifics. While the fly-and-forage hypothesis for diurnal migration has primary been applied to raptors, many diurnal migrants forage socially, and the presence of foraging conspecifics and/or heterospecifics may be a significant benefit in locating food resources and ultimately migrating during the day.     

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.     

Influence of body condition and weather on departures of first-year European robins, <Emphasis Type="Italic">Erithacus rubecula</Emphasis>, from an autumn migratory stopover site

How migratory birds decide when to leave a stopover site is important to the understanding of bird migration strategies. Our study looks at how body condition and the weather affect the decision to depart on nocturnal migratory flight. During two autumn migration seasons (2002–2003), we radio tracked 51 first-year European robins, Erithacus rubecula, at a stopover site on the Courish Spit (Eastern Baltic) from the first day after landing until their migratory departure. The tagged robins stopped over for 1–14 days. There was no clear relationship between stopover duration and energetic condition on arrival. Weather conditions (wind, precipitation, and cloud cover) on departure differed measurably between years. In 2002, robins took off mainly under following winds and clear skies. In 2003, there were mainly light head winds and partially cloudy or overcast skies. This could be explained by the year-specific role of weather factors in making the decision to depart. In both years, robins making short (1–2 days) stopovers took off in more varied weather situations than those individuals with long stopovers. This suggests that robins from the former group were more inclined to continue with migration than longer-stay birds that, apart from re-fuelling, could be waiting for favourable weather. The lack of a relationship between stopover duration and body condition and some departures under unfavourable weather conditions suggest that endogenous spatiotemporal programmes may play an important role in controlling stopover duration in robins.     

Persistent bimodal activity patterns in wild and captive black-tailed godwit <Emphasis Type="Italic">Limosa limosa</Emphasis> under different environmental conditions

There is a large literature dealing with daily foraging routines of wild birds during the non-breeding season. While different laboratory studies have showed that some bird activity patterns are a persistent property of the circadian system, most of field studies preclude the potential role of an endogenous circadian rhythm in controlling bird’s foraging routines. In this study we compared the patterns of diurnal foraging activity and intake rates of migrating black-tailed godwits, Limosa limosa (radio-tagged and non-tagged individuals) at two stopover sites (habitats) with different environmental characteristics, aiming at identifying proximate factors of bird activity routines. To gain insights into the role of food availability in control of such foraging routines, we also estimated foraging activity patterns in captive godwits subjected to constant food availability. Captive and wild black-tailed godwits showed a persistent bimodal activity pattern through daylight period. Food availability had a significant effect on the intake rates, but had a subtler effect on foraging and intake rate rhythms. Temperature and wind speed (combined in a weather index) showed non-significant effects on both rhythms. Although we could not discard a role for natural diurnal changes in light intensity, an important timing cue, our findings support the idea that an endogenous circadian rhythm could be an important proximate factor regulating foraging activity and food items taken per unit time of wild black-tailed godwits during migration.     

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.     

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     

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     

Energy budgets reveal equal benefits of varied migration strategies in northern gannets

We investigated migration and wintering of adult northern gannets (Morus bassanus) breeding in east Scotland, North Sea, by deploying geolocation loggers over three winters. The wintering ranges of these birds varied from the North Sea to the Atlantic off West Africa. Flight time was taken as a proxy for migration and foraging effort. Gannets wintering off Africa had higher total flight times during migration than birds wintering further north. Total flight times in different wintering regions were generally low. Birds off West Africa consistently spent < 25 % of daylight hours in flight, but birds further north showed more variable values that may reflect more variable weather or food availability. Winter sea surface temperatures ranged from 9 °C (North Sea) to 16 °C (West Africa). Thermostatic costs in winter as estimated by measuring thermal conductance in carcasses in still air and water were 28 % higher in North Sea than off West Africa. This effect is aggravated by higher thermostatic costs caused by stronger wind chills in the North Sea compared to the conditions off West Africa. Birds wintering close to the UK arrived at the colony on average 12 days earlier than birds wintering off West Africa. We conclude that the net cost-benefit analysis may be similar for all wintering areas investigated.     

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     

Evidence of food-based competition among passerine migrants during stopover

Summary Local concentrations of migrating, fat-depleted birds with similar diets can lead to increased competition for food at a time when energy demand is high. Results of a predator-exclosure experiment indicate that intercontinental passerine migrants depress food abundance during stopover following migration across the Gulf of Mexico. Moreover, migrants that stop when a high number of potential competitors are present replenish energy reserves more slowly than migrants that stop under low density conditions. Competition increase the rate of food depletion and may decrease the probability that a migrant will meet its energetic requirements and complete a successful migration.     

Integration of distance, direction and habitat into a predictive migratory movement model for blue-winged teal (Anas discors)

Historically, the migration of birds has been poorly understood in comparison to other life stages during the annual cycle. The goal of our research is to present a novel approach to predict the migratory movement of birds. Using a blue-winged teal case study, our process incorporates not only constraints on habitat (temperature, precipitation, elevation, and depth to water table), but also approximates the likely bearing and distance traveled from a starting location. The method allows for movement predictions to be made from unsampled areas across large spatial scales. We used USGS’ Bird Banding Laboratory database as the source of banding and recovery locations. We used recovery locations from banding sites with multiple within-30-day recoveries were used to build core maximum entropy models. Because the core models encompass information regarding likely habitat, distance, and bearing, we used core models to project (or forecast) probability of movement from starting locations that lacked sufficient data for independent predictions. The final model for an unsampled area was based on an inverse-distance weighted averaged prediction from the three nearest core models. To illustrate this approach, three unsampled locations were selected to probabilistically predict where migratory blue-wing teals would stopover. These locations, despite having little or none data, are assumed to have populations. For the blue-winged teal case study, 104 suitable locations were identified to generate core models. These locations ranged from 20 to 228 within-30-day recoveries, and all core models had AUC scores greater than 0.80. We can infer based on model performance assessment, that our novel approach to predicting migratory movement is well-grounded and provides a reasonable approximation of migratory movement.     

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.     

The effect of reproductive condition on the foraging behavior of female hoary bats,Lasiurus cinereus

Summary Female mammals experience larg changes in time and energy budgets associated with reproduction and these may influence the foraging strategies of individuals. I studied the changes in foraging behavior associated with reproduction in female hoary bats, Lasiurus cinereus. As lactation progressed, individuals departed to forage earlier in the evening and spent more time foraging per night and less time roosting with their young. Foraging time increased by at least 73% between early lactation and fledging and then declined as the young became independent. Females with two young foraged for longer than did those with one and females with pre- and postfledging young foraged in different habitats. The changes in foraging time suggest that foraging activity of female L. cinereus is constrained and individuals act as time minimizers, adjusting their foraging behavior to meet current energy demand. Predation risk is unlikely to constrain the behavior of these bats. However, maximizing energy intake throughout lactation may not be the optimal strategy because storing excess energy increases flight cost and may reduce foraging efficiency. The need to keep newborn young warm may also influence foraging time. Such constraints, causing changes in foraging activity, may alter the availability of habitats and prey and must be considered when modelling foraging strategies. In addition, changes in flight time may significantly alter the energy budgets of bats in different stages of reproduction.