Differential migration of the sexes cannot be explained by the body size hypothesis in Teal

The “body size hypothesis” predicts that if individuals of a population migrate different distances from the breeding to the wintering grounds, the distance should be related to the differential ability to cope with adverse conditions, with larger individuals wintering further north. Data collected over a 40-year period in Essex, UK and the Camargue, southern France, revealed that the average body mass of Teal ringed in Essex during these years was actually not greater than that of Teal ringed in the Camargue. A higher proportion of males were included in the UK ringing catch than in the French catch, but we found no support for the body size hypothesis to explain such differential migration of the sexes.     

Seasonal variation in migration strategies used to cross ecological barriers in a nearctic migrant wintering in Africa

Ecological barriers such as oceans, mountain ranges or glaciers can have a substantial influence on the evolution of animal migration. Along the migration flyway connecting breeding sites in the North American Arctic and wintering grounds in Europe or Africa, nearctic species are confronted with significant barriers such as the Atlantic Ocean and the Greenland icecap. Using geolocation devices, we identified wintering areas used by ringed plovers nesting in the Canadian High‐Arctic and investigated migration strategies used by these nearctic migrants along the transatlantic route. The main wintering area of the ringed plovers (n = 20) was located in western Africa. We found contrasting seasonal migration patterns, with ringed plovers minimizing continuous flight distances over the ocean in spring by making a detour to stop in Iceland. In autumn, however, most individuals crossed the ocean in one direct flight from southern Greenland to western Europe, as far as southern Spain. This likely resulted from prevailing anti‐clockwise winds associated with the Icelandic low‐pressure system. Moreover, the plovers we tracked largely circumvented the Greenland icecap in autumn, but in spring, some plovers apparently crossed the icecap above the 65°N. Our study highlighted the importance of Iceland as a stepping‐stone during the spring migration and showed that small nearctic migrants can perform non‐stop transatlantic flights from Greenland to southern Europe.     

Loop migration in adult marsh harriers Circus aeruginosus,as revealed by satellite telemetry

Loop migration among birds is characterized by the spring route lying consistently west or east of the autumn route. The existence of loops has been explained by general wind conditions or seasonal differences in habitat distribution. Loop migration has predominantly been studied at the population level, for example by analysing ring recoveries. Here we study loop migration of individual marsh harriers Circus aeruginosus tracked by satellite telemetry. We show that despite a generally narrow migration corridor the harriers travelled in a distinct clockwise loop through Africa and southern Europe, following more westerly routes in spring than in autumn. We used the Normalized Difference Vegetation Index (NDVI) to identify potential feeding habitat in Africa. Suitable habitat seemed always more abundant along the western route, both in spring and autumn, and no important stopover site was found along the eastern route. Observed routes did thus not coincide with seasonal variation in habitat availability. However, favourable habitat might be more important during spring migration, when the crossing of the Sahara seems more challenging, and thus habitat availability might play an indirect role in the harriers’ route choice. Grid‐based wind data were used to reconstruct general wind patterns, and in qualitative agreement with the observed loop marsh harriers predominantly encountered westerly winds in Europe and easterly winds in Africa, both in autumn and in spring. By correlating tail‐ and crosswinds with forward and perpendicular movement rates, respectively, we show that marsh harriers are partially drifted by wind. Thus, we tentatively conclude that wind rather than habitat seems to have an overriding effect on the shape of the migration routes of marsh harriers. General wind conditions seem to play an important role also in the evolution of narrow migratory loops as demonstrated for individual marsh harriers.     

Migration patterns of Hoopoe Upupa epops and Wryneck Jynx torquilla: an analysis of European ring recoveries

For many bird species, recovery of ringed individuals remains the best source of information about their migrations. In this study, we analyzed the recoveries of ringed European Hoopoe (Upupa epops) and the Eurasian Wryneck (Jynx torquilla) from 1914 to 2005 from all European ringing schemes. The aim was to define general migration directions and to make inferences about the winter quarters, knowing that hardly any recoveries are available from sub-Saharan Africa. For the autumn migration, there is evidence of a migratory divide for the Hoopoe in Central Europe, at approximately 10–12°E. Autumn migration directions of Wrynecks gradually change from SW to SE depending on the longitude (west to east) of the ringing place. In both species, only a few recoveries were available indicating spring migration directions, but they showed similar migration axes as for autumn migration, and hence no evidence for loop-migration. Due to a paucity of recoveries on the African continent, we can make only limited inferences about wintering grounds: extrapolating migration directions are only indicative of the longitude of the wintering area. The directions of autumn migration indicate a typical pattern observed in European long-distance migrants: west-European Hoopoes and Wrynecks are likely to winter in western Africa, while central- and east-European birds probably winter more in the east. Due to the migratory divide, for the Hoopoe, this phenomenon is more pronounced.     

Circannual variation in blood parasitism in a sub‐Saharan migrant passerine bird,the garden warbler

Knowing the natural dynamics of pathogens in migratory birds is important, for example, to understand the factors that influence the transport of pathogens to and their transmission in new geographical areas, whereas the transmission of other pathogens might be restricted to a specific area. We studied haemosporidian blood parasites of the genera Plasmodium, Haemoproteus and Leucocytozoon in a migratory bird, the garden warbler Sylvia borin. Birds were sampled in spring, summer and early autumn at breeding grounds in Sweden, on migration at Capri, Italy and on arrival and departure from wintering staging areas in West Africa: mapping recoveries of garden warblers ringed in Fennoscandia and Capri showed that these sites are most probably on the migratory flyway of garden warblers breeding at Kvismaren. Overall, haemosporidian prevalence was 39%, involving 24 different parasite lineages. Prevalence varied significantly over the migratory cycle, with relatively high prevalence of blood parasites in the population on breeding grounds and at the onset of autumn migration, followed by marked declines in prevalence during migration both on spring and autumn passage. Importantly, we found that when examining circannual variation in the different lineages, significantly different prevalence profiles emerged both between and within genera. Our results suggest that differences in prevalence profiles are the result of either different parasite transmission strategies or coevolution between the host and the various parasite lineages. When separating parasites into common vs. rare lineages, we found that two peaks in the prevalence of rare parasites occur; on arrival at Swedish breeding grounds, and after the wintering period in Africa. Our results stress the importance of appropriate taxonomic resolution when examining host‐parasite interactions, as variation in prevalence both between and within parasite genera can show markedly different patterns.     

European flyway permeability and abmigration in Teal Anas crecca, an analysis based on ringing recoveries

Abmigration, the switch between flyways, is thought to be frequent in dabbling ducks, and supposedly results from high female philopatry combined with winter pairing: a male pairing in winter with a female from another flyway may follow her to her breeding area and thus abmigrate. On this basis, the frequency of abmigration should be much higher amongst males than amongst females, should increase with time since ringing, and be higher if environmental conditions force birds from different flyways into the same geographical area in winter (in this case the Iberian peninsula in the case of cold spells). Analysing more than 9000 recoveries of Teal ringed in the Camargue, southern France, showed that even with the most conservative flyway definition, 15% of Camargue (Mediterranean flyway) individuals (both males and females) were subsequently recovered in the North-west European flyway. Moving a flyway boundary closer to the Camargue (i.e. to the south-east) increased this percentage to c . 32%. None of the predicted patterns were supported by the data: there was no systematic sex difference in abmigration rate; abmigration showed no consistent increase with time from ringing, and was not significantly more frequent after cold spells. These results suggest that there is permeability between these two flyways for all classes of individuals, which might better be considered as a continuum. This should be taken into account in the development of management and conservation policies for Teal in Europe.     

Migration and moult of Dunlin Calidris alpina wintering in the central Mediterranean

Dunlin migration in northeast Italy is described. An attempt to identify the main routes and staging areas used by birds wintering in the central Mediterranean is presented. The results of monthly counts from 1990–1995 revealed that the bulk of the population occupied the wintering area in October and left for the breeding grounds in April and May. The analysis of 342 Italian recoveries of foreign ringed birds showed that 65% were ringed during post-breeding migration through the Baltic Sea, whereas just a few birds had been ringed in western Europe. First-year birds arrived in autumn with a single migratory wave, peaking in October. Two categories of adults were identified during post-breeding migration: birds which directly reached Italian wintering sites and birds which arrived after they had suspended their migration for moulting: the Azov/Black Sea wetlands are suggested as possible moulting areas. Out of 2444 adults and 1627 first-years ringed between 1989 and 1996 at our study area, we obtained a total of 42 recoveries abroad and evidence of direct links between Azov/Black Sea and N Adriatic wetlands, both during autumn and spring migrations. Primary moult was observed only in adults arriving early, the second migratory wave being composed of moulted birds. Locally moulting adults adopted a moult strategy characterized by high raggedness scores, typical of resident moulters. Body mass was not affected by primary moult stage or intensity, winter mass values being reached two weeks after the average date of primary moult completion.     

Around the Mediterranean: an extreme example of loop migration in a long‐distance migratory passerine

An important issue in migration research is how small‐bodied passerines pass over vast geographical barriers; in European–African avian migration, these are represented by the Mediterranean Sea and the Sahara Desert. Eastern (passing eastern Mediterranean), central (passing Apennine Peninsula) and western (via western Mediterranean) major migration flyways are distinguished for European migratory birds. The autumn and spring migration routes may differ (loop migration) and there could be a certain level of individual flexibility in how individuals navigate themselves during a single migration cycle. We used light‐level loggers to map migration routes of barn swallows Hirundo rustica breeding in the centre of a wide putative contact zone between the northeastern and southernwestern European populations that differ in migration flyways utilised and wintering grounds. Our data documented high variation in migration patterns and wintering sites of tracked birds (n = 19 individuals) from a single breeding colony, with evidence for loop migration in all but one of the tracked swallows. In general, two migratory strategies were distinguished. In the first, birds wintering in a belt stretching from southcentral to southern Africa that used an eastern route for both the spring and autumn migration, then shifted their spring migration eastwards (anti‐clockwise loops, n = 12). In the second, birds used an eastern or central route to their wintering grounds in central Africa, shifting the spring migration route westward (clockwise loops, n = 7). In addition, we observed an extremely wide clockwise loop migration encompassing the entire Mediterranean, with one individual utilising both the eastern (autumn) and western (spring) migratory flyway during a single annual migration cycle. Further investigation is needed to ascertain whether clockwise migratory loops encircling the entire Mediterranean also occur other small long‐distance passerine species.     

Seasonal migrations of four individual bar-headed geese Anser indicus from Kyrgyzstan followed by satellite telemetry

The Kyrgyz population of the bar-headed goose Anser indicus has declined dramatically during the past decades. Human persecution during migration and habitat loss at stopover and wintering sites are commonly regarded as most serious threats. However, little is known about seasonal movements, migration routes, and wintering sites of the bar-headed geese from Kyrgyzstan, which represent the westernmost geographical population of the species. As part of a conservation project, which also included reinforcement of the wild population by the release of hand-reared juveniles, in late summer of 1998, five bar-headed geese, three wild adults and two hand-reared goslings, were fitted with sun-powered satellite transmitters in order to track their movements from Lake Son Kul and Lake Chatyr Kul in Kyrgyzstan. The five individuals contributed very unevenly to the more than 5,000 signals in total that were received from the French ARGOS system: one failed after 8 weeks, while another one was tracked for more than 2 years. The four geese contributing to this study followed three completely different migration routes leading to their wintering areas in Pakistan, India and Uzbekistan, while stopover areas were situated in southern Tajikistan and in western Tibet. Both in autumn and spring the adult birds migrated distances of 1,280–1,550 km in two steps, with stopover periods of 32–46 days (autumn) and 16–23 days (spring). Flight speeds of up to 680 km per actual migration day were recorded regularly, even during crossings of very high summits. A hand-reared juvenile flew non-stop for 790 km to southern Uzbekistan and even visited southernmost Turkmenistan, where the species is very rarely seen. The timing of migration varied considerably between individuals but also for the same individual between years. We compare our tracking results with previous findings (field observations, ring recoveries, and satellite tracking results) and discuss them with respect to migration over high-mountain habitats and a general migration strategy of the species.     

Modelling the Progression of Bird Migration with Conditional Autoregressive Models Applied to Ringing Data

Migration is a fundamental stage in the life history of several taxa, including birds, and is under strong selective pressure. At present, the only data that may allow for both an assessment of patterns of bird migration and for retrospective analyses of changes in migration timing are the databases of ring recoveries. We used ring recoveries of the Barn Swallow Hirundo rustica collected from 1908–2008 in Europe to model the calendar date at which a given proportion of birds is expected to have reached a given geographical area (‘progression of migration’) and to investigate the change in timing of migration over the same areas between three time periods (1908–1969, 1970–1990, 1991–2008). The analyses were conducted using binomial conditional autoregressive (CAR) mixed models. We first concentrated on data from the British Isles and then expanded the models to western Europe and north Africa. We produced maps of the progression of migration that disclosed local patterns of migration consistent with those obtained from the analyses of the movements of ringed individuals. Timing of migration estimated from our model is consistent with data on migration phenology of the Barn Swallow available in the literature, but in some cases it is later than that estimated by data collected at ringing stations, which, however, may not be representative of migration phenology over large geographical areas. The comparison of median migration date estimated over the same geographical area among time periods showed no significant advancement of spring migration over the whole of Europe, but a significant advancement of autumn migration in southern Europe. Our modelling approach can be generalized to any records of ringing date and locality of individuals including those which have not been recovered subsequently, as well as to geo-referenced databases of sightings of migratory individuals.     

Migration of red‐backed shrikes from the Iberian Peninsula: optimal or sub‐optimal detour?

The current Northern Hemisphere migration systems are believed to have arisen since the last glaciation. In many cases, birds do not migrate strait from breeding to non‐breeding areas but fly via a detour. All western European populations of red‐backed shrikes Lanius collurio are assumed to reach their southern African wintering grounds detouring via southeast Europe. Based on theoretical considerations under an optimality framework this detour is apparently optimal. Here, we use individual geolocator data on red‐backed shrikes breeding in Spain to show that these birds do indeed detour via southeast Europe en route to southern Africa where they join other European populations of red‐backed shrikes and return via a similar route in spring. Disregarding potential wind assistance, the routes taken for the tracked birds in autumn were not optimal compared to crossing the barrier directly. For spring migration the situation was quite different with the detour apparently being optimal. However, when considering potential wind assistance estimated total air distances during autumn migration were overall similar and the barrier crossing shorter along the observed routes. We conclude that considering the potential benefit of wind assistance makes the route via southeast Europe likely to be less risky in autumn. However, it cannot be ruled out that other factors, such as following a historical colonisation route could still be important.     

Fuelling rates of garganey ( Anas querquedula) staging in the Camargue, southern France, during spring migration

Most species of long-distance migratory birds put on energy stores to fuel their travels. However, recent studies have highlighted the potential costs associated with carrying too much fuel, either through increased predation risk or decreased flight efficiency. Consequently, it is now widely accepted that migratory birds should carry optimal rather than maximum fuel loads. Information from 372 garganey (Anas querquedula) ringed and recaptured at least once during the same spring in the Camargue, southern France, was used to document fuelling rates of individual ducks in relation to environmental variation and individual variation in condition. On average, garganey added very little fuel stores in the Camargue (mean gain per day=0.33 g, less than 0.5% of mean body-mass in total over an average stay of 5 days). Fuelling rates were negatively correlated with body mass at capture, but it cannot be excluded that this pattern was a statistical artefact. Given their body-mass at ringing, garganey could potentially still fly long distances when they stop in the Camargue. It is therefore likely that the aim of their stay in southern France is more for resting than refuelling, a finding that may have implications for the proper management of stop-over sites.Communicated by F. Bairlein     

Real-time weather analysis reveals the adaptability of direct sea-crossing by raptors

Many animals seasonally travel between their breeding and wintering grounds. With their advanced mobility, birds often migrate over thousands of kilometres. Recently, satellite-tracking studies have revealed peculiar migration routes for some avian species at a global scale. However, the adaptability of such migration routes has not been clearly demonstrated. Using satellite-tracking data for 33 individuals, we show that the Japanese population of Oriental honey-buzzards (Pernis ptilorhynchus) directly crosses the 650-km-wide East China Sea during their autumn migration, although they fly a longer route around the sea rather than directly crossing it during their spring migration. By applying aerodynamic theory, we show that the buzzards could cross the sea by soaring and gliding flight. Moreover, using a high-resolution meteorological-prediction analysis, we demonstrate that the migratory trajectory of the birds strongly depends on the wind direction at their estimated locations. In the area, northeastern tailwinds blow stably only during autumn. Thermals were abundant ca. 500–1,000 m over the East China Sea in autumn, but that was not the case in spring. We suggest that the autumn-migration route across the East China Sea is likely to have evolved in response to the specific weather conditions over the sea. Animations showing movements of Oriental honey-buzzards and temporal change in weather conditions are available at: , , , , , and .     

Wind conditions facilitate the seasonal water‐crossing behaviour of Oriental Honey‐buzzards Pernis ptilorhynchus over the East China Sea

Migratory raptors rarely fly over stretches of water larger than 25 km, although different species undertake water crossings of varying lengths, depending mainly on their wing morphology. Oriental Honey‐buzzards fly c. 680 km over the East China Sea in autumn from breeding areas in Japan to wintering areas in Southeast Asia, but avoid this long water crossing in spring. We investigated the effects of weather on this exceptional migratory behaviour and its seasonality through a maximum entropy niche modelling approach. We used data collected through satellite tracking of 31 adult birds as presence points and a set of variables related to wind, precipitation and convective condition as environmental predictors. Results of modelling showed very different, almost non‐overlapping, areas suitable for migration over the East China Sea region in autumn and spring. Suitable migration routes in autumn mostly occurred over the sea, whereas suitable areas for spring migration mostly occurred over land, suggesting that circumnavigating the East China Sea is preferable in spring. At the regional scale, wind conditions facilitate water‐crossing behaviour of Oriental Honey‐buzzards in autumn, but not in spring. Specifically, suitable tailwinds over the sea enable water‐crossing in autumn, whereas in spring, wind support and convective conditions are best over land. Our modelling did not suggest any importance of convective conditions for autumn migration. However, we expect that at smaller temporal scales, convective conditions would be a considerable facilitator of the water‐crossing behaviour in this species.     

Determining the boundaries of migratory bird flyways: a Bayesian model for Eurasian teal Anas crecca in western Europe

Accurate flyway delineation is a prerequisite for effective conservation and management of migratory bird populations, yet such limits have so far mostly been set subjectively. We present a statistical method to infer population boundaries from the analysis of ring recoveries, using a Bayesian framework. The approach was applied to Eurasian teal Anas crecca ringed in Camargue, southern France, and Abberton Reservoir, Essex, eastern England. The results presented show the boundaries of the two teal flyways in western Europe, with a zone of overlap, broadly matching those previously defined. The percentage of teal switching flyways (abmigration rate) was 2.4–2.6%, greater in birds ringed as juveniles than as adults. Abmigrants ended up at sites within the other flyway where the density of local birds was lower than expected by chance, suggesting abmigration resulted from exploratory or aberrant behaviour. The methodology presented here can be used to infer flyway boundaries of any bird with an adequate ring‐re‐encounter dataset, which has crucial consequences for the evaluation of their trends in abundance and hence conservation status, and the management of sustainable harvests.     

A flyway perspective on food resource abundance in a long-distance migrant, the Eurasian teal (Anas crecca)

Two frequent assumptions about the evolution of long-distance migration in birds are that they travel long distances annually to reach food-rich areas for breeding, and that they time their migratory journey to be at staging sites when the latter provide the best feeding conditions. These assumptions have rarely been properly tested, and there is no study in which a species’ major food types have been measured by standardized methods throughout a flyway and over a large part of the year. We here present such data for Eurasian teal (Anas crecca), converted to a common energetic currency, and collected at wintering, spring staging and breeding sites. Teal did not time migration to maximize local food abundance; most birds left wintering and spring staging sites before a sharp increase in invertebrate food abundance occurred. On the other hand, hatching of ducklings coincided with a peak in invertebrate food abundance on boreal breeding lakes. Mean overall food abundance (invertebrates and seeds combined) did not differ between wintering sites in southern France and breeding sites in northern Sweden at the time of breeding. Our results are inconsistent with the hypothesis that long-distance migration in dabbling ducks has evolved because adult birds gain an immediate pay-off in increased food abundance by flying north in spring. However, our data confirm a selective advantage for breeding at higher latitudes, because hatching of ducklings may coincide with a peak in invertebrate emergence and because longer days may increase the duration of efficient foraging.     

Discovering the migration and non‐breeding areas of sand martins and house martins breeding in the Pannonian basin (central‐eastern Europe)

The central‐eastern European populations of sand martin and house martin have declined in the last decades. The drivers for this decline cannot be identified as long as the whereabouts of these long distance migrants remain unknown outside the breeding season. Ringing recoveries of sand martins from central‐eastern Europe are widely scattered in the Mediterranean basin and in Africa, suggesting various migration routes and a broad non‐breeding range. The European populations of house martins are assumed to be longitudinally separated across their non‐breeding range and thus narrow population‐specific non‐breeding areas are expected. By using geolocators, we identified for the first time, the migration routes and non‐breeding areas of sand martins (n = 4) and house martins (n = 5) breeding in central‐eastern Europe. In autumn, the Carpathian Bend and northern parts of the Balkan Peninsula serve as important pre‐migration areas for both species. All individuals crossed the Mediterranean Sea from Greece to Libya. Sand martins spent the non‐breeding season in northern Cameroon and the Lake Chad Basin, within less than a 700 km radius, while house martins were widely scattered in three distinct regions in central, eastern, and southern Africa. Thus, for both species, the expected strength of migratory connectivity could not be confirmed. House martins, but not sand martins, migrated about twice as fast in spring compared to autumn. The spring migration started with a net average speed of > 400 km d^–1 for sand martins, and > 800 km d^–1 for house martins. However, both species used several stopover sites for 0.5–4 d and were stationary for nearly half of their spring migration. Arrival at breeding grounds was mainly related to departure from the last sub‐Saharan non‐breeding site rather than distance, route, or stopovers. We assume a strong carry‐over effect on timing in spring.     

Population differentiation of migratory directions in birds: comparison between ringing results and orientation behaviour of hand-raised migrants

Summary Blackcaps (Sylvia atricapilla) from five areas in Central Europe were hand-raised. Their autumn migratory orientation was tested in funnel-shaped cages. Their directional choices were compared to recoveries of conspecifics ringed during the breeding season in the same areas, which are situated on a transect across a migratory divide between southeastward and southwestward migrating populations. Results from ringing and orientation tests were in good agreement with respect to mean direction and dispersion of flight directions. An exception is the area around Linz (NW Austria), right on the migratory divide, where ringing yielded a strong scatter, but hand-raised birds chose westerly directions. The recent establishment of a novel migration route toward the British Isles was reflected in both data sets: in southern Germany the percentage of northwestward migrants is 6.8% according to orientation tests and 11.8% according to ringing recoveries. Testing the orientation of young passerines in captivity can yield valuable information about population differentiation of migratory behaviour. It is more efficient than ringing in this respect, because it circumvents the low recovery rates and is free of biases affecting ringing data. In the blackcap, geographic differentiation of migratory directions occurs on a finer scale than previously recognized and can change significantly within 2–3 decades.     

MIGRATION OF THE NORTHWESTERN EUROPEAN POPULATION OF THE CUCKOO CUCULUS CANORUS, AS SHOWN BY RINGING

The migration of the Cuckoo Cuculus canorus is described from recoveries of ringed birds. Young birds begin to migrate in August and most go southwards by September—those in continental Europe move southeast, south or southwest, while those from the British Isles move southeast. In spring, migrants arrive in Europe in April/May, second-(calendar) year birds doing so later than post-second-year birds. In autumn old birds depart in advance of first-year birds; and the individual bird may move stepwise through Europe. Old birds showed a marked tendency to return to the same places inhabited in previous breeding seasons, and birds ringed as nestlings subsequently tended to return to the general vicinity of the natal site.     

The large-scale detoured migration route and the shifting pattern of migration in Oriental honey-buzzards breeding in Japan

We describe the detoured migration route of the Oriental honey-buzzard Pernis ptilorhyncus , showing differences between autumn and spring migration, using data from 10 adult individuals marked with satellite transmitters. In autumn, the migration routes were very similar from Japan to the south end of the Malay Peninsula. The wintering sites were distributed within the Philippines, Borneo and the Malay Archipelago. During autumn, migration of the birds had few long-term stopover sites, instead, sometimes decidedly slowing their migration rate while proceeding in a consistent direction. During spring migration, the honey-buzzards penetrated into southern China, moving north to the base of the Korean Peninsula. The birds then went south through the Korean Peninsula to reach Japan. Before travelling to China, all spring migrants stopped for several weeks in south-east Asia. The slow rate of travel in the autumn suggests that migrants were foraging and replenishing their energy reserves. Instead of a migration strategy that uses only a few long-term stopover sites, honey-buzzards may adopt a strategy based on a number of short-term stay sites.