Animal movements in air and water can be strongly affected by experienced flow. While various flow-orientation strategies have been proposed and observed, their performance in variable flow conditions remains unclear. We apply control theory to establish a benchmark for time-minimizing (optimal) orientation. We then define optimal orientation for movement in steady flow patterns and, using dynamic wind data, for short-distance mass movements of thrushes (Turdus sp.) and 6000 km non-stop migratory flights by great snipes, Gallinago media. Relative to the optimal benchmark, we assess the efficiency (travel speed) and reliability (success rate) of three generic orientation strategies: full compensation for lateral drift, vector orientation (single-heading movement) and goal orientation (continually heading towards the goal). Optimal orientation is characterized by detours to regions of high flow support, especially when flow speeds approach and exceed the animal's self-propelled speed. In strong predictable flow (short distance thrush flights), vector orientation adjusted to flow on departure is nearly optimal, whereas for unpredictable flow (inter-continental snipe flights), only goal orientation was near-optimally reliable and efficient. Optimal orientation provides a benchmark for assessing efficiency of responses to complex flow conditions, thereby offering insight into adaptive flow-orientation across taxa in the light of flow strength, predictability and navigation capacity.
flow orientation; animal navigation; migration; lateral drift; optimization; movement ecology
Narrow migration corridors known in diurnal, social migrants such as raptors, storks and geese are thought to be caused by topographical leading line effects in combination with learning detailed routes across generations. Here, we document narrow-front migration in a nocturnal, solitary migrant, the common cuckoo Cuculus canorus, using satellite telemetry. We tracked the migration of adult cuckoos from the breeding grounds in southern Scandinavia (n = 8), to wintering sites in south-western Central Africa (n = 6) and back to the breeding grounds (n = 3). Migration patterns were very complex; in addition to the breeding and wintering sites, six different stopover sites were identified during the 16,000 km annual route that formed a large-scale clockwise loop. Despite this complexity, individuals showed surprisingly similar migration patterns, with very little variation between routes. We compared observed tracks with simulated routes based on vector orientation (with and without effects of barriers on orientation and survival). Observed distances between routes were often significantly smaller than expected if the routes were established on the basis of an innate vector orientation programme. Average distance between individuals in eastern Sahel after having migrated more than 5,000 km for example, was merely 164 km. This implies that more sophisticated inherent guiding mechanisms, possibly involving elements of intermediate goal area navigation or more elaborate external cues, are necessary to explain the complex narrow-front migration pattern observed for the cuckoos in this study.
Recent technological advancements now allow us to obtain geographical position data for a wide range of animal movements. Here we used light-level geolocators to study the annual migration cycle in great reed warblers (Acrocephalus arundinaceus), a passerine bird breeding in Eurasia and wintering in sub-Saharan Africa. We were specifically interested in seasonal strategies in routes and schedules of migration. We found that the great reed warblers (all males, no females were included) migrated from the Swedish breeding site in early August. After spending up to three weeks at scattered stopover sites in central to south-eastern Europe, they resumed migration and crossed the Mediterranean Sea and Sahara Desert without lengthy stopovers. They then spread out over a large overwintering area and each bird utilised two (or even three) main wintering sites that were spatially separated by a distinct mid-winter movement. Spring migration initiation date differed widely between individuals (1-27 April). Several males took a more westerly route over the Sahara in spring than in autumn, and in general there were fewer long-distance travels and more frequent shorter stopovers, including one in northern Africa, in spring. The shorter stopovers made spring migration on average faster than autumn migration. There was a strong correlation between the spring departure dates from wintering sites and the arrival dates at the breeding ground. All males had a high migration speed in spring despite large variation in departure dates, indicating a time-minimization strategy to achieve an early arrival at the breeding site; the latter being decisive for high reproductive success in great reed warblers. Our results have important implications for the understanding of long-distance migrants’ ability to predict conditions at distant breeding sites and adapt to rapid environmental change.
The small size of the billions of migrating songbirds commuting between temperate breeding sites and the tropics has long prevented the study of the largest part of their annual cycle outside the breeding grounds. Using light-level loggers (geolocators), we recorded the entire annual migratory cycle of the red-backed shrike Lanius collurio, a trans-equatorial Eurasian-African passerine migrant. We tested differences between autumn and spring migration for nine individuals. Duration of migration between breeding and winter sites was significantly longer in autumn (average 96 days) when compared with spring (63 days). This difference was explained by much longer staging periods during autumn (71 days) than spring (9 days). Between staging periods, the birds travelled faster during autumn (356 km d–1) than during spring (233 km d–1). All birds made a protracted stop (53 days) in Sahelian sub-Sahara on southbound migration. The birds performed a distinct loop migration (22 000 km) where spring distance, including a detour across the Arabian Peninsula, exceeded the autumn distance by 22 per cent. Geographical scatter between routes was particularly narrow in spring, with navigational convergence towards the crossing point from Africa to the Arabian Peninsula. Temporal variation between individuals was relatively constant, while different individuals tended to be consistently early or late at different departure/arrival occasions during the annual cycle. These results demonstrate the existence of fundamentally different spatio-temporal migration strategies used by the birds during autumn and spring migration, and that songbirds may rely on distinct staging areas for completion of their annual cycle, suggesting more sophisticated endogenous control mechanisms than merely clock-and-compass guidance among terrestrial solitary migrants. After a century with metal-ringing, year-round tracking of long-distance migratory songbirds promises further insights into bird migration.
avian migration; geolocator; passerine; migration speed; phenology; migration routes
Migratory land birds perform extreme endurance flights when crossing ecological barriers, such as deserts, oceans and ice-caps. When travelling over benign areas, birds are expected to migrate by shorter flight steps, since carrying the heavy fuel loads needed for long non-stop flights comes at considerable cost. Here, we show that great snipes Gallinago media made long and fast non-stop flights (4300–6800 km in 48–96 h), not only over deserts and seas but also over wide areas of suitable habitats, which represents a previously unknown migration strategy among land birds. Furthermore, the great snipes achieved very high ground speeds (15–27 m s−1), which was not an effect of strong tailwind support, and we know of no other animal that travels this rapidly over such a long distance. Our results demonstrate that some migratory birds are prepared to accept extreme costs of strenuous exercise and large fuel loads, even when stopover sites are available along the route and there is little tailwind assistance. A strategy of storing a lot of energy before departure, even if migration is over benign habitats, may be advantageous owing to differential conditions of fuel deposition, predation or infection risk along the migration route.
avian migration; endurance exercise; geolocators; stopover ecology; wind assistance
Vast numbers of insects and passerines achieve long-distance migrations between summer and winter locations by undertaking high-altitude nocturnal flights. Insects such as noctuid moths fly relatively slowly in relation to the surrounding air, with airspeeds approximately one-third of that of passerines. Thus, it has been widely assumed that windborne insect migrants will have comparatively little control over their migration speed and direction compared with migrant birds. We used radar to carry out the first comparative analyses of the flight behaviour and migratory strategies of insects and birds under nearly equivalent natural conditions. Contrary to expectations, noctuid moths attained almost identical ground speeds and travel directions compared with passerines, despite their very different flight powers and sensory capacities. Moths achieved fast travel speeds in seasonally appropriate migration directions by exploiting favourably directed winds and selecting flight altitudes that coincided with the fastest air streams. By contrast, passerines were less selective of wind conditions, relying on self-powered flight in their seasonally preferred direction, often with little or no tailwind assistance. Our results demonstrate that noctuid moths and passerines show contrasting risk-prone and risk-averse migratory strategies in relation to wind. Comparative studies of the flight behaviours of distantly related taxa are critically important for understanding the evolution of animal migration strategies.
seasonal migration; songbirds; Lepidoptera; Autographa gamma; flight speed; orientation
The exploration of animal migration has entered a new era with individual-based tracking during multiple years. Here, we investigated repeated migratory journeys of a long-distance migrating bird, the marsh harrier Circus aeruginosus, in order to analyse the variation within and between individuals with respect to routes and timing. We found that there was a stronger individual repeatability in time than in space. Thus, the annual timing of migration varied much less between repeated journeys of the same individual than between different individuals, while there was considerable variation in the routes of the same individual on repeated journeys. The overall contrast in repeatability between time and space was unexpected and may be owing to strong endogenous control of timing, while short-term variation in environmental conditions (weather and habitat) might promote route flexibility. The individual variation in migration routes indicates that the birds navigate mainly by other means than detailed route recapitulation based on landmark recognition.
avian migration; satellite-tracking; repeatability; routes; timing
Performance of migrating birds can be affected by a number of intrinsic and extrinsic factors like morphology, meteorological conditions and migration strategies. We compared travel speeds of four raptor species during their crossing of the Sahara desert. Focusing the analyses on this region allows us to compare different species under equivalent conditions in order to disentangle which factors affect migratory performance.
We tracked raptors using GPS satellite transmitters from Sweden, Spain and Italy, and evaluated their migratory performance at both an hourly and a daily scale. Hourly data (flight speed and altitude for intervals of two hours) were analyzed in relation to time of day, species and season, and daily data (distance between roosting sites) in relation to species, season, day length and tailwind support.
Despite a clear variation in morphology, interspecific differences were generally very small, and did only arise in spring, with long-distance migrants (>5000 km: osprey and Western marsh-harrier) being faster than species that migrate shorter distances (Egyptian vulture and short-toed eagle). Our results suggest that the most important factor explaining hourly variation in flight speed is time of day, while at a daily scale, tailwind support is the most important factor explaining variation in daily distance, raising new questions about the consequences of possible future changes in worldwide wind patterns.
Wind and ocean currents may potentially have important effects on travelling animals, as an animal which does not respond to lateral flow will be drifted from its intended direction of movement. By analysing daily movements of migrating ospreys Pandion haliaetus and marsh harriers Circus aeruginosus, as recorded by satellite telemetry, in relation to global wind data, we showed that these raptors allow on average 47 per cent drift. Furthermore, our analyses revealed significant geographical and temporal variation in the response to crosswinds. During some parts of the migration, the birds drifted and in other parts they compensated or even overcompensated. In some regions, the response of marsh harriers depended on the wind direction. They drifted when the wind came from one side and (over)compensated when the wind came from the opposite side, and this flexible response was different in different geographical regions. These results suggest that migrating raptors modulate their response to crosswinds at different places and times during their travels and show that individual birds use a much more varied repertoire of behavioural responses to wind than hitherto assumed. Our results may also explain why contrasting and variable results have been obtained in previous studies of the effect of wind on bird migration.
bird orientation; migration; satellite tracking; wind drift
We investigated the risk associated with crossing the Sahara Desert for migrating birds by evaluating more than 90 journeys across this desert by four species of raptors (osprey Pandion haliaetus, honey buzzard Pernis apivorus, marsh harrier Circus aeruginosus and Eurasian hobby Falco subbuteo) recorded by satellite telemetry. Forty per cent of the crossings included events of aberrant behaviours, such as abrupt course changes, slow travel speeds, interruptions, aborted crossings followed by retreats from the desert and failed crossings due to death, indicating difficulties for the migrants. The mortality during the Sahara crossing was 31 per cent per crossing attempt for juveniles (first autumn migration), compared with only 2 per cent for adults (autumn and spring combined). Mortality associated with the Sahara passage made up a substantial fraction (up to about half for juveniles) of the total annual mortality, demonstrating that this passage has a profound influence on survival and fitness of migrants. Aberrant behaviours resulted in late arrival at the breeding grounds and an increased probability of breeding failure (carry-over effects). This study also demonstrates that satellite tracking can be a powerful method to reveal when and where birds are exposed to enhanced risk and mortality during their annual cycles.
avian migration; barrier crossings; carry-over effects; mortality
Autumn migration of adult Eurasian hobbies Falco subbuteo from Europe to southern Africa was recorded by satellite telemetry and observed routes were compared with randomly simulated routes. Two non-random features of observed routes were revealed: (i) shifts to more westerly longitudes than straight paths to destinations and (ii) strong route convergence towards a restricted area close to the equator (1° S, 15° E). The birds migrated south or southwest to approximately 10° N, where they changed to south-easterly courses. The maximal spread between routes at 10° N (2134 km) rapidly decreased to a minimum (67 km) close to the equator. We found a striking relationship between the route convergence and the distribution of continuous rainforest, suggesting that hobbies minimize flight distance across the forest, concentrating in a corridor where habitat may be more suitable for travelling and foraging. With rainforest forming a possible ecological barrier, many migrants may cross the equator either at 15° E, similar to the hobbies, or at 30–40° E, east of the rainforest where large-scale migration is well documented. Much remains to be understood about the role of the rainforest for the evolution and future of the trans-equatorial Palaearctic-African bird migration systems.
bird migration; Eurasian hobby Falco subbuteo; equatorial rainforest; route convergence; ecological barrier