Evolution of leapfrog migration: A test of competition-based hypotheses.

Leapfrog migration is a common migration pattern in birds where the breeding and wintering latitudes between populations are in reversed latitudinal sequence. Competition for wintering and breeding sites has been suggested to be an ultimate factor, and several competitor-based hypotheses have been proposed to explain this pattern. If wintering sites close to the breeding sites are favored, competitive exclusion could force subdominant individuals to winter further away. Competitive exclusion could be mediated either through body size or by prior occupancy. The alternative "spring predictability" hypothesis assumes competition for sufficiently close wintering areas, allowing the birds to use autocorrelated weather cues to optimally time spring migration departure. To test predictions and assumptions of these hypotheses, we combined morphometrics, migration, and weather data from four populations of common ringed plover breeding along a latitudinal (56-68° N) and climatic gradient (temperate to Arctic). Critical for our evaluation was that two populations were breeding on the same latitude in subarctic Sweden with the same distance to the closest potential wintering site, but differed in breeding phenology, and wintered in West Africa and Europe, respectively. Thus, while breeding on the same latitude, their winter distribution overlapped with that of an Arctic and temperate population. Body size was largest within the temperate population, but there was no size difference between the two subarctic populations. Populations wintering in Europe arrived there before populations wintering in Africa. The largest variation in the arrival of meteorological spring occurred at the temperate breeding site, while there was almost no difference among the other sites. In general, temperatures at the northernmost wintering area correlated well with each breeding site prior to breeding site-specific spring arrival. Based on these observations, we conclude that competitive exclusion through body-size-related dominance cannot explain leapfrog migration. Furthermore, the assumptions on which the "spring predictability" hypothesis is based did not match the observed wintering ranges either. However, we could not reject the hypothesis that competitive exclusion mediated by prior occupancy in the wintering area could lead to leapfrog migration, and therefore, this hypothesis should be retained as working hypothesis for further work.

Seasonal patterns and processes of migration in a long-distance migratory bird: energy or time minimization?

Optimal migration theory prescribes adaptive strategies of energy, time or mortality minimization. To test alternative hypotheses of energy- and time-minimization migration we used multisensory data loggers that record time-resolved flight activity and light for positioning by geolocation in a long-distance migratory shorebird, the little ringed plover, Charadrius dubius. We could reject the hypothesis of energy minimization based on a relationship between stopover duration and subsequent flight time as predicted for a time minimizer. We found seasonally diverging slopes between stopover and flight durations in relation to the progress (time) of migration, which follows a time-minimizing policy if resource gradients along the migration route increase in autumn and decrease in spring. Total flight duration did not differ significantly between autumn and spring migration, although spring migration was 6% shorter. Overall duration of autumn migration was longer than that in spring, mainly owing to a mid-migration stop in most birds, when they likely initiated moult. Overall migration speed was significantly different between autumn and spring. Migratory flights often occurred as runs of two to seven nocturnal flights on adjacent days, which may be countering a time-minimization strategy. Other factors may influence a preference for nocturnal migration, such as avoiding flight in turbulent conditions, heat stress and diurnal predators.

Consequences of migratory distance, habitat distribution and season on the migratory process in a short distance migratory shorebird population.

BACKGROUND: The migratory process in birds consists of alternating periods of flight and fueling. Individuals of some populations make few flights and long stopovers, while others make multiple flights between short stopovers. Shorebirds are known for executing marathon flights (jumps), but most populations studied are long distance migrants, often crossing major barriers and thus forced to make long-haul flights. The sub-division of migration in short/medium distance migratory populations, where the total migration distance is shorter than documented non-stop flight capacity and where routes offer more homogenous stopover landscape, is little explored. METHODS: Here we combine data based on conventional light level geolocators and miniaturized multi sensor loggers, comprising acceleration and light sensors, to characterize the migratory routes and migration process for a short/medium distance (~ 1300 to 3000 km) migratory population of common ringed plover (Charadrius hiaticula) breeding in southern Sweden. We were specifically interested in the variation in number and duration (total and individual) of flights/stopovers between seasons and in relation to migration distance. RESULTS: Most stopovers were located along the European Atlantic coast. On average 4.5 flights were made during autumn migration irrespective of migration distance, but in spring the number of flights increased with distance. The equal number of flights in autumn was explained by that most individuals migrating farther performed one longer flight (all but one lasting > 20 h), likely including crossing of the Bay of Biscay. Median duration of single flights was 8.7 h in autumn and 5.5 h in spring, and median stopover duration was ~ 1 day in both seasons. There was a positive relationship between total flight duration and migration distance, but total flight duration was 36% lower in spring compared to autumn. CONCLUSIONS: Our results suggest that when suitable stopovers are abundant common ringed plovers prefer making shorter flights even if longer flights are within the capacity of the species. This behaviour is predicted under both time and energy minimizing strategies, although the variable flight distances suggest a policy of time selected migration. Even if populations using several stopovers seem to be more resilient for environmental change along the route, these results are informative for conservation efforts and for predicting responses to future environmental change.

Measuring power input, power output and energy conversion efficiency in un-instrumented flying birds.

Cost of flight at various speeds is a crucial determinant of flight behaviour in birds. Aerodynamic models, predicting that mechanical power (Pmech) varies with flight speed in a U-shaped manner, have been used together with an energy conversion factor (efficiency) to estimate metabolic power (Pmet). Despite few empirical studies, efficiency has been assumed constant across flight speeds at 23%. Ideally, efficiency should be estimated from measurements of both Pmech and Pmet in un-instrumented flight. Until recently, progress has been hampered by methodological constraints. The main aim of this study was to evaluate recently developed techniques and estimate flight efficiency across flight speeds. We used the 13C-labelled sodium bicarbonate method (NaBi) and particle image velocimetry (PIV) to measure Pmet and Pmech in blackcaps flying in a wind tunnel. We also cross-validated measurements made by NaBi with quantitative magnetic resonance (QMR) body composition analysis in yellow-rumped warblers. We found that Pmet estimated by NaBi was ∼12% lower than corresponding values estimated by QMR. Pmet varied in a U-shaped manner across flight speeds in blackcaps, but the pattern was not statistically significant. Pmech could only be reliably measured for two intermediate speeds and estimated efficiency ranged between 14% and 22% (combining the two speeds for raw and weight/lift-specific power, with and without correction for the ∼12% difference between NaBi and QMR), which were close to the currently used default value. We conclude that NaBi and PIV are viable techniques, allowing researchers to address some of the outstanding questions regarding bird flight energetics.

The migration pattern of a monogamous shorebird challenges existing hypotheses explaining the evolution of differential migration.

Differential migration by sex, where one sex migrates further than the other, occurs in many bird species. How this pattern evolves is however little understood. The first aim of this study was to investigate the extent of differential migration in the common ringed plover Charadrius hiaticula, breeding in southeast Sweden, and test three main hypotheses (the social dominance, body size and arrival time hypothesis) regarding the evolution of differential migration. Geolocators were used to derive spatiotemporal data and morphometrics were collected from the studied population. Males migrated 800 km farther compared to females, were marginally larger and there was no statistical difference in spring arrival between the sexes. In contrast to other studies none of the previously proposed hypotheses could account for the observed pattern. An additional finding was that both sexes arrive up to 1.5 months before egg laying, but males initiate territorial behavior upon arrival. Based on these observations we suggest that males have a higher energetic demand, and challenges to meet those, early in the breeding season. Therefore we hypothesize that males arrive to the breeding site with residual fuel reserves accumulated at the wintering site to cover at least parts of these demands. Based on this hypothesis we present a simple model to explain the longer migration by males. The model is contingent on a trade-off between site specific fueling rates (which we assume to increase with decreasing latitude), cost of the extra migration distance and predation risk during fueling. This framework may be applicable to other cases of differential migration, especially in temperate breeding species which exhibit long pre-egg laying periods.