High altitude bird migration at temperate latitudes: a synoptic perspective on wind assistance.

At temperate latitudes the synoptic patterns of bird migration are strongly structured by the presence of cyclones and anticyclones, both in the horizontal and altitudinal dimensions. In certain synoptic conditions, birds may efficiently cross regions with opposing surface wind by choosing a higher flight altitude with more favourable wind. We observed migratory passerines at mid-latitudes that selected high altitude wind optima on particular nights, leading to the formation of structured migration layers at varying altitude up to 3 km. Using long-term vertical profiling of bird migration by C-band Doppler radar in the Netherlands, we find that such migration layers occur nearly exclusively during spring migration in the presence of a high-pressure system. A conceptual analytic framework providing insight into the synoptic patterns of wind assistance for migrants that includes the altitudinal dimension has so far been lacking. We present a simple model for a baroclinic atmosphere that relates vertical profiles of wind assistance to the pressure and temperature patterns occurring at temperate latitudes. We show how the magnitude and direction of the large scale horizontal temperature gradient affects the relative gain in wind assistance that migrants obtain through ascending. Temperature gradients typical for northerly high-pressure systems in spring are shown to cause high altitude wind optima in the easterly sectors of anticyclones, thereby explaining the frequent observations of high altitude migration in these synoptic conditions. Given the recurring synoptic arrangements of pressure systems across temperate continents, the opportunities for exploiting high altitude wind will differ between flyways, for example between easterly and westerly oceanic coasts.

Quantifying flow-assistance and implications for movement research.

The impact that flows of air and water have on organisms moving through these environments has received a great deal of attention in theoretical and empirical studies. There are many behavioral strategies that animals can adopt to interact with these flows, and by assuming one of these strategies a researcher can quantify the instantaneous assistance an animal derives from a particular flow. Calculating flow-assistance in this way can provide an elegant simplification of a multivariate problem to a univariate one and has many potential uses; however, the resultant flow-assistance values are inseparably linked to the specific behavioral strategy assumed. We expect that flow-assistance may differ considerably depending on the behavioral strategy assumed and the accuracy of the assumptions associated with that strategy. Further, we expect that the magnitude of these differences may depend on the specific flow conditions. We describe equations to quantify flow-assistance of increasing complexity (i.e. more assumptions), focusing on the behavioral strategies assumed by each. We illustrate differences in suggested flow-assistance between these equations and calculate the sensitivity of each equation to uncertainty in its particular assumptions for a range of theoretical flow conditions. We then simulate trajectories that occur if an animal behaves according to the assumptions inherent in these equations. We find large differences in flow-assistance between the equations, particularly with increasing lateral flow and increasingly supportive axial flow. We find that the behavioral strategy assumed is generally more influential on the perception of flow-assistance than a small amount of uncertainty in the specification of an animal's speed (i.e. <5 ms(-1)) or preferred direction of movement (i.e. <10°). Using simulated trajectories, we show that differences between flow-assistance equations can accumulate over time and distance. The appropriateness and potential biases of an equation to quantify flow-assistance, and the behavioral assumptions the equation implies, must be considered in the context of the system being studied, particularly when interpreting results. Thus, we offer this framework for researchers to evaluate the suitability of a particular flow-assistance equation and assess the implications of its use.