Keywords:GWI index GDD jerk index Barnacle geese Stopover site Breeding site Mixed effect linear regression a b s t r a c t According to the green wave hypothesis, herbivores follow the flush of spring growth of forage plants during their spring migration to northern breeding grounds. In this study we compared two green wave indices for predicting the timing of the spring migration of avian herbivores: the satellite-derived green wave index (GWI), and an index of the rate of acceleration in temperature (GDDjerk). The GWI was calculated from MODIS normalized difference vegetation index (NDVI) satellite imagery and GDDjerk from gridded temperature data using products from the global land data assimilation system (GLDAS). To predict the timing of arrival at stopover and breeding sites, we used four years (2008-2011) of tracking data from 12 GPS-tagged barnacle geese, a long-distance herbivorous migrant, wintering in the Netherlands, breeding in the Russian Arctic. The stopover and breeding sites for these birds were identified and the relations between date of arrival with the date of 50% GWI and date of peak GDDjerk at each site were analyzed using mixed effect linear regression. A cross-validation method was used to compare the predictive accuracy of the GWI and GDDjerk indices. Significant relationships were found between the arrival dates at the stopover and breeding sites for the dates of 50% GWI as well as the peak GDDjerk (p < 0.01). The goose arrival dates at both stopover and breeding sites were predicted more accurately using GWI (R 2 cv = 0.68, RMSD cv = 5.9 and R 2 cv = 0.71, RMSD cv = 3.9 for stopover and breeding sites, respectively) than GDDjerk. The GDDjerk returned a lower accuracy for prediction of goose arrival dates at stopover ( R 2 cv = 0.45, RMSD cv = 7.79) and breeding sites (R 2 cv = 0.55, RMSD cv = 4.93). The positive correlation between the absolute residual values of the GDDjerk model and distance to the breeding sites showed that this index is highly sensitive to latitude. This study demonstrates that the satellite-derived green wave index (GWI) can accurately predict the timing of goose migration, irrespective of latitude and therefore is suggested as a reliable green wave index for predicting the timing of avian herbivores spring migration.
Migratory birds need considerable energy reserves to fuel long-distance flights to their breeding grounds in spring. To attain sufficient energy deposits before departure, birds require high daily intake rates, which can be reached by utilizing high-quality food. During such periods of high energy demand, animals often track changes in the nutritious value of their food, for example by switching to a more profitable habitat or diet. Pre-migratory Barnacle Geese Branta leucopsis staging along the Wadden Sea coast are known to switch from pastures to salt marshes during spring. Previous studies have suggested that Barnacle Geese switch habitat to track changes in protein levels, which drop in pastures as spring progresses, and to avoid high levels of disturbance. Here we made use of detailed information on pre-migratory habitat use of individual Barnacle Geese tracked by GPS/accelerometer tags to assess which factors may drive a habitat switch. We analysed habitat use and time budgets of individual birds and combined this with data on food quality in two habitats to analyse differences in food intake. We found large individual variation in pre-migratory habitat use, both in the extent of salt marsh use, as well as in the timing of a switch from pastures to salt marshes. In salt marshes, geese spent more time grazing and made fewer flight movements, potentially as they experienced lower levels of disturbance compared to geese in pastures. By increasing grazing time and reducing flight movements, geese in salt marshes may compensate for reduced food quality. Our results show that Barnacle Geese trade-off high intake rates and high costs in pastures with low intake rates and low costs in salt marshes.
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