Geographic variation in onset of singing among populations of two migratory birds

Even though singing plays a major role in bird communication, environmental variables affecting the geographic patterns observed in the variation of singing onset within large areas have not previously been studied. The singing phenology of two long-distance migrants, the Cuckoo Cuculus canorus and the Nightingale Luscinia megarhynchos, recorded in thousands of sites throughout Spain, was related to a set of 51 explanatory variables by partial least squares regression. Observed variability in singing onset among localities follows spatial trends on an Iberian scale at 10 × 10 km resolution, and was well modelled by some environmental variables. Of these variables, climate was the most influential. Males of both species sang earlier in warmer and drier sites. Males who were near the Straits of Gibraltar and in the southern Iberian basins, which are geographic gradients closely related with climate in the Iberian Peninsula, were also heard earlier. Most environmental variables affected the singing activity of both species equally, i.e., the same environmental gradients are linked to singing onset. However, the models' predictions indicated differences in the geographical pattern in singing onset between both species, mainly related to their rates of progression during spring migration through Spain. Local variables, such as land uses, vegetation productivity or species abundance, played a minor role on singing onset spatial patterns. Therefore, models' composition supports the hypothesis that singing onset is a good proxy of arrival dates in migratory species, since those variables related to indirect effects on singing onset phenology through migratory progression and arrival to the breeding grounds showed the highest influence.