Each cuckoo's journey was unique. Their departure dates were weeks apart as were arrival dates. Two birds chose westerly flight paths, through Spain and along the coast of North-West Africa, before continuing in a south-easterly direction on the final legs. The other three chose more direct routes, through Italy, across the Mediterranean into North Africa, then across the Sahara. Each journey was punctuated by stopovers of several weeks, at the end of flights which could well have been gruelling, covering up to 2200 Km over a two day period.
It is amazing that all five birds should reach their destination without mishap, and that five birds with breeding grounds around the same area should find themselves close together after having been separated by up to 3600 Km. I used to think that bird behaviour was governed entirely by instinct and that instinct was a property of the species followed blindly by each individual. Free will was not something I associated with birds.
It is hard to imagine such a journey being made without knowledge, at least knowledge, conscious or unconscious, of the destination. I suspect each bird also has knowledge of the hazardous regions to be crossed. It avoids very long flights over the sea, and rests and refuels before attempting a desert crossing. This knowledge cannot have been acquired through learning because the cuckoo's surrogate parents know nothing of migration to another continent. It is possible that this knowledge is resident in the bird brain having been put there through reactions initiated by cuckoo DNA. It is also possible that that the knowledge is resident elsewhere, and cuckoo DNA initiates reactions that result in the cuckoo being able to access it at the appropriate time.
Fascinating!