For at least the last 2 years there have been flightless bees crawling around outside Hive A. These bees were clearly suffering from Deformed Wing Virus or another disabling virus. This year there are none.
The cause was explained to me by a researcher at University of Salford. All winter the Varroa mites feed from worker bees. They accumulate DWV virus particles. In spring, when brood production increases they rush in to infest the new brood and transfer more virus to them. These bees show visible signs of DWV -- deformed wings or an inability to fly.