Overwintering honey bees: biology and management

In temperate climates, honey bees (Apis mellifera) survive the winter by entering a distinct physiological and behavioral state. In recent years, beekeepers are reporting unsustainably high colony losses during the winter, which have been linked to parasitization by Varroa mites, virus infections, geographic location, and variation across honey bee genotypes. Here, we review literature on environmental, physiological, and social factors regulating entrance, maintenance, and exit from the overwintering state in honey bees in temperate regions and develop a testable model to explain how multiple factors may be acting synergistically to regulate this complex transition. We also review existing knowledge of the factors affecting overwintering survival in honey bees and providing suggestions to beekeepers aiming to improve their colonies' overwintering success.