migratory beekeeping, queen imports, drone reduction, and various other alterations of the bee's natural biology. These apiculture practices tend to limit natural selection and to disrupt the hard‐won adaptations of A. mellifera; they impact both the genes and the lifestyle of the honey bee (Neumann and Blacquière 2016). Now, what can be done from an animal husbandry and animal health perspective to reverse such trends?
The bee doctor must be prepared to examine honey bee health through a new lens that takes a holistic approach to medicine – one that features an understanding of and appreciation for the health of honey bees living in nature. In some parts of the world, beekeepers are already looking at beekeeping less as a process of domestication that forces the production of honey, wax, propolis, and pollination at great cost to colonies and more as the stewardship of a natural living system. The global decline in bee health is a direct consequence of man's disruption of this system: the introduction of exotic parasites and pathogens, the rise in disease virulence driven by beekeeping practices, and the evolution of drug resistance caused by indiscriminate treatments of colonies. Indeed, it is the pharmaceutical‐centric approach to preventative care for honey bees that is the fundamental reason behind the inclusion of honey bees among the food‐producing animals in North America that now fall under FDA regulations requiring the services of a veterinarian for antibiotic use. A key feature of a healthy system is achieving a balance between the host and the pathogen that promotes host resistance and pathogen avirulence – we can find this balance by promoting good genes and a good lifestyle in the bees.
Figure 1.7 Polyandry, or the multiple matings of a queen with drones from different patrilines, has been associated with colony vigor and improved winter survival. The health benefits of polyandry are linked to improved foraging rates, greater brood production, lower mite infestations, and the possession of rare alleles important for control of infectious disease.
Promoting Good Genes
The idea that honey bees have been domesticated by mankind remains a matter of debate. What is clear is that across North America there are populations of wild colonies of A. mellifera that thrive independent of beekeeping activities and that do not require the regular input of new colonies from honey bee swarms arising from managed colonies (Oliver 2014; Seeley 2017b; Radcliffe and Seeley 2018). Furthermore, the wild colonies tend to be genetically distinct from those that queen breeders produce for commercial purposes; the former are both more diverse genetically and they show strong evidence of regional adaptation (Figure 1.7) (reviewed in Seeley 2019a,b). Evidently, the honey bee colonies managed by beekeepers are semi‐domesticated, since their genes are influenced somewhat by queen breeders and their lifestyle is strongly influenced by their owners (Chapman et al. 2008; Oliver 2014).
An important lesson can be learned from the many animals that man has domesticated over the past thousand years: domestication carries with it a reliance on humans and generally a loss of the ability to survive in the wild. Here we can take some guidance from Charles Darwin:
One of the most remarkable features in our domesticated races is that we see in them adaptation, not indeed to the animal's or plant's own good, but to man's use or fancy. Some variations have probably arisen suddenly, or by one step. However, we cannot suppose that all the breeds were suddenly produced as perfect and useful as we now see them… . The key is man's power of accumulative selection: Nature gives successive variations: man adds then up in certain directions useful to him.
(Darwin 1868)
Among the honey bee traits that are known to have a genetic basis, resistance to disease has shown to be a strong component of colony fitness (Tarpy and Seeley 2006). With this in mind, we believe that both the beekeeper and the bee doctor will be wise to consider the following items when it comes to managing the genetics of honey bees.
Goal 1: Select Locally Adapted, Survivor Stock
Bait hives are a ready method for beekeepers to incorporate wild honey bees and their genes into their apiaries (Figure 1.8). A queen honey bee of local origin is well suited to an ecoregion or ecotype and has genes that provide a good fit with the local floral diversity, regional environmental conditions (including extremes of temperature, humidity, drought, etc.), and agents of disease.
A wonderful example of the adaptation of honey bees to their locale is the ecotype of A. mellifera that lives in the Landes heathlands of southwestern France (Louveaux 1973). The Landes bees have evolved to have a brood cycle with an unusual, second peak of brood production in August, just in time for the bloom of ling heather (Calluna vulgaris) in the Landes landscape. It is interesting to note that when Louveaux moved Paris honey bees to Landes, the Paris bees kept ahead (in colony weight gain, pollen collection, and brood production) of the Landes bees until the middle of July. Up to that point, the Landes bees had trailed behind the Paris bees because the Paris bees had reared more brood in May and June. In August, however, all the colonies of the Landes bees had a second burst of young bees emerging shortly before the heather bloom and by the end of summer these colonies had collected an astonishing 14 kg more honey than the colonies of Paris bees (Louveaux 1973).
Figure 1.8 Bait hives are small nest boxes that are filled with empty comb and sometimes lures or attractants (lemon grass oil or Nasanov pheromone). During the swarm season (May to June in the northeastern United States), scout bees search out and select bait hives during house hunting behaviors. Wild honey bees are well adapted to Varroa and often fare much better than managed bees. Therefore, bait hives are a simple means for the acquisition of locally adapted honey bee stock when they are used in places where there are few beekeepers.
Goal 2: Promote Drone Comb Building and Drone Mating in Congregation Areas
Modern apiarists work to limit the amount of drone comb produced by honey bees because beekeepers have learned that by preventing their colonies from producing drones, they can increase honey production (Seeley 2002). Furthermore, drone comb is the preferred site for Varroa reproduction. Limiting it, however, partially “castrates” a colony by reducing the ability of a colony to contribute to the population of drones in a region, an important driver of honey bee diversity and fitness (Seeley 2017b). It is now known that a colony's health and productivity is enhanced by its having high genetic diversity among its worker bees, which arises from the multiple mating (polyandry) strategy of queen honey bees (Tarpy and Seeley 2006; Seeley and Tarpy 2007; Mattila and Seeley 2007). On average, a queen honey bee mates with, and acquires sperm from 10 to 20 drones. Inhibiting drone production in colonies hinders the maintenance of genetic diversity within a region, including the genes that may hold resistance to mites (Rosenkranz et al. 2010).
Goal 3: Cull Failing Colonies Before Collapse
Some veterinarians with experience in honey bee disease and/or epidemiology have campaigned against the emergence of Treatment‐Free Beekeeping or Natural Beekeeping because of the risk of spreading disease through the collapse of colonies. Perhaps most alarming is the phenomenon
