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Bee declines have been partly attributed to the impacts of invasive or emerging parasite outbreaks. For western honeybees,Apis mellifera, major losses are associated with the virus-vectoring mite,Varroa destructor. In response, beekeepers have focused breeding efforts aimed at conferring resistance to this key parasite. One method of many is survival-based beekeeping where colonies that survive despite significantVarroainfestations produce subsequent colonies. We argue that this ‘hands-off’ approach will not always lead toVarroaresistance evolving but rather tolerance. Tolerance minimizes host fitness costs of parasitism without reducing parasite abundance, whereas resistance either prevents parasitism outright or keeps parasitism intensity low. With clear epidemiological distinctions, and as honeybee disease dynamics impact other wild bees owing to shared pathogens, we discuss why tolerance outcomes in honeybee breeding have important implications for wider pollinator health. Crucially, we argue that unintentional selection for tolerance will not only lead to more spillover from honeybees but may also select for pathogens that are more virulent in wild bees leading to ‘tragedies of tolerance’. These tragedies can be avoided through successful breeding regimes that specifically select for lowVarroa. We emphasize how insights from evolutionary ecology can be applied in ecologically responsible honeybee management. 

There is increasing interest in the role that evolution may play in current and future pandemics, but there is often also considerable confusion about the actual evolutionary predictions. This may be, in part, due to a historical separation of evolutionary and medical fields, but there is a large, somewhat nuanced body of evidence-supported theory on the evolution of infectious disease. In this review, we synthesize this evolutionary theory in order to provide a framework for clearer understanding of the key principles. Specifically, we discuss the selection acting on zoonotic pathogens' transmission rates and virulence at spillover and during emergence. We explain how the direction and strength of selection during epidemics of emerging zoonotic disease can be understood by a three Ts framework: trade-offs, transmission, and time scales. Virulence and transmission rate may trade-off, but transmission rate is likely to be favoured by selection early in emergence, particularly if maladapted zoonotic pathogens have ‘no-cost’ transmission rate improving mutations available to them. Additionally, the optimal virulence and transmission rates can shift with the time scale of the epidemic. Predicting pathogen evolution, therefore, depends on understanding both the trade-offs of transmission-improving mutations and the time scales of selection.