Do linden trees kill bees? Reviewing the causes of bee deaths on silver linden (Tilia tomentosa)

For decades, linden trees (basswoods or lime trees), and particularly silver linden (Tilia tomentosa), have been linked to mass bee deaths. This phenomenon is often attributed to the purported occurrence of the carbohydrate mannose, which is toxic to bees, in Tilia nectar. In this review, however, we conclude that from existing literature there is no experimental evidence for toxicity to bees in linden nectar. Bee deaths on Tilia probably result from starvation, owing to insufficient nectar resources late in the tree's flowering period. We recommend ensuring sufficient alternative food sources in cities during late summer to reduce bee deaths on silver linden. Silver linden metabolites such as floral volatiles, pollen chemistry and nectar secondary compounds remain underexplored, particularly their toxic or behavioural effects on bees. Some evidence for the presence of caffeine in linden nectar may mean that linden trees can chemically deceive foraging bees to make sub-optimal foraging decisions, in some cases leading to their starvation.