How and why do bees buzz? Implications for buzz pollination.

Buzz pollination encompasses the evolutionary convergence of specialized floral morphologies and pollinator behaviour in which bees use vibrations (floral buzzes) to remove pollen. Floral buzzes are one of several types of vibrations produced by bees using their thoracic muscles. Here I review how bees can produce these different types of vibrations and discuss the implications of this mechanistic understanding for buzz pollination. I propose that bee buzzes can be categorized according to their mode of production and deployment into: (i) thermogenic, which generate heat with little mechanical vibration; (ii) flight buzzes which, combined with wing deployment and thoracic vibration, power flight; and (iii) non-flight buzzes in which the thorax vibrates but the wings remain mostly folded, and include floral, defence, mating, communication, and nest-building buzzes. I hypothesize that the characteristics of non-flight buzzes, including floral buzzes, can be modulated by bees via modification of the biomechanical properties of the thorax through activity of auxiliary muscles, changing the rate of activation of the indirect flight muscles, and modifying flower handling behaviours. Thus, bees should be able to fine-tune mechanical properties of their floral vibrations, including frequency and amplitude, depending on flower characteristics and pollen availability to optimize energy use and pollen collection.