Why are There 'Lazy' Ants? How Worker Inactivity can Arise in Social Insect Colonies

Abstract

"All cold-blooded animals and a large number of warm-blooded ones spend an unexpectedly large proportion of their time doing nothing at all, or at any rate, nothing in particular." (Elton 1927) Many animals are remarkably "lazy", spending >50% of their waking hours "resting" . This is common across all taxa, ecologies, and life histories, including what are commonly considered to be highly industrious animals: the social insects (e.g., Aesop's Fable 'The Grasshopper and the Ant'). This dissertation broadly seeks to explain a phenomenon that has long been observed, but never adequately addressed, by asking: 'why are there 'lazy' ants?' First, I established that inactivity was a real and ecologically relevant phenomenon in the ant Temnothorax rugatulus by testing whether inactivity was a lab artifact. I then showed that inactive workers comprise a behaviorally distinct group of workers that are commonly overlooked in studies looking at colony function, though they typically represent at least half of the individuals within social insect colonies. I then tested a set of mutually non-exclusive hypotheses explaining inactivity in social insects: that (1) inactivity is a form of social "cheating" in which egg-laying workers selfishly invest in their own reproduction rather than contribute to colony fitness, (2) inactive workers comprise a pool of reserve workers used to mitigate the effects of fluctuations in colony workload, (3) inactivity is the result of physiological constraints on worker age such that young and old workers may less active due to inexperience/physical vulnerability, and physiological deterioration respectively, (4) inactive workers are performing an as-yet unidentified function, such as playing a role in communication and acting as food stores, or repletes, and that (5) inactive workers represent the 'slow' end of intra-nest variation in worker 'pace-of-life'. Inactivity is linked to worker age, reproduction, and a potential function as food stores for the colony. These hypotheses are not mutually exclusive, and in fact, likely form a 'syndrome' of behaviors common to inactive social insect workers. Their simultaneous contribution to worker inactivity may explain the difficulty in finding a simple answer to this deceptively simple question.