The energetics of running stability: costs of transport in grass-cutting ants depend on fragment shape.

Grass-cutting ants (Atta vollenweideri) carry fragments that can be many times heavier and longer than the ants themselves and it is important for them to avoid falling over during load transport. To investigate whether the energetic costs of transport are affected by the need to maintain stability, the rate of CO(2) production was measured in both unladen workers and workers carrying standardized paper fragments of different size and shape. We tested: (1) the effect of mass by comparing workers carrying either light or heavy fragments of the same size, and (2) the effect of shape by comparing short and long fragments of the same mass. Consistent with previous studies, metabolic rate increased but running speed remained constant when ants carried heavier fragments. The net cost of transport (normalized to the total mass of ant and fragment) was the same for heavy and light fragments, and did not differ from the costs of carrying a unit body mass. Ants carrying long fragments showed similar metabolic rates but ran significantly slower than ants carrying short fragments. As a consequence, net cost of transport was significantly higher for long fragments than for short ones, and higher than the costs of carrying a unit body mass. The observed reduction in running speed is likely a result of the ants' need to maintain stability. When the absolute costs of transport were compared, smaller ants required more energy to carry heavier and longer fragments than larger workers, but the opposite was found for lighter and shorter fragments. The absolute costs of transport per unit fragment mass suggest that it is energetically advantageous for a colony to allocate smaller workers for the transport of small fragments and larger workers for large fragments. The present results underline the importance of biomechanical factors for the understanding of leaf-cutting ant foraging strategies.