BackgroundForaging workers of grass-cutting ants (Atta vollenweideri) regularly carry grass fragments larger than their own body. Fragment length has been shown to influence the ants’ running speed and thereby the colony’s food intake rate. We investigated whether and how grass-cutting ants maintain stability when carrying fragments of two different lengths but identical mass.Principal FindingsAnts carried all fragments in an upright, backwards-tilted position, but held long fragments more vertically than short ones. All carrying ants used an alternating tripod gait, where mechanical stability was increased by overlapping stance phases of consecutive steps. The overlap was greatest for ants carrying long fragments, resulting in more legs contacting the ground simultaneously. For all ants, the projection of the total centre of mass (ant and fragment) was often outside the supporting tripod, i.e. the three feet that would be in stance for a non-overlapping tripod gait. Stability was only achieved through additional legs in ground contact. Tripod stability (quantified as the minimum distance of the centre of mass to the edge of the supporting tripod) was significantly smaller for ants with long fragments. Here, tripod stability was lowest at the beginning of each step, when the center of mass was near the posterior margin of the supporting tripod. By contrast, tripod stability was lowest at the end of each step for ants carrying short fragments. Consistently, ants with long fragments mainly fell backwards, whereas ants carrying short fragments mainly fell forwards or to the side. Assuming that transporting ants adjust neither the fragment angle nor the gait, they would be less stable and more likely to fall over.ConclusionsIn grass-cutting ants, the need to maintain static stability when carrying long grass fragments has led to multiple kinematic adjustments at the expense of a reduced material transport rate.

Grass-cutting ants (Atta vollenweideri) carry leaf fragments several times heavier and longer than the workers themselves over considerable distances back to their nest. Workers transport fragments in an upright, slightly backwards-tilted position. To investigate how they maintain stability and control the carried fragment's position, we measured head and fragment positions from video recordings. Load-transporting ants often fell over, demonstrating the biomechanical difficulty of this behavior. Long fragments were carried at a significantly steeper angle than short fragments of the same mass. Workers did not hold fragments differently between the mandibles, but performed controlled up and down head movements at the neck joint. By attaching additional mass at the fragment's tip to load-carrying ants, we demonstrated that they are able to adjust the fragment angle. When we forced ants to transport loads across inclines, workers walking uphill carried fragments at a significantly steeper angle, and downhill at a shallower angle than ants walking horizontally. However, we observed similar head movements in unladen workers, indicating a generalized reaction to slopes that may have other functions in addition to maintaining stability. Our results underline the importance of proximate, biomechanical factors for the understanding of the foraging process in leaf-cutting ants.

SUMMARYGrass-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 antsʼ 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.

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