Desert ants, Cataglyphis, navigate in their vast desert habitat by path integration. They continuously integrate directions steered (as determined by their celestial compass) and distances traveled, gauged by as-yet-unknown mechanisms. Here we test the hypothesis that navigating ants measure distances traveled by using some kind of step integrator, or "step counter." We manipulated the lengths of the legs and, hence, the stride lengths, in freely walking ants. Animals with elongated ("stilts") or shortened legs ("stumps") take larger or shorter strides, respectively, and concomitantly misgauge travel distance. Travel distance is overestimated by experimental animals walking on stilts and underestimated by animals walking on stumps.

High-speed video analysis was used to examine the actual changes in stride length, stride frequency and walking speed caused by the manipulations of leg length. Unexpectedly, quantitative characteristics of walking behaviour remained almost unaffected by imposed changes in leg length, demonstrating remarkable robustness of leg coordination and walking performance.These data further allowed normalisation of homing distances displayed by manipulated animals with regard to scaling and speed effects. The predicted changes in homing distance are in quantitative agreement with the experimental data, further supporting the pedometer hypothesis.

Path integration, although inherently error-prone, is a common navigation strategy in animals, particularly where environmental orientation cues are rare. The desert ant Cataglyphis fortis is a prominent example, covering large distances on foraging excursions. The stride integrator is probably the major source of path integration errors. A detailed analysis of walking behaviour in Cataglyphis is thus of importance for assessing possible sources of errors and potential compensation strategies. Zollikofer (J Exp Biol 192:95–106, 1994a) demonstrated consistent use of the tripod gait in Cataglyphis, and suggested an unexpectedly constant stride length as a possible means of reducing navigation errors. Here, we extend these studies by more detailed analyses of walking behaviour across a large range of walking speeds. Stride length increases linearly and stride amplitude of the middle legs increases slightly linearly with walking speed. An initial decrease of swing phase duration is observed at lower velocities with increasing walking speed. Then it stays constant across the behaviourally relevant range of walking speeds. Walking speed is increased by shortening of the stance phase and of the stance phase overlap. At speeds larger than 370 mms−1, the stride frequency levels off, the duty factor falls below 0.5, and Cataglyphis transitions to running with aerial phases.Electronic supplementary materialThe online version of this article (doi:10.1007/s00359-015-0999-2) contains supplementary material, which is available to authorized users.

Air-cushioned spheres are widely used as treadmills to study behavioural and neurophysiological questions in numerous species. We describe an improved spherical treadmill design that reliably registers the path and walking behaviour of an animal walking on top of the sphere. The simple and robust set-up consists of a very light hollowed styrofoam ball supported by an air stream in a hollow half sphere and can be used indoors and outdoors. Two optical mouse sensors provided with lenses of 4.6 mm focal length detect the motion of the sphere with a temporal resolution of more than 200 frames sand a spatial resolution of less than 0.2 mm. The treadmill can be used in an open-or closed-loop configuration with respect to yaw of the animal. The tethering allows animals to freely adjust their body posture and in the closed-loop configuration to quickly rotate around their yaw axis with their own moment of inertia. In this account, we present the first evidence of naturalistic homing navigation on a spherical treadmill for two species of Cataglyphis desert ants. We were able to evaluate with good precision the walking speed and angular orientation at any time. During homing the ants showed a significant difference in walking speed between the approach and search phases; moreover, they slowed down significantly as soon as they reached zero vector state, the fictive nest position.