) of the signal without decrement over distances of several metres.

Although several behavioural studies demonstrate the ability of insects to localise the source of vibrations, it is still unclear how insects are able to perceive directional information from vibratory signals on solid substrates, because time-of-arrival and amplitude difference between receptory structures are thought to be too small to be processed by insect nervous systems. The termite Macrotermes natalensis communicates using vibrational drumming signals transmitted along subterranean galleries. When soldiers are attacked by predators, they tend to drum with their heads against the substrate and create a pulsed vibration. Workers respond by a fast retreat into the nest. Soldiers in the vicinity start to drum themselves, leading to an amplification and propagation of the signal. Here we show that M. natalensis makes use of a directional vibration sensing in the context of colony defence. In the field, soldiers are recruited towards the source of the signal. In arena experiments on natural nest material, soldiers are able to localise the source of vibration. Using two movable platforms allowing us to vibrate the legs of the left and right sides of the body with a time delay, we show that the difference in time-of-arrival is the directional cue used for orientation. Delays as short as 0.2 ms are sufficient to be detected. Soldiers show a significant positive tropotaxis to the platform stimulated earlier, demonstrating for the first time perception of time-of-arrival delays and vibrotropotaxis on solid substrates in insects.

Highlights d Acacia ants discriminate browser-induced vibrations from wind-induced vibrations d Browser-induced vibrations serve as a long-distance alarm cue for ants d Acacia ants make use of tropotactic directional vibration sensing Correspondence felix.hager@rub.de In BriefHager and Krausa show that the acacia ant Crematogaster mimosae defends its host tree by exploiting plant-borne vibrations caused by browsers feeding on the tree. Browser-induced vibrations serve as a long-distance alarm cue, and the ants make use of tropotactic directional vibration sensing to orient to the attacked part of the tree. SUMMARYLiving in the African savanna is dangerous, especially for plants. Many plants therefore engage in mutualism with ants, in which plants provide food and shelter in exchange for protection against herbivores. Ants become alarmed when the plant takes on some sort of damage. They immediately emerge from their plant shelter and aggressively defend the plant. Mammalian herbivores can have devastating effects on trees by browsing, breaking tree branches, stripping bark, and pushing over entire trees. However, mutualistic ants substantially reduce the amount of damage. To efficiently protect the tree, ants need to rapidly react together when the tree is under attack. Here, we show that the acacia ant Crematogaster mimosae defends its host tree by exploiting plant-borne vibrations caused by browsers feeding on the tree. Experiments with controlled vibrations show that ants discriminate browser-induced vibrations from those induced by wind, become alarmed, and patrol on the branches. Browser-induced vibrations serve as a long-distance alarm cue. The vibrations propagate through the whole acacia tree and trigger ants' defensive behavior, even on the other side of the tree. Furthermore, the ants make use of tropotactic directional vibration sensing to orient to the attacked part of the tree and fight back the attacker.

No abstract

Leafcutter ants communicate with the substrate-borne component of the vibratory emission produced by stridulation. Stridulatory signals in the genus Atta have been described in different behavioural contexts, such as foraging, alarm signalling and collective nest building. Stridulatory vibrations are employed to recruit nestmates, which can localize the source of vibration, but there is little information about the underlying mechanisms. Our experiments reveal that time-of-arrival delays of the vibrational signals are used for tropotactic orientation in Atta sexdens. The detected time delays are in the same range as the time delays detected by termites. Chemical communication is also of great importance in foraging organization, and signals of different modalities may be combined in promoting the organization of collective foraging. Here we show that the tropotactic orientation to vibrational signals interacts with chemical communication signals.