Drilling into solid substrates with slender beam-like structures is a mechanical challenge, but is regularly done by female parasitic wasps. The wasp inserts her ovipositor into solid substrates to deposit eggs in hosts, and even seems capable of steering the ovipositor while drilling. The ovipositor generally consists of three longitudinally connected valves that can slide along each other. Alternative valve movements have been hypothesized to be involved in ovipositor damage avoidance and steering during drilling. However, none of the hypotheses have been tested in vivo. We used 3D and 2D motion analysis to quantify the probing behavior of the fruit-fly parasitoid Diachasmimorpha longicaudata (Braconidae) at the levels of the ovipositor and its individual valves. We show that the wasps can steer and curve their ovipositors in any direction relative to their body axis. In a soft substrate, the ovipositors can be inserted without reciprocal motion of the valves. In a stiff substrate, such motions were always observed. This is in agreement with the damage avoidance hypothesis of insertion, as they presumably limit the overall net pushing force. Steering can be achieved by varying the asymmetry of the distal part of the ovipositor by protracting one valve set with respect to the other. Tip asymmetry is enhanced by curving of ventral elements in the absence of an opposing force, possibly due to pretension. Our findings deepen the knowledge of the functioning and evolution of the ovipositor in hymenopterans and may help to improve man-made steerable probes.Diachasmimorpha longicaudata | ovipositor kinematics | buckling avoidance | spatial probing | minimally invasive probe F rom a mechanical perspective, it is very difficult to drill into a solid substrate with a very thin probe, because it can easily bend and break. Parasitic wasps, however, do this regularly when they use their slender ovipositors to search for hosts in solid substrates, such as fruits or even wood (1-3).The general morphology of the ovipositor is similar across all wasp species (4, 5); it consists of four elements, called valves, of which two are often merged such that three functional valves remain (Fig. 1). In most species, the distal part of the ovipositor is morphologically distinct (3, 6), which we will refer to as the tip. The valves can slide along each other (5, 7) and do not get dislocated under natural conditions, because they are longitudinally connected via a tongue-and-groove mechanism (5, 8-10). The ovipositor and the "wasp waist," a constriction of the body between the first and second abdominal segment (11), are essential in probing behavior and are therefore considered to be instrumental in the evolution of the order (11-15). The shape, structure, and mechanical properties of the ovipositors are putatively adapted to the substrates into which the animals need to probe (6,(16)(17)(18), and because both substrates and hosts are so diverse, this might have resulted in high species diversification of the hymenopterans (13,14). Howeve...