Anti-predator defences of a bombardier beetle: is bombing essential for successful escape from frogs?

Some animals, such as the bombardier beetles (Coleoptera: Carabidae: Brachinini), have evolved chemical defences against predators. When attacked, bombardier beetles can discharge noxious chemicals at temperatures of approximately 100 °C from the tip of their abdomens, "bombing" their attackers. Although many studies to date have investigated how bombardier beetles discharge defensive chemicals against predators, relatively little research has examined how predators modify their attacks on bombardier beetles to avoid being bombed. In this study, I observed the black-spotted pond frog (Anura: Ranidae) attacking the bombardier beetle under laboratory conditions. In Japan, is a generalist predator in grasslands where the bombardier beetle frequently occurs. Almost all the frogs (92.9%) observed rejected live bombardier beetles; 67.9% stopped their attacks once their tongues touched the beetles, and 25.0% spat out the beetles immediately after taking the beetles into their mouths. No beetle bombed a frog before being taken into a frog's mouth. All beetles taken into mouths bombed the frogs. Only 7.1% of the frogs swallowed live bombardier beetles after being bombed in the mouth. When dead beetles were provided instead, 85.7% of the frogs rejected the dead beetles, 71.4% stopped their attacks after their tongues touched the beetles, and 14.3% spat out the beetles. Only 14.3% of the frogs swallowed the dead beetles. The results suggest that the frogs tended to stop their predatory attack before receiving a bombing response from the beetles. Therefore, bombing was not essential for the beetles to successfully defend against the frogs. Using its tongue, may be able to rapidly detect a deterrent chemical or physical characteristics of its potential prey and thus avoid injury by stopping its predatory attack before the beetle bombs it.