Bombardier beetles repel invasive bullfrogs.

Invasive non-native predators negatively affect native species; however, some native species can survive the predation pressures of invasive species by using pre-existing antipredator strategies or evolving defenses against invasive predators. The American bullfrog Lithobates catesbeianus (Anura: Ranidae) has been intentionally introduced to many countries and regions, and has impacted native animals through direct predation. Bombardier beetles (Coleoptera: Carabidae: Brachininae: Brachinini) discharge chemicals at a temperature of approximately 100 °C from the tip of the abdomen when they are attacked by predators. This "bombing" can successfully repel predators. However, adults of a native bombardier beetle Pheropsophus (Stenaptinus) occipitalis jessoensis have been reportedly found in the gut contents of the introduced bullfrog L. catesbeianus in Japan. These records suggest that the invasive bullfrog L. catesbeianus attacks the native bombardier beetle P. occipitalis jessoensis under field conditions in Japan; however, the effectiveness of the bombing defense against invasive bullfrogs is unclear. To test the effectiveness of the bombing defense against bullfrogs, we investigated the behavioral responses of L. catesbeianus juveniles to P. occipitalis jessoensis adults under laboratory conditions. Contrary to previous gut content results, almost all the bullfrogs (96.3%) rejected bombardier beetles before swallowing them; 88.9% rejected the beetles after being bombed, and 7.4% stopped attacking the beetles before being bombed. Only 3.7% successfully swallowed and digested the beetle. All of the beetles collected from non-bullfrog-invaded sites could deter bullfrogs, suggesting that the pre-existing defenses of bombardier beetles played an essential role in repelling bullfrogs. When treated beetles that were unable to discharge hot chemicals were provided, 77.8% of bullfrogs successfully swallowed and digested the treated beetles. These results indicate that bombing is important for the successful defense of P. occipitalis jessoensis against invasive bullfrogs. Although invasive bullfrogs have reportedly impacted native insect species, P. occipitalis jessoensis has an existing defense mechanism strong enough to repel the invasive predators.

Sugiura S ，Date T 《PeerJ》

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

Sugiura S 《PeerJ》

Beetle bombing always deters praying mantises.

Some animals have evolved chemical weapons to deter predators. Bombardier beetles (Coleoptera: Carabidae: Brachininae: Brachinini) can eject toxic chemicals at temperatures of 100 °C from the tips of their abdomens, 'bombing' the attackers. Although some bombardier beetles can reportedly deter predators, few studies have tested whether bombing is essential for successful defence. Praying mantises (Mantodea) are ambush predators that attack various arthropods. However, it is unclear whether bombardier beetles deter mantises. To test the defensive function of bombing against praying mantises, I observed three mantis species, Tenodera sinensis, Tenodera angustipennis, and Hierodula patellifera (Mantidae), attacking the bombardier beetle Pheropsophus jessoensis (Carabidae: Brachininae: Brachinini) under laboratory conditions. All mantises easily caught the beetles using their raptorial forelegs, but released them immediately after being bombed. All of the counterattacked mantises were observed to groom the body parts sprayed with hot chemicals after releasing the beetles. When treated P. jessoensis that were unable to eject hot chemicals were provided, all mantises successfully caught and devoured the treated beetles. Therefore, bombing is essential for the successful defence of P. jessoensis against praying mantises. Consequently, P. jessoensis can always deter mantises.

Sugiura S 《PeerJ》

Defenses of whirligig beetles against native and invasive frogs.

Many native insects have evolved defenses against native predators. However, their defenses may not protect them from non-native predators due to a limited shared history. The American bullfrog, Aquarana catesbeiana (Anura: Ranidae), which has been intentionally introduced to many countries, is believed to impact native aquatic animals through direct predation. Adults of whirligig beetles (Coleoptera: Gyrinidae), known for swimming and foraging on the water surface of ponds and streams, reportedly possess chemical defenses against aquatic predators, such as fish. Although whirligig beetles potentially encounter both bullfrogs and other frogs in ponds and lakes, the effectiveness of their defenses against frogs has been rarely studied. To assess whether whirligig beetles can defend against native and non-native frogs, we observed the behavioral responses of the native pond frog, Pelophylax nigromaculatus (Anura: Ranidae), and the invasive non-native bullfrog, A. catesbeiana, to native whirligig beetles, Gyrinus japonicus and Dineutus orientalis, in Japan. Adults of whirligig beetles were provided to frogs under laboratory conditions. Forty percent of G. japonicus and D.orientalis were rejected by P. nigromaculatus, while all whirligig beetles were easily consumed by A. catesbeiana. Chemical and other secondary defenses of G. japonicus and D. orientalis were effective for some individuals of P. nigromaculatus but not for any individuals of A. catesbeiana. These results suggest that native whirligig beetles suffer predation by invasive non-native bullfrogs in local ponds and lakes in Japan.

Sugiura S ，Hayashi M 《PeerJ》

Bombardiers and assassins: mimetic interactions between unequally defended insects.

In defensive mimicry, resemblance between unequally defended species can be parasitic; this phenomenon has been termed quasi-Batesian mimicry. Few studies have used real co-mimics and their predators to test whether the mimetic interactions were parasitic. Here, we investigated the mimetic interaction between two well-defended insect species, the bombardier beetle Pheropsophus occipitalis jessoensis (Coleoptera: Carabidae) and the assassin bug Sirthenea flavipes (Hemiptera: Reduviidae), using their potential predator, the pond frog Pelophylax nigromaculatus (Anura: Ranidae), which coexists with these insect species in the same habitat in Japan. We observed behavioural responses of this frog species (adults and juveniles) to adult Ph. occipitalis jessoensis and adult S. flavipes under laboratory conditions. Among the frogs, 100% and 75% rejected Ph. occipitalis jessoensis and S. flavipes, respectively, suggesting that, compared with the assassin bug S. flavipes, the bombardier beetle Ph. occipitalis jessoensis is more well-defended against frogs. An assassin bug or a bombardier beetle was provided to a frog that had encountered the other insect species. Frogs with a history of assassin bug encounter demonstrated a lower rate of attack toward bombardier beetles. Similarly, frogs with a history of bombardier beetle encounter demonstrated a lower rate of attack toward assassin bugs. Therefore, both the bombardier beetle Ph. occipitalis jessoensis and the assassin bug S. flavipes benefit from the mimetic interaction.

Sugiura S ，Hayashi M 《PeerJ》