Geographical distribution and phoretic associations of the viviparous nematode Tokorhabditis atripennis with Onthophagus dung beetles in Japan

Viviparity is generally considered to be rare in animals. In nematodes, only six species of Rhabditida are viviparous. Five of these species have been identified in association with Onthophagus dung beetles, with Tokorhabditis atripennis being repeatedly isolated from the dung beetle Onthophagus atripennis in Japan. T. atripennis is easy to culture in a laboratory setting, and its host, O. atripennis, is distributed all over Japan. Therefore, T. atripennis is an ideal candidate for ecological and evolutionary studies on viviparity. However, the extent of their distribution and relationship with dung beetles, as well as habitats, remain unclear. In the present study, we conducted field surveys and successfully isolated 27 strains of viviparous nematodes associated with tunneler dung beetles from various regions of Japan, all of which were identified as T. atripennis. T. atripennis exhibited a strong association with Onthophagus dung beetles, especially O. apicetinctus and O. atripennis. And it was predominantly found in specific anatomical locations on the beetle bodies, such as the ‘groove between pronotum and elytron’ and the ‘back of the wings’. Our findings suggest that Onthophagus species are the primary hosts for T. atripennis, and T. atripennis exhibits a close relationship with the living environments of tunneler beetles. This association may play a significant role in the evolution of viviparity in nematodes.