Life history, systematics and flight ability of the Early Permian stem-mayflies in the genus Misthodotes Sellards, 1909 (Insecta, Ephemerida, Permoplectoptera)

Abstract

Abstract Background The stem-group of Ephemeroptera is phylogenetically important for understanding key steps in evolutionary history of early pterygote insects. However, these taxa have been mostly studied from the taxonomy point of view focused on the pattern of wing venation and often using only classical optical microscopy devices. In-depth studies on detailed morphology of the different body structures are scarcely performed, although the results are critical for elucidation of life history traits and their evolutionary pattern among the basal pterygotes. Results New information is presented on the morphology of two species of Misthodotes, which are stem-mayflies from the Early Permian. Based on new results obtained from a re-examination of the type specimens and supplementary material, we infer the life history traits of both the adult and larval stages of these Palaeozoic insects and reconsider previous interpretations. For the first time, we report the structure of the thoracic pleura and the articulation at the base of the wing in a stem-group of Ephemeroptera and compare them with those of extant mayflies. We also provide additional support for the systematic placement of investigated taxa and an amended diagnosis of the genus Misthodotes. Conclusions Adult Misthodotes sharovi and Misthodotes zalesskyi had chewing mouthparts, which enabled them to scavenge or feed on plants. The wing apparatus was adapted for slow powered flapping flight and gliding, using long caudal filaments for steering. The wing base does not have rows of articulary sclerites as previously hypothesized for some Palaeozoic taxa but inflexible axilla similar to that found in modern mayflies. The structure of the thoracic pleura is also similar to that in the crown group of Ephemeroptera, while differences in the course of sutures may be explained by an evolutionary trend towards more powerful dorsoventral flying musculature and forewing-based flight (anteromotorism) in modern taxa. There is no evidence for swarming behaviour and mating in the air as occurs in modern mayflies as they had none of the associated morphological adaptations. Putative larvae of Misthodotes can not be unambiguously associated with the adults. They also exhibit some morphological specializations of Protereismatidae like 9 pairs of abdominal tracheal gills supporting their benthic lifestyle with legs adapted to burrowing.