On the value of Burmese amber for understanding insect evolution: Insights from †Heterobathmilla – an exceptional stem group genus of Strepsiptera (Insecta)

Burmese amber and amber from other periods and regions became a rich source of new extinct insect species and yielded important insights in insect evolution in the dimension of time. Amber fossils have contributed to the understanding of the phylogeny, biology, and biogeography of insects and other groups, and have also gained great importance for dating molecular trees. Another major potential is the documentation of faunal, floral and climatic shifts. Evolutionary transitions can be well-documented in amber fossils and can reveal anatomical transformations and the age of appearance of structural features. Here, using a new stem group species of Strepsiptera from Burmite, we evaluate this potential of amber insect fossils to assess the current phylogeny of Strepsiptera, with the main emphasis on the early splitting events in the stem group. Amber fossils have greatly contributed to the understanding of the evolution of Strepsiptera in the late Mesozoic and the Cenozoic. †Heterobathmilla kakopoios Pohl and Beutel gen. et sp. n. described here is placed in the stem group of the order, in a clade with †Kinzelbachilla (†Kinzelbachillidae) and †Phthanoxenos (†Phthanoxenidae). †Phthanoxenidae has priority over †Kinzelbachillidae, and the latter is synonymised. The superb details available from this new fossil allowed us to explore unique features of the antennae, mouthparts, and male copulatory apparatus, and to provide a phylogenetic hypothesis for the order. The younger †Protoxenos from Eocene Baltic amber was confirmed as sister to all remaining extinct and extant groups of Strepsiptera, whereas the position of the Cretaceous †Cretostylops in the stem group remains ambivalent. While the value of Burmite and amber from other periods has a recognized impact on our knowledge of the evolution in various lineages, this new fossil does not fundamentally change our picture of the phylogeny and evolution of early Strepsiptera. However, it offers new insights into the morphological diversity in the early evolution of the group.