A winged relative of ice crawlers in amber bridges the cryptic extant Xenonomia and a rich fossil record

Until the advent of phylogenomics, the atypical morphology of extant representatives of the insect orders Grylloblattodea (ice crawlers) and Mantophasmatodea (gladiators) had confounding effects on efforts to resolve their placement within Polyneoptera. This recent research has unequivocally shown that these species‐poor groups are closely related and form the clade Xenonomia. Nonetheless, divergence dates of these groups remain poorly constrained, and their evolutionary history debated, as the few well‐identified fossils, characterized by a suite of morphological features similar to that of extant forms, are comparatively young. Notably, the extant forms of both groups are wingless, whereas most of the pre‐Cretaceous insect fossil record is composed of winged insects, which represents a major shortcoming of the taxonomy. Here, we present new specimens embedded in Early Cretaceous amber from Myanmar and belonging to the recently described species Aristovia daniili. The abundant material and pristine preservation allowed a detailed documentation of the morphology of the species, including critical head features. Combined with a morphological data set encompassing all Polyneoptera, these new data unequivocally demonstrate that A. daniili is a winged stem Grylloblattodea. This discovery demonstrates that winglessness was acquired independently in Grylloblattodea and Mantophasmatodea. Concurrently, wing apomorphic traits shared by the new fossil and earlier fossils demonstrate that a large subset of the former “Protorthoptera” assemblage, representing a third of all known insect species in some Permian localities, are genuine representatives of Xenonomia. Data from the fossil record depict a distinctive evolutionary trajectory, with the group being both highly diverse and abundant during the Permian but experiencing a severe decline from the Triassic onwards.

Methods
The RTI file composing this dataset was derived from a set of photographs obtained using a light dome of about 30 cm in diameter and equipped with 54 LEDs, and a camera Canon EOS 5DS equipped with a MP-E 65 mm macro lens, both driven by a control box (dome and control box, Flydome, Paris, France; camera body and lens, Canon, Tokyo, Japan). The 45 usable photographs (9 were excluded due to improper exposure) were batch-optimized, including a ‘horizontal flipping’ step, using Adobe Photoshop CS6 and were further compiled into an RTI file using the RTI Builder software v. 2.0.2 using the HSH fitter (software freely available from Cultural Heritage Imaging, San Francisco, CA, USA).