After terrestrialization, the diversification of arthropods and vertebrates is thought to have occurred in two distinct phases1, the first between the Silurian and the Frasnian stages (Late Devonian period) (425–385 million years (Myr) ago), and the second characterized by the emergence of numerous new major taxa, during the Late Carboniferous period (after 345 Myr ago). These two diversification periods bracket the depauperate vertebrate Romer’s gap (360–345 Myr ago) and arthropod gap (385–325 Myr ago)1, which could be due to preservational artefact2,3. Although a recent molecular dating has given an age of 390 Myr for the Holometabola4, the record of hexapods during the Early–Middle Devonian (411.5–391 Myr ago, Pragian to Givetian stages) is exceptionally sparse and based on fragmentary remains, which hinders the timing of this diversification. Indeed, although Devonian Archaeognatha are problematic5,6, the Pragian of Scotland has given some Collembola and the incomplete insect Rhyniognatha, with its diagnostic dicondylic, metapterygotan mandibles5,7. The oldest, definitively winged insects are from the Serpukhovian stage (latest Early Carboniferous period)8. Here we report the first complete Late Devonian insect, which was probably a terrestrial species. Its ‘orthopteroid’ mandibles are of an omnivorous type, clearly not modified for a solely carnivorous diet. This discovery narrows the 45-Myr gap in the fossil record of Hexapoda, and demonstrates further a first Devonian phase of diversification for the Hexapoda, as in vertebrates, and suggests that the Pterygota diversified before and during Romer’s gap.

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We thank O. Béthoux who discovered and prepared most of the arthropod material from Strud including the specimen described herein. We also thank G. Budd and G. Edgecombe for discussion on the fossil material and improving the first version of the paper, Gesves local council staff and field workers of the Strud expeditions, G. Odebert and S. Fernandez for preparing illustrations, and C. Lemzaouda and O. Béthoux for photographs of the associated arthropod fauna. Thanks are due to A. Folie for our request of a catalogue number for the specimen described herein (requests for materials can be sent to afolie@naturalsciences.be). This work was partly supported by the French National Agency under the TERRES project (number ANR-2010-BLAN-607). Support for M.S.E. was provided by US National Science Foundation grant DEB-0542909.

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  1. UMR CNRS 7205, CP 50, Entomologie, Muséum national d’Histoire naturelle, 45 rue Buffon, F-75005 Paris, France

    • Romain Garrouste
    • , Patricia Nel
    • , Philippe Grandcolas
    • , Cyrille D’Haese
    •  & André Nel
  2. UMR CNRS 7207, Paléontologie, Muséum national d’Histoire naturelle, 8 rue Buffon, F-75005 Paris, France

    • Gaël Clément
    •  & Pierre Gueriau
  3. Division of Entomology, Natural History Museum, and Department of Ecology and Evolutionary Biology, 1501 Crestline Drive – Suite 140, University of Kansas, Lawrence, Kansas 66045, USA

    • Michael S. Engel
  4. Department of Earth Sciences, Uppsala University, Villavägen 16, SE-752 36 Uppsala, Sweden

    • Linda Lagebro
  5. Service de Paléontologie animale et humaine, Département de Géologie, Université de Liège, Bât. B.18, Allée du Six-Août, Sart Tilman, B-4000 Liège, Belgium

    • Julien Denayer
    •  & Sébastien Olive
  6. IPANEMA, USR 3461 CNRS - Ministère de la Culture et de la Communication, F-91190, Saint-Aubin, France

    • Pierre Gueriau
  7. CNRS – MNHN DICAP, Service Multimédia, CP 27, 57 rue Cuvier, F-75005 Paris, France

    • Patrick Lafaite
  8. Royal Belgian Institute of Natural Sciences Paleontology Department, 29, Rue Vautier, B-1000 Brussels, Belgium

    • Sébastien Olive
    •  & Cyrille Prestianni


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R.G., P.N. and G.C. are first authors with equal rank; R.G., A.N., P.N., P.G., C.D’H., L.L., M.S.E., J.D., C.P., P.G. and S.O. drafted the manuscript and prepared figures. A.N. and P.N. coordinated the manuscript; G.C. coordinated and participated in fieldwork at the Strud locality and contributed to the draft manuscript; L.L., J.D., C.P., P.G. and S.O. also participated in fieldwork.

Competing interests

The authors declare no competing financial interests.

Corresponding authors

Correspondence to Romain Garrouste or André Nel.

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