Skip to main content

Thank you for visiting You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

A complete insect from the Late Devonian period


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.

This is a preview of subscription content, access via your institution

Relevant articles

Open Access articles citing this article.

Access options

Rent or buy this article

Prices vary by article type



Prices may be subject to local taxes which are calculated during checkout

Figure 1
Figure 2: General habitus of Strudiella devonica gen. et sp. nov.
Figure 3: Strudiella devonica gen. et sp. nov., counterpart details.
Figure 4: Phylogeny of basal hexapod clades.


  1. Ward, P., Labandeira, C., Laurin, M. & Berner, R. A. Confirmation of Romer’s gap as a low oxygen interval constraining the timing of initial arthropod and vertebrate terrestrialization. Proc. Natl Acad. Sci. USA 103, 16818–16822 (2006)

    Article  ADS  CAS  PubMed  PubMed Central  Google Scholar 

  2. Smithson, T. R., Wood, S. P., Marshall, J. E. A. & Clack, J. A. Earliest Carboniferous tetrapod and arthropod faunas from Scotland populate Romer’s gap. Proc. Natl Acad. Sci. USA 109, 4532–4537 (2012)

    Article  ADS  CAS  PubMed  PubMed Central  Google Scholar 

  3. Retallack, G. J. Woodland hypothesis for Devonian evolution of tetrapods. J. Geol. 119, 235–258 (2011)

    Article  ADS  Google Scholar 

  4. Rehm, P. et al. Dating the arthropod tree based on large-scale transcriptome data. Mol. Phyl. Evol. 61, 880–887 (2011)

    Article  Google Scholar 

  5. Grimaldi, D. & Engel, M. S. Evolution of the Insects (Cambridge Univ. Press, 2005)

    Google Scholar 

  6. Shear, W. A. & Selden, P. A. in Plants Invade the Land. Evolutionary & Environmental Perspectives (eds Gensel, P. G. & Edwards, D. ) 29–51 (Columbia Univ. Press, 2001)

    Google Scholar 

  7. Engel, M. S. & Grimaldi, D. New light shed on the oldest insect. Nature 427, 627–630 (2004)

    Article  ADS  CAS  PubMed  Google Scholar 

  8. Prokop, J., Nel, A. & Hoch, I. Discovery of the oldest known Pterygota in the Lower Carboniferous of the Upper Silesian Basin in the Czech Republic (Insecta: Archaeorthoptera). Geobios 38, 383–387 (2005)

    Article  Google Scholar 

  9. Clément, G. et al. Devonian tetrapod from Western Europe. Nature 427, 412–413 (2004)

    Article  ADS  PubMed  Google Scholar 

  10. Blieck, A., Clément, G. & Streel, M. in The Terrestrialization Process: Modelling Complex Interactions at the Biosphere–Geosphere Interface (eds Vecoli, M., Clément, G. & Meyer-Berthaud, B. ) 129–138 (Geological Society of London, 2010)

    Google Scholar 

  11. Prestianni, C., Streel, M., Thorez, J. & Gerrienne, P. Strud: old quarry, new discoveries. Preliminary report. Carnets Géol. Mémoir. 2007, 43–47 (2007)

    Google Scholar 

  12. Martínez-Delclòs, X., Briggs, D. E. G. & Peñalver, E. Taphonomy of insects in carbonates and amber. Palaeogeogr. Palaeoclimateol. Palaeoecol. 3225, 1–46 (2004)

    Google Scholar 

  13. Bitsch, C. & Bitsch, J. Phylogenetic relationships of basal hexapods among the mandibulate arthropods: a cladistic analysis based on comparative morphological characters. Zool. Scr. 33, 511–550 (2004)

    Article  Google Scholar 

  14. McLaughlin, P. A. Comparative Morphology of Recent Crustacea ( W. H., Freeman, 1980)

    Google Scholar 

  15. Imms, A. D. On the antennal musculature in insects and other arthropods. Q. J. Microsc. Sci. 81, 273–320 (1939)

    Google Scholar 

  16. Yack, J. E. The structure and function of auditory chordotonal organs in insects. Microsc. Res. Tech. 63, 315–337 (2004)

    Article  PubMed  Google Scholar 

  17. Sturm, H. & Machida, R. Archaeognatha. Handbook of Zoology, Vol. 4 of Arthropoda: Insecta, i–viii, 1–213 (Berlin: Walter de Gruyter, 2001)

    Google Scholar 

  18. Furst Von Lieven, A. The transformation from monocondylous to dicondylous mandibles in the Insecta. Zool. Anz. 239, 139–146 (2000)

    Google Scholar 

  19. Kukalová-Peck, J. Phylogeny of higher taxa in Insecta: finding synapomorphies in the extant fauna and separating them from homoplasies. Evol. Biol. 35, 4–51 (2008)

    Article  Google Scholar 

  20. Staniczek, A. H. The mandible of silverfish (Insecta: Zygentoma) and mayflies (Ephemeroptera): its morphology and phylogenetic significance. Zool. Anz. 239, 147–178 (2000)

    Google Scholar 

  21. Bitsch, J. The arthropod mandible: morphology and evolution. Phylogenetic implications. Ann. Soc. Entomol. Fr. (N.S.) 37, 305–321 (2001)

    Google Scholar 

  22. Snodgrass, R. E. Comparative studies on the jaws of mandibulate arthropods. Smithson. Misc. Coll. 116, 1–85 (1952)

    Google Scholar 

  23. Kukalová-Peck, J. Carboniferous protodonatoid dragonfly nymphs and the synapomorphies of Odonatoptera and Ephemeroptera (Insecta: Palaeoptera). Palaeodiversity 2, 169–198 (2009)

    Google Scholar 

  24. Bechly, G., Brauckmann, C., Zessin, W. & Gröning, E. New results concerning the morphology of the most ancient dragonflies (Insecta: Odonatoptera) from the Namurian of Hagen-Vorhalle (Germany). Z. Zool. Syst. Evol. 39, 209–226 (2001)

    Article  Google Scholar 

  25. Gangwere, S. K. The structural adaptations of mouthparts in Orthoptera and allies. Eos . Rev. Esp. Entomol. 41, 67–85 (1965)

    Google Scholar 

  26. Guthrie, D. M. & Tindall, A. R. The Biology of the Cockroach (Edward Arnold, 1968)

    Google Scholar 

  27. Brauckmann, C., Schöllmann, L. & Sippel, W. Die fossilen Insekten, Spinnentiere und Eurypteriden von Hagen-Vorhalle. Geol. Paläontol. Westfalen 59, 1–89 (2003)

    Google Scholar 

  28. Labandeira, C. C. Silurian to Triassic plant and hexapod clades and their associations: new data, a review, and interpretations. Arthr. Syst. Phyl. 64, 53–94 (2006)

    Google Scholar 

  29. Labandeira, C. C. The origin of herbivory on land: initial patterns of plant tissue consumption by arthropods. Insect Sci. 14, 259–275 (2007)

    Article  Google Scholar 

  30. Stein, W. E., Berry, C. M., VanAller Hernick, L. & Mannolini, F. Surprisingly complex community discovered in the mid-Devonian fossil forest at Gilboa. Nature 483, 78–81 (2012)

    Article  ADS  CAS  PubMed  Google Scholar 

Download references


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 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.

Author information

Authors and Affiliations



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.

Corresponding authors

Correspondence to Romain Garrouste or André Nel.

Ethics declarations

Competing interests

The authors declare no competing financial interests.

Supplementary information

Supplementary Information

This file contains Supplementary Text and Supplementary Figures 1-4. This file was replaced on 5 October 2012. (PDF 7116 kb)

PowerPoint slides

Rights and permissions

Reprints and Permissions

About this article

Cite this article

Garrouste, R., Clément, G., Nel, P. et al. A complete insect from the Late Devonian period. Nature 488, 82–85 (2012).

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI:

This article is cited by


By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.


Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing