Organisms Diversity & Evolution

, Volume 13, Issue 1, pp 55–66 | Cite as

The head anatomy of Epiophlebia superstes (Odonata: Epiophlebiidae)

  • Alexander Blanke
  • Felix Beckmann
  • Bernhard Misof
Original Article


The relic dragonfly family Epiophlebiidae is recovered as sistergroup of Anisoptera (= Epiprocta) by most molecular and morphological analyses. However, in a recent study it was placed within Anisoptera as sister group of Cordulegastridae. In another study, several affinities to Zygoptera in the morphology of the ovipositor and the egg-laying behaviour were pointed out. Here, we present a detailed study of the outer, as well as the inner, head morphology of Epiophlebia superstes. Compared with the last detailed literature account, three additional mandibular muscles were discovered, as well as additional buccal and pharyngeal muscles. The results are compared with the anatomic features of Zygoptera and Anisoptera. A formal character evaluation focused on head characters confirms the sistergroup relationship of Epiophlebiidae and Anisoptera.


Muscle equipment Zygoptera Anisoptera Anisozygoptera Epiophlebiidae Phylogeny Anatomy MicroCT 3D-reconstruction Epiprocta 



Fabian Wilde (HZG, Geesthacht), as well as Sebastian Thieß and Wolfgang Drube (DESY, Hamburg), provided excellent support at the DESY synchrotron facilities. The scanning was done in the course of proposals no. I-20080169 and I-20090211. Marco Stampanoni and Peter Modregger are sincerely thanked for their support at the PSI for SLS TOMCAT beamline analysis (proposals 20100137 and 20110069). Irina Ruf (Steinmann Institute, Bonn, Germany) made microCT investigations at the device of the Steinmann Institute possible. We would like to express our gratitude to Professor Ryuichiro Machida (University of Tsukuba, Japan) who kindly provided specimens of E. superstes. Benjamin Wipfler and Rolf Beutel (University of Jena, Germany) are thanked for useful advice. Furthermore, we thank two anonymous reviewers for their valuable comments. The laboratory members of the ZFMK are sincerely thanked for their support.

Supplementary material

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  1. Ando, H. (1962). The comparative embryology of Odonata with special reference to a relic dragonfly. Epiophlebia superstes. Tokyo: The Japan Society for the Promotion of Science.Google Scholar
  2. Asahina, S. (1954). A morphological study of a relic dragonfly Epiophlebia superstes Selys (Odonata, Anisozygoptera). Tokyo: The Japan Society for the Promotion of Science.Google Scholar
  3. Bechly, G. (1996). Morphologische Untersuchungen am Flügelgeäder der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata) unter besonderer Berücksichtigung der phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, 2, 1–402.Google Scholar
  4. Beckmann, F., Herzen, J., Haibel, A., Müller, B., & Schreyer, A. (2008). High density resolution in synchrotron-radiation-based attenuation-contrast microtomography. Proceedings of SPIE, 7078, 70781D–70783D.CrossRefGoogle Scholar
  5. Betz, O., Wegst, U., Weide, D., Heethoff, M., Helfen, L., Lee, W.-K., et al. (2007). Imaging applications of synchrotron X-ray phase-contrast microtomography in biological morphology and biomaterials science. I. General aspects of the technique and its advantages in the analysis of millimetre-sized arthropod structure. Journal of Microscopy, 227, 51–71.PubMedCrossRefGoogle Scholar
  6. Blanke, A., Wipfler, B., Letsch, H., Koch, M., Beckmann, F., Beutel, R. G., & Misof, B. (2012). Revival of Palaeoptera – head characters support a monophyletic origin of Odonata and Ephemeroptera (Insecta). Cladistics (in press).Google Scholar
  7. Bybee, S. M., Ogden, T. H., Branham, M. A., & Whiting, M. F. (2008). Molecules, morphology and fossils: a comprehensive approach to odonate phylogeny and the evolution of the odonate wing. Cladistics, 23, 1–38.Google Scholar
  8. Carle, F. L. (1982). The wing vein homologies and phylogeny of the Odonata: a continuing debate. Societas Internationalis Odonatologica Rapid Communications, 4, 1–66.Google Scholar
  9. Dumont, H. J., Vierstraete, A., & Vanfleteren, J. R. (2010). A molecular phylogeny of the Odonata (Insecta). Systematic Entomology, 35, 6–18.CrossRefGoogle Scholar
  10. DuPorte, E. M. (1946). Observations on the morphology of the face in insects. Journal of Morphology, 79, 371–417.PubMedCrossRefGoogle Scholar
  11. DuPorte, E. M. (1957). The comparative morphology of the insect head. Annual Review of Entomology, 2, 55–70.CrossRefGoogle Scholar
  12. Fiala, J. C. (2005). Reconstruct: a free editor for serial section microscopy. Journal of Microscopy, 218, 52–61.PubMedCrossRefGoogle Scholar
  13. Fleck, G., Ullrich, B., Brenk, M., Wallnisch, C., Orland, M., Bleidissel, S., et al. (2008). A phylogeny of anisopterous dragonflies (Insecta, Odonata) using mtRNA genes and mixed nucleotide/doublet models. Journal of Zoological Systematics and Evolutionary Research, 46, 310–322.CrossRefGoogle Scholar
  14. Gade, G., Simek, P., & Fescemyer, H. W. (2011). Adipokinetic hormones provide inference for the phylogeny of Odonata. Journal of Insect Physiology, 57, 174–178.PubMedCrossRefGoogle Scholar
  15. Goloboff, P. A., Farris, J. S., & Nixon, K. C. (2008). TNT, a free program for phylogenetic analysis. Cladistics, 24, 774–786.CrossRefGoogle Scholar
  16. Gorb, S. N. (1999). Evolution of the dragonfly head-arresting system. Proceedings of the Royal Society of London. Series B: Biological Sciences 266, 525–535.Google Scholar
  17. Klass, K.-D. (2008). The female abdomen of ovipositor-bearing Odonata (Insecta: Pterygota). Arthropod Systematics & Phylogeny 66, 45–142.Google Scholar
  18. Letsch, H. O., Greve, C., Kück, P., Fleck, G., Stocsits, R. R., & Misof, B. (2009). Simultaneous alignment and folding of 28 S rRNA sequences uncovers phylogenetic signal in structure variation. Molecular Phylogenetics and Evolution, 53, 758–771.PubMedCrossRefGoogle Scholar
  19. Li, J.-K., Nel, A., Zhang, X.-P., Fleck, G., Gao, M.-X., Lin, L. I. N., et al. (2012). A third species of the relict family Epiophlebiidae discovered in China (Odonata: Epiproctophora). Systematic Entomology, 37, 408–412.CrossRefGoogle Scholar
  20. Lohmann, H. (1996). Das phylogenetische System der Anisoptera (Odonata). Entomologische Zeitschrift, 106, 209–266.Google Scholar
  21. Mathur, P. N., & Mathur, K. C. (1961). Studies on the cephalic musculature of adult Ictinus angulosus Selys (Odonata, Anisoptera, Gomphidae, Ictinae). Journal of Morphology, 109, 237–249.CrossRefGoogle Scholar
  22. Matushkina, N. A. (2008). The ovipositor of the relic dragonfly Epiophlebia superstes: a morphological re-examination (Odonata: Epiophlebiidae). International Journal of Odonatology, 11, 71–80.CrossRefGoogle Scholar
  23. Misof, B., Rickert, A. M., Buckley, T. R., Fleck, G., & Sauer, K. P. (2001). Phylogenetic signal and its decay in mitochondrial SSU and LSU rRNA gene fragments of Anisoptera. Molecular Biology and Evolution, 18, 27–37.PubMedCrossRefGoogle Scholar
  24. Pass, G. (1991). Antennal circulatory organs in Onychophora, Myriapoda and Hexapoda – Functional morphology and evolutionary implications. Zoomorphology, 110, 145–164.CrossRefGoogle Scholar
  25. Pohl, H. (2010). A scanning electron microscopy specimen holder for viewing different angles of a single specimen. Microscopy Research and Technique, 73, 1073–1076.PubMedCrossRefGoogle Scholar
  26. Rehn, A. C. (2003). Phylogenetic analysis of higher-level relationships of Odonata. Systematic Entomology, 28, 181–239.CrossRefGoogle Scholar
  27. Romeis, B. (1989). Mikroskopische Technik (17). Munich: Urban & Schwarzenberg.Google Scholar
  28. Seifert, G. (1995). Entomologisches Praktikum (3rd ed.). Stuttgart: Georg Thieme Verlag.Google Scholar
  29. Short, J. R. T. (1955). The morphology of the head of Aeshna cyanea (Müller) (Odonata, Anisoptera). Transactions of the Royal Entomological Society London, 106, 197–211.CrossRefGoogle Scholar
  30. Snodgrass, R. E. (1935). Principles of insect morphology. McGraw-Hill, New York, XXX.Google Scholar
  31. Snodgrass, R. E. (1947). The insect cranium and the “epicranial suture”. Smithsonian Miscellaneous Collections, 107, 1–52.Google Scholar
  32. Staniczek, A. H. (2000). The mandible of silverfish (Insecta: Zygentoma) and mayflies (Ephemeroptera): Its morphology and phylogenetic significance. Zoologischer Anzeiger, 239, 147–178.Google Scholar
  33. Staniczek, A. H. (2001). Der Larvenkopf von Oniscigaster wakefieldi McLachlan, 1873 (Insecta: Ephemeroptera: Oniscigastridae). Ein Beitrag zur vergleichenden Anatomie und Phylogenie der Eintagsfliegen. PhD thesis, Eberhard-Karls-Universität Tübingen.Google Scholar
  34. Strenger, A. (1952). Die funktionelle und morphologische Bedeutung der Nähte am Insektenkopf. Zoologische Jahrbücher, 72, 468–521.Google Scholar
  35. Tillyard, R. J. (1921). On an anisozygopterous larva from the Himalayas (Order Odonata). Records of the Indian Museum, 22, 93–107.Google Scholar
  36. Trueman, J. W. H. (1996). A prelimnary cladistic analysis of odonate wing venation. Odonatologica, 25, 59–72.Google Scholar
  37. Wipfler, B., Machida, R., Müller, B., & Beutel, R. G. (2011). On the head morphology of Grylloblattodea (Insecta) and the systematic position of the order, with a new nomenclature for the head muscles of Dicondylia. Systematic Entomology, 36, 241–266.CrossRefGoogle Scholar

Copyright information

© Gesellschaft für Biologische Systematik 2012

Authors and Affiliations

  • Alexander Blanke
    • 1
  • Felix Beckmann
    • 2
  • Bernhard Misof
    • 1
  1. 1.Zentrum für molekulare BiodiversitätZoologisches Forschungsmuseum Alexander KoenigBonnGermany
  2. 2.Institute of Materials ResearchHelmholtz-Zentrum GeesthachtGeesthachtGermany

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