Eur. J. Entomol. 111 (4): 575-583, 2014 | DOI: 10.14411/eje.2014.073

Variation in the shape of the wings and taxonomy of Eurasian populations of the Calopteryx splendens complex (Odonata: Calopterygidae)

Saber SADEGHI1, Henri J. DUMONT2
1 Department of Biology, Faculty of Sciences, Shiraz University, Shiraz, Iran; e-mails:;
2 Department of Biology, Gent University, Ledeganckstraat 35, 9000-Gent, Belgium; e-mail:

We used geometric morphometrics to determine variation in the morphology of the forewings of individuals in 20 populations of Calopteryx splendens s.l. in Eurasia and related these to the circum-specific taxonomy of this taxon. We found differences in shape, with the largest and smallest centroid size of the wings in adjacent northern (orientalis) and western (intermedia) populations in Iran, respectively, so isolation and relationship are not necessarily determined by distance, but often associated with the stream basin inhabited. The variation in wing shape, however, was much greater. with Populations at the eastern edge of the range (Tajikistan, Kyrgyzstan and East Kazakhstan) uniquely different. Oddly, no taxonomic name is associated with them, although they may be among the oldest representatives of the splendens complex. The European and Asian populations are in two separate clades. One of these includes insects with no to a medium-sized wing spot, which does not reach the tip of the wing (waterstoni-group), while the other includes insects with very broad wing spots, or, when short, it extends to the very tip of the wings and most females are androchrome (ancilla or intermedia group). Turkmenistan and northern Iranian population form a separate line inside this clade, which we equate with Calopteryx orientalis. South Albanian and Greek populations are in a separate branch corresponding to ssp. balcanica; two populations from Ireland and Italy form a branch that has no equivalent in traditional taxonomy, while Calopteryx xanthostoma was not identified by its wing shape. Understanding this multitude of phenotypes and the enormous amount of variation within certain populations but not in others becomes easy if one assumes there were two probable late Pleistocene waves of migration, the first of insects lacking spots that migrated east and westwards from the South Black Sea basin, the second, perhaps from the west Caspian, composed of heavily spotted insects with androchromic females. The hybridization between these two waves resulted in the current plethora of colour forms and accounts for why similar phenotypes may turn up independently in widely distant locations.

Keywords: Odonata, Calopterygidae, Calopteryx splendens, Eurasia, wing shape, taxonomy

Received: November 9, 2013; Accepted: May 9, 2014; Revised: May 9, 2014; Prepublished online: August 29, 2014; Published: October 1, 2014Show citation

ACS AIP APA ASA Harvard Chicago IEEE ISO690 MLA NLM Turabian Vancouver
SADEGHI, S., & DUMONT, H.J. (2014). Variation in the shape of the wings and taxonomy of Eurasian populations of the Calopteryx splendens complex (Odonata: Calopterygidae). Eur. J. Entomol.111(4), 575-583. doi: 10.14411/eje.2014.073.
Download citation


  1. Adams D.C., Rohlf F.J. & Slice D. 2004: Geometric morphometrics: ten years of progress following the 'revolution'. - Ital. J. Zool. 71: 5-16 Go to original source...
  2. Asahina S. 1976: Indian paddy field Odonata taken by Miss I. Hattori. - Mushi 46: 115-127
  3. Bookstein F.L. 1991: Morphometric Tools for Landmark Data: Geometry and Biology. Cambridge Univ. Press, Cambridge, 435 pp
  4. Bookstein F.L. 1996: Combining the tools of geometric morphometrics. In Marcus L.F., Corti M., Loy A., Naylor G.J.P. & Slice D.E. (eds): Advances in Morphometrics. Plenum Press, New York, pp. 131-152
  5. Bookstein F.L. 1997: Shape and the information in medical images: a decade of the morphometric synthesis. - Comput. Vision Image Under. 66: 97-118 Go to original source...
  6. Chaput-Bardy A., Lemaitre C., Picard D. & Secondi J. 2008: In-stream and overland dispersal across a river network influences gene flow in a freshwater insect, Calopteryx splendens. - Mol. Ecol. 17: 3496-3505 Go to original source...
  7. Claude J., Paradis E., Tong H. & Auffray J.-C. 2002: A geometric morphometric assessment of the effects of environment and cladogenesis on the evolution of the turtle shell. - Biol. J. Linn. Soc. 79: 485-501 Go to original source...
  8. Cordoba-Aguilar A. & Cordero-Rivera A. 2005: Evolution and ecology of Calopterygidae (Zygoptera: Odonata): Status of knowledge and future research perspectives. - Neotrop. Entomol. 34: 861-879 Go to original source...
  9. Dumont H.J. 1972: The taxonomic status of Calopteryx xanthostoma (Charpentier, 1825) (Zygoptera: Calopterygidae). - Odonatologica 1: 21-29
  10. Dumont H.J., Demirsoy A. & Verschuren D. 1987: Breaking the Calopteryx-bottleneck: taxonomy and range of Calopteryx splendens waterstoni Schneider, 1984 and C. splendens tschaldirica Bartenef, 1909 (Zygoptera: Calopterygidae). -Odonatologica 16: 239-247
  11. Dumont H.J., Mertens J. & De Coster W. 1993: The Calopteryx splendens cline in southwestern France, analyzed by quantitative wingspot analysis (Zygoptera: Calopterygidae). - Odonatologica 22: 345-351
  12. Dumont H.J., Weekers P.H.H., Vanfleteren J.R. & Jonckheere D.E. 2005: Phylogenetic relationships, divergence time estimation and global biogeographic patterns of calopterygoid damselflies (Odonata, Zygoptera) inferred from ribosomal DNA sequences. - Syst. Biol. 54: 347-362 Go to original source...
  13. Goodall C.R. 1991: Procrustes methods in the statistical analysis of shape. - J. R. Stat. Soc. (B) 53: 285-339
  14. Hammer O.Y. & Harperd A.T. 2007: PAST - Paleontological Statistics, version 1.62. (
  15. Kendall D.G. 1997: The diffusion of shape. - Adv. Appl. Probab. 9: 428-430 Go to original source...
  16. Koskimaki J., Markus K.I., Rantala J., Taskinen J., Tynkkynen K. & Suhonen K. 2004: Immunocompetence and resource holding potential in the damselfly, Calopteryx virgo L. - Behav. Ecol. 15: 169-173 Go to original source...
  17. Kosterin O.E. & Sivtseva L.V. 2009: Odonata of Yakutia (Russia), with description of Calopteryx splendens njuja ssp. nov. (Zygoptera: Calopterygidae). - Odonatologica 38: 113-132
  18. Mcpeek M.A. 1990: Behavioral differences between Enallagma species (Odonata) influencing differential vulnerability to predators. - Ecology 71: 1714-1726 Go to original source...
  19. Milliken G.A. & Johnson D.E. 1984: Analysis of Messy Data. Vol. 1. Designed Experiments. Van Nostrand Reinhold, New York, NY, 473 pp
  20. Misof B., Anderson C.L. & Hadrys H. 2000: A phylogeny of the damselfly genus Calopteryx (Odonata) using mitochondrial 16S rDNA markers. - Mol. Phylogen. Evol. 15: 5-14 Go to original source...
  21. Rantala M.J., Koskimaki J., Taskinen J., Tynkkynen K. & Suhonen J. 2000: Immunocompetence, developmental stability and wingspot size in the damselfly Calopteryx splendens. - Proc. R. Soc. Lond. (B) 267: 2453-2457 Go to original source...
  22. Ricklefs R.E. & Miles D.B. 1994: Ecological and evolutionary inferences from morphology: an ecological perspective. In Wainwright P.C. & Reilly S.M. (eds): Ecological Morphology. Integrative Organismal Biology. University of Chicago Press, Chicago, IL, pp. 13-41
  23. Ris F. 1918: Der Artbegriff, insbesondere in der Entomologie. - Mitt. Entomol. (Zuerich) 4: 261-278
  24. Rohlf F.J. 1998: On applications of geometric morphometrics to studies of ontogeny and phylogeny. - Syst. Biol. 47: 147-158 Go to original source...
  25. Rohlf F.J. 2000: NTSYSpc Numerical Taxonomy and Multivariate Analysis System. Version 2.1 User Guide. Dept Ecol. & Evol., St. Univ. New York, Stony Brook, NY
  26. Rohlf F.J. 2002: Geometric morphometrics and phylogeny. In MacLeod N. & Forey P.L. (eds): Morphology, Shape and Phylogeny. Systematics Association Special Volume Series 64. Taylor and Francis, London, pp. 175-193
  27. Rohlf F.J. 2003: tpsSmall, Version 1.20 ed. Dept Ecol. & Evol., St. Univ. New York, Stony Brook, NY
  28. Rohlf F.J. 2004: Thin-plate Spline, Digitize Landmarks and Outlines, Version 1.20. Dept Ecol. & Evol., St. Univ. New York, Stony Brook, NY
  29. Rohlf F.J. 2006: tpsDig2, Digitize Landmarks and Outlines, Version 2.10. SUNY at Stony Brook
  30. Rohlf F.J. 2007: tpsRelw, Relative Warps Analysis, Version5/13/05. Department of Ecology and Evolution, State University of New York, Stony Brook, NY
  31. Rohlf F.J. 2010: tpsDig2, Digitize Landmarks and Outlines, Version 2.10. Dept Ecol. & Evol., St. Univ. New York, Stony Brook, NY
  32. Rueppell G., Hilfert-Rueppell D., Rehfeldt G. & Schuette C. 2005: Die Prachtlibellen Europas. Gattung Calopteryx. Die Neue Brehm-Buecherei 654. Westarp, Magdeburg, 256 pp
  33. Sadeghi S., Adriaens D. & Dumont H.J. 2009: Geometric morphometric analysis of wing shape variation in ten European populations of Calopteryx splendens Harris (1782) (Zygoptera: Odonata). - Odonatologica 38: 343-360
  34. Sadeghi S., Kyndt T. & Dumont H.J. 2010: Genetic diversity, population structure and taxonomy of Calopteryx splendens (Odonata: Calopterygidae) an AFLP analysis. - Eur. J. Entomol. 107: 137-146 Go to original source...
  35. Schmidt E. 1961: Ergebnisse det Deutschen Afghanisatn-Expedition 1956 der Landessammlungen fuer Naturkunde Karlsruhe sowie der Expeditionen J. Klapperich, Bonn 1952-53 und Dr. K. Lindberg, Lund (Schweden) 1957-60. - Beitr. Naturk. Forsch. SuedwDtl. 19: 399-435
  36. Seidenbusch R. 1994: Anmerkungen zur Variabilitaet des dunklen Fluegelfleckes bei Calopteryx splendens splendens Harris. - Acta Albert. Ratisb. 49: 209-212
  37. Sheets H.D. 2000: Integrated Morphometrics Package (IMP). Available at
  38. Siva-Jothy M.T. 1999: Male wing pigmentation may affect reproductive success via female choicein a calopterygid damselfly (Zygoptera). - Behaviour 136: 1365-1377 Go to original source...
  39. Siva-Jothy M.T. 2000: A mechanistic link between parasite resistance and expression of sexually selected trait in a damselfly. - Proc. R. Soc. Lond. (B) 267: 2523-2527 Go to original source...
  40. Slice D.E. & Bookstein F.L. 2007: A glossary for geometric morphometrics. Online,
  41. Sokal R.R. & Rohlf F.J. 1995: Biometry: The Principles and Practice of Statistics in Biological Research. 3rd. ed. W.H. Freeman, San Francisco, 887 pp
  42. Tynkkynen K., Rantala M.J. & Suhonen J. 2004: Interspecific aggression and character displacement in the damselfly. - J. Evol. Biol. 17: 259-267 Go to original source...
  43. Weekers P.H.H., De Jonckheere J.F. & Dumont H.J. 2001: Phylogenetic relationships inferred from ribosomal ITS sequences and biogeographic patterns in representatives of the genus Calopteryx (Insecta: Odonata) of the West Mediterranean and adjacent West European zone. - Mol. Phylogen. Evol. 20: 89-99 Go to original source...
  44. Zelditch M.L., Swiderski D.L., Sheets H.D. & Fink W.L. 2004: Geometric Morphometrics for Biologists: A Primer. Elsevier Acad. Press, London, 437 pp

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY), which permits use, distribution, and reproduction in any medium, provided the original publication is properly cited. No use, distribution or reproduction is permitted which does not comply with these terms.