Contributions to Zoology, 85 (1) – 2016Mei Liu; Na Ma; Bao-Zhen Hua: Intraspecific morphological variation of the scorpionfly Dicerapanorpa magna (Chou) (Mecoptera: Panorpidae) based on geometric morphometric analysis of wings

To refer to this article use this url:


Adams DC, Rohlf FJ, Slice DE. 2013. A field comes of age: geometric morphometrics in the 21st century. Hystrix, the Italian Journal of Mammalogy 24: 7-14.

Allen CE, Zwaan BJ, Brakefield PM. 2011. Evolution of sexual dimorphism in the Lepidoptera. Annual Review of Entomology 56: 445-464.

An ZS, Kutzbach JE, Prell WL, Porter SC. 2001. Evolution of Asian monsoons and phased uplift of the Himalaya-Tibetan plateau since Late Miocene times. Nature 411: 62-66.

Aytekin MA, Terzo M, Rasmont P, Çağatay N. 2007. Landmark based geometric morphometric analysis of wing shape in Sibiricobombus Vogt (Hymenoptera: Apidae: Bombus Latreille). Annales de la Société Entomologique de France (N.S.) 43: 95-102.

Baracchi D, Dapporto L, Turillazzi S. 2011. Relevance of wing morphology in distinguishing and classifying genera and species of Stenogastrinae wasps. Contributions to Zoology 80: 191-199.

Barão KR, Gonçalves GL, Mielke OH, Kronforst MR, Moreira GR. 2014. Species boundaries in Philaethria butterflies: an integrative taxonomic analysis based on genitalia ultrastructure, wing geometric morphometrics, DNA sequences, and amplified fragment length polymorphisms. Zoological Journal of the Linnean Society 170: 690-709.

Benítez HA, Bravi R, Parra LE, Sanzana MJ, Sepúlveda-Zúñiga E. 2013. Allometric and non-allometric patterns in sexual dimorphism discrimination of wing shape in Ophion intricatus: Might two male morphotypes coexist? Journal of Insect Science 13: 1-10.

Blanckenhorn WU. 2000. The evolution of body size: what keeps organisms small? Quarterly Review of Biology 75: 385-407.

Bookstein FL. 1991. Morphometric tools for landmark data. Cambridge: Cambridge University Press.

Bookstein FL. 1996. Combining the tools of geometric morphometrics. Pp. 131-151 in: Marcus LF, Corti M, Loy A, Nayulor GJP, Slice DE, eds, Advances in morphometrics. New York: Plenum Press.

Byers GW, Thornhill R. 1983. Biology of the Mecoptera. Annual Review of Entomology 28: 203-228.

Chou I, Ran RB, Wang SM. 1981. Study on the classification of Chinese Mecoptera (I, II). Entomotaxonomia 3: 1-22.

Dong YP, Zhang GW, Neubauer F, Liu XM, Genser J, Hauzenberger C. 2011. Tectonic evolution of the Qinling orogen, China: review and synthesis. Journal of Asian Earth Sciences 41: 213-237.

Dryden IL, Mardia KV. 1998. Statistical analysis of shape. New York: Wiley.

Fairbairn DJ, Preziosi RF. 1976. Sexual selection and the evolution of the sexual size dimorphism in the waterstrider, Aquarius remigis. Evolution 50: 1549-1559.

Francuski LJ, Vujić A, Kovačević A, Ludoški J, Milankov V. 2009. Identification of the species of the Cheilosia variabilis group (Diptera, Syrphidae) from the Balkan Peninsula using wing geometric morphometrics, with the revision of status of C. melanopa redi Vujić, 1996. Contributions to Zoology 78: 129-140.

Gavrilets S, Li H, Vose MD. 2000. Patterns of parapatric speciation. Evolution 54: 1126-1134.

Gerard M, Michez D, Fournier D, Maebe K, Smagghe G, Biesmeijer JC, De Meulemeester T. 2015. Discrimination of haploid and diploid males of Bombus terrrestris (Hymenoptera; Apidae) based on wing shape. Apidologie 46: 644-653.

Gidaszewski NA, Baylac M, Klingenberg CP. 2009. Evolution of sexual dimorphism of wing shape in the Drosophila melanogaster subgroup. BMC Evolutionary Biology 9: 110.

Gilchrist GW. 1990. The consequences of sexual dimorphism in body size for butterfly flight and thermoregulation. Functional Ecology 4: 475-487.

Grimaldi D, Engel MS. 2005. Evolution of the insects. Cambridge: Cambridge University Press.

Grodnitsky DL. 1999. Form and function of insect wings: the evolution of biological structures. Baltimore, USA: The Johns Hopkins University Press.

Haas HL, Tolley KA. 1998. Geographic variation of wing morphology in three Eurasian populations of the fruit fly, Drosophila lummei. Journal of Zoology 245: 197-203.

Honěk A. 1993. Intraspecific variation in body size and fecundity in insects: a general relationship. Oikos 66: 483-492.

Hou XY, Hua BZ. 2008. Structures of the female reproductive systems in Panorpidae (Mecoptera) with remarks on their taxonomic significance. Acta Zootaxonomica Sinica 33: 427-434.

Hu GL, Yan G, Xu H, Hua BZ. 2015. Molecular phylogeny of Panorpidae (Insecta: Mecoptera) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 85:22-31.

Hua BZ, Chou I. 1997. The Panorpidae (Mecoptera) of Funiu Mountain in Henan Province. Entomotaxonomia 19: 273-278.

Huberty CJ, Olejnik S. 2006. Applied MANOVA and discriminant analysis. 2nd ed. Hoboken (NJ): Wiley.

IBM Corporation. 2010. IBM SPSS Statistics for Windows, version 19.0. Armonk, NY: IBM Corporation.

Jarošík V, Honek A. 2007. Sexual differences in insect development time in relation to sexual size dimorphism. Pp. 205-211 in Fairbairn DJ, Blanckenhorn WU, Székely T, eds, Sex, size and gender roles: Evolutionary studies of sexual size dimorphism. Oxford: Oxford University Press.

Jones JR. 2010. Intraspecific variation in a scorpionfly newly recorded from Texas and the state of taxonomy of North American Panorpidae (Mecoptera). Psyche 2010: 152981.

Jorge LR, Cordeiro-Estrela P, Klaczko LB, Moreira GRP, Freitas AVL. 2011. Host-plant dependent wing phenotypic variation in the neotropical butterfly Heliconius erato. Biological Journal of the Linnean Society 102: 765-774.

Klingenberg CP. 2011. MorphoJ: an integrated software package for geometric morphometrics. Molecular Ecology Resources 11: 353-357.

Klingenberg CP, McIntyre GS. 1998. Geometric morphometrics of developmental instability: analyzing patterns of fluctuating asymmetry with Procrustes methods. Evolution 52: 1363-1375.

Liang ZX. 2002. The natural environment change of Qin-Ba Mountains during the period of history. Collections of Essays on Chinese Historical Geography 17: 39-47.

Ma N, Hu GL, Zhang JX, Hua BZ. 2014. Morphological variation of the scorpionfly Panorpa obtusa Cheng (Mecoptera: Panorpidae) with a new synonym. PLoS ONE 9: e108545.

Ma N, Liu SY, Hua BZ. 2011. Morphological diversity of male salivary glands in Panorpidae (Mecoptera). European Journal of Entomology 108: 493-499.

Mayr E. 1963. Animal species and evolution. Cambridge: Harvard University Press.

McCulloch, GA, Wallis, GP, Waters, JM. 2009. Do insects lose flight before they lose their wings? Population genetic structure in subalpine stoneflies. Molecular Ecology, 18: 4073-4087.

McLachlan AJ. 1986. Sexual dimorphism in midges: strategies for flight in the rain-pool dweller Chironomus imicola (Diptera: Chironomidae). The Journal of Animal Ecology 55: 261-267.

McPeek MA. 1990. Behavioral differences between Enallagma species (Odonata) influencing differential vulnerability to predators. Ecology 71: 1714-1726.

Michaux B. 1989. Morphological variation of species through time. Biological Journal of the Linnean Society 38: 239-255.

Milliken GA, Johnson DE. 2009. Analysis of messy data. Vol. 1: Designed Experiments. 2nd ed. New York: CRC Press.

Nedeljković Z, Ačanski J, Vujić A, Obreht D, Ðan M, Ståhls G, Radenković S. 2013. Taxonomy of Chrysotoxum festivum Linnaeus, 1758 (Diptera: Syrphidae) – an integrative approach. Zoological Journal of the Linnean Society 169: 84-102.

Neto JM, Gordinho L, Belda EJ, Marín M, Monrós JS, Fearon P, Crates R. 2013. Phenotypic divergence among West European populations of reed bunting Emberiza schoeniclus: the effects of migratory and foraging behaviours. PLoS ONE 8: e63248.

Nie XN, Hua BZ. 2004. Panorpidae (Mecoptera) from Huoditang in Qinling Mountains. Entomotaxonomia 26: 187-192.

Palmer M. 2002. Landmark-based morphometric analysis of two sibling species of the genus Asida (Coleoptera, Tenebrionidae). Contributions to Zoology 70: 1-10.

Pepinelli M, Spironello M, Currie DC. 2013. Geometric morphometrics as a tool for interpreting evolutionary transitions in the black fly wing (Diptera: Simuliidae). Zoological Journal of the Linnean Society 169: 377-388.

Perrard A, Baylac M, Carpenter JM, Villemant C. 2014. Evolution of wing shape in hornets: why is the wing venation efficient for species identification? Journal of Evolutionary Biology 27: 2665-2675.

R Core Team. 2013. R: A language and environment for statistical computing. R Foundation for Statistical Computing. Vienna, Austria. Available at

Riedel A, Sagata K, Suhardjono YR, Tänzler, R, Balke M. 2013. Integrative taxonomy on the fast track – towards more sustainability in biodiversity research. Frontiers in Zoology 10: 15.

Rohlf FJ. 2004. tpsDig2, version 2.0. Available at

Rohlf FJ. 2006. tpsRelw, version 1.44. Available at

Rohlf FJ, Loy A, Corti M. 1996. Morphometric analysis of Old World Talpidae (Mammalia, Insectivora) using partial-warp scores. Systematic Biology 45: 344-362.

Rohlf FJ, Slice D. 1990. Extensions of the Procrustes method for the optimal superimposition of landmarks. Systematic Zoology 39: 40-59.

Sheets HD. 2012. IMP software series. Canisius College, Buffalo, New York.

Singh BN. 2012. Concepts of species and modes of speciation. Current Science 103: 784-790.

Stillwell RC, Blanckenhorn WU, Teder T, Davidowitz G, Fox CW. 2010. Sex differences in phenotypic plasticity affect variation in sexual size dimorphism in insects: from physiology to evolution. Annual Review of Entomology 55: 227-245.

Sun JH, Ren D, Huang JD. 2007. Current knowledge of research on Mecoptera fossils in China. Acta Zootaxonomica Sinica 32: 881-889.

Teder T, Tammaru T. 2005. Sexual size dimorphism within species increases with body size in insects. Oikos 108: 321-334.

Thornhill R. 1979. Adaptive female-mimicking behavior in a scorpionfly. Science 205: 412-414.

Thornhill R. 1980. Competition and coexistence among Panorpa scorpionflies (Mecoptera: Panorpidae). Ecological Monographs 50: 179-197.

Velzen RV, Wahlberg N, Sosef MSM, Bakker FT. 2013. Effects of changing climate on species diversification in tropical forest butterflies of the genus Cymothoe (Lepidoptera: Nymphalidae). Biological Journal of the Linnean Society 108: 546-564.

Wang Y, Yan JP. 2011. Evolution of the Qinling Mountain and the impact factors. Journal of Northwest University (Natural Science Edition) 41: 163-169.

Wang ZC, Zou CN, Tao SZ, Li J, Wang SQ, Zhao CY. 2004. Analysis on tectonic evolution and exploration potential in Dabashan foreland basin. Acta Petrolei Sinica 25: 23-28.

Ward PH. 1979. Structural variation in the genitalia of the Panorpa alpina-complex (Mecoptera). Systematic Entomology 4: 71-79.

Ward PH. 1983. Scorpion-flies of the Panorpa cognata-complex in the western Palaearctic region (Mecoptera). Journal of Natural History 17: 627-645.

White MJD. 1978. Modes of speciation. San Francisco: WH Freeman.

Ying T. 1994. An analysis of the flora of Qinling Mountain range: its nature, characteristics and origins. Acta Phytotaxonomica Sinica 32: 389-410.

Yuan JH, Cheng FY, Zhou SL. 2012. Genetic structure of the tree peony (Paeonia rockii) and the Qinling Mountains as a geographic barrier driving the fragmentation of a large population. PLoS ONE 7: e34955.

Zelditch ML, Swiderski DL, Sheets HD, Flink WL. 2004. Geometric morphometrics for biologists: A primer. San Diego: Elsevier Academic Press.

Zhang PZ, Zheng DW, Yin GM, Yuan DY, Zhang GL, Li CY, Wang ZC. 2006. Discussion on late Cenozoic growth and rise of northeastern margin of the Tibetan Plateau. Quaternary Science 26: 5-13.

Zhong W, Hua BZ. 2013. Dicerapanorpa, a new genus of East Asian Panorpidae (Insecta: Mecoptera: Panorpidae) with descriptions of two new species. Journal of Natural History 47: 1019-1046.