Morphological discrimination of the genera Binodoxys Mackauer and Trioxys Haliday (Hymenoptera: Braconidae: Aphidiinae) on the basis of the general shape of forewings

Maja Lazarević, Marijana Ilić Milošević, Saša Stojan Stanković, Vladimir Žikić


The genera Binodoxys and Trioxys, members of the Aphidiinae subfamily, are classified in the same tribe Trioxini and the subtribe Trioxina. These two genera are morphologically very similar but differ ecologically. Generally, Binodoxys species are specialized in parasitizing aphids on herbaceous plants unlike Trioxys species which have co-evolved with the tree attacking aphids. For this study 80 right forewings of four Binodoxys and two Trioxys species were analyzed using the geometric morphometrics method. The sixteen specific landmarks were digitalized on each wing and the results demonstrated statistically significant differences of wing shape between the genera. By visualizing using the CVA method, the wing structure that was the most variable was the radial sector vein – which is much shorter in Binodoxys than in Trioxys species.


Trioxina, geometric morphometrics, Binodoxys, Trioxys, parasitoids

Full Text:



Akhtar, M.S., Dey, D., Usmani, M.K. 2011: A catalogue of aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae) from India. Insecta Mundi, 151: 1-31.

Bookstein, F.L. 1991: Morphometric Tools for Landmark Data: Geometry and Biology. Cambridge University Press, New York, 435 pp.

Ilić Milošević, M., Petrović, A., Stanković, S.S., Čkrkić, J., Starý, P., Žikić, V., Tomanović, Ž. 2015: Taxonomic Position and Phylogenetic Relationships of the Genera and Species Euaphidius and Remaudierea (Hymenoptera: Braconidae: Aphdiinae) Analyzed Using Molecular Markers and Geometric Morphometrics. Annals of Entomological Society of America, 108: 435-445.

Kavallieratos, N.G., Tomanović, Ž., Starý, P., Žikić, V., Petrović-Obradović, O. 2010: Parasitoids (Hymenoptera: Braconidae: Aphidiinae) attacking aphids feeding on Solanaceae and Cucurbitaceae crops in Southeastern Europe: Aphidiine aphid- plant associations and key. Annals of Entomological Society of America, 103: 153–164.

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

Kos, K., Petrović, A., Starý, P., Kavallieratos, N.G., Ivanović, A., Toševski, I., Jakše, J., Trdan, S., Tomanović, Ž. 2011: On the Identity of Cereal Aphid Parasitoid Wasps Aphidius uzbecistanicus, Aphidius rhopalosiphi, and Aphidius avenaphis (Hymenoptera: Braconidae: Aphidiinae) by Examination of COI Mitochondrial Gene, Geometric Morphometrics, and Morphology. Annals of the Entomological Society of America, 104: 1221-1232.

Mackauer, M. 1959: Die mittel-, west- und nordeuropäischen Arten der Gattung Trioxys Haliday (Hymenoptera: Braconidae: Aphidiinae). Beiträgezur Entomologie, 9: 144-179.

Mackauer, M. 1960: Zur Systematik der Gattung Trioxys Haliday (Hymenoptera: Braconidae: Aphidiinae). Beiträgezur Entomologie, 10: 137-160.

Mescheloff, E., Rosen, D. 1993: Biosystematic studies of the Aphidiidae of Israel (Hymenoptera: Ichneumonoidea). 5. The genera Trioxys and Binodoxys. Israel Journal of Entomology, XXVII: 31-47.

Mitrovski-Bogdanović, A., Ivanović, A., Tomanović, Ž., Žikić, V., Starý, P., Kavallieratos, N.G. 2009: Sexual dimorphism in Ephedrus persicae (Hymenoptera: Braconidae: Aphidiinae): intraspecific variation in size and shape. The Canadian Entomologist, 141: 550-560.

Mitrovski-Bogdanović, A., Tomanović, Ž., Mitrović, M., Petrović, A., Ivanović, A., Žikić, V., Starý, P., Voburger, C. 2014: The Praon dorsale – yomenae s.str. complex (Hymenoptera, Braconidae, Aphidiinae): Species discrimination using morphometrics and molecular makers with description of a new species. Zoologischer Anzeiger, 253: 270-282.

Pike, K.S., Starý, P., Miller, R., Allison, D., Boydston, L., Graf, G., Miller, T. 1996: New species and host records of aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae) from the Pacific Northwest, U.S.A. Proceeding of Entomological Society of Washington, 98: 570-591.

Pike, K.S., Starý, P., Miller, T., Graf, G., Allison, D., Boydston, L. & Miller, R. 2000: Aphid parasitoid (Hymenoptera: Braconidae: Aphidiinae) of Northwest USA. Proceeding of Entomological Society of Washington, 102: 688-740.

Rohlf, F.J. 2005: TpsDig, digitize landmarks and outlines, version 2.05. Department of Ecology and Evolution, State University of New York at Stony Brook (

Sheets, H.D. 2003: IPM Suite (Integrated Morphometrics Package). Canisius College, Buffalo, NY.

Stanković, S.S., Petrović, A., Ilić Milošević, M., Starý, P., Kavalllieratos, N.G., Žikić, V., Tomanović, Ž. 2015: Morphological and molecular characterization of common European species of Adialytus (Hymenoptera: Braconidae: Aphidiinae) based on the mtCOI barcoding gene and geometric morphometrics of forewings. European Journal of Entomology, 112: 165-174.

Starý, P. 1981: Biosystematical clasification of Trioxys Haliday and related genera (Hymenoptera, Aphidiidae). Bollettino del Laboratorio di Entomologia Agraria Portici, XXXVIII: 85-93.

StatSoft Inc. 2004. Statistica (data analysis software system), version 7,

Tomanović, Ž., Petrović, A., Mitrović, M., Kavallieratos, N.G., Starý, P., Rakhshani, E., Rakhshanipour, M., Popović, A. Shukshuk, A.H., Ivanović, A. 2014: Molecular and morphological variability within the Aphidius colemani group with redescription of Aphidius platensis Brethes (Hymenoptera: Braconidae: Aphdiinae). Bulletin of Entomological Research, 104: 552-565.

Yu, D.S., Van Achterberg, C., Horstmann, K. 2012: World Ichneumonoidea 2011. Taxonomy, Biology, Morphology and Distribution. Taxapad (Scientific Names for Information Management), Interactive Catalogue, Ottawa Available on:

Zelditch, M.L., Swiderski, D.L., Sheets, H.D. 2012: Geometric morphometrics for biologist: a primer. Academic Press, London, UK, 488 pp.

Žikić, V., Ilić - Milošević, M., Stanković, S., Petrović, A., Petrović-Obradović, O., Kavallieratos, N. G., Starý, P. & Tomanović, Ž., 2012: Aphidiinae (Braconidae: Hymenoptera) of Serbia and Montenegro – tritrophic interactions. Acta Entomologica Serbica, 17: 83-105.

Žikić, V., Petrović, A., Ivanović, A. 2014: Allometric shape changes indicate siginficant divergence in the wing shape between asexual and sexual lineages of Lysiphlebus fabarum (Marshall) (Hymenoptera: Braconidae: Aphidiinae). Acta Entomologica Serbica, 19: 53-62.

Žikić, V., Tomanović, Ž., Ivanović, A., Kavallieratos, N.G., Starý, P., Stanisavljević, Lj.Ž., Rakhshani, E. 2009: Morphological Characterization of Ephedrus persicae Biotypes (Hymenoptera: Braconidae: Aphdiinae) in the Palaearctic. Annals of the Entomological Society of America, 102: 1-11.


  • There are currently no refbacks.