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Courtship and mating behaviour in the parasitoid wasp Cotesia urabae (Hymenoptera: Braconidae): mate location and the influence of competition and body size on male mating success

  • G.A. Avila (a1) (a2), T.M. Withers (a2) (a3) and G.I. Holwell (a4)


Cotesia urabae is a solitary larval endoparasitoid that was introduced into New Zealand in 2011 as a classical biological control agent against Uraba lugens. A detailed knowledge of its reproductive biology is required to optimize mass rearing efficiency. In this study, the courtship and mating behaviour of C. urabae is described and investigated from a series of experiments, conducted to understand the factors that influence male mating success. Cotesia urabae males exhibited a high attraction to virgin females but not mated females, whereas females showed no attraction to either virgin or mated males. Male mating success was highest in the presence of a male competitor. Also, the time to mate was shorter and copulation duration was longer when a male competitor was present. Larger male C. urabae had greater mating success than smaller males when paired together with a single female. This knowledge can now be utilized to improve mass rearing methods of C. urabae for the future.


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Allen, G.R. (1990 a) Uraba lugens Walker (Lepidoptera: Noctuidae): larval survival and parasitoid biology in the field in South Australia. Journal of the Australian Entomological Society 29, 301312.
Allen, G.R. (1990 b) The phenologies of Cotesia urabae, Dolichogenidea eucalypti (Hymenoptera: Braconidae) and their host Uraba lugens (Lepidoptera: Noctuidae) in the Adelaide region. Australian Journal of Zoology 38, 347362.
Allen, G.R. & Keller, M.A. (1991) Uraba lugens (Lepidoptera: Noctuidae) and its parasitoids (Hymenoptera: Braconidae): temperature, host size and development. Environmental Entomology 20, 458469.
Avila, G.A., Berndt, L.A. & Holwell, G.I. (2013 a) First releases and monitoring of the biological control agent Cotesia urabae Austin and Allen (Hymenoptera: Braconidae). New Zealand Entomologist 36, 6572.
Avila, G.A., Berndt, L.A. & Holwell, G.I. (2013 b) Dispersal behavior of the parasitic wasp Cotesia urabae (Hymenoptera: Braconidae): a recently introduced biocontrol agent for the control of Uraba lugens (Lepidoptera: Nolidae) in New Zealand. Biological Control 66, 166172.
Avila, G.A., Withers, T.M. & Holwell, G.I. (2015) Host testing of the parasitoid Cotesia urabae (Austin & Allen, 1989) (Hymenoptera: Braconidae) to assess the risk posed to the New Zealand nolid moth Celama parvitis (Howes, 1917) (Lepidoptera: Nolidae): do host deprivation and experience influence acceptance of non-target hosts? Austral Entomology 54, 270277.
Avila, G.A., Withers, T.M. & Holwell, G.I. (2016 a) Olfactory cues used in host-habitat location and host location by the parasitoid Cotesia urabae (Hymenoptera: Braconidae). Entomologia Experimentalis et Applicata 158, 202209.
Avila, G.A., Withers, T.M. & Holwell, G.I. (2016 b) Laboratory odour-specificity testing of Cotesia urabae to assess potential risks to non-target species. BioControl 61, 365377.
Benelli, G., Bonsignori, G., Stefanini, C. & Canale, A. (2012) Courtship and mating behaviour in the fruit fly parasitoid Psyttalia concolor (Szépligeti) (Hymenoptera: Braconidae): the role of wing fanning. Journal of Pest Science 85, 5563.
Berndt, L.A. (2010) Will competition from Meteorus pulchricornis (Wesmael) (Hymenoptera: Braconidae) limit the success of the potential biocontrol agent Cotesia urabae Austin & Allen (Hymenoptera: Braconidae)? Australian Journal of Entomology 49, 369376.
Berndt, L.A., Withers, T.M., Mansfield, S. & Hoare, R.J.B. (2009) Non-target species selection for host range testing of Cotesia urabae . New Zealand Plant Protection 62, 168173.
Berndt, L.A., Sharpe, A.M., Gresham, B.A., Hayes, D. & Allen, G.R. (2013) Laboratory rearing of Cotesia urabae Austin and Allen (Hymenoptera: Braconidae), a biological control agent of Uraba lugens Walker (Lepidoptera: Nolidae). New Zealand Entomologist 36, 17.
Blanckenhorn, W.U., Mühlhäuser, C., Morf, C., Reusch, T. & Reuter, M. (2000) Female choice, female reluctance to mate and sexual selection on body size in the dung fly Sepsis cynipsea . Ethology 106, 577593.
Blaul, B. & Ruther, J. (2012) Body size influences male pheromone signals but not the outcome of mating contests in Nasonia vitripennis . Animal Behaviour 84, 15571563.
Bouchard, Y. & Cloutier, C. (1985) Role of olfaction in host finding by aphid parasitoid Aphidius nigripes (Hymenoptera: Aphidiidae). Journal of Chemical Ecology 11, 801808.
Bourdais, D. & Hance, T. (2009) Lack of behavioural evidence for kin avoidance in mate choice in a hymenopteran parasitoid (Hymenoptera: Braconidae). Behavioural Processes 81, 9294.
Bredlau, J.P., Mohajer, Y.J., Cameron, T.M., Kester, K.M. & Fine, M.L. (2013) Characterization and generation of male courtship song in Cotesia congregata (Hymenoptera: Braconidae). PloS ONE 8, e62051.
Conner, W., Roach, B., Benedict, E., Meinwald, J. & Eisner, T. (1990) Courtship pheromone production and body size as correlates of larval diet in males of the arctiid moth, Utetheisa ornatrix . Journal of Chemical Ecology 16, 543552.
De Freitas, M.D.R.T., De Lima Mendonça, A., Nascimento, R.R.D. & Sant’ Ana, A.E.G. (2004) Behavioural evidence for a female sex pheromone in Cotesia flavipes (Hymenoptera: Braconidae). Physiological Entomology 29, 183187.
Drosopoulos, S.A. & Claridge, M.F. (2006) Insect Sounds and Communication: Physiology, Behaviour, Ecology and Evolution. Boca Raton, FL, CRC Press.
Durocher-granger, L., Martel, V. & Boivin, G. (2011) Gamete number and size correlate with adult size in the egg parasitoid Trichogramma euproctidis . Entomologia Experimentalis et Applicata 140, 262268.
Field, S. & Keller, M. (1993 a) Courtship and intersexual signaling in the parasitic wasp Cotesia rubecula (Hymenoptera: Braconidae). Journal of Insect Behavior 6, 737750.
Field, S. & Keller, M. (1993 b) Alternative mating tactics and female mimicry as post-copulatory mate-guarding behaviour in the parasitic wasp Cotesia rubecula . Animal Behaviour 46, 11831189.
Godfray, H.C.J. (1994) Parasitoids: Behavioral and Evolutionary Ecology. Princeton, N.J., Princeton University Press.
Greenfield, M.D. (2002) Signalers and Receivers: Mechanisms and Evolution of Arthropod Communication. Oxford, New York, Oxford University Press.
Hardy, I.C.W., Ode, P.J. & Siva-Jothy, M.T. (2005) Mating behaviour. pp. 219260 in Jervis, M. (Ed.) Insects as Natural Enemies: A Practical Perspective. Dordrecht, Netherlands, Springers.
Henry, L. (2008) Assortative mating and the role of phenotypic plasticity in male competition: implications for gene flow among host-associated parasitoid populations. Biology Letters 4, 508.
Jahn-Eimermacher, A., Lasarzik, I. & Raber, J. (2011) Statistical analysis of latency outcomes in behavioral experiments. Behavioural Brain Research 221, 271275.
Jang, E.B. (1995) Effects of mating and accessory gland injections on olfactory-mediated behavior in the female mediterranean fruit-fly, Ceratitis capitata . Journal of Insect Physiology 41, 705710.
Johansson, B.G. & Jones, T.M. (2007) The role of chemical communication in mate choice. Biological Reviews 82, 265289.
Joyce, A., Bernal, J., Vinson, S. & Lomeli-Flores, R. (2009) Influence of adult size on mate choice in the solitary and gregarious parasitoids, Cotesia marginiventris and Cotesia flavipes . Journal of Insect Behavior 22, 1228.
Joyce, A.L., Hunt, R.E., Bernal, J.S. & Bradleigh Vinson, S. (2008) Substrate influences mating success and transmission of courtship vibrations for the parasitoid Cotesia marginiventris . Entomologia Experimentalis et Applicata 127, 3947.
Kleinbaum, D.G. & Klein, M. (2012) Survival analysis a self-learning text. New York, NY, Springer.
Kugimiya, S., Shimoda, T., Wajnberg, E., Uefune, M. & Takabayashi, J. (2010) Host-searching responses to herbivory-associated chemical information and patch use depend on mating status of female solitary parasitoid wasps. Ecological Entomology 35, 279286.
McClure, M., Whistlecraft, J. & McNeil, J.N. (2007) Courtship behaviour in relation to the female sex pheromone in the parasitoid Aphidius ervi (Hymenoptera: Braconidae). Journal of Chemical Ecology 33, 19461959.
McNeil, J.N. & Brodeur, J. (1995) Pheromone-mediated mating in the aphid parasitoid, Aphidius nigripes (Hymenoptera: Aphididae). Journal of Chemical Ecology 21, 959972.
Morbey, Y.E. (2013) Protandry, sexual size dimorphism, and adaptive growth. Journal of Theoretical Biology 339, 9399.
Painting, C.J., & Holwell, G.I. (2014) Exaggerated rostra as weapons and the competitive assessment strategy of male giraffe weevils. Behavioral Ecology 25, 12231232.
Péter, A. (2014) Solomon Coder (version beta 14.10.04): A simple solution and free solution for behavior coding. Available online at (accessed 9 November 2014).
Quicke, D.L.J. (2014) The Braconid and Ichneumonid Parasitoid Wasps Biology, Systematics, Evolution and Ecology. Hoboken, Wiley Blackwell.
Quinn, G.P. & Keough, M.J. (2002) Experimental Design and Data Analysis for Biologists. Cambridge, UK, Cambridge University Press.
Ramadan, M.M., Wong, T.T.Y. & Wong, M.A. (1991) Influence of parasitoid size and age on male mating success of opiinae (Hymenoptera: Braconidae), larval parasitoids of fruit flies (Diptera: Tephritidae). Biological Control 1, 248255.
Rowbottom, R., Allen, G., Walker, P. & Berndt, L. (2013) Phenology, synchrony and host range of the Tasmanian population of Cotesia urabae introduced into New Zealand for the biocontrol of Uraba lugens . BioControl 58, 625633.
Ruther, J. (2013) Novel insights into pheromone-mediated communication in parasitic hymenopterans. pp. 112144 in Wajnberg, E. & Colazza, S. (Eds) Chemical Ecology of Insect Parasitoids. Chichester, UK, John Wiley & Sons, Ltd.
Teder, T. (2005) Male-biased size dimorphism in ichneumonine wasps (Hymenoptera: Ichneumonidae) – the role of sexual selection for large male size. Ecological Entomology 30, 342349.
Vinson, S.B. (1972) Courtship behavior and evidence of a sex pheromone in the parasitoid Campoletis sonorensis (Hymenoptera: Ichneumonidae). Environmental Entomology 1, 409414.
Wajnberg, E. (2010) Genetics of the behavioral ecology of egg parasitoids. pp. 149165 in Consoli, F. L., Parra, J. R. & Zucchi, R. A. (Eds) Egg Parasitoids in Agroecosystems with Emphasis on Trichogramma. Dordrecht, London, Springer.
Wong, B.B.M. & Candolin, U. (2005) How is female mate choice affected by male competition? Biological Reviews 80, 559571.
Wyatt, T.D. (2003) Pheromones and Animal Behaviour: Communication by Smell and Taste. Cambridge, Cambridge University Press.
Xu, H., Veyrat, N., Degen, T. & Turlings, T.C.J. (2014) Exceptional use of sex pheromones by parasitoids of the genus Cotesia: males are strongly attracted to virgin females, but are no longer attracted to or even repelled by mated females. Insects 5, 499512.


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Courtship and mating behaviour in the parasitoid wasp Cotesia urabae (Hymenoptera: Braconidae): mate location and the influence of competition and body size on male mating success

  • G.A. Avila (a1) (a2), T.M. Withers (a2) (a3) and G.I. Holwell (a4)


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