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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

Published online by Cambridge University Press:  15 December 2016

G.A. Avila*
The New Zealand Institute for Plant & Food Research Limited, Mt Albert, Private Bag 92169, Auckland 1142, New Zealand Better Border Biosecurity, New Zealand
T.M. Withers
Better Border Biosecurity, New Zealand Scion (New Zealand Forest Research Institute), Private Bag 3020, Rotorua 3046, New Zealand
G.I. Holwell
School of Biological Sciences, The University of Auckland, Private Bag 92019, Auckland 1142, New Zealand
*Author for correspondence Phone: +64 9 925 7198 Fax: +64 9 925 7001 E-mail:


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.

Research Papers
Copyright © Cambridge University Press 2016 

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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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.Google Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
Bouchard, Y. & Cloutier, C. (1985) Role of olfaction in host finding by aphid parasitoid Aphidius nigripes (Hymenoptera: Aphidiidae). Journal of Chemical Ecology 11, 801808.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle ScholarPubMed
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.CrossRefGoogle Scholar
Drosopoulos, S.A. & Claridge, M.F. (2006) Insect Sounds and Communication: Physiology, Behaviour, Ecology and Evolution. Boca Raton, FL, CRC Press.Google Scholar
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.CrossRefGoogle Scholar
Field, S. & Keller, M. (1993 a) Courtship and intersexual signaling in the parasitic wasp Cotesia rubecula (Hymenoptera: Braconidae). Journal of Insect Behavior 6, 737750.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
Godfray, H.C.J. (1994) Parasitoids: Behavioral and Evolutionary Ecology. Princeton, N.J., Princeton University Press.Google Scholar
Greenfield, M.D. (2002) Signalers and Receivers: Mechanisms and Evolution of Arthropod Communication. Oxford, New York, Oxford University Press.Google Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle ScholarPubMed
Jahn-Eimermacher, A., Lasarzik, I. & Raber, J. (2011) Statistical analysis of latency outcomes in behavioral experiments. Behavioural Brain Research 221, 271275.CrossRefGoogle ScholarPubMed
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.CrossRefGoogle Scholar
Johansson, B.G. & Jones, T.M. (2007) The role of chemical communication in mate choice. Biological Reviews 82, 265289.CrossRefGoogle ScholarPubMed
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
Kleinbaum, D.G. & Klein, M. (2012) Survival analysis a self-learning text. New York, NY, Springer.Google Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
McNeil, J.N. & Brodeur, J. (1995) Pheromone-mediated mating in the aphid parasitoid, Aphidius nigripes (Hymenoptera: Aphididae). Journal of Chemical Ecology 21, 959972.CrossRefGoogle Scholar
Morbey, Y.E. (2013) Protandry, sexual size dimorphism, and adaptive growth. Journal of Theoretical Biology 339, 9399.CrossRefGoogle ScholarPubMed
Painting, C.J., & Holwell, G.I. (2014) Exaggerated rostra as weapons and the competitive assessment strategy of male giraffe weevils. Behavioral Ecology 25, 12231232.CrossRefGoogle Scholar
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).Google Scholar
Quicke, D.L.J. (2014) The Braconid and Ichneumonid Parasitoid Wasps Biology, Systematics, Evolution and Ecology. Hoboken, Wiley Blackwell.Google Scholar
Quinn, G.P. & Keough, M.J. (2002) Experimental Design and Data Analysis for Biologists. Cambridge, UK, Cambridge University Press.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
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. CrossRefGoogle Scholar
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.CrossRefGoogle Scholar
Vinson, S.B. (1972) Courtship behavior and evidence of a sex pheromone in the parasitoid Campoletis sonorensis (Hymenoptera: Ichneumonidae). Environmental Entomology 1, 409414.CrossRefGoogle Scholar
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.Google Scholar
Wong, B.B.M. & Candolin, U. (2005) How is female mate choice affected by male competition? Biological Reviews 80, 559571.CrossRefGoogle Scholar
Wyatt, T.D. (2003) Pheromones and Animal Behaviour: Communication by Smell and Taste. Cambridge, Cambridge University Press.CrossRefGoogle Scholar
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.CrossRefGoogle ScholarPubMed
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