Journal of Insect Behavior

, Volume 18, Issue 2, pp 171–186 | Cite as

Patch Time Allocation by the Parasitoid Diadegma semiclausum (Hymenoptera: Ichneumonidae). II. Effects of Host Density and Distribution

  • X. G. WangEmail author
  • M. A. Keller


This study investigated the effects of host density and distribution on the patch-leaving behavior of Diadegma semiclausum (Hymenoptera: Ichneumonidae), a solitary endoparasitoid of larval Plutella xylostella (Lepidoptera: Plutellidae). Individual female wasps were released onto an experimental plant infested with host larvae at different densities and distributions, and were allowed to freely leave for an alternative host plant placed upwind of the experimental plant in a wind tunnel. The influence of host density and distribution, as well as within-patch foraging experience, on the parasitoid’s patch-leaving tendency was analyzed by means of the proportional hazards model. This study aimed to test the predictions of a number of patch-leaving models, including the Marginal Value Theorem, “rules of thumb”, and incremental or countdown mechanisms. The parasitoid’s patch-leaving tendency decreased with increased host density, more clustered host distribution, and unsuccessful host encounter as a result of host defense, but increased with successful oviposition. None of the simple rules of thumb such as fixed time, fixed number of hosts parasitized, or fixed giving-up time was employed by this parasitoid. The results agreed with the general predictions of the Marginal Value Theorem that patch residence time and numbers of ovipositions by the parasitoid increased with increasing host density. The decreasing influence of oviposition on the parasitoid’s patch-leaving tendency, regardless of host density or distribution, was consistent with the prediction of a countdown mechanism.


Diadegma semiclausum foraging behavior host density and distribution parasitoids patch-leaving behavior patch time allocation proportional hazards model 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Allison, P. D. (1997). Survival Analysis Using the SAS System, A Practical Guide, SAS Institute Inc., Cary, NC, USA.Google Scholar
  2. Charnov, E. L. (1976). Optimal foraging, the Marginal Value Theorem. Theor. Popul. Biol. 9: 129–136.Google Scholar
  3. Collett, D. (1994). Modeling Survival Data in Medical Research, Chapman and Hall, London.Google Scholar
  4. Cox, D. R. (1972). Regression models and life tables. Biometrics 38: 67–77.Google Scholar
  5. Driessen, G., and Bernstein, C. (1999). Patch departure mechanisms and optimal host exploitation in an insect parasitoid. J. Anim. Ecol. 68: 445–459.Google Scholar
  6. Driessen, G., Bernstein, C., van Alphen, J. J. M., and Kacelnik, A. (1995). A countdown mechanism for host search in the parasitoid Venturia canescens. J. Anim. Ecol. 64: 117–125.Google Scholar
  7. Godfray, H. C. J. (1994). Parasitoids: Behavioural and Evolutionary Ecology, Princeton University Press, Princeton, NJ.Google Scholar
  8. Green, R. F. (1984). Stopping rules for optimal foragers. Am. Nat. 123: 30–43.Google Scholar
  9. Haccou, P., and Meelis, E. (1994). Statistical Analysis of Behavioural Data, Oxford University Press, New York.Google Scholar
  10. Haccou, P., De Vlas, S. J., van Alphen, J. J. M., and Visser, M. E. (1991). Information processing by foragers: Effects of intra-patch experience on the leaving tendency of Leptopilina heterotoma. J. Anim. Ecol. 60: 93–106.Google Scholar
  11. Hemerik, L., Driessen, G., and Haccou, P. (1993). Effects of intra-patch experiences on patch time, search time and searching efficiency of the parasitoid Leptopilina clavipes. J. Anim. Ecol. 62: 33–44.Google Scholar
  12. Iwasa, Y., Higashi, M., and Yamamura, N. (1981). Prey distribution as a factor determining the choice of optimal foraging strategy. Am. Nat. 117: 710–723.Google Scholar
  13. Keller, M. A. (1990). Responses of the parasitoid Cotesia rubecula to it s host Pieris rapae in a flight tunnel. Entomol. Exp. Appl. 57: 243–249.Google Scholar
  14. Keller, M. A., and Tenhumberg, B. (2000). New insight into the foraging behavior of parasitic wasps. In Austin, A. D., and Dowton, M. D. (eds.), Hymenoptera: Evolution, Biodiversity and Biological Control, CSIRO Publishing, Victoria, Australia, pp. 247-257.Google Scholar
  15. McNair, J. N. (1982). Optimal giving-up time and the marginal value theorem. Am. Nat. 119: 511–529.Google Scholar
  16. Nelson, J. M., and Roitberg, B. D. (1995). Flexible patch time allocation by the leafminer parasitoid, Opius dimidiatus. Ecol. Entomol. 20: 245–252.Google Scholar
  17. Noldus, L. P. J. J. (1991). The observer: A software system for collection and analysis of observational data. Behav. Res. Meth. Instru. Comp. 23: 415–429.Google Scholar
  18. Stephens, D. W., and Krebs, J. R. (1986). Foraging Theory, Princeton University Press, Princeton, New Jersey.Google Scholar
  19. Tenhumberg, B., Keller, M. A., Possingham, H. P., and Tyre, A. J. (2001). Optimal patch leaving behaviour: A case study using the parasitoid Cotesia rubecula. J. Anim. Ecol. 70: 683–691.Google Scholar
  20. van Alphen, J. J. M., and Vet, L. E. M. (1986). An evolutionary approach to host finding and selection. In Waage, J. K., and Greathead, D. (eds.), Insect Parasitoids, Academic Press, London, pp. 23–61.Google Scholar
  21. van Alphen, J. J. M., and Jervis, M. A. (1996). Foraging behaviour. In Jervis, M., and Kidd, N. (eds.), Insect Natural Enemies, Practical Approaches to Their Research and Evaluation, Chapman and Hall, UK, pp. 1–62.Google Scholar
  22. van Alphen, J. J. M., Bernstein, C., and Driessen G. (2003). Information acquisition and time allocation in insect parasitoids. Trend Ecol. Evol. 18: 81–87.Google Scholar
  23. van Roermund, H. J. W., Hemerik, L., and van Lenteren, J. C. (1994). Influence of intrapatch experiences and temperature on the time allocation of the whitefly parasitoid Encarsia formosa (Hymenoptera: Aphelinidae). J. Insect Behav. 7: 483–501.Google Scholar
  24. van Steenis, M. J., El-Khawass, K. A. M. H., Hemerik, L., and van Lenteren, J. C. (1996). Time allocation of the parasitoid Aphidius colemani (Hymenoptera: Aphidiidae) foraging for the Aphis gossypii (Homoptera: Aphididae) on cucumber leaves. J. Insect Behav. 9: 283–295.Google Scholar
  25. Vos, M., Hemerik, L., and Vet, L. E. M. (1998). Patch exploitation by the parasitoids Cotesia rubecula and Cotesia glomerata in multi-patch environments with different host distributions. J. Anim. Ecol. 67: 774–783.Google Scholar
  26. Waage, J. K. (1979). Foraging for patchily-distributed hosts by the parasitoid, Nemeritis canescens. J. Anim. Ecol. 48: 353–371.Google Scholar
  27. Wajnberg, E., Rosi, M. C., and Colazza, S. (1999). Genetic variation in patch time allocation in a parasitic wasp. J. Anim. Ecol. 68: 121–133.Google Scholar
  28. Wajnberg, E., Fauvergue, X., and Pons, O. (2000). Patch leaving decision rules and the Marginal Value Theorem: An experimental analysis and a simulation model. Behav. Ecol. 11: 577–586.Google Scholar
  29. Wajnberg, E., Gonsard, P. A., Tabone, E., Curty, C., Lezcano, N., and Colazza, S. (2003). A comparative analysis of patch-leaving decision rules in a parasitoid family. J. Anim. Ecol. 72: 618–626.Google Scholar
  30. Wang, X. G. (2001). Patch Exploitation by the Parasitoids of Plutella xylostella (L.): From Individual Behavior to Population Dynamics. Ph.D. Thesis, University of Adelaide, Australia.Google Scholar
  31. Wang, X. G., and Keller, M. A. (2002). A comparison of the host-searching efficiency of two larval parasitoids of Plutella xylostella. Ecol. Entomol. 27: 105–114.Google Scholar
  32. Wang, X. G., and Keller, M. A. (2003). Patch time allocation by the parasitoid Diadegma semiclausum (Hymenoptera: Ichneumonidae). I. Effect of inter-patch distance. J. Insect Behav. 16: 279–294.Google Scholar
  33. Wang, X. G., and Keller, M. A. (2004). Patch time allocation by the parasitoid Diadegma semiclausum (Hymenoptera: Ichneumonidae). III. Effects of kairomone sources and previous parasitism. J. Insect Behav. 17: 761–776.Google Scholar
  34. Wang, X. G., and Messing, R. H. (2003). Forging behavior and patch time allocation by Fopius arisanus, an egg-larval parasitoid of tephritid fruit flies. J. Insect Behav. 16: 593–612.Google Scholar

Copyright information

© Springer Science + Business Media, Inc. 2005

Authors and Affiliations

  1. 1.Plant and Pest Science, School of Agriculture and WineUniversity of AdelaideAustralia
  2. 2.University of Hawaii, College of Tropical Agriculture and Human ResourcesKapaaHawaii

Personalised recommendations