Advertisement

Journal of Insect Behavior

, Volume 10, Issue 6, pp 783–804 | Cite as

Swarming Behavior, Sexual Dimorphism, and Female Reproductive Status in the Sex Role-Reversed Dance Fly Species Rhamphomyia marginata

  • Bo G. Svensson
Article

Abstract

Dance flies are predaceous insects which often form male mating swarms. In many species males prior to swarming catch an insect prey, which is presented to the female at mating. In Rhamphomyia marginata, females in contrast to males gather to swarm, while males carrying a prey visit swarms for mating. Here I describe the swarming and courtship behavior in R. marginata and provide data on sexual dimorphism and swarming female reproductive status. Females swarm in small clearings in the forests. There was no specific swarm-maker. The swarming period lasted for 2–3 h and peaked around sunset. Identical swarm sites were used each evening and for several years. The mean number of females in swarms (swarm sites with at least one female) was 9.9 ± 9.1 (range, 1–40; n = 107) in 1993 and 7.1 ± 7.0 (range, 1–35; n = 68) in 1994. No obvious competition between females in swarms was observed. The operational sex ratio in swarms was extremely female biased (all swarms, 0.04). Less than one-third of male visits to swarms resulted in mating and males were found more often in larger swarms. Nuptial prey consisted of male midges. Females seem to mate more than once. Swarming females had undeveloped eggs, whereas mated females in swarms had further developed eggs than unmated females. Amount of sperm in the spermatheca was correlated with egg size. Amount of sperm and egg size did not correlate with wet weight, wing length, or wing load, except for egg size and weight. The wing coloration pattern and shape in R. marginata females are unique among dance flies, being greatly enlarged (1.6 times larger than that of males) and bicolored (gray part, 60% of wing area). When females, instead of males, possess extravagant secondary sexual characters, it is predicted from sexual selection theory that females should compete for males and that males should be selective in their choice of partner. A sex-role reversal will evolve when assess to males limit female reproductive success. The dance fly species R. marginata, like Empis borealis, another dance fly species studied earlier and discussed here, seems to fit these predictions.

swarm sex-role reversal nuptial gift dance fly Rhamphomyia marginata Empididae 

Preview

Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.

REFERENCES

  1. Andersen, T., and Fjellberg, A. (1976). New records of Rhamphomyia marginata Fabr. (Dipt., Empididae) from East and West Norway. Norw. J. Entomol. 23: 205–206.Google Scholar
  2. Andersson, M. (1994). Sexual Selection, Princeton University Press, Princeton, NJ.Google Scholar
  3. Andersson, H., and Iwasa, Y. (1996). Sexual selection. Trends Ecol. Evol. 11: 53–58.Google Scholar
  4. Barták, M. (1982). The Czechoslovak species of Rhamphomyia (Diptera, Empididae), with descriptions of a new species from central Europe. Acta Univ. Carol. Biol. 1980(1982): 381–461.Google Scholar
  5. Bateson, P. (1983). Mate Choice, Cambridge University Press, Cambridge.Google Scholar
  6. Bayfield, N. G. (1994). Effects of prey abundance on swarming Empis borealis L. (Diptera: Empididae) in north-east Scotland. Entomol. Gazette 45: 59–67.Google Scholar
  7. Björklund, M. (1990). A phylogenetic interpretation of sexual dimorphism in body size and ornament in relation to mating system in birds. J. Evol. Biol. 3: 171–183.Google Scholar
  8. Blum, M. S., and Blum, N. A. (1979). Sexual Selection and Reproductive Competition in Insects, Academic Press, New York.Google Scholar
  9. Bradbury, J. W., and Andersson, M. B. (1987). Sexual Selection: Testing the Alternatives, Wiley, Chichester.Google Scholar
  10. Chvála, M. (1976). Swarming, mating and feeding habits in Empididae (Diptera), and their significance in evolution of the family. Acta Entomol. Bohemoslov. 73: 353–366.Google Scholar
  11. Chvála, M. (1983). The Empidoidea (Diptera) of Fennoscandia and Denmark. II. General Part. The Families Hybotidae, Atelestidae and Microphoridae. Fauna Entomol. Scand. 12: 1–279.Google Scholar
  12. Chvála, M. (1994). The Empidoidea of Fennoscandia and Denmark. III. Genus Empis. Fauna Entomol. Scand. 29: 1–187.Google Scholar
  13. Chvála, M., and Wagner, R. (1989). Family Empididae. In Soós, A., and Papp, L. (eds.), Catalogue of Palearctic Diptera, Vol. 6. Therevidae-Empididae, Akadémiai Kiadó, Budapest, pp. 228–336.Google Scholar
  14. Cumming, J. M. (1994). Sexual selection and the evolution of dance fly mating systems (Diptera: Empididae; Empidinae). Can. Entomol. 126: 907–920.Google Scholar
  15. Darwin, C. (1871). The Descent of Man, and Sexual Selection in Relation to Sex, Murry, London.Google Scholar
  16. Downes, J. A. (1969). The swarming and mating flight of Diptera. Annu. Rev. Entomol. 14: 271–298.Google Scholar
  17. Downes, J. A. (1970). The feeding and mating behavior of the specialized Empidinae (Diptera); Observations on four species of Rhamphomyia in the high artic and general discussion. Can. Entomol. 102: 769–791.Google Scholar
  18. Frey, R. (1956). Empididae. In Linden, E. (ed.), Die Fliegen der Palaearktischen Region, IV(4), Schweizerbart, Stuttgart, pp. 400–639.Google Scholar
  19. Funk, D. (1992). Sex, flies and videotape. In Gelhaus, J. K. (ed.), Society Meeting of October 23, 1991. Entomol. News 103: 59–64.Google Scholar
  20. Gruhl, K. (1924). Paarungsgewohnheiten der Dipteren. Z. Wiss. Zool. 122: 205–280.Google Scholar
  21. Gruhl, K. (1955). Neue Beobachtungen an Schwarm-und Tanzgesellschaften der Dipteren. Dt. Entomol. Z. (N.F.) 2: 332–353.Google Scholar
  22. Gruhl, K. (1963). Sind Tänze von Weibchen bei Empis und Rhamphomya (Diptera) an gewisse Sondermerkmale gebunden? Mitt. Deut. Entomol. Ges. 22: 88–96.Google Scholar
  23. Gwynne, D. T. (1988). Courtship feeding and the fitness of female katydids (Orthoptera: Tettigoniidae). Evolution 42: 545–555.Google Scholar
  24. Gwynne, D. T. (1991). Sexual competition among females: what causes courtship-role reversal? Trends Ecol. Evol. 6: 118–121.Google Scholar
  25. Gwynne, D. T., and Simmons, L. W. (1990). Experimental reversal of courtship in an insect. Nature 346: 172–174.Google Scholar
  26. Hackman, W. (1968). Rhamphomyia (Pararhamphomyia) marginata Fabricius funnen i Finland. Notulae Entomol. 48: 90.Google Scholar
  27. Hanks, L. M., Millar, J. G., and Paine, T. D. (1996). Mating behavior of the eucalyptus longhorned borer (Coleoptera: Cerambycidae) and the adaptive significance of long “horns.” J. Insect Behav. 9: 383–393.Google Scholar
  28. Hunter, F. F., Sutcliffe, J. F., and Stratton, C. (1992). Subcostal incomplete, A new genetic mutant of Stomoxys calcitrans L. (Diptera: Muscidae). J. Hered. 83: 453–455.Google Scholar
  29. Höglund, J., and Alatalo, R. V. (1995). Leks, Princeton University Press, Princeton, NJ.Google Scholar
  30. Janetos, A. C. (1980) Strategies of female choice: A theoretical analysis. Behav. Ecol. Sociobiol. 7: 107–112.Google Scholar
  31. Kessel, E. L. (1955). The mating activities of balloon flies. Syst. Zool. 4: 97–105.Google Scholar
  32. Krebs, J. R., and Davies, N. B. (1993). An Introduction to Behavioural Ecology, Blackwell Scientific, Oxford.Google Scholar
  33. Kruse, K. C. (1990). Male backspace availability in the giant waterbug (Belostoma flumineum Say). Behav. Ecol. Sociobiol. 26: 281–289.Google Scholar
  34. Kvarnemo, C., and Ahnesjö, I. (1996). The dynamics of operational sex ratios and competition for mates. Trends Ecol. Evol. 11: 404–408.Google Scholar
  35. Lack, D. (1968). Ecological Adaptations for Breeding in Birds, Chapman and Hall, London.Google Scholar
  36. Lundbeck, W. (1910). Empididae. In Diptera Danica, Genera and Species of Flies Hitherto Found in Denmark, Vol. 3, Gad, Copenhagen.Google Scholar
  37. Marden, J. (1987). Maximum lift production during takeoff in flying animals. J. Exp. Biol. 130: 235–258.Google Scholar
  38. Newkirk, M. R. (1970). Biology of the long-tailed dance fly, Rhamphomyia longicauda (Diptera: Empididae); A new look at swarming. Ann. Entomol. Soc. Am. 63: 1407–1412.Google Scholar
  39. Opheim, M. (1973). Rhamphomyia platyptera Panz. (Dipt., Empididae) ny for Norge. Norw. J. Entomol. 20: 334.Google Scholar
  40. Poulton, E. B. (1913). Empididae and their prey in relation to courtship. Entomol. Month. Mag. 49: 177–180.Google Scholar
  41. Reynolds, J. D. (1996). Animal breeding systems. Trends Ecol. Evol. 11: 68–72.Google Scholar
  42. Richards, O. W. (1927). Sexual selection and allied problems in insects. Biol. Rev. 11: 298–364.Google Scholar
  43. Selander, R. K. (1972). Sexual selection and dimorphism in birds. In Campbell, B. (ed.), Sexual Selection and the Descent of Man 1871–1971, Aldine, Chicago, pp. 180–230.Google Scholar
  44. Simmons, L. W. (1995). Relative parental expenditure, potential reproductive rates, and the control of sexual selection in katydids. Am. Nat. 145: 797–808.Google Scholar
  45. Sing, B. N., and Chatterjee, S. (1987). Greater mating success of Drosophila biarmipes males possessing an apical dark black wing patch. Ethology 75: 81–83.Google Scholar
  46. Sivinski, J. (1988). Unusual female-aggregated mating systems in phorid flies. J. Insect Behav. 1: 123–128.Google Scholar
  47. Smith, K. G. V. (1969). The Empididae of Southern Africa (Diptera). Ann. Natal Mus. 19: 1–347.Google Scholar
  48. Smith, R. L. (1979). Paternity assurance and altered roles in the mating behavior of a giant water bug, Abedus herberti (Heteroptera: Belostomatidae). Anim. Behav. 27: 716–725.Google Scholar
  49. Sullivan, R. T. (1981). Insect swarming and mating. Fla. Entomol. 64: 44–65.Google Scholar
  50. Svensson, B. G., and Petersson, E. (1987). Sex-role reversed courtship behavior, sexual dimorphism and nuptial gifts in the dance fly Empis borealis (L.) Ann. Zool. Fenn. 24: 323–334.Google Scholar
  51. Svensson, B. G., and Petersson, E. (1988). Non-random mating in the dance fly Empis borealis: The importance of male choice. Ethology 79: 307–316.Google Scholar
  52. Svensson, B. G., and Petersson, E. (1992). Why insects swarm: Testing the models for lek mating systems on swarming Empis borealis females. Behav. Ecol. Sociobiol. 31: 253–261.Google Scholar
  53. Svensson, B. G., and Petersson, E. (1994). Mate choice tactics and swarm size: A model and a test in a dance fly. Behav. Ecol. Sociobiol. 35: 161–168.Google Scholar
  54. Svensson, B. G., and Petersson, E. (1995). Diurnal and seasonal variations in swarming and mating behavior of the dance fly Empis borealis (Diptera; Empididae). Ann. Zool. Fenn. 32: 403–409.Google Scholar
  55. Svensson, B. G., Petersson, E., and Forsgren, E. (1989). Why do males of the dance fly Empis borealis refuse to mate? The importance of female age and size. J. Insect Behav. 2: 387–395.Google Scholar
  56. Svensson, B. G., Petersson, E., and Frisk, M. (1990). Nuptial gift size prolongs copulation duration in the dance fly Empis borealis. Ecol. Entomol. 15: 225–229.Google Scholar
  57. Thornhill, R., and Alcock, J. (1983). The Evolution of Insect Mating Systems, Harvard University Press, Cambridge, MA.Google Scholar
  58. Trivers, R. L. (1972). Parental investment and sexual selection. In Campell, B. G. (ed.), Sexual Selection and the Descent of Man, Heinemann, London, pp. 136–179.Google Scholar
  59. Tuomikoski, R. (1939). Beobachtungen über das Schwärmen und die Kopulationen einiger Empididen (Dipt.). Ann. Entomol. Fenn. 5: 1–30.Google Scholar
  60. van der Goot, V. S., and de Vos, R. (1988). Zwermen van Rhamphomyia marginata in de schemering (Diptera: Empididae). Entomol. Ber. Amst. 48: 49–52.Google Scholar
  61. Väsinen, R. (1982). Vanishing and vulnerable Diptera in Finland. Notulae Entomol. 62: 111–121.Google Scholar
  62. Wahlgren, E. (1910). Svensk Insektsfauna. Diptera 1. Första underordningen Orthorapha. Andra gruppen flugor Brachycera. 24. Fam. Dansflugor. Empididae. Entomol. Tidskr. 11: 41–95.Google Scholar

Copyright information

© Plenum Publishing Corporation 1997

Authors and Affiliations

  • Bo G. Svensson
    • 1
  1. 1.Department of ZoologyUppsala UniversityUppsalaSweden

Personalised recommendations