Behavioral Ecology and Sociobiology

, Volume 34, Issue 3, pp 217–223 | Cite as

Song as a pursuit-deterrent signal, and its occurrence relative to other anti-predation behaviours of skylark (Alauda arvensis) on attack by merlins (Falco columbarius)

  • Will Cresswell


Skylarks show a range of anti-predation behaviours including flocking, refuge-seeking and song. Responses by skylarks to merlin attack were recorded over three winters on a Scottish estuary to determine the effectiveness of song as a pursuit-deterrent signal, and its use with respect to other anti-predation options such as flocking. Mortality due to merlin predation was high. Skylarks used song as a pursuit-deterrent signal. Merlins chased non- or poorly singing skylarks for longer periods compared to skylarks that sang well. A merlin was more likely to catch a non-singing than a poorly singing than a full-singing skylark. Temperature did not affect chase lengths, song types or success rates of attacks. Larger flocks of skylarks were preferentially attacked so that the individual probability of being attacked within some larger flock sizes was greater than in a smaller flock. Success rate did not vary with flock size. Merlins chose skylarks before any song was heard, so there was no cost for non-singing skylarks in joining flocks. The frequency of capture on merlin attack depended on the escape response used by the skylark. Non-singing skylarks were probably more likely to escape by seeking a refuge or staying on the ground on attack, while singing skylarks were more likely to escape if they flew. The optimal escape option available to a skylark on merlin attack was probably dependent on its condition, as indicated by its ability to sing on attack.

Key words

Skylark Merlin Pursuit-deterrence Antipredation 


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  1. Baker RR, Parker GA (1979) The evolution of bird colouration. Phil Trans R Soc London B 287:63–130Google Scholar
  2. Bauwens D, Thoren C (1981) Escape tactics and vulnerability to predation associated with reproduction in the lizard (Lacerta vivipara) J Anim Ecol 50:733–743Google Scholar
  3. Brown LH (1976) British birds of prey. Collins, LondonGoogle Scholar
  4. Buchanan JB, Schick CT, Brennan LA, Herman SG (1988) Merlin predation on wintering dunlins: hunting success and dunlin escape tactics. Wilson Bull 100:108–118Google Scholar
  5. Cramp S (1988) The birds of the Western Polearctie, Vol. 5. Oxford: Oxford University PressGoogle Scholar
  6. Cramp S, Simmons KEL (1980) The birds of the Western Polearctie, Vol. 2. Oxford: Oxford University PressGoogle Scholar
  7. Cresswell W (1993) Escape responses by redshank (Tringa totanus) on attack by avian predators. Anim Behav 46:609–611Google Scholar
  8. Cresswell W (1994) Flocking is an effective anti-predation strategy in redshank (Tringa totanus). Anim Behav in pressGoogle Scholar
  9. Cresswell W, Whitfield DP (in press) The effects of raptor predation on wintering wader populations at the Tyninghame estuary south-east Scotland. IbisGoogle Scholar
  10. Dekker D (1988) Peregrine falcon and merlin predation on small shore-birds and passerines in Alberta. Can J Zool 66:925–928Google Scholar
  11. Fitzgibbon CD, Fanshawe JH (1988) Stotting in Thomson's gazelles: an honest signal of condition. Behav Ecol Sociobiol 23:69–74Google Scholar
  12. Foster WA, Treherne JE (1981) Evidence for the dilution effect in the selfish herd from fish predation on a marine insect. Nature 295:466–467Google Scholar
  13. Grafen A (1991) Modelling in behavioural ecology In: Krebs JR, Davies NB (eds) Behavioural ecology: an evolutionary approach. Blackwell Scientific, OxfordGoogle Scholar
  14. Hamilton WD (1971) Geometry for the selfish herd. J Theor Biol 31:295–311Google Scholar
  15. Harper DGC (1991) Communication. In: Krebs JR, Davies NB (eds) In: Behavioural ecology: an evolutionary approach. Blackwell Scientific, OxfordGoogle Scholar
  16. Hasson O (1991) Pursuit-deterrent signals: communication between prey and predator. Trends Ecol Evol 6:325–329Google Scholar
  17. Krebs JR, Davies NB (eds) (1987) An introduction to behavioural ecology, 2nd edn. Blackwell Scientific, OxfordGoogle Scholar
  18. Lima SL, Dill LM (1990) Behavioural decisions made under the risk of predation: a review and prospectus. Can J Zool 68:619–640Google Scholar
  19. Norusis MJ (1990) SPSS/PC+ advanced statistics 4.0. SPSS Inc, GorinchemGoogle Scholar
  20. Page G, Whitacre DF (1975) Raptor predation on wintering shorebirds. Condor 77:73–83Google Scholar
  21. Rice WR (1989) Analyzing tables of statistical tests. Evolution 43:223–225Google Scholar
  22. Ritchison G (1983) The function of singing in female black-headed grosbeaks (Pheucticus melanocephalus): family group maintenance. Auk 100:105–116Google Scholar
  23. Rudebeck G (1950) The choice of prey and modes of hunting of predatory birds with special reference to their selective effect Oikos 2:65–88Google Scholar
  24. Rudebeck G (1951) The choice of prey and modes of hunting of predatory birds with special reference to their selective effect. Oikos 3:200–231Google Scholar
  25. Simms E (1992) British larks, pipits and wagtails. HarperCollins, LondonGoogle Scholar
  26. Whitfield DP (1985) Raptor predation on wintering waders in southeast Scotland. Ibis 127:554–558Google Scholar
  27. Woodland DJ, Jaafer Z, Knight ML (1980) The “pursuit deterrent” function of alarm signals. Am Nat 115:748–753Google Scholar
  28. Ydenberg RC, Dill LM (1986) The economics of fleeing from predators. Adv Study Behav 16:229–249Google Scholar
  29. Zahavi A (1975) Mate selection — a selection for a handicap. J Theor Biol 53:205–214Google Scholar

Copyright information

© Springer-Verlag 1994

Authors and Affiliations

  • Will Cresswell
    • 1
  1. 1.Institute of Cell, Animal, and Population BiologyEdinburgh UniversityEdinburghUK

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