Behavioral Ecology and Sociobiology

, Volume 26, Issue 3, pp 181–190 | Cite as

The effects of nesting stage, sex, and type of predator on parental defense by killdeer (Charadrius vociferous): testing models of avian parental defense

  • Dianne H. Brunton


Two models predicting the temporal patterns of parental investment in offspring defense over the nesting cycle were tested. The first is based on offspring age, the other on the vulnerability of offspring to predation. Both models make very similar predictions for altricial species after eggs have hatched, i.e., increases in intensity of parental defense until fledging. For precocial species, however, the post-hatching predictions of each model are different: the offspring age model predicts a continued increase in defense intensity, while the vulnerability model predicts a decline. I examined the temporal patterns of parental defense of a precocial shorebird, the killdeer (Charadrius vociferus), and determined which model was supported. Killdeer responses to human and natural predators were observed. Killdeer were less willing to leave the nest, responded most intensely, and displayed closest to a potential predator around hatching. Defense intensity increased from early to late incubation as predicted by the offspring age model. However, after hatching killdeer parental defense declined for both males and females, thus supporting the vulnerability model for this stage. Males and females responded significantly differently to all types of predators. Males took greater risks, remained on the nest longer, defended offspring more intensely, and displayed closer to the predator than females at the approach of a potential predator. Responses to natural predators depended on the type of predator and the approach made by the predator; a greater range of defense behavior was used for predators approaching on the ground compared to aerial predators. In general, killdeer responses to humans were more intense and less variable than their responses to natural predators. This was most likely because the human intruder approached nests and chicks more directly and closer than natural predators.


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Andersson M, Wiklund CG, Rundgren H (1980) Parental defense of offspring: a model and an example. Anim Behav 28:536–542Google Scholar
  2. Armstrong EA (1956) Distraction display and the human predator. Ibis 98:641–654Google Scholar
  3. Ashkenazie S, Safriel UN (1979) Time-energy budget of the Semipalmated Sandpiper (Calidris pusilla) at Barrow, Alaska. Ecology 60:783–799Google Scholar
  4. Barash DP (1975) Evolutionary aspects of parental behavior: distraction display of the alpine accentor. Wilson Bull 87:367–373Google Scholar
  5. Bengtson SA (1970) Breeding behaviour of the Purple Sandpiper Calidris maritima in West Spitsbergen. Ornis Scand 1:17–25Google Scholar
  6. Bent AC (1929) Life histories of North American shorebirds. Part 2, US Natl Mus Bull 142Google Scholar
  7. Biermann GC, Robertson RJ (1981) An increase in parental investment during the breeding season. Anim Behav 29:487–489Google Scholar
  8. Brown RG (1962) The aggressive and distraction display behavior of the Western Sandpiper Ereunetes mauri. Ibis 104:1–12Google Scholar
  9. Brunton DH (1986) Fatal antipredator behavior by Killdeer. Wilson Bull 98(4):605–607Google Scholar
  10. Brunton DH (1987) Reproductive effort of male and female Killdeer (Charadrius vociferus). PhD dissertation University Michigan, Ann Arbor, MichiganGoogle Scholar
  11. Brunton DH (1988a) Sexual differences in time budgets of Killdeer during the breeding season. Anim Behav 36:705–717Google Scholar
  12. Brunton DH (1988b) Energy expenditure in reproductive effort: the reproductive strategies of male and female Killdeer (Charadrius vociferus). Auk 105:553–564Google Scholar
  13. Buitron D (1983) Variability in the responses of Black-billed Magpies to natural predators. Behaviour 78:209–236Google Scholar
  14. Burger J (1981) Sexual differences in parental activities of breeding Black Skimmers. Am Nat 117:975–984Google Scholar
  15. Curio E (1975) The functional organization of antipredator behaviour in the Pied Flycatcher: a study of avian visual perception. Anim Behav 23:1–115Google Scholar
  16. Curio E, Regelmann K, Zimmerman U (1984) The defence of first and second broods by Great Tit (Parus major) parents: a test of predictive sociobiology. Z Tierpsychol 66:101–127Google Scholar
  17. East M (1981) Alarm calling and parental investment in the Robin, Erithacus rubecula. Ibis 123(2):223–230Google Scholar
  18. Erpino MJ (1968) Nest-related activities of Black-billed Magpies. Condor 70:154–165Google Scholar
  19. Gibson F (1971) The breeding biology of the American Avocet Recurvirostra americana in central Oregon. Condor 73:444–454Google Scholar
  20. Gochfeld M (1984) Antipredator behavior: aggressive and distraction displays of shorebirds. In: Burger J, Olla BL (eds) Shorebirds: breeding behavior and populations. Plenum Publ. Corp., New York, pp 289–377Google Scholar
  21. Gottfried BM (1979) Anti-predator aggression in birds nesting in old field habitats: an experimental analysis. Condor 81:251–257Google Scholar
  22. Gramza A (1967) Responses of brooding Nighthawks to a disturbance stimulus. Auk 84:72–86Google Scholar
  23. Greig-Smith PW (1980) Parental investment in nest defense by Stonechats (Saxicola torquata). Anim Behav 28:604–619Google Scholar
  24. Harvey PH, Greenwood PJ (1978) Antipredator defence by Stonechats (Saxicola torquata). Anim Behav 28:604–619Google Scholar
  25. Hobbs JN (1972) Breeding of Red-capped Dotterel at Fletcher's Lake Dareton, NSW. Emu 72:121–125Google Scholar
  26. Howe MA (1982) Social organization in a nesting population of eastern Willets (Catoptrophorus semipalmatus). Condor 84:88–102Google Scholar
  27. Hussell D, Page GW (1976) Observations on the breeding biology of the Black-bellied Plovers on Devon Island, N.W.T., Canada. Wilson Bull 88:632–653Google Scholar
  28. Jones A (1979) Notes on the behaviour of the Variable Oystercatcher. Notornis 26:47–52Google Scholar
  29. Jourdain FC (1936) The so-called injury feigning in birds. Oologists Rec 16:25–37Google Scholar
  30. Komeda S (1983) Nest attendance of parent birds in the Painted Snipe. Auk 100:48–55Google Scholar
  31. Kruuk H (1964) Predators and anti-predator behavior of the Black-headed Gull (Larus ridibundus L.). Behav Suppl 11:1–129Google Scholar
  32. Larson S (1960) On the influence of the Arctic fox (Alopex lagopus) on the distribution of Arctic birds. Oikos 11:277–305Google Scholar
  33. Lazarus J, Inglis IR (1986) Shared and unshared parental investment, parent-offspring conflict and brood size. Anim Behav 34:1791–1804Google Scholar
  34. Lemmetyinen R (1971) Nest defence behaviour of Common and Arctic Terns and its effects on the success achieved by predators. Ornis Fenn 48:13–24Google Scholar
  35. Lenington S (1980) Bi-parental care in Killdeer: an adaptive hypothesis. Wilson Bull 92(1):8–20Google Scholar
  36. Maclean GL (1973) A review of the biology of the Australian Desert Waders, Stilta and Peltohyas. Emu 73:61–70Google Scholar
  37. Maxson SJ, Oring LW (1980) Breeding season time and energy budgets of the polyandrous Spotted Sandpiper. Behaviour 74:200–263Google Scholar
  38. Merritt PG (1984) Observer recognition by the Northern Mockingbird. J Field Ornithol 55:252–253Google Scholar
  39. Montevecchi W, Porter J (1980) Parental care and related behavior before fledging in Northern Gannets (Morus bassanus) with reference to other marine birds. In: Burger J, Olla B, Winn H (eds) Behavior of marine animals. Vol. VI. Marine birds. Plenum Press, New York, pp 323–366Google Scholar
  40. Montgomerie RD, Weatherhead PJ (1988) Risks and rewards of nest defence by parental birds. Q Rev Biol 63:167–187Google Scholar
  41. Mundahl JT (1982) Role specialization in the parental and territorial behavior of the Killdeer. Wilson Bull 94(4):515–530Google Scholar
  42. Myers JP (1978) One deleterious effect of mobbing in the Southern Lapwing (Vanellus chilensis). Auk 95:419Google Scholar
  43. Nol E (1980) Factors affecting the nesting success of the Killdeer (Charadrius vociferus) on Long Point, Ontario. M.S. thesis, University Guelph, Guelph, OntarioGoogle Scholar
  44. Nol E, Lambert A (1984) Comparison of Killdeer, Charadrius vociferus, breeding in Mainland and peninsula sites in Southern Ontario. Can Field Nat 98:7–11Google Scholar
  45. Patterson TL, Petrinovich L, James DK (1980) Reproductive value and appropriateness of response to predators by White-crowned Sparrows. Behav Ecol Sociobiol 7:227–231Google Scholar
  46. Phillips RE (1972) Sexual and agonistic behavior in the Killdeer (Charadrius vociferus). Anim Behav 20:1–9Google Scholar
  47. Pierotti R (1981) Male and female parental roles in the Western Gull under different environmental conditions. Auk 98:532–549Google Scholar
  48. Ratcliffe DA (1977) Observations on the breeding of the Golden Plover in Great Britain. Bird Study 23:63–116Google Scholar
  49. Regelmann K, Curio E (1983) Determinants of brood defence in the Great Tit Parus major L. Behav Ecol Sociobiol 13:131–145Google Scholar
  50. Shields WM (1984) Barn Swallow mobbing: self-defence, collateral kin defence, group defence, or parental care? Anim Behav 32:132–148Google Scholar
  51. Simmons KEL (1955) The nature of the predator reactions of waders towards humans, with special reference to the role of the aggressive-, escape- and brooding-drives. Behaviour 8:130–173Google Scholar
  52. Skutch AF (1949) Do tropical birds rear as many young as they can nourish? Ibis 91:430–455Google Scholar
  53. Trivers RL (1972) Parental investment and sexual selection. In: Campbell B (ed) Sexual selection and the descent of man, 1871–1971. Aldine Press, Chicago, pp 136–179Google Scholar
  54. Vogt W (1938) Preliminary notes on the behavior and ecology of the eastern Willet. Proc Linn Soc NY 49:8–42Google Scholar
  55. Walters JR (1982) Parental Behavior in Lapwings (Charadriidae) and its relationships with clutch sizes and mating systems. Evolution 36:1030–1040Google Scholar
  56. Weatherhead PJ (1979) Do Savannah Sparrows commit the Concorde Fallacy? Behav Ecol Sociobiol 5:373–381Google Scholar
  57. Weatherhead PJ (1982) Risk-taking by red-winged blackbirds and the Concorde Fallacy. Z Tierpsychol 60:199–208Google Scholar

Copyright information

© Springer-Verlag 1990

Authors and Affiliations

  • Dianne H. Brunton
    • 1
  1. 1.School of Natural ResourcesUniversity of MichiganAnn ArborUSA

Personalised recommendations