, Volume 159, Issue 4, pp 893–901 | Cite as

Nest construction by a ground-nesting bird represents a potential trade-off between egg crypticity and thermoregulation

  • Paul M. Mayer
  • Levica M. Smith
  • Robert G. Ford
  • Dustin C. Watterson
  • Marshall D. McCutchen
  • Mark R. Ryan
Behavioral Ecology - Original Paper


Predation selects against conspicuous colors in bird eggs and nests, while thermoregulatory constraints select for nest-building behavior that regulates incubation temperatures. We present results that suggest a trade-off between nest crypticity and thermoregulation of eggs based on selection of nest materials by piping plovers (Charadrius melodus), a ground-nesting bird that constructs simple, pebble-lined nests highly vulnerable to predators and exposed to temperature extremes. Piping plovers selected pebbles that were whiter and appeared closer in color to eggs than randomly available pebbles, suggesting a crypsis function. However, nests that were more contrasting in color to surrounding substrates were at greater risk of predation, suggesting an alternate strategy driving selection of white rocks. Near-infrared reflectance of nest pebbles was higher than randomly available pebbles, indicating a direct physical mechanism for heat control through pebble selection. Artificial nests constructed of randomly available pebbles heated more quickly and conferred heat to model eggs, causing eggs to heat more rapidly than in nests constructed from piping plover nest pebbles. Thermal models and field data indicated that temperatures inside nests may remain up to 2–6°C cooler than surrounding substrates. Thermal models indicated that nests heat especially rapidly if not incubated, suggesting that nest construction behavior may serve to keep eggs cooler during the unattended laying period. Thus, pebble selection suggests a potential trade-off between maximizing heat reflectance to improve egg microclimate and minimizing conspicuous contrast of nests with the surrounding substrate to conceal eggs from predators. Nest construction behavior that employs light-colored, thermally reflective materials may represent an evolutionary response by birds and other egg-laying organisms to egg predation and heat stress.


Anti-predation Charadrius melodus Egg crypsis Nest construction Thermoregulation 



We are grateful for constructive remarks by K. Forshay, C. Lehman, A. Person, R. Phillips, B. Root, G. Schnell, D. Siems, A. Strong, B. Woodworth, and two astute anonymous reviewers. We thank B. Root for assistance in the field. A. Person and J. Dean provided piping plover eggs from the Sam Noble Oklahoma Museum of Natural History and the Smithsonian Institution/National Museum of Natural History, respectively. The US Environmental Protection Agency (Agency) through its Office of Research and Development partially funded and collaborated in the research described here under cooperative agreement (CR832885) with East Central University. The research described here has not been subjected to Agency review and therefore does not necessarily reflect the views of the Agency, and no official endorsement should be inferred. Mention of trade names or commercial products does not constitute endorsement or recommendation for use. This research adhered to the Association for the Study of Animal Behavior/Animal Behavior Society Guidelines for the Use of Animals in Research, the legal requirements of the country in which the work was carried out, and all institutional guidelines.


  1. Amat JA, Fraga RM, Arroyo GM (1999) Reuse of nesting scrapes by Kentish plovers. Condor 101:157–159CrossRefGoogle Scholar
  2. Ankney CD, Hopkins J (1985) Habitat selection by roof-nesting killdeer. J Field Ornithol 56:284–286Google Scholar
  3. Bakken GS, Vanderbilt VC, Buttemer WA, Dawson WR (1978) Avian eggs: thermoregulatory value of very high near-infrared reflectance. Science 200:321–323PubMedCrossRefGoogle Scholar
  4. Bent AC (1929) Life histories of North American shorebirds. Part II. U.S. Natl Mus Bull 146Google Scholar
  5. Beissinger SR, Cook MI, Arendt WJ (2005) The shelf life of bird eggs: testing egg viability using a tropical climate gradient. Ecology 86:2164–2175CrossRefGoogle Scholar
  6. Bergstrom PW (1989) Incubation temperatures of Wilson’s plovers and killdeers. Condor 91:634–641CrossRefGoogle Scholar
  7. Blanco G, Bertellotti M (2002) Differential predation by mammals and birds: implications for egg-colour polymorphism in a nomadic breeding seabird. Biol J Linn Soc 75:137–146CrossRefGoogle Scholar
  8. Brua RB (2002) Parent-embryo interactions. In: Deeming DC (ed) Avian incubation: behaviour, environment, and evolution. Oxford ornithological series. Oxford University Press, Oxford, pp 88–99Google Scholar
  9. Bunni MK (1959) Killdeer, Charadrius vociferus Linnaeus, in the breeding season: ecology, behavior, and the development of homiothermism. Sc.D. thesis, University of Michigan, Ann ArborGoogle Scholar
  10. Cairns WE (1982) Biology and behavior of breeding piping plovers. Wilson Bull 94:531–545Google Scholar
  11. Caro T (2005) The adaptive significance of coloration in mammals. Bioscience 55:125–136CrossRefGoogle Scholar
  12. Castilla AM, Dhondt AA, Díaz-Uriarte R, Westmoreland D (2007) Predation in ground nesting birds: an experimental study using natural egg-color variation. Avian Conserv Ecol 2(1):2. http://www.ace-eco.org/vol2/iss1/art2/ Google Scholar
  13. Clark RN (1999) Spectroscopy of rocks and minerals and principles of spectroscopy. In: Rencz AN (ed) Manual of remote sensing, vol 3. Remote sensing for the earth sciences. Wiley, New York, pp 3–58Google Scholar
  14. Collias NE (1997) On the origin and evolution of nest building by passerine birds. Condor 99:253–270CrossRefGoogle Scholar
  15. Collias NE, Collias EC (1984) Nest building and bird behaviour. Princeton University Press, PrincetonGoogle Scholar
  16. Cooper CB, Hochachka WM, Butcher G, Dhondt AA (2005) Seasonal and latitudinal trends in clutch size: thermal constraints during laying and incubation. Ecology 86:2018–2031CrossRefGoogle Scholar
  17. Cott HB (1940) Adaptive coloration in animals. Methuen, LondonGoogle Scholar
  18. Courtney JC, Martin TE (2000a) Effects of ambient temperature on avian incubation behavior. Behav Ecol 11:178–188CrossRefGoogle Scholar
  19. Courtney JC, Martin TE (2000b) Evolution of passerine incubation behavior: influence of food, temperature, and nest predation. Evolution 54:670–685Google Scholar
  20. Darwin C (1859) The origin of species. Murray, LondonGoogle Scholar
  21. Diamond J (1982) Evolution of bowerbirds’ bowers: animal origins of the aesthetic sense. Nature 297:99–102CrossRefGoogle Scholar
  22. Donnelly WA, Whoriskey FG Jr (1991) Background-color acclimation of brook trout for crypsis reduces risk of predation by hooded mergansers Lophodytes cucullatus. N Am J Fish Manage 11:206–211CrossRefGoogle Scholar
  23. Fitzpatrick LC, Guerra CG (1988) Microclimatic features of gray gull, Larus modestus, nests in the Atacama Desert. Gerfaut 78:421–428Google Scholar
  24. Flemming SP, Chiasson RD, Austin-Smith PJ (1992) Piping plover nest site selection in New Brunswick and Nova Scotia. J Wildl Manage 56:578–583CrossRefGoogle Scholar
  25. Franklin DC (1995) Helmeted honeyeaters build bulker nests in cold weather. Auk 112:247–248Google Scholar
  26. Götmark F (1992) Blue eggs do not reduce nest predation in the song thrush, Turdus philomelos. Behav Ecol Sociobiol 30:245–252CrossRefGoogle Scholar
  27. Gore JA, Kinnison MJ (1991) Hatching success in roof and ground colonies of least terns. Condor 93:759–762CrossRefGoogle Scholar
  28. Grant GS (1982) Avian incubation: egg temperature, nest humidity, and behavioral thermoregulation in a hot environment. Ornithological monographs no. 30. American Ornithologists Union, WashingtonGoogle Scholar
  29. Graul WD (1973) Possible function of head and breast markings in Charadriinae. Wilson Bull 85:60–70Google Scholar
  30. Haig SM (1992) Piping plover: In: Poole A, Stettenheim P, Gill F (eds) The birds of North America, no. 2. The Academy of Natural Science, The American Ornithologists’ Union, PhiladelphiaGoogle Scholar
  31. Haig SM, Ferland CL, Amirault D, Cuthbert F, Dingledine J, Goossen P, Hecht A, McPhillips N (2005) The importance of complete species censuses and evidence for regional declines in piping plovers. J Wildl Manage 69:160–173CrossRefGoogle Scholar
  32. Hansell M (2000) Bird nests and construction behavior. Cambridge University Press, LondonGoogle Scholar
  33. Haskell DG (1996) Do bright colors at nests incur a cost due to predation? Evol Ecol 10:285–288CrossRefGoogle Scholar
  34. Håstad O, Victorsson J, Ödeen A (2005) Differences in color vision make passerines less conspicuous in the eyes of their predators. Proc Natl Acad Sci 102:6391–6394PubMedCrossRefGoogle Scholar
  35. Hosmer DW Jr, Lemeshow S (1989) Applied logistic regression. Wiley, New YorkGoogle Scholar
  36. Hozumi S, Yamane S (2001) Effects of surface darkening on cell temperature in paper wasp nests: measurements using paper model nests. Entomol Sci 4:251–256Google Scholar
  37. Jackson RW (1924) Attracting grasshopper sparrows, meadowlarks, and killdeers to use artificial nesting sites. Oologists’ Rec 4:2–5Google Scholar
  38. Kilner RM (2006) The evolution of egg colour and patterning in birds. Biol Rev 81:383–406. doi: 10.1017/S1464793106007044 PubMedCrossRefGoogle Scholar
  39. Korhonen K (1991) Heat flux density on egg surface and incubation rhythm of willow grouse (Lagopus l. lagopus L.). J Therm Biol 16:65–70CrossRefGoogle Scholar
  40. Kull RC Jr (1977) Color selection of nesting material by killdeer. Auk 94:602–604Google Scholar
  41. Lack D (1958) The significance of the colour of turdine eggs. Ibis 100:145–166CrossRefGoogle Scholar
  42. Leader N, Yom-Tov Y (1998) The possible function of stone ramparts at the nest entrance of the blackstart. Anim Behav 56:207–217PubMedCrossRefGoogle Scholar
  43. Littlejohns RT (1932) Notes on four species of dotterels. Emu 31:15–20Google Scholar
  44. Lloyd P, Plaganyi E, Lepage D, Little RM, Crowe TM (2000) Nest-site selection, egg pigmentation and clutch predation in the ground-nesting Namaqua sandgrouse Pterocles namaqua. Ibis 142:123–131CrossRefGoogle Scholar
  45. Mayer PM (1991) Conservation biology of piping plovers in the northern Great Plains. Thesis, University of Missouri-ColumbiaGoogle Scholar
  46. Mayer PM (1993) Conservation of least terns and piping plovers on the Missouri River in North Dakota: management implications of the relationship between breeding population sizes and Garrison Dam operations. In: Higgins KF, Brashier MR (eds) Proceedings, the Missouri River and its tributaries: piping plover and least tern symposium. South Dakota State University, Brookings, pp 47–59Google Scholar
  47. Mayer PM, Ryan MR (1991a) Survival rates of artificial piping plover nests in American avocet colonies. Condor 93:753–755CrossRefGoogle Scholar
  48. Mayer PM, Ryan MR (1991b) Electric fences reduce mammalian predation on piping plover nests and chicks. Wildl Soc Bull 19:59–63Google Scholar
  49. Montevecchi WA (1976) Field experiments on the adaptive significance of avian eggshell pigmentation. Behaviour 58:26–39CrossRefGoogle Scholar
  50. Nguyen LP, Nol E, Abraham KF (2007) Using digital photographs to evaluate the effectiveness of plover egg crypsis. J Wildl Manage 71:2084–2089CrossRefGoogle Scholar
  51. Nicolai CA, Sedinger JS, Wege ML (2004) Regulation of development time and hatch synchronization in black brant (Branta bernicla nigricans). Funct Ecol 18:475–482CrossRefGoogle Scholar
  52. Nol E, Humphrey RC (1994) American oystercatcher (Haematopus palliatus). In: Poole A, Gill F (eds) The birds of North America, no. 82. The Academy of Natural Sciences, The American Ornithologists’ Union, PhiladelphiaGoogle Scholar
  53. Oniki Y (1985) Why robin eggs are blue and birds build nests: statistical tests for Amazonian birds. Ornithol Monogr 36:536–545Google Scholar
  54. Poulton EB (1890) The colours of animals: their meaning and use, especially considered in the case of insects. Appleton, New YorkGoogle Scholar
  55. Prindiville EM (1986) Habitat selection and productivity of piping plovers in central North Dakota. MS thesis, University of Missouri-ColumbiaGoogle Scholar
  56. Prindiville Gaines E, Ryan MR (1988) Piping plover habitat use and reproductive success in North Dakota. J Wildl Manage 52:266–273CrossRefGoogle Scholar
  57. Reid JM, Cresswell W, Holt S, Mellanby RJ, Whitfield DP, Ruxton GD (2002) Nest scrape design and clutch heat loss in pectoral sandpipers (Calidris melanotos). Funct Ecol 16:305–312CrossRefGoogle Scholar
  58. Ricklefs RE, Hainsworth FR (1969) Temperature regulation in nestling cactus wrens: the nest environment. Condor 71:32–37CrossRefGoogle Scholar
  59. Root BG (1996) Alkaline wetland vegetation dynamics at North Dakota piping plover nesting beaches. Dissertation, University of Missouri-ColumbiaGoogle Scholar
  60. Ryan MR, Root BG, Mayer PM (1993) Status of piping plovers in the Great Plains of North America: a demographic simulation model. Conserv Biol 7:581–585CrossRefGoogle Scholar
  61. Sabo JL (2003) Hot rocks or no hot rocks: overnight retreat availability and selection by a diurnal lizard. Oecologia 136:329–335PubMedCrossRefGoogle Scholar
  62. Sánchez JM, Corbacho C, del Viejo AM, Parejo D (2004) Colony-site tenacity and egg color crypsis in the gull-billed tern. Waterbirds 27:21–30CrossRefGoogle Scholar
  63. Schneider EG, McWilliams SR (2007) Using nest temperature to estimate nest attendance of piping plovers. J Wildl Manage 71:1998–2006CrossRefGoogle Scholar
  64. Shaffer TL (2004) A unified approach to analyzing nest success. Auk 121:526–540CrossRefGoogle Scholar
  65. Skutch AF (1976) Parent birds and their young. University of Texas Press, AustinGoogle Scholar
  66. Soler JJ, Moeller AP, Soler M (1998) Nest building, sexual selection and parental investment. Evol Ecol 12:427–441CrossRefGoogle Scholar
  67. Solís JC, de Lope F (1995) Nest and egg crypsis in the ground-nesting stone curlew Burhinus oedicnemus. J Avian Biol 26:135–138CrossRefGoogle Scholar
  68. SSI (2004) Systat software, RichmondGoogle Scholar
  69. Staus NL, Mayer PM (1999) Arthropods and predation of artificial nests in the Bahamas: implications for subtropical avifauna. Wilson Bull 111:561–564Google Scholar
  70. Stoleson SH, Beissinger SR (1999) Egg viability as a constraint on hatching synchrony at high ambient temperatures. J Anim Ecol 68:951–962. doi: 10.1046/j.1365-2656.1999.00342.x CrossRefGoogle Scholar
  71. Szentirmai I, Székely T (2002) Do Kentish plovers regulate the amount of their nest material? An experimental test. Behaviour 139:847–859CrossRefGoogle Scholar
  72. Szentirmai I, Székely T, Liker A (2005) The influence of nest size on heat loss in penduline tit eggs. Acta Zool Acad Sci Hung 51:59–66Google Scholar
  73. Tinbergen N (1963) The shell menace. Nat Hist 72(7):28–35Google Scholar
  74. Underwood TJ, Sealy SG (2002) Adaptive significance of egg colouration. In: Deeming DC (ed) Avian incubation: behaviour, environment, evolution. Oxford University Press, Oxford, pp 280–289Google Scholar
  75. U.S. Fish and Wildlife Service (1985) Endangered and threatened wildlife and plants: determination of endangered and threatened status for the piping plover: a final rule. Fed Regist 50:50726–50734Google Scholar
  76. Wallace AR (1889) Darwinism. New York, HumboldtGoogle Scholar
  77. Weidinger K (2001) Does egg colour affect predation rate on open passerine nests? Behav Ecol Sociobiol 49:456–464CrossRefGoogle Scholar
  78. West PM, Packer C (2002) Sexual selection, temperature and the lion’s mane. Science 297:1339–1343PubMedCrossRefGoogle Scholar
  79. Westmoreland D, Best LB (1986) Incubation continuity and the advantage of cryptic egg coloration to mourning doves. Wilson Bull 98:297–300Google Scholar
  80. Westmoreland D, Kiltie RA (1996) Egg crypsis and clutch survival in three species of blackbirds (Icteridae). Biol J Linn Soc 58:159–172CrossRefGoogle Scholar
  81. Westmoreland D, Schmitz M, Burns KE (2007) Egg color as an adaptation for thermoregulation. J Field Ornithol 787:176–183CrossRefGoogle Scholar
  82. Wilcox L (1939) Notes on the life history of the piping plover. Birds Long Isl 1:3–13Google Scholar
  83. Wilcox L (1959) A twenty year banding study of the piping plover. Auk 76:129–152Google Scholar

Copyright information

© GovernmentEmployee: United States Environmental Protection Agency 2009

Authors and Affiliations

  • Paul M. Mayer
    • 1
  • Levica M. Smith
    • 2
    • 4
  • Robert G. Ford
    • 1
    • 5
  • Dustin C. Watterson
    • 2
  • Marshall D. McCutchen
    • 1
  • Mark R. Ryan
    • 3
  1. 1.Office of Research and DevelopmentUS Environmental Protection AgencyAdaUSA
  2. 2.McNair Scholars ProgramEast Central UniversityAdaUSA
  3. 3.School of Natural ResourcesUniversity of MissouriColumbiaUSA
  4. 4.Physics DepartmentTexas A&M UniversityCollege StationUSA
  5. 5.Office of Research and DevelopmentUS Environmental Protection AgencyCincinnatiUSA

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