Behavioral Ecology and Sociobiology

, Volume 62, Issue 7, pp 1069–1078 | Cite as

Obligatory barrier crossing and adaptive fuel management in migratory birds: the case of the Atlantic crossing in Northern Wheatears (Oenanthe oenanthe)

  • Julia DelingatEmail author
  • Franz Bairlein
  • Anders Hedenström
Original Paper


Behaviour on migration was often suggested to be selected for time-minimising strategies. Current optimality models predict that optimal fuel loads at departure from stopover sites should increase with increasing fuel deposition rates. We modified such models for the special case of the east Atlantic crossing of the Northern Wheatear (Oenanthe oenanthe). From optimality theory, we predict that optimal time-minimising behaviour in front of such a barrier should result in a positive correlation between fuel deposition rates and departure fuel loads only above a certain threshold, which is the minimum fuel load (f min) required for the barrier crossing. Using a robust range equation, we calculated the minimum fuel loads for different barrier crossings and predict that time-minimising wheatears should deposit a minimum of 24% fuel in relation to lean body mass (m 0 ) for the sea crossing between Iceland and Scotland. Fuel loads of departing birds in autumn in Iceland reached this value only marginally but showed positive correlation between fuel deposition rate (FDR) and departure fuel load (DFL). Birds at Fair Isle (Scotland) in spring, which were heading towards Iceland or Greenland, were significantly heavier and even showed signs of overloading with fuel loads up to 50% of lean body mass. Departure decisions of Icelandic birds correlated significantly with favourable wind situations when assuming a migration direction towards Spain; however, the low departure fuel loads contradict a direct non-stop flight.


Barrier crossing Flight costs Optimal migration Oenanthe oenanthe Fuel loads 



This study was financially supported by the ESF BIRD-Program and the Deutsche Forschungsgemeinschaft (BA 816/15-1). Deryk Shaw and the Fair Isle Bird Observatory kindly supported the field work, as well as Aevar Petersen and the Icelandic Institute of Natural History. The authors acknowledge the use of National Centers for Environmental Prediction Reanalysis data and the NOAA-CIRES Climate Diagnostic Centre for providing wind data. We thank Gudmundur Gudmundsson, Heiko Schmaljohann and two anonymous referees for very helpful comments on the manuscript.


  1. Åkesson S, Hedenström A (2000) Wind selectivity of migratory flight departures in birds. Behav Ecol Sociobiol 47:140–144CrossRefGoogle Scholar
  2. Åkesson S, Alerstam T, Hedenström A (1996a) Flight initiation of nocturnal passerine migrants in relation to celestial orientation conditions at twilight. J Avian Biol 27:95–102CrossRefGoogle Scholar
  3. Åkesson S, Karlsson L, Walinder G, Alerstam T (1996b) Bimodal orientation and the occurrence of temporary reverse bird migration during autumn in south Scandinavia. Behav Ecol Sociobiol 38:293–302CrossRefGoogle Scholar
  4. Alerstam T (1981) The course and timing of bird migration. In: Aidley DJ (ed) Animal migration. Society for Experimental Biology Seminar Series, vol 13. Cambridge University Press, Cambridge, pp 9–54Google Scholar
  5. Alerstam T (1990) Bird migration. Cambridge University Press, CambridgeGoogle Scholar
  6. Alerstam T, Lindström Å (1990) Optimal bird migration: the relative importance of time, energy and safety. In: Gwinner E (ed) Bird migration: the physiology and ecophysiology. Springer, Berlin, Heidelberg, New York, pp 331–335Google Scholar
  7. Alerstam T (2001) Detours in bird migration. J Theor Biol 209:319–331PubMedCrossRefGoogle Scholar
  8. Alerstam T, Hedenström A (1998) The development of bird migration theory. J Avian Biol 29:343–369CrossRefGoogle Scholar
  9. Alerstam T, Hedenström A, Åkesson S (2003) Long-distance migration: evolution and determinants. Oikos 103:247–260CrossRefGoogle Scholar
  10. Bairlein F (1998) The European–African songbird migration network: new challenges for large-scale study of bird migration. Biol Conserv Fauna 102:13–27Google Scholar
  11. Bauchinger U, Biebach H (2001) Differential catabolism of muscle protein in Garden Warblers (Sylvia borin): flight and leg muscle act as a protein source during long. distance migration. J Comp Physiol B 171:293–301PubMedCrossRefGoogle Scholar
  12. Bayly NJ (2006) Optimality in avian migratory fuelling behaviour: a study of a trans-Saharan migrant. Anim Behav 71:173–182CrossRefGoogle Scholar
  13. Bruderer B, Boldt A (2001) Flight characteristics of birds: I. radar measurements of speeds. Ibis 143:178–204CrossRefGoogle Scholar
  14. Bruderer B, Underhill L, Liechti F (1995) Altitude choice of night migrants in a desert area predicted by meteorological factors. Ibis 137:44–45CrossRefGoogle Scholar
  15. Cramp S (1988) Handbook of the birds of Europe, the Middle East and North Africa. Oxford University Press, OxfordGoogle Scholar
  16. Dänhardt J, Lindström Å (2001) Optimal departure desicions of songbirds from an experimental stopover site and the significance of weather. Anim Behav 62:235–243CrossRefGoogle Scholar
  17. Delingat J, Dierschke V, Schmaljohann H, Mendel B, Bairlein F (2006) Daily stopovers as optimal migration strategy in a long-distance migrating passerine: The Northern Wheatear Oenanthe oenanthe. Ardea 94:593–605Google Scholar
  18. Dierschke V, Mendel B, Schmaljohann H (2005) Differential timing of spring migration in Northern Wheatears: hurried males or weak females? Behav Ecol Sociobiol 57:470–480CrossRefGoogle Scholar
  19. Drent R, Daan S (1980) The prodent parent: energetic adjustment in avian breeding. Ardea 68:225–252Google Scholar
  20. Erni B, Liechti F, Underhill LG, Bruderer B (2002) Wind and rain govern the intesity of nocturnal bird migration in central Europe—a log-linear regression analysis. Ardea 90:155–166Google Scholar
  21. Fransson T (1998) A feeding experiment on migratory fuelling in whitethroats, Sylvia communis. Anim Behav 55:153–162PubMedCrossRefGoogle Scholar
  22. Gosler AG, Greenwood JJD, Baker JK, Davidson NC (1998) The field determination of body size and condition in passerines: a report to the British Ringing Committee. Br Birds 45:92–103Google Scholar
  23. Gudmundsson F (1970) Bird migration studies on Surtsey in the Spring of 1968. Surtsey Research Progress Report 1968 Field Season V: 30–38. ReykjavikGoogle Scholar
  24. Gudmundsson GA, Lindström Å, Alerstam T (1991) Optimal fat loads and long-distance flights by migrating Knots Calidris canutus, Sanderlings C. alba and Turnstones Arenaria interpres. Ibis 133:140–152CrossRefGoogle Scholar
  25. Hedenström A (2002) Aerodynamics, evolution, and ecology of bird flight. Trends Ecol Evol 7:415–422CrossRefGoogle Scholar
  26. Hedenström A (2006) Scaling of migration and the annual cycle in birds. Ardea 94:399–408Google Scholar
  27. Hedenström A (2007) Adaptations to migration in birds: behavioural strategies, morphology and scaling effects. Phil Trans R Soc Lond B DOI  10.1098/rstb.2007.2140
  28. Hedenström A, Pettersson J (1986) Differences in fat deposits and wing pointedness between male and female Willow Warblers caught on spring migration at Ottenby, SE Sweden. Ornis Scand 17:182–185CrossRefGoogle Scholar
  29. Hedenström A, Alerstam T (1997) Optimum fuel loads in migratory birds: distinguishing between time and energy minimization. J Theor Biol 189:227–234PubMedCrossRefGoogle Scholar
  30. Hussell DJT, Lambert AB (1980) New estimates of weight loss in birds during nocturnal migration. Auk 97:547–558Google Scholar
  31. Imboden C, Imboden D (1972) Formel für Orthodrome und Loxodrome bei der Berechnung von Richtung und Distanz zwischen Beringungs- und Wiederfundort. Vogelwarte 26:336–346Google Scholar
  32. Kaiser A (1993) A new multi-category classification of subcutaneous fat deposits of song birds. J Field Ornithol 64:246–255Google Scholar
  33. Klaassen M (2003) The relationships between migration and breeding strategies in arctic breeding birds. In: Berthold P, Gwinner E, Sonnenschein E (eds) Avian migration. Springer, Berlin Heidelberg New York, pp 237–249Google Scholar
  34. Komenda-Zehnder S, Liechti F, Bruderer B (2002) Is reverse migration a common feature of nocturnal bird migration?—an analysis of radar data from Israel. Ardea 90:325–334Google Scholar
  35. Kvist A, Klaaseen M, Lindström Å (1998) Energy expenditure in relation to flight speed: what is the power of mass loss rate estimates? J Avian Biol 29:485–498CrossRefGoogle Scholar
  36. Liechti F (2006) Birds: blowin' by the wind? J Ornithol 147:202–211CrossRefGoogle Scholar
  37. Liechti F, Schaller E (1999) The use of low-level jets by migrating birds. Naturwissenschaften 86:549–551PubMedCrossRefGoogle Scholar
  38. Liechti F, Hedenström A, Alerstam T (1994) Effects of sidewinds on optimal flight speed of birds. J Theor Biol 170:219–225CrossRefGoogle Scholar
  39. Lindström Å, Alerstam T (1992) Optimal fat loads in migrating birds: a test of the time-minimization hypothesis. Am Nat 140:477–491CrossRefPubMedGoogle Scholar
  40. Lindstöm Å, Piersma T (1993) Mass changes in migrating birds: the evidence for fat and protein storage re-examined. Ibis 135:70–78CrossRefGoogle Scholar
  41. Luttik R, Wattel J (1979) Observation of land birds on weather ships in the North Atlantic. Limosa 52:191–208Google Scholar
  42. Mc Williams SR, Guglielmo C, Pierce B, Klaassen M (2004) Flying, fasting, and feeding in birds during migration: a nutritional and physiological ecology perspective. J Avian Biol 35:377–393CrossRefGoogle Scholar
  43. Morrison RIG, Hobson KA (2004) Use of body stores in shorebirds after arrival on high-arctic breeding grounds. Auk 121:333–344CrossRefGoogle Scholar
  44. Nisbet ICT (1963) Weight-loss during migration. Part II: review of other estimates. Bird Banding 34:107–159Google Scholar
  45. Ottosson U, Sandberg R, Petterson J (1990) Orientation cage and release experiments with migratory Wheatears (Oenanthe oenanthe) in Scandinavia and Greenland: the importance of visual cues. Ethology 86:57–70CrossRefGoogle Scholar
  46. Pennycuick CJ (1975) Mechanics of flight. In: Farner DS, King JR (eds) Avian biology. Academic Press, New York, pp 1–75Google Scholar
  47. Pennycuick CJ (1989) Bird flight performance: a practical calculation manual. Oxford University Press, OxfordGoogle Scholar
  48. Penycuick CJ (1978) Fifteen testable predictions about bird flight. Oikos 30:165–176CrossRefGoogle Scholar
  49. Piersma T (1998) Phenotypic flexibility during migration: optimization of organ size contingent on the risks and rewards of fuelling and flight? J Avian Biol 29:511–520CrossRefGoogle Scholar
  50. Richardson WJ (1990) Timing of bird migration in relation to weather: updated review. In: Gwinner E (ed) Bird migration. Springer, Berlin Heidelberg New York, pp 78–101Google Scholar
  51. Salomonsen F (1934) La variation géographique et la migration de la Traquet motteux. L’oiseau 2:222–225Google Scholar
  52. Sandberg R (1996) Fat reserves of migrating passerines at arrival on the breeding grounds in Swedish Lapland. Ibis 138:514–524CrossRefGoogle Scholar
  53. Sandberg R, Moore FR (1996) Fat stores and arrival on the breeding grounds: reproductive consequences for passerine migrants? Oikos 77:577–581CrossRefGoogle Scholar
  54. Schmaljohann H, Dierschke V (2005) Optimal migration and predation risk: a field experiment with Northern Wheatears (Oenanthe oenanthe). J Anim Ecol 74:131–138CrossRefGoogle Scholar
  55. Smith RJ, Moore FR (2003) Arrival fat and reproductive performance in a long-distance passerine migrant. Oecologia 134:325–331PubMedGoogle Scholar
  56. Snow DW (1953) The migration of the Greenland Wheatear. Ibis 95:377–378Google Scholar
  57. Svensson L (1992) Identification guide to European passerines, 4th edn. Naturhistoriska Riksmuseet, StockholmGoogle Scholar
  58. Thorup K, Ortvad TE, Rabøl J (2006) Do Nearctic Northern Wheatears (Oenanthe oenanthe leucorhoa) migrate nonstop to Africa? Condor 108:446–451CrossRefGoogle Scholar
  59. Weber TP, Hedenström A (2000) Optimal stopover decisions under wind influence: the effects of correlated winds. J Theor Biol 205:95–104PubMedCrossRefGoogle Scholar
  60. Weber TP, Ens BJ, Houston AI (1998) Optimal avian migration: a dynamic model of fuel stores and site use. Evol Ecol 12:377–401CrossRefGoogle Scholar
  61. Weber TP, Fransson T, Houston AI (1999) Should I stay or should I go? Testing optimality models of stopover decisions in migrating birds. Behav Ecol Sociobiol 46:280–286CrossRefGoogle Scholar
  62. Weber TP, Houston AI, Ens BJ (1994) Optimal departure fat loads and site use in avian migration: an analytical model. Proc R Soc Lond B Biol Sci 258:29–34CrossRefGoogle Scholar
  63. Williamson K (1958) Bergmann’s rule and obligatory overseas migration. Br Birds LI:209–232Google Scholar

Copyright information

© Springer-Verlag 2007

Authors and Affiliations

  • Julia Delingat
    • 1
    Email author
  • Franz Bairlein
    • 1
  • Anders Hedenström
    • 2
  1. 1.Institute of Avian ResearchWilhelmshavenGermany
  2. 2.Department of Theoretical EcologyEcology BuildingLundSweden

Personalised recommendations