Optimal Migration Theory

  • Nikita Chernetsov


This chapter is devoted to the critical analysis of the optimal migration theory. I suggest that this theory has multiple issues, mainly because some of its basic assumptions are not supported by the empirical evidence. The fundamental assumption of the U-shaped flight power curve is not supported by the data. In the birds with high aerodynamic quality (most passerine migrants belong to this category) the flight costs are independent of flight speed, within a rather broad range of flight speed routinely employed. Nevertheless, the optimal migration theory played a very important role in the attempts to find quantitative relationships between the main ecological parameters of migratory stopovers, i.e. between stopover duration, fuel deposition rate and departure fuel load. It has advanced stopover studies greatly, but I claim that its critical revision is necessary.


Fuel Load Flight Speed Migratory Speed Stopover Site Fuel Store 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


  1. Åkesson S, Hedenström A (2000) Wind selectivity of migratory flight departures in birds. Behav Ecol Sociobiol 47:140–144CrossRefGoogle Scholar
  2. Alerstam T, Hedenström A (1998) The development of bird migration theory. J Avian Biol 29:343–369CrossRefGoogle Scholar
  3. Alerstam T, Lindström Å (1990) Optimal bird migration: the relative importance of time, energy, and safety. In: Gwinner E (ed) Bird migration. Springer, BerlinGoogle Scholar
  4. Bairlein F (1987) The migratory strategy of the garden warbler: a survey of field and laboratory data. Ring Migr 8:59–72CrossRefGoogle Scholar
  5. Bairlein F (1991) Body mass of Garden Warblers (Sylvia borin) on migration: a review of field data. Vogelwarte 36:48–61Google Scholar
  6. Bayly NJ (2006) Optimality in avian migratory fuelling behaviour: a study of a trans-Saharan migrant. Anim Behav 71:173–182CrossRefGoogle Scholar
  7. Bayly NJ (2007) Extreme fattening by sedge warblers, Acrocephalus schoenobaenus, is not triggered by food availability alone. Anim Behav 74:471–479CrossRefGoogle Scholar
  8. Begon M, Townsend CR, Harper JL (2006) Ecology: from individuals to ecosystems, 4th edn. Blackwell, OxfordGoogle Scholar
  9. Berger M (1985) Sauerstoffverbrauch von Kolibris (Colibri coruscans und C. thalassinus) beim Horizontalflug. In: Nachtigall W (ed) BIONA report 3. G Fischer, StuttgartGoogle Scholar
  10. Bernstein MH, Thomas SP, Schmidt-Nielsen K (1973) Power input during flight in the fish crow Corvus ossifragus. J Exp Biol 58:401–410Google Scholar
  11. Blem CR (2000) Energy balance. In: Whittow GC (ed) Sturkie’s avian physiology. Academic Press, San DiegoGoogle Scholar
  12. Bowlin MS, Bisson I-A, Shamoun-Baranes J, Reichard JD, Sapir N, Marra PP, Kunz TH, Wilcove DS, Hedenström A, Guglielmo CG, Åkesson S, Ramenofsky M, Wikelski M (2010) Grand challenges in migration biology. Integr Comp Biol 50:261–279PubMedCrossRefGoogle Scholar
  13. Bulyuk VN, Tsvey A (2006) Timing of nocturnal autumn migratory departures in juvenile European robins (Erithacus rubecula) and endogenous and external factors. J Ornithol 147:298–309CrossRefGoogle Scholar
  14. Bundle MW, Hansen KS, Dial KP (2007) Does the metabolic rate-flight speed relationship vary among geometrically similar birds of different mass? J Exp Biol 210:1075–1083PubMedCrossRefGoogle Scholar
  15. Butler PJ, Bishop CM (2000) Flight. In: Whittow GC (ed) Sturkie’s avian physiology. Academic Press, San DiegoGoogle Scholar
  16. Charnov EL (1976) Optimal foraging, the marginal value theorem. Theor Popul Biol 9:129–136PubMedCrossRefGoogle Scholar
  17. Chernetsov N (2010) Recent experimental data on the energy costs of avian flight call for a revision of optimal migration theory. Auk 127:232–234CrossRefGoogle Scholar
  18. Chernetsov NS, Skutina EA, Bulyuk VN, Tsvey AL (2004) Optimal stopover decisions of migrating birds under variable stopover quality: model predictions and the field data. Zh Obschei Biol 65:211–217Google Scholar
  19. Cimprich DA, Woodrey MS, Moore FR (2005) Passerine migrants respond to variation in predation risk during stopover. Anim Behav 69:1173–1179CrossRefGoogle Scholar
  20. Dänhardt J, Lindström Å (2001) Optimal departure decisions of songbirds from an experimental stopover site and the significance of weather. Anim Behav 62:235–243CrossRefGoogle Scholar
  21. Delingat J, Bairlein F, Hedenström A (2008) Obligatory barrier crossing and adaptive fuel management in migratory birds: the case of the Atlantic crossing in northern wheatears (Oenanthe oenanthe). Behav Ecol Sociobiol 62:1069–1078CrossRefGoogle Scholar
  22. Dial KP, Biewener AA, Tobalske BW, Warrick DR (1997) Mechanical power output of bird flight. Nature 390:67–70CrossRefGoogle Scholar
  23. Dierschke V (2003) Predation hazard during migratory stopover: are light or heavy birds under risk? J Avian Biol 24:24–29CrossRefGoogle Scholar
  24. Dolnik VR (1995) Resursy energii i vremeni u ptits v prirode (Energy and time resources in free-living birds). Nauka, St PetersburgGoogle Scholar
  25. Egevang C, Stenhouse IJ, Phillips RA, Petersen A, Fox JW, Silk JRD (2010) Tracking of Arctic terns Sterna paradisaea reveals longest animal migration. Proc Natl Acad Sci USA 107:2078–2208PubMedCrossRefGoogle Scholar
  26. Ellington CP (1991) Limitations on animal flight performance. J Exp Biol 160:71–91Google Scholar
  27. Engel S, Biebach H, Visser GH (2006) Metabolic cost of avian flight in relation to flight velocity: a study in rose-coloured starlings (Sturnus roseus Linnaeus). J Comp Physiol B 176:415–427PubMedCrossRefGoogle Scholar
  28. Engel S, Bowlin MS, Hedenström A (2010) The role of wind-tunnel studies in integrative research on migration biology. Integr Comp Biol 50:323–335PubMedCrossRefGoogle Scholar
  29. Erni B, Liechti F, Bruderer B (2002) Stopover strategies in passerine bird migration: a simulation study. J Theor Biol 219:479–493PubMedCrossRefGoogle Scholar
  30. Fransson T (1998) Patterns of migratory fuelling in whitethroats Sylvia communis in relation to departure. J Avian Biol 29:569–573CrossRefGoogle Scholar
  31. Harrison JF, Roberts SP (2000) Flight respiration and energetics. Annu Rev Physiol 62:179–205PubMedCrossRefGoogle Scholar
  32. Hedenström A (2002) Aerodynamics, evolution and ecology of bird flight. Trends Ecol Evol 17:415–422CrossRefGoogle Scholar
  33. Hedenström A (2008) Adaptations to migration in birds: behavioural strategies, morphology and scaling effects. Phil Trans R Soc B 363:287–299PubMedCrossRefGoogle Scholar
  34. Hedenström A (2012) Recent experimental data on the energy costs of avian flight do not call for a revision of optimal migration theory. Auk 129 (in press)Google Scholar
  35. Hedenström A, Alerstam T (1997) Optimum fuel loads in migratory birds: distinguishing between time and energy minimization. J Theor Biol 189:227–234PubMedCrossRefGoogle Scholar
  36. Houston AI (1998) Models of optimal avian migration: state, time and predation. J Avian Biol 29:395–404CrossRefGoogle Scholar
  37. Klaassen M, Lindström Å (1996) Departure fuel loads in time-minimizing migrating birds can be explained by the energy costs of being heavy. J Theor Biol 183:29–34CrossRefGoogle Scholar
  38. Kullberg C, Fransson T, Jakobsson S (1996) Impaired predator evasion in fat blackcaps (Sylvia atricapilla). Proc R Soc Lond B 263:1671–1675CrossRefGoogle Scholar
  39. Kullberg C, Jakobsson S, Fransson T (2000) High migratory fuel load impair predator evasion in sedge warblers. Auk 117:1034–1038CrossRefGoogle Scholar
  40. Kvist A, Lindström Å, Green M, Piersma T, Visser GH (2001) Carrying large fuel loads during sustained bird flight is cheaper than expected. Nature 413:730–732PubMedCrossRefGoogle Scholar
  41. Lind J (2004) What determines probability of surviving predator attacks in bird migration? The relative importance of vigilance and fuel load. J Theor Biol 231:223–227PubMedCrossRefGoogle Scholar
  42. Lind J, Cresswell W (2006) Anti-predation behaviour during bird migration; the benefit of studying multiple behavioural dimensions. J Ornithol 147:310–316CrossRefGoogle Scholar
  43. Lind J, Fransson T, Jakobsson S, Kullberg C (1999) Reduced take-off ability in robins (Erithacus rubecula) due to migratory fuel load. Behav Ecol Sociobiol 46:65–70CrossRefGoogle Scholar
  44. Lindhe Norberg UM (2004) Bird flight. Acta Zool Sinica 50:921–935Google Scholar
  45. Lindström Å, Alerstam T (1992) Optimal fat loads in migrating birds: a test of the time-minimization hypothesis. Am Nat 140:477–491PubMedCrossRefGoogle Scholar
  46. Loria DE, Moore FR (1990) Energy demands of migration on red-eyed vireos, Vireo olivaceus. Behav Ecol 1:24–35CrossRefGoogle Scholar
  47. Moore FR (1994) Resumption of feeding under risk of predation: effect of migratory condition. Anim Behav 48:975–977CrossRefGoogle Scholar
  48. Norberg UM (1990) Vertebrate flight. Mechanics, physiology, morphology, ecology and evolution. Springer, BerlinGoogle Scholar
  49. Norberg UM (1996) Energetics of flight. In: Carey C (ed) Avian energetics and nutritional ecology. Chapman & Hall, NYGoogle Scholar
  50. Payevsky VA (1985) Demografiya ptits (Avian demography). Nauka, LeningradGoogle Scholar
  51. Pennycuick CJ (1975) Mechanics of flight. In: Farner DS, King JR, Parkes KC (eds) Avian biology, vol 5. Academic Press, NYGoogle Scholar
  52. Pennycuick CJ (1989) Bird flight performance: a practical manual. Oxford University Press, NYGoogle Scholar
  53. Rayner JMV (1990) The mechanics of flight and bird migration performance. In: Gwinner E (ed) Bird migration. Springer, BerlinGoogle Scholar
  54. Robinson WD, Bowlin MS, Bisson I, Shamoun-Baranes J, Thorup K, Diehl R, Kunz TH, Mabey S, Winkler DW (2010) Integrating concepts and technologies to advance the study of bird migration. Front Ecol Environ 8:354–361CrossRefGoogle Scholar
  55. Salewski V (1999) Untersuchungen zur Überwinterungsökologie paläarktischer Singvögel in Westafrika unter besonderer Berücksichtigung der Wechselwirkungen zu residenten Arten. W&T Verlag, BerlinGoogle Scholar
  56. Salewski V, Jones P (2006) Palearctic passerines in Afrotropical environments: a review. J Ornithol 147:192–201CrossRefGoogle Scholar
  57. Schmidt-Nielsen K (1997) Animal physiology: adaptation and environment, 5th edn. Cambridge University Press, NYGoogle Scholar
  58. Schmidt-Wellenburg CA, Biebach H, Daan S, Visser GH (2007) Energy expenditure and wing beat frequency in relation to body mass in free flying Barn Swallows (Hirundo rustica). J Comp Physiol B 177:327–337PubMedCrossRefGoogle Scholar
  59. Schmidt-Wellenburg CA, Engel S, Visser GH (2008) Energy expenditure during flight in relation to body mass: effects of natural increases in mass and artificial load in rose coloured starlings. J Comp Physiol B 178:767–777PubMedCrossRefGoogle Scholar
  60. Stutchbury BJM, Tarof SA, Done T, Gow E, Kramer PM, Tautin J, Fox JW, Afanasyev V (2009) Tracking long-distance songbird migration by using geolocators. Science 323:896PubMedCrossRefGoogle Scholar
  61. Stutchbury BJM, Gow E, Done T, MacPherson M, Fox JW, Afanasyev V (2011) Effects of post-breeding moult and energetic condition on timing of songbird migration into the tropics. Proc R Soc Lond B 278:131–137CrossRefGoogle Scholar
  62. Tobalske BW, Hedrick TL, Dial KP, Biewener AA (2003) Comparative power curves in bird flight. Nature 421:363–366PubMedCrossRefGoogle Scholar
  63. Torre-Bueno JR, LaRochelle J (1978) The metabolic cost of flight in unrestrained birds. J Exp Biol 75:223–229PubMedGoogle Scholar
  64. Tucker VA (1968) Respiratory exchange and evaporative water loss in the flying budgerigar. J Exp Biol 48:67–87Google Scholar
  65. Tucker VA (1972) Metabolism during flight in the laughing gull Larus atricilla. Am J Physiol 222:237–245PubMedGoogle Scholar
  66. Videler JJ (2005) Avian flight. Oxford University Press, OxfordGoogle Scholar
  67. von Stünzner-Karbe D (1996) Territorialität, Habitatnutzung und Furagierverhalten überwinternder Trauerschnäpper (Ficedula hypoleuca) in West-Afrika. Diplomarbeit, Universität BayreuthGoogle Scholar
  68. Vrugt JA, van Belle J, Bouten W (2007) Pareto front analysis of flight time and energy use in long-distance bird migration. J Avian Biol 38:432–442CrossRefGoogle Scholar
  69. Ward S, Bishop CM, Woakes AJ, Butler PJ (2002) Heart rate and the rate of oxygen consumption of flying and walking barnacle geese (Branta leucopis) and bar-headed geese (Anser indicus). J Exp Biol 205:3347–3356PubMedGoogle Scholar
  70. Ward S, Möller U, Rayner JMV, Jackson DM, Nachtigall W, Speakman JR (2004) Metabolic power of European starlings Sturnus vulgaris during flight in a wind tunnel, estimated from heat transfer modelling, doubly labelled water and mask respirometry. J Exp Biol 207:4291–4298PubMedCrossRefGoogle Scholar
  71. Weber TP, Houston AI (1997a) Flight costs, flight range and the stopover ecology of migrating birds. J Anim Ecol 66:297–306CrossRefGoogle Scholar
  72. Weber TP, Houston AI (1997b) A general model for time-minimising avian migration. J Theor Biol 185:447–458CrossRefGoogle Scholar
  73. Weber TP, Houston AI, Ens BJ (1994) Optimal departure fat loads and stopover site use in avian migration: an analytical model. Proc R Soc Lond B 258:29–34CrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2012

Authors and Affiliations

  1. 1.Biological Station RybachyZoological InstituteSt. PetersburgRussia

Personalised recommendations