Wader species wintering on the Banc d'Arguin increased their body mass by about 40% during the 4–6 weeks before their departure in spring. This estimate is based on 1) the empirical fact that most waders which had finished or suspended their body moult into summer plumage were heavy and thus ready to take off, and 2) the assumption that only the heaviest birds in the population left, which allowed the mass of disappearing waders to be estimated from counts and the frequency distribution of body masses in samples of captured birds. The mass gain on the Banc d'Arguin is just over 1% per day, when expressed as a proportion of winter mass. A review of studies on waders preparing for migration shows that 1) the total migratory reserve adds 20–80% to winter mass, 2) the rate of mass gain is 0.1–4% per day and 3. the period of mass increase lasts four weeks on average, but longer if waders prepare for spring migration on the wintering areas. We suggest that all wader species leaving the B anc d'Arguin at the end of April and the beginning of May are able to reach SE. and NW. Europe without refuelling. This seems only possible if current equations to predict flight range systematically under-estimate this range, even when the energetic benefits of favourable winds at high altitude are taken into account.
How to translate text using browser tools
1 January 1990
Moult, Mass and Flight Range of Waders Ready to Take Off for Long-Distance Migrations
Leo Zwarts,
Bruno J. Ens,
Marcel Kersten,
Theunis Piersma
T Alerstam
,
G.A. Gudmundsson
,
P.E. Jönsson
,
J. Karlsson
&
Å. Lindström
1990. Orientation, migration routes and flight behaviour of Knots, Turnstones and Brent geese departing from Iceland in spring. Arctic: in press. Google Scholar
M. Barter
1984. Weight variations in Red-necked Stint (Calidris ruficollis) whilst wintering in Tasmania. Occasional Stint 3: 69–80. Google Scholar
M. Barter
1986. Sex-related differences in adult Curlew Sandpipers Calidris ferruginea caught in Victoria. Stilt 8: 2–8. Google Scholar
M. Barter
1989. Bar-tailed Godwit Limosa lapponica in Australia Part 2: weight, moult and breeding success. Stilt 14: 49–53. Google Scholar
M. Barter
,
A. Jessop
&
C. Minton
1988a. Red Knot Calidris canutus rogersi in Australia. Part 1: Subspecies confirmation, distribution and migration. Stilt 12: 29–32. Google Scholar
M. Barter
,
A. Jessop
&
C. Minton
1988b. Red Knot Calidris canutus rogersi in Australia. Part 2: Biometrics and moult in Victoria and north-western Australia. Stilt 13: 20–27. Google Scholar
J.J. Beukema
1974. Seasonal changes in the biomass of the macro-benthos of a tidal flat area in the Dutch Wadden Sea. Neth. J. Sea Res. 8: 94–107. Google Scholar
G.C. Boere
1976. The significance of the Dutch Waddenzee in the annual life cycle of arctic, subarctic and boreal waders. Part I. The function as a moulting area. Ardea 64: 210–291. Google Scholar
N.J.B.A. Branson
&
C.D.T. Minton
1976. Moult, measurements and migrations of the Grey Plover. Bird Study 23: 257–266. Google Scholar
N.J.B.A. Branson
,
E.D. Ponting
&
C.D.T. Minton
1979. Turnstone populations on the Wash. Bird Study 26: 47–54. Google Scholar
N.E. Buxton
1989. Observations on the low weights of Knot in the western isles of Scotland. Ring. & Migr. 10: 9–12. Google Scholar
G. Castro
&
J.P. Myers
1988. A statistical method to estimate the cost of flight in birds. J. Field Ornith. 59: 369–380. Google Scholar
G. Castro
&
J.P. Myers
1989. Flight range estimates for shorebirds. Auk 106: 474–476. Google Scholar
M.R. Chambers
&
H. Milne
1979. Seasonal variation in the condition of some intertidal invertebrates of the Ythan estuary, Scotland. Estuar. Coast. Mar. Sc. 8: 411–419. Google Scholar
C. Clapham
1978. The Ringed Plover populations of Morecambe Bay. Bird Study 25: 175–180. Google Scholar
C. Clapham
1979. The Turnstone populations of Morecambe Bay. Ring. & Migr. 2: 144–150. Google Scholar
N.A. Clark
,
B.S. Turner
&
J.F. Young
1982. Spring passage of Sanderlings Calidris alba on the Solway Firth. Wader Study Group Bull. 36: 10–11. Google Scholar
S. Cramp
&
K.E.L. Simmons
(eds.)
1983. The birds of the western Palearctic, Vol. III. Oxford University Press, Oxford. Google Scholar
N.C. Davidson
1983. Formulae for estimating the lean weight and fat reserves of live shorebirds. Ring. & Migr. 4: 159–166. Google Scholar
N.C. Davidson
1984a. How valid are flight range estimates for waders? Ring. & Migr. 5: 49–64. Google Scholar
N.C. Davidson
1984b. Changes in the condition of Dunlins and Knots during short-term captivity. Can. J. Zool. 62: 1724–1731. Google Scholar
N.C. Davidson
&
P.R. Evans
(eds.)
1986. The ecology of migrant Knots in North Norway during May 1985. Report SRG 86/1. Shorebird Research Group, Univ. of Durham, Durham. Google Scholar
W.R.J. Dean
1977. Moult of the Curlew Sandpiper at Barberspan. Ostrich, Suppl. 12: 97–101. Google Scholar
W.J.A. Dick
&
M.W. Pienkowski
1979. Autumn and early winter weights of waders in north-west Africa. Ornis Scand. 10: 117–123. Google Scholar
W.J.A. Dick
,
T. Piersma
&
P. Prokosch
1987. Spring migration of the Siberian Knots Calidris canutus canutus: results of a co-operative Wader Study Group project. Ornis Scand. 18: 5–16. Google Scholar
J. Dominguez
&
J.L. Rabuñal
1989. Migrating waders on the Atlantic coast of Galicia (NW. Spain). Miscel. Lania Zoologica 13 (in press). Google Scholar
R. Drent
&
T. Piersma
1990. An exploration of the energetics of leap-frog migration in arctic breeding waders. In:
E. Gwinner
(ed.).
Bird migration: the physiology and ecophysiology: 399–412. Springer-Verlag, Berlin. Google Scholar
P.O. Dunn
,
T.A. May
,
M.A. McCollough
&
M.A. Howe
1988. Length of stay and fat content of migrant Semipalmated Sandpipers in eastern Maine. Condor 90: 824–835. Google Scholar
A.J. van Dijk
,
F.E. de Roder
,
E.C.L. Marteijn
&
H. Spiekman
1990. Summering waders on the Banc d'Arguin, Mauritania: a census in June 1988. Ardea 78: 145–156. Google Scholar
R.A. Eades
&
J.D. Okill
1976. Weight variations of Ringed Plovers on the Dee estuary. Ring. & Migr. 1: 92–97. Google Scholar
R.A. Eades
&
J.D. Okill
1977. Weight changes of Dunlins on the Dee estuary in May. Bird Study 24: 62–63. Google Scholar
C.C.H. Elliott
,
M. Waltner
,
L.G. Underhill
,
J.S. Pringle
&
W.J.A. Dick
1976. The migration system of the Curlew Sandpiper Calidris ferruginea in Africa. Ostrich 47: 191–213. Google Scholar
M. Engelmoer
,
T. Piersma
,
W. Altenburg
&
R. Mes
1984. The Banc d'Arguin (Mauritania). In:
P.R. Evans
,
J.D. Goss-Custard
&
W.G. Hale
(eds.)
Coastal waders and wildfowl in winter: 293–310. Cambridge University Press, Cambridge. Google Scholar
B.J. Ens
,
T. Piersma
,
W.J. Wolff
&
L. Zwarts
(eds.)
1989. Report of the Dutch-Mauritanian project Banc d'Arguin 1985–1986. WIWO-report 25/RIN-report 89/6, Texel. Google Scholar
B.J. Ens
,
P. Duiven
,
C.J. Smit
&
T.M. van Spanje
1990. Spring migration of turnstones from the Banc d'Arguin, Mauritania. Ardea 78: 301–314. Google Scholar
P. Esselink
&
L. Zwarts
1989. Seasonal variation in burrow depth and tidal trends in surface activity of Nereis diversicolor. Mar. Ecol. Prog. Ser. 56: 243–254. Google Scholar
P.R. Evans
&
M.W. Pienkowski
1984. Population dynamics of shorebirds. Behavior of Marine Animals 6: 83–123 Google Scholar
A.A. Goede
&
E. Nieboer
1983. BWeight variation of Dunlins Calidris alpina during post-nuptial moult, after application of weight data transformations. Bird Study 30: 157–163. Google Scholar
A. A. Goede
,
E. Nieboer
&
P.M. Zegers
1990. Body mass increase, migration pattern and breeding grounds of Dunlins Calidris a. alpina, staging in the Dutch Wadden Sea in spring. Ardea 78: 135–144. Google Scholar
L.G. Grimes
1974. Radar tracks of palaearctic waders departing from the coast of Ghana in spring. Ibis 116: 165–171. Google Scholar
P.W. Hicklin
1983a. The feeding ecology of Semipalmated Sandpipers in the Bay of Fundy. Wader Study Group Bull. 39: 48. Google Scholar
P.W. Hicklin
1983b. Influence of prey density and biomass on migration timing of Semipalmated Sandpipers in the Bay of Fundy. Wader Study Group Bull. 39: 59. Google Scholar
H. Insley
&
L. Young
1981. Autumn passage of Ringed Plovers through Southampton Water. Ring. & Migr. 3: 157–164. Google Scholar
C. Johnson
1985. Patterns of seasonal weight variation in waders on the Wash. Ring. & Migr. 6: 19–32. Google Scholar
C. Johnson
&
C.D.T. Minton
1980. The primary moult of the Dunlin Calidris alpina at the Wash. Ornis Scand. 11: 190–195. Google Scholar
O.W. Johnson
&
P.M. Johnson
1983. Plumage-molt-age relationships in “over-summering” and migratory Lesser Golden Plovers. Condor 85: 406–419. Google Scholar
O.W. Johnson
,
M. L. Morton
,
P.L. Bruner
&
P.M. Johnson
1989. Fat cyclicity, predicted migratory flight ranges, and features of wintering behavior in Pacific Golden Plovers. Condor 91: 156–177. Google Scholar
D.W. Johnston
&
R.W. McFarlane
1967. Migration and bioenergetics of flight in the Pacific Golden Plover. Condor 69: 156–168. Google Scholar
J. Jukema
1986. De voorjaarsrui bij Goudplevieren Pluvialis apricaria in Friesland. Limosa 59: 111–113. Google Scholar
M. Kersten
1989. Dunlin. In:
B.J. Ens
,
T. Piersma
,
W.J. Wolff
&
L. Zwarts
(eds.)
Report of the Dutch-Mauritanian project Banc d'Arguin 1985–1986: 57–67. WIWO-report 25 / RIN-report 89/6, Texel. Google Scholar
M. Kersten
&
T. Piersma
1987. High levels of energy expenditure in shorebirds; metabolic adaptations to an energetically expensive way of life. Ardea 75: 175–187. Google Scholar
M. Kersten
&
C.J. Smit
1984. The Atlantic coast of Morocco. In:
P.R. Evans
,
J.D. Goss-Custard
&
W.G. Hale
(eds.)
Coastal waders and wildfowl in winter: 276–292. Cambridge University Press, Cambridge. Google Scholar
J.K. Kirkwood
1983. A limit to metabolisable energy intake in mammals and birds. Comp. Biochem. Physiol. 75A: 1–3. Google Scholar
M. Klaassen
,
M. Kersten
&
B.J. Ens
1990. Energetic requirements for maintenance and premigratory body mass gain of waders wintering in Africa. Ardea 78: 209–220. Google Scholar
B. Lane
&
A. Jessop
1985. Tracking of migrating waders in north-western Australia using meteorological radar. Stilt 6: 17–28. Google Scholar
D. Lank
1979. Dispersal and predation rate of wing-tagged Semipalmated Sandpipers Calidris pusilla and an evaluation of the technique. Wader Study Group Bull. 27: 41–46. Google Scholar
D. Lank
1985. Migratory behavior of Semipalmated Sandpipers at inland and coastal areas. Ph.D. Thesis, Cornell University, Ithaca. Google Scholar
S.L. Lima
1986. Predation risk and unpredictable feeding conditions: determinants of body mass in birds. Ecology 67: 377–385. Google Scholar
R.J. Longstreet
1930. Notes on speed of flight of certain water birds. Auk 47: 428–429. Google Scholar
J.W. Mascher
1966. Weight variations in resting Dunlins (Calidris a. alpina) on autumn migration in Sweden. Bird-Banding 37: 1–34. Google Scholar
J.W. Mascher
&
V. Marcström
1976. Measures, weights, and lipid levels in migrating Dunlins Calidris a. alpina L. at the Ottenby Bird Observatory, South Sweden. Ornis Scand. 7: 49–59. Google Scholar
D. Masman
&
M. Klaassen
1987. Energy expenditure during free flight in trained and free-living Eurasian Kestrels (Falco tinnunculus). Auk 104: 603–616. Google Scholar
E.H. McEwan
&
P.M. Whitehead
1984. Seasonal changes in body weight and composition of Dunlin (Calidris alpina). Can. J. Zool. 62: 154–156. Google Scholar
R. McNeil
1969. La détermination du continu lipidique et de la capacité de vol chez quelques espèces d'oiseaux de rivage (Charadriidae et Scolopacidae). Can. J. Zool. 47: 525–536. Google Scholar
R. McNeil
1970. Hivernage et estivage d'oiseaux aquatiques nord-américains dans le Nord-Est du Vénézuéla (Mue, accumulation de graisse, capacité de vol et routes de migration). Oiseau et R.F.O. 40: 185–302. Google Scholar
R. McNeil
&
F. Cadieux
1972. Numerical formulae to estimate flight range of some north American shorebirds from fresh weight and wing length. Bird-Banding 43: 107–113. Google Scholar
R. Meinertzhagen
1955. The speed and altitude of bird flight (with notes on other animals). Ibis 97: 81–117. Google Scholar
N.B. Metcalfe
&
R.W. Furness
1985. Survival, winter population stability and site fidelity in the Turnstone Arenaria interpres. Bird Study 32: 207–214. Google Scholar
E. Middlemiss
1961. Biological aspects of Calidris minuta while wintering in south-west Cape. Ostrich 32: 107–121. Google Scholar
R.I.G. Morrison
1975. Migration and morphometrics of European Knot and Turnstone on Ellesmere Island, Canada. Bird-Banding 46: 290–301. Google Scholar
R.I.G. Morrison
1984. Migration systems of some New World shorebirds. Behav. of Marine Animals 6: 125–202. Google Scholar
M Moser
&
M. Carrier
1983. Patterns of population turnover in Ringed Plovers and Turnstones during their spring migration through the Solway Firth in 1983. Wader Study Group Bull. 39: 37–41. Google Scholar
K.A. Nagy
&
C.C. Peterson
1989. Scaling of water flux rate in animals. Univ. of California Press, Berkeley. Google Scholar
H. Noer
1979. Speeds of migrating waders Charadriidae. Dansk orn. Foren. Tidsskr. 73: 215–224. Google Scholar
GAG Münster
1989. Beobachtungen zur Heimzugstrategie des Kampfläufers Philomachus pugnax. J. Orn. 130: 175–182. Google Scholar
G. Page
&
A.L.A. Middleton
1972. Fat deposition during autumn migration in the Semipalmated Sandpiper. Bird-Banding 43: 85–96. Google Scholar
D.C. Paton
&
B.J. Wykes
1978. Re-appraisal of moult of Red-necked Stints in southern Australia. Emu 78: 54–60. Google Scholar
D.J. Pearson
1987. The status, migrations and seasonality of the Little Stint in Kenya. Ring. & Migr. 8: 91–108. Google Scholar
M.W. Pienkowski
&
W.J.A. Dick
1975. The migration and wintering of Dunlin Calidris alpina in northwest Africa. Ornis Scand. 6: 151–167. Google Scholar
M.W. Pienkowski
&
P.J. Knight
,
D.J. Stanyard
&
F.B. Argyle
1976. The primary moult of waders on the Atlantic coast of Morocco. Ibis 118: 347–365. Google Scholar
M.W. Pienkowski
,
C.S. Lloyd
&
C.D.T. Minton
1979. Seasonal and migrational weight changes in Dunlins. Bird Study 26: 134–148. Google Scholar
T. Piersma
1987. Hink, stap of sprong? Reisbeperkingen van arctische steltlopers door voedselzoeken, vetopbouw en vliegsnelheid. Limosa 60: 185–191. Google Scholar
T. Piersma
&
N.E. van Brederode
1990. The estimation of fat reserves in coastal waders before their departure from northwest Africa in spring. Ardea 78: 221–236. Google Scholar
T. Piersma
&
J. Jukema
1990. Budgeting the flight of a long-distance migrant: changes in nutrient reserve levels of Bar-tailed Godwits at successive spring staging sites. Ardea 78: 315–337. Google Scholar
T. Piersma
,
M. Klaassen
,
J.H. Bruggemann
,
A-M. Blomert
,
A. Gueye
,
Y. Ntiamoa-Baidu
&
N.E. van Brederode
1990. Seasonal timing of the spring departure of waders from the Banc d'Arguin, Mauritania. Ardea 78: 123–134. Google Scholar
T Prater
&
J. Wilson
1972. Spring migration of Knot in Morecambe Bay. Wader Study Group Bull. 5: 9–12. Google Scholar
P. Prokosch
1988. Das Schleswig-Holsteinische Wattenmeer als Frühjahrs-Aufenthaltsgebiet arktischer Watvogel-Populationen am Beispiel von Kiebitzregenpfeifer (Pluvialis squatarola, L. 1758), Knutt (Calidris canutus, L. 1758) und Pfuhlschnepfe (Limosa lapponica, L. 1758). Corax 12: 273–442. Google Scholar
J.M.V. Rayner
1990. The mechanics of flight and bird migration performances. In:
E. Gwinner
(ed.)
Bird migration: the physiology and ecophysiology. Springer-Verlag, Berlin. Google Scholar
W.J. Richardson
1979. Southeastward shorebird migration over Nova Scotia and New Brunswick in autumn: a radar study. Can. J. Zool. 57: 107–124. Google Scholar
M.B. Schmitt
&
P.J. Whitehouse
1976. Moult and mensural data of Ruff on the Witwatersrand. Ostrich 47: 179–190. Google Scholar
G.D. Schnell
&
J.J. Hellack
1978. Flight speeds of Brown Pelicans, Chimney Swifts, and other birds. Bird-Banding 49: 108–112. Google Scholar
C.J. Smit
&
T. Piersma
1989. Numbers, mid-winter distribution and migration of wader populations using the East Atlantic Flyway. In:
H. Boyd
&
J-Y. Pirot
(eds.)
Flyways and reserve networks for water birds: 24–63. IWRB Spec. Publ. 9, Slimbridge. Google Scholar
P.I. Stanley
&
C.D.T. Minton
1972. The unprecedented westward migration of Curlew sandpipers in autumn 1969. Brit. Birds. 65: 365–380. Google Scholar
D.J. Steventon
1977. Dunlin in Portsmouth, Langstone and Chichester Harbours. Ring. & Migr. 1: 141–147. Google Scholar
P.K. Stoddard
,
J.E. Marsden
&
T.C. Williams
.
Computer simulation of autumnal bird migration over the western north Atlantic. Anim. Behav. 31: 173–180. Google Scholar
E. Stresemann
&
V. Stresemann
1966. Die Mauser der Vögel. J. Orn. 107, Sonderheft. Google Scholar
R.W. Summers
,
L.G. Underhill
,
M. Waltner
&
D.A. Whitelaw
1987. Population, biometrics and movements of the Sanderling Calidris alba in southern Africa. Ostrich 58: 24–39. Google Scholar
R.W. Summers
,
L.G. Underhill
,
C.F. Clinning
&
M. Nicols
1989. Populations, migrations, biometrics and moult of the Turnstone Arenaria i. interpres on the east Atlantic coastline, with special reference to the Siberian population. Ardea 77: 145–168 Google Scholar
R.W. Summers
&
M. Waltner
1978. Seasonal variations in the mass of waders in southern Africa, with special reference to migration. Ostrich 50: 21–37. Google Scholar
C. Swennen
,
M. F. Leopold
,
L.L.M. de Bruijn
1989. Time-stressed oystercatchers, Haematopus ostralegus, can increase their intake rate. Anim. Behav. 38: 8–22. Google Scholar
D.G. Thomas
&
A.J. Dartnall
1971a. Moult of the Red-necked Stint. Emu 71: 49–53. Google Scholar
D.G. Thomas
&
A.J. Dartnall
1971b. Moult of the Curlew Sandpiper in relation to its annual cycle. Emu 71: 153–158. Google Scholar
Thompson M.C.
1974. Migratory patterns of Ruddy Turnstones in the Central Pacific region. Living Bird 12: 5–23. Google Scholar
A.J. Tree
1979. Biology of the Greenshank in southern Africa. Ostrich 50: 240–251. Google Scholar
V.A. Tucker
&
K. Schmidt-Koenig
1971. Flight speeds of birds in relation to energetics and wind directions. Auk 88: 91–107. Google Scholar
G.C. Whittow
1986. Energy metabolism. In:
P.D. Sturkie
(ed.)
Avian Physiology, third edition: 253–268. Springer-Verlag, New York. Google Scholar
T.C. Williams
1985. Autumnal bird migration over the Windward Caribbean Islands. Auk 102: 163–167. Google Scholar
T.C. Williams
&
J.M. Williams
1988. Radar and visual observations of autumnal (southward) shorebird migration on Guam. Auk 105: 460–466. Google Scholar
W.H. Wilson
1990. Relationship between prey abundance and foraging site selection by Semipalmated Sandpipers on a Bay of Fundy mudflat. J. Field Ornith. 61: 9–19. Google Scholar
J.R. Wilson
,
M.A. Czajkowski
&
M.W. Pienkowski
1980. The migration through Europe and wintering in West Africa of Curlew Sandpipers. Wildfowl 31: 107–122. Google Scholar
E. Wymenga
,
M. Engelmoer
,
C.J. Smit
&
T.M. van Spanje
1990. Geographical breeding origin and migration of waders wintering in West Africa. Ardea 78: 83–112. Google Scholar
L. Zwarts
1988 Numbers and distribution of coastal waders in Guinea-Bissau. Ardea 76: 42–55. Google Scholar
L. Zwarts
1990. Increased prey availability drives premigration hyperphagia in Whimbrels and allows them to leave the Banc d'Arguin, Mauritania, in time. Ardea 78: 279–300. Google Scholar
L. Zwarts
,
A-M. Blomert
,
B.J. Ens
,
R. Hupkes
&
T.M. van Spanje
1990a. Why do waders reach high density on the tidal flats of the Banc d'Arguin Mauritania?, Ardea 78: 39–52. Google Scholar
L. Zwarts
,
A-M. Blomert
&
R. Hupkes
1990b. Increase of feeding time in waders preparing their spring departure from the Banc d'Arguin, Mauritania. Ardea 78 237–256. Google Scholar
L. Zwarts
&
S. Dirksen
1990. Digestive bottleneck limits the increase in food intake of Whimbrels preparing their spring migration from the Banc d'Arguin, Mauritania. Ardea 78: 257–278. Google Scholar
L. Zwarts
&
T. Piersma
1990. How important is the Banc d'Arguin, Mauritania, as a temporary staging area for waders in spring? Ardea 78: 113–122. Google Scholar