Absolute Consistency: Individual versus Population Variation in Annual-Cycle Schedules of a Long-Distance Migrant Bird

Flexibility in scheduling varies throughout an organism’s annual cycle, reflecting relative temporal constraints and fitness consequences among life-history stages. Time-selection can act at different scales, either by limiting the range of alternative strategies in the population, or by increasing the precision of individual performance. We tracked individual bar-tailed godwits Limosa lapponica baueri for two full years (including direct observation during non-breeding seasons in New Zealand and geolocator tracking of round-trip migrations to Alaska) to present a full annual-cycle view of molt, breeding, and migration schedules. At both population and individual scales, temporal variation was greater in post-breeding than pre-breeding stages, and greater in molts than in movements, but schedules did not tighten across successive stages of migration toward the breeding grounds. In general, individual godwits were quite consistent in timing of events throughout the year, and repeatability of pre-breeding movements was particularly high (r = 0.82–0.92). However, we demonstrate that r values misrepresent absolute consistency by confounding inter- and intra-individual variation; the biological significance of r values can only be understood when these are considered separately. By doing so, we show that some stages have considerable tolerance for alternative strategies within the population, whereas scheduling of northbound migratory movements was similar for all individuals. How time-selection simultaneously shapes both individual and population variation is central to understanding and predicting adaptive phenological responses to environmental change.

[1]  A. Dawson Control of the annual cycle in birds: endocrine constraints and plasticity in response to ecological variability , 2008, Philosophical Transactions of the Royal Society B: Biological Sciences.

[2]  A. Møller Phenotype-dependent arrival time and its consequences in a migratory bird , 1994, Behavioral Ecology and Sociobiology.

[3]  C. Both,et al.  Adjustment to climate change is constrained by arrival date in a long-distance migrant bird , 2001, Nature.

[4]  A. Hedenström,et al.  The scheduling of molt in migratory birds , 1995, Evolutionary Ecology.

[5]  Thomas Alerstam,et al.  Temporal and spatial patterns of repeated migratory journeys by ospreys , 2006, Animal Behaviour.

[6]  A. Helbig,et al.  Phylogeny and species limits in the Palaearctic chiffchaff Phylloscopus collybita complex: mitochondrial genetic differentiation and bioacoustic evidence† , 1996 .

[7]  P. Lourenço,et al.  Repeatable timing of northward departure, arrival and breeding in Black-tailed Godwits Limosa l. limosa, but no domino effects , 2011, Journal of Ornithology.

[8]  Felix Liechti,et al.  Birds: blowin’ by the wind? , 2006, Journal of Ornithology.

[9]  L. Underhill,et al.  The Primary Moult of Curlew Sandpipers Calidris ferruginea in North-Western Australia Shifts According to Breeding Success , 2011 .

[10]  T. Alerstam,et al.  Individuality in bird migration: routes and timing , 2011, Biology Letters.

[11]  Colin E. Studds,et al.  Rainfall-induced changes in food availability modify the spring departure programme of a migratory bird , 2011, Proceedings of the Royal Society B: Biological Sciences.

[12]  E. Gwinner,et al.  Circannual clocks in avian reproduction and migration , 2008 .

[13]  Alasdair I. Houston,et al.  The timing of migration within the context of an annual routine , 1998 .

[14]  E. Gwinner Bird Migration: Physiology and Ecophysiology , 2011 .

[15]  Kate L. Laskowski,et al.  The repeatability of behaviour: a meta-analysis , 2009, Animal Behaviour.

[16]  R. Furness,et al.  Short-Lived Repeatabilities in Long-Lived Great Skuas: Implications for the Study of Individual Quality , 1999 .

[17]  P. Battley Consistent annual schedules in a migratory shorebird , 2006, Biology Letters.

[18]  Alasdair I. Houston,et al.  Optimal avian migration: A dynamic model of fuel stores and site use , 1998, Evolutionary Ecology.

[19]  Nils Warnock,et al.  Spring Stopover Ecology of Migrant Western Sandpipers , 1998 .

[20]  C. Guglielmo,et al.  Short- and long-term costs of reproduction in a migratory songbird , 2012 .

[21]  J. Conklin Extreme migration and the annual cycle : individual strategies in New Zealand Bar-tailed Godwits : a thesis presented in partial fulfilment of the requirements for the degree of Doctor of Philosophy in Ecology at Massey University, Palmerston North, New Zealand , 2011 .

[22]  Erik Postma,et al.  Selection on Heritable Phenotypic Plasticity in a Wild Bird Population , 2005, Science.

[23]  S. Hinsley,et al.  Rate of moult affects feather quality: a mechanism linking current reproductive effort to future survival , 2000, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[24]  David C. Douglas,et al.  Contrasting extreme long-distance migration patterns in bar-tailed godwits Limosa lapponica , 2012 .

[25]  H. Kokko Competition for early arrival in migratory birds , 1999 .

[26]  David C. Douglas,et al.  Extreme endurance flights by landbirds crossing the Pacific Ocean: ecological corridor rather than barrier? , 2008, Proceedings of the Royal Society B: Biological Sciences.

[27]  J. Madsen Spring Migration Strategies in Pink-Footed Geese Anser brachyrhynchus and Consequences for Spring Fattening and Fecundity , 2001 .

[28]  P. Battley,et al.  Impacts of wind on individual migration schedules of New Zealand bar-tailed godwits , 2011 .

[29]  P. Battley,et al.  Contour-feather moult of Bar-tailed Godwits (Limosa lapponica baueri) in New Zealand and the northern hemisphere reveals multiple strategies by sex and breeding region , 2011 .

[30]  E. Rees Consistency in the timing of migration for individual Bewick's swans , 1989, Animal Behaviour.

[31]  Jean-Louis Martin,et al.  Inter-annual variation in the breeding chronology of arctic shorebirds: effects of weather, snow melt and predators. , 2010 .

[32]  P. Battley,et al.  Do body condition and plumage during fuelling predict northwards departure dates of Great Knots Calidris tenuirostris from north‐west Australia? , 2003 .

[33]  E. Gwinner,et al.  S40-2 Timing of molt as a buffer in the avian annual cycle , 2006 .

[34]  J. Lyons,et al.  Fat Content and Stopover Ecology of Spring Migrant Semipalmated Sandpipers in South Carolina , 1995 .

[35]  J. Wingfield,et al.  ENDOCRINE CONTROL OF LIFE-CYCLE STAGES: A CONSTRAINT ON RESPONSE TO THE ENVIRONMENT? , 2000 .

[36]  C. M. Lessells,et al.  Unrepeatable repeatabilities: a common mistake , 1987 .

[37]  J. Wingfield,et al.  Adjustments of the prebasic molt schedule in birds , 1992 .

[38]  O. Hoegh‐Guldberg,et al.  Ecological responses to recent climate change , 2002, Nature.

[39]  Shinichi Nakagawa,et al.  Repeatability for Gaussian and non‐Gaussian data: a practical guide for biologists , 2010, Biological reviews of the Cambridge Philosophical Society.

[40]  J. Wiens,et al.  MODELS AND REALITY: TIME–ENERGY TRADE‐OFFS IN PECTORAL SANDPIPER (CALIDRIS MELANOTOS) MIGRATION , 1999 .

[41]  G. Gauthier,et al.  Individual variation in timing of migration: causes and reproductive consequences in greater snow geese (Anser caerulescens atlanticus) , 2004, Behavioral Ecology and Sociobiology.

[42]  W. Becker Manual of Quantitative Genetics , 1992 .

[43]  F. Pulido,et al.  Phenotypic changes in spring arrival: evolution, phenotypic plasticity, effects of weather and condition , 2007 .

[44]  Theunis Piersma,et al.  Pay-offs and penalties of competing migratory schedules , 2003 .

[45]  J. Takekawa,et al.  Migration and stopover strategies of individual Dunlin along the Pacific coast of North America , 2004 .

[46]  P. Battley,et al.  Carry-over effects and compensation: Late arrival on non-breeding grounds affects wing moult but not plumage or schedules of departing bar-tailed godwits Limosa lapponica baueri , 2012 .

[47]  Charles M. Francis,et al.  The influence of climate on the timing and rate of spring bird migration , 2004, Oecologia.

[48]  Michael R. Dohm,et al.  Repeatability estimates do not always set an upper limit to heritability , 2002 .

[49]  S. Baird,et al.  The birds of North America , 1974 .

[50]  T. Fransson Timing and speed of migration in North and West European populations of Sylvia warblers , 1995 .

[51]  Thomas Alerstam,et al.  Optimal Bird Migration: The Relative Importance of Time, Energy, and Safety , 1990 .

[52]  Vsevolod Afanasyev,et al.  Spring stopover routines in Russian Barnacle Geese Branta leucopsis tracked by resightings and geolocation , 2006 .

[53]  A. Dawson Control of molt in birds: association with prolactin and gonadal regression in starlings. , 2006, General and comparative endocrinology.

[54]  S. Bearhop,et al.  Carry-over effects as drivers of fitness differences in animals. , 2011, The Journal of animal ecology.

[55]  Aevar Petersen,et al.  Tracking of Arctic terns Sterna paradisaea reveals longest animal migration , 2010, Proceedings of the National Academy of Sciences.

[56]  Peter P. Marra,et al.  Linking fluctuations in rainfall to nonbreeding season performance in a long-distance migratory bird, Setophaga ruticilla , 2007 .

[57]  P. Battley,et al.  Breeding latitude drives individual schedules in a trans-hemispheric migrant bird. , 2010, Nature communications.