Kronfeld-Schor, N. & Dayan, T. Partitioning of time as an ecological resource. Annu. Rev. Ecol. Evol. Syst. 34, 153–181 (2003).
Google Scholar
Tauber, E. & Kyriacou, C. P. Review: Genomic approaches for studying biological clocks. Funct. Ecol. 22, 19–29 (2008).
White, E. R. & Hastings, A. Seasonality in ecology: Progress and prospects in theory. Ecol. Complex. 44, 100867 (2020).
Google Scholar
Ko, C. H. & Takahashi, J. S. Molecular components of the mammalian circadian clock. Hum. Mol. Genet. 15, R271–R277 (2006).
Google Scholar
Cassone, V. M. Avian circadian organization: A chorus of clocks. Front. Neuroendocrinol. 35, 76–88 (2014).
Google Scholar
Kyriacou, C. P., Peixoto, A. A., Sandrelli, F., Costa, R. & Tauber, E. Clines in clock genes: Fine-tuning circadian rhythms to the environment. Trends Genet. 24, 124–132 (2008).
Google Scholar
Partch, C. L., Green, C. B. & Takahashi, J. S. Molecular architecture of the mammalian circadian clock. Trends Cell Biol. 24, 90–99 (2014).
Google Scholar
Helm, B. et al. Two sides of a coin: ecological and chronobiological perspectives of timing in the wild. Philos. Trans. R. Soc. B Biol. Sci. 372, 20160246 (2017).
Google Scholar
Kalmbach, D. A. et al. Genetic basis of chronotype in humans: Insights from three landmark GWAS. Sleep https://doi.org/10.1093/sleep/zsw048 (2017).
Google Scholar
Takahashi, J. S., Shimomura, K. & Kumar, V. Searching for genes underlying behavior: Lessons from circadian rhythms. Science 322, 909–912 (2008).
Google Scholar
Yoshimura, T. et al. Molecular analysis of avian circadian clock genes11Published on the World Wide Web on 23 May 2000. Mol. Brain Res. 78, 207–215 (2000).
Google Scholar
Gekakis, N. et al. Role of the CLOCK Protein in the Mammalian circadian mechanism. Science 280, 1564–1569 (1998).
Google Scholar
Saleem, Q., Anand, A., Jain, S. & Brahmachari, S. K. The polyglutamine motif is highly conserved at the Clock locus in various organisms and is not polymorphic in humans. Hum. Genet. 109, 136–142 (2001).
Google Scholar
Darlington, T. K. et al. Closing the circadian loop: CLOCK-induced transcription of its own inhibitors per and tim. Science 280, 1599–1603 (1998).
Google Scholar
King, D. P. et al. Positional cloning of the mouse circadian clock gene. Cell 89, 641–653 (1997).
Google Scholar
Follett, B. Rhythms and photoperiodism in birds. Biological rhythms and photoperiodism in plants (1998).
Hazlerigg, D. G. & Wagner, G. C. Seasonal photoperiodism in vertebrates: from coincidence to amplitude. Trends Endocrinol. Metab. 17, 83–91 (2006).
Google Scholar
Gwinner, E. Circadian and circannual programmes in avian migration. J. Exp. Biol. 199, 39–48 (1996).
Google Scholar
Stirland, J. A., Mohammad, Y. N. & Loudon, A. S. I. A mutation of the circadian timing system (tau gene) in the seasonally breeding Syrian hamster alters the reproductive response to photoperiod change. Proc. R Soc. London Ser. B Biol. Sci. 263, 345–350 (1996).
Google Scholar
Bradshaw, W. E. & Holzapfel, C. M. Evolution of animal photoperiodism. Annu. Rev. Ecol. Evol. Syst. 38, 1–25 (2007).
Google Scholar
Graham, J. L., Cook, N. J., Needham, K. B., Hau, M. & Greives, T. J. Early to rise, early to breed: A role for daily rhythms in seasonal reproduction. Behav. Ecol. 28, 1266–1271 (2017).
Google Scholar
Rittenhouse, J. L., Robart, A. R. & Watts, H. E. Variation in chronotype is associated with migratory timing in a songbird. Biol. Lett. 15, 20190453 (2019).
Google Scholar
O’Malley, K. G., Ford, M. J. & Hard, J. J. Clock polymorphism in Pacific salmon: Evidence for variable selection along a latitudinal gradient. Proc. R. Soc. B Biol. Sci. 277, 3703–3714 (2010).
Google Scholar
O’Malley, K. G. & Banks, M. A. A latitudinal cline in the Chinook salmon (Oncorhynchus tshawytscha) Clock gene: Evidence for selection on PolyQ length variants. Proc. R. Soc. B Biol. Sci. 275, 2813–2821 (2008).
Google Scholar
Peterson, M. P. et al. Variation in candidate genes CLOCK and ADCYAP1 does not consistently predict differences in migratory behavior in the songbird genus Junco. F1000Research 2, 115 (2013).
Google Scholar
Saino, N. et al. Polymorphism at the Clock gene predicts phenology of long-distance migration in birds. Mol. Ecol. 24, 1758–1773 (2015).
Google Scholar
Saino, N. et al. Timing of molt of barn swallows is delayed in a rare Clock genotype. PeerJ 1, e17 (2013).
Google Scholar
Johnsen, A. et al. Avian Clock gene polymorphism: Evidence for a latitudinal cline in allele frequencies. Mol. Ecol. 16, 4867–4880 (2007).
Google Scholar
Liedvogel, M., Szulkin, M., Knowles, S. C. L., Wood, M. & Sheldon, B. C. Phenotypic correlates of Clock gene variation in a wild blue tit population: Evidence for a role in seasonal timing of reproduction. Mol. Ecol. 18, 2444–2456 (2009).
Google Scholar
Caprioli, M. et al. Clock gene variation is associated with breeding phenology and maybe under directional selection in the migratory barn swallow. PLoS ONE 7, e35140 (2012).
Google Scholar
Dor, R. et al. Clock gene variation in Tachycineta swallows. Ecol. Evol. 2, 95–105 (2012).
Google Scholar
Dor, R. et al. Low variation in the polymorphic Clock gene poly-Q region despite population genetic structure across barn swallow (Hirundo rustica) populations. PLoS ONE 6, e28843 (2011).
Google Scholar
O’Brien, C. et al. Geography of the circadian gene clock and photoperiodic response in western North American populations of the three-spined stickleback Gasterosteus aculeatus. J. Fish Biol. 82, 827–839 (2013).
Google Scholar
Mueller, J. C., Pulido, F. & Kempenaers, B. Identification of a gene associated with avian migratory behaviour. Proc. R. Soc. B Biol. Sci. 278, 2848–2856 (2011).
Google Scholar
Liedvogel, M. & Sheldon, B. C. Low variability and absence of phenotypic correlates of Clock gene variation in a great tit Parus major population. J. Avian Biol. 41, 543–550 (2010).
Google Scholar
Lugo-Ramos, J. S., Delmore, K. E. & Liedvogel, M. Candidate genes for migration do not distinguish migratory and non-migratory birds. J. Comp. Physiol. A 203, 383–397 (2017).
Google Scholar
Majoy, S. B. & Heideman, P. D. Tau differences between short-day responsive and short-day nonresponsive white-footed mice (Peromyscus leucopus) do not affect reproductive photoresponsiveness. J. Biol. Rhythms 15, 501–513 (2000).
Google Scholar
O’Brien, C. et al. Geography of the circadian gene clock and photoperiodic response in western North American populations of the threespine stickleback Gasterosteus aculeatus. J. Fish Biol. 82, 827–839 (2013).
Google Scholar
Contina, A., Bridge, E. S., Ross, J. D., Shipley, J. R. & Kelly, J. F. Examination of clock and Adcyap1 gene variation in a neotropical migratory passerine. PLoS ONE 13, e0190859 (2018).
Google Scholar
Herzog, E. D. Neurons and networks in daily rhythms. Nat. Rev. Neurosci. 8, 790–802 (2007).
Google Scholar
Chahad-Ehlers, S. et al. Expanding the view of clock and cycle gene evolution in Diptera. Insect Mol. Biol. 26, 317–331 (2017).
Google Scholar
Denlinger, D. L., Hahn, D. A., Merlin, C., Holzapfel, C. M. & Bradshaw, W. E. Keeping time without a spine: What can the insect clock teach us about seasonal adaptation?. Philos. Trans. R. Soc. B Biol. Sci. 372, 20160257 (2017).
Google Scholar
van Noordwijk, A. J. et al. A framework for the study of genetic variation in migratory behaviour. J .Ornithol. 147, 221–233 (2006).
Google Scholar
Newton, I. The Migration Ecology of Birds (Academic Press, 2008).
Gohli, J., Lifjeld, J. T. & Albrecht, T. Migration distance is positively associated with sex-linked genetic diversity in passerine birds. Ethol. Ecol. Evol. 28, 42–52 (2016).
Google Scholar
Bazzi, G. et al. Clock gene polymorphism, migratory behaviour and geographic distribution: A comparative study of trans-Saharan migratory birds. Mol. Ecol. 25, 6077–6091 (2016).
Google Scholar
Doren, B. M. V., Liedvogel, M. & Helm, B. Programmed and flexible: Long-term Zugunruhe data highlight the many axes of variation in avian migratory behaviour. J. Avian Biol. 48, 155–172 (2017).
Google Scholar
Helm, B., Gwinner, E. & Trost, L. Flexible seasonal timing and migratory behavior: Results from stonechat breeding programs. Ann. N. Y. Acad. Sci. 1046, 216–227 (2005).
Google Scholar
Helm, B. & Gwinner, E. Migratory restlessness in an equatorial nonmigratory bird. PLoS Biol. 4, e110 (2006).
Google Scholar
Helm, B. Geographically distinct reproductive schedules in a changing world: Costly implications in captive Stonechats. Integr Comp Biol 49, 563–579 (2009).
Google Scholar
Dhondt, A. A. Variations in the number of overwintering stonechats possibly caused by natural selection. Ringing Migr. 4, 155–158 (1983).
Google Scholar
Brown, C. R. & Brown, M. B. Weather-mediated natural selection on arrival time in cliff swallows (Petrochelidon pyrrhonota). Behav. Ecol. Sociobiol. 47, 339–345 (2000).
Google Scholar
GOUDET, J. FSTAT, a program to estimate and test gene diversities and fixation indices, version 2.9.3. http://www2.unil.ch/popgen/softwares/fstat.htm (2001).
Van Doren, B. M. et al. Correlated patterns of genetic diversity and differentiation across an avian family. Mol. Ecol. 26, 3982–3997 (2017).
Google Scholar
Illera, J. C., Richardson, D. S., Helm, B., Atienza, J. C. & Emerson, B. C. Phylogenetic relationships, biogeography and speciation in the avian genus Saxicola. Mol. Phylogenet. Evol. 48, 1145–1154 (2008).
Google Scholar
Illera, J. C. & Díaz, M. Reproduction in an endemic bird of a semiarid island: A food-mediated process. J. Avian Biol. 37, 447–456 (2006).
Google Scholar
Illera, J. C. & Díaz, M. Site fidelity in the Canary Islands stonechat Saxicola dacotiae in relation to spatial and temporal patterns of habitat suitability. Acta Oecol. 34, 1–8 (2008).
Google Scholar
Gwinner, E. & Dittami, J. Endogenous reproductive rhythms in a tropical bird. Science 249, 906–908 (1990).
Google Scholar
Dittami, J. & Gwinner, E. Annual cycles in the African stonechat Saxicola torquata axillaris and their relationship to environmental factors. J. Zool. 207, 357–370 (1985).
Google Scholar
Gwinner, E. Circannual rhythms in tropical and temperate-zone stonechats: A comparison of properties under constant conditions. Ökologie der Vögel 13, 5–14 (1991).
Gwinner, E. Circannual Rhythms: Endogenous Annual Clocks in the Organization of Seasonal Processes (Springer, 2012).
Helm, B., Fiedler, W. & Callion, J. Movements of European stonechats Saxicola torquata according to ringing recoveries. ARDEA-WAGENINGEN- 94, 33 (2006).
Opaev, A., Red’kin, Y., Kalinin, E. & Golovina, M. Species limits in Northern Eurasian taxa of the common stonechats, Saxicola torquatus complex (Aves: Passeriformes, Muscicapidae). Vertebr.ate Zool. 68, 199 (2018).
Gwinner, E. & Czeschlik, D. On the significance of spring migratory restlessness in caged birds. Oikos 30, 364–372 (1978).
Google Scholar
Krist, M., Munclinger, P., Briedis, M. & Adamík, P. The genetic regulation of avian migration timing: combining candidate genes and quantitative genetic approaches in a long-distance migrant. Oecologia https://doi.org/10.1007/s00442-021-04930-x (2021).
Google Scholar
Berthold, P. & Pulido, F. Heritability of migratory activity in a natural bird population. Proc. R. Soc. London Ser. B Biol. Sci. 257, 311–315 (1994).
Google Scholar
Pulido, F. & Berthold, P. Current selection for lower migratory activity will drive the evolution of residency in a migratory bird population. Proc. Natl. Acad. Sci. 107, 7341–7346 (2010).
Google Scholar
Liedvogel, M. & Lundberg, M. The Genetics of Migration. In Animal Movement Across Scales (eds Hansson, L.-A. & Åkesson, S.) 219–231 (Oxford University Press, 2014). https://doi.org/10.1093/acprof:oso/9780199677184.003.0012.
Google Scholar
Åkesson, S. & Helm, B. Endogenous programs and flexibility in bird migration. Front. Ecol. Evol. 8, 78 (2020).
Google Scholar
Stevenson, T. J. & Kumar, V. Neural control of daily and seasonal timing of songbird migration. J. Comp. Physiol. A 203, 399–409 (2017).
Google Scholar
Verhagen, I. et al. Genetic and phenotypic responses to genomic selection for timing of breeding in a wild songbird. Funct. Ecol. 33, 1708–1721 (2019).
Google Scholar
Helm, B. & Gwinner, E. Timing of Postjuvenal molt in African (Saxicola Torquata Axillaris) and European (Saxicola Torquata Rubicola) stonechats: Effects of genetic and environmental factors. Auk 116, 589–603 (1999).
Google Scholar
Zink, R. M., Pavlova, A., Drovetski, S., Wink, M. & Rohwer, S. Taxonomic status and evolutionary history of the Saxicola torquata complex. Mol. Phylogenet. Evol. 52, 769–773 (2009).
Google Scholar
Flinks, H. & Pfeifer, F. Brutzeit, Gelegegröße und Bruterfolg beim Schwarzkehlchen (Saxicola torquata). Charadrius 23, 128–140 (1987).
Urquhart, E. Stonechats (Christopher Helm, 2002).
Glutz von Blotzheim, U. Bauer Handbuch der Vögel Mitteleuropas KM: Bd. 11. Aula, Wiesbaden (1988).
Yamaura, Y. et al. Tracking the Stejneger’s stonechat Saxicola stejnegeri along the East Asian-Australian Flyway from Japan via China to southeast Asia. J. Avian Biol. 48, 197–202 (2017).
Google Scholar
Gwinner, E., Neusser, V., Engl, D., Schmidl, D. & Bals, L. Haltung, Zucht und Eiaufzucht afrikanischer und europäischer Schwarzkehlchen Saxicola torquata. Gefiederte Welt 111, 118–120 (1987).
Flinks, H., Helm, B. & Rothery, P. Plasticity of moult and breeding schedules in migratory European Stonechats Saxicola rubicola. Ibis 150, 687–697 (2008).
Google Scholar
Humphrey, P. S. & Parkes, K. C. An approach to the study of molts and plumages. Auk 76, 1–31 (1959).
Google Scholar
Berthold, P. Bird Migration: A General Survey (Oxford University Press, 2001).
RStudio | Open source & professional software for data science teams. https://rstudio.com/.
R Core Team. R: A language and environment for statistical computing. (R Foundation for Statistical Computing, 2013).
Bates, D., Mächler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. http://arxiv.org/abs/1406.5823 (2014).
Lüdecke, D. & Lüdecke, M. D. Package ‘sjPlot’. (2015).
del Hoyo, J., Elliott, A., Sargatal, J., Christie, D. A. & de Juana, E. Handbook of the Birds of the World Alive (Lynx Edicions, 2018).
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