Springer Nature is making SARS-CoV-2 and COVID-19 research free. View research | View latest news | Sign up for updates

Phenology, variation in habitat use, and daily activity patterns of Eleonora’s falcon overwintering in Madagascar



The vast majority of the global population of Eleonora’s falcon overwinters in Madagascar, where the natural environment is threatened by human-induced habitat changes, particularly intensive forest degradation.


We described Eleonora’s falcon phenology and habitat use based on fine-scale telemetry data and field surveys, and investigated which environmental parameters shape the observed patterns, to obtain a better insight into the species’ ecological requirements during the wintering season.


We used high resolution GPS telemetry and remotely sensed data to establish bird–habitat associations and investigate spatiotemporal activity. We also verified habitat composition through ground surveys.


Eleonora’s falcon exploits a variety of habitats, exhibiting a distinct phenological pattern in their use. The species exhibits high site fidelity, moving progressively from more open areas towards landscapes with denser tree cover, possibly as a response to spatiotemporal patterns in food abundance. Time budget analysis revealed that Eleonora’s falcon dedicates just about 15% of its daily activity to foraging, which is performed almost exclusively during daylight hours, with a greater tendency to forage at higher elevations with denser tree cover. On-site assessment of habitat composition revealed that existing habitat maps overlook small fragments of habitat and land use. However, the species’ overall preference for humid forest suggests ongoing deforestation would adversely affect the availability of preferred habitat.


Our findings suggest that GPS logger technology, when combined with ground surveys, can enhance our understanding of long-distance migratory bird species’ ecology at finer and more ecologically relevant scales.

This is a preview of subscription content, log in to check access.

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5


  1. Agafonkin V, Thieurmel B (2018) Suncalc: compute sun position, sunlight phases, moon position and lunar phase. R package version 0.4

  2. Ausden M (2004) Habitat management. In: Sutherland WJ, Newton I, Green R (eds) Bird ecology and conservation: a handbook of techniques. Oxford University Press, Oxford, pp 329–369

  3. Barov B, Derhé M (2011) Review of the implementation of species action plans of threatened birds in the European Union (2004–2010). BirdLife International for the European Commission

  4. Barron DG, Brawn JD, Weatherhead PJ (2010) Meta-analysis of transmitter effects on avian behaviour and ecology. Methods Ecol Evol 1:180–187

  5. Beyer HL (2012) Geospatial modelling environment Accessed 19 Apr 2019

  6. Bibby CJ, Burgess ND, Hill DA, Mustoe SH (2000) Bird census techniques, 2nd edn. Academic, London

  7. Bildstein KL (2006) Migrating raptors of the world: their ecology and conservation. Cornell University Press, Ithaca

  8. Brandt MJ, Cresswell W (2009) Diurnal foraging routines in a tropical bird, the rock finch Lagonosticta sanguinodorsalis: how important is predation risk? J Avian Biol 40:90–94

  9. Brown DR, Sherry TW (2006) Food supply controls the body condition of a migrant bird wintering in the tropics. Oecologia 149:22–32

  10. Buij R, Gschweng M (2017) Nocturnal hunting by Eleonora’s Falcons Falco eleonorae on their breeding and non-breeding grounds. Acta Ornithol 52:35–49

  11. Calenge C (2006) The package “adehabitat” for the R software: a tool for the analysis of space and habitat use by animals. Ecol Model 197:516–519

  12. Catry I, Dias MP, Catry T, Afanasyev V, Fox J, Franco A, Sutherland WJ (2011) Individual variation in migratory movements and winter behaviour of Iberian Lesser Kestrels Falco naumanni revealed by geolocators. Ibis 153:154–164

  13. CLS (2013) Argos user’s manual. Collecte Localisation Satellites, Paris

  14. Cody ML (1985) Habitat selection in birds. Academic, San Diego

  15. Craig RJ, Mitchell ES, Mitchell JE (1988) Time and energy budgets of bald eagles wintering along the Connecticut River. J Field Ornithol 59:22–32

  16. Cramp S, Simmons K, Gillmor R, Hollom P, Hudson R, Nicholson E, Ogilvie M, Olney P, Roselaar C, Voous K, Wallace D, Wattel J (1980) Handbook of the birds of Europe, the Middle East and North Africa. In: The birds of the Western Palearctic. Hawks to bustards, vol 2. Oxford University Press, London

  17. Cresswell W (2014) Migratory connectivity of Palaearctic-African migratory birds and their responses to environmental change: the serial residency hypothesis. Ibis 156:493–510

  18. Del Hoyo J, Elliott A, Sargatal J (1994) Handbook of the birds of the World. New world vultures to guinea fowl, vol II. Lynx Edicions, Barcelona

  19. ESRI (2012) ArcGIS ver. 10.1. ESRI, Inc., Redlands

  20. Fieberg J, Kochanny CO (2005) Quantifying home-range overlap: the importance of the utilization distribution. J Wildl Manag 69:1346–1359

  21. Garshelis DL (2000) Delusions in habitat evaluation: Measuring use, selection, and importance. In: Boitani L, Fuller TK (eds) Research techniques in animal ecology: controversies and consequences. Columbia University Press, New York, pp 111–164

  22. Ghimire CP, Bruijnzeel LA, Lubczynski MW, Ravelona M, Zwartendijk BW, Van Meerveld HI (2017) Measurement and modelling of rainfall interception by two differently aged secondary forests in upland eastern Madagascar. J Hydrol 545:212–225

  23. Global Forest Watch (2019) World Resources Institute. Accessed 3 Sep 2019

  24. Google, Inc. (2013). Google Earth Pro. Google, Inc.

  25. Gottschalk T, Huettmann F, Ehlers M (2005) Thirty years of analysing and modelling avian habitat relationships using satellite imagery data: a review. Int J Remote Sens 26:2631–2656

  26. Green GM, Sussman RW (1990) Deforestation history of the eastern rain forests of Madagascar from satellite images. Science 248:212–215

  27. Grinand C, Rakotomalala F, Gond V, Vaudry R, Bernoux M, Vieilledent G (2013) Estimating deforestation in tropical humid and dry forests in Madagascar from 2000 to 2010 using multi-date Landsat satellite images and the random forests classifier. Remote Sens Environ 139:68–80

  28. Gschweng M, Kalko EK, Berthold P, Fiedler W, Fahr J (2012) Multi-temporal distribution modelling with satellite tracking data: predicting responses of a long-distance migrant to changing environmental conditions. J Appl Ecol 49:803–813

  29. Guilford T, Åkesson S, Gagliardo A, Holland RA, Mouritsen H, Muheim R, Wiltschko R, Wiltschko W, Bingman VP (2011) Migratory navigation in birds: new opportunities in an era of fast-developing tracking technology. J Exp Biol.

  30. Guilford T, Meade J, Freeman R, Biro D, Evans T, Bonadonna F, Boyle D, Roberts S, Perrins C (2008) GPS tracking of the foraging movements of Manx Shearwaters Puffinus puffinus breeding on Skomer Island, Wales. Ibis 150:462–473

  31. Harper GJ, Steininger MK, Tucker CJ, Juhn D, Hawkins F (2007) Fifty years of deforestation and forest fragmentation in Madagascar. Environ Conserv 34:325–333

  32. Jacobsen LB, Jensen NO, Willemoes M, Hansen L, Desholm M, Fox AD, Tøttrup AP, Thorup K (2017) Annual spatiotemporal migration schedules in three larger insectivorous birds: European nightjar, common swift and common cuckoo. Anim Biotelem.

  33. Kassara C, Fric J, Gschweng M, Sfenthourakis S (2012) Complementing the puzzle of Eleonora’s Falcon (Falco eleonorae) migration: new evidence from an eastern colony in the Aegean Sea. J Ornithol 153:839–848

  34. Kassara C, Fric J, Sfenthourakis S (2014) Distribution modelling of Eleonora’s falcon Falco eleonorae Géné, 1839 occurrence in its wintering grounds: a niche-based approach with satellite telemetry data. Bird Conserv Int 24:100–113

  35. Kassara C, Gangoso L, Mellone U, Piasevoli G, Hadjikyriakou TG, Tsiopelas N, Giokas S, López-López P, Urios V, Figuerola J, Silva R, Bouten W, Kirschel ANG, Virani MZ, Fiedler W, Berthold P, Gschweng M (2017) Current and future suitability of wintering grounds for a long-distance migratory raptor. Sci Rep.

  36. Kenward RE (2000) A manual for wildlife radio tagging. Academic, London

  37. Kerlinger P (2008) How birds migrate. Stackpole Books, Mechanicsburg

  38. Kotzerka J, Garthe S, Hatch SA (2010) GPS tracking devices reveal foraging strategies of Black-legged Kittiwakes. J Ornithol 151:459–467

  39. Kremen C, Razafimahatratra V, Guillery RP, Rakotomalala J, Weiss A, Ratsisompatrarivo J (1999) Designing the Masoala National Park in Madagascar based on biological and socioeconomic data. Conserv Biol 13:1055–1068

  40. Kuznetsova A, Brockhoff PB, Christensen RHB (2017) lmerTest package: tests in linear mixed effects models. J Stat Softw 82:1–26

  41. Lihu X, Jianjian L, Chunfu T, Wenshan H (2007) Foraging area and hunting technique selection of common kestrel (Falco tinnunculus) in winter: the role of perch sites. Acta Ecol Sin 27:2160–2166

  42. Limiñana R, Romero M, Mellone U, Urios V (2012) Mapping the migratory routes and wintering areas of Lesser Kestrels Falco naumanni: new insights from satellite telemetry. Ibis 154:389–399

  43. Mack E, Firbank L, Bellamy P, Hinsley S, Veitch N (1997) The comparison of remotely sensed and ground-based habitat area data using species-area models. J Appl Ecol 34:1222–1228

  44. Mellone U, López-López P, Limiñana R, Urios V (2012) Wintering habitats of Eleonora’s Falcons Falco eleonorae in Madagascar. Bird Study 59:29–36

  45. Meyburg B, Howey PW, Meyburg C, Dietrich Fiuczynski K (2011) Two complete migration cycles of an adult Hobby tracked by satellite. Br Birds 104:2–15

  46. Moat J, Smith PP (2007) Atlas of the vegetation of Madagascar. Royal Botanic Gardens, Kew

  47. NASA (2018) Reverb/ECHO Earth data. Accessed 8 Feb 2018

  48. Newton I (2004) Population limitation in migrants. Ibis 146:197–226

  49. Norris DR, Marra PP, Kyser TK, Sherry TW, Ratcliffe LM (2004) Tropical winter habitat limits reproductive success on the temperate breeding grounds in a migratory bird. Proc R Soc Lond B 271:59–64

  50. Otis DL, White GC (1999) Autocorrelation of location estimates and the analysis of radiotracking data. J Wildl Manag 63:1039–1044

  51. Plumpton DL, Andersen DE (1997) Habitat use and time budgeting by wintering ferruginous hawks. Condor 99:88–93

  52. Pulido F (2007) The genetics and evolution of avian migration. AIBS Bull 57:165–174

  53. Rappole JH, King DI, Diez J (2003) Winter- vs. breeding-habitat limitation for an endangered avian migrant. Ecol Appl 13:735–742

  54. Ristow D (1999) International species action plan Eleonora’s falcon (Falco eleonorae). BirdLife International, Council of Europe, Cambridge

  55. Rosén M, Hedenström A, Badami A, Spina F, Åkesson S (1999) Hunting flight behaviour of the Eleonora’s falcon Falco eleonorae. J Avian Biol 30:342–350

  56. Sanderson FJ, Donald PF, Pain DJ, Burfield IJ, Van Bommel FP (2006) Long-term population declines in Afro-Palearctic migrant birds. Biol Conserv 131:93–105

  57. Sherry TW, Holmes RT (1996) Winter habitat quality, population limitation, and conservation of Neotropical–Nearctic migrant birds. Ecology 77:36–48

  58. Styger E, Rakotondramasy HM, Pfeffer MJ, Fernandes EC, Bates DM (2007) Influence of slash-and-burn farming practices on fallow succession and land degradation in the rainforest region of Madagascar. Agric Ecosyst Environ 119:257–269

  59. Sutherland WJ, Newton I, Green R (2004) Bird ecology and conservation: a handbook of techniques. Oxford University Press, Oxford

  60. Tapia L, Kennedy PL, Mannan RW (2007) Habitat sampling. In: Bird DM, Bildstein KL (eds) Raptor research and management techniques. Hancock House, Surrey, pp 153–169

  61. R Core Team (2018) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna

  62. Thorup K, Alerstam T, Hake M, Kjellén N (2003) Can vector summation describe the orientation system of juvenile ospreys and honey buzzards? An analysis of ring recoveries and satellite tracking. Oikos 103:350–359

  63. USGS (2016) Global multi-resolution terrain data. Accessed 2 Feb 2018

  64. Walter H (1979) Eleonora’s falcon: adaptations to prey and habitat in a social raptor. The University of Chicago Press, Chicago

  65. Wolda H (1978) Seasonal fluctuations in rainfall, food and abundance of tropical insects. J Anim Ecol 47:369–381

  66. Wolda H (1988) Insect seasonality: why? Annu Rev Ecol Evol Syst 19:1–18

  67. Worton BJ (1989) Kernel methods for estimating the utilization distribution in home-range studies. Ecology 70:164–168

  68. Xenophontos M, Blackburn E, Cresswell W (2017) Cyprus wheatears Oenanthe cypriaca likely reach sub-Saharan African wintering grounds in a single migratory flight. J Avian Biol 48:529–535

  69. Zefania S (2001) Observation of sooty and Eleonora’s falcons in Madagascar. Wings over Africa. In: Proceedings of an international seminar on bird migration: research, conservation, education and flight safety, Israel, pp 151–159

  70. Zuur AF, Leno EN, Walker EN, Saveliev AA, Smith GM (2009) Mixed effects models and extensions in ecology with R. Springer, New York

Download references


We would like to thank W. Cresswell and two anonymous reviewers for helpful comments on a previous version of the manuscript. We also thank P. Charilaou, for his support on the project. We are especially grateful to the late A. Crabtree for his invaluable guidance in the field in Cyprus. We would like to thank D. Randrianjafiniasa and R. Loukman for their assistance in the field in Madagascar. Work in Cyprus and in Madagascar was funded by the A. G. Leventis Foundation, The Peregrine Fund, the William A. Burnham Memorial Fund, the Joint Nature Conservation Committee and the Sovereign Base Areas Administration (SBAA). The 2009–2010 Telemetry Project was funded by the A.G. Leventis Foundation through the “Survey and Conservation of Seabirds in Greece” Project, while the 2015–2018 telemetry project was funded by the European Union LIFE Instrument and the Green Fund through the “LIFE ElClimA LIFΕ13 NAT/GR/000909” Project, which also partly-funded fieldwork in Madagascar. A Bird Ringing License and a Research License to attach bird transmitters in Cyprus were obtained from the SBAA Environment Department. The Ministry of Environment and Energy (Greece) kindly granted permission for capturing and tagging Eleonora’s falcon in breeding areas in Greece and all field surveys and bird handling complied with current laws in Greece. Access to National Parks in Madagascar was arranged in cooperation with The Peregrine Fund Madagascar team.

Author information

Correspondence to Alexander N. G. Kirschel.

Additional information

Publisher's Note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Electronic supplementary material

Below is the link to the electronic supplementary material.

Supplementary material 1 (DOCX 396 kb)

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Hadjikyriakou, T.G., Kassara, C., de Roland, L.R. et al. Phenology, variation in habitat use, and daily activity patterns of Eleonora’s falcon overwintering in Madagascar. Landscape Ecol 35, 159–172 (2020).

Download citation


  • Falco eleonorae
  • Telemetry
  • Ground-truthing
  • Fidelity
  • Humid forest
  • NDVI