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Foraging ecology of Red-billed Tropicbird Phaethon aethereus in the Caribbean during early chick rearing revealed by GPS tracking



1Marine Animal Ecology Group, Wageningen University, Droevendaalsesteeg 1, 6708 PB, Wageningen, The Netherlands (hannah.madden@wur.nl)
2Caribbean Netherlands Science Institute, PO Box 65, St. Eustatius, Caribbean Netherlands, NIOZ Royal Netherlands Institute for Sea Research, and Utrecht University, PO Box 59, 1790 AB Den Burg, Texel, The Netherlands
3Department of Forestry and Environmental Conservation, Clemson University, Clemson, South Carolina, USA
4US Geological Survey - South Carolina Cooperative Fish and Wildlife Research Unit and Department of Forestry and Environmental Conservation, Clemson University, Clemson, South Carolina, USA

Received 20 January 2022, accepted 31 May 2022

Date Published: 2022/10/15
Date Online: 2022/09/22

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Appendix 1


MADDEN, H., SATGÉ, Y., WILKINSON, B. & JODICE, P.G.R. 2022. Foraging ecology of Red-billed Tropicbird Phaethon aethereus in the Caribbean during early chick rearing revealed by GPS tracking. Marine Ornithology 50: 165-175.

Key words: diet, EEZ, foraging area, MPA, oceanographic variables, Phaethon aethereus, seabird


Investigating the foraging patterns of tropical seabirds can provide important information about their ocean habitat affinities as well as prey choice. Foraging studies of Red-billed Tropicbird Phaethon aethereus populations in the Caribbean are lacking. We sought to rectify this by opportunistically sampling regurgitates at nest sites on the island of St. Eustatius, Lesser Antilles, and by linking the GPS tracks of foraging adults to remotely sensed environmental variables. Diet samples were dominated by Exocoetidae (59.5%) and Belonidae (14.9%), although we were unable to identify 25.5% of samples due to digestion. Tropicbirds nesting on St. Eustatius exhibited diurnal foraging patterns, foraged in deeper waters with higher chlorophyll concentration, and consumed fewer Exocoetidae species compared to travelling behaviour. The maximum distance travelled from the colony was 953.7 km, with an average trip length of 176.8 (± 249.8) km. The biologged birds crossed multiple exclusive economic zones and marine protected areas, and on that basis, we suggest that efforts to protect and conserve this species may require transboundary collaboration throughout the wider Caribbean.


AINLEY, D.G. 1977. Feeding methods in seabirds: a comparison of polar and tropical nesting communities in the eastern Pacific Ocean. In: LLANO, G.A. (Ed.). Adaptations within Antarctic Ecosystems. Proceedings of the 3rd SCAR Symposium on Antarctic Biology. Washington, USA: Smithsonian Institute.

AKAIKE, H. 1974. A new look at the statistical model identification. IEEE Transactions on Automatic Control 19: 716-723.

ALONSO, H., GRANADEIRO, J.P., DIAS, M.P., CATRY, T. & CATRY, P. 2018. Fine-scale tracking and diet information of a marine predator reveals the origin and contrasting spatial distribution of prey. Progress in Oceanography 162: 1-12.

ARNOLD, T.W. 2010. Uninformative parameters and model selection using Akaike's Information Criterion. The Journal of Wildlife Management 74: 1175-1178.

AU, D.W.K. & PITMAN, R.L. 1986. Seabird interactions with dolphins and tuna in the eastern tropical Pacific. The Condor 88: 304−317.

BALLANCE, L.T. & PITMAN, R.L. 1999. Foraging ecology of tropical seabirds. In: ADAMS, N.J. & SLOTOW, R.H. (Eds.) Proceedings of the 22nd International Ornithological Congress. Durban, South Africa: BirdLife South Africa Johannesburg.

BALLANCE, L.T., PITMAN, R.L. & REILLY, S.B. 1997. Seabird community structure along a productivity gradient: importance of competition and energetic constraint. Ecology 78: 1502−1518.

BARTOŃ, K. 2016. MuMIn: Multi-Model Inference. R package, version 1.43.17. [Manual accessed at https://CRAN.R-project.org/package=MuMIn on 05 May 2020.]

BATES, D., MÄCHLER, M., BOLKER, B. & WALKER, S. 2015. Fitting linear mixed-effects models using lme4. Journal of Statistical Software 67: 1-48. doi:10.18637/jss.v067.i01

BEAL, M., OPPEL, S. & DIAS, M. 2020. track2KBA: Identifying Important Areas from Tagged-animal Tracking Data. R package version [Manual accessed at https://github.com/BirdLifeInternational/track2kba on 31 March 2020.]

BIRDLIFE INTERNATIONAL. 2020. Important Bird Areas factsheet: Boven, Sint Eustatius. Cambridge, UK: Birdlife International. [Accessed at http://www.birdlife.org/ on 23 October 2020.]

BIRDLIFE INTERNATIONAL. 2021. Species factsheet: Phaethon aethereus. Cambridge, UK: Birdlife International. [Accessed at http://www.birdlife.org/ on 22 April 2021.]

BOEKEN, M. 2016. Breeding Success of Red-Billed Tropicbirds Phaethon aethereus on the Caribbean Island of Saba. Ardea 104: 263-271. doi:10.5253/arde.v104i3.a8

BURNHAM, K.P. & ANDERSON, D.R. 2002. Model Selection and Multimodel Inference: A Practical Information-Theoretic Approach, 2nd Edition. New York, USA: Springer.

CASTILLO-GUERRERO, J.A., GUEVARA-MEDINA, M.A. & MELLINK, E. 2011. Breeding ecology of the Red-Billed Tropicbird Phaethon aethereus under contrasting environmental conditions in the Gulf of California. Ardea 99: 61-71. doi:10.5253/078.099.0108

CATRY, T., RAMOS, J.A., JAQUEMET, S. ET AL. 2009a. Comparative foraging ecology of a tropical seabird community of the Seychelles, western Indian Ocean. Marine Ecology Progress Series 374: 259−272.

CATRY, T., RAMOS, J.A., LE CORRE, M. & PHILLIPS, R.A. 2009b. Movements, at-sea distribution and behaviour of a tropical pelagic seabird: the Wedge-tailed Shearwater in the western Indian Ocean. Marine Ecology Progress Series 391: 231−242.

COOK, T.R., CHEREL, Y. & TREMBLAY, Y. 2006. Foraging tactics of chick-rearing Crozet Shags: individuals display repetitive activity and diving patterns over time. Polar Biology 29: 562-569.

DIAMOND, A.W. 1975. The biology of tropicbirds at Aldabra Atoll, Indian Ocean. The Auk 92: 16-39.

DIOP, N., ZANGO, L., BEARD, A. ET AL. 2018. Foraging ecology of tropicbirds breeding in two contrasting marine environments in the tropical Atlantic. Marine Ecology Progress Series 607: 221-236.

DODGE, S., BOHRER, G., WEINZIERL, R. ET AL. 2013. The environmental-data automated track annotation (Env-DATA) system: linking animal tracks with environmental data. Movement Ecology 1: 3.

FAUCHALD, P., ERIKSTAD, K.E. & SKARSFJORD, H. 2000. Scale-dependent predator-prey interactions: The hierarchical spatial distribution of seabirds and prey. Ecology 81: 773-783.

FLANDERS MARINE INSTITUTE. 2020. Union of the ESRI Country shapefile and the Exclusive Economic Zones (version 3). Ostend, Belgium: Vlaams Instituut voor de Zee. doi:10.14284/403 [Accessed at https://www.marineregions.org/ on 24 November 2020.]

FLEISHMAN, A.B., ORBEN, R.A. & GILMOUR, M.E. 2019. trakR: Basic animal tracking data analysis tools. Version 0.0.9. GitHub repository. [Accessed at https://github.com/abfleishman/trakR on 23 April 2021.] doi:10.5281/zenodo.4391403

FURNESS, B.L., LAUGKSCH, R.C. & DUFFY, D.C. 1984. Cephalopod beaks and studies of seabird diets. The Auk 101: 619-620.

GEBCO BATHYMETRIC COMPILATION GROUP. 2020. The GEBCO 2020 Grid - a continuous terrain model of the global oceans and land. London, UK: British Oceanographic Data Centre, National Oceanography Centre, Natural Environment Research Council. doi:10.5285/a29c5465-b138-234d-e053-6c86abc040b9

GRÉMILLET, D. & BOULINIER, T. 2009. Spatial ecology and conservation of seabirds facing global climate change: a review. Marine Ecology Progress Series 391: 121-137.

HARRISON, C.S., HIDA, T.S. & SEKI, M.P. 1983. Hawaiian seabird feeding ecology. Wildlife Monographs 85: 3-71.

HEBSHI, A.J. 2008. The Importance of Food and Scale in the Ecology of Tropical Seabirds. PhD dissertation. Honolulu, USA: University of Hawaii at Manoa.

HENNICKE, J.C. & WEIMERSKIRCH, H. 2014. Coping with variable and oligotrophic tropical waters: foraging behaviour and flexibility of the Abbott's Booby Papasula abbotti. Marine Ecology Progress Series 499: 259-273.

HUMANN, P. & DELOACH, N. 2002. Reef Fish Identification: Florida, Caribbean and Bahamas, 3rd Edition. Jacksonville, USA: New World Publications Inc.

HUNT, G.L., JR. & SCHNEIDER, D.C. 1987. Scale-dependent processes in the physical and biological environment of marine birds. In: CROXALL, J.P. (Ed.). Seabirds: Feeding Ecology and Role in Marine Ecosystems. London, UK: Cambridge University Press.

JAQUEMET, S., LE CORRE, M., MARSAC, F., POTIER, M. & WEIMERSKIRCH, H. 2005. Foraging habitats of the seabird community of Europa Island (Mozambique Channel). Marine Biology 147: 573-582.

JODICE, P.G.R., RONCONI, R.A., RUPP, E., WALLACE, G.E. & SATGÉ, Y. 2015. First satellite tracks of the Endangered Black-capped Petrel. Endangered Species Research 29: 23-33. doi:10.3354/esr00697

JODICE, P.G.R. & SURYAN, R.M. 2010. The transboundary nature of seabird ecology. In: TROMBULAK, S.C. & BALDWIN, R.F. (Eds.). Landscape-scale Conservation Planning. Dordrecht, The Netherlands: Springer.

JOVANI, R., LASCELLES, B., GARAMSZEGI, L.Z., MAVOR, R., THAXTER, C.B. & ORO, D. 2016. Colony size and foraging range in seabirds. Oikos 125: 968-974.

KASCHNER, K., KESNER-REYES, K. GARILAO, C. ET AL. 2019. AquaMaps: Predicted range maps for aquatic species. Version 10/2019. [Accessed at www.aquamaps.org on 08 September 2020.]

LAMB, J.S., SATGÉ, Y.G. & JODICE, P.G.R. 2017. Influence of density‐dependent competition on foraging and migratory behavior of a subtropical colonial seabird. Ecology and Evolution 7: 6469-6481.

LEE, D.S. & MACKIN, W.A. 2008. Red-billed Tropicbird. West Indian Breeding Seabird Atlas. Natick, USA: BirdsCaribbean. [Accessed at www.wicbirds.net/rbtr.html on 18 December 2020.]

LEE, D.S. & WALSH-MCGEHEE, M. 2000. Population estimates, conservation concerns, and management of tropicbirds in the western Atlantic. Caribbean Journal of Science 36: 267-279.

LEWALLEN, E.A., BOHONAK, A.J., BONIN, C.A., VAN WIJNEN, A.J., PITMAN, R.L. & LOVEJOY, N.R. 2017. Phylogenetics and biogeography of the Two‐wing Flyingfish (Exocoetidae: Exocoetus). Ecology and Evolution 7: 1751-1761.

LONGHURST, A.R. & PAULY, D. 1987. Ecology of Tropical Oceans. San Diego, USA: Elsevier Inc.

LOWRIE, K., LOWRIE, D. & COLLIER, N. 2012. Seabird Breeding Atlas of the Lesser Antilles. Green Cove Springs, USA: Environmental Protection in the Caribbean.

MACINTYRE, I.G., KINSMAN, D.J.J. & GERMAN, R.C. 1975. Geological reconnaissance survey of Saba Bank, Caribbean Sea. Caribbean Journal of Science 15: 11-20.

MADDEN, H. 2019. Reproductive performance, mate fidelity and nest cavity fidelity in red-billed tropicbirds Phaethon aethereus mesonauta on St. Eustatius, Caribbean Netherlands. Ardea 107: 227-237.

MCDUIE, F. & CONGDON, B.C. 2016. Trans-equatorial migration and non-breeding habitat of tropical shearwaters: implications for modelling pelagic Important Bird Areas. Marine Ecology Progress Series 550: 219-234.

MCDUIE, F., WEEKS, S.J. & CONGDON, B.C. 2018. Oceanographic drivers of near-colony seabird foraging site use in tropical marine systems. Marine Ecology Progress Series 589: 209-225.

MCDUIE, F., WEEKS, S.J., MILLER, M.G.R. & CONGDON, B.C. 2015. Breeding tropical shearwaters use distant foraging sites when self-provisioning. Marine Ornithology 43: 123-129.

MEESTERS, H.W.G., SLIJKERMAN, D.M.E., DE GRAAF, M. & DEBROT, A.O. 2010. Management Plan for the Natural Resources of the EEZ of the Dutch Caribbean. Report no. C100/10. Wageningen, The Netherlands: Institute for Marine Resources & Ecosystem Studies (IMARES).

MONTICELLI, D., RAMOS, J.A. & QUARTLY, G.D. 2007. Effects of annual changes in primary productivity and ocean indices on breeding performance of tropical Roseate Terns in the western Indian Ocean. Marine Ecology Progress Series 351: 273-286.

NAKAGAWA, S. & SCHIELZETH, H. 2013. A general and simple method for obtaining R2 from generalized linear mixed-effects models. Methods in Ecology and Evolution 4: 133-142.

NIETO, P., MÜCHER, C.A., MEESTERS, H.W.G. & CLEVERS, J.G.P.W. 2013. Classifying benthic habitats and deriving bathymetry at the Caribbean Netherlands using multispectral imagery. Case study of St. Eustatius. Alterra report no. 2467. Wageningen, The Netherlands: IMARES.

OPPEL, S., WEBER, S., WEBER, N. ET AL. 2017. Seasonal shifts in foraging distribution due to individual flexibility in a tropical pelagic forager, the Ascension Frigatebird. Marine Ecology Progress Series 585: 199-212.

ORTA, J. 1992. Family Phaethontidae. In: DEL HOYO, J., ELLIOTT, A. & SARGATAL, J. (Eds.) Handbook of the Birds of the World, Volume I: Ostrich to Ducks. Barcelona, Spain: Lynx Edicions.

ORTA, J., JUTGLAR, F., GARCIA, E.F.J. & KIRWAN, G.M. 2020. Red-billed Tropicbird (Phaethon aethereus), version 1.0. In: DEL HOYO, J., ELLIOTT, A., SARGATAL, J., CHRISTIE, D.A. & DE JUANA, E. (Eds.) Birds of the World. Ithaca, USA: Cornell Laboratory of Ornithology. [Accessed at https://doi.org/10.2173/bow.rebtro.01 on 09 April 2020.]

PETERSON, M.S., BROWN-PETERSON, N.J. & FRANKS, J.S. 2007. Large pelagic fishes in the Caribbean Sea and the Gulf of Mexico: Current status and integrated management. Gulf and Caribbean Research 19: 1-3. doi: 10.18785/gcr.1902.01

PHILLIPS, R.A., LEWIS, S., GONZÁLEZ-SOLÍS, J. & DAUNT, F. 2017. Causes and consequences of individual variability and specialization in foraging and migration strategies of seabirds. Marine Ecology Progress Series 578: 117-150.

PIONTKOVSKI, S.A. & WILLIAMS, R. 1995. Multiscale variability of tropical ocean zooplankton biomass. ICES Journal of Marine Science 52: 643-656.

POLI, C.L., HARRISON, A.-L., VALLARINO, A., GERARD, P.D. & JODICE, P.G.R. 2017. Dynamic oceanography determines fine scale foraging behavior of Masked Boobies in the Gulf of Mexico. PLoS One 12: e0178318.

R CORE TEAM. 2020. R: A Language and Environment for Statistical Computing. Vienna, Austria: R Foundation for Statistical Computing.

RICHARDSON, P.L. 1983. Eddy kinetic energy in the North Atlantic from surface drifters. Journal of Geophysical Research - Oceans 88: 4355-4367.

SANTOS, C.D., CAMPOS, L.F.A.S. & EFE, M.A. 2019. Foraging habitat choice of White-tailed Tropicbirds revealed by fine-scale GPS tracking and remote sensing. PeerJ 7: e6261.

SCHAFFNER, F.C. 1988. The Breeding Biology and Energetics of the White-tailed Tropicbird (Phaethon lepturus) at Culebra, Puerto Rico. PhD dissertation. Coral Gables, USA: University of Miami.

SCHAFFNER, F.C. 1991. Nest-site selection and nesting success of White-tailed Tropicbirds (Phaethon lepturus) at Cayo Luís Peña, Puerto Rico. The Auk 108: 911-922. doi:10.1093/auk/108.4.911

SOANES, L.M., BRIGHT, J.A., BOLTON, M., MILLETT, J., MUKHIDA, F. & GREEN, J.A. 2014. Foraging behaviour of Brown Boobies Sula leucogaster in Anguilla, Lesser Antilles: Preliminary identification of at-sea distribution using a time-in-area approach. Bird Conservation International 25: 87-96.

SOANES, L.M., BRIGHT, J.A., BRODIN, G., MUKHIDA, F. & GREEN, J.A. 2015. Tracking a small seabird: first records of foraging movements in the Sooty Tern Onychoprion fuscatus. Marine Ornithology 43: 235-239.

SOANES, L.M., BRIGHT, J.A., CARTER, D. ET AL. 2016. Important foraging areas of seabirds from Anguilla, Caribbean: Implications for marine spatial planning. Marine Policy 70: 85-92.

SOANES, L.M., GREEN, J.A., BOLTON, M., MILLIGAN, G., MUKHIDA, F. & HALSEY, L.G. 2021. Linking foraging and breeding strategies in tropical seabirds. Journal of Avian Biology 52: e02670 doi:10.1111/jav.02670

SOMMERFELD, J. & HENNICKE, J.C. 2010. Comparison of trip duration, activity pattern and diving behaviour by Red-tailed Tropicbirds (Phaethon rubricauda) during incubation and chick-rearing. Emu 110: 78-86.

SPEAR, L.B., AINLEY, D.G. & WALKER, W.A. 2007. Foraging dynamics of seabirds in the eastern Tropical Pacific Ocean. Studies in Avian Biology No. 35. Norman, USA: Cooper Ornithological Society.

STENAPA (ST. EUSTATIUS NATIONAL PARKS FOUNDATION). 2022. Our Parks. St. Eustatius, Dutch Caribbean: St. Eustatius National Parks. [Accessed at https://www.statiapark.org/our-park/ on 03 August 2022.]

STUDWELL, A.J., HINES, E., ELLIOTT, M.L. ET AL. 2017. Modeling nonresident seabird foraging distributions to inform ocean zoning in Central California. PLoS One 12: e0169517.

SUGIURA, N. 1978. Further analysts of the data by Akaike's information criterion and the finite corrections. Communications in Statistics - Theory and Methods 7: 13-26.

TAKARINA, N.D., NURLIANSYAH, W. & WARDHANA, W. 2019. Relationship between environmental parameters and the plankton community of the Batuhideung Fishing Grounds, Pandeglang, Banten, Indonesia. Biodiversitas 20: 171-180.

UNEP-WCMC & IUCN (United Nations Environment Programme World Conservation Monitoring Centre and International Union for Conservation of Nature). 2020. Protected Planet: The World Database on Protected Areas. Cambridge, UK: UNEP-WCMC and IUCN. [Accessed at www.protectedplanet.net on 24 July 2020.]

WAKEFIELD, E.D., PHILLIPS, R.A. & MATTHIOPOULOS, J. 2009. Quantifying habitat use and preferences of pelagic seabirds using individual movement data: a review. Marine Ecology Progress Series 391: 165-182.

WEIMERSKIRCH, H. 2007. Are seabirds foraging for unpredictable resources? Deep-Sea Research Part II 54: 211−223.

WEIMERSKIRCH, H., LE CORRE, M., JAQUEMET, S. & MARSAC, F. 2005a. Foraging strategy of a tropical seabird, the Red-footed Booby, in a dynamic marine environment. Marine Ecology Progress Series 288: 251−261.

WEIMERSKIRCH, H., LE CORRE, M., ROPERT-COUDERT, Y., KATO, A. & MARSAC, F. 2005b. The three-dimensional flight of Red-footed Boobies: Adaptations to foraging in a tropical environment? Proceedings of the Royal Society B 272: 53−61.

WILKINSON, B.P., HAYNES-SUTTON, A.M., MEGGS, L. & JODICE, P.G.R. 2020. High spatial fidelity among foraging trips of Masked Boobies from Pedro Cays, Jamaica. PLoS One 15: e0231654.

WOOD, A.G., NAEF‐DAENZER, B., PRINCE, P.A. & CROXALL, J.P. 2000. Quantifying habitat use in satellite‐tracked pelagic seabirds: application of kernel estimation to albatross locations. Journal of Avian Biology 31: 278−286.

WOOD, S. & SCHEIPL, F. 2020. gamm4: Generalized Additive Mixed Models using 'mgcv' and 'lme4'. R package version 0.2-6. [Manual accessed at https://CRAN.R-project.org/package=gamm4 on 27 May 2020.]

ZAVALA-HIDALGO, J., GALLEGOS-GARCÍA, A., MARTÍNEZ-LÓPEZ, B., MOREY, S.L. & O'BRIEN, J.J., 2006. Seasonal upwelling on the western and southern shelves of the Gulf of Mexico. Ocean Dynamics 56: 333−338.

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