Volume 48, No. 2

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Spatiotemporal characterization of non-breeding Great Shearwaters Ardenna gravis within their wintering range


1Stellwagen Bank National Marine Sanctuary, NOAA National Ocean Service, Scituate, MA 02066, USA *(
2University of Rhode Island, Graduate School of Oceanography, Narragansett, RI 02882, USA
3University of New England, Animal Behavior Program, Department of Psychology, Biddeford, ME 04005, USA
4United States Fish and Wildlife Service, Milbridge, ME 02658, USA
5Boston University Marine Program, Department of Biology, Boston University, Boston, MA 02215, USA


POWERS, K.D., WILEY, D.N., ROBUCK, A.R., OLSON, Z.H., WELCH, L.J., THOMPSON, M.A. & KAUFMAN, L. 2020. Spatiotemporal characterization of non-breeding Great Shearwaters Ardenna gravis within their wintering range. Marine Ornithology 48: 215 - 229

Received 22 January 2020, accepted 17 June 2020

Date Published: 2020/10/15
Date Online: 2020/07/26
Key words: Great Shearwaters, utilization distribution, wintering range, satellite tracking, fecal DNA analysis, sand lance, bycatch necropsies


Movements of Great Shearwaters Ardenna gravis wintering in the Northwest Atlantic showed age-based spatial and temporal flexibility, with foraging tactics linked to a defined physical preference of their primary prey. From 2013 to 2018, we tracked 58 Great Shearwaters during their wintering season using platform terminal transmitters deployed in the same area of the southwest Gulf of Maine. Utilization distributions (UDs) for individual birds were created from convex hulls, which were then combined for spatial and temporal analyses. Of the 95% kernel UDs, 55% were contained within the Gulf of Maine and the remainder extended to the Scotian Shelf off Nova Scotia and the Grand Banks off Newfoundland. Analysis of fecal DNA from tagged birds and others captured with them indicated that Northern sand lance Ammodytes dubius were the primary prey while in the Gulf of Maine. This relationship was supported by the overlap of UDs and sand lance habitat. The spatial occurrence of sand lance from bottomfish trawl survey data demonstrated that these fish preferred shallow water (< 100 m deep) with substrates consisting of high sand content (> 50%) and grain sizes ranging from 0.35-2.00 mm in diameter. These same properties were associated and spatially aligned with the collective 25% kernel UD of Great Shearwaters. Necropsy of bycaught Great Shearwaters from an area that overlapped in space and time with tagged individuals and sand lance habitat demonstrated that most birds (89%) were young (0-2 years), based on gonadal development, molt score, and/or bursa of Fabricius. Coupling demographic information from necropsies with spatial habits and movement timing of tagged birds suggests this region serves as a winter “nursery” for Great Shearwaters.


ANSTEAD, K.A., SCHAFFLER, J.J. & JONES, C.M. 2016. Coast-wide nursery contribution of new recruits to the population of Atlantic Menhaden. Transactions of the American Fisheries Society 145: 627-636. doi:10.1080/00028487.2016.1150345

ASMFC ATLANTIC MENHADEN STOCK ASSESSMENT SUBCOMMITTEE (ASMFC) 2017. 2017 Atlantic Menhaden Stock Assessment Update. Arlington, USA: Atlantic States Marine Fisheries Commission.

BRADLEY, J.S., GUNN, B.M., SKIRA, I.J., MEATHREL, C.E. & WOOLLER, R.D. 1999. Age-dependent prospecting and recruitment to a breeding colony of Short-tailed Shearwaters Puffinus tenuirostris. Ibis 141: 277-285.

BROUGHTON, J.M. 1994. Size of the bursa of Fabricius in relation to gonad size and age in Laysan and Black-footed Albatrosses. The Condor 96: 203-207.

BROWN, R.G.B. 1986. Revised Atlas of Eastern Canadian Seabirds. Dartmouth, Canada: Canadian Wildlife Service, Bedford Institute of Oceanography.

BROWN, R.G.B. 1988. The wing-moult of fulmars and shearwaters (Procellariidae) in Canadian Atlantic waters. The Canadian Field-Naturalist 102: 203-208.

BROWN, R.G.B., BARKER, S.P., GASKIN, D.E. & SANDEMAN, M.R. 1981. The foods of Great and Sooty Shearwaters Puffinus gravis and P. griseus in eastern Canadian waters. Ibis 123: 19-30.

BUGONI, L., NAVES, L.C. & FURNESS, R.W. 2015. Moult of three Tristan da Cunha seabird species sampled at sea. Antarctic Science 27: 240-251. doi:10.1017/S0954102014000583

CAMPIONI, L., DIAS, M.P., GRANADEIRO, J.P. & CATRY, P. 2020. An ontogenetic perspective on migratory strategy of a long-lived pelagic seabird: Timings and destinations change progressively during maturation. Journal of Animal Ecology 89: 29-43. doi:10.1111/1365-2656.13044

CAPORASO, J.G., KUCZYNSKI, J., STOMBAUGH, J. ET AL. 2010. QIIME allows analysis of high-throughput community sequencing data. Nature Methods 7: 335-336.

CARVALHO, P.C. & DAVOREN, G.K. 2019. Associations of non-breeding shearwater species on the northeastern Newfoundland coast. Marine Ecology Progress Series 627: 1-12. doi:10.3354/meps13112

CORTES, V., GARCIA-BARCELONA, S. & GONZÁLEZ-SOLÍS, J. 2018. Sex- and age-biased mortality of three shearwater species in longline fisheries of the Mediterranean. Marine Ecology Progress Series 588: 229-241. doi:10.3354/meps12427

CROXALL, J.P., BUTCHART, S.H.M., LASCELLES, B. ET AL. 2012. Seabird conservation status, threats and priority actions: A global assessment. Bird Conservation International 22: 1-34. doi:10.1017/S0959270912000020

DAVOREN, G.K., MONTEVECCHI, W.A. & ANDERSON, J.T. 2003. Search strategies of a pursuit-diving marine bird and the persistence of prey patches. Ecological Monographs 73: 463-481. doi:10.1890/02-0208

DAVOREN, G.K., PENTON, P., BURKE, C. & MONTEVECCHI, W.A. 2012. Water temperature and timing of capelin spawning determine seabird diets. ICES Journal of Marine Science 69: 1234-1241. doi:10.1093/icesjms/fss032

DEAGLE, B.E., KIRKWOOD, R. & JARMAN, S.N. 2009. Analysis of Australian fur seal diet by pyrosequencing prey DNA in faeces. Molecular Ecololgy 18: 2022-2038.

DIAS, M.P., MARTIN, R., PEARMAIN, E.J. ET AL. 2019. Threats to seabirds: A global assessment. Biological Conservation 237: 525-537.

FADER, G.B., KING, L.H. & JOSENHANS, H.W. 1982. Surficial Geology of the Laurentian Channel and the Western Grand Banks of Newfoundland. Paper 81-22. Ottawa, Canada: Geological Survey of Canada. doi:10.4095/119807

FAUCHALD, P. 2009. Spatial interaction between seabirds and prey: Review and synthesis. Marine Ecology Progress Series 391: 139-151.

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.

FISHPOOL, L.D.C. & EVANS, M.I (Eds.) 2001. Important Bird Areas in Africa and Associated Islands: Priority Sites for Conservation. Birdlife International Conservation Series No. 11. Newbury and Cambridge, UK: Pisces Publications and BirdLife International. 

GETZ, W.M., FORTMANN-ROE, S., CROSS, P.C., LYONS, A.J., RYAN, S.J. & WILMERS, C.C. 2007. LoCoH: Nonparametric kernel methods for constructing home ranges and utilization distributions. PLoS One 2: e207. doi:10.1371/journal.pone.0000207

GLICK, B. 1983. Chapter 6: Bursa of Fabricius. In: FARNER, D.S., KING, J.R. & PARKES, K.C. (Eds.) Avian Biology, 1st Edition. New York, USA: Academic Press.

GINN, H.B. & MELVILLE, D.S. 1983. Moult in Birds. BTO Guide 19. Bangor, Wales: British Trust for Ornithology.

GONZÁLEZ-SOLÍS, J., CROXALL, J.P., ORO, D., & RUIZ, X. 2007. Trans-equatorial migration and mixing in the wintering areas of a pelagic seabird. Frontiers in Ecology and the Environment 5: 297-301.

GONZÁLEZ-SOLÍS, J. & SHAFFER, S.A. 2009. Introduction and synthesis: Spatial ecology of seabirds at sea. Marine Ecology Progress Series 391: 117-120. doi:10.3354/meps08282

GULKA, J., CARVALHO, P.C., JENKINS, E., JOHNSON, K., MAYNARD, L. & DAVOREN, G.K. 2017. Dietary niche shifts of multiple marine predators under varying prey availability on the northeast Newfoundland coast. Frontiers in Marine Science 4: 324.

HAMAN, K.H., NORTON, T.M., RONCONI, R.A. ET AL. 2013 Great Shearwater (Puffinus gravis) mortality events along the eastern coast of the United States. Journal of Wildlife Diseases 49: 235-245.

HAZEN, E.L., MAXWELL, S.M., BAILEY, H. ET AL. 2012. Ontogeny in marine tagging and tracking science: Technologies and data gaps. Marine Ecology Progress Series 457: 221-240. doi:10.3354/meps09857

HECTOR, J.A.L., CROXALL, J.P. & FOLLETT, B.K. 1986. Reproductive endocrinology of the Wandering Albatross Diomedea exulans in relation to biennial breeding and deferred sexual maturity. Ibis 128: 9−22.

HEDD, A., MONTEVECCHI, W.A., OTLEY, H., PHILLIPS, R.A. & FIFIELD, D.A. 2012. Trans-equatorial migration and habitat use by Sooty Shearwaters Puffinus griseus from the South Atlantic during the nonbreeding season. Marine Ecology Progress Series 449: 277−290.

HONG, P., WILEY, D.N., POWERS, K.D., MICHENER, R.H., KAUFMAN, L. & HATCH, K.A. 2019. Stable isotope analyses of multiple tissues of Great Shearwaters (Ardenna gravis) reveals long-term dietary stability, short-term changes in diet, and can be used as a tool to monitor food webs. Diversity 11: 163. doi:10.3390/d11090163

HUETTMANN, F. & DIAMOND, A.W. 2000. Seabird migration in the Canadian northwest Atlantic Ocean: Moulting locations and movement patterns of immature birds. Canadian Journal of Zoology 78: 624−647.

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

JONSEN, I.D., FLEMMING, J.M. & MYERS, R.A. 2005. Robust state-space modeling of animal movement data. Ecology 86: 2874-2880.

JONSEN, I.D., MYERS, R.A. & JAMES, M.C. 2007. Identifying leatherback turtle foraging behaviour from satellite telemetry using a switching state-space model. Marine Ecology Progress Series 337: 255-264.

JOUVENTIN, P., CAPDEVILLE, D., CUENOT-CHAILLET, F. & BOITEAU, C. 1994. Exploitation of pelagic resources by a non-flying seabird: Satellite tracking of the King Penguin throughout the breeding cycle. Marine Ecology Progress Series 106: 11-19.

KRUMBEIN, W.C. & SLOSS, L.L. 1963. Stratigraphy and Sedimentation, 2nd Edition. San Francisco, USA: W.H. Freeman.

LYONS, A. 2014. T-LoCoH for R: Tutorial and Users Guide. Vienna, Austria: R-Forge. [Accessed online at in Aug 2014.]

LYONS, A.J., TURNER, W.C. & GETZ, W.M. 2013. Home range plus: A space-time characterization of movement over real landscapes. Movement Ecology 1: 2. doi:10.1186/2051-3933-1-2

MACLEOD, C.J., ADAMS, J. & LYVER, P. 2008. At-sea distribution of satellite-tracked Grey-faced Petrels, Pterodroma macroptera gouldi, captured on the Ruamaahua (Aldermen) Islands, New Zealand. Papers and Proceedings of the Royal Society of Tasmania 142: 73-88.

MANNOCCI, L., BOUSTANY, A.M., ROBERTS, J.J. ET AL. 2017. Temporal resolutions in species distribution models of highly mobile marine animals: Recommendations for ecologists and managers. Diversity and Distributions 23: 1098-1109. doi:10.1111/ddi.12609

MASE, Y. & OISHI, T. 1991. Effects of castration and testosterone treatment on the development and involution of the bursa of Fabricius and the thymus in the Japanese Quail. General and Comparative Endocrinology 84: 426-433.

MASELLA, A.P., BARTRAM, A.K., TRUSZKOWSKI, J.M., BROWN, D.G. & NEUFELD, J.D. 2012. PANDAseq: Paired-end assembler for illumina sequences. BMC Bioinformatics 13: 31.

MCMURDIE, P.J. & HOLMES, S. 2013. phyloseq: An R package for reproducible interactive analysis and graphics of microbiome census data. PLoS One 8: e61217.

MOORE, J.K., FU, W., PRIMEAU, F. ET AL. 2018. Sustained climate warming drives declining marine biological productivity. Science 359: 1139-1143. 

NEFSC FISHERIES SAMPLING BRANCH (NEFSC FSB) 2016. Observer Data Entry Manual. Woods Hole, USA: National Oceanic and Atmospheric Administration, Northeast Fisheries Science Center.

NELSON, G.A. & ROSS, M.R. 1991. Biology and population changes of Northern Sand Lance (Ammodytes dubius) from the Gulf of Maine to the Middle Atlantic Bight. Journal of Northwest Atlantic Fishery Science 11: 11-27.

NORTHEAST FISHERIES SCIENCE CENTER (NEFSC) 2018a. 64th Northeast Regional Stock Assessment Workshop (64th SAW) Assessment Summary Report. Northeast Fisheries Science Center Reference Document 18-03. Woods Hole, USA: National Oceanic and Atmospheric Administration. 

NORTHEAST FISHERIES SCIENCE CENTER (NEFSC) 2018b. 65th Northeast Regional Stock Assessment Workshop (65th SAW) Assessment Summary Report. Northeast Fisheries Science Center Reference Document 18-08. Woods Hole, USA: National Oceanic and Atmospheric Administration.

NYE, J.A., LINK, J.S., HARE, J.A. & OVERHOLTZ, W.J. 2009 Changing spatial distribution of fish stocks in relation to climate change and population size on the northeast United States continental shelf. Marine Ecology Progress Series 393: 111-129. doi:10:3354/meps08220

PALECZNY, M., HAMMILL, E., KARPOUZI, V. & PAULY, D. 2015. Population trend of the world's monitored seabirds, 1950-2010. PLoS One 10: e0129342. doi:10.1371/journal.pone.0129342 

PHILLIPS, R.A., XAVIER, J.C. & CROXALL, J.P. 2003. Effects of satellite transmitters on albatrosses and petrels. The Auk 120: 1082-1090.

PINET, P., JAQUEMET, S., PINAUD, D., WEIMERSKIRCH, H., PHILLIPS, R.A. & LE CORRE, M. 2011. Migration, wintering distribution and habitat use of an endangered tropical seabird, Barau's Petrel Pterodroma baraui. Marine Ecology Progress Series 423: 291-302. doi:10.3354/meps08971

POLITIS, P.J., GALBRAITH, J.K., KOSTOVICK, P. & BROWN, R.W. 2014. Northeast Fisheries Science Center Bottom Trawl Survey Protocols for the NOAA Ship Henry B. Bigelow. Northeast Fisheries Science Center Reference Document 14-06. Woods Hole, USA: National Oceanic and Atmospheric Administration.

POWERS, K.D. 1983. Pelagic Distributions of Marine Birds Off the Northeastern United States. NOAA Technical Memorandum NMFS-F/NEC-27. Woods Hole, USA: National Oceanic and Atmospheric Administration.

POWERS, K.D. & BACKUS, E.H. 1987. Chapter 35: Energy transfer to seabirds. In: BACKUS E.H. (Ed.) Georges Bank. Cambridge, USA: MIT Press.

POWERS, K.D., WILEY, D.N., ALLYN, A.J., WELCH, L.J. & RONCONI, R.A. 2017. Movements and foraging habitats of Great Shearwaters Puffinus gravis in the Gulf of Maine. Marine Ecology Progress Series 574: 211-226. doi:10.3354/meps12168

PYLE, P. 2008. Identification Guide to North American Birds. Part II. Anatidae to Alcidae. Point Reyes Station, USA: Slate Creek Press.

R DEVELOPMENT CORE TEAM 2018. R: A language and environment for statistical computing. Vienna, Austria: The R Foundation for Statistical Computing.

RIOTTE-LAMBERT, L. & WEIMERSKIRCH, H. 2013. Do naive juvenile seabirds forage differently from adults? Proceedings of the Royal Society B: Biological Sciences 280: 20131434.

RONCONI, R.A., KOOPMAN, H.N., MCKINSTRY, C.A.E., WONG, S.N.P. & WESTGATE, A.J. 2010. Inter-annual variability in diet of non-breeding pelagic seabirds Puffinus spp. at migratory staging areas: Evidence from stable isotopes and fatty acids. Marine Ecology Progress Series 419: 267-282. doi:10.3354/meps08860

RONCONI, R.A., SCHOOMBIE, S., WESTGATE, A.J., WONG, S.N.P., KOOPMAN, H.N. & RYAN, P.G. 2018. Effects of age, sex, colony and breeding phase on marine space use by Great Shearwaters Ardenna gravis in the South Atlantic. Marine Biology 165: 58. doi:10.1007/s00227-018-3299-x

RYAN, P.G. (Ed.) 2007. Field Guide to the Animals and Plants of Tristan da Cunha and Gough Island. Newbury, USA: Pisces Publications.

SCHAFFER, S.A., TREMBLAY, Y., WEIMERSKIRCH, H. ET AL. 2006. Migratory shearwaters integrate oceanic resources across the Pacific Ocean in an endless summer. Proceedings of the National Academy of Sciences 103: 12799-12802.

SCHNEIDER, D.C. & PIATT, J.F. 1986. Scale-dependent correlation of seabirds with schooling fish in a coastal ecosystem. Marine Ecology Progress Series 32: 237-246.

SCOPEL, L., DIAMOND, A., KRESS, S. & SHANNON, P. 2019. Varied breeding responses of seabirds to a regime shift in prey base in the Gulf of Maine. Marine Ecology Progress Series 626: 177-196. doi:10.3354/meps13048

SCOTT, J.S. 1982. Selection of bottom type by groundfishes of the Scotian Shelf. Canadian Journal of Fisheries and Aquatic Sciences 39: 943-947.

SHEPPARD, S.K., BELL, J., SUNDERLAND, K.D., FENLON, J., SKERVIN, D. & SYMONDSON, W.O.C. 2005. Detection of secondary predation by PCR analyses of the gut contents of invertebrate generalist predators. Molecular Ecology 14: 4461-4468. doi:10.1111/j.1365-294X.2005.02742.x

STAUDINGER, M.D., GOYERT, H., SUCA, J.J. ET AL. 2020. The role of sand lances (Ammodytes sp.) in the Northwest Atlantic ecosystem: A synthesis of current knowledge with implications for conservation and management. Fish and Fisheries. 21: 522-556. doi:10.1111/faf.12445

SURYAN, R.M., KULETZ, K.J., PARKER-STETTER, S.L. ET AL. 2016. Temporal shifts in seabird populations and spatial coherence with prey in the southeastern Bering Sea. Marine Ecology Progress Series 549: 199-215. doi:10.3354/meps11653

THOMAS, A.C., JARMAN, S.N., HAMAN, K.H., TRITES, A.W. & DEAGLE, B.E. 2014. Improving accuracy of DNA diet estimates using food tissue control materials and an evaluation of proxies for digestion bias. Molecular Ecology 23: 3706-3718.

TREMBLAY, Y., BERTRAND, S., HENRY, R.W., KAPPES, M.A., COSTA, D.P. & SHAFFER, S.A. 2009. Analytical approaches to investigating seabird-environment interactions: A review. Marine Ecology Progress Series 391: 153-163.

VAN FRANEKER, J.A. 2004. Save the North Sea—Fulmar-Litter-EcoQO Manual Part 1: Collection and Dissection Procedures. Alterra-rapport 672. Wageningen, Netherlands: Alterra.

WATSON, G.E. 1971. Molting Great Shearwaters (Puffinus gravis) off Tierra del Fuego. The Auk 88: 440-442.

WEIMERSKIRCH, H., SALAMOLARD, M., SARRAZIN, F. & JOUVENTIN, P. 1993. Foraging strategy of Wandering Albatrosses through the breeding-season: A study using satellite telemetry. The Auk 110: 325-342.

WEIMERSKIRCH, H., DONCASTER, C.P. & CUENOT-CHAILLET, F. 1994. Pelagic seabirds and the marine environment: Foraging patterns of Wandering Albatrosses in relation to prey availability and distribution. Proceedings of the Royal Society B: Biological Sciences 255: 91-97.

WINTERS, G.H. 1983. Analysis of the biological and demographic parameters of the Northern Sand Lance, Ammodytes dubius, from the Newfoundland Grand Bank. Canadian Journal of Fisheries and Aquatics Sciences 40: 409-419.

WONG, S.N.P., RONCONI, R.A. & GJERDRUM, C. 2018. Autumn at-sea distribution and abundance of phalaropes Phalaropus and other seabirds in the lower Bay of Fundy, Canada. Marine Ornithology 46: 1-10.

WINSHIP, A.J., KINLAN, B.P., WHITE, T.P., LEIRNESS, J.B. & CHRISTENSEN, J. 2018. Modeling At-Sea Density of Marine Birds to Support Atlantic Marine Renewable Energy Planning: Final Report. OCS Study: BOEM 2018-010. Sterling, USA: Bureau of Ocean Energy Management, Office of Renewable Energy Programs.

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