Key words: stable isotope analysis, foraging ecology, trophic dynamics, diet, seabird ecology
References
Barrett, R. T., Camphuysen, C. J., Anker-Nilssen, T., Chardine, J. W., Furness, R. W., Garthe, S., Hüppop, O., Leopold, M. F., Montevecchi, W. A., & Veit, R. R. (2007). Diet studies of seabirds: A review and recommendations.
ICES Journal of Marine Science, 64(9), 1675-1691.
https://doi.org/10.1093/icesjms/fsm152
Bates, D., Mächler, M., Bolker, B., & Walker, S. (2015). Fitting linear mixed-effects models using lme4.
Journal of Statistical Software, 67(1), 1-48.
https://doi.org/10.18637/jss.v067.i01
Bearhop, S., Waldron, S., Votier, S. C., & Furness, R. W. (2002). Factors that influence assimilation rates and fractionation of nitrogen and carbon stable isotopes in avian blood and feathers.
Physiological and Biochemical Zoology, 75(5), 451-458.
https://doi.org/10.1086/342800
Becker, B. H., Newman, S. H., Inglis, S., & Beissinger, S. R. (2007). Diet-feather stable isotope (δ
15N and δ
13C) fractionation in Common Murres and other seabirds.
The Condor, 109(2), 451-456.
https://doi.org/10.1093/condor/109.2.451
Bennett, S. (2017).
An investigation of the predation of Atlantic Puffins, Fratercula arctica,
by Great Black-backed Gulls, Larus marinus [Unpublished master's thesis]. Imperial College London.
http://hdl.handle.net/10044/1/52359
Bodin, N., Le Loc'h, F., & Hily, C. (2007). Effect of lipid removal on carbon and nitrogen stable isotope ratios in crustacean tissues.
Journal of Experimental Marine Biology and Ecology, 341(2), 168-175.
https://doi.org/10.1016/j.jembe.2006.09.008
Bolnick, D. I., Svanbäck, R., Fordyce, J. A., Yang, L. H., Davis, J. M., Hulsey, C. D., & Forister, M. L. (2003). The ecology of individuals: Incidence and implications of individual specialization.
The American Naturalist, 161(1), 1-28.
https://doi.org/10.1086/343878
Borrmann, R. M., Phillips, R. A., Clay, T. A., & Garthe, S. (2019). High foraging site fidelity and spatial segregation among individual Great Black-backed Gulls.
Journal of Avian Biology, 50(12), 1-10.
https://doi.org/10.1111/jav.02156
Calado, J. G., Paiva, V. H., Ceia, F. R., Gomes, P., Ramos, J. A., & Velando, A. (2020). Stable isotopes reveal year-round sexual trophic segregation in four Yellow-legged Gull colonies.
Marine Biology, 167, Article 65.
https://doi.org/10.1007/s00227-020-3676-0
Coulson, J. C. (2019). Gulls (Collins New Naturalist Library, Book 139). William Collins.
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(7), 1234-1241.
https://doi.org/10.1093/icesjms/fss032
Dehnhard, N., Achurch, H., Clarke, J., Michel, L. N., Southwell, C., Sumner, M. D., Eens, M., & Emmerson, L. (2020). High inter- and intraspecific niche overlap among three sympatrically breeding, closely related seabird species: Generalist foraging as an adaptation to a highly variable environment?
Journal of Animal Ecology, 89(1), 104-119.
https://doi.org/10.1111/1365-2656.13078
Demongin, L. (2016). Identification guide to birds in the hand. (H. Lelièvre & G. Candelin, Trans.). Laurent Demongin.
Dias, M. P., Martin, R., Pearmain, E. J., Burfield, I. J., Small, C., Phillips, R. A., Yates, O., Lascelles, B., Borboroglu, P. G., & Croxall, J. P. (2019). Threats to seabirds: A global assessment.
Biological Conservation, 237(September), 525-537.
https://doi.org/10.1016/j.biocon.2019.06.033
Elliott, K. H., Roth, J. D., & Crook, K. (2017). Lipid extraction techniques for stable isotope analysis and ecological assays. In S. K. Bhattacharya (Ed.),
Lipidomics (pp. 9-24). Springer New York.
https://doi.org/10.1007/978-1-4939-6996-8_2
Enners, L., Schwemmer, P., Corman, A.-M., Voigt, C. C., & Garthe, S. (2018). Intercolony variations in movement patterns and foraging behaviors among Herring Gulls (
Larus argentatus) breeding in the eastern Wadden Sea.
Ecology and Evolution, 8(15), 7529-7542.
https://doi.org/10.1002/ece3.4167
Furness, R. W., & Tasker, M. L. (2000). Seabird-fishery interactions: Quantifying the sensitivity of seabirds to reductions in sandeel abundance, and identification of key areas for sensitive seabirds in the North Sea.
Marine Ecology Progress Series, 202, 253-264.
https://doi.org/10.3354/meps202253
Greig, S. A., Coulson, J. C., & Monaghan, P. (1985). Feeding strategies of male and female adult Herring Gulls (
Larus argentatus).
Behaviour, 94(1-2), 41-59.
https://doi.org/10.1163/156853985X00262
Grémillet, D., Ponchon, A., Paleczny, M., Palomares, M.-L. D., Karpouzi, V., & Pauly, D. (2018). Persisting worldwide seabird-fishery competition despite seabird community decline.
Current Biology, 28(24), 4009-4013.e2.
https://doi.org/10.1016/j.cub.2018.10.051
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, Article 324.
https://doi.org/10.3389/fmars.2017.00324
Harris, M. P., & Wanless, S. (1996). Differential responses of Guillemot
Uria aalge and Shag
Phalacrocorax aristotelis to a late winter wreck.
Bird Study, 43(2), 220-230.
https://doi.org/10.1080/00063659609461014
Harris, M. P., Beare, D., Toresen, R., Nøttestad, L., Kloppmann, M., Dörner, H., Peach, K., Rushton, D. R. A., Foster-Smith, J., & Wanless, S. (2007). A major increase in snake pipefish (
Entelurus aequoreus) in northern European seas since 2003: Potential implications for seabird breeding success.
Marine Biology, 151, 973-983.
https://doi.org/10.1007/s00227-006-0534-7
Hebert, C. E., Weseloh, D. V. C., Idrissi, A., Arts, M. T., O'Gorman, R., Gorman, O. T., Locke, B., Madenjian, C. P., & Roseman, E. F. (2008). Restoring piscivorous fish populations in the Laurentian Great Lakes causes seabird dietary change.
Ecology, 89(4), 891-897.
https://doi.org/10.1890/07-1603.1
Hobson, K. A., & Clark, R. G. (1992). Assessing avian diets using stable isotopes II: Factors influencing diet-tissue fractionation.
The Condor, 94(1), 189-197.
https://doi.org/10.2307/1368808
Hobson, K. A., Piatt, J. F., & Pitocchelli, J. (1994). Using stable isotopes to determine seabird trophic relationships.
Journal of Animal Ecology, 63(4), 786-798.
https://doi.org/10.2307/5256
Jackson, A. L., Inger, R., Parnell, A. C., & Bearhop, S. (2011). Comparing isotopic niche widths among and within communities: SIBER—Stable Isotope Bayesian Ellipses in R.
Journal of Animal Ecology, 80(3), 595-602.
https://doi.org/10.1111/j.1365-2656.2011.01806.x
Jessopp, M., Arneill, G. E., Nykänen, M., Bennison, A., & Rogan, E. (2020). Central place foraging drives niche partitioning in seabirds.
Oikos, 129(11), 1704-1713.
https://doi.org/10.1111/oik.07509
Jiménez, S., Domingo, A., Brazeiro, A., Defeo, O., Wood, A. G., Froy, H., Xavier, J. C., & Phillips, R. A. (2016). Sex-related variation in the vulnerability of wandering albatrosses to pelagic longline fleets.
Animal Conservation, 19(3), 281-295.
https://doi.org/10.1111/acv.12245
Käkelä, A., Furness, R. W., Kelly, A., Strandberg, U., Waldron, S., & Käkelä, R. (2007). Fatty acid signatures and stable isotopes as dietary indicators in North Sea seabirds.
Marine Ecology Progress Series, 342, 291-301.
https://doi.org/10.3354/meps342291
Kasinsky, T., Yorio, P., Dell'Arciprete, P., Marinao, C., & Suárez, N. (2021). Geographical differences in sex-specific foraging behaviour and diet during the breeding season in the opportunistic Kelp Gull (
Larus dominicanus).
Marine Biology, 168, Article 14.
https://doi.org/10.1007/s00227-020-03812-9
Kazama, K., Nishizawa, B., Tsukamoto, S., González, J. E., Kazama, M. T., & Watanuki, Y. (2018). Male and female Black-tailed Gulls
Larus crassirostris feed on the same prey species but use different feeding habitats.
Journal of Ornithology, 159(4), 923-934.
https://doi.org/10.1007/s10336-018-1565-9
Kowalczyk, N. D., Chiaradia, A., Preston, T. J., & Reina, R. D. (2015). Fine-scale dietary changes between the breeding and non-breeding diet of a resident seabird.
Royal Society Open Science, 2(4), Article 140291.
https://doi.org/10.1098/rsos.140291
Langlois Lopez, S., Clewley, G. D., Johnston, D. T., Daunt, F., Wilson, J. M., O'Hanlon, N. J., & Masden, E. (2024). Reduced breeding success in Great Black-backed Gulls (
Larus marinus) due to harness-mounted GPS device.
Ibis, 166(1), 69-81.
https://doi.org/10.1111/ibi.13247
Langlois Lopez, S., Daunt, F., Wilson, J., O'Hanlon, N. J., Searle, K. R., Bennett, S., Newell, M. A., Harris, M. P., & Masden, E. (2023). Quantifying the impacts of predation by Great Black-backed Gulls
Larus marinus on an Atlantic Puffin
Fratercula arctica population: Implications for conservation management and impact assessments.
Marine Environmental Research, 191(June), Article 105994.
https://doi.org/10.1016/j.marenvres.2023.105994
Lato, K. A., Madigan, D. J., Veit, R. R., & Thorne, L. H. (2021). Closely related gull species show contrasting foraging strategies in an urban environment.
Scientific Reports, 11, Article 23619.
https://doi.org/10.1038/s41598-021-02821-y
Lenth, R., Singmann, H., Love, J., Buerkner, P., & Herve, M. (2018).
emmeans: Estimated marginal means, aka least-squares means. Version 1.7.0.
https://doi.org/10.32614/CRAN.package.emmeans
Mancini, P. L., Bond, A. L., Hobson, K. A., Duarte, L. S., & Bugoni, L. (2013). Foraging segregation in tropical and polar seabirds: Testing the intersexual competition hypothesis.
Journal of Experimental Marine Biology and Ecology, 449(November), 186-193.
https://doi.org/10.1016/j.jembe.2013.09.011
Mawhinney, K., & Diamond, T. (1999). Sex determination of Great Black-Backed Gulls using morphometric characters.
Journal of Field Ornithology, 70(2), 206-210.
https://www.jstor.org/stable/4514402
Maynard, L. D., Gulka, J., Jenkins, E., & Davoren, G. K. (2021). Different individual-level responses of Great Black-backed Gulls (
Larus marinus) to shifting local prey availability.
PLOS One, 16(10), Article e0252561.
https://doi.org/10.1371/journal.pone.0252561
McInnes, J. C., Jarman, S. N., Lea, M.-A., Raymond, B., Deagle, B. E., Phillips, R. A., Catry, P., Stanworth, A., Weimerskirch, H., Kusch, A., Gras, M., Cherel, Y., Maschette, D., & Alderman, R. (2017). DNA metabarcoding as a marine conservation and management tool: A circumpolar examination of fishery discards in the diet of threatened albatrosses.
Frontiers in Marine Science, 4, 277.
https://doi.org/10.3389/fmars.2017.00277
Meier, R. E., Votier, S. C., Wynn, R. B., Guilford, T., McMinn Grivé, M., Rodríguez, A., Newton, J., Maurice, L., Chouvelon, T., Dessier, A., & Trueman, C. N. (2017). Tracking, feather moult and stable isotopes reveal foraging behaviour of a critically endangered seabird during the non-breeding season.
Diversity and Distributions, 23(2), 130-145.
https://doi.org/10.1111/ddi.12509
Michalik, A., McGill, R. A. R., Furness, R. W., Eggers, T., van Noordwijk, H. J., & Quillfeldt, P. (2010). Black and white - Does melanin change the bulk carbon and nitrogen isotope values of feathers?
Rapid Communications in Mass Spectrometry, 24(7), 875-878.
https://doi.org/10.1002/rcm.4462
Mitchell, P. I., Newton, S. F., Ratcliffe, N., & Dunn, T. E. (2004). Seabird populations of Britain and Ireland. T. & A. D. Poyser.
Mizutani, H., Fukuda, M., & Kabaya, Y. (1992). δ13C and δ15N enrichment factors of feathers of 11 species of adult birds.
Ecology, 73(4), 1391-1395.
https://doi.org/10.2307/1940684
Navarro, J., Moreno, R., Braun, L., Sanpera, C., & Hennicke, J. C. (2014). Resource partitioning between incubating and chick-rearing Brown Boobies and Red-tailed Tropicbirds on Christmas Island.
Zoological Studies, 53, Article 27.
https://doi.org/10.1186/s40555-014-0027-1
Newsome, S. D., Martinez del Rio, C., Bearhop, S., & Phillips, D. L. (2007). A niche for isotopic ecology.
Frontiers in Ecology and the Environment, 5(8), 429-436.
https://doi.org/10.1890/060150.1
Nielsen, J. M., Clare, E. L., Hayden, B., Brett, M. T., & Kratina, P. (2018). Diet tracing in ecology: Method comparison and selection.
Methods in Ecology and Evolution, 9(2), 278-291.
https://doi.org/10.1111/2041-210X.12869
O'Hanlon, N. J., Clewley, G. D.., Johnston, D. T.., Thaxter, C. B., Langlois Lopez, S., Quinn, L. R., Boersch-Supan, P. H.., Masden, E. A.., Daunt, F., Wilson, J., Burton, N. H. K.., & Humphreys, E. M. (2025). Partial niche partitioning in three sympatric gull species through foraging areas and habitat selection.
Ecology and Evolution, 15(7), e71577.
https://doi.org/10.1002/ece3.71577
O'Hanlon, N. J., McGill, R. A. R., & Nager, R. G. (2017). Increased use of intertidal resources benefits breeding success in a generalist gull species.
Marine Ecology Progress Series, 574, 193-210.
https://doi.org/10.3354/meps12189
Owen, E., Daunt, F., Moffat, C., Elston, D. A., Wanless, S., & Thompson, P. (2013). Analysis of fatty acids and fatty alcohols reveals seasonal and sex-specific changes in the diets of seabirds.
Marine Biology, 160, 987-999.
https://doi.org/10.1007/s00227-012-2152-x
Paritte, J. M., & Kelly, J. F. (2009). Effect of cleaning regime on stable-isotope ratios of feathers in Japanese Quail (
Coturnix japonica).
The Auk, 126(1), 165-174.
https://doi.org/10.1525/auk.2009.07187
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.
https://doi.org/10.3354/meps12217
Phillips, R. A., McGill, R. A. R., Dawson, D. A., & Bearhop, S. (2011). Sexual segregation in distribution, diet and trophic level of seabirds: Insights from stable isotope analysis.
Marine Biology, 158, 2199-2208.
https://doi.org/10.1007/s00227-011-1725-4
Post, D. M., Layman, C. A., Arrington, D. A., Takimoto, G., Quattrochi, J., & Montaña, C. G. (2007). Getting to the fat of the matter: Models, methods and assumptions for dealing with lipids in stable isotope analyses.
Oecologia, 152, 179-189.
https://doi.org/10.1007/s00442-006-0630-x
R Core Team. (2021).
R: A language and environment for statistical computing. R Foundation for Statistical Computing.
https://www.R-project.org/
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.
https://doi.org/10.3354/meps08860
Ronconi, R. A., Steenweg, R. J., Taylor, P. D., & Mallory, M. L. (2014). Gull diets reveal dietary partitioning, influences of isotopic signatures on body condition, and ecosystem changes at a remote colony.
Marine Ecology Progress Series, 514, 247-261.
https://doi.org/10.3354/meps10980
Searle, K. R., Regan, C. E., Perrow, M. R., Butler, A., Rindorf, A., Harris, M. P., Newell, M. A., Wanless, S., & Daunt, F. (2023). Effects of a fishery closure and prey abundance on seabird diet and breeding success: Implications for strategic fisheries management and seabird conservation.
Biological Conservation, 281(May), 109990.
https://doi.org/10.1016/j.biocon.2023.109990
Sherley, R. B., Barham, B. J., Barham, P. J., Campbell, K. J., Crawford, R. J. M., Grigg, J., Horswill, C., McInnes, A., Morris, T. L., Pichegru, L., Steinfurth, A., Weller, F., Winker, H., & Votier, S. C. (2018). Bayesian inference reveals positive but subtle effects of experimental fishery closures on marine predator demographics.
Proceedings of the Royal Society B, 285(1886), 20172443.
https://doi.org/10.1098/rspb.2017.2443
Smith, J. A., Mazumder, D., Suthers, I. M., & Taylor, M. D. (2013). To fit or not to fit: Evaluating stable isotope mixing models using simulated mixing polygons.
Methods in Ecology and Evolution, 4(7), 612-618.
https://doi.org/10.1111/2041-210X.12048
St John Glew, K., Espinasse, B., Hunt, B. P. V., Pakhomov, E. A., Bury, S. J., Pinkerton, M., Nodder, S. D., Gutiérrez-Rodríguez, A., Safi, K., Brown, J. C. S., Graham, L., Dunbar, R. B., Mucciarone, D. A., Magozzi, S., Somes, C., & Trueman, C. N. (2021). Isoscape models of the Southern Ocean: Predicting spatial and temporal variability in carbon and nitrogen isotope compositions of particulate organic matter.
Global Biogeochemical Cycles, 35(9), e2020GB006901.
https://doi.org/10.1029/2020GB006901
Steenweg, R. J., Ronconi, R. A., & Leonard, M. L. (2011). Seasonal and age-dependent dietary partitioning between the Great Black-backed and Herring Gulls.
The Condor, 113(4), 795-805.
https://doi.org/10.1525/cond.2011.110004
Stenhouse, I., & Montevecchi, W. A. (1999). Indirect effects of the availability of capelin and fishery discards: Gull predation on breeding storm-petrels.
Marine Ecology Progress Series, 184, 303-307.
https://doi.org/10.3354/meps184303
Stock, B. C., Jackson, A. L., Ward, E. J., Parnell, A. C., Phillips, D. L., & Semmens, B. X. (2018). Analyzing mixing systems using a new generation of Bayesian tracer mixing models.
PeerJ, 6, e5096.
https://doi.org/10.7717/peerj.5096
Sweeting, C. J., Polunin, N. V. C., & Jennings, S. (2006). Effects of chemical lipid extraction and arithmetic lipid correction on stable isotope ratios of fish tissues.
Rapid Communications in Mass Spectrometry, 20(4), 595-601.
https://doi.org/10.1002/rcm.2347
Wernham, C., Toms, M., Marchant, J., Clark, J., Siriwardena, G., & Baillie, S. (Eds.). (2002). The migration atlas: Movements of the birds of Britain and Ireland. T. & A. D. Poyser.
Westerberg, K., Brown, R., Eagle, G., & Votier, S. C. (2019). Intra-population variation in the diet of an avian top predator: Generalist and specialist foraging in Great Black-backed Gulls
Larus marinus. Bird Study, 66(3), 390-397.
https://doi.org/10.1080/00063657.2019.1693961
Williams, C. T., & Buck, C. L. (2010). Using fatty acids as dietary tracers in seabird trophic ecology: Theory, application and limitations.
Journal of Ornithology, 151, 531-543.
https://doi.org/10.1007/s10336-010-0513-0