Volume 47, No. 2

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Survival and recruitment dynamics of Black-legged Kittiwakes Rissa tridactyla at an Alaskan colony


1School of Biodiversity Conservation, Unity College, Unity, ME 04988, USA (
2Department of Wildlife, Fisheries, and Conservation Biology, University of Maine, Orono, ME 04469, USA
3US Fish and Wildlife Service, Migratory Bird Management Office, Anchorage, AK 99503, USA
4US Geological Survey, Maine Cooperative Fish and Wildlife Research Unit, Orono, ME 04469, USA


MCKNIGHT, A., BLOMBERG, E.J., IRONS, D.B., LOFTIN, C.S. & MCKINNEY, S.T. 2019. Survival and recruitment dynamics of Black-legged Kittiwakes Rissa tridactyla at an Alaskan colony. Marine Ornithology 47: 209 - 222

Received 03 May 2018, accepted 16 May 2019

Date Published: 2019/10/15
Date Online: 2019/09/26
Black-legged Kittiwake, Rissa tridactyla, colonial breeding, survival, recruitment, capture-mark-recapture analysis, demographic modeling


Most seabirds breed colonially and exhibit considerable site fidelity over the course of their long lifespans. Initial colony selection can therefore have substantial fitness consequences, but factors contributing to recruitment into colonies and subsequent fidelity remain unclear. We used multi-state capture-recapture models to test several hypotheses related to apparent fledgling survival, probability of recruitment to natal colonies, and apparent post-recruitment survival in Black-legged Kittiwakes Rissa tridactyla, using data from individuals banded as chicks and subsequently resighted at a colony in south-central Alaska over a 20-year period. Competitive models suggested that apparent fledgling survival declined throughout our study. This decline was likely driven by intrinsic, cohort-specific processes and was not explainable by post-fledging weather or climate conditions. Independent resightings at other colonies suggest the apparent decline may have been at least partially influenced by permanent emigration (natal dispersal), which occurred more frequently when the colony size was large. Recruitment was primarily age-dependent, with no detectable effects from early life experience or from annual changes in four factors: colony size, colony productivity, climate, or average weather conditions. We estimated an average recruitment age of seven years, which is older than typically reported for Atlantic kittiwake populations and which supports a more conservative life history strategy for kittiwakes in the Pacific. Variation in the apparent survival of recruits was cohort-specific and did not correlate with age or annual changes in the factors listed above. Instead, apparent survival of recruits was best explained by colony size during a cohort's second year, suggesting a degree of negative density dependence in post-recruitment survival or fidelity. This information could prove useful to managers deciding how to allocate resources among small, growing colonies and large, well-established colonies.


AEBISCHER, N.J. 1995. Philopatry and colony fidelity of Shags Phalacrocorax aristotelis on the east coast of Britain. Ibis 137: 11-18.

AEBISCHER, N.J. & COULSON, J.C. 1990. Survival of the kittiwake in relation to sex, year, breeding experience and position in the colony. Journal of Animal Ecology 59: 1063-1071.

AINLEY, D.G., FORD, R.G., BROWN, E.D., SURYAN, R.M. & IRONS, D.B. 2003. Prey resources, competition, and geographic structure of kittiwake colonies in Prince William Sound. Ecology 84: 709-723.

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

ASHBROOK, K., WANLESS, S., HARRIS, M.P. & HAMER, K.C. 2010. Impacts of poor food availability on positive density dependence in a highly colonial seabird. Proceedings of the Royal Society B: Biological Sciences 277: 2355-2360.

ATWOOD, J.L. & MASSEY, B.W. 1988. Site fidelity of Least Terns in California. The Condor 90: 389-394.

AUBRY, L.M., CAM, E., KOONS, D.N., MONNAT, J.-Y. & PAVARD, S. 2011. Drivers of age-specific survival in a long-lived seabird: Contributions of observed and hidden sources of heterogeneity. Journal of Animal Ecology 80: 375-383.

AUBRY, L.M., KOONS, D.N., MONNAT, J.-Y. & CAM, E. 2009. Consequences of recruitment decisions and heterogeneity on age-specific breeding success in a long-lived seabird. Ecology 90: 2491-2502.

BOSMAN, D.S., VERCRUIJSSE, H.J., STIENEN, E.W., VINCX, M., & LENS, L. 2013. Age of first breeding interacts with pre-and post-recruitment experience in shaping breeding phenology in a long-lived gull. PLoS One 8: e82093.

BOULINIER, T., MCCOY, K.D., YOCCOZ, N.G., GASPARINI, J. & TVERAA, T. 2008. Public information affects breeding dispersal in a colonial bird: Kittiwakes cue on neighbours. Biology Letters 4: 538-540.

BOULINIER, T. & DANCHIN, E. 1997. The use of conspecific reproductive success for breeding patch selection in terrestrial migratory species. Evolutionary Ecology 11: 505:517.

BRIED, J. & JOUVENTIN, P. 2002. Site and mate choice in seabirds: An evolutionary approach. In: SCHREIBER, E.A. & BURGER, J. (Eds.) Biology of Marine Birds. Boca Raton, USA: CRC Press.

BUCKLEY, N.J. 1997. Spatial-concentration effects and the importance of local enhancement in the evolution of colonial breeding in seabirds. The American Naturalist 149: 1091-1112.

BURNHAM, K.P. & ANDERSON, D.R. 1998. Model Selection and Inference: A Practical Information-Theoretic Approach. New York, USA: Springer-Verlag.

BYRD, G.V., SCHMUTZ, J.A., & RENNER, H.M. 2008. Contrasting population trends of piscivorous seabirds in the Pribilof Islands: A 30-year perspective. Deep-Sea Research Part II 55: 1846-1855.

BYRD, G.V. & WILLIAMS, J.C. 1993. Red-legged Kittiwake (Rissa brevirostris), version 2.0. In: POOLE, A.F. & GILL, F.B. (Eds.) The Birds of North America Online. Ithaca, USA: Cornell Lab of Ornithology. [Available online at, accessed February 2017.]

CADIOU, B. 1999. Attendance of breeders and prospectors reflects the quality of colonies in the Kittiwake Rissa tridactyla. Ibis 141: 321-326.

CAIRNS, D.K. 1988. Seabirds as indicators of marine food supplies. Biological Oceanography 5: 261-271.

CAIRNS, D.K. 1989. The regulation of seabird colony size: A hinterland model. The American Naturalist 134: 141-146.

CAM, E., CADIOU, B., HINES, J.E. & MONNAT, J.-Y. 2002. Influence of behavioral tactics on recruitment and reproductive trajectory in the Kittiwake. Journal of Applied Statistics 29: 163-185.

CAM, E., COOCH, E.G. & MONNAT, J.-Y. 2005. Earlier recruitment or earlier death? On the assumption of equal survival in recruitment studies. Ecological Monographs 75: 419-434.

CAM, E., MONNAT, J.-Y. & HINES, J.E. 2003. Long-term fitness consequences of early conditions in the kittiwake. Journal of Animal Ecology 72: 411-424.

CHAPDELAINE, G. & BROUSSEAU, P. 1989. Size and trends of Black-legged Kittiwake (Rissa tridactyla) populations in the Gulf of St. Lawrence (Quebec) 1974-1985. American Birds 43: 21-24.

CLANCY, C.F., O'CALLAGHAN, M.J.A. & KELLY, T.C. 2006. A multi-scale problem arising in a model of avian flu virus in a seabird colony. Journal of Physics: Conference Series 55: 45-54.

CLODE, D. 1993. Colonial breeding seabirds: Predators or prey? Trends in Ecology and Evolution 8: 336-338.

COULSON, J.C. 1966. The influence of the pair-bond and age on the breeding biology of the Kittiwake Gull Rissa tridactyla. Journal of Animal Ecology 35: 269-279.

COULSON, J.C. 2002. Colonial breeding in seabirds. In: SCHREIBER, E.A. & BURGER, J. (Eds.) Biology of Marine Birds. New York, USA: CRC Press.

COULSON, J.C. & OUELLET, H. 1988. Lifetime reproductive success in the black-legged kittiwake (Rissa tridactyla). In: OUELLET, H. (Ed.) Acta XIX Congressus Internationalis Ornithologici, Volume 2. Ottawa, Canada: University of Ottawa Press.

COULSON, J.C. & COULSON, B.A. 2008. Measuring immigration and philopatry in seabirds; recruitment to Black-legged Kittiwake colonies. Ibis 150: 288-299.

COULSON, J.C. & NÈVE DE MÉVERGNIES, G. 1992. Where do young kittiwakes Rissa tridactyla breed, philopatry or dispersal. Ardea 80: 187-197.

COULSON, J.C. & THOMAS, C.S. 1985. Changes in the biology of the kittiwake Rissa tridactyla: A 31-year study of a breeding colony. Journal of Animal Ecology 54: 9-26.

COULSON, J.C. & WOOLLER, R.D. 1984. Incubation under natural conditions in the kittiwake gull, Rissa tridactyla. Animal Behavior 32: 1204-1215.

CRESPIN, L., HARRIS, M.P., LEBRETON, J.-D., FREDERIKSEN, M. & WANLESS, S. 2006. Recruitment to a seabird population depends on environmental factors and on population size. Journal of Animal Ecology 75: 228-238.

DAHDUL, W.M. & HORN, M.H., 2003. Energy allocation and postnatal growth in captive Elegant Tern (Sterna elegans) chicks: Responses to high- versus low-energy diets. The Auk 120: 1069-1081.

DANCHIN, E. & MONNAT, J.-Y. 1992. Population-dynamics modeling of two neighboring Kittiwake Rissa tridactyla colonies. Ardea 80: 171-180.

DANCHIN, E., BOULINIER, T. & MASSOT, M. 1998. Conspecific reproductive success and breeding habitat selection: Implications for the study of coloniality. Ecology 79: 2415-2428.

DELANY, S. & SCOTT, D. 2006. Waterbird Population Estimates, 4th Edition. Wageningen, Netherlands: Wetlands International.

DESCAMPS, S., LE BOHEC, C., LE MAHO, Y., GENDNER, J.-P. & GAUTHIER-CLERC, M. 2009. Relating demographic performance to breeding-site location in the King Penguin. The Condor 111: 81-87.

EGGERS, S., GRIESSER, M., NYSTRAND, M. & EKMAN, J. 2006. Predation risk induces changes in nest-site selection and clutch size in the Siberian Jay. Proceedings of the Royal Society B: Biological Sciences 273: 701-706.

EARTH SYSTEM RESEARCH LABORATORY (ESRL) 2016a. Niño 3.4 SST Index (anomaly; 1981-2010 mean removed): Standard PSD Format. Boulder, USA: Earth System Research Laboratory, National Oceanic and Atmospheric Administration. [Available online at, accessed July 2016.]

ESRL 2016b [Online]. NCEP/NCAR Reanalysis Monthly Means and Other Derived Variables, Daily Global Analyses data. Boulder, USA: Earth System Research Laboratory, National Oceanic and Atmospheric Administration. [Available online at, accessed July 2016.]

ESRL 2016c. NOAA Optimum Interpolation (OI) Sea Surface Temperature (SST) V2 data. Boulder, USA: Earth System Research Laboratory, National Oceanic and Atmospheric Administration. [Available online at, accessed July 2016.]

FERNÁNDEZ-CHACÓN, A., GENOVART, M., PRADEL, R. ET AL. 2013. When to stay, when to disperse and where to go: Survival and dispersal patterns in a spatially structured seabird population. Ecography 36: 1117-1126.

FINNEY, S.K., HARRIS, M.P., KELLER, L.F., ELSTON, D.A., MONAGHAN, P. & WANLESS, S. 2003. Reducing the density of breeding gulls influences the pattern of recruitment of immature Atlantic Puffins Fratercula arctica to a breeding colony. Journal of Applied Ecology 40:545-552.

FORBES, L.S. & KAISER, G.W. 1994. Habitat choice in breeding seabirds: When to cross the information barrier. Oikos 70: 377-384.

FORERO, M.G., TELLA, J.L., HOBSON, K.A., BERTELLOTTI, M. & BLANCO, G. 2002. Conspecific food competition explains variability in colony size: A test in Magellanic Penguins. Ecology 83: 3466-3475 

FREDERIKSEN, M. & BREGNBALLE, T. 2001. Conspecific reproductive success affects age of recruitment in a great cormorant, Phalacrocorax carbo sinensis, colony. Proceedings of the Royal Society B: Biological Sciences 268: 1519-1526.

GILL, V.A & HATCH, S.A. 2002. Components of productivity in Black-legged Kittiwakes Rissa tridactyla: Response to supplemental feeding. Journal of Avian Biology 33: 113-126.

GOLET, G.H., SCHMUTZ, J.A., IRONS, D.B. & ESTES, J.A. 2004. Determinants of reproductive costs in the long-lived Black-legged Kittiwake: A multiyear experiment. Ecological Monographs 74: 353-372.

GOODMAN, D. 1984. Risk spreading as an adaptive strategy in iteroparous life histories. Theoretical Population Biology 25: 1-20.

GRÉMILLET, D., DELL'OMO, G., RYAN, P.G., PETERS, G., ROPERT-COUDERT, Y. & WEEKS, S.J. 2004. Offshore diplomacy, or how seabirds mitigate intra-specific competition: A case study based on GPS tracking of Cape Gannets from neighboring colonies. Marine Ecology Progress Series 268: 265-279.

HADLEY, G.L., ROTELLA, J.J. & GARROTT, R.A. 2007. Evaluation of reproductive costs for Weddell seals in Erebus Bay, Antarctica. Journal of Animal Ecology 76: 448-458.

HAMILTON, W.D. 1971. Geometry for the selfish herd. Journal of Theoretical Biology 31: 295-311.

HARDING, A., PAREDES, R., SURYAN, R.M. ET AL. 2013. Does location really matter? An inter-colony comparison of seabirds breeding at varying distances from productive oceanographic features in the Bering Sea. Deep-Sea Research Part II 94: 178-191.

HASSELL, M.P. 1975. Density-dependence in single-species populations. Journal of Animal Ecology 44: 283-295.

HATCH, S.A., ROBERTS, B.D. & FADELY, B.S. 1993. Adult survival of Black-legged Kittiwakes Rissa tridactyla at a Pacific colony. Ibis 135: 247-254.

HERNANDEZ-MATIAS, A., JOVER, L. & RUIZ, X. 2003. Predation on Common Tern eggs in relation to sub-colony size, nest aggregation and breeding synchrony. Waterbirds 26: 280-289.

HERRING RESEARCH AND MONITORING TEAM (HRMT) 2014. Pacific herring in Prince William Sound: A synthesis of recent findings. Science synthesis report for the Herring Research and Monitoring Program. Project 14120111 report for the Exxon Valdez Oil Spill (EVOS) Trustee Council. Anchorage, USA: EVOS. [Available online at, accessed November 2016.]

HUNT, G.L., JR., EPPLEY, Z.A. & SCHNEIDER, D.C. 1986. Reproductive performance of seabirds: The importance of population and colony size. The Auk 103: 306-317.

JOINT INSTITUTE FOR THE STUDY OF THE ATMOSPHERE AND OCEAN (JISAO) 2016 [Online]. University of Washington. [Available online at, accessed May 2016.]

KHARITONOV, S.P. & SIEGEL-CAUSEY, D. 1988. Colony formation in seabirds. In: JOHNSTON, R.F. (Ed.) Current Ornithology. Volume 5. Boston, USA: Springer.

KILDAW, S.D., IRONS, D.B., NYSEWANDER, D.R. & BUCK, C.L. 2005. Formation and growth of new seabird colonies: The significance of habitat quality. Marine Ornithology 33: 49-58.

KLOMP, N.I. & FURNESS, R.W. 1990. Variations in numbers of nonbreeding Great Skuas attending a colony. Ornis Scandinavica 21: 270-276.

KOKKO, H., HARRIS, M.P. & WANLESS, S. 2004. Competition for breeding sites and site-dependent population regulation in a highly colonial seabird, the common guillemot Uria aalge. Journal of Animal Ecology 73: 367-376.

LAAKE, J.L. 2013. RMark: An R Interface for Analysis of Capture-Recapture Data with MARK. AFSC Processed Report 2013-01. Seattle, USA: Alaska Fisheries Science Center, National Oceanic and Atmospheric Administration, National Marine Fisheries Service.

LAWSON, A.J., SEDINGER, J.S. & TAYLOR, E.J. 2017. Life history patterns, individual heterogeneity, and density dependence in breeding common goldeneyes of the northern boreal forest. Journal of Avian Biology 48: 597-610.

LEWIS, S., SHERRATT, T.N., HAMER, K.C. & WANLESS, S. 2001. Evidence of intra-specific competition for food in a pelagic seabird. Nature 412: 816-819.

LINK, W.A., COOCH, E.G. & CAM, E. 2002. Model-based estimation of individual fitness. Journal of Applied Statistics 29: 207-224.

MANGIN, S., GAUTHIER-CLERC, M., FRENOT, Y., GENDNER, J.-P. & LE MAHO, Y. 2003. Ticks Ixodes uriae and the breeding performance of a colonial seabird, king penguin Aptenodytes patagonicus. Journal of Avian Biology 34: 30-34.

MARTIN, E.T. 1995. Avian life history evolution in relation to nest sites, nest predation, and food. Ecological Monographs 65: 101-127.

MARTINEZ-ABRAIN, A., ORO, D., FORERO, M.G. & CONESA, D. 2003. Modeling temporal and spatial colony-site dynamics in a long-lived seabird. Population Ecology 45: 133-139.

MCKNIGHT, A. 2017. Population ecology of colonially breeding seabirds: How intrinsic processes, mediating influences, and individual heterogeneity affect population vital rates. PhD dissertation. Orono, USA: University of Maine.

MCKNIGHT, A., BLOMBERG, E.J., IRONS, D.B., LOFTIN, C.S. & MCKINNEY, S.T. 2019. Data from: Survival and recruitment dynamics of Black-legged Kittiwakes Rissa tridactyla at an Alaskan colony. Durham, USA: Dryad Digital Repository. doi:10.5061/dryad.ch2740k 

MCKNIGHT, A., IRONS, D.B., ALLYN, A.J., SULLIVAN, K.M. & SURYAN, R.M. 2011. Winter dispersal and activity patterns of post-breeding black-legged kittiwakes Rissa tridactyla from Prince William Sound, Alaska. Marine Ecology Progress Series 442: 241-253.

MCKNIGHT, A., SULLIVAN, K.M., IRONS, D.B., STEPHENSEN, S.W. & HOWLIN, S. 2008. Marine bird and sea otter population abundance of Prince William Sound, Alaska: Trends following the T/V Exxon Valdez oil spill, 1989-2007. Exxon Valdez Oil Spill Restoration Project Annual Report (Restoration Project 070751). Anchorage, USA: US Fish and Wildlife Service.

MINIAS, P., KACZMAREK, K. & JANISZEWSKI, T. 2012. Distribution of pair quality in a tree-nesting waterbird colony: Central-periphery model vs. satellite model. Canadian Journal of Zoology 90: 861-867.

MOFFITT, S. 2016. Documentation of Pacific Herring (Clupea pallasii) milt from aerial surveys in the Prince William Sound Management Area, Alaska, 1973-2016. Anchorage, USA: Alaska Department of Fish and Game, Commercial Fisheries Division. [Available online at, accessed November 2016.]

MØLLER, A.P. 1987. Advantages and disadvantages of coloniality in the swallow, Hirundo rustica. Animal Behavior 35: 819-832.

NICHOLS, J.D. & KENDALL, W.L. 1995. The use of multi-state capture-recapture models to address questions in evolutionary ecology. Journal of Applied Statistics 22: 835-846.

ORELL, M. & BELDA, E.J. 2002. Delayed cost of reproduction and senescence in the Willow Tit Parus montanus. Journal of Animal Ecology 71: 55-64.

ORO, D. & FURNESS, R.W. 2002. Influences of food availability and predation on survival of kittiwakes. Ecology 83: 2516-2528.

ORO, D. & RUXTON, G.D. 2001. The formation and growth of seabird colonies: Audouin's Gull as a case study. Journal of Animal Ecology 70: 527-535.

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

PAPINEAU, J.M. 2001. Wintertime temperature anomalies in Alaska correlated with ENSO and PDO. International Journal of Climatology 21: 1577-1592.

PAREDES, R., HARDING, A.M.A., IRONS, D.B. ET AL. 2012. Proximity to multiple foraging habitats enhances seabirds' resilience to local food shortages. Marine Ecology Progress Series 471: 253-269.

PIATT, J.F., HARDING, A.M.A., SHULTZ, M. ET AL. 2007. Seabirds as indicators of marine food supplies: Cairns revisited. Marine Ecology Progress Series 352: 221-234.

PONCHON, A., CHAMBERT, T., LOBATO, E., TVERAA, T., GRÉMILLET, D. & BOULINIER, T. 2015. Breeding failure induces large scale prospecting movements in the black-legged kittiwake. Journal of Experimental Marine Biology and Ecology 473: 138-145.

PONCHON, A., ILISZKO, L., GRÉMILLET, D., TVERAA, T. & BOULINIER, T. 2017. Intense prospecting movements of failed breeders nesting in an unsuccessful breeding subcolony. Animal Behaviour 124: 183-191.

PONCHON, A., CHOQUET, R., TORNOS, J., MCCOY, K.D., TVERAA, T. & BOULINIER, T. 2018. Survival estimates strongly depend on capture-recapture designs in a disturbed environment inducing dispersal. Ecography 41: 1-12.

PORTER, J.M. & COULSON, J.C. 1987. Long-term changes in recruitment to the breeding group, and the quality of recruits at a Kittiwake Rissa tridactyla colony. Journal of Animal Ecology 56: 675-689.

PYK, T.M., WESTON, M.A., BUNCE, A. & NORMAN, F.I. 2013. Establishment and development of a seabird colony: Long-term trends in phenology, breeding success, recruitment, breeding density and demography. Journal of Ornithology 154: 299-310.

PYLE, P., SYDEMAN, W.J. & HESTER, M. 2001. Effects of age, breeding experience, mate fidelity and site fidelity on breeding performance in a declining population of Cassin's auklets. Journal of Animal Ecology 70: 1088-1097.

R CORE TEAM 2016. R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing. [Available online at, accessed March 2017.)

REED, T.E., KRUUK, L.E., WANLESS, S., FREDERIKSEN, M., CUNNINGHAM, E.J. & HARRIS, M.P. 2008. Reproductive senescence in a long-lived seabird: Rates of decline in late-life performance are associated with varying costs of early reproduction. American Naturalist 171: E89-E101.

REGEHR, H.M. & MONTEVECCHI, W.A. 1997. Interactive effects of food shortage and predation on breeding failure of black-legged kittiwakes: Indirect effects of fisheries activities and implications for indicator species. Marine Ecology Progress Series 155: 249-260.

RENNER, H.M., MUETER, F., DRUMMOND, B.A., WARZYBOK, J.A. & SINCLAIR, E.H. 2012. Patterns of change in diets of two piscivorous seabird species during 35 years in the Pribilof Islands. Deep-Sea Research Part II 65-70: 273-291.

ROLLAND, C., DANCHIN, E. & DE FRAIPONT, M. 1998. The evolution of coloniality in birds in relation to food, habitat, predation, and life-history traits: A comparative analysis. American Naturalist 151: 514-529.

SÆTHER, B.-E., RINGSBY, T.H. & RØSKAFT, E. 1996. Life history variation, population processes and priorities in species conservation: Towards a reunion of research paradigms. Oikos 77: 217-226.

SCHJØRRING, S., GREGERSEN, J. & BREGNBALLE, T. 1999. Prospecting enhances breeding success of first-time breeders in the great cormorant, Phalacrocorax carbo sinensis. Animal Behaviour 57: 647-654.

STEARNS, S.C. 1992. The Evolution of Life Histories. Oxford, UK: Oxford University Press.

STEINER, U.K. & GASTON, A.J. 2005. Reproductive consequences of natal dispersal in a highly philopatric seabird. Behavioral Ecology 16: 634-639.

SZOSTEK, K.L., SCHAUB, M. & BECKER, P.H. 2014. Immigrants are attracted by local pre-breeders and recruits in a seabird colony. Journal of Animal Ecology 83: 1015-1024.

TELLA, J.L., FORERO, M.G., BERTELLOTTI, M., DONÁZAR, J.A., BLANCO, G. & CEBALLOS, O. 2001. Offspring body condition and immunocompetence are negatively affected by high breeding densities in a colonial seabird: A multiscale approach. Proceedings of the Royal Society B: Biological Sciences 268: 1455-1461.

THIBAULT, J.-C. 1993. Natal philopatry in the Cory's Shearwater (Calonectris d. diomedea) on Lavezzi Island, Corsica. Colonial Waterbirds 16: 77-82.

VINCENZI, S.  HATCH, S., MANGEL, M. & KITAYSKY, A. 2013. Food availability affects onset of reproduction in a long-lived seabird. Proceedings of the Royal Society B: Biological Sciences 280: 20130554.

VINE, I. 1971. Risk of visual detection and pursuit by a predator and the selective advantage on flocking behavior. Journal of Theoretical Biology 30: 405-422.

WHITE, G.C. & BURNHAM, K.P. 1999. Program MARK: Survival estimation from populations of marked animals. Bird Study 46: S120-S139.

WITTENBURGER, J.F. & HUNT, G.L., JR. 1985. The adaptive significance of coloniality in birds. In FARNER, D.S., KING, J.R. & PARKES, K.C. (Eds.) Avian Biology (Volume VIII). New York, USA: Academic Press, Inc.

WHITTHAM, R.M. & LEONARD, M.L. 1999. Predation and breeding success in roseate terns (Sterna dougallii). Canadian Journal of Zoology 77: 851-856.

WOOLLER, R.D. & COULSON, J.C. 1977. Factors affecting the age of first breeding of the kittiwake Rissa tridactyla. Ibis 119: 339-349.

YOUNG, T.P. 1990. Evolution of semelparity in Mount Kenya lobelias. Evolutionary Ecology 4: 157-172.

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