Section: Ecology
Topic: Ecology, Population biology

Sexual segregation in a highly pagophilic and sexually dimorphic marine predator

10.24072/pcjournal.75 - Peer Community Journal, Volume 1 (2021), article no. e71.

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Sexual segregation is common in many species and has been attributed to intra-specific competition, sex-specific differences in foraging efficiency or in activity budgets and habitat choice. However, very few studies have simultaneously quantified sex-specific foraging strategies, at sea distribution, habitat use, and trophic ecology. Moreover, these studies come from low latitude areas reflecting a lack of evidence for polar species. We investigated sexual segregation in snow petrels Pagodroma nivea and combined movement, foraging trip efficiency, stable isotope and oceanographic data to test whether sexual segregation results from sex-specific habitat use. Breeding birds foraging in the Dumont d'Urville sea, Antarctica, were tracked during incubation. Some similarities between males and females foraging characteristics did not support the sexual segregation hypothesis. Indeed, space-use sharing and utilization distribution, d13C values and foraging trip performances (trip duration, length, speed and directions, mass gain, proportion mass gain) were similar between males and females. However, there was support for sexual segregation in foraging characteristics linked to foraging habitats. Females foraged less than males in areas with higher sea ice concentration (SIC 70%) and had lower d15N values in plasma, blood cells and feathers. Foraging efficiency (proportionate daily mass gain while foraging), was greater for females than for males, and was greater for larger females with deeper bills. Females were more efficient than males during short (2 days) foraging trips, and for females, but not for males, mass gain, proportion mass gain and body condition at return from a foraging trip were positively correlated to SIC of the foraging areas. Together, these results suggest an absence of sexual segregation at large spatial scales in snow petrels during incubation, but strongly support habitat segregation between high (70%) more profitable SIC (males) and low SIC areas (females), probably driven by intraspecific competition. Therefore, male and female snow petrels segregate at small spatial scales mainly determined by habitat (SIC) characteristics.

Published online:
DOI: 10.24072/pcjournal.75
Type: Research article

Barbraud, Christophe 1; Delord, Karine 1; Kato, Akiko 1; Bustamante, Paco 2; Cherel, Yves 1

1 Centre d’Etudes Biologiques de Chizé, UMR7372 CNRS-La Rochelle Université, 79360 Villiers-en-Bois, France
2 Littoral Environnement et Sociétés (LIENSs), UMR7266 CNRS-La Rochelle Université, 2 rue Olympe de Gouges, 17000 La Rochelle, France
License: CC-BY 4.0
Copyrights: The authors retain unrestricted copyrights and publishing rights
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Barbraud, Christophe; Delord, Karine; Kato, Akiko; Bustamante, Paco; Cherel, Yves. Sexual segregation in a highly pagophilic and sexually dimorphic marine predator. Peer Community Journal, Volume 1 (2021), article  no. e71. doi : 10.24072/pcjournal.75.

PCI peer reviews and recommendation, and links to data, scripts, code and supplementary information: 10.24072/pci.ecology.100025

Conflict of interest of the recommender and peer reviewers:
The recommender in charge of the evaluation of the article and the reviewers declared that they have no conflict of interest (as defined in the code of conduct of PCI) with the authors or with the content of the article.

[1] Ainley, D.; Fraser, W.; Smith, W.; Hopkins, T.; Torres, J. The structure of upper level pelagic food webs in the Antarctic: Effect of phytoplankton distribution, Journal of Marine Systems, Volume 2 (1991) no. 1-2, pp. 111-122 | DOI

[2] Ainley, D. G.; Fraser, W. R.; Sullivan, C. W.; Torres, J. J.; Hopkins, T. L.; Smith, W. O. Antarctic Mesopelagic Micronekton: Evidence from Seabirds That Pack Ice Affects Community Structure, Science, Volume 232 (1986) no. 4752, pp. 847-849 | DOI

[3] Ainley, D. G.; O'Connor, E. F.; Boekelheide, R. J. The Marine Ecology of Birds in the Ross Sea, Antarctica, Ornithological Monographs (1984) no. 32 | DOI

[4] Ainley, D.; Ribic, C.; Fraser, W. Does prey preference affect habitat choice in Antarctic seabirds?, Marine Ecology Progress Series, Volume 90 (1992), pp. 207-221 | DOI

[5] Ainley, D. G.; Ribic, C. A.; Spear, L. B. Species-Habitat Relationships among Antarctic Seabirds: A Function of Physical or Biological Factors?, The Condor, Volume 95 (1993) no. 4, pp. 806-816 | DOI

[6] Baduini, C. L.; Hyrenbach, K. D. Biogeography of procellariiform foraging strategies: does ocean productivity influence provisioning, Marine Ornithology, Volume 31 (2003)

[7] Barboza, P. S.; Bowyer, R. T. Sexual segregation in dimorphic deer: a new gastrocentric hypothesis, Journal of Mammalogy, Volume 81 (2000) no. 2, pp. 473-489 | DOI

[8] Barbraud, C. Aspects écologiques et évolutifs de la variation de la taille corporelle chez le pétrel des neiges, Pagodroma nivea. PhD Thesis, Université de Tours, France. (1999)

[9] Barbraud, C. Subspecies-Selective Predation of Snow Petrels by Skuas, Oikos, Volume 86 (1999) no. 2 | DOI

[10] Barbraud, C.; Chastel, O. Early body condition and hatching success in the snow petrel Pagodroma nivea, Polar Biology, Volume 21 (1999) no. 1, pp. 1-4 | DOI

[11] Barbraud, C.; Mariani, A.; Jouventin, P. Variation in call properties of the snow petrel, Pagodroma nivea, in relation to sex and body size, Australian Journal of Zoology, Volume 48 (2000) no. 4 | DOI

[12] Barbraud, C.; Weimerskirch, H.; Robertson, G. G.; Jouventin, P. Size-related life history traits: insights from a study of snow petrels (Pagodroma nivea), Journal of Animal Ecology, Volume 68 (1999) no. 6, pp. 1179-1192 | DOI

[13] Baylis, A.; Orben, R.; Costa, D.; Arnould, J.; Staniland, I. Sexual segregation in habitat use is smaller than expected in a highly dimorphic marine predator, the southern sea lion, Marine Ecology Progress Series, Volume 554 (2016), pp. 201-211 | DOI

[14] Bearhop, S.; Waldron, S.; Votier, S. C.; Furness, R. W. Factors That Influence Assimilation Rates and Fractionation of Nitrogen and Carbon Stable Isotopes in Avian Blood and Feathers, Physiological and Biochemical Zoology, Volume 75 (2002) no. 5, pp. 451-458 | DOI

[15] Beck, J. Breeding seasons and moult in some smaller Antarctic petrels In: Antarctic Ecology ( MW Holdgate Ed.) (1970), pp. 542-550

[16] Beier, P. Sex Differences in Quality of White-Tailed Deer Diets, Journal of Mammalogy, Volume 68 (1987) no. 2, pp. 323-329 | DOI

[17] Benjamini, Y.; Hochberg, Y. Controlling the False Discovery Rate: A Practical and Powerful Approach to Multiple Testing, Journal of the Royal Statistical Society: Series B (Methodological), Volume 57 (1995) no. 1, pp. 289-300 | DOI

[18] Breed, G. A.; Bowen, W.; McMillan, J.; Leonard, M. Sexual segregation of seasonal foraging habitats in a non-migratory marine mammal, Proceedings of the Royal Society B: Biological Sciences, Volume 273 (2006) no. 1599, pp. 2319-2326 | DOI

[19] Brown, D. Breeding biology of the Snow petrel Pagodroma nivea (Forster), Anare Scientific Reports, Volume 89 (1966)

[20] Calenge, C.; Dray, S.; Royer-Carenzi, M. The concept of animals' trajectories from a data analysis perspective, Ecological Informatics, Volume 4 (2009) no. 1, pp. 34-41 | DOI

[21] Carravieri, A.; Cherel, Y.; Blévin, P.; Brault-Favrou, M.; Chastel, O.; Bustamante, P. Mercury exposure in a large subantarctic avian community, Environmental Pollution, Volume 190 (2014), pp. 51-57 | DOI

[22] Cavalieri, D. J.; Crawford, J. P.; Drinkwater, M. R.; Eppler, D. T.; Farmer, L. D.; Jentz, R. R.; Wackerman, C. C. Aircraft active and passive microwave validation of sea ice concentration from the Defense Meteorological Satellite Program special sensor microwave imager, Journal of Geophysical Research, Volume 96 (1991) no. C12 | DOI

[23] Ceia, F. R.; Phillips, R. A.; Ramos, J. A.; Cherel, Y.; Vieira, R. P.; Richard, P.; Xavier, J. C. Short- and long-term consistency in the foraging niche of wandering albatrosses, Marine Biology, Volume 159 (2012) no. 7, pp. 1581-1591 | DOI

[24] Chastel, O.; Weimerskirch, H.; Jouventin, P. High annual variability in reproductive success and survival of an Antarctic seabird, the snow petrel Pagodroma nivea, Oecologia, Volume 94 (1993) no. 2, pp. 278-285 | DOI

[25] Chaurand, T.; Weimerskirch, H. Incubation routine, body mass regulation and egg neglect in the Blue Petrel Halobaena caerulea, Ibis, Volume 136 (1994) no. 3, pp. 285-290 | DOI

[26] Cherel, Y. Isotopic niches of emperor and Adélie penguins in Adélie Land, Antarctica, Marine Biology, Volume 154 (2008) no. 5, pp. 813-821 | DOI

[27] Cherel, Y.; Fontaine, C.; Richard, P.; Labatc, J.-P. Isotopic niches and trophic levels of myctophid fishes and their predators in the Southern Ocean, Limnology and Oceanography, Volume 55 (2010) no. 1, pp. 324-332 | DOI

[28] Cherel, Y.; Hobson, K. Geographical variation in carbon stable isotope signatures of marine predators: a tool to investigate their foraging areas in the Southern Ocean, Marine Ecology Progress Series, Volume 329 (2007), pp. 281-287 | DOI

[29] Cherel, Y.; Hobson, K. A.; Bailleul, F.; Groscolas, R. Nutrition, physiology, and stable isotopes: new information from fasting and molting penguins, Ecology, Volume 86 (2005) no. 11, pp. 2881-2888 | DOI

[30] Cherel, Y.; Hobson, K. A.; Weimerskirch, H. Using stable isotopes to study resource acquisition and allocation in procellariiform seabirds, Oecologia, Volume 145 (2005) no. 4, pp. 533-540 | DOI

[31] Cherel, Y.; Koubbi, P.; Giraldo, C.; Penot, F.; Tavernier, E.; Moteki, M.; Ozouf-Costaz, C.; Causse, R.; Chartier, A.; Hosie, G. Isotopic niches of fishes in coastal, neritic and oceanic waters off Adélie land, Antarctica, Polar Science, Volume 5 (2011) no. 2, pp. 286-297 | DOI

[32] Chesson, P. Mechanisms of Maintenance of Species Diversity, Annual Review of Ecology and Systematics, Volume 31 (2000) no. 1, pp. 343-366 | DOI

[33] Clarke, J.; Manly, B.; Kerry, K.; Gardner, H.; Franchi, E.; Corsolini, S.; Focardi, S. Sex differences in Adélie penguin foraging strategies, Polar Biology, Volume 20 (1998) no. 4, pp. 248-258 | DOI

[34] Cleasby, I.; Wakefield, E.; Bodey, T.; Davies, R.; Patrick, S.; Newton, J.; Votier, S.; Bearhop, S.; Hamer, K. Sexual segregation in a wide-ranging marine predator is a consequence of habitat selection, Marine Ecology Progress Series, Volume 518 (2015), pp. 1-12 | DOI

[35] Clutton‐Brock, T. H.; Iason, G. R.; Guinness, F. E. Sexual segregation and density‐related changes in habitat use in male and female Red deer ( Cerrus elaphus ), Journal of Zoology, Volume 211 (1987) no. 2, pp. 275-289 | DOI

[36] Conradt, L. Could asynchrony in activity between the sexes cause intersexual social segregation in ruminants?, Proceedings of the Royal Society of London. Series B: Biological Sciences, Volume 265 (1998) no. 1403, pp. 1359-1368 | DOI

[37] Conradt, L. Definitions, hypotheses, models and measures in the study of animal segregation In: Sexual segregation in vertebrates: ecology of the two sexes ( KE Ruckstuhl and P Neuhaus , eds). (2005), pp. 11-34

[38] Croxall, J. Sexual dimorphism in Snow petrels Pagodroma nivea, Notornis, Volume 29 (1982)

[39] Croxall, J. P.; Steele, W. K.; McInnes, S. J.; Prince, P. A. Breeding distribution of the snow petrel Pagodroma nivea, Marine Ornithology, Volume 23 (1995)

[40] Delord, K.; Pinet, P.; Pinaud, D.; Barbraud, C.; De Grissac, S.; Lewden, A.; Cherel, Y.; Weimerskirch, H. Species-specific foraging strategies and segregation mechanisms of sympatric Antarctic fulmarine petrels throughout the annual cycle, Ibis, Volume 158 (2016) no. 3, pp. 569-586 | DOI

[41] Elliott, K. H.; Gaston, A. J.; Crump, D. Sex-specific behavior by a monomorphic seabird represents risk partitioning, Behavioral Ecology, Volume 21 (2010) no. 5, pp. 1024-1032 | DOI

[42] Fairbairn, J.; Shine, R. Patterns of Sexual Size Dimorphism in Seabirds of the Southern Hemisphere, Oikos, Volume 68 (1993) no. 1 | DOI

[43] Ferretti V, S. G. C. R. C. N. Diet of the Snow Petrel Pagodroma nivea at Laurie Island, Antarctica, during the 1997/98 breeding season, Marine Ornithology, Volume 29 (2001)

[44] Fieberg, J.; Kochanny, C. O. Quantifying home-range overlap: the importance of the utilization distribution, Journal of Wildlife Management, Volume 69 (2005) no. 4, pp. 1346-1359 | DOI

[45] Field, S. A.; Tyre, A. J.; Jonzen, N.; Rhodes, J. R.; Possingham, H. P. Minimizing the cost of environmental management decisions by optimizing statistical thresholds, Ecology Letters, Volume 7 (2004) no. 8, pp. 669-675 | DOI

[46] Giraldo, C.; Cherel, Y.; Vallet, C.; Mayzaud, P.; Tavernier, E.; Moteki, M.; Hosie, G.; Koubbi, P. Ontogenic changes in the feeding ecology of the early life stages of the Antarctic silverfish (Pleuragramma antarcticum) documented by stable isotopes and diet analysis in the Dumont d’Urville Sea (East Antarctica), Polar Science, Volume 5 (2011) no. 2, pp. 252-263 | DOI

[47] González-Solís, J.; Croxall, J. P.; Wood, A. G. Sexual dimorphism and sexual segregation in foraging strategies of northern giant petrels, Macronectes halli, during incubation, Oikos, Volume 90 (2000) no. 2, pp. 390-398 | DOI

[48] Griffiths, A. Factors affecting the distribution of the Snow Petrel (Pagodroma nivea) and the Antarctic Petrel (Thalassoica antarctica), Ardea, Volume 71 (1983)

[49] Gross, J. E. Sexual Segregation in Ungulates: A Comment, Journal of Mammalogy, Volume 79 (1998) no. 4, pp. 1404-1409 | DOI

[50] Guillotin, M.; Jouventin, P. Le Pétrel des Neiges à Pointe Géologie, Gerfaut, Volume 70 (1980)

[51] Harder, L. D.; Barrett, S. C. H.; Cole, W. W. The mating consequences of sexual segregation within inflorescences of flowering plants, Proceedings of the Royal Society of London. Series B: Biological Sciences, Volume 267 (2000) no. 1441, pp. 315-320 | DOI

[52] Harper, P. C.; Croxall, J. P.; Cooper, J. A guide to foraging methods used by marine birds in antarctic and subantarctic seas, Biomass Handbook, Volume 24 (1985)

[53] Hastie, T. J.; Tibshirani, R. J. Generalized additive models, volume 43 of Monographs on Statistics and Applied Probability., 1990

[54] Hennicke, J. C.; James, D. J.; Weimerskirch, H. Sex-Specific Habitat Utilization and Differential Breeding Investments in Christmas Island Frigatebirds throughout the Breeding Cycle, PLOS ONE, Volume 10 (2015) no. 6 | DOI

[55] Hijmans, R. raster: Geographic Data Analysis and Modeling. R package version 2.6-7, 2018

[56] Hobson, K. A.; Clark, R. G. Assessing Avian Diets Using Stable Isotopes I: Turnover of13C in Tissues, The Condor, Volume 94 (1992) no. 1, pp. 181-188 | DOI

[57] Hobson, K. A.; Clark, R. G. Turnover of13C in Cellular and Plasma Fractions of Blood: Implications for Nondestructive Sampling in Avian Dietary Studies, The Auk, Volume 110 (1993) no. 3, pp. 638-641 | DOI

[58] Hochkirch, A.; Gröning, J.; Krause, S. Intersexual niche segregation in Cepero’s Ground-hopper, Tetrix ceperoi, Evolutionary Ecology, Volume 21 (2007) no. 6, pp. 727-738 | DOI

[59] Hodum, P.; Hobson, K. Trophic relationships among Antarctic fulmarine petrels:insights into dietary overlap and chick provisioning strategies inferred from stable-isotope (d15N and d13C) analyses, Marine Ecology Progress Series, Volume 198 (2000), pp. 273-281 | DOI

[60] Isenmann, P. Contribution à la biologie de reproduction du pétrel des neiges (Pagodroma nivea Forster). Le problème de la petite et de la grande forme. , L'Oiseau et la RFO, Volume 40 (1970)

[61] Jackson, A. L.; Inger, R.; Parnell, A. C.; Bearhop, S. Comparing isotopic niche widths among and within communities: SIBER - Stable Isotope Bayesian Ellipses in R, Journal of Animal Ecology, Volume 80 (2011) no. 3, pp. 595-602 | DOI

[62] Jaeger, A.; Connan, M.; Richard, P.; Cherel, Y. Use of stable isotopes to quantify seasonal changes of trophic niche and levels of population and individual specialisation in seabirds, Marine Ecology Progress Series, Volume 401 (2010), pp. 269-277 | DOI

[63] Jeltsch, F.; Bonte, D.; Pe'er, G.; Reineking, B.; Leimgruber, P.; Balkenhol, N.; Schröder, B.; Buchmann, C. M.; Mueller, T.; Blaum, N.; Zurell, D.; Böhning-Gaese, K.; Wiegand, T.; Eccard, J. A.; Hofer, H.; Reeg, J.; Eggers, U.; Bauer, S. Integrating movement ecology with biodiversity research - exploring new avenues to address spatiotemporal biodiversity dynamics, Movement Ecology, Volume 1 (2013) no. 1 | DOI

[64] Kernaléguen, L.; Arnould, J. P. Y.; Guinet, C.; Cazelles, B.; Richard, P.; Cherel, Y. Early-life sexual segregation: ontogeny of isotopic niche differentiation in the Antarctic fur seal, Scientific Reports, Volume 6 (2016) no. 1 | DOI

[65] Lewis, S.; Benvenuti, S.; Dall–Antonia, L.; Griffiths, R.; Money, L.; Sherratt, T. N.; Wanless, S.; Hamer, K. C. Sex-specific foraging behaviour in a monomorphic seabird, Proceedings of the Royal Society of London. Series B: Biological Sciences, Volume 269 (2002) no. 1501, pp. 1687-1693 | DOI

[66] Maher, W. J. Breeding Biology of the Snow Petrel near Cape Hallett, Antarctica, The Condor, Volume 64 (1962) no. 6, pp. 488-499 | DOI

[67] Main, M. B.; Weckerly, F. W.; Bleich, V. C. Sexual Segregation in Ungulates: New Directions for Research, Journal of Mammalogy, Volume 77 (1996) no. 2, pp. 449-461 | DOI

[68] Mancini, P. L.; Bond, A. L.; Hobson, K. A.; Duarte, L. S.; Bugoni, L. Foraging segregation in tropical and polar seabirds: Testing the Intersexual Competition Hypothesis, Journal of Experimental Marine Biology and Ecology, Volume 449 (2013), pp. 186-193 | DOI

[69] Martin, A. R.; Silva, V. M. F. River dolphins and flooded forest: seasonal habitat use and sexual segregation of botos ( Inia geoffrensis ) in an extreme cetacean environment, Journal of Zoology, Volume 263 (2004) no. 3, pp. 295-305 | DOI

[70] Mougin, J. Etude écologique de quatre espèces de pétrels antarctiques, L'Oiseau et la RFO, Volume 38 (1968)

[71] Neuhaus, P.; Ruckstuhl, K. Can the Activity Budget Hypothesis Explain Sexual Segregation in Desert Bighorn Sheep?, Behaviour, Volume 141 (2004) no. 5, pp. 513-520 | DOI

[72] Paiva, V. H.; Pereira, J.; Ceia, F. R.; Ramos, J. A. Environmentally driven sexual segregation in a marine top predator, Scientific Reports, Volume 7 (2017) no. 1 | DOI

[73] Parnell, A. C.; Inger, R.; Bearhop, S.; Jackson, A. L. Source Partitioning Using Stable Isotopes: Coping with Too Much Variation, PLoS ONE, Volume 5 (2010) no. 3 | DOI

[74] Peig, J.; Green, A. J. New perspectives for estimating body condition from mass/length data: the scaled mass index as an alternative method, Oikos, Volume 118 (2009) no. 12, pp. 1883-1891 | DOI

[75] Peig, J.; Green, A. J. The paradigm of body condition: a critical reappraisal of current methods based on mass and length, Functional Ecology, Volume 24 (2010) no. 6, pp. 1323-1332 | DOI

[76] Pereira, J. M.; Paiva, V. H.; Phillips, R. A.; Xavier, J. C. The devil is in the detail: small-scale sexual segregation despite large-scale spatial overlap in the wandering albatross, Marine Biology, Volume 165 (2018) no. 3 | DOI

[77] Phillips, R. A.; McGill, R. A. R.; Dawson, D. A.; Bearhop, S. Sexual segregation in distribution, diet and trophic level of seabirds: insights from stable isotope analysis, Marine Biology, Volume 158 (2011) no. 10, pp. 2199-2208 | DOI

[78] Phillips, R. A.; Silk, J. R. D.; Phalan, B.; Catry, P.; Croxall, J. P. Seasonal sexual segregation in twoThalassarchealbatross species: competitive exclusion, reproductive role specialization or foraging niche divergence?, Proceedings of the Royal Society of London. Series B: Biological Sciences, Volume 271 (2004) no. 1545, pp. 1283-1291 | DOI

[79] Phillips, R.; Wakefield, E.; Croxall, J.; Fukuda, A.; Higuchi, H. Albatross foraging behaviour: no evidence for dual foraging, and limited support for anticipatory regulation of provisioning at South Georgia, Marine Ecology Progress Series, Volume 391 (2009), pp. 279-292 | DOI

[80] Phillips, R. A.; Xavier, J. C.; Croxall, J. P. Effects of satellite transmitters on albatrosses and petrels, The Auk, Volume 120 (2003) no. 4 | DOI

[81] Pinet, P.; Jaquemet, S.; Phillips, R. A.; Le Corre, M. Sex-specific foraging strategies throughout the breeding season in a tropical, sexually monomorphic small petrel, Animal Behaviour, Volume 83 (2012) no. 4, pp. 979-989 | DOI

[82] Pinkerton, M. H.; Forman, J.; Bury, S. J.; Brown, J.; Horn, P.; O'Driscoll, R. L. Diet and trophic niche of Antarctic silverfishPleuragramma antarcticumin the Ross Sea, Antarctica, Journal of Fish Biology, Volume 82 (2013) no. 1, pp. 141-164 | DOI

[83] R Development Core Team R: A language and environment for statistical computing. Vienna: R Foundation for statistical computing,, 2015

[84] Rau, G.; Ainley, D.; Bengtson, J.; Torres, J.; Hopkins, T. 15N/14N and 13C/12C in Weddell Sea birds, seals, and fish: implications for diet and trophic structure, Marine Ecology Progress Series, Volume 84 (1992), pp. 1-8 | DOI

[85] Rayner, N. A. Global analyses of sea surface temperature, sea ice, and night marine air temperature since the late nineteenth century, Journal of Geophysical Research, Volume 108 (2003) no. D14 | DOI

[86] Ridoux, V.; Offredo, C. The diets of five summer breeding seabirds in Ad�lie Land, Antarctica, Polar Biology, Volume 9 (1989) no. 3, pp. 137-145 | DOI

[87] Roback, P. J.; Askins, R. A. Judicious Use of Multiple Hypothesis Tests, Conservation Biology, Volume 19 (2005) no. 1, pp. 261-267 | DOI

[88] Romey, W. L.; Wallace, A. C. Sex and the selfish herd: sexual segregation within nonmating whirligig groups, Behavioral Ecology, Volume 18 (2007) no. 5, pp. 910-915 | DOI

[89] Ruckstuhl, K. E. Sexual segregation in vertebrates: proximate and ultimate causes, Integrative and Comparative Biology, Volume 47 (2007) no. 2, pp. 245-257 | DOI

[90] Ruckstuhl, K.; Neuhaus, P. Sexual Segregation in Vertebrates, Cambridge University Press, Cambridge, 2005 | DOI

[91] Selander, R. K. Sexual Dimorphism and Differential Niche Utilization in Birds, The Condor, Volume 68 (1966) no. 2, pp. 113-151 | DOI

[92] Shaffer, S. A.; Costa, D. P.; Weimerskirch, H. Behavioural factors affecting foraging effort of breeding wandering albatrosses, Journal of Animal Ecology, Volume 70 (2001) no. 5, pp. 864-874 | DOI

[93] Shine, R. Ecological Causes for the Evolution of Sexual Dimorphism: A Review of the Evidence, The Quarterly Review of Biology, Volume 64 (1989) no. 4, pp. 419-461 | DOI

[94] Sims, D. W. Differences in habitat selection and reproductive strategies of male and female sharks. In: Sexual segregation in vertebrates: ecology of the two sexes ( KE Ruckstuhl and P Neuhaus , eds) In: Sexual segregation in vertebrates: ecology of the two sexes (KE Ruckstuhl and P Neuhaus, eds) (2005), pp. 127-147

[95] Spear, L. B.; Ainley, D. G. Morphological Differences Relative to Ecological Segregation in Petrels (Family: Procellariidae) of the Southern Ocean and Tropical Pacific, The Auk, Volume 115 (1998) no. 4, pp. 1017-1033 | DOI

[96] Tartu, S.; Bustamante, P.; Goutte, A.; Cherel, Y.; Weimerskirch, H.; Bustnes, J. O.; Chastel, O. Age-Related Mercury Contamination and Relationship with Luteinizing Hormone in a Long-Lived Antarctic Bird, PLoS ONE, Volume 9 (2014) no. 7 | DOI

[97] Thaxter, C. B.; Daunt, F.; Hamer, K. C.; Watanuki, Y.; Harris, M. P.; Grémillet, D.; Peters, G.; Wanless, S. Sex-specific food provisioning in a monomorphic seabird, the common guillemotUria aalge: nest defence, foraging efficiency or parental effort?, Journal of Avian Biology, Volume 40 (2009) no. 1, pp. 75-84 | DOI

[98] Tveraa, T.; Lorensten, S.-H.; Sæther, B.-E. Regulation of foraging trips and costs of incubation shifts in the Antarctic petrel (Thalassoica antarctica), Behavioral Ecology, Volume 8 (1997) no. 5, pp. 465-469 | DOI

[99] Van Franeker, J.; Williams, R. Diet of fulmarine petrels in the Windmill Islands, Wilkes Land, Antarctica. Preliminary results, Circumpolar Journal, Volume 7 (1992)

[100] Votier, S. C.; Bearhop, S.; Witt, M. J.; Inger, R.; Thompson, D.; Newton, J. Individual responses of seabirds to commercial fisheries revealed using GPS tracking, stable isotopes and vessel monitoring systems, Journal of Applied Ecology, Volume 47 (2010) no. 2, pp. 487-497 | DOI

[101] Wakefield, E. D.; Phillips, R. A.; Matthiopoulos, J.; Fukuda, A.; Higuchi, H.; Marshall, G. J.; Trathan, P. N. Wind field and sex constrain the flight speeds of central-place foraging albatrosses, Ecological Monographs, Volume 79 (2009) no. 4, pp. 663-679 | DOI

[102] Wearmouth, V. J.; Sims, D. W. Chapter 2 Sexual Segregation in Marine Fish, Reptiles, Birds and Mammals, Advances in Marine Biology, Elsevier, 2008, pp. 107-170 | DOI

[103] Weimerskirch, H. Are seabirds foraging for unpredictable resources?, Deep Sea Research Part II: Topical Studies in Oceanography, Volume 54 (2007) no. 3-4, pp. 211-223 | DOI

[104] Weimerskirch, H.; Lallemand, J.; Martin, J. Population sex ratio variation in a monogamous long-lived bird, the wandering albatross, Journal of Animal Ecology, Volume 74 (2005) no. 2, pp. 285-291 | DOI

[105] Weimerskirch, H.; Mougey, T.; Hindermeyer, X. Foraging and provisioning strategies of black-browed albatrosses in relation to the requirements of the chick: natural variation and experimental study, Behavioral Ecology, Volume 8 (1997) no. 6, pp. 635-643 | DOI

[106] Weimerskirch, H.; Salamolard, M.; Sarrazin, F.; Jouventin, P. Foraging strategy of wandering albatrosses through the breeding season: a study using satellite telemetry, Auk, Volume 110 (1993)

[107] Weimerskirch, H.; Shaffer, S.; Tremblay, Y.; Costa, D.; Gadenne, H.; Kato, A.; Ropert-Coudert, Y.; Sato, K.; Aurioles, D. Species- and sex-specific differences in foraging behaviour and foraging zones in blue-footed and brown boobies in the Gulf of California, Marine Ecology Progress Series, Volume 391 (2009), pp. 267-278 | DOI

[108] Widmann, M.; Kato, A.; Raymond, B.; Angelier, F.; Arthur, B.; Chastel, O.; Pellé, M.; Raclot, T.; Ropert-Coudert, Y. Habitat use and sex-specific foraging behaviour of Adélie penguins throughout the breeding season in Adélie Land, East Antarctica, Movement Ecology, Volume 3 (2015) no. 1 | DOI

[109] Wood, S.; Scheipl, F.; Wood, M. S. Package 'gamm4', American Statistician, Volume 45 (2017)

[110] Worton, B. J. Kernel Methods for Estimating the Utilization Distribution in Home-Range Studies, Ecology, Volume 70 (1989) no. 1, pp. 164-168 | DOI

[111] Yoccoz, N. G. Overuse, and misuse of significance tests in evolutionary biology and ecology, Bulletin of the Ecological Society of America, Volume 72 (1991)

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