In unpredictable environments, any tactic that enables avian parents to adjust brood size and, thus, energy expenditure to environmental conditions should be favoured. Hatching asynchrony (HA), which occurs whenever incubation commences before clutch completion, may comprise such a tactic. For instance, the sibling rivalry hypothesis states that the hierarchy among chicks, concomitant to HA, should both facilitate the adjustment of brood size to environmental conditions and reduce several components of sibling competition as compared to synchronous hatching, at both brood and individual levels. We thus predicted that brood aggression, begging and feeding rates should decrease and that older chick superiority should increase with HA increasing, leading to higher growth and survival rates. Accordingly, we investigated the effects of an experimental upward and downward manipulation of HA magnitude on behaviour, growth and survival of black-legged kittiwake (Rissa tridactyla) chicks. In line with the sibling rivalry hypothesis, synchronous hatching increased aggression and tended to increase feeding rates by parents at the brood level. Begging rates, however, increased with HA contrary to our expectations. At the individual level, as HA magnitude increased, the younger chick was attacked and begged proportionally more often, experienced a slower growth and a higher mortality than its sibling. Overall, the occurrence of energetic costs triggered by synchronous hatching both for parents and chicks, together with the lower growth rate and increased mortality of the younger chick in highly asynchronous broods suggest that natural HA magnitude may be optimal.
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We thank J.-B. Ferdy and F. Helfenstein for their statistical advice. We also thank J. White, an anonymous referee, and the associate editor Ian Hartley for their critical comments on earlier versions of the manuscript. The study was financed within a 4-year grant from the French Polar Institute Paul-Emile Victor (IPEV ‘Programme 1162 SexCoMonArc’). This work originated in the lab EDB, part of the ‘Laboratoire d'Excellence’ (LABEX) entitled TULIP (ANR-10-LABX-41).
This experiment was conducted under the approval of the USGS Alaska Science Center Animal Care and Use Committee, the IPEV Ethical Committee, in accordance with U.S. laws and under permits from the U.S. Fish and Wildlife Service and the State of Alaska. Any use of trade names is for descriptive purposes only and does not imply endorsement of the U.S. Government.
Benowitz-Fredericks ZM, Kitaysky AS, Welcker J, Hatch SA (2013) Effects of food availability on yolk androgen deposition in the black-legged kittiwake (Rissa tridactyla), a seabird with facultative brood reduction. PLoS ONE 8:e62949PubMedCentralPubMedCrossRefGoogle Scholar
Blanchard P, Hanuise N, Dano S, Weimerskirch H (2007) Offspring sex ratio in relation to parental structural size and body condition in the long-lived wandering albatross (Diomedea exulans). Behav Ecol Sociobiol 61:767–773CrossRefGoogle Scholar
Box G, Cox D (1964) An analysis of transformations. J R Stat Soc B Met 26:211–252Google Scholar
Braun BM, Hunt GLJ (1983) Brood reduction in black-legged kittiwakes. Auk 100:469–476Google Scholar
Drent RH, Daan S (1980) The prudent parent: energetic adjustments in avian breeding. Ardea 68:225–252Google Scholar
Lack D (1954) The natural regulation of animal numbers. Oxford University Press, OxfordGoogle Scholar
Leclaire S, Helfenstein F, Degeorges A, Wagner RH, Danchin É (2010) Family size and sex-specific parental effort in black-legged kittiwakes. Behaviour 147:13–14CrossRefGoogle Scholar
Leclaire S, Bourret V, Wagner RH, Hatch SA, Helfenstein F, Chastel O, Danchin É (2011) Behavioral and physiological responses to male handicap in chick-rearing black-legged kittiwakes. Behav Ecol 22:1156–1165CrossRefGoogle Scholar
Mainwaring MC, Blount JD, Hartley IR (2012) Hatching asynchrony can have long-term consequences for offspring fitness in zebra finches under captive conditions. Biol J Linn Soc 106:430–438CrossRefGoogle Scholar
Maynard-Smith J (1982) Evolution and the theory of games. Cambridge University Press, CambridgeCrossRefGoogle Scholar
Maynard-Smith J, Parker GA (1976) The logic of asymmetric contests. Anim Behav 24:159–175CrossRefGoogle Scholar
Merkling T, Leclaire S, Danchin E, Lhuillier E, Wagner RH, White J, Hatch SA, Blanchard P (2012) Food availability and offspring sex in a monogamous seabird: insights from an experimental approach. Behav Ecol 23:751–758CrossRefGoogle Scholar
Roulin A, Dreiss AN (2012) Sibling competition and cooperation over parental care. In: Royle NJ, Smiseth PT, Kölliker M (eds) The evolution of parental care. Oxford University Press, Oxford, pp 133–149CrossRefGoogle Scholar
Royle NJ, Hamer KC (1998) Hatching asynchrony and sibling size hierarchies in gulls: effects on parental investment decisions, brood reduction and reproductive success. J Avian Biol 29:266–272CrossRefGoogle Scholar
Siegel RB, Weathers WW, Beissinger SR (1999) Hatching asynchrony reduces the duration, not the magnitude, of peak load in breeding green-rumped parrotlets (Forpus passerinus). Behav Ecol Sociobiol 45:444–450CrossRefGoogle Scholar
Smiseth PT, Morgan K (2009) Asynchronous hatching in burying beetles: a test of the peak load reduction hypothesis. Anim Behav 77:519–524CrossRefGoogle Scholar
Stearns S (1992) The evolution of life histories. Oxford University Press, OxfordGoogle Scholar
Stenning MJ (1996) Hatching asynchrony, brood reduction and other rapidly reproducing hypotheses. Trends Ecol Evol 11:243–246PubMedCrossRefGoogle Scholar
Vallarino A, Evans N, Daunt F, Wanless S, Nager R (2012) Egg components vary independently of each other in the facultative siblicidal black-legged Kittiwake Rissa tridactyla. J Ornithol:1–11Google Scholar
White J, Leclaire S, Kriloff M, Mulard H, Hatch SA, Danchin E (2010) Sustained increase in food supplies reduces broodmate aggression in black-legged kittiwakes. Anim Behav 79:1095–1100CrossRefGoogle Scholar
Wiebe KL, Bortolotti GR (1994a) Food supply and hatching spans of birds: energy constraints or facultative manipulation. Ecology 75:813–823CrossRefGoogle Scholar
Wiebe KL, Bortolotti GR (1994b) Energetic efficiency of reproduction: the benefits of asynchronous hatching for American kestrels. J Anim Ecol:551–560Google Scholar
Wiebe KL, Bortolotti GR (2000) Parental interference in sibling aggression in birds: what should we look for? Ecoscience 7:1–9Google Scholar
Wiehn J, Ilmonen P, Korpimäki E, Pahkala M, Wiebe KL (2000) Hatching asynchrony in the Eurasian kestrel Falco tinnunculus: an experimental test of the brood reduction hypothesis. J Anim Ecol 69:85–95CrossRefGoogle Scholar
Williams GC (1966) Natural selection, the costs of reproduction, and a refinement of Lack's principle. Am Nat 100:687–690CrossRefGoogle Scholar