Abstract
Hatching asynchrony of nestling birds leads to weight asymmetries, which in turn affect the nestlings’ relative success when competing for feedings brought to the nest. We present a game theoretic model that predicts how weight asymmetry influences the nestlings’ energy on securing feedings, thus determining the caloric value remaining for weight gain as well as the distribution of feedings obtained. The model has a unique Evolutionary Stable Strategy (ESS) profile, in which nestlings in more asymmetric nests exhibit less aggression and hence achieve larger weight gain per feeding. The impetus for this model was data from a long-term study of Arabian babblers (Turdoides squamiceps) that showed a surprising negative correlation between the number of feedings that a nest received and the overall weight gain in the nest. This finding is, however, entirely consistent with our model—in more symmetric nests, the individuals fight more and are consequently hungrier and beg for additional food, are fed more, but still gain less weight due to the higher energetic costs of fighting. The model provides a fundamental explanation also for related findings in other species, in which chicks in asynchronous broods were found to be heavier than those from synchronous broods. In addition, it supports the sibling rivalry hypothesis by which brood asynchrony may diminish aggressive interactions among nestlings, leading to more efficient use of resources.
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References
Amundsen T, Slagsvold T (1991) Hatching asynchrony: facilitating adaptive or maladaptive brood reduction. Acta XX Int Ornithol Cong III:1707–1719
Amundsen T, Slagsvold T (1998) Hatching asynchrony in great tits: a bet-hedging strategy? Ecology 79:295–304
Clark AB, Wilson DS (1981) Avian breeding adaptations: hatching asynchrony, brood reduction, and nest failure. Q Rev Biol 56:253–277
Cook MI, Monaghan P, Burns MD (2000) Effects of short-term hunger and competitive asymmetry on facultative aggression in nestling black guillemots Cepphus grille. Behav Ecol 11:282–287
Corchon LC (2007) The theory of contests: a survey. Rev Econ Des 11:69–100
Cornes R, Hartley R (2005) Asymmetric contests with general technologies. Econ Theor 26:923–946
Forbes LS (1991) Hunger and food allocation among nestlings of facultatively siblicidal ospreys. Behav Ecol Sociobiol 29:189–195
Forbes S (2011) Social rank governs the effective environment of siblings. Biol Lett 7:346–348
Forbes S, Glassey B (2000) Asymmetric sibling rivalry and nestling growth in red-winged blackbirds (Agelaius phoeniceus). Behav Ecol Sociobiol 48:413–417
Fujioka M (1987) Sibling competition and siblicide in asynchronously-hatching broods of the cattle egret Bubulcus ibis. Anim Behav 33:1228–1242
Godfray HCJ (1991) The signalling of need by offspring to their parents. Nature 353:328–330
Godfray HCJ (1995) Signalling of need between parents and young: parent–offspring conflict and sibling rivalry. Am Nat 146:1–24
Godfray HCJ, Parker GA (1992) Sibling competition, parent–offspring conflict, and clutch size. Anim Behav 43:473–490
Hanh DC (1981) Asynchronous hatching in the laughing gull: cutting losses and reducing rivalry. Anim Behav 29:421–427
Haskell D (1994) Experimental evidence that nestling begging behaviour incurs a cost due to nest predation. Proc R Soc Lond B 257:161–164
Hussell DJT (1972) Factors affecting clutch size in Arctic passerines. Ecol Monogr 42:881–892
Kilner R (2001) A growth cost of begging in captive canaries. Proc Natl Acad Sci U S A 98:11394–11398
Kilner R, Johnstone RA (1997) Begging the question: are offspring solicitation behaviours signals of need? Trends Ecol Evol 12:11–15
Konrad KA (2009) Strategy and dynamics in contests. Oxford Univ Press, Oxford
Konrad KA (2010) Dynamic contests. Discussion Paper SP II 2010, Max Planck Institute, available online at http://ssrn.com/abstract=1695939
Lack D (1947) The significance of clutch-size. Parts 1 and 2. Ibis 89:302–352
Leech SM, Leonard ML (1997) Begging and the risk of predation in nestling birds. Behav Ecol 8:644–646
Macnair M, Parker GA (1979) Models of parent-offspring conflict. III. Intra-brood conflict. Anim Behav 27:1202–1209
Mock DW, Parker GA (1997) The evolution of sibling rivalry. Oxford Univ Press, New York
Mock DW, Ploger BJ (1987) Parental manipulation of optimal hatch asynchrony in cattle egrets: an experimental study. Anim Behav 35:150–160
Mock DW, Schwagmeyer PL (1990) The peak load reduction hypothesis for avian hatching asynchrony. Evol Ecol 4:249–260
Mock DW, Dugas MB, Strickler SA (2011) Honest begging: expanding from signal of need. Behav Ecol 22:909–917
O’Connor RJ (1978) Brood reduction in birds: selection for fratricide, infanticide and suicide? Anim Behav 26:79–96
Osorno JL, Drummond H (1995) The function of hatching asynchrony in the blue-footed booby. Behav Ecol Sociobiol 37:265–273
Ostreiher R (1997) Food division in the Arabian babbler nest: adult choice or nestling competition? Behav Ecol 8:233–238
Ostreiher R (1999) Nestling feeding space strategy. Auk 116:651–657
Ostreiher R (2001) The importance of nestling location for obtaining food in open cup-nests. Behav Ecol Sociobiol 49:340–347
Parker GA, Macnair M (1979) Models of parent-offspring conflict. IV. Suppression: evolutionary retalization by the parent. Anim Behav 27:1210–1235
Parker GA, Mock DW, Lamey TC (1989) How selfish should stronger sibs be? Am Nat 133:846–868
Parker GA, Royle NJ, Hartley IR (2002a) Begging scrambles with unequal chicks: interactions between need and competitive ability. Ecol Lett 5:206–215
Parker GA, Royle NJ, Hartley IR (2002b) Intrafamilial conflict and parental investment: a synthesis. Philos T Roy Soc B 357:295–307
Porkert J, Spinka M (2006) Begging in common redstart nestlings: scramble competition or signalling of need? Ethology 112:398–410
Rodríguez-Gironés MA (1999) Sibling competition stabilizes signaling resolution models of parent–offspring conflict. Proc R Soc Lond B 266:2399–2402
Rodríguez-Gironés MA, Enquist M, Lachmann M (2001a) Role of begging and sibling competition in foraging strategies of nestlings. Anim Behav 61:733–745
Rodríguez-Gironés MA, Zuniga JM, Redondo T (2001b) Effects of begging on growth rates of nestling chicks. Behav Ecol 12:269–274
Royle NJ, Hartley IR, Owens IPF, Parker GA (1999) Sibling competition and the evolution of growth rates in birds. Proc R Soc Lond B 266:923–932
Schwagmeyer PL, Mock DW (2008) Parental provisioning and offspring fitness: size matters. Anim Behav 75:291–298
Stein WE (2002) Asymmetric rent-seeking with more than two contestants. Public Choice 113:325–336
Stenning MJ (1996) Hatching asynchrony, brood reduction and other rapidly reproducing hypotheses. Trends Ecol Evol 11:243–246
Vinuela J (2000) Opposing selective pressures on hatching asynchrony: egg viability, brood reduction, and nestling growth. Behav Ecol Sociobiol 48:333–343
Weibull JO (1995) Evolutionary game theory. Mass Inst Technol, Cambridge
Wiebe KL, Bortolotti GR (1994) Energetic efficiency of reproduction: the benefits of asynchronous hatching of American kestrels. J Anim Ecol 63:551–560
Wright J (1997) Helping-at-the-nest in Arabian babblers: signalling social status or sensible investment in chicks? Anim Behav 54:1439–1448
Wright J (1998) Helpers-at-the-nest have same provisioning rule as parents: experimental evidence from playbacks of chick begging. Behav Ecol Sociobiol 42:423–429
Zahavi A (1989) The Arabian babbler. In: Newton I (ed) Lifetime reproduction in birds. Acad Press, London, pp 253–275
Zahavi A (1990) Arabian babbler. In: Stacey PB, Koenig WD (eds) Cooperative breeding in birds. Camb Univ Press, London, pp 103–130
Acknowledgments
We wish to thank Amotz and Avishag Zahavi, Avner Anava, Yoel Perel, and Yuval Dagay for their help during fieldwork as well as for fruitful discussions, the Society for the Protection of Nature in Israel (SPNI) and Hazeva Field Study Center for the long-term hospitality, and Scott Forbes and an anonymous reviewer for critical comments. We also wish to thank the MEDS Department at Kellogg School of Management, Northwestern University for their hospitality and collegiality to A.H. during a 2-year visit there when the theoretical portions of this study were completed. S.P-J. was supported, in part, by The Women’s Board at University of Chicago.
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Communicated by M. Leonard
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Ostreiher, R., Pruett-Jones, S. & Heifetz, A. Asymmetric contests at the nest. Behav Ecol Sociobiol 66, 1237–1246 (2012). https://doi.org/10.1007/s00265-012-1377-5
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DOI: https://doi.org/10.1007/s00265-012-1377-5