Clutch size determination in shorebirds: revisiting incubation limitation in the pied avocet (Recurvirostra avosetta)


  • Szabolcs Lengyel,

    Corresponding author
    1. Department of Ecology, University of Debrecen, 4032 Debrecen, Egyetem tér 1., Hungary;
    Search for more papers by this author
    • Present addresss: Department of Zoology, North Carolina State University, 100 Brooks Avenue, Raleigh, NC 27695-7617, USA

  • Béla Kiss,

    1. BioAquaPro Environmental Consulting LLC, 4032 Debrecen, Soó R. u. 21., Hungary; and
    Search for more papers by this author
  • C. Richard Tracy

    1. Ecology, Evolution, and Conservation Biology MS-315, University of Nevada, Reno, NV 89557, USA
    Search for more papers by this author

*Correspondence author. E-mail:


  • 1Traits strongly related to fitness, such as offspring number, are expected to show intraspecific variation among individuals. However, offspring number is invariant in several reptiles, birds, and mammals. Most shorebirds (210+ species), for example, have an invariant clutch size of four eggs, which is unexpected in such an ecologically, behaviourally and socially diverse group.
  • 2The incubation-limitation hypothesis (ILH) suggests that shorebird clutch size is limited by the inability of adults to incubate clutches larger than four eggs. Several recent studies reported no overall costs of incubating experimentally enlarged clutches and concluded no support for the traditional ILH. However, most studies have not measured all potential costs, and none has quantified costs beyond egg hatching. We conducted a clutch-enlargement experiment and measured potential costs both during incubation and chick rearing in pied avocets (Recurvirostra avosetta L.).
  • 3Hatching was more asynchronous and egg hatchability was marginally lower in enlarged clutches than in controls. Nonetheless, more young hatched from enlarged clutches (mean: 4·2 ± 0·17 SE) than from controls (3·4 ± 0·09), and the two groups did not differ in incubation period, complete or partial clutch failure, or hatchling body size, apparently refuting the ILH.
  • 4However, pairs incubating enlarged clutches occupied poorer feeding territories during chick rearing, experienced higher chick mortality, and eventually raised fewer young to independence (mean adjusted for season: 0·7 ± 0·16 SE juveniles) than did control pairs (1·2 ± 0·13). Chick survival was primarily associated with prey availability, and predation risks were not higher in larger broods.
  • 5Our results provide evidence that incubating unusually large clutches can affect post-hatching performance and lead to lower annual reproductive success in shorebirds. This study, therefore, supports the ILH and points to the importance of monitoring reproductive success beyond the hatching of the chicks.