title: Helpers at the nest improve late-life offspring performance: evidence from a long-term study and a cross-foster experiment.
abstract: BACKGROUND
  Conditions during an individual's rearing period can have far reaching consequences for its survival and reproduction later in life. Conditions typically vary due to variation in parental quality and/or the environment, but in cooperative breeders the presence of helpers adds an important component to this. Determining the causal effect of helpers on offspring fitness is difficult, since high-quality breeders or territories are likely to produce high-quality offspring, but are also more likely to have helpers because of past reproductive success. This problem is best resolved by comparing the effect of both helping and non-helping subordinates on offspring fitness, however species in which both type of subordinates commonly occur are rare.
  
  
  RESULTS
  We used multi-state capture-recapture models on 20 years of data to investigate the effect of rearing conditions on survival and recruitment in the cooperatively breeding Seychelles warbler (Acrocephalus sechellensis), with both helping and non-helping subordinates. The number of helpers in the rearing territory, but not territory quality, group- or brood size, was positively associated with survival of offspring in their first year, and later in life. This was not a result of group size itself since the number of non-helpers was not associated with offspring survival. Furthermore, a nestling cross-foster experiment showed that the number of helpers on the pre-foster territory was not associated with offspring survival, indicating that offspring from territories with helpers do not differ in (genetic) quality.
  
  
  CONCLUSIONS
  Our results suggest that the presence of helpers not only increase survival of offspring in their first year of life, but also subsequent adult survival, and therefore have important fitness consequences later in life. This means that when calculating the fitness benefits of helping not only short-term but also the late-life benefits have to be taken into account to fully understand the evolution of cooperative breeding.
authors.affiliation: Animal Ecology group, Centre for Ecological and Evolutionary Studies, University of Groningen, Groningen, The Netherlands. L.Brouwer@MyScience.eu
authors.forename: Lyanne
authors.initials: L
authors.lastname: Brouwer
authors.forename: David S
authors.initials: DS
authors.lastname: Richardson
authors.forename: Jan
authors.initials: J
authors.lastname: Komdeur
mesh_terms: D000818:Animals; D001947:Breeding; D006379:Helping Behavior; D008018:Life Cycle Stages; D008136:Longevity; D009425:Nesting Behavior / Q000502:physiology*; D020308:Songbirds / Q000254:growth & development*