Studies on the evolution of cooperative behaviour are typically confined to understanding its adaptive value. It is equally essential, however, to understand its potential to evolve, requiring knowledge about the phenotypic consistency and genetic basis of cooperative behaviour. While previous observational studies reported considerably high heritabilities of helping behaviour in cooperatively breeding vertebrates, experimental studies disentangling the relevant genetic and non-genetic components of cooperative behaviour are lacking. In a half-sibling breeding experiment, we investigated the repeatability and heritability of three major helping behaviours performed by subordinates of the cooperatively breeding fish Neolamprologus pulcher. To experimentally manipulate the amount of help needed in a territory, we raised the fish in two environments differing in egg predation risk. All three helping behaviours were significantly repeatable, but had very low heritabilities. The high within-individual consistencies were predominantly due to maternal and permanent environment effects. The perceived egg predation risk had no effect on helping, but social interactions significantly influenced helping propensities. Our results reveal that developmentally plastic adjustments of provided help to social context shape cooperative phenotypes, whereas heritable genetic variation plays a minor role.

Helping behaviour in the cichlid Neolamprologus pulcher540 observations of 3 major alloparental brood care behaviours of the cooperatively breeding cichlid fish Neolamprologus pulcher bred in a paternal half-sibling breeding design. Includes dependent variables egg cleaning, digging, and defence against egg predator Telmatochromis vittatus of individuals during a test or a repeated test, covariates that predicted helping behaviours and random effects for quantitative genetic analyses.data_cichlid_helping.xlsx