Helpers in primitively eusocial and cooperatively breeding animal societies forfeit their own reproduction to rear the offspring of a queen or breeding pair, but may eventually attain breeding status themselves. Kin selection¹ provides a widely accepted theoretical framework for understanding these societies, but differences in genetic relatedness do not explain a universal societal feature: the huge variation between individuals in helping effort^{2,3,4,5,6,7,8,9,10}. An alternative explanation for this variation lies in a fundamental trade-off faced by helpers: by working harder, they increase the indirect component of their fitness, but simultaneously decrease their own future survival and fecundity^2,4,8. Here, we show that individuals work less hard when they stand to lose more future fitness through working. We experimentally manipulated two components of future fitness in social queues of hover wasps (Stenogastrinae): a helper's chance of inheriting an egg-laying position, and the workforce available to rear her offspring should she inherit. After each manipulation, helpers increased or decreased their effort as appropriate to the change in expected future fitness that they experienced. Although helping provides significant indirect fitness benefits for hover wasps¹¹, our study shows that variation in the costs associated with helping is the major determinant of helping effort.

1. Department of Biology, University College London, Wolfson House, 4 Stephenson Way, London NW1 2HE, UK

Correspondence to: Jeremy Field¹ Correspondence and requests for materials should be addressed to J.F. (Email: jeremy.field@ucl.ac.uk).

Received 26 October 2005 |Accepted 5 January 2006

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