“…Correspondence between predictions of this theory and foraging behaviour in social insects has been investigated in bumblebees (see e.g., Pyke, 1978;Cartar and Dill, 1990;Harder, 1990;Waddington, 1995), honeybees (see e.g., Schmid-Hempel et al, 1985;Schmid-Hempel and Schmid-Hempel, 1987) and ants (see e.g., Traniello et al, 1984;Holder Bailey and Polis, 1987;Fewell, 1988;Crist and MacMahon, 1991;Ferster and Traniello, 1995;Fewell et al, 1996). Optimality models have recently been challenged by observations of "sub-optimal" performance at the individual level, in bees (Nunez, 1982) and ants (Kacelnik, 1993;Roces andNunez, 1993, Detrain et al, 1999).…”