The signaling function of an extra-floral display: what selects for signal development?

The vertical inflorescences of the Mediterranean annual Salvia viridis carry many small, colorful flowers, and are frequently terminated by a conspicuous tuft of colorful leaves ("flags") that attracts insect pollinators. Insects may use the flags as indicators of the food reward in the inflorescences, as long-distance cues for locating and choosing flowering patches, or both. Clipping of flags from patches of inflorescences in the field significantly reduced the number of pollinators that arrived at the patches, but not the total number of inflorescences and flowers visited by them. The number of flowers visited per inflorescence significantly increased with inflorescence size, however. Inflorescence and flower visits rates signific antly increased with patch size when flags were present, but not after flag removal. 6% of the plants in the study population did not develop any flag during blooming, yet suffered no reduction in seed set as compared to flag-bearing neighboring individuals. These results suggest that flags signal long-distance information to pollinators (perhaps indicating patch location or size), while flower-related cues may indicate inflorescence quality. Plants that do not develop flags probably benefit from the flag signals displayed by their neighbors, without bearing the costs of flag production. Thus, flagproducing plants can be viewed as altruists that enhance their neighbors' fitness. Greenhouse-grown S. viridis plants allocated = 0.5% of their biomass to flag production, and plants grown under water stress did not reduce their biomass allocation to flags as compared to irrigated controls. These findings suggest that the expenses of flag production are modest, perhaps reducing the cost of altruism. We discuss additional potential evolutionary mechanisms that may select for the maintenance of flag production.