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 ('flag') that attracts insect visitors. Insects may use the flags as indicators of food rewards in the inflorescences below, as long-distance cues for locating and choosing flowering patches, or both. Clipping of all flags from patches of inflorescences in the field reduced the number of arriving insects, but not the total number of inflorescences and flowers visited by them. The number of flowers visited per inflorescence increased with inflorescence size, and inflorescence and flower visits rates significantly increased with patch size. Six percent 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. Removal of flags from all inflorescences in a patch reduced seed set in comparison with untreated controls, while flag clipping from ten randomly selected inflorescences in a patch did not decrease seed production. These results suggest that flags signal long-distance information to potential pollinators (possibly 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 signal production. Greenhouse-grown S. viridis plants allocated a low proportion of their biomass to flags. Plants grown under water stress did not reduce biomass allocation to flags as compared to irrigated controls. Water loss rates of picked flags were lower than those of picked leaves. These findings suggest that the expenses of flag production and maintenance are modest, reducing the selective advantage of individuals that do not carry flags. We discuss additional potential evolutionary mechanisms that may select for flag production.