Selection toward shorter flowers by butterflies whose probosces are shorter than floral tubes

Darwin's meticulous observations on the function of oral shape led to his famous prediction of a long-tongued pollinator, which he believed to be the evolutionary trigger for the long-spurred flowers of the Madagascar star orchid. Although tubular flowers are common, long tubes or spurs are an exception, suggesting that selection maintaining short flowers is widespread. Using the butterfly-pollinated carnation Dianthus carthusianorum and two butterfly species differing in proboscis length (Melanargia galathea and Inachis io) as model organisms, we experimentally demonstrate a reduction in pollinator efficiency with an increasing difference between proboscis length and floral tube length. Such a relationship is a prerequisite for the evolution of floral shape in response to pollinator morphology.