We hypothesize that pollination efficiency selects for equal distances between the pollinator reward and the anthers, and the stigmas, creating an adaptive ridge. We predict that this fitness surface governs the divergence of many plant species. We use the theory of adaptive accuracy, precision and mean optimality to assess how close populations lie to the hypothesized adaptive ridge and which factors contribute to departure from the optimum. Patterns of accuracy of pollen placement and receipt were compared across species in three study systems, Dalechampia (Euphorbiaceae), Collinsieae (Plantaginaceae) and Stylidium (Stylidiaceae), in order to assess the roles of stamen/stigma imprecision and population mean departure from the optimum in the generation of floral inaccuracy. We found that population mean departure from the optimum was the most important factor in Dalechampia, female imprecision and departure from the optimum were about equally important factors in Collinsieae, and stamen and stigma imprecision were equally important in Stylidium, with virtually no departure from the optimum. Possible reasons for imprecision and departure from the optimum were assessed using phylogenetically informed methods, indicating important roles of limited floral integration in the generation of imprecision, and conflicting selective pressures, associated with outcrossing, in the generation of departure from the optimum. New Phytologist (2009) 183: 600-617doi: 10.1111/j.1469-8137.2009.02930.x.