Optimal sugar concentrations of floral nectars -dependence on sugar intake efficiency and foraging costs.

A model is developed to elucidate the determinants of sugar concentrations in flower nectars. This model analyses the efficiency of sugar intake, or energy flux, which for nectarivores closely approximates the rate of net energy gain. For both steady state and some non-steady flows of nectars, this energy flux is shown to be maximal at particular sugar concentrations referred to here as the maximum flux concentration. Higher concentrations actually yield lower energy intake rates because the concomitant rapid increase in viscosity sharply reduces the rate of fluid intake. For pure sucrose solutions, the maximum flux concentration is 22%. For flower nectars, which are chemically more complex, the maximum flux concentration is predicted to be closer to 26%, using the first viscosity measures obtained for flower nectars. This concentration is shown to be essentially independent of the pollinator's feeding organ morphology and of the type of potential inducing nectar flow. It is proposed that this concentration applies for virtually all pollinators that select nectars with maximal energy flux.However not all pollinators are expected to select such nectars because this 26% concentration is not necessarily "optimal". The model predicts that optimal sugar concentrations vary for particular pollinators as a function of two primary factors: (1) the energy flux derived from the nectar, as discussed above, as well as (2) the relative contribution of transit costs to overall foraging costs. Relatively "dilute" nectars, with sugar concentrations close to the maximal flux value, are predicted for flowers pollinated by organisms that minimize feeding time to reduce high feeding costs, such as that of hovering or of exposure to enhanced predation while feeding. More concentrated nectars are predicted for flowers pollinated by nectarivores that incur high foraging transit costs relative to feeding costs.Flowers pollinated by hovering pollinators, including many hummingbirds, hawkmoths and bats, have nectars with mean sugar concentrations in close accord with the 26% maximum flux concentration predicted. Moreover, these nectars have relatively low concentrations of nonsugar constituents, which increase viscosity and thereby decrease sugar flux. Over 75% of the flowers examined in this study, which are pollinated primarily by territorial hummingbird species, provide nectars that allow sugar uptake with an efficiency of 90% or greater of the maximal value. According to the model, these data suggest that feeding costs of these pollinators far outweigh foraging transit costs. In contrast, the model suggests that flower nectars taken by traplining hummingbirds and by bees, with sugar concentrations significantly above the maximum flux value, reflect the higher costs of foraging flight relative to costs of feeding for these pollinators.Increasing temperature decreases nectar viscosity, and thereby increases absolute nectar uptake rates sharply. This leads to a number of predictions regarding foraging behavior as well as flower location, orientation, and color. However, the maximum flux concentration is shown to be practically invariable over a wide range of temperatures-increasing by only 2% sugar from 10°C to 30°C. Thus, contrary to previous expectations, little change in average sugar concentrations of flowers pollinated by particular groups of nectarivores is expected from cooler to warmer regions.