Optimal foraging, plant density and the marginal value theorem.

The stochastic, discrete analogue of the marginal value theorem predicts that as the cost of moving between plants increases, bees should increase the percentage of the available flowers which they visit per plant. This prediction was tested using two populations of Polemonium foliosissimum and their primary pollinators Bombus flavifrons and B. bifarius. The results of these tests were equivocal. Bees did not perform exactly as the marginal value theorem predicted they should to maximize their rate of net energy intake. Instead of visiting more flowers per plant as movement costs increased bees were observed to alter their behavior in other ways in an attempt to maximize their rate of net energy intake. They were demonstrated to be flying randomly with respect to direction, flying short flight distances relative to the plant spacing distances encountered, flying predominately between nearest neighbor plants, and to be visiting flowers of other plant species while enroute from one P. foliosissimum flower to another P. foliosissimum flower. Such behavioral flexibility strongly implies that optimal foraging models which predict a shift in any particular behavior in response to environmental conditions are too simplistic to accurately predict foraging behavior.