Conflicting constraints on male mating success shape reward size in pollen-rewarding plants

Premise: Pollen-rewarding plants face two conflicting constraints: They must prevent consumptive emasculation while remaining attractive to pollen-collecting visitors. Small pollen packages (the quantity of pollen available in a single visit) may discourage visitors from grooming (reducing consumptive loss) but may also decrease a plant's attractiveness to pollen-collecting visitors. What package size best balances these two constraints? Methods: We modeled the joint effects of pollinators' grooming behaviors and package size preferences on the optimal package size (i.e., the size that maximizes pollen donation). We then used this model to examine Darwin's conjecture that selection should favor increased pollen production in pollen-rewarding plants. Results: When package size preferences are weak, minimizing package size reduces grooming losses and should be favored (as in previous theoretical studies). Stronger preferences select for larger packages despite the associated increase to grooming loss because loss associated with nonremoval of smaller packages is even greater. Total pollen donation increases with production (as Darwin suggested). However, if floral visitation declines or packages size preference increases with overall pollen availability, the fraction of pollen donated may decline as per-plant pollen production increases. Hence, increasing production may result in diminishing returns. Conclusions: Pollen-rewarding plants can balance conflicting constraints on pollen donation by producing intermediate-sized pollen packages. Strictly pollen-rewarding plants may have responded to past selection to produce