Very low fruit set in milkweeds and other flowering plants often has b
een attributed to greater sexual selection on inflorescence size via m
ale, rather than female, reproductive success. Although this explanati
on has been generally accepted, alternate explanations have been prese
nted, and recently the ''male function'' or ''pollen donation'' hypoth
esis has been sharply criticized. In this paper, we make the distincti
on between selection on total flower number and on the size of inflore
scence units, both of which have been termed ''inflorescence size.'' W
e present an ESS model for the evolution of inflorescence design that
considers reproductive success through male and female function. The m
odel predicts that selection will balance the proportional changes in
female and male reproductive success resulting from changes in inflore
scence-unit size. We conducted a field study of selection on the size
of inflorescence units (umbers) by manipulating umbel size and number
in a natural population of Asclepias tuberosa, in southeastern Arizona
, during two reproductive seasons. We found that the male fitness func
tion reached a maximum at an intermediate umbel size in both years (al
though not significantly different from the smallest umbel size in eit
her year), whereas the female fitness function was highest for the sma
llest umbel size in one year, but was constant across umbel sizes in t
he other year. We also found that pollinator visitation rate correspon
ded well with male, but not female, function, and that between-year Va
riation in the male reproductive success of different umbel sizes corr
esponded with variation in the composition of the pollinator pool. Our
empirical results, when inserted in the model, predict ESS umbel size
s similar to those observed in the study population and the species th
roughout its range.