North American Black-Fruited Hawthorns. I. Variation in Floral Construction, Breeding System Correlates, and Their Possible Evolutionary Significance in Crataegus sect. Douglasii Loudon

Differences in the numbers of stamens and styles per flower are conspicuous features of variation in North American hawthorns (Crataegus L.). Variation in stamen number between individuals is discontinuous, with modes of approximately 20 and 10 (or fewer). In North American black-fruited section Douglasii Loudon the 10-stamen morphotype is exclusively polyploid, whereas the 20-stamen morphotype comprises both diploids and polyploids. Polyploidy is associated with apospory, self-fertility, and varying degrees of pollen sterility. Variation in stamen number may also be correlated with variation in distribution, phenology, leaf shape, and other features of floral morphology, leading to recognition of taxa at the specific or infraspecific level. Comparable variation in stamen number is virtually unknown in Eurasian hawthorns, as in the majority of flowering plants. In section Douglasii stamen number morphotypes have been recognized as either varieties or species; although correlations between stamen number and other features are not as straightforward as was previously surmised, the higher rank appears to be appropriate. These data on breeding system and morphological variation may be explainable in terms of hypotheses linking gametophytic apomixis, polyploidization, and optimal strategies for pollen dispersal.