Pseudodiplostemony, and its implications for the evolution of the androecium in the Caryophyllaceae

The androecium of the Caryophyllaceae is varied, ranging from a two-whorled condition to a single stamen. A number of species belonging to the three subfamilies, Caryophyl-loideae, Alsinoideae and Paronychioideae have been studied ontogenetically with the SEM to understand their peculiar androecial development in the broader context of the Caryophyllales alliance. Although patterns of initiation are highly variable among species, there are three ontogenetic modes of stamen initiation: all stamens simultaneous within a whorl, the antepetalous stamens simultaneous and the antesepalous sequentially with a reversed direction, or both whorls sequentially with or without a reversed direction. The most common floral (ontogenetic) sequence of the Caryophyllaceae runs as follows: five sepals (in a 2/5 sequence), the stamens in front of the three inner sepals successively, stamens opposite the two outermost sepals, five antepetalous stamens (simultaneously or in a reversed spiral superimposed on the spiral of the antesepalous stamens), five outer sterile (petaloid) organs arising before, simultaneously or after the antesepalous stamens, often by the division of common primordia. A comparison with the floral configurations of the Phytolaccaceae and Molluginaceae indicates that the outer petaline whorl of the Caryophyllaceae corresponds positionally to the alternisepalous stamens of somePhytolacca, such asP. dodecandra. The difference withP. dodecandra lies in the fact that an extra inner or outer whorl is formed in the Caryophyl-laceae, in alternation with the sepals. A comparable arrangement exists in the Molluginaceae, though the initiation of stamens is centrifugal. A comparison of floral ontogenies and the presence of reduction series in the Caryophyllaceae support the idea that the pentamerous arrangement is derived from a trimerous prototype. Petals correspond to sterillized stamens and are comparable to two stamen pairs opposite the outer sepals and a single stamen alternating with the third and fifth sepals. Petals are often in a state of reduction; they may be confused with staminodes and they often arise from common stamenpetal primordia. The antesepalous stamen whorl represents an amalgamation of two whorls: initiation is reversed with the stamens opposite the fourth and fifth formed sepals arising before the other, while the stamens opposite the first and second formed sepals are frequently reduced or lost. Reductive trends are correlated with the mode of initiation of the androecium, as well as changes in the number of carpels, and affect the antesepalous and antepetalous whorls in different proportions. It is concluded that the androecium of the Caryophyllaceae is pseudodiplos-temonous and is not comparable to diplostemonous forms in the Dilleniidae and Rosidae. The basic floral formula of Caryophyllaceae is as follows: sepals 5—petals 5 (sterile stamens)—antesepalous stamens 3+2—antepetalous stamens 5 gynoecium 5.