The Embryology and Relationships of Dactylocladus (Crypteroniaceae) and a Discussion of the Family

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Abstract
Dactylocladus shares with Crypteronia and Axinandra the basic ordinal embryological features found in other Myrtales. The three genera also have and ephemeral endothecium and ephemeral nucellar tissue, a combination of embryological features unknown elsewhere in Myrtales and even in possibly related families such as Oliniaceae, Penaeaceae, Rhynchocalycaceae, and Alzateaceae. On the basis of embryology, Crypteroniaceae are sharply distinct from Melastomataceae, which are most often suggested as relatives, and from the four families above. Embryological evidence thus supports a recognition of Crypteroniaceae sensu stricto, composed of Crypteronia, Dactylocladus, and Axinandra. Within the family, the embryological features of Crypteronia are similar to those of Dactylocladus but differ from those of Axinandra. Neither Dactylocladus nor Crypteronia has an endothelium or the associated secondary multiplication of the inner integument, and both also differ from Axinandra in seed morphology. Embryologically, Axinandra seems to be the most specialized member of the family. The principal embryological difference between Crypteronia and Dactylocladus is in seed-coat structure. Therefore, in contrast with some earlier suggestions, embryological evidence suggests a relatively close affinity between Crypteronia and Dactylocladus and a more isolated position for Axinandra.

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