Sphaerocrystals in seeds of Euphorbia subgenus Esula (Euphorbiaceae): morphology, occurrence, and relationship with phylogeny

The term ‘sphaerocrystals’ (also sphaerites) in classic plant morphology was usually applied to spherical bodies visible in polarized light with a characteristic ‘Maltese cross’ and consisting of both inorganic and organic compounds. Initially sphaerocrystals were found in algal cells but were later also discovered in various organs of vascular plants (including seeds). Several authors noticed spherical particles on the seed surface and in testa cells of some Euphorbia species but a comprehensive study of this character in the genus (including the morphological origin of such particles) has not been carried out. We studied 72 species belonging to all 21 sections of Euphorbia subgenus Esula for presence of spherical particles on seed surface and in the testa using scanning, transmission, confocal laser electron microscopy, and light microscopy; histochemical and element analyses were also involved. We found that these particles have the same morphological features (including the demonstration of ‘Maltese cross’ in polarized light) as the ‘classic’ sphaerocrystals. We have recorded sphaerocrystals in 53 sampled species belonging to 20 sections of Euphorbia subgenus Esula; they are localized mostly in mesotesta cells and in some species are exuded onto the seed surface where they are often aggregated into clusters. Sphaerocrystals were absent in the sampled species (with one exception) of one of the major clades of the subgenus, formed by E. sect. Helioscopia and E. sect. Holophyllum. The absence of sphaerocrystals in this early divergent lineage of E. subgen. Esula may be related with adaptation of species of this lineage to more mesic habitats.