Functional morphology and the evolution of the cystoid Echinosphaerites

Echinosphaerites, known from the Lower and Middle Ordovician, has branched biserial brachioles. Such features are so far unknown in the Rhombifera. Echinosphaerites had a skeletal meshwork like that of other blastozoan echinoderms, with a fine outer mesh layer and an inner coarse mesh layer. During evolution the number and location of brachioles, including the pattern of brachiole branching, changed by increase in the number of brachioles and increased complexity of the branching pattern. The exothecal pore structures increased in the complexity of patterns of tangential canals. The pattern of skeletal growth in Echinosphaerites is discussed. The Echinosphaeritidae and Caryocystitidae families are closely related and show parallel development.