Two speciation events occurred in the Eocene lineage beginning with Echinocythereis isabenana Oertli (Aragon, Spain). After a long period of stasis in ornament and shape/size, this species underwent a relatively rapid decrease in size, accompanied by ornamental changes. The transition required about 30,000 yr. The new species E. aragonensis Oertli is significantly smaller (20% decrease in size). The lateral papillation is denser and less regular, the papillae being significantly smaller than in the ancestral form. This ornament gradually yields to increasing frequencies of individuals with reduced papillae superimposed on a progressively better developed network of filaments. This change, which is regionally valid, is ultimately replaced by fully expressed reticulations, on which minute pustules may occur; this development typifies the third species, E. posterior Oertli. Some ornamental characteristics of E. aragonensis are anticipated in rare individuals of E. isabenana. Ornamental features of E. posterior appear in late larvae of E. isabenana. Generally, E. aragonensis and E. posterior do not differ significantly in size. A trend toward smaller carapaces in the upper samples of E. posterior could be related to ecophenotypic effects.
All species are highly polymorphic for both shape and ornament; the morphs are not sharply bounded and are mostly of uncertain evolutionary status. The transition from isabenana to aragonensis is heralded by a sharp rise in the coefficient of variation for length measures of the carapace. There is a pronounced multivariate morphometric discontinuity between isabenana and the descendant forms, which are largely multivariately homogeneous. The isabenana-aragonensis transition could have occurred by selection or by random genetic drift. Both models can be accommodated by the data; however, the weight of evidence favors the former hypothesis. Evolution in the Echinocythereis lineage seems to have occurred by two different mechanisms. The first transition was rather rapid, with many disjunct features. The second was slow, the changes being more of degree than of kind.