Shell asymmetry in Cretaceous Cyclothyrididae (Brachiopoda): variability, ontogeny and terminology

The asymmetry observed in rhynchonellid brachiopod shells has been discussed for decades and continues to attract attention. This noteworthy modification of the anterior margin morphology during the ontogeny has evolved several times in rhynchonellids, and seems to reflect a genetic basis. First, we try to clarify the terminology regarding asymmetrical, dissymmetrical and symmetrical shells that has existed since the beginning of the 20th century. The Cretaceous populations observed clearly exhibit antisymmetry (also called random asymmetry). During the Cretaceous, some populations of Cyclothyris McKoy, 1844 include a mixture of truly asymmetrical specimens and others that exhibit an intermediate degree of asymmetry, herein called atypical morphologies. Shapes of specimens coming from two different locations in France were captured using geometric morphometrics. We used the range of different morphologies: 1) to test alternative hypotheses about the ontogeny of asymmetry; 2) to test for the possibility of several morphogroups; and 3) to discuss the determinism of the asymmetry.