Sendino, C., Zágoršek, K. & Taylor, P.D. 2012: Asymmetry in an Ordovician conulariid cnidarian. Lethaia, Vol. 45, pp. 423–431.

Conulariids are fossils of the presumed polyps of an extinct scyphozoan cnidarian group. Their cone-shaped skeletons normally show perfect tetraradial symmetry. However, in the Ordovician species Metaconularia anomala (Barrande 1867) from Drabov (Czech Republic), tetraradial symmetry is compromised in three ways: (1) the skeleton often shows torsion; (2) the four sides may vary in width at the same level within one individual; and (3) one side may be deleted to give a triradial skeleton. Almost 2000 specimens were studied in museum collections. About 56% of analysed specimens are twisted in an anticlockwise direction (sinistral) when viewed from the apex towards the aperture, 28% show no torsion, 1% exhibit clockwise torsion (dextral) and the remaining 15% cannot be classified. Maximum measured torsion rate was 1.5°/mm. A significant negative correlation between torsion rate and length suggests that more highly torted individuals may have survived less well. Almost 5% of individuals show loss of one side for at least part of their lengths. Although many individuals have four sides of equal width, in a significant proportion the sides are of unequal width, up to a maximum/minimum side width ratio of 2 (i.e. widest face twice the width of the narrowest). In the absence of a satisfactory taphonomic model to explain the asymmetries, they are regarded as mirroring asymmetries in the living conulariids, with the strong preference for sinistral torsion interpreted as an example of a fixed asymmetry that was genetically controlled and heritable. It is speculated that the signalling protein Nodal as well as Hox-like genes were involved in controlling the asymmetries described in M. anomala.