Unusual pitted Ordovician brachiopods from the East Baltic: the significance of coarsely pitted ornamentations in linguliforms

A characteristic coarsely pitted ornamentation (with pits larger than 0.015 mm across) can be found on some very enigmatic and rare Lower Ordovician phosphatic-shelled linguliform brachiopods, including the paterinid Lacunites balaschovae Gorjansky and the obolid Foveola maarduensis Gorjansky. Both of these species are poorly understood and known from only a few specimens from the Floian of the East Baltic. Here we describe the Ordovician species Lacunites ivantsovi sp. nov. from the lower Darriwilian of the St Petersburg region and Foveola ivari sp. nov. from the Sandbian of Estonia, representing the last known survivors of these genera in the area. The new Darriwilian species of Lacunites is also one of the last members of the Class Paterinata, which most importantly includes some of the earliest known brachiopods from the early Cambrian. The coarsely pitted ornamentation of Foveola ivari is associated with the formation of asymmetrical so-called terrace lines in the umbonal area. The latter may indicate a burrowing life style. The pits in Lacunites and Foveola are directly comparable to the more or less identical coarse pits in the discinoid Trematis. Unlike the microscopic pits found in many fossil linguliforms, it is most likely that the coarse pits were empty in life; the minute surface ornamentation of irregular wrinkles in the outer primary layer continues across the bottom of the depressions in Foveola and Trematis; in the latter the cementing attachment discs of an attached phosphatic tubular Byronia-like organism also extend down into the pits.