Organophosphatic brachiopods: Patterns of biodiversification and extinction in the early Palaeozoic

From a database of 226 Cambrian - Ordovician genera of organophosphatic-sheIled brachiopods comprising the Subphylum Linguliformea, 11 sets of morphological characters typify all orders and superfamilies. Seven sets of these large-scale evolutionary novelties were established already by the end of the mid Cambrian, 2 more by the end of the late Cambrian, and the remaining 2 before the end of the Arenig. The earliest linguliformeans are of Tommotian age and represent some of the oldest known benthic organisms with a mineralised skeleton. Major diversification at the generic level took place during the mid and late Cambrian, by which time members of the Order Lingulida spread from near-shore to deep-water environments and became dominant in low diversity benthic assemblages that inhabited mobile sandy bottoms. There was a significant decline in diversity of linguliformeans during the latest Cambrian. Following recovery in the late Tremadoc-early Arenig, they then became one of the most distinctive components of benthic assemblages inhabiting marginal environments, e.g. eutrophic basins, shallow mobile sands, and abyssal depths. During the Llanvirn, there was a significant, worldwide turnover in linguliformean brachiopod faunas, when the majority of epibenthic lingulides of the families Obolidae, Zhanatellidae and Elkaniidae became extinct and were replaced, in shallow marine biofacies, by assemblages dominated by bivalved molluscs and burrowing lingulides. There is no trace until the mid Ashgill of a significant decline of micromorphic taxa in linguliformean assemblages characteristic of outer shelf environments. However, all siphonotretides, paterinides and most acrotretide genera disappeared towards the end of the late Ordovician pre-Hirnantian Dicellograptus anceps Biozone.