Middle Ordovician harknessellid brachiopods (Dalmanellidina) from the Mediterranean margin of Gondwana

Abstract

The family Harknessellidae Bancroft, 1928 (Orthida, Dalmanellidina) was designed to embrace an assemblage of species referred previously to Harknessella Reed, 1917, and included five genera known mainly from the Middle and Upper Ordovician of England. Herein, we suggest reassigning to this family the genus Cacemia Mitchell, 1974, widespread in the middle Darriwilian (upper Middle Ordovician) of the Iberian and Armorican massifs. Since its designation, Cacemia was placed among the dalmanellidin heterorthids, in spite of its strongly mucronate hinge line, which is totally unknown within this Mediterranean family. A new harknessellid has been identified from the upper Darriwilian beds of the Central Iberian Zone (Central Spain): Isabelella fascicostellata Reyes-Abril Villas gen. et sp. nov. It is similar to Horderleyella Bancroft, 1928 for its coarsely fascicostellate radial ornamentation and obtuse cardinal angles, although its convexoplane to convexoconcave profile allows discrimination from the typically dorsibiconvex Horderleyella. A phylogenetic analysis of the family places both Cacemia and Isabelella in basal positions of their clades, which fits with their early stratigraphic record. Based on our study, the family Harknessellidae appears to have originated in the high latitude Mediterranean margins of Gondwana during pre-Darriwilian times, before the detachment of Avalonia from Gondwana. The family reached its highest diversification in Avalonia throughout the Late Ordovician, keeping connections with the Mediterranean and Proto-Andean margins of Gondwana, as well as with the mid-latitude palaeocontinents of Baltica and South China.