Hake (Merluccius Rafinesque, 1810) and forkbeard (Phycis Walbaum, 1792) otoliths in the European Cenozoic – a review and evaluation in time and space

The gadiform genera Merluccius Rafinesque, 1810 and Phycis Walbaum, 1792 have rather different otolith morphologies. However, their otoliths share one feature in the fossil record: they are both dominated by small otolith specimens from putative juvenile fishes and otolith fragments. This effect has resulted in difficulties in the recognition of the diversity in the fossil record pertaining to the two genera. Here, I address these complexities by reviewing a substantial set of large Merluccius and Phycis otoliths. Eight species each are identified in the two genera beginning with the Oligocene. Six species are being described as new: Merluccius aequipar, M. kokeni, M. leptus, Phycis harzhauseri, P. pericarpathicus, and P. tortoniensis. The extinct genus Lagophycis, which was first mentioned by Gaemers in 1988, is defined here. Otoliths of the extant Merluccius, Phycis, and Urophycis Gill, 1863 species are documented, and the evolution of the genera Merluccius and Phycis in the European seas is outlined. The palaeobiogeographic history of the two genera is discussed in the context of the development of the geosphere and biosphere in Europe.