Exceptionally preserved hyolithids from the middle Cambrian of North China

Hyoliths are extinct enigmatic organisms of early lophotrochozoan affinity known globally from the Palaeozoic Era and were especially diverse and abundant in the Cambrian Period. However, the commonly incomplete preservation of hyolith exoskeletons and our limited knowledge of their soft anatomy makes their ecological and biological aspects unclear. Konservat-Lagerstatte are crucial windows to unlock the mysteries of hyoliths. Here we report a new occurrence of exceptionally preserved hyolithid hyoliths from the middle Cambrian Mantou Formation (Miaolingian, Wuliuan) in Shandong Province, North China. The preserved soft organs of the new species Novakotheca weifangensis sp. nov. include a U-shaped gut and possible pharynx, oesophagus, stomach and digestive gland, which provide significant new information for the reconstruction of the digestive system of hyolithids. Two taphonomic modes of hyoliths described herein are recognized: soft tissue preservation through pyritization and three-dimensional shell preservation through phosphatization. Morphological variations due to different preservational pathways in the same species are revealed, highlighting the taphonomic bias on taxonomy. The ecological association between hyoliths and small brachiopod epibionts is a direct example of species interactions, providing insights into the ecological structures and adaptability of early animals during Cambrian time.